Amptliflication Reage'rJrts

Re'deffning FastPCR
,Bfo-Rad's unique iProof'" f"Jigh·ffOOlity DNA. poiymemse cJeli'v€rs lJTIsurp@ssed fk!JeJity,'Mlh higher yields ~nd fewer rescUon f'aillJfes. riOt aJI your PCR needs,

spero amJ

!!II

Ex"l.ir€fne Sf:) Glee! - rflcr>eased proc.:oosMly dl"amattc allly reduces extsns ion ste,ps (to 15-30 sec.tkb) and overall run times (t'O 25-35 min) Unmatched nde'lity - Novel pn:lO'mading enzymo is the most accurate thermostable: potym6rrase (5:2-Iok:l mom ac:GLlr~e than Taq~
Minimal enzyme usage -

III

ill

Ellhlanoed proce:ss~vity €nablGls '!he use of less !3ru':yme (0.,25-1 U/reaction}

F'I"iISi!1gl1.i1e l:)yl:l!lH,MaflcJied DNA ~CI~ d protEin S:007d I!O iF'rooi po1y~ gj;ves,ii, a p.l\'A'lrtu~Sllidirl!til W oolhe iOOpllcaLOO IL»JA

F()( more infOrrr'l6atior'1, IJisit us on ~ha Wall at.

'lNW'W,b1o"raJd.ICOff'llladltiP!"OOfl

Visit US on the \Neb a! disoover.bfo-road.loofMl CBlI t.ollrree C1Jt 1-aOO"4Blo'RAD ("1-000"424-6723): outside th~ US.cootact yeur local sales,offtlc€"

No,w Availabll,e,~ R,I&DISy:s,tems, 2007 Catalog

1

Oner~ng MOire than '110,,000 pmdurts!

, Millitl~t~il: S~lI~ Sltoor:a.~ A , [Il1~A t:it$ >& ~>ge~~ , Ct!I·B.II~e!!I ~I<I~:& IEI.I5~ot A
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R&D Systems

R~'S.~U!rn,i ~ij[lt>~ ~ ~0l123S S2'»11!lo Inl'Oi!'.ltnOS)!'Stem~[OJilk;

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IEnoocrinol,ogy IImlmuno,lolgy frc)'~e,a.s.es, Nleuros(ience

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ItW 'SIj!$~!ru,E!:!rQ;p!ii

D,ev,ellop'lm,e'~t S:te'm (e,lIs Si'QirnaJlTransduct:~on G~y'oobioiogy

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!II!~~p.!!>I!'IJIj ~ "'I!!~"Q!I! ~T~~ ~PI"IIki>.

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Seletti on expan ding week~y-vi:sit www.RnDSystems.com

GE Heolthcore

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working with proteins b ~lieve in AKTA, UNICORN and wizards.

To 100,000 scienrists worldwJde, AKTA • sets the standard in protein purification. All systems in the AKTAd@sign'" f(Jrnil~ work with inoielligent UNl(ORN'M softwore, which rnckes It easy to CM trol ~e ry stoge 01 yOu r pu rificotion process. 131..1t we're never CMien l to Slond sti Il. The

result Ls products rike AKT~press"', vmich con solve low expresSIQi''1 and double-togged pn;aein pu rifi co tion Cno IIeli ges, 0 n d AM Apurifie r 0 time-so vi 1'19c utomo ted proteirl pu r~fI o lion c
H.

system then con be configured

to suit your personal cpplic(ltion

Of'LdwOfldlaw needs,

By cOI'lEinuolly d@veloping technology the! con turn your scienbfic idees. imo re<Jlily, we're bringmg science to liFe ond helplrLg trcrsform heclihccre, We corl it Protein SeparOlions 'WWw.gehealthco re,caml
~kl{l

~-imogined.

Discover (he Jegel1dary purlficcnon power of UNICORN and AKTA., VISit

C2l!106~,~!~C~IXIIJ)I·l\jI!!iiIH~~.
GE.li'!MI~~~ ~!i'c~rKli!!lti.l!.o ~i!!!I0o!f'i.o--= (~

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431

433
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440 441, 443 473

535 536

~~itoir~ .(h~u:e Contact SEie/l(,fl ~:arll:l1DrI1! S,am.pi~ Na1,ls,ma~~rs AAAS News &. Nol'e~
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C~ta~ytijli:Conn.ections

IL,E.TIE,R:S
rtIi:l'W(on gress Ma.y IBe Watrmil'lgt Up

to Pta n1~

444 445

fOIF fa ppirM] [llIli,~~km s ~ UNAt~-A MO<In Mis'5toll (j,elS;;IFI 'F'

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459

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445< 446

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Report Saas NSf' Prize~o Spur I'lI1lIl1loll!'ation

ue Balks

p,o bl~!iI1! wj:~h ~ ~ill1.1rs~tyH~ri~g if! S !)~~r! s

at (.amplJs" 1Mde Sa n on Tbh<l([o ,Mo~l'y fOIF R~~e,ju{1m

447
447

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n~e lor the TIe'.cI1l11

448

449

'C:l!Jd~boak
R,~dre:sh M'uhonthapjXJ OU'orl~t

462
463

nl~ Icndall1gtu::d
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NI!!WSIFOCUS
Lab Chimp'

450 Ull in
453

The God D:~~ILIi~ie:l'l R. O{Jwkil1$. revifiVi~ by M. Shermer

~l'i3trl'i:I Quagga MIJs.~nlJr:ns

W!21i't1i!rnUilited Slale-s

IEDUCATION

IFOIRUIM
464

n As!,roll(lIll1ic(l1 Sociely Me e'~jng Tlm~~Iii!!adoo qua~r Pramises !II Shed Li!iht ~I!IJ tljlniYl;lr~"5POlS'
rl!iidclle~Ealilih D<E'Ilil~nsMolllh!! Milly \!JIay

AI$lt(i.FilIIiiIifi~ttlid'e,~ 11'l1tii~icilILMlijzedi R!~;:!t:iiI'l91~filru.cticn ,f._ 1M, Con no!'" et at

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456 SNP~ .•Sihuit EltI(NQJtlnv!si!)lIl I Ali. KUmOf
>::> lIr:[X1rt

5oci!';ly rO:r:lfu~ral~\Ie and

B;i~t(igy~elil'!gl

466

Fllotei;f!ls, P,rlWi~ Sq,wjdt Wilh 1E:;:~!!Ul\i"i~ E)IIe:5i91iU Mu~i,(! IHbf!fS Slilih 1111(10 Iii [gM'h~"!r Whal~ Wfi I'I'I1i Sp&m I\ar:l.O:ri~.~

p. PS
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drl!llifil

467'

M. .Po Ba.I'dLIYin, P:. 8., R'/1iRf!S, Ii, -p:. Hl1orrg. M. f. Mf lniy,i:'

Tile .Mothe. of AU Stelll1i Ce!lro? /1, C 5pradJing ana Y. litem)
>:> R~P'Orf p, 518

467

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425

.1 Leu'gelf 'pof'Ho'io of IPlJlriifi'Clatiion Ipr,olooolls,

• • Plu:g-and-p'la,y slpin-c,olu.m!ll' Qutomatiion wii'lh Q,tA'cube Low- to lh'iglh,-thlfOl!Shpul'Clwtoma'tilon
S:YS,"IeI1IS"

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WVi/w.sCqencexpress. org, PHIYSICS 'Ql!lanttll'fl'l H,;II~ur~Lt~n Polarr OKlilie H],e'~m5:~rU!ttme5, Jl Tsukmaki f,t a L Th~ qU~'lIIlUm Hilil dfl1d', IJISI1Idll~~WJ1 ill ~ffili(Qndlud!Jr5"us, OOw <ilisOSll@ll ~n .a Ilayeil1d,lqn(-magIllfMlIIlI oil:nde. 10. 1126i.scienct!. ] n71130 CEU, BlOLOG,Y
AM,cmlb.r<1l1ll1ll Ri(!(;I!!Jilor
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CONTENTS

MIEDiCllNE

Cadherlil"l~llinr SYilo'liial LilrlilflglForTllatiolil it M. leI! f.?t el,

and Patho'lo.Qy 1m! rthritl,$ A

A mouse ~ersiOril <of rheu l'I'I'Olitoi,d<l!~trrlfitis cOln be amelijof<lll1'd b~ 'iflh ibiiti,o I'! or ,~liminlation or a (,@ll!>IJ!'face ,il(Hll~~ionI'll otfful»!' found wi'~nin j;0iIlIS. ~lJgg!1'$'1in9 ,a llilfrap~llI~i( approach ror litumalils.

10.11261sd~m;e.tB7306 Prohlin M.e,ciliahls Cl!llLJlalr

A p~~tefll 13 62 <'ILl BmCHiElMlS'M"l' EmlU!laitfln'9 MellIibr.iine Pf")l:e'irlIE.lj1o~L!tiolrl~y Ra,tiona~ D'~i9n' M. Rapp, S. Sep'piilii, E. Gfar:lsel'h .. ,G. VOIJ Hc1jlil!! 'Stell~ise mod ifk<!i!io~5 tlJ i! dru!lj efflul;I; pump stlow h@~ theoocteilial dillleri!: PU!!1IP rfllily h~~ ~ol'l'(ld~ihfQU'.gh d~p~kO!'li~ <Inti fnversiOIll of ili1J iI~~sl!1! llPr(1!ein d~mi!i n,

Uptake

R. ,K:wiof}'Uchi e~at
A rn~mbr<!n:e ~ll1:~I~ wi~ha prEViously Ulilkll!ow!'l h!'"ttloo bind'sthe,cOIJllier ~Qf villlmin/t uu~s ~>elei'!~ ,o~the 'o'i!~mrn. arnod lro1~j)Ofts it into ~U~,

Qr Villamim

10.112 6ISciellce.]

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IBREVIA.
GoiITI memt en ~1E\lolllJti ~l'Iary Pa~ns Ul"ltI'edyin gl Flo~\ler (olorVlU'iat1~:!:rI ill ,Antiffhirmm ~ oM .. D. Ramller
frJiJ r~l
Q~

EVOILUT~ON

ECOLOGY

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(j JfI

d' If ~lJ!e~,~)(@ 1fI('~ Si'Q ma'ls! mJ

481

J WI If!1!P~119 Spidhl.lrs 1\[ L M, Urn, AIt- F. to/it( D" ~{

llIltfa"'io~.e~, m.g hl i 5 rEiqU Ered l'Qr c))unsl1ip in !>lOme ~Ulmp,in9' spl!llers; I'lt(It~s rWil]ni~i! lemOiles' UlV'.~lki1ed nu(mm.enu; ~emale5 rfl:og11li1~e ma ~s by IUV"r,~I,~dti,1{~!P'<!!()ll s Ofll tl'\@ir oodi~~" ~

ww.:\HdefKfffl/Jg.1Jf!JfffjilrlJIlU~llIfulllJ

151581114 61a

Re-sponse ,~~, Olnment on "EvolutiMary !Paths C Un;d'e~ ng "lowe r Color variatlmilinl Ai!1UrmiMm" ~.yi II. Co Whibley et a~.
faU fM m iWfm sr:irHrremo!l.6'9'{~ilwnretJVfut1l31.515S1114 rHo

IRE,$IE.ARCH AIRiIICL.E.
DEVELOPMENTAL 'BIOl[J(iY IHlst:ocol1ll plll~iboo Emlbrvonri(' SUll1ll C@llls, 482 by lP.1!rU'Htrlo~~ne~ls KiWi Kim Ell a l. ~YS!li! em bcry@slholt d'~eto~ by p~,!1he!10geDi!'sis (,,1'1 ~ J WUfro 'fJl !li!1!IiIbryQni1; st~m ,c@l~, immYI'1;(lIogicol1ly com!li~tubk witll the d@lMJf~

R'IEV~leW'
MJCROBJOLOGV Recomb. rloIlliQriJami! th,~ IMatllJre 'of

Ba,cleria I Sl)edi1!t;il!Jfi' (. rrYlser. W. Po NCI1I;JfW, 8. G. Sprmt

476

IRIE.P10RTS,
MATiERiAl.,s SOIE:NCE IRe1Ilmi~'blleSwih;:lhilllg of Hydlr'G)Qe~~Adu(lt.ed 1Ma no $ltUJl(ltulr~ bno CC!I1lPW'lI, Mr(J~'p.!iU~n'ils

487

A. ShIOI'€r.ko

$iii CQO n,lf1IocollJ rnf!S !li!mbed'\ied ~n O! pg,1yme'f hydrogel fapidt~ cha F!ge' tt.eir til!: in re5P()lnw tQ chOll1i'!Jes[n humidiw, ~QmJIl!ii!:l:p~ems C~.11i be m~ d'e b¥ a dijlJ~inrg th~ ~Im~ fMd r)~ Ih~ g~ L M,PUED PHVSICS IE~E!ct~orne(halntGail es(l!llau)'r~ from G!"aph~ni! She@t$ 49.0 R j, S. 8tJ1flCh eu~. A gr.aphell!li! stJoo~ QneOlt·om thick,. slJl5~l'1ded <!1U(l;~ i:I r!l\!n~Coll!! l~~lil(h. wm b~ i11l:lu.;~d to vi bra.t~ {I,p'lkillty ~,ooetmCllrk(llly, prQll'idingl Um U:lrilDilmlwtl-dimm!si{jna,llr~wna1.t1r. C:HEMISVAY

~:lifIpr!l)r~oo OXY.ll~iIil Redl:.l>C.Uol"! i1:·t'illih~ Oli), ~t~m(ilU} A vjaJ Illnf;I'l~$>ed:SUlrraoe S,he A.lla1.labiUly v. R. Slulnenkewic e l at.

493

TIn~ IPh! n rkbed outer. su rifero la~1N o:r UII! dQ~~-pack.oo1 il) 'Suf~~e' O has alii altered e'l~clJ(l11Ii( ~l.Ju(EuJfe lilat.la,wfs ();l: <lid ~JPtl0n (I1,r.er s,ptcl'eSi1J ell as OH.

427

Science
R:IEpiOR1S,
CONTINUED ...
NT At t, 1B:IOlCl GY As)!1iI'I etric ~nherit,lnIu! 'o~ Mo~her\i!er.sL!lS IIi)i!!,gihte,r m 51 B (en~r®solilile! iin5tern C~IIIDi'!li'sion Y. M. Yal.llosbi!'o, A. P. MQfmrt~id,). R, Pellin. M. 1. furler liii li.h'E! ,ro<!ess 0'1 prllc:llicing dilferei'illiatii'lg p ila,~h'tm, li1e mofher C!'~lrosome of DroJop,.f'iil,(J m~le g,erm ,ce~ls is,!!!JU:M1ed Io'laa lIlitroWblJlea fr.!Y near the fl'idua. ll~aoOOd 1'!9 <iSllm~etric diYisioill, D:EVElOI?MIE
>~ PBr;Mtli~te- Jj6~

CONTENTS

CHEMISTRY Water CCltii!lY!i~S !Dr aJ IFlad'ical·M(J:le~ 1I1i(!! 497 Ga.s~Phas>~IRea.ct!>ol1 E Vi,ihfiflger·M.(Jrrirrel' et .1IL M~Ylfli!l"Inenl~ 01'rt';1lcti[)nl mw!O1L~dsimi!IL~li:®M ~O!,N~hi!t wtIl~'l!'r m®l@cuWS(<ln<lIt as (,a!:al~~!i ill 1.IiJ~ gi;ls j}:tMsl!and may pntigpatI! ill [DII'i'-~(lm fl@mlure ~BilI(~On5ill tM atmuipMre"

:>" FffJ{J&WtnJ.

47()

PLANETARY SCIENCE: M~rtiallil Atlll'losplil,~rt( E~o~1.Gl,nRo:irie5 :5:, J3ambosh el al, Mea5uJel"lilents from Mar5IE~1~SS ~h[J'fI mat. the 'sular 'llriml is f~m01iltn9 omy iii ~lIIiIall amC!unt o~ ,Mi!llr~' almo"pn~rl!', imJI~ying thai. ~he formally all'IlJlild.an'~IH~O~OO'C:O~rems i"iIiI underground. GEOPHYSICS

:501

D:EVELOPMIE f'ITALBIOUJ IKinf'llh:s M()lrlJ~Olg~'1IJ A, Kicheva .1! I.

or

GY

Gradtl?rlt F(m1f!ati(l~

521

I~~

Tine ~ij n~tic pa rameters ~h~I:e5~a Dti~h and mOltintairn morphog;entK !ijtlOldil'flts",rlO: d~li1liedl.md mll?tIslllred 'lor ~W() ~ey m(llfphog:ells il'l Dro.mph''/(}, 503
GENETICS

S~e\" Earlih(l ~.aJki!!'S GtliflC:I'Qh!nt witih EIl]wdi Ii: 'Tr@ll!lors

a~,d Slow Sli~ IEv@lfJts y. IUJ, K Oboro. K. SI'lJomt S. xJiif}(;, It Hirose

A seriE!~ QI'we.lk ilClW"I,re!ljueriC}' !!ii!nn~y~kes, 'Qf ~Es:p!lnd ~th ~'eismltc tre nl(lr j)ntd Sili!) t!?p~~des Qn ~ $ubodI!ld~n~i)ne' benlNllh l\ilp~ n. p~,rhil ~ in.U'w!;in~ av!2r<l~1 'S\f@!iS, J!>~ P!'!Ipm~ "" .>171 OCEAN SCiENCE

A ~Sile'l1It" P~I.ymo;rp,i;risITrl ~f1 the MORt Gene 525 Ch<!rr1ges Substrat.e' Spet:Hidty (, Kimrlli·SQl!a1r el aL A ril~e,bnIt 5ijIiiO:i'I~ii'iOi:JS. ((;t!-,olli in allillM oJ a ,d!rug·refl'!ta~i:!e!ii~ne' calli c;hang,e uai'i~~l.ion ~in,el~C$.i!lM so produce' a ((iii'rtormMionillUy
dis:l'iilKt~rateii'l

s:pu~e~,~~ Pf~«ff~

p. 466

S~ro:l'!gRelatio1f!lshif:) ~til!,U'ffl,e!1t [l'M's'i'lod the Sol~r R:~di<iilli(l~Oo:se Qlj~r~he Gl(llb~l $ i,lf'~9I(eO~'~ ) .. M 'V(lrJina {}rJ'rj R, SimQ
fhUtl'lilQullIt ,(]If drm~,thyb~lidQ
ill

506
Ii!

IMMUNDtl-OGY

pmdULQd by m~1il1l!i!®rgant~InS oo[~

W form doi!OCl~!Jut d~pend15on i.In@ lImOOlllihl'f [oc<ll. $t1111i9h~, bmingl n~aii7l'e~oodbad

Illllilgin'g of! Gerlilllili1al tCe'ntte,r Se~e(tion Even't~ 101Irilllgi A~Hnnity MlIItu ra lion C II C. AJlen; t. Okod(!, H. l. :nmg. j. G. Cysl'ef "",ithin ~mpht firlctes:. the tCelh are hi911ly ~olJ.ile' OLnd seem (()r help If()m olher immwne ce ~s..
NEUROSC~EN C!E

52B

s~Si:em.

Ourj!il!i se~Ktio!l of i~muF!'e ,cells 'ttia'l !TN:!~e higtNlfinily alltiboodies 'to! compete

MICROBIOLOGY

A ft.al~illlilnogeiil·.A!ti1;fatlng Protea,se' SlledricaUV 509 COIil~roh the IDe'!ol'e'I.!),~ment ,oi IPrimh!llrY P~,elJnHmlk INil'gue IW' W. latliem, P. A. Price, II:: L MiHef" w: E. Golt/mull IP~ag!JIe ilcmrra cant infecD. the 1JJll9Jbees u~il!of '~111!' b iPr~~~nle
(!II il5:pe,cil i:c proI.e8GJ!?,. wlflilh pro!J.illl~ h!?ll!l5i~ol.illliitii~ host (ie'lenses9!1o may 00 iltlJ 5efiUllherapeutlc ta rget. A. Vir us ]111a IFllmglus,j n 11 N<lllIt: lFhree':W,atv .sYrIllbiosi:s . :s 13 IlI~Nltliredl for n'~rr'llli'lil oieroll1lC!e T L. M..Mcif'tjlilill':l, R. :5. ,/leliman, R. .J. RodrifjlJflZ, M . .J, Roos,sinck A 11 eat·t~eral1t gra5,s in 65~ 'l'eUDi,',i~!.>orul· soil!: (~i'I grclI.l ol1lly ~u!nl jl1i'Kbedi boy a hmgu~ ~hi"h is,protetti't~ Gilly when ~I it'Sel~ is irdect'ed, b~a myCllJ'o'i NS'. Whe Ne lim~~Ba5~~ ,of toss Ave ~$;iQiill!il S, 5 li]te(isiiil!il·.iMaki~g Urli:!;er Rislk 5, oM. :rom; C R. fOx, c. r.~P'E'~ R., Ii. Po/drGck OtrorllilpF'i,ng brain lI~twlJi[b ro~oncllTl!'J~ to g,jJmhling [1)S'm lRilm to ~ail1~, (orl1! liiili1li9 witlii b@b!lliioril[ o !m!'IViI ion 5 ,ailw~.tni:sk.a~tf~i®ll. I
NEUROSCliE NeE MICROB:IClLOGV

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EXTR:EMIE PR:E!CISUJlN wmrH UN PAIRAI!.IL.ElIEDSIPIEE[I ,A!\![1 ROIBUSTNES:S
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ci'~i(e.rrec~s.,silIchas col,lisione.l aC:li~ UOr'l, dirert p,8lrticipalion of wOller as a catalyst. V'Qhrhfl @)eIH~,lJjrUf]eg~U Qr.
(p', The Jole of VI.'a'le ~ in maf1lV a! mQ:SIP~eril!: Ir~ ~li~ll~ r'e~h1S. poorily lUf!der:st(lod at the In'lo~e(U~lr Iev~t bela Ulse it is d~~WkuLt. d~~tnnglill ~ilno:nsp eto i

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Early UriHli hi.'!ltory, M.m ~\\aiS OIlC@ 'I!«!,t and hlJlmid all'!d had a deMeJ' atn10sphe;r,e lh QiFl it na,s LO~y.

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llris at!TW~ilhere· wars $uppoo.sed ly ll:iattt~N~dby ~hil!

:5>Glcllrwund: nd Io'>l iI1la space. S,liIralbaslil a ol ~lp, (1:lJ lj rid tl1i!ll the' I:lStil'po\l f~t~l,Qdav [Q[ "'OIl~li 5 in the rrna,r1IiaII1i allll10isphere is \!'~r'l"lQ\l\'. 'bll~ed on measul"~m ellt~ffHJ'm lIhe oribilingl Mars B:pr€55 S!pa(ecr(l~L Prol1~.gating the,;;!! raJl.!i!~O<lci: mrd over a [period (lr 3.5 GY'!'!,Io'lltd re 5!1 II ill the relitIDI(all Oi~' 0.2: tQ 4 mbar {If (0,2 anda r~ centimel'e~ OIrwLer. Ra~iler I han ~avhlllg 1~ltthe'lll!alm,et,(02i3!nd ~\fal<8\F rd I c:o~ in ~teoo be tocked away Imlil~all'nitS5!Ulrf:;tc'~,
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l,o waMr vapor, the nenocelumns are lined up and s~lld II P (In tne SlI.Irlfece. This. ac'tq"lationl ij~
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ord,e:r or ~emnds) 1!lmdre'ller5ib!e. HVii' hydrlJg~l (an ~lso be ~e!1!)plgM.!dI so thQit ~~~ motion of till e silicon nan ecelu 11lI1m~ (ICC U rs 111 ill

'wul.n qjua ntum

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tiens til implij(.H~ a (il!l~l"tk role fo:r individu a~lM;Ilt~r ml}lec:~ ~!t'5.1lil1e gals-pha:5e n reaction of OH radi(~~) \~ itn a,oe't.a ~~ F!hyd,(!',il1 ~~r Je~llU:s sugg~s.l ~hat wale', c:o:l1'lpl'~XQliol'ltOo

[jj

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or such

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liillere h mn-sirierable UI1(ertll~my .about wha~ oonsljl!yt@,s.1! sp,e:.ci@"S Ifor iJacli@:nia b@c(!us<!? ,orlhl! <llp,lJlIrenrly ,~:uhtll1g;e Ifla~ocur:s be'l\veell! mkroOa"gaI1i5lrns. whii(~ :S'@'!llm to rre~utl. 1m mkmbiaII]OIl!UlOiticlI1:; beill1g
ralillp,il nt

i '~ 1.0119 periods of tens

lrremo:r, klnlg· peri od volGal1i{ fVeml!S, and ~w€'t al. (p, 503, pu bl ish sd ~ oll[flil'~ 30 Nlo'l,l'emb;!rr; see l~e l?erspecUve by ~ Dlia!ge!'~) ~d:ef!ltnry yet anot~er type of s,e'is.m~( ~ r~'iI,erberration. ~hiltor venl·lo'i'!.l·~ir,eQlUi!iuy evoents ~ ~ulh 'eq'lIiv~le!1l m~gl':l.itlid~ ~f 3. '10 3.5 \ii1!1d
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.~ wi:[h ~Qlwol'ci:lIlictn~ml;lr (InC. ~~O~\l P 'eve I'll's,. Soli g, nl,e ~O,ifl(:idefl(e Q~ thes!e ~hr~ pheil1Qmena imWllrvt1; the dtl:ed~on o1ll~d, ,har(!cteriila~.iolll of 'uo ~low ea,rl~'qI.Hlkes, 'ilIhkh ar\€' thought «I) ifLtrea5e

res;onator vi Ibra les in reO;;j:l0m;e t.O din a p!l~ied !,;d~~al farc~. Ci!llldmakin~I..il1 r~oriat~Jj" 'th~I1m1~r a!illd le~ ma!>~~w 'gen \?ra ~I.y ~.ause ~t o QflI~rale 'I'oIi~l L !:It li1iglhe{ r~(jIJ~luies and havoe bette r se IlSJli 'iitll'" B,g rncl1ilet '(po 490~ e;.d\lliated IJlI.tri3thinshel'~:5 o'~'grraphene homgrapllite and suspeFlded ~he hm·di lIIlellisiiona~lay.en <lUOS:5lremn . They then a

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"lel'id~.Fr"ser e'l 0.1', (~7'6)hH,1IS. Oil, the disc Iributqon arid e'f~e,el ,af Illllltlltiol'l an d U!oombil'la· 'km hqu ~Iildt's, and, by combinirqg a nmoe ~in9 alpJPro.=!(nl wilh 11rllview of t~t t};i~tin9 dala, re~1 the C;:Q'nditij(lI1lSoII"lillderwhk:~ ba,cleriaL spoo·
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• Throughput

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-1-ligll1 . .'j~ins~C'-r~~ld'~I,(ura'Y ,gn::ah;'[ than 99.5:t}h ov~r2nO bases

• Consensus-read accuracy gl'e~1 than 99'.9'IJC}f~ tcr

Visit www:.,genome,--seqHleil',JC;IJg'.()O',in to learn ubout the t:xP<ll~TI ng, nu mber o{p0cr-re'l" iewed p II.Ih~k;;~ s ppeari ng \veek~.y. di tkm

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Diagnostics
Roche Di:agnO:Mics GI'iI1lbH AoctlC Appllioo Science
68298 M~Mheim Gellrti ... lI:y

~Hi.lOMIL SED;lJEN~R _."~,IJI' ~~ lie ~"".~l!Hn ~~n1 ,[;1 U:i'!. ~i1Il

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lfis5Ue'$ d(trriV~dI rr(l.m embryOflh: :Sliem «(tI~'Srnay p:r,ove lherapeul~ki'lUy Ur~(thd•. blJ~ a Ink€ly problem li\iill[ b'l?~ih@ n!j«li~n of stem (@lll5 thiU C.1:! S!!J~fa(@' a IfIlig ens di'.ssilrni htllt'llrhoS@ ~f 'th@ reci p i~f!t, In rry ~aw order to (reate emb nyoll1lksle m {ell llines lhat (arry ~1'lat(hed surf.;!{~ Q llltig,en~,K.im el at '(p, til82" publijshed onl~l1I~ 1<1 De'cltllllbillr;:fi!e tl1@'"t:)lI~~) I~v@! analy.-ted pl'!rth@flo9:~m:rUc d@rivalilJln or ,~mbJyIOnk nem ,ceb il'l ~Ile mO!:l~~. U~fl9 {lor;ytes h(l~n t~'II0sep::lIr,au~ phases ~I mitosis, they llf11du(.ed pa rln'!2inogel18't.i(. dllli'E!lJlpm,enl iii! wh kh lhe tells Co'! r~y t.WCI copies o:~'~h8' l mat!m~ I g,eno(iJ1J8'. Stem e@ l~s. d'E1,ri'llloo hum Ulli!!~eemblY0:5 I,'llolJlld bea~pedrk match ler ~he d enor ,of thl1oo(y~. Such ptl rlhene'\Jer!@lkaH¥ de Fi\ltd :it~DI1ulls oolJ'ld ilf~dQ,J:C~T!ilflY,but !lot atl, lir~!,J:~S. iJ ~s:"S,bly b~(a~ s.eof the I illbseme Gr the p.a1~ernaHy imprinted geno:me ..

Pla g uin 9 the Lu ngls

I? ~11Ig'ueis not j !J.!s.l r,ul~m,i t Ued! [lly rb:li!lf,; hy~er\lir!J.!bm~. P neumon lc I1l~g ~H~can ItlE! tran ~l1IliUoo di IT!K~ily ,almOFig ~erOi"~e by wllghil!lg .•LiiiI'thlE:!I11I,et lit" {p.5n~nshQ'l'!,i t~at a specHi.c. v;jrLJlenrCe' fact,or rrom Yel':~ifliopflsfi£, pli!sl1l~lnog,en o'lcti,vtllm, is illlie(lif2din~.J1l<host (el~s.1!nd pmmot!!'S. p,mlifertlltiolll nl the b<l,delFia and n1l8~r;ive lUlIlg in~Lammatl{1ln. n ~spossilbie that ~hir5,baeterial p,rolea~e convens hest p ~;:mJfliIllQg ~1fiI int,C) plaSIiI'llli'1. 'th@tE~)' r@i@as~g~lf'app@d ba ~l€'r ia ho m t ~brl n (~QK Thus, in hib ~li fig n P ~[lsmi 1'10gelili .iutiVdilor d uri ng therapy mig h l S~O\lll di.SletllS;e pr{l£lr,~r;~ilolili [ll'ldlllkiw nli b tot tCS ti m.e ~o ~.!1!f!:1! !n,~t

Stop Smoki ng
lfhe ~,(tgforloi t h~ b r~i n G~Lh::dth e insu la ~,a!llreooi\f~dllrel~tiv'€ly Uule attentien iln dmg addktio'n ~H,eratllre.IN'HilI!ld 'E'hd. (p. 531) nnw r~port ~.h@ f@!lutts of .1! r~nOs,p~ct~l.I@ ,jiln:ato mi I!:€!L~I1€!llysisof a lri!,r,g~ coBfNlfl or brain-damag;l?d ~,ti@iflts" 1ihe aflal.y)is ~\las promptf!d' lJylh@firQb)erVilItiOfil(llf~ sifllgle Pc~lielrll ~vi~hil1isul~1an~age,,,,,,hQ qUlii~ a ~e"lIere Oi(otM1E" d .alddi(~ionimme,diat!~ly upen rewvery hom his .acute! lIl11unlID9iCi!~ danIl,a9 e wit hout any app3 r.el1:t di Ukta ~lY0 rrrellaipse. Damage to ~he ]~sol,p ,alppears to rlEtdlJc:e'tin", mg,!!! to smoke rath e-r Ult1n, ~or ualillple, redlJ!:illlglhe H'r..rard m reil'lro:r,C!emenit aSliodaled wil.h '~mJj)~king. '

Thrlee-Way Partuership
A three ·'i'I}ay en dO.5.ymbi(l5is iml'oJlving a gra~s, il [1I,UIl gu,s~i;ll1da virus ~as be en di~(mre;ed in the hot s~!n:ts of ¥e~li()\'M'Stol'le INiiliol\g,II~.a rk by M~.rqUI'E:z et 01. '(p" 5 Bl. A vi railly' infec'ted i~mg illS; inf,e<ts.~i1e .f"lQ~t.g.ri'!:5~·s rooh and mn'len; heal ~olerame to both glF.~s and fungl!!!s. FU'f~lhermlll,re .• it us the virus in ~,htl'fU1n-gY!l~~C!a!. allD~~s it ~~ eifl(:!<lW lhiis. 't(l[era~c.e'. ~ ung1us lad:iililg U1.-el,Iin.l~ not glrovide he~t resisidid time ~o its h OS[ 'gra ss I!In:tn L'iI. wa~ rei n(,edBd.

HWI,ds,g'II1I~.AlIP,ha Inst~tlUlte fo,[r l,ii,otecllilmJ olLo'gy 18ri' n 9i ngl i nd i\i' nd u ali ze d t herapi

es fo!" diseases by d'evelopnng tnighthrouqhput rasesrch toots" For further details about HAlE, \I,isil www.h,;ij lib .or'~

H €!·,adis, I Wmn
o rm of the end uri ng ob~!ml'a!iom '~Il5!UliJlPJjJof IUIITl'il nirr a'HQrJalily i~ lMi'iIl are d1l:spropJ1l.nti rt tha orn,Le<ly s'ensiUve t~ lesses reWli;,retQ g~i!ns, W~'eFi o'~~ered \;I )O·~O g~~ble, the gJo~e~lia'lgai!n needs ~o be
lwiceas lalrgJll as the 1l9tnlia'll.oss. in ordewFQF slJbjed!i to IlxhibH an equtl.1 ~vilUn~l'Iess. t'OCi([ept Of to rejel:t. the !;I'tI m'b le, Tom ,et ,t'li. (p. '.)·1'J,) Ihave map ped 11 s.elof brain regions. that rer.pmndi po rametrically to I,he :lii!i!!S ~,!' Olt~l1tr~l9~~lrtS amidlosses, an d :shO~\i th a~~ih@s;@ reQli:ii)m!:!r~ m~n.· S@Ifls,itiv!! to P losses, Wurth ermOrl:!', the belwe en-slLlbjeCl d inie~e;n.:es. flm neu r<lli.respo nses r'e'!le(ll~,e be~we;en ~5'n1bje(l d ~rrer@im:.E! sin t n@i r b@hav~oJi!l ,lJiv@rsiollJt(), lasses.

Operon'5
Q

5!'rV~(P5

~nclude:

[Hi.glt'i-thiFOi;ligltiput 0[1igllo nUl deat ide :syrnth esls .AROSu.,

.Arr,f,I'lI' Fl:eady ClI~m,go!Sets

Inside IB (en Central.

B c'~lh be,c:01lfle· eflhwive rai::t(l~ies.~Cll!"::I!'!tibO<dy prodli(liiOmJ o:rdy ahe~ they 11~~~ gorne tlH01,lg!~ a :5'~ri~~0 r lIIl tltu rati on 5tep~ that 5e~e(~.den es ,of B (,~115{llIrryin,g ~mi1!'ti ( m u t.a lID li1I~ for hi g[ll-aUi nily ~ ,;jli'ilib~die~. Thi!s PJOO~!isti! ~~~ IP~ ci!fii!l lh'e ~erl!Ti Ceii'ief', ~\1 h~f'e B (.ellure also tli olUlgM ~.o ('0 im. ~ p~t~ vi 9ft! ~o!J~IV for avalla b le anli91!in, U~in9 j 1Il~lFavita~ IIIl i(:rwA;iIJlPY, AU~'P.1et or, (p. 52 S·; p:ublh h,ed ~ (I ~liflie 21 [)~'oembe'r) ~h$e~ 'tha tlhe benQlviQ r or 9 errrnnna I'cenie ~ B celis. ~s m or~ ((Ili1Is.i~ten~with a ti tom Il!lllitkm ror the ,('Itl'ent~on h e-~I]Elrl' QIi1Ill!Ihil n for~,a rc;e il I1Ug ~n. lihft~ fi IIlIGingl CDuld pNI'II\~ u~,~.~ fu l imJ co nSlde'rl Fig hO'l'l.t be:sUo 'sti'U!lulate uo bu 5.t imlf1i"lun e reSPQflJ5es wit~ 'valcdflle$,

IOIP,Arra/y'l\'I lP're~[prir!lt,edl Microalrrrarys

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www.Olteron.com

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QOilIr;tal1ies.

iII\cmica lB:radIQr<I.l~ ~'h~ ~,ecudve edlitor or Sciffi.f.

WE ALIL KNOW THAT Al C:ONFERENICE5. THE REAl UCHANG E OF IDEAS HAIPIPEN5 IOU RING d~l;.' reaksaadmeals. Throl~ghiur~lmli~~c~1m~l~~mk:iiJti{Jn.S. b ,,\,~di~vv~r Or)fi'ilmo:~bo!~d~ ~ml
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ftUI;1ilr~ 'ci;)lh~bm;;uli!Jn, Thelaternarioaal

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OfW!JI¥Ii!;!'11

and El1guneeri:rag 11m Kuw;<1h this, month, whjch bmugjlmt logQthel· ,Arollb scienti srs and eaugig,l«U'~ from the Middl,r; Em';I,1JIul North Africa {M E NA} with u.s, women '~old~ng [:j.imlhHpo,5.ili.{)~ilS.wa:s all efforl [.0 :sp~m. sueh cermecrions, Ag~im;:t the hm::kdmp of du~~re.sent pol it'OC:'[I~ l inmw:.c::h,uge(11W:~thc()nce[ll:Sabout It::l,'mrUs.!Ill. nuc.letll: p:D'O~i c fe:l:<l~ion. Ollild economic \lu~llera1)i!ily .. 1 W<'lS ned to see business C:::<III:d:s bei:L1;gswapped a!.an .aswni.sh~ng mle. tiui The r~dlllt;::.;l'.O'I!1idim';llrity onl1ll;~~ il'!lt~nie~hm!' s pl!\!vidGd t.-.ng~bl~~vid:l(!~~ur~~~CllHdy,ic ~1T~Cl ~11l:l[ \!o'Orn.en could hmfC OJ:lt~~.eregli.cm. The conference "v,,~(~f:'Jl1m,ized the KWI~1j,"1 fbumlmioII by i~ for ~.~.e- (hf'.ilnOOim:m~lfSc~l;lt~.C'e. the A KlI\\,'<Ii.l. lnsdune for Scienti:Wic Re.Iie<lrclh,tllcArnb Fund lor :&';oml.o:m:ics and Social Devclo;pmenl.,
lua Sclenoe.ledulOlogy d~e American All$(lCi~liml fol' I.hit Ad\l~Il.<'!;;liii1~l'1tef'Seience (publisher ofSciellce). ;nll,d the u.s. Deperuneru OrS~<LI.e. Iil.~~.d~" n~l~pklGs; ~)f~h~prime!m~.lis.h;,:rof Kt~w i~,.A C;:~fqltn'll~ ~hc .. p~lrpOse of~lin.e1,1.1{!ctiilllg 'I'!'<IS~DIDsH:r~eadersh:ip skins b:y ereatung Ilelw'Od,;sanliloM_g female E>del1i!.:ists in the regkll:l and betwees these scientists and U.S. \\iUn1LeTi.A'S ¥.t:'n!lIril.l A. AI mlllYiI.m.k}\ It\w~j ri, ~nl p'ide:n~ !o.rogisl.~\OIn Sandli Arabia q
';OUF .nK-rep~n;f. .. 'together there, Cre<Ui~dll i.i~oglie e d ~11!'ll \vil.l!!'ty dO\\Ji!1 the fo~mootim'i!l for Mum cojJ~iboJC:Mions~" CI~~ri.y.[~\,fib~orum cummu~ily (If \!;'(l!l1i~rl $!cu~!l!lisuis eJ.nergi 11~i:lm.he region. At.the OO:l1fell.'enoe.mmily Ar.,.D '\\l'Ome~,l t

Leaders

wrore.

mtll $i;;£.~~ln~]~ euum~rp~:rl$ fUi! Ihe fifl'T lhVle'.As personal ::;,~mies\V('re $~..red, J$-e!l~lt'dlb~Jt~nf)Js qlll.icb:~ V'd~lislli.ed" Sorae oft~e auerdees 'wt~re ilu:erl]gtiCl~~llly];::uao\!"~~. hereas (M:hel'SII~.,d~.,we!f'ld w {)I~tside '[heir h~1!i.ne C(miiltliY fm' th6Firl1)ll,in.'!Ie: !\i~1Lne \l,~re <I\~im-gmtre,,000heitSlt.ii.dil~om~ I: i~l)f ~ had ohi:l!drelllbefore p~n~mi:l,g dl.eir deg~«::!'~rorn.e h~~dreanned honm,el,ropil1lg to make a di ffi;r,enoe anleJ YC~I.'S Oililrnfid \\"~1ilc others had fQr~fl:dO[lpC!rl:l.u~it~.C's lhdr (~\Vll 'co~ml]'ics. Vel ~nrclTl;lliml.C'd in fOCl~S!,"\I on makinl£.H C!DIlt1l'ilJln'io~,I]]jmugJ1,ll'ile~Jr w\JJik even \\i.heil,l societal IKl:I:'Um.15 p:~1I!;'leilled OOOI~~' cles, C01!illidJ1;llt a:lild fU·'!UClUlfLlG. these \liI'(m~.cnlUldl;lffil~Uld 'tlh;!!l S(~ci(l~'GOlfmomfJ; de\l'~~lopml;'lnilof [he region, dCl~lds o.m gJobal (.~pHcity:·i)(Lil~din~ aad 5l1<iJ:eg~c~nvestmelll .in science and technology. Hefore the meetirng •. ll1yo\~FJ;l i:~mlge ofAmb women W~IS ofa:ua, oppressed and mm·.w.ll!A~ize,d g,w!!JP,Yt'~ SI:,llist~Jl;S prre.\H,l:I'l~,dduring themeet i[lg by Safl'ilJi bl.ui!lll pi~ intedamore rmlJ11Ced ra picture .. Drm"rimlg:~m a 200.5 Un~~ed Nm ions :[,duC<IIIIOliliEll, Sc i~nli fic and Cu~lmi!J Ol·g.:~m f !2:m!I.lon repen; she lloted~:~'l[ 74% gf seience gmduoi~in HOlhf<lin arewomen. R." ,eornpared to 4.3% ~n
Dl~.cthmiLcd SUlU~s.. Unfo.rl.~I.~l~tdy. this pnti:i[ivc ~mI{JUC~'IlOI' O:WS(.':lby I.~U: orld B~uk. Group is W G~!l(h;(S~!uS dat~ fgr2:0M. t'!l'hk~h pt'iint (J.ut thn[\MOln~~~!U~l~,('liJ;p:lIi!ly 19% of lilt;': h)lid~abtlr o force in, lB.d'ir:Ji.nl.\\~Jil~el~.ey OI(;CO'LII~t .:j&t~ the Un:Lled States, T~.lf~\i\brl.dBank sectorbrief for un

Gl:ndcr in MI'iNA f~)c~~;;; '(10 thi ~r.<~r.tdlt»; 1,nd~(Jt1:$ th~l!t"U!~likr: in (lthr;;:r Ngitms ,~:rhe world, r sigulifli,cmll progress :illlreducing gender gaps, in edncatieu and inlo\.\''eil:.ing fef~mly rates has not cm;.i~d 'tha:ougtl~mu ME.NA's l<:!oornltl;rk1;ll," These oh,~en'.l(iol'l:;; \',;ere buu.4: O~t~ by 5~n~'I..e f u d~eanendeesJor whom <I. scieuce e'l!t~l.c;ltion ~isill. re-<I~ity. burfull panici~lJ'<lliulii iml~he \~:ork fOliC!: is still elusive ..As the 200{)lntcrAcach:::lli1IY CO~IIH~~~ report. ~J'in'te.n}br Selena: SI:lTIitl£d"·~!.Ob(1l]1 c<lpm::iry b~l iiding ... is :ilmpo ssi b~,e\,\1 i~lml1.~ fu ~I. eng<Lg,ement of women m: lhegJ<l:s:sIL'OOt::;." [-I'OPL1f~ll1y. oonr~reua~ will sig!!l;;l~. to the r~~ionIJlfl[ [11(; lhlii!~ for Jull ~J'l:WlgCIm:~nlL"l I1m'l.'. n~i~ In :Sleppi]mc out of In}" Oi!i~'l,l comfQin zone 1iO~~ne:l.ldhe cQ{I,1fereIlGe. I canied some uaease .:~\'V~<LS, t note-ager to Uy ewer Baghdad snd feh a ~im:lledkllmvllilge ,)[1£nn the An.'ab \\,u:D'Jd. My dlli~.ywurld .~ an ed~l{lrprnvides im'l;lllec'liu,,~l:it imu~.~llion 1d~tm~ht "I-suITiciemle,'d gf e'L1~e·L~~em.8m : ,1m! tJl,e eyes ,of a Yt'!·IlIl,e.ua \VtllHatn-;,,'1U lh~u ~. eu ld see oil: .ber~!lhQ\l!I'ed me 1~:l'I[ there i1> Inll:L<'!: to i e
ergCII,J'l!litelt.

mid her gaze :t,pO:kevolume5o: a shared U1 il1i~ Ih:r kIl1O\,'!.Ilecig:e. prQ!l=SSIDJ.l.,.1 achievemem; [Inti the chanee 1.0 mal\;c a diffcl"Cl:'trlCoe, On the .m~i~ng·s iilst {lidlY, she ~oke' of herg:m,tilmle fOlr

'~.~~,e (,~mrr4;lre'l~~

;)\IiI,d.

nno:w it ll'ilfldliber.;~l~dh~r, H~rey~!'i. her presence•.h;;uJ \:~I~I<llly liht;;r.u~d

nUl and, no dO~llbt. o~lhers,

437'

EDITED, BY GIllBERT CH ~N AN D ]A,t(E YE5TON

c EU,

1):1OILOGY

Caught,hll 'lillie IN! E'

WIl Em(iln;ula'~iJng mmtrophils !il re (1haIIlelll~led Iby mierebes, trn'eJ GlI'I~ra 1iI~~'erlil1l th:~!1!1'SIllh'~, ~f1t,(l ~'~lJtoo,phi l il'~tr.a,~~'lb,d,~ r traps. er I~ ETs, vfhkh a ~e.il:1)1f!sulJct;ed Oillt of dm:II"111i1UIli Urom tme l'Iudeus} and gram!l~e proteil1l~ Ummllne ql'locIP~,aiSml.mh~Sl~ N Us grab l1o~d 'of ~!1!d1SEN~Ule~t~fbaJ!l:lleri!l!r--']!!i1,d ilfild~i!lg ~.

the<y keep them \mtMn the range o:f .antlmkwbi,al emymes ana pe:illide~. Ilfuchs er ,al. :llltl1JW thalt~'hi$ pjQ(:es~ ~nvo'lve$ a 1'1 ay typ,e !)If cell ~ellth prog~i!m" An~r li1eUilNJpmil:sa re :Mim!..li;[1t@id, thl! llio:rRl:~~, ~eg'alielli or d'lrorl1ii!tilil s@Q ilil~i!lregl~oUl~ t~alt are acllv.elytra !l~(rilbiJ1lgl gefles.~l'Id th(K~e tl1~t a ~ ij I:!adive di$.tlIPD€'ars" oilIlmJI he 1'Hlld~ bscornlE! dl'l'f!o~I1ilEN:i.SUlbs€'qu@l'I~ly,iWltr.,ilH!llll!'u t

mernbralfle:!O, iilll(JmHngl UH1! nuderlr ,elWl!!~®pll ,11in1[! a~so granule' rrrlembral1le:s,_ di~il'lt@Q ra,~e;''~Ms lleadst(l UiI'8: mixin galla .a:;se:mMy of the N EJ 'tlOimp(I nCriB. ~l.Ihm(h are' them ~etMs.ed rimnil ~,he dying (eWa~itslPllal!>mal membr.me njpllJ~e~. lh]s, :s.e:qliJefi.:Oe' cell d:r,athis distlni:.t hom !iJther knoW',nrorrns of tl[il~911iIf'l'imec el cell ~eatn .;;IIl'IId :apl[lea~$ to invobe Ir1e,;,(:lwe oxygen ~~d,~ bu,pe~(Dxide and

by INADtPH o~ida:se.Chrol:!:l( gra:rud~malt'ol,i,s d~s' !E!\'I~eIpatil:ll1t5are defic](1)111 ]11 Ihi~ ell1lzyme am~ GIII1III'Hltl'IilaI~e NI Eir5, wihi(hi 1llI1IY e~p!ai!'ll ill p~rl why they s~n'e!, reWlo€J1Jt irdectlom;, nUlt~V@~~with thi!!' tast ibreath,. ne Lltmp hils wntrib!i~e~o~nf1' ~ighldg[flim;lin\!ldliillle' lII1liuobe:5.,_, ScMH }. Cell Sio'. 1.1,6,231 (2mm. lPe'r@~ide') UiJ.!lIt. are
!)i1Od uc:ed
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i ...
N

Aminoac.iclse'xist IllredQm~llaJlltly .:riS lisomers, alAhotighi i'he lTIinor-1mag,e D iSQmer$ rigu~ in ~Hl'\!'er.:ll. iochtllllh;all~aJtl'IWily5\, ifiyriidoj((j I ph esb j)hate-d~pendeFil emJ~)fi1'les ",j;u:enl~reilli1ilif1o adds 'f!l)rthir~ :purposrt b'( 1:I~l'id~,rI~them tfuJ~ouglMla hydrogen I:mndl--~tabil.il.lE!d illflU!1le~lnk<i ge. IPairk rt

Un~ ~.ril1l~rk aim lI1Io:nia chal'lFne.t ArmtB. TWeI crystal. ~Uu'liJf\!s or baaefi'~I,AmtEl (:lind ,On.eor .archa~a~ ,Alii'll].} ~e'i;'e;;r1e that eaeh rrnonomler ~eatllrE'!>~ d p.iI'S~g~vay for ~mmlJllija (NHI~) end ill ps.eud!l)5Yllllliflel~k Ill~'ir or live tr al'lsm em br al'le helix dorm(;Ii F1$ (riOf !lI1or:e on h (ali'!' t~ is du a~topology
allToilingelllie'nt might have evolved, see RlIp p er 01-. Re:polls. ScienC,e'IE.xpr€'S5. 25 J~llllua!y 20(7). When i I1llri!(e:lllll~ar n~tmgel'l l:5li miling, the prim~ry ArrUIB Rlgu~€iLQrthe lrfr:Jil~rk IPI'uleirr

A!iI'ItB ,aJFidP!fly~LcaUly f!illlled~~ ~~epas.X!ge of alrifUlmni,tII. AJllriVi!? hy.drog,ens: o~ th~ g,lJal'l1diniuFl1l (~llo!'i n lfid pa!"I,!'iill!f'S,)~ l~~~ ilUi!t~UijOF rl~::iCieor I h~ I SU ArntB pore, which seernu ritftil'lg beea U:lie the I;irrall'llil h !iil~r ~br C! ~1.ada ~t~d~Q nITld~~,t el
(lIIillITHmii'a,. IHQW

t
~ ~

~ ~
:l!!

i~ uch s

(lIIlIUII1I1['nate' i3$50J;;!atiol'l

disr'l!Ipled? Asifltr(lJ!l:elhda ~ lfIiuo.gell drops. the' '511l0l11mol,!2cllIle :2-m.og'lutarate (I:C,(U Fruda I.es ,1Ind

~ :;f
ilL

bifld~

00 G~flIK (nI1fl'(~Se'111where

1'SnQ)!t-:auite dear

aI, lla'f!?

IJ~IP~['@'(!

a S rIII€i!I.~I1(1l~,l!lf r~(@p:torthaJt.

IJ~Ads aJmino i!dd~ lhr10ugh a sirnilar m(lllif but also ~nCiorp~rat~ C! c:h1~a~.1l!'ilf1C!iJMh~l iJackOOIl'E!.lNh,@im Li3!lIIiIino i!ddsi3!re bound illl

Gl~ K-d iss.oda l~

t IJ ~Uow .1I11I1i'n,onia efiilry; an lumeg!,llaltt1d dltlnnel, '~'lIl1d persi'Jlen~. ammou~i" iflnU:d l,I\'cL!I~1 en'~l on pH horneo5t<lsi5.
GUUSl;'i~n: l e
O! nd

~hepres.en«: or 1Ji~l;yl.am~n e ba~. 'the~

an!' !i@~~Ii\!l~lll'~pil'll!l!riled Ily proton eXd1li'!.f1gi~~(Id1(: I!);!s()'mers il'! on~er l(l reljev.e sterlc (omkain~s impMedi by the fr:[llnecr.rorlk. file ,sluthors. could eUide-nH~ fn,.e'rt the !l:Otli~i9m~tiol1~ H c:hlFl'Ill'lmlno of add'rS, imJw:lung those wi'thaJ} die. or basle s.irl.f' ch1Jf1I'IS. S@I'@l:ti~uli@:5 ITIfII'!,Istirrorn 7:1 Uor .ata ni 11 e)
t~ 20~1 UO~HlJ,ro~if1e)" Add II1Ydf(ll~i:s liberated th-e!'lilpilil~~rl:l!ii.l(l prodllC(!! hom lh€ h.aIJU\~W~~~, ahel( Io'I.Ihk.n the re~E'Jlftm (Oiuld be r~~dedl. - ]S¥ 1. A!m. Chem, Soc. 129" 10.lO2 Yj.. 067 ';1'249~200 1~.

yeO along Io\IithM!!l'Aif'~ ~he:geilnll2iFa;(;iol1) iliAd perhaps ATP hydro~y5h may Ibeinvol.ved in 'the l'.idfl'lIl:sio!ll{!lnd r'!:!.tra(tiiUll iJf the 11" [oop. - GJ( Ptoc, N(JI/, lM,ad; SCI:. USA l.oiil, 42; 12 13 (2:orl7J: [MBO). lCIt UI3 8151.emi!loj. 7i6{11492 ~2'007).

tli ~ ~

~
~
lZ

2 M I

be fatal given its

01.

COlliro~

et al have

~~~~d!l1e s~ru c:lt!re(.a l neAIfiI'l~B~b!~u.@')-GI,!lIK 2.0 i! nd Vi A, r!25~e[(redl) C::O:tlllI"tex,with li~@oty) or ~jhe 1:!.1!il:t~!"ia~ Dl:!lie'rgerntt hr~I.h:DW). Al'IltB-GlnK COPll,pkix,

'l'me- (yl.ok1 ne inlt!r leu k~,n.;2 '0 l- 2)1,5, 1:IIlI.h prodU!i:fIJ by .c!lfI,d ex~~ts :Stf'OlfiIg errects on T (ellis. lDe:spi~e it') fund1lm~nt.il1 imp,rutame lor r ~en d!wllJ,p!ltlenHllld immUJil@ r@gll..!l~liM.IL-2 is, only rte ~~igly e..'([I:IrI~$~dI. n ((msisl.e:m wilh a need 1.0 ke.ep thoe' potent eHi~m, of l~is growth fai(lo,
!lind!!! r tilglli1t(On~j['oL Vi Ua riM el' of. l1ave ilnveni91'!lt,il'dthe il'H'!fi1ag ~fII1IeL of Ib2 ~re~si()n hy helper T n~~· n (!ell~ and fitld!ihal ~~rall~e€!db::i(k !TI~ha!'ii:sms ~xi,s.t ~!ll

.. .. ~
fI!
r;;

~
!:;

I
~
1.-,

and Vi ~diz el cl, h~'

mnlril)Ulteci
'Mrlrogi~f!lVs. or <iOIJrS<~r (ll!llil'ss,~rUial eLel'Q1j@~~i!J:r ~ tile, and the prumarYlllilode bywhi{lh mkmbf'5 a(qu irre nitrogen is by a III 1J11(ll'li!~ III ptake Uuo~gh
til III

~
Q

eled ron

I1Tl icro5(;QIlJ

( s-tl!dy O'rl,h~
[1111

alrdlaealilrnUl·GIf1IKl complex," They Ifi l'IeI lhal

e!'lt.ended regioll

to ,H lh!l r loop, wtrkh mntaun!> a 111 a m~l'imli ne rfSidu e a~ 'uts liP. ir'!:S!?rts iLrioo til Hy~(lplils!i'!iCV,@!lt'il:lu t~,(Ii

or Gil III K referred:

ilJr@vily or 11'5 e:-:pre-ssi'llFl. Sell@dio:ri (1{~okin es,.....jnc.~liI d~,n9 s,ome that !!Ise re<eptors bear r mig ~he c.Orlllmm "f chaTua-elhi bolted Ihe' abilily 1;0 s~ppre'S& !L·,2 erl.![)rres.~lof1l. MO'St pol'€Hl
le;rlS!U~ tlh~

5
'" s
~ ~ ~ ti

438

2:6 JArN'UARY 2007

VOL31S

SCI~NC:!

w'l,iVVl,!'.sclen(;:emaJg,org~

EDITOR'S 'CHOICE
am(H1 gth~!l'e ~~,asIl-2 iIS@K~nd it was at 11t~ 1Je's,t in wmbin.alioll1 wi!lh em!? or mOil! (lr~lh~ olbr tyt()~in>es. If!lJeglalivelEedbad:: wa~ ,a1so 1!1ft~iated by li1'E1~I1I~rtl(lilllLl~arSlil'lTIamily o'r 'IranS{ription far:l'Ors. Io'I,ihichas~isl wHl1ll!ne diU,ere!:lliatijon or Till a nd I M,2 cells, irnlilfie' in vitro observi!lion'S ~'ile-resllppmted liJyin vivo eXI~e;rim~nts, in which U!l'"lit!!l~nt Q[ mice wilifl rE\c~mbimmt !Il -2 n~:s!.Jrab1y reduu!d the Ilevels or Il-2 prodaced aHer imifIlu niziiltion. lit \!flU be Qr interest to in1lt!stig)aJt~ 'i'illletb erlhese IpathwilrY"i mo perale< ilil d:nslilnc:l ~'II;J!jS loc,ontJrol e~rily I (eUiIfl111l11fn ily. reg ~la~~r'll T [ells, i!l'Id~lhe Irorm<l~ion o,r T cell mE1mmy. - SIS J. tIp. !Wed. ID.I0841lem., 20 06,n 98 (.2mm.
MArEIR:IU,$

R1.(ll:ls 'rw,e'S flw~,l:er

add~t ion h a5 valriQU5~Y be ell ~silOO a~ the lriglg:er cl'fthe B'!!1l;n~ warn1lilt>g
fJ'E! mod 14,600 '~.a r:5 .ago or .as the (oil use of the Older l'l,ryas colDl e1len t lmlat 'lE!flmiEIICl'ted Ute B~Uillg 14,000 years agel; um:e'~lnitv ab(!luUiIle
r

From primates
to p roteoml cs

ag,e or m~\ilP"ra 'ha:; pJ@'i'erUec r~50lu! ion of 'Ml~~ ctiJ'I1Hid.i1119 a il'llS. Stan rtm:l e!' Qr. p rres.e lilt a d r'ecold of North AUaln/tk [h;e;p Wau~r (N,A,DW) [,low int.n~ityhom Eirik Orin 'Ilul, wm vi ned ~1Jith p.reviQ~::dy pl/lblishoo lJ!',proved di9llif19ffQ~ IIIlimp-la, esta blhhe~a Ire la~.iw ly 1II1ilnm 200-ye.ar 'wakenilllg of NAD\IV 14,600 year~ ago.
r

research
for careers in science tu rn to Science

c~in(~(h::11 whh mwp- :l1~.The data C!1~o~h(Jw that 1

no d is.<:emible sea-level rise at5com,jl<lflied ~.he ,b~al" NAlDW s[owdffi'j'J1! (j1J.! rill'lg t~@ 'YQU !1g'~r Orya~. The51~~s1JIUS dem(m~lra~ r[hal NAI[lW i[o.rml!lli(m 31ld d1ll]J'1at!l' al'! net (CI1~u(lUed i?i(,lus~iI~ly by the' IIlld gnitude< or Ir,Il'e of RIl!1.~lwa~.e'r Uon~ ,i:liddi beeause l~'o,el~f9~Sl me tlwat@!' pu lse ~@~u~ted ~1!1 tille relal]ve'l~ mi~or mder Dry.as \Vh~rleiil'> the dijlf'l'i;;itDlo.Qi(:3 ~Iymuc h Imore dr:li !l1C1ti( ¥Oi.).ltgef Drf3"S wail mot a~~tiJdalte-dl,\lith oiM.~rni:lb~~ rise) in '$ieil levell. TAU$, the lO(;ilJiiQI'I I)i~mett~j,1ateraddii'lions rnay b~ hnl[lOr~anl. - H]IS ~1t!(lCt!(iflrm'fr.1plly.Zl; PM103 (20OcM.
GENETICS

now

SCII

E~CE

W~'ggl.~ng into Scmethilrnrg lhiill1lFIIlerlr

If!lJallCl'5~ale rei:ll~.Utre:5 be f,[ll:irkated in polymer (an thi n r-ilmo:> bry rQDm-~ell1 pe ITali,lrre fQ.rgi FIg or eilloossing, ill whkh the m:aleri,lil !lImlergQEls in~I4)s.li(UO'~hoof! be~'~lh a die to cr~l~ en impr'eos'S 'rille llhinne'5s of lhe 1.<I,yer remaflllingi 1011. oo10\v the die is, 1i111f1iled by friaioll, t~~ shr~arr S'lr~1I19th or !lI~t~~iata!1~ th@ pr~S~lUi! rrested by materia ~lnri ld ing Llprollll~i{je til e rd~e,a) i;iI~U as by the a mo U~HDr rOlle !.h~ di@' il~eH calli ~o!le;r.amebe fore 'it d'efforlill!L Cro~ ,el al. ~Jmlolf tha llll~ pl.ying .11 s maill-a rrnplitude s hear! nrg motion {.a ppro\l>i:im ate~y 10 11'I11llI~ the plan eof iin

ln~

N,o·t Cheapler

by the Do,z e l111

l
r

thelibn ·In;ltivat@!i :5mffa~e f~ow. Thl!!' shear mlJli~1'I net on~y ,alllQ1I'l5nlmJil lid rmer ~aye~ tQ boe·lorlll1leo

~ ~~m~l({J'I. d11!l1~I'!SI()!l~ for a rgoile~ a~iOlm!l~~IiI, .~ IFlmill:! !!ll@irnl@n~:5nnu~tlOmwerre us@a to rtj(mfUFIII ~ the' lJ!'Irlr~rLyin9 me( l1an1r5:1I11.- PI[IIS i lIlalloi.elt. 7, ::m.l021Jnl0624'566 {2cum.

i or'~en l!~il_l~~ yieokl_of stru ctulr~e1iwi~l1t~~e = 5!lIJI!"l'!bJ:r

but abo decrea ses tht:

Qr d~feii:ts,thin

Fill!n'esS. d\efil1ed as an 1f1dividiu:al's rep.roduclive ~'lIC:OO$~, ~:s!i!'ie;:i$U ~dl by Iii e ~i.ilTiber or rlln~,pr iugl, alilld p8rr e-flitswiil.h more oMs,plrirngare 9 elillera ~Iy cDn~id~~ rjtt~r, Hl1w~wu, ~ V11l"mel :Smi~h:i!hQW that i'flll m.arried ooulJ'les iI'l prein cUr'S,1 rial. soc ieiiie'5. the sourYival of I~QI!htil ~ mQ(n~'lIl'ld the father depe~ded o~ l~e nruIllI ber or orf'5prill1g!lhe li'I.Iife had (ill'ld!'amill~e~ wi~h iewe-r dlHd~enl had mlor~ surviving 9 r,i:lindchil.dlrem. ln .arni:lrtCl.l.ys,ls0[ ,2],000 fE!WrdS. o'f SllIr-lfilli'liI ffll.a 1;I'l'll91,h -Ci2ffi!U!fY L1 taih. ~unrivi3iL r ates on:he, lPi:lirel'it~ and ehi loren were ~lgfilifikarlHy im'!le~ in the ~<lI~i:!$tlall1iiLie~. lhat is" r thGse \~th 1.2 or mOf'e c.hildr@rI. Til!!! n!l!galiv@ ~He(l Oi~ bear; 119(hi Id~Em ofj,iitll'les;s \lias slrlinge r in r~ma ~t"S, la~t~ aR!!!rme napa US@, a ~d' \Va's,evident even in wome.n Nth ,emly 110 3: on~prilll~l H'Or,!!\e'''~Ir. aheu the b1rlh of lh~ fOtJ n~~'~hilld,ltl~ risk of· de.ath becarne5,igllilki:llntl.y gre.alterin
1 r

Don't gl1l't I.ost: ln the career iUI!lg~~+

At

s.,i~!lC~careefs,o~ we .lre comhu~lpirqg lfOIJ~"d i:5ffiirmlyFo~liiIded

mined~o knowl:edge experience

till: ~iglM~
on the

,job, O1!J!d@~i'll@ring WI ~fliJ ~,adllk>e. O!iJ!~' d

experli!'ieof Science. and ttlle Il.ol'Ig

of Af:J.,Pi$ in adiva.tiIcil'lg Sc~!~J'I ce·
s.'C ie·nc € alwl)!Jnd~hE!' world,

Cc:u't!~{S ICMg Ii s ti:1~ !'1ahu al s@~~ectio~.

www.s(iiencecareers.org
Fll!iatl!J'es include:

• Theusa nos 'of job postii FIgs,

• Ca reef togols from Next. Wave .~Grant r~furrn<llilion • Re$tHt1le/(V' Dataoos-e ,~Career itQnHlfil

r.a/tilers·, too. In a ddJition, 'tlhos~ ti:hiklr@n with tire

lar!j;est nu mber Or[ :sibl1llg~ were Less ~iitely to SoU rvive to the age of 18. with th~ YOl!lnge~l ~ilbling:s 111. rl'!\i!t€!;t risk The .abiti~y humalll rrE1lrn1ll!l') to g or Sl,m,rive ~ong aherthe los-s of l'1e,prodl.d ve hl ~(~ lion h i:li5previously be.E11'1 e::KlJlllal IJy lhe pmifned live e'~~~lt ~ mo~h~r'sliNiv~l hs O~ll~,eSuc(e~ s c'~ h~.r ~:~WfJP rung; wi~h~Ult.a liIlC!th~r, s.u rvi'fQrs.hiIJll of the thUdr'Bn h ~o~led. 1N~'lJerlheles~. lhh;
p;re'!l$U re ror r'emalle$

~ ~.

~ CLIMATE

.~ Po S'IP,~as,hO~(0

..
(5

$.[IIENn

ld Water
ili ~~Fge ;;md rapid

;i liJurin gth~ last dle~a lci,Q'tijij~,

; inje!;t1QIii of glla~ial ITIe~tlj\\flterhoml1he L.aQ.gr~m-

tide lice S~eet to the IMQrlthAtlaWitilc kalled melit-

~ w1ile'r p,~dsre h, 0 r 1I1IWP-ta) Iraf,s,oo 5O~a l~1ll1by [5 -2'0 m if1I only a[bClul two (entu ries. TIll s enou-

stUldiyd~I'lilcllsba~~:i that th'i!re Inay' [be-']!,~~'i;liv:eto ·~top lleprodu:dnrg ber(ln~ they ,c:oml~iele' ~hfld In~arin g..- LMZ IWe, Nail. Ar:(Ni. S(i U.5..IIi. ]lO~, 553 {2:00J1..

439

SCience
• W\lfIlI.S(l,el'l(illmaJg.iO rQl UOO! Ntw1tOfi~ A'lttMIJI!!, rIIr'"
'1N.a~inglloor" IE :OllmD.!l Ecf.1Dfia1l: ~"~~~5[il1). fA'( ~2·~N5o!'i~

E~ITI;la'I!HJ;!~!W

lXrlu~i;Hmll~ D[PiJIl1i~~I~Cl~S.

It. B!li!liu H!lr[J!lOOl" IBar.gii Km'1f,i;l! L,IItIIn'H

1~1!i.iU. ~'1_o)i!~'Ui!i!)di!

!)~!!::!!~I !nnnec!)i' 1Ml.I!iiidfard illl'~ [i[IItOA. R. )ii!lD.!t. r~oUilllll!JmaiI

M:1l!iWrii lpt\jiU~fI

r~:J:~OiI]l'o'!i ~R!SllrR

,!!l!J!1n !. ~!';rn'It"
(rintln;jlmi!~p@it,3~~.rnrg'1111UHlilllJ

~1;J1I!l~r;RB!llflrl~N

NI~Il~:I! !ilfl1i1l~ !II,!.1!M'I1i Waoj.!.;!R Willir; ,5lrIfIl:lU ~1't;~1I<1lI~'ABrn-1\I

Vdr!Pll~ (!l5Il!!!Ia ~EI!I'KI !1IJ'fi1MSQIIPil.l

fIILHI1IIIUIT'& MfIMl_1lI

!'lBI!IIlI~

N~ :WH~6'Sro,

r.I!.'(

:;:~~.,)71~~7

lHiDLltit..,1I2~:a: linl!! R/OOICi C~nlbri!l1~Il', I~U!! 1!lQ' (liZ ·d4 (01 ~:~ 326S.~o,.l~ ."~ (0) ~n 3i.26S0j,
B:1~tl:lun

_fli

!'i', .mr~;wiiO!t ~-.olltl I'fIS1""'HII,E!iIlilial 11 (iIIlXlg~ ~IIIilIU IJ:rrIJilU ~Ue[[ j. (11;11,; Pilmela !. H1n~. P'.:rrJlaA.~31B; {8oll!lri1~ f.tln. S. Wi'll! n~root@, ~1'l!1ly:A. P1wl rjl L Bryd'llrlbii'. ~lJy Ailtlillauo;[h.\ H, .jI!~~ Smilli,. \I'~!r!aVi~:strfl, D\Jri~ lj\oo~ ."'~~(!(Irm '~l!ruIa Ia·'k~'i, 'rfrI!5II!J1\, tau r,~Nb l.1l1rii; !ilIIUIIE I~IIR; S:1~"'rl!l·~ w~tl;' ,ljj1';l~L~tE ~!!II!IEII_Rr~f~ S, Mar~t:I~~: nil!! ~'i!I'nnI!t!; 5rnmnoo JI. SuE~~;.!l!iiSlHll,!lrE IJEII1Mr~(jj_ Ettl ~n~: lilFOIlUJDI .ifil~\It!fI]Ol!lf!t l1:~; IIIllIJlMt!IM~ '/kIll TIll!,; :~III!: cO!'u~~d~~f(J,~ t (~~. Qrrth!~ '""!li!!itl, IH nrfl' J,:!~h. ~!b;:IrO'I' P. Oro~ Itt'll'li.~ )nh.Tfll.t~, W~!IOjj(lr; ";.,,,. ~1!lt_;I, IWliJllllF! ~. flctl'1 Hi(lOlll'j,'!i:I\c; (I)!lO!a;g(O~!~i!:I Ci!rOlW! ~!ll, ~l~r 'S1,~U:l; ~~1:!!I1! tii'!,;j~l!!iI!"', !t.'!m<!~yi!ir\'!l (IiQr1j, (M~ Fi!.i~!J'~!!', [kIi !i.
r!l!i1i).1!

I!U'~Ilf.:URlI,wsfll.il;mril!

TIIlfiiiril.AtrliJll.. Kiir!!ll~r Srnilli. \tl~m1JOO)n.I~OO~ _~om(hi~iI[,Oprnl!l[ R{;tiD~; ~UIIll'i'~J:!""'rllI!OII. :iPHVoIJ!;T!I TMidtii liliW~ J:j~m~, iii.. Erin ~Jm
~'lit~i$~

(a~1!!d.~(1Ilrfl!1lI.'IIII1I (l'lIihia JDIl~~;

Bam,

'!I,i'ritlli~o!oi!; rniiiii~M,iiIiJII~r~ , trl<rlli!l~

f"91l c~:t!!~~ ~I oiIddf~j.J,. IYII~jIlfl9~~It~J, rlt'!'!' ~1l:I, ~m:I!i~bn!: ;e:b:l''''~'1!.!iAS ~l~Z:n Of ~!2-326·04fl\' ~~02-842-l.06S. ~lUf!i,J ,~cId!'t~t:!: ~AIlS, !ku: 9&179. ~~mr.g~l)II1<, I!>(; ri!0090"~!7* I1i' A.!ViS ~mb~ :stJl""i~~ ~g~{_i.~J'!!>O!! ~~!!!i!U

.~ulooncw~

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Wl\ri~t, AR~ Gui~, f:alli~ia ~.~

p.o.
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26 JANiUARY20Q7 VOL315

SCIENCE w'!NW.sclencem!,g.org

L2 J fib
IE'DITIE D BV (0 N SUN C E HaIL DEN

S(ient i~ts have found !l:o(,aine en ,[I samp,~jng ,Dr OI!!IbI~1Ilcalli!:. I'lot~,. ,addiil'l9 ~o ,ccm::~rl1
ll'l.alll'r,el(!ll'ldi~ brecQoJlllil'lglhe of E'l!Irop!!. ((t~illl1!! capitall

Snow' Ov,elr Ireland

Jest ~Tlh.1!tSi.t~Jg~~L'S that l.hQ~ bilts ~'l'~r~~oU~d lip and u 'l.e d for sl'lorting, '" he says. "The re-st mav h'tJ';'@' b~@!n conl.alrl'lilfl~t~ du rin~ IlJr(lC~S;Sllll~ lin b~I1k.~. ~

A le1,t1m 1e1!dby Br;e,U P'aiu l~, an e\Il\Ii ronunental.

(~~m~Slat Dub~il'l CiW 1IJ!1!~rsiry, a!1i1ly.n:d

<15biliis ,,"~Uecl€d hom a r,(JIU the cily, lJ!5in~ nd
'ilirOj!11l,,~ggfC!J)~Y (!(JIke residuf'S aJlfld !'!H!r'SS, on <lU (ti'

Apo(:,aILypiU, Adj ustrnent

The
r,]!11i110U~ [Joe IfIliSday

Clock of ~he Su/l.e"f;1 oj

hd'J

tlieAfomic

Scllmlisrs (BAS)

begun to

speotmJmetry. and lound [h!1Frn.Ih ree biU:;; elso nad trsees of heroin. ~iIih!!! C;h"i!IlI'I1ci!1:!i produce aJ I'li(i(l'. dear $iglil,~l dOWIll to pi cog{a1m ,C~JiI(!~ntil'ati~m," say!> Fritz

.~
ti
'

I
i ..
0-

ia{l(lr anothr globall rCOiloem i'lmlo lts t il1llillg: •.~Jn~l!!!ad or r~necliflg tl.nllythe UH~at ,(If Ill!ldeaf alml'lliihil.a'lion, iils eontl·ofthe oIjj'j!(lrlei r.aiculLJs f;ft{!iW iI'Idu ae:5 dimal@ chang!!. ~Fb::u::ling, de:smJdiv~ storln~,
im;r,~asl'!d drought. ~Ilid PQI,/nice

Siirglil!l, iii (hem ~st fil, Ut~

In ~mnlle rail"BiQFI1I,edi(.i;lIL and Ph!lIrma~e"l!ti!;al Re'5e.mh in NlJrembelFgI,

rlllet! ~reCi;;l!,lsing Loss 0 f IHie a rqd IprOpeil'ly, ~ BASsaySllf lht Iris.~'S lh<tt slP~rked the Iml<m/eo, 'l'hal ami !illd€(Jf prrom@r.aJlio!1l, including
I

A:rt~rs:pelf'lding •~ e:x:ol1(: ku::;ates.

Li.I

i dreems up an e:l:.p!.anationrQ.r
~s:pecles

mn~Cltiflg SP@ChlHlfllSif1l a 'yo~n:g! Britis h !Ilia ~li!r,alij5t

yga,Fs

Ge rf1l~f!rY. This iSIi! 't the rim s[udll' or its; kind •

A 2:001 U.5 . .s~udy ,o~ S1 blills, ror exa I'I1Iple,. r("lmd
(!(l~a~!H~,

l'/iJ'onhKr(lre~'s rre(eill~ I:II:m~hle'iil, th~ mim!!Hl':! iI1~H1!(j from'! '7 to 5 minutes belore mid'ni'ginL

me.!v@pusMd

how aile

Ition rl'ts Cha rles [)arw1n~, bu t. ltalso m.alrne.s

,~ fUrred lRu~e~WaUacce (1823,-1913).
·Ii ...

lll',all1l!j~'Qfms, another. The descr1lpinto

hemin, m,f;'t.haI'l] phelam~

ilE",

and

the co-d~scwerer of rl;a~Uiral se~eclicHll 8rJd on@ ()if ~h@ 19UI[~llmlry' s g leading bio~ogisl5. Wallace was
rf
::l!

,~

'~

rOl!.!lnd'ers bioge-ogr.aphy, l'h~ ~tlidy of ~ [lrgallisliTl~' dislrn buli;o n. ~ A, ne\l!I online exhibitkom

also one offthe

or

But ltls thillate'\it ill a r~C;E!nt burSl or fep(m~,~y European re'Sear(h,~r~ thl suggest!; in(rre.lIsi:ng d m'g a bu ~ a n~lhe colllllilVIi'erll. ~Spafi'lis! hotlght to 'be ~~e iil!1Iii'lber-Ofle l !;Q1lInlry for ~'~(':!!~Jl~ ~bus~," :S~V~Sor9,~t ~bi!!~, I reland apJ)eiilrs to be ri~ l:ng lo the lop,." P,aull i1l~ ~h:ill in S% 'Of th~X!!1iIplre's, parltc:lliarily €20am:1 f50 mnes, the ('oncelill" tral'lon wa~ about 100 t1 me'S niglher lhalrli~ihe' phemydi.dinll.

WhaJl d~, ~he' s;t,,[warL'S of w.eal;lOf!lS (Onlrol [hink ,tD~ thi!i.~UZlier doom? ~I understalnd the~lr coneern abolM bro~u:fel)im~ the a p pea ~~ says IVi! n Oellri eh Oir thE! Fed eratio:n Orr Aml<E~ri(an Sde'{I~n:5tsin 'Wa5hingtol'l, [).(., bUil "I hope ~tdlo~:i 1101 di ~b <'I' ~ aUe-llition" hOffillhe FHJdea.r-we;;lipons ~hrei~t. ~t'~ 14lin time thl.? dod;; ma~:5hiHe1I ~rFI'il the ~ts1947 debut;; thes ~re·sl we''Ii'e been W~IS ill 1991. ~ti~ iII'~pon~ fed lll:liOrrlS movoo ~he docik rtf} ba dk tOr ].7 1111 iJ'lul~.e~.

i
fiI'
!

the Nalural

liis~ei'l'l,Mu.seum in tondOl'l1 documents

WaUa.ce's work and

me 'With anno'l;ated

l;omlbs Olpen Up »
~.s:arre about te gll'l lOllgl~SOlig hl pl'ek~ into, the '~m!lbs I)f~Ih(lli:auI!11iY':S '~.1!r~v!i!!111~~I;)rs, In )1JI~y2005, ~heo];tlipanese #l.NJhaeological. A~'ii{j!(i:altlmll 15 academle ted :SC(!~UE:~ M.~jnJ;! JI~~;1!n's, lr!1p~rial H~ms!i!h.ohjjAge-n'f;,Y ror lPe-rrnissiornl W Tn ~m'!l~tfga~f 111:1miell1~ ~mpellia~ ~Dmlbsrha~ h~E! '~O'Men'Only um.~ry, if gil}l, illlsp,eCliolll bysde'llltis!S" Ind~de,d(llill lfuie li$1l i$ the tomb ilf Emp€\rnr Mnlot..!lI, IiIlli'JOiruled fill the 4till 'or '5~h um:tmy i!nd is :ll.1idto' rest in one 01
]01 p 111'1alf(llliH~Ij)I09is \

allrtides. You can brows:e some or hi,$ tral'Jel

indu din 9 Ht@ ~eUI@ j n which he r desmbes the destructlon of aU his South Amel"i(:an spedmelris in ~I s,hipbo~rd 'fife. O[h@iI' Qin'@Jrings ~ndicate Ithat Wal~iJ(~didrll't r~)efll. b<l'lngo'rler)hado,~d Iby~he o~.der

dis!lal:cnes"

sde!1last for example WaUace Mo~e·(I

friend thal~.he was "thalli!kh,l~ llhat. it: has
nCit be,@n t.e<U HI rIli@ to t.o~he pUlblk. ~ »
WtV'l'Ol

give the tileo:ry

'~hrt wor!!!1's largesllbllrf~ I COIlI1p[e-J«!S, f1te~ ~ PJ'lrt:Srt!lQt·day Osaka. LaS.1lH.mth. ~h:e J d~:~m.y'ecfeedllhlll o:ne!reprl!!s!llilt.1!Hve! hom ~dh sociely will be alLrnlll!!d to enter oorm~ Onllll:! IOiITlIl5. d .I nthe pil~l, eli lk~ Iil<tV!l;! a~(used~he Imlpe rial. H I) L1~.e~ol.d1~gi1'n{y, w:hklil ifiw;r5,~S 5ilJ;IfI.e900 b arI'M~i~@'s.. ~~ !b,locking del'ail.'oo i Ilve sli'ga~1QIiI$ iOl1Ilth@ $I rOUJlflds ~h:a t SCleilfisU m,ig hi!. 'tum UiP Elillde<IilC,'@' 1]11[11 a IiIceslors (lir tfld~y's royal lanlilly re;!lll~y ~ame rl~Gm K~~elil, 'Ihe Japaln~~ 'lregel'ld' wra~e!l [he 1~llie.tigre blllcik~hro!l!~ h 12 5 IlITlIIJe-,rOrs lQ 'the S,hirlto s!Jn 9tJddess Ama~era~l! Omi k.1l1T1i. Myths a~itll!!, ther~ '~~aCOOe'mic @ebate over \!Ihiid~ oWer ~it)mb5 ItOll s;e. IJM hkh j fnperia I bof! e'S.

h 1111 kinatu r~on li net ,i)ICU

(ol!~(lions~.r3l· Lheo-lfIlJ~euf'l1/waUilj(@[ol[ediofllilhern es~ls'tj51l

'Koji Taik.aihaS.ih1, fo)'a ~1i!1UIi11ver:s~lY archaeQlogiS!t O~l Ute associ~~!~fI' s bo~rd, says ,re:sead'~rs. a aHowlld tiO lOlJl{ h a liIy~hfmg. :)0 they i'Hf' m~ely to (lj):n!;e'ntrat~ ~l'nIllilappiim'!I to.lII1l:1 l.a~muLs. and de,taHs of~l~ WiOf5. "U'luil Ji! o~". we' (0 u ki UQ' l g~t if'lCli~, ell~, so thh. is i!IS!ep f o~he rlgih't d i~~(,lion, ~ ~ays idikiahas.hi.
'1'010II, llbll I

W'W'W,scien cemaq,!J rq

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For news and

with impact, turn to Science

There's only (Ine source f'(l,rnews and liese'a rrh wnlh the gre:~i,t\i;:5l im pact - Sci!l;nce. '!III itlii ever 7QO,OO 0 week! 'II print !le,adelrs., (li fld m illio ns !f!!Ofe onl in.;:" 5cieijc€ ranks es one ohh€ most !hIghly read !'lii!Ullltidi:s.dpHf:1lary iOUln'Nil~S iii! line world. And for imp,a,ct. Scfence ,carn"t be beat Acco,rdi OS to the' n~cen l ty released Tho m $.C'I'! ~Sl ,~IIlU mal Cit1itio n Rep ert 20005, Sdem::,(! ra lilIlk.>edi!ist he NO.1 lilriost·(Tted mu ~tidisci pi inmy [eurnal with a eli t<lJi ollractll r of 31,. FaUrided in 1880 by ill\!le ntor Tho m~s Ed lson, and PII.I I,~she by t~e nen prom AAAS. Sdem:e's rr1l;! lati on als b d pu the leadi f1lgSQ~H,;efor news" researc 11,and lead in g edge presenta lio'n of co ntent CQriltinues ~(\I g,r,ow'.l.:oo'k~!'lg for news and ressarcn that will imp.i.'!ctthe world t'OlII'lo,rrow? "Vile I1i too k ~I'll Science.

www.scie.ncemag ... org

TO ioi~AA!!.S and Jecehre :!ICU~ow!'! poerSO!~al~py ofS,imc,e iM!1Ywee'k go !o ~iliiIi!.SdQ<rgtiilll:Jil '

EDIUD

BY VUDHI]I1IBHAHACHAR.jIH

M ave rs
I

U PWARIlJ; (i,AZIE. The

Senthern
ObSep'tLUJf)'

~ol;\\'Ile~~d {1nh~(uropeai:~

(ESo.) hopes to

grn]1!k~

the 45.~'ear.ol.d orgallmfz!llifm

even mere of 11

pl<lyer on the internarioual as·t'ro;L~Qmical! 5Ce.U1C. Dutch ~sn:onomcr Tim de Zecuw. nO\\I sclemlflc director of Leiden OhservatOi.ry in the Nel~er~allds. 5r1)iS his biggest c~.<1llenge will be 10. 'pcrslL<Lde ESO's [3 member states te hll~,i.d 42·:ImH::iter the Eur()i)~,HiI Exmolmli;;ly Large Telescope, Till;; Ob&1;;~''!,I;jJ:~(lf!'Y isalso il partner W.~tlln, ll.~.elhmued Slates in the Atae ~u:m,1Large M~.~~:Ln~u:ter r.ray . A b[;:dn~bui~l hil Chi le, Man. IVlmmtait'n., (hre:clo.r oflhe SlRIce l'elc;r.;col~e SC.ie'~.oelmLs:tltute ir!,ll B~hh1l'iOh;. M1Ir)lh!ind. sily~de LctlUW i~~i "·f9[ilJtas.liGehai~·· far ESO. De Zeeuw, 50 .. 5l1cceedshen:ch astrenomer C<llh,eTine C ~f::arKky, 'who is k"vi I~g[0 become pres id~!ilt (I r the l.m~r:~~_ltio~{ii1 ASlroll(~nm,~.c<l1 UJ~ion{Sdeuct3 .. 8 September 2006.p. US:)),

:WOLF [PiRIZ~S;.George Feher has alway~ had a kl'l(lck for (F;,ulking tough pmblems, Grow~ng upin (~echoslovalkja ill'1~he 1I.'9'30s,he talU9 hit himir.~U ftldio ~l~(trooic). 11111941, he and eight Je,\,,lj.sh friends. escaped INoihl:]perserution .and fle-d to, B.riti)h-n.lk~d IP.tlLe$tjm~., .Ahl'1l'a $l~!'it il'l Briti ~h det~nlioli, Fe her \llorked OIU·t til

H:ei:lll"e1ltIJI oiwrshy of]~rysa lem (m arhem a~ks), aJnd Albert fert Qr ClNR51n OrSilY •.
Fram.e, (lindl P~~~I'G~unb~rg of tne]rilt(ih
R'e5f!Q1:: h Center in Ge r Fn~ ny (p hysics) " n

find cornt:el'nlp:~.lJou~!:ow,ud the uloiver51ily (Scient,e, 5 ·S.epu~mbe-~ 1997,. P·.U41). On 15, .l[an~ary, 5ool!!~high ((lurtjll.lstke Park Hoag Wl)Cl LJph~M the IJlI'i1]versity's ded

liN TM E COURTS,
D'E.VASllAIED. PImatnrtma'lhi:!l professor who
r~sfi red froma Kon~.il1'II 1'l1v€'F$ity 11 ye~lrs I,J iIIg1t.? shot .1Iil:'l1'lrrowat a Seoul high court judge: last ~i!1kfifl~f losi r1I9~i) .ii ppea I. of t he<dsn1issaL Ihejvdge suffered ojl:lV miflorilljl!J~. and the resBan::ner has been charged with ,*Umpledl mUlrdef. MyIJJng 'Ho Kim lost hiis; iob ;;J l Sung kyunkrcl.arl 1I1l1iversityin JL 996 af~err t'e~lilllg ]II1Stij tuliorJliI,i ilmho,rj~.h.lli about a n error ill tI math problem on the I,Iniv'~rsily':s enilr~mc€€':r.arn,He d.QIin~ed retaliation, bUil.m:lm~flist.r.nors ~l;'!iidhi\! WtlS saeked b~~~ui5ehe had been rude to' students

sion, ";AHho'!.!gh we re(ogt1li~e Kim ,f),!i ~ lfIIlalfl OIl decent sch.oLnship who 11 a de,all (OrnscirulC::l1!, a'S.
\I'I'e

way l~ demde radio, traF1-s,mls's;ions b€'lweet1l

,it;:
iii

the IBrnHs;h IIi]gh (orum i:551'o ner and ]0 DQ~\!f1lfng $t,thill la te r proved pilfol'Qll lin the [[stl'aeli\~alrf,o IT

~ Z ~ ~ .~
'1!1

ill dep endien ce. F'ehe r

nUlved!o UII!~ U'nllte.d

S~~ res 11'1 1946 ~I rna eV~i'iI'HlaHy \'l'O'lJl"ld lip (.~'lih')j'ni~, San Dl~go"
alt the U IIlfversU'I1.. '.1 of

zr

did !loUm;l h~m:a$il ql,l'~lilried me:mberol th~ academic m('!mbeuhfp· •.~ ti'l(! ]iJI,dg·(li ~aid'. Ki~ then whipp'eo outa crossbow and fi red til SIO n.e .ii r rO~ij t11~he judge, \.,'Omldh'i9! hiis m~dlrUt "W[lenl nl1iy~pp.eal1ll!flS dismissed, I felti~la t my I.ife was over. ~ lKim told repor~eir~, deniyling Ihat he had tried to klU th~ jlliclgl~., Th@ Kor{!(! I'll Prof>e<s50r:s union issued ~! )l<lIt~mentafter the indd ent decrying the JllIl]!lg as biClsed, Kim. who's been lJlJ1;emp!.o,~di,orthe j)t1 st s®er aI ye-.1Ilrs., wuldl now even race i! deal.h sentence iflhe aUe rnptedi murde-r chawgf! ls jJwo1ve<d,oHirials; stly.

.~l!'\Ihere

he tllm eel his;

~·~!.eulthin9 to il:!ncoV'ering 'ihe ~!IOrkil~gl$ efthe .~ reCliclf01'11 cenlter:~he rno re(Ullii r mll(:iline that

InBrief'>
eft AfOI() RI[]! PI!!:1:2:11::5. Earth sdenHs:[ Wilillac::e Bweckell' has ''IOn the 2006< Craf.oord P~nz€''ffrom lh~ Royal Swe-d'ish .I\!l:ademy or ScienC:·f/s.
Broecker (h;~fU.a researcherat
'$,SOO~OOO a~valdfor his pioooeriF1~ ~IIJd'i~) ·()If~he link biltl\'l~e rII ocean c~~mist:ry ~rnd (arbon di(u;:idek~Y@ls if) the iltmosp,here. Jhle ~ ..:aclelf!V [has, also ~ln!lQl,llnced lhe willner of the 2007 p:r~z:e: glltll~oPQlogislIR(lber! Trivers (above. rigllt) of RlLJt,ge!f'S Uln~\i'~rs]ty ~n th~w [BrlLlIllSWkk,. INew ]IQ~s~y, who reoowe;&~he honor
Colililmbia Un~ver:sity,. r~(!~~~s tille

f2 d ri~s [photosYllth esls.

~

last r-l'eek, Feher's; d~te(t]ve work earned .r;{ him a shalre oi th~ 2007 Wolr PriZ€ 'i 11

Gle-m~stf~,. Feher (flboveo) shares. th(! g s :IJ)O,OOO prize wiU'! Ada Yonath, ill (he!mlist '~fl n,d ~,~r etur al b]ol09]~t a Ilh @ \'l[Ie~ u zraann .~. lnstirute of Sde!ue ill Reho:vot tsrsel, who piorlleeroo \'la~} to use x·rdYS,to map HH~ :~ three-disn e nsf;Qn~l s·~n.u:;ture Gf Hne rib os em e, ~ mb~r VII·oUPn2i~S th lS y@m '90' to5tQph e 111 .~ 51i1fl~!eof lh eoU l!liver$Hy of (iii liIli)rI~]g. ~, Be-wk~l~y. al1d Harry Fu~nel"lb~rgf 1~1Q o

I
i

.~ Got a IIp rllrthis pagfll.·maH

~--------------------------propte@aaB,s,Oorg

for hi~ (Oflltri b utio·n~to the

understs ~d ~l~~

(lf~.ooperilili

Oil a FII conf d

lkt.i n tile

illl'li!l1fli:lII. "'IIorld..

W'W'W,scien cemaq,!J rq

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from Ind ~LSlr.y, H ~dm~t,p lUlU gfo\~"th Imilllell {ielfl' and WR.r~ l(lI1·~n(ln L~sh M~p@d of 'lI.S.. emissions w it~ 0\ decade iHHI in o'rgOl'niz'~OJm industry· MYirorlllTl.ei1lillist pa rtl1'N~ tQ lower eurrera :~evels b}' 60 % to S 1}1It:, b)' sir"P Oli'l di nlilt~ e oollg~. P'ad mJielill. IE's JeUn1!Y G
1

~OSO..Thegml~)'\~"'Il~

11)eomrol the cost of e.l1l]ssion permits, PCIi.il,rlPS w itlh. <I price I~mi t, bm not lin ;J. way thar

woulduuderutinc l!'le: system, ilnd [0 bar old-fashjened coal plaats from getti ng rl'ee

cre.d1 Coal ilion membe n; say l:~lCY",',un to its. .uvo:id p iecemeal stone ~~,]iSS]OI1S C<LpS,. sue h
tough newCI!I:~fQrnii.!I.!>yslt!nL :.'Iltd more d!mC(},lmimlmeasures down the road, Another set of{mcc~odd b~drdlo'\iv:s 1I1s;o
~~3

Idbbi~d C~mgr~ss last we~k, ··W~ d~~f1;! t~1

im ,lgi ne <I\"~{}r~dull ,,,'hi c:I'n science <l.~ulIl:I ~,~

N,ew COlogress M;ay IBe War'mingl Up'IO Plans f,D,r IC,aplp1iln'g' IE,mi:ssioln:s,
The first winter
~toIFII:J.% ,ofdu:

gioill.c(!(l,p~I'1i.~e·' reraek II;': l im;.'l~eC'h:f!!lg1;! [Ill d c other problems. said Reverend RidHI.rd Ozi k of the NiI~ional AS:i<OC]clli un of [lv<XrI"' gelicals (NAEt one of sev em I evangelieal Ch:risth[l!l 1c~~d,erS\l!/i'loave joined with h
Harvard emJ.tomologist E. O.Wi:~son <IIul M i~$~uri B ~J!=.'t!mical Gard.tll'l! b(JI!) Ilis l Pctl;'ir RO!\~en<lnd ollrn.el":JIle~e<!l,rc~.ers "n'::spm.lsible for

season. long

mre;'"

due. b~ew l.iln,m ugh W~l:sl.li.ng ten, D. C .• las l

weekend, B\u' ]J«.l!]xJwems 01·tk;di!f<11 h;gi~l;nliQn to linl~l '(\1l'boflcl'l'1li~I'Q:linsoontinued I)] bask in the glu',,' ofincrt1m s ing support {OF'theiH·m~S\l. The new Den~,ocr,atuc :unajodty in bath houses orCOl1!grre~" l~af>made it IT\{)iI'e lilk.£!)1
that a bi]~ c<lpp:ung caroon ennssienswillland on Prosidem {i~{Jf<d.eW. Bu~1l'~desk b~ftlf(; he ~e<wcsofit::e :iln J\:UUl3UY ,2:009. Of course, tlhe lini:ll!'OlI"~l', fflnlj' ]cgbdmion \\I~IIbe sh!l:J."M.'dhy O

slash enml:ilssions steeply j)~ 205ft AIl.1don 12 January, the newly formed United S:lntes

care for C reat ion." NAE has agreed to dit>~

('~ irll!;ue ACli'o:i1 P'l,rtml.~f;.;hip.'l group th.u Includes HI ID~.ajm·U..S•.companies suchas

General Eleocllic ,Hl,d B,P American as well m; \~fRI and the Naul~I~Resoun:es. Defense OHindl (NRDC). WJ\'Ifl(~ll[~d f(];r;). ·~!l11111iil.d~~ory [and] flexible" emissionstrading sySi"
~l;;V'!I~Ihil~

hib~I~~ .~"r-il:SWIr's '~(](~I kif' \,!.iith ill.lfo:llwn.nion

0.'111

the w~lrInil:lg at1uos.p:nere and other cl1ivi~ rom,n:::ma~. problems to eounter \li"h<Jit Cizik caUs'junk scieaee" fmm d~m[lle skepdcs ..
Sach c01!lliHoliils;~t'emll~ke]~1
~rl rre-~g.(h~n s

has impressed !l:';I!l~rgy 'In~IY\'Ol;';.

~he hand of lawmakers such as, Senator B<lfl1,O!['a BOl'!:.I;;f ~D'--'C A )1. ~J;,':ml of rhe E'I:'!JVlm.flime:mi.t :mtd.I?u,bh,cWor.ks committee. a : who [0 I~i~i;:sacarbon cap ":;lS soon as pos:sib~.e.'·I-iI,ollse Speaker N,lIle), felos:i (O--CA}. mcanwhlle. l~~~ereatcd a spce.la:1 cemarirree 1;0 explore and lP'llhlic~ze clilnl,lIe ~h;;u!lg~ l~gisl<utiOl), Hnlis~ ~n~rg)' commit'W::LIU:S

the demmul'l of variou S ]jO""'erlll~ i~:l m.~i~,es. d :;lll!ti il'l~{'itn:M well encounter (I, pi'~'sid(;jHi!l1
veto, But R.~vquestion the ne\'i'follindpo~ itic<~! momentum bchh!!d some typ~ ~)f r~d~r;}i action, "The tide i~hnmi~.lg." ~<4:y':; Jonathsn Lash ofthcmi.olD,pmffh WorldRleS<{llll;\Ce,s lnsri-

"You (;QuMn'[ .~2IJVe imagined :2 yem.~s,i:1Igo a grnu p u f such eempan ies gcu.~.l1g beh i~HI such ~"lr!ilel:S:' s."ys, Jason Gru:l.ne'l of the
W3S~1~IlBt'OIl_

N~li(infil. Con~mis:si.nll O~I En~rgy ]loli.cy in D.C., <Ii similar consensus-

b~1Id~ g rOtlp dm~i I~1004 reeu mn~';)!ldl,;;~1 i ng 11;l:;$'ISgJ~:;:_i.ivegnO'!I",. N R DC',"; lDavi.d Doniger says the convenl.iQ!Il.al wnsd!JiIl1~, ill! rhe

nne {WRI) in \VtI;.;hingtonl).C.. which has je i!l,ed <'l1l:l,e'Wrn1 ut~] e.l'ship between CIlVJ.r:tT~l= :ll.1e.l1t~~;i "Lndindm~toy <lmlmmced this week. :s't!) Tille preside !:Il'S, State () f Ih.e III n ~OIlJ address, d~ih"~r.;:d a F~~r ci(.'tn·l?\v~n~to S press, \/oI',IS expecred to fm:m;, Oil ethanol <IS
psrt uf a I1m~m~II'ld tHory el ~lmUle :;;mHI;l~y. But most observers expect CO:lllgr~:s:sto. be more~gg~ssi\'le inlill!d:Jilllg greenhouse g1'ls

time was tll.ll Ihe g.mul:J;s; pri:riIciple:->

'''reiU ,;~5

~e I;.~h';lir<lnd auto .indu!).l:ry :mppyrtl;r JO~!1 DiugeH {D--MI:} opposes the Il~W ecmmirree, wh ich \VO(Im.'I: have leg is~"ui\'e <'!IU~:tli.(lrUly.
13m its cI:C!nkm. miiuee structure"
S<'I),S,Om!lr1CI, dcnl0IlS~'m~L':S

··Fal'as,the politieal ct::mi.terof gl:<liviJy would

pres,idcnt.

em:us,sion::>. T''Ii\'Q :sem~:wrs hQP:ing w become Al'iz(lll.1 RClf.H~hlk;;1Il Jo:lm.n M.cc.~:iln .uu.l; [II illoi~ Detn OCI,',1l :Barack
0lI!

Ob~m~. have cosponsored

444

bil~.'~:h~u ouM w

go:' ln 2005. forexm:J,1p~e.. then eJlle"dlYcom~1J1JJ:Uu.ee SemtWilf [):ete Dominici {R-NM l chair ur:g{ld O!l~reSS to "~~'!"fJI~Cl~"'i,;l~Yfu.ldreS$I .'" e M>1lU: clmlllge .. Ehllli1e best thaJ the Rellubl i:C<1mi," " hid l'mdy ti('juld do \\I(~:S m~l!k~a '!.iag~.e.lHI!lbinding statement c,IHullg $o:r ;·ma.ndalory steps" to cap grccnbeuse gas cmt,is~ial1ls. ~:-Im ..I::~eRep.~blie'lIl=:; sl~oCk. to yohl.lu:ary 'i'lpp~,dl.cs Ie promO'! illg ~'ed~l~. ""hi ~~.-~reh. The:; snn~mem by D1l'U: new CI:ill1,~ate piilrt~ ~ership iseflsily [h{l~lmSl ag.g:r~s~.:;.i'\t~ yl,)C[ VOL315

dli..n Pelosi "]S ~u)tgoimlgw

defeJi~o lmte com(~fISy_ Soml:l

Y..,-r aggl\1s~ivtlaerion \wn"lOO

iIII :mlTIo\! ing her mg:encb.

~ ~ ~
if

of the 49 .Repub~:i:c"I1."illil the :l OO"s.e.n. Senaie

wi II b~n!I;'t.~~~d ~~wrdd';il by_ . 'the Wh~~A;l~ ~l) 'i."I;)~U ii!

UmJS-.e ohmi:Y emissioas legislarion, notes ~ S'elllalcEn~~:~y cotnmlttec s~:~~f:udirector ~ Roboen Silm,l:OJl.. McClI illl and OlJ.ml1l"<'l,·=-; :i~:1is a ~ b

Jl'O£i'tJ'I'e step. sa}':> {IllOlhc!l" ;lide. blil ~o\\i:un the S,~TPlfl'Ort of <I majowity of

..:m 11!I11N1I'11S(;liiI

is; ~~1l1 ilk~ll¥ ~ :senators" ~

~

26 JANiU~RY2007

SCIENCE w'INW_sclencem!,g.org

SPACE

8CIENICIE

fOKY'O-A m~:ssioll 10 explore the moon's ori gi n and eve I U~iOUll, h .J:> eras hl;al before 1~llll'lcll. ilrlier t~i5 month, Japan's space E
agency leril'1lil1(1i~~d~ts :lUNAR,·j\ probe

because its ,nub.it iou S h~chnolo~~f \Ji.':IS still ~(Dl r'~Olldy R'I g'o I;1v~nllf[~r~h~I.-.~meh hf~d been d,e:~ayedibr 12, years, The C<ln:l!eUatiO:~l of the m-saarrcd mission hn, se nt ~f11! mor s Ih rowLg~,llm,e ,~I ba ~ 0 spaee science 00I1uul.mity,. "The t'Ge]ing is
one gfLn'ilifo:f:~,1I1egrerlm,'I:t Ihi~l1i,liss~onis not 11y.ifIJg~" !>~j"l!: Cliw NC~IL :fL goo.!Qgi.Sllll the

scientists sketched out. :key uu resol ved ques~j;on s ,l.boul the n'llOOr:l'~,~uteri er, I,;U~J!e gm;:~ a 011 the eD1J,guneering si de hlldd~ed over how in mel insiCl~ l:ilC moon. 1I".~,e}l opted m dcsi,~n a probe to drop frmn (l~bit <JimJembed inslrtl~
~)!:)\:1Ii!l'S f!nm.l~y" aml ]u!act

something

!!IiI" ~h~ hU1!Olf ~oilIhili]! has never been tried before.

ex treme d!lY- ni ghl teflm,peratllreI"11Lctu;fIiliom. h'lstnln.'lem-s left 01;1the surfuce b)' previous missions, By I 99.5-lhe p lalmed h:lliitchdfl~C-I~C o.rbiH:1f \\~<1S ,ready to ga, TJine peneuaiars were aot, Tlu~y'N:S~!nnet.even !lf~~~12 addil~[)nf!l years ,omde\,'e~[lpmeD1J:~;In1.d ,es;czll.nillig costs: dll.c O1~git1l<l~ priee •fig, far the sa.tcll.liH~'I~m'U: ·W ..IS $: I 06 mi]lio:Iil ... and $ [32 Inn Uion ~lfl~been spent so f.. r;

d~.\!fried

\lV'~.lIiljwellt

WITl'Ilg~)'

K:'I[o

U!litUv"'fsi.ty of Notf!.;;: Df!!l)!;l in SU'~Q,th B'~!1dL Ind i,nllliQ. Desphel:l~,e ~etb,u;:k, Jap3~, :s,space sci e~niti S01.y ~hey fel'l,l <lin cOrLlilm iued '!O p'Ulr" ~,'s
3U [ng~edm i,ci:llily cllllileng.ung VCI1HI.U'eS.

:·mys lh~ team V;)r:;t~y l~~.cl~f·' estimated l:liu;: challenges of gird.ilil.g insrrumerns and
electronics
j ffil'HH.::t

to survive

an

Planued by the Institute ofSp.u:e .IW1,d Astronautica ~Science (ISJ-\.S~ in Sagam~~

hara, 1I1I~M Tolyo.LUNAR-A called for;;lln orbitung spacecraft W' dmp two torpedo" ~l~ aru."d~~n) el' CllUit) th~ 111fKliIl1'S surl::t~~~~ b on the near side and one !OfI the f;<Lrside. These "penetrsrors" wml,ld lodge], meters bel ow the su~Jm;e. acli \,.Jti ~;lg sei ~:lI1I~nme~er::i to detect fa illtm.oonqu[lkes Ullild other sensors m measuee hear nO\'f.th~J,l g:eia)' '~111.e data W E<ulh" Scientists l],OIdhoped 10 deduce deta ills
of the

~'"ltg.~ly ~mlk,tIIuwn I,Uilil~~1f interior, such

as whet~,l.erthe .moon has an iron cose. PI. bettel! picm re ,nflmi!<!it .~!m~ttlflllyWUl! Id Ot)!1S~:~i!~1l models o:f':lmO\1I1 rhe moon :[onned" M iS'i>~m~phu1ll1 ing beg;!ll:ll1. ilh 1:1 ig h hepes w in the C"'lr~y i 99'Os" Wh:~le ]SAS phllll.et;Jry

in p•. :~jng ,I force of n ILlP ~;o 10.00'0 times ~ll~'J~ of g~'I!i[y. ··h,1.odding h~,d iruJic.1ll:ed: thaI; just ~(l:Im.e hardenill,g of Ctlmp(l11,e!U$ wtluld b~ sufficient,' Kala says. 13m resting proved otherwise, The fir:>t smlg\,\1u;.;findillg .•] proper tc!>t site. Only <l:lkr l~.C SidJ~Lij:m:!d, llh@ HlI NAR:41. orlJiliJF 9{1S,b~!l ri~o'!d:.' lor YNrS, b ullpJ<O'b~~m5 fii:l:i'~;.l.am;u:;hdate passed did eagineers realize tb:iU I.SAS with the p'<lyload ledm its (an(~UlatiQJlI. lacked :i:l!J!iwMe ~q:~tiplm~e:n!t oIlnd "flley thought this would be mose p~';;Ic:tiica] racilities. In 1997. with the project already 2 ye~1n' b~hi~d sclm6'dlUI~, ~~I;;y !'!!,uUyf[~und :f1 Oil!'! rban develaping a rover to traverse the adequate testing ground: Sasdia N~n~.o:Im;111 11I1IOon"& expanses, :;~y:!i Nb:~.ab~ Kaw. an Lqlbon!]o!:"i~i> .in A~bufILlcrqw::. Nc\\,' Mc;r.:i!cu" [SAS Il~~.nC'la:rjl eientlsr. And bl~ry.ing the s There, <I <device aki [Ito <I gi.U],t, up!lii:de~bwn lie iustruments would S:~,~e.id rhem fromthe m~ss rOr! to !i1e ml10ln ca ned CIi!~l'Idray'fla iil- :JL.fh@~ rsl Ini:la,a JIl uli! 20141. w~th a~n'OllIaIJIS o,fl~he 1110011by 202Q.ISRO f~ ~.eekiflg~250 !l!Iimolll im SMln.·Up~ullld~ ~O'I.ofll~e fil'S'U;I$UlOJl4!1!lt. OIl'll' Oi~ Une Ikey t'il!'cilllloh:lglies en '~e5;lduring Ih~ I1ecCl!very ~xpeln11i11~l'It W<iI'!; thermal pmlediol'l. an '5,hil'M !polen liCil i31s~ml'l'<llJll.s mn line in lense h heat (I n1S1!i!dII:nyrrktiml du ring ree IIllrry",~ ~.BElIIIHDf40 re'5.eCl1ichers here prsdti(ed~he IIQ IillweiQ ht lleillt.·re~r:5li!ni~. fI1alteri:!!ll!> made hom ~Pll118' silk a that
.J]rM
's,e I'lsi n g

~[Id iiI

Inches Closerto' 'Crlw,od SpaclIJ' Fllight

1IlIIIRllMJ:A:NlANTHIAIPIIJJ RAM" KfRAIUI.-J ndlia toe k a lIllother s.ffila U. ~[t'p' lllliis li'I.Ie~kIIl'lll'Hd a place at '~hetop table oI5pa(e·~ining oolltom,. Tine ~ndi<ln SPifI(.'e Riese-arch O~ganiilaliDri (iSIRO} t,u!lfu;hed (I o<e~'I'~e5() ~pacen.illlt into orbi:l en 11.0]a II1lt.! and om 22: ]anualiY. (oll1lnJ~I!II'591U~ded ba (k to Ea r~1h ,ny, it

ts 1il~~f!J!'ledi for

'ror a sp!.tlS,h(lOWll in the Bay~'1 Bengal"lifO

kilolflil!l!ters h'llm its.l.aulrlch 'site"

Tille f@~t o'r lilUf~(:hif!g ,~r'ld fWi'e'v~ngl a probe 'ha's ,t,C) date olll~Y be-@Il ad~~evedJby Rldssia, the lhllited Stat'e't arid Chllt'la. and ls ~rl ~:S~!lnf~1 step

towa Jld ~e~dim:J1asu(tfllams iI1tO !:l:rbi l. "!Both th ~ I,all ifIch ,~IJI'IdSllli!5hd(lw~ (iiI tine' spacle capslJ.le ~\lere'~omplel~d w~thlexlbQQk prred:s.lon:' 5.i;l'YS
~1lOl1e'5~e~ClIll ~, Krls !:u:I,ul1ll,llraihy, The Space C.apliule RemvelY ExljIl~rii'n'Bfll is par10f <I n ,er~ort1.0 €'I.I'l'lIllu.ilIW ~1JI1lI"h ~mnalllniltielll<ll~ ifllD space, ~O'I'IIin~llhi!lt!JQiill. llileSOOO Ire§eiJIrdl~r) IhIlNl a:lllile Vf~r.illin s'alrablilari SpilIc;e'Cenlll r hope L"O lImh .[Illol.h Elf ('1<~Ilt'r" la 111O,efllilll. ~al:J.5UI!Iii' Hlon" 1111 2,008, the plalll ~.cll~ for lh'~ lat.!ln{~of iI remote,I S litO

'GQ.iI~edi~It~0 uLlicae ,of tlile So S.O-ki logrrram 1:011 ~~. ~TIiIl! llilerllfl.-J l PI'tDt!!(;t~OIll pliU ~syslel]fJ il\1aS on@' or the ~flO por;t~nl ~.xIP@rill'iemt:s bei n9 cs rri@ ~ eut 1111 h@ ~ 'SPii!O~ f1@'(O!,I@rymodlul@" This is aJ p'fewlf:s~'r r~:r a'll forlhtOnlljf]!iI r"~L!s<ilbl@ l"UUl(h \!@!llid~-al!1d ifllllie IoIllQI rUIiI, 00 tliili! maf'!liledll'!i!iss~o~ OO()'.~says, 'spa C~ (IMoor ~ ir~U(lr III filii" S t1resh. lilile miss,i@n alsO' carried tW(llill(lu~rria~!l sd€i1lc€ e.xperimernl'S in li\ihidll ,crystals were glr!o~~1'I1 nde<r JileC!r-~e-~o""9Ia\l'i'~.yCOil! ~ij lism;. R~seClr(~ ~rs arE u r Ii!!lW s'UJdyfrrgllh@ calp~.IlI~.eNl s'eo'~ wheol~er o1IH the e;(p@rilfrn@!us Wi!!lt .[I.t.oudlil1glQ Il'hm" -PAULAVA IBiAGI!A

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445

N'EWS OF TI-IE WEEK

exp~os~ ves ~f)fi re~lil!\l p~Ul£tlrii~(l~rs, SCn'fpp~d lhe m~~hUl.A JAXA spokesperson t~u;:S<Jlldy C'dn1~" due to schedule oo.UII~ says the dJe.cis,ioJl \~'-.:!I,:S based 01'1 the orbiter's Em detC'.r1Oirnt i.o.UI.in storage and tl~e [Ret t1'n<!11 the s11ri>1:ints, lSAS could ij$~I.1'i1.~S!l~d.ia.fi1lci~[lies, p,elrte~·r.n()r.;. <I we ,IibCit re.m:iy fur prime time. o[l~,yonce every' 10 mOIDI,lh~, resultingin 'HlI Nm ::l]~hopeis IOL.st. Taka~li Nakaj ililln. cur ~ cxcl1,tda.timlgly slew process off trial and error; AI 1!1l:Sl.aUiier the ir1S~1[LUi1l,l,elll!'iWen: hanle:l1I,ed rem LUNAR-A m,n~<lger. !l~~)1'!ii dKH the penes u lfiei;elul.y. IS-AS cngi rI(;"CrJ>determi m::d Il~~1 Uililur' projc:e~ is nut bcing!l~ndil)i~ed.IS:AS d~\rC'I(jp~n~~l[ \iJtlrk i;li~dthen try ~h~elt'!~~~ric~'ll wiring \~S 'll'lU rn]~L O!iD.~~ lh<'l~ t'llm !;~}I'!~pl~~~ loaded 0111<tno~]m.er moon w.m;: fixed, they found tH~<'lllhey IKld to, die\"ei,o:p [0 ge~,thepenetraters
C,U'D1JiI(l'L'i1 uses

illl.!!)

derermlne '!i\lh~d~~rhe ~WOi~ hills an iron r C ere, prov ill!;! til a tt~l.,epeuetnuorswork, S uhseq uent miss ions; oeml d Il(~VCI~~lenespanded

their

US,eI

on the

mOOiL~

.md, perhaps,

MOl~.

a circuit to reset the 'PU .uf~~fim,pill;t h~ T

launch date slipped again
3]l,d,

~lIi~i\J~"

"Researchers

k iud ofim:esl:i~atli,QI'I:' ~ys ~,~Ul.o:dru NlizlIlallli .• E [Igi.ll!;;::eltii,~.g 'E1\.p I Of!;: r ,~SI! LEN.12:}. S et for a ~p<!C:'i;l :ilCiii:iUi::.l \'\i~o h1;l.~d,,'<lI1lue is,~im:l lImil m Dmil,g the most recent rO~Jlld one-sting. launch illl la'te March,. SELENE comprises an in June 2006. the seiemi ric il.1MmLr:L~,e[lt~. c~r~ t'Bis retirement 2 ye",II'S ~llg,o, cults, and sonW,are;;)'~~ passed n.msl£l". K{l(o fhll1C'rary seicntists o~1Xsidc Japan agiJ1i:C" orbit~r llnd a pziir OIf ~mli'lncrsatel Iioc;sUmt wi iii aim illstrmneul:!>;n lhe mooll~Qooll.a:~ lih.eGnon .says. BtH. fu:nlh,'!':'f work is necessm'y to llilll<lke "The comlmllll~~Y would very 1.1ULci:llike to ~h\;t:,(ln'n!!lltl1]!!~c,l[ixH1S, ys~~ms more !:,Qbi)!]$l, s SI.;;Ca g.,;lop~~)'sict~1 n.clwoflii~m the ilIlOO$l (0 d~tuiliild dmOllJ'Ct em its hlp(],gra,!,l~Y; dl£ til~!~l\)lm,·' he says. Ancther rourrd of te s~i ng is sdlile<!l~ slud}lil1ite'rlll;)_~ structure and hear fluw, but tal makeup of its sllrf:u:e~ Its rmlgneri.sm,;mCl its grav.u·I~Muol1ial ndd, "\Ii,le'.re confident." says two penetraters do 11.01 make a .lmcnvork," u led for til i!; s:mUl1i'lle:I!", 111m's roo late to Si"l\ie LUNAR-,A" AfI:er Ol :mys AI1I.dre'l'\;' C~I':TIS. "phll1lr.:!!'tOl.l'y scientist ,n ](~~O.'·We '\,1;: ~eMedi.1. ,nK~\"'e'refew:!y ro s,:o." series o f iute 1"IlI3 I reviews [S;'\.S':\l pareat Johns; Hopkins University i.~m B~I[imore. 1r ~O.•tlu~l W()Iii1Id. Sillne i()flh:e sri ng 'om of lake the hll!ll'iM~nsdemise of LUNAR-A. m,'g"Iib~Z1I1:ioQlli. Jap;,m Aemsp.n:e [1>\,pl.o· M,aryLaf'ld, K<l.lo says lSA.s had hoped t~.e' ~I!N N 1:5,NIO.RMIILE rali(J'~, Age~cy lJf\XA), 01'1 .~:5 Jia l1Ju:!Iry LUNA.R~A would galhcr enough d]ata to
agaiJ11.

w~n still roqm;:;~this

13m he VO\VS tluulSAS wm.uclnain at the forefreut of space engiueeriag and space sdencc_ "Jfyoo dbfll 'I d.evdo'p ~l~W tl.tc~:lIn.CIlogit'~. y{)~ \'!IO~'t m .. k'l;; !l~\'\f .sdeD'!![ific d b:cnv~ eries," Ka:lo says. ISAS next ,n~]rfj,'lg~ ~ngilW'~dilg .,]iIHI Qf science is its S300 mi]~iolll Sele'D'l.ologicJlI. and
t~

(NSF) S:l~ouldtest rhewaiers befere plm.l~~n!;!! irll'O Ol, COSIly p:rOlJf<lm to award.prizes ill e~oo!,lr:"~.gt' UI1lIfii{N~tioUl. Tih31i: tlhe:\fflIdiC:l ofa[ll expert p.:;ill!iei of the: R"Ilional A.cadle<~lues· 1\1<111011(1]' Resc<lreh CQnmcil'1i.1'hichh~:s c.mtiml:-JlIY endorsed an idea Congress h.c81drdered NSF to consider; o I mDlk,1Hy., ~ti:giskllOI:l'lmny have kYStilllttl.~I.
[011

Th (: N ati una ~ S'C ieucc Fotmd~t

~hhlk I!lll:lomof:rerillg megaprlzcs for soluticns re impanam seieuriflc challenges with :S(ldeudiltlplk~~i.n~~.A~one hearing, fair e'~~lmple.Wol r speculated 'lbol'll[ how a $.~. bi~Uon prize could y.iekl in} deg.,.nt tech!m~,{lgi,c'il~ sclutien W l~,~ nation's dependenee Om! forei.gn Qul.l<lst s~milmcr. NSF' oW:i,cials turned for guidam,ce :to the Nmio:L'iLid Academ ies, which thi s week issued a 44~p;!lg~ report lIrgi~g NSF m .f1,dOpD "an experimemal ,;lpproioliCh" b;V p i.lot~:Ulg a .~21nd· fUll of !'m.~Uprizesbefore cn~· s.uderill£ spending big bucks.

uoncd areas such as nano self-assembly ..

clflernl.1.islU'y. low-eacbon ene~y tech. nolo-1:.~~s, ~!ld [~ach:i!lg Stll'1!\vare. The f~l'Jort green suggesls IhPlt NSF shcm:ld, ~m'lrt with a h<lmlfu:ll

of prizes I<mgung I'ru:m, $])00..000 1.0:$2

lienand

raise the payout t,o asmuch

mm-

as

In 2:005 .. Repliesentn~ve IIFm:~lk \'¥o]l' ( R- VA J. t~e~ eh;!!jr of the spe ndi ng psne ~ that se ts NS F's budget. asked ~llii,e:agem:y to

S1J1ei~prizes are useful, S:'IY!ip<.ne~i:iit and former NSf Oimetor Eriehlll~)ch. "because I:hey bring .illi new p(':op~!.;i\,\1'h(l,don', !l.)r.m~Hy parl~c.~p~~M~ goverU'lm~m ill, progr~uns and be'~<I'I.ts.e lit

$.30 1li1lil:lliol'l inlle coneept proves succc&'1..rill. Myer::; ~'lYS Congre~~ should increase NS.:F·~ b'wl.d.gellie! make room rorlhe pl.ugrmn 50 that its 'Cllrst doesn 'I WiV1ldt::rmi!ll;: (}ID!goi!~ytlCtiv~lic:s.. The report recommends st~]lrl~:Uilg the rriz;~$ tlli!' year, bm wi 'h NSF f,H;i~lg ~! budget fi'eez'e (Sden[·e. 5 J.am~~~:~"y, 24). p, that's no~ likely to l~.app(~n., "It 's a solid report. ,u1.d P3, lot iriS ~S' .dw<ly~~:1ia,e fiSh~: W<'I!j'
to g(i."' j:lfl.ys senlor NSF !!dmi~dl!:lr~l{)r Nath •. iI,iel Pius. 'i'l.'b,o h<ls m.u::ked dill: issue. u
"BtLi' ~.ii:.'~am! ~xp~!l[l:iv~pr!l)~,o~~tkm,

<lil~d

,III oW:,> you

U.l

H~CU(;:COllltr'll'·

",e]'S~:l'I11 ide.qs. But I ~MtHl.1dllJ"t sp.end RI bill ion 00 it:' f'.!I.mJ,ellaiar Mark. M.yers. a r~t~~'d resG<l!reh e1'\.cculiv~ ~u Xerox. be I ieve~ tim,at NSF ··l~$Ii;'d$ l() d~'!,'tdQP Olml;;'lhtldlology" 031 hO'!,1I''Wl [lIn gu{'h a pro" gralll arid thC'l~ evni.~I.~Heeilich Me~) of the W<lY. The pH 11t1 I dud Ilot r~~~llI1il';;lbd~mytopk·.5 10.1" NSF to' t<l.c.~Je. <lJ~:hou.gh rOlf

[ d_ml"t W~ ,ftO\,'"\'>'e~ould al¥tlnJ i~ is y'~<~r:' til An c\rcn blggcc;:r obsHlde. hrnl;'e'lrer. may he ~:liUe~I(;lcemC~]f_Ulg,~ on Cap'ilO~.Hin.\i\hlf:is r!(~ lomllgcr un 111£ spending subp:a1~d, fLmll,~I,ouse ~ RepuM:i(.<ms .Ire i:cn the mi nori~y; A De:DJi1L()(:l:lnic~ • <!: aide 0111 the ~JmlSe('ennni ~:I:ee' 0[1 Scie~.U1e ~lIld i

lCl;hnoh)g~l. wh~('h ,ov~!:'Sc~!> SF.. $;]}'5-lh,u N

~~zes probably

~hUSHf~tive pu rp()s{~si I

l !ll(lU-

make mOle sense ,un mj s.,>iQn~ Orit;:lD'1~l;-d ~k~m~y M.Lcl~m. NASA ~1if' th~ D~re:~tl !i< Department. And the .. ide quesl iairn"O: ide'<Ii~ the t11l!ii a 'pI:i2:eWm aUT~clllew pl!l~I'~ ~Ilto lbe ~ rese'lrd,1 g~n~.e: "11.1ave <1 tmroclltilrn~ebel iev.ii',g ::;; dHM. [111~rc fWC IlHi'1my P~Ol~k doing bask ~ re.se:"drcj~ ''>'liiI.Ci' ,lre'mI,'t<lII,'e't'Ldy r:hm~,I~)[lr'Mil!b NSF ~ 'L!ld il];SpmW:f~!US~" "']IElfFIII:IIl¥ MEIFOJI'S ~

446

26 JANiU~RY2007

VOL315

SCIENCE w!NW_Sclencem!,g.org

~I II

, I I

R:ESEA.RCIH

fUNtlJiNG

IUC: IB~alllk:s t C~ampus;-WidlI8Ban en a Tolba.ccto, IMloneV'for IRel:s"ea.rch

Ceneernedabout ~~adClmlic fn:·cdoID~. Ihc ~he !il ra I1t do nOl v~o:~:f~len lve rsil'ypo.lic:r u

Kans:as, S~amlltards E"Iolllve Again

E~9hlee I'l menths a nelf it~s~ate ed'uYlllio:n

board adopted ~(ielll(e slimda rcls pr-Onlo!i rig

tl'leteachilillg or iJllt~lHgeiiu design ODt KansaiS is. ~e't to tess Ihem out N~:d 11'I(]Il'Ilh, il newly

lJiI\~v.~ y of C~~li rsit l~'ir!~~aUC) h~~~ ( d~luj'~~

\lQtung
QilmapbuiI,

(Iiu" e;"~impl~.by t'Kclll.:Jdi!lgfi)reig!Nj;"$)..

f\:;'

of last year, th.ere were 19 acl.:ilv·e g:W<Ll'lts. t a UC ~ll pp(l:r~ed by the l:ob;,\oc()i]igd~!~try. IQli! llfappmved, the ban would make LIe the ing $. t5. 31lffi,U hO:Im (DC 'steral grants and .. Ol.l~yu.s .. m~ivel.-gity to forbid tohaceo dolcUlJimu::I's· added 'IP lo $,4 bim:OUl t~~ t )leon. ~ .. la:~s. campus-wlde. Facul:ty members Before the regenrswould cons idera alll!lJI is,h0d OWl' the issue for 4 years belfure ~obllccn.ftnmding bail,. l!iJrn.~'(;"V~r. ~lsked the they caU:ung (J1Iil He's go\.·,erlli[lg: body-lh,e :>ena~e for :unpLlt Th~ll:~ when l!l.ringS!;Olc::oJ.n~ il"Cgem~~:fl~:ake ;.'! S1.~nd. [~J::~",:a~l. n,;g~n ~~~ r:licrI[ed."The 'mbaoo{l issue put our princiH~J;;
to impese 3biamlel

h."I11 0111

re search IUl1d ir]g from t~~b" CCQ

C('J!ll·1i·rml~.~s.

ples :i,mI conflicr," sayssenate mcmherand He Santa CI~iI,IZ

!l.OC i.1I

p:lo yeh 0 .~o~ ist :F'l.y·e

Cr.·asby. "'J\I.iOtM of us agreed th,l[ acadlemic freedom tHIHill,L:lS a n ot~l.;rllliinci.plC's."·· Bl~~ then

the ~!"!ut()ff' !i~\ri'!.i:\'\fGddoeumented

~!i.lbIiCOO

ev idence t:~l the .• n

!ndl,!:s~ryhad mnnipu-

lated researchers irno publi.h-

align~ bo.ard @ii:pe<.ts to adc,Pt sL:a,nd.ard~ Ih~~ emphasize ewluticm. The (h~mge fOI[tli'WS el~(tjolli s 'thal Hipped the bO\,lrd's 6-4 {Qm;fr"'i!li\!'·~ 11l1ljoritv to.1II 61'-4 ''!il~rgif!~QF m~d'e~ates (Scie,'lC'e', 11 AJu:g!JIs.l~()06, p.. 7~3). As. a r'es.ull, llrn~ board r,eplo'3l(ed the former (hair, iO proponent St~E A1bfams.. wl~h B~UWagnon, '\lho has lou gh~ <'Ig<limt the I,[HaulI'I!Jeci stllllrl<lriCI5 sin ce their ~dopliorl in AugllJiS~ 200S,. Ihe p~ncling $t1!~darrd:'5. ave ~eorlli'llriUei!l h by a (,Of'f'l!i!i! illee .(IIPpofFiled by l~~ boo ~d!ih::l!t d l"li,;,oe;r,~ iii prmiud dleenned u Il'.1I WlP l<Ilble by cO!,!$'~!'¥.alli~·I2'S"~W@'ll be gl.;!d to Mng b~(k standsrd s !:hat d (I nro't mntai nsu pe,lIliaiturall ~xplanCllio"$andaw~ il11tir1~ with ~ali()riaJl a!'!di inti?maUonal.llivflns •.~ S<tys Siuoe Galmble. a ~noderate 00llrd me;lnlibef. "nlest s.t~Fldards wH!lh~lp te.1u;hens to fit~tnSlthem tht le'achin9 of i;voh.lt!oFlary content."
..i'N'lIllDHI~t~li tBtIlllAilTA.€HARJEE

~ng biased l:'ie:-ouh:S.....-.1by l\.\ltmklil.1i! manuscripts or linemel!ilillg

tncut b~<!ll'lkel baJ1l Qn tob<1cco, rUl1.dil111al Lie. bu~.]a rm1'5 En~~IQII1l (rigJlr) ~ys ;!;1II:h" ban would Yi(lOla~e iI;cadfemi.r:lreedom.
fiss;iQ!i!I. S~~~I

ofrl1mclj~lg .. G:~:.'InIZ charges, "We rea Ii ze d <!.C3.~

IN.DR,oviingl fOnr IMmtn ROiverS

Budglellrouble'5 at NASA r,~~nlikely nb: plaFlS 00 se-nda seres of sophi~ li,cilltOOrobo~~c l'O¥ers to the moon after Ihe .1Iger:l{Y sends sn ol"bi~e:r 1.ll1eJ'~!l~,d yeClw. NASA omcia ls blame oil Hght e'xpror.atioln budget almd~he rising {Os! of the rov~rs, 1,~hi(11W@'re mf?c.nt tc~rtd p05s.ibl@ hmllan l.andilrn'g sltli!f. and gather :sdetrUil1<: d.a~(L IIUl'ustry ~n d a ge".~y5(l~H(~~ Ul.1lt up say
lo half ~.~'hl:! mugtlm~y '5800 miUion s~l.lIsid ~ t ·o'lle~~h~' f!exl 3 yearsllor rove!" dev,elopm;e·f!t at .Ill~ars.hell Spa{ e Fng ht (enter 1n HlJIlIllwill.e,. Alall:l.aIl'flBr, could go to !l'j,oddard Spil('~ ~lighl (.erUe-r in GrEe r1b~~t. tM:aI)'[,a,nd. arndAliI]'~5 R~e";I·(~ (en~er in Mollll1itlliin View, C[llirmnhll, w desiign SlnilU@ffllnd ch@ap@r :!ipaoC~W!rtt thai ,colild dl) simi La rjobs, 'l'h~ deds.]oiil 1:0 (,alf!(e·lthe o~grmlal set of r(!/ll\e;ro; mmceo; JIlI~ta~iI1!wIIlilOIlM.~aHer NASA Am.e

Fi!C'!!JI.ty·

St!1l!ll'O!lGL1!nt~: (h1U wO!uld Ii~Jetc

deli!l le Ih:tldll~l eM be iIl'lils;al~Y." $.l'~ys:CliO:S~>JI Forn~aIliY. ..

<Jill.ow.i~g the Iill.n:il\lers.i.ty to
.~!CCepl lobuc~(l

moneywas

punted h b.<lcl~ to the lacu:l.ty last week, "It's ;,1 very good. decision,' S''lOYS James EI1I,s~m:I.n cpldie:l.lliol.ow~tm He los Angele.s ..ml \i\jlln.o uses Phi~ip Morri'S mn!'Ut')t ··AC"l(ltemi!i.:: freedom nuik.cs.l:hi s a g~'e',Jt [mi ''e~.''Sity.and the rm;:I.ilI~)i ~,!I;~d ~~ml' ~or;fl~lSid!;'ir~h~~~s.S·LU~ ore m
rhoroughjy."
I] ut

a.llowinlg 3.1:iltreilll toacademic

freedom,

s.umt.on

En~rom ,tiis<1,gwe:i. '''I. do not feell~tt a:I.lY tebaece fum.ding ;l'lY'rml~remenlsl've had ha...e \ ~~l(jrtlip~il~I~'d <lc[md~~nie Ih1l;:d()ffiI~:' he S<~s. my 1[1;1.10013. nslmm published <I. study inthe E !J~"ilishi'l.trtdtr:ul.ml(J"J'Mrtdl'H f(]U91d no ~ermi(m
between seccudhand :lin'lill.ok!e <liml luung C'f1nCe\I' deaths, UIL'pmj'CCL \~~ prurr~~I)' I'umledlby rhe Cearer fbrllKl:OOr AUf Research, 'livhic:h ilse~ fis n.LIl~~.,iIby lob.fL!:,'.¢(l e(imp~llniG;!i;..~~~li\lllJij~. ~Yy8: the s;~lldyw<lsrn.ethodl.(iI]og~c.\ny sound <ll:lid

m.'IIHZ. :1 bin-

c:rugi~m~Ir'and antiwbae-co crusader . ~ hlimcc:o

m UC Sa~l

~ F"'1.Il;t::il~'1;o. U:liI.s t~.e .~r1,lml.eJ1tilm~H1<!: "Tille h

hH~U~lry fu~ds r~;B;C~lreJ:iJto c{lI!liifi~!O~

~ the public."·I.le says. "This manipulation of aii th.;: :>d~n~Wc pn)~e&~ $\m.bv~rt[.; '~~'Hh;mi!;; I.. f~~.Qln

rhat it

Irtdd",'d1!~11 inlil~}.!iil[Ii!.lt ~~.i!mf.i[y

(Jpilm.ion t~J

i
~

OII,uli;j ~lll,t~l~~~~C~~dle li~fidmn~~l~rd to

~.bclJll~Vl.:rs.~ly.

~.nll&o;;IOJiIJ .1 0

MO~mllemUm for <~ UC-\Mid.e bi!sed QI~,lhe!lr,a(kmic I"'rni:dom~.·\.~cad." h § tobacco rumlil~g ha~ .l~ecil b~LiI.dil1g:Since The selJi;llc's rt:cOI1UneIHhuJo.iiI W lbe ~ lQ03-. ;SeVeJJlllm:~~ W]~llnllll~.e mnveTS.I.l)' :lly.~~ tern. inc.II[J.d~I.~gUC Bcrkel,ey':S School of re;geDllts reHe'cted lhe div~ sivene:ss 0 f the !l:i Pub] '_Ie H~al·t~'.'ll~dIhe UC S~11Dh.H!O C~~('e~ i$$li!.(:. On O!~L;; llHuld. I.hol;;facul[y ~S$~ne;;d~lfl,r I ~- - - ti:I - - ' - --- -, .. ..... '·growl:: is;:S'LH!!i: of OJ.cadcmic freed.om M,,'ould :,::;: Ce:ruc·r. have sh~n tbeir door!> to tobacco
~ llllU!1;~y.

~he :;mo~i~lg dt:bm~""t;~]<'lIlddi:JIJly. d(le5afl: UC :lI.How only (~crt.'liI1J points of Vie\~f tab,e b.u~ 0111 fm:tdt:d."~e$1v~. "~vIy emire C.lfeer h~!j oor;:11

chi.e·(Mkha~l GriUVn I!m~v~d~h~ pro.gram fro.m 5 ~1I M.a rsllalL lim at. derisi(ln,50lJrl.lE!s '>i!Iy~ 'i!i.U Imade !.o pAli!a5~(O:rlgrE~'s Repubhca n~
dOl'lilinal~d Ali!!bama d,~LegaUo.l1, bUi~tihe re'etl'll

i Inie
Ii

But UC"s f3Cl!Hy-c{)nli!Jpm'1;:d

AC~lc1~-

Sel1JlIle voi.~tod these' mll1lS in M<ly:WO.:ij. cl(,';(~hlr.i~.g thi'lt ()Ili~.y the rlilgel1Jish[~dth.e

.~O ded :irm:f~uH.ling_"dn.d Unn t~1ey ~ II~ad 1.0d·o It far thG: c:nrUro SY51.~~].(I.r no[ at.flR ~ Cm:relU:ly. UC resear'Chers !l;.m t:ik,e ~ rtllll fiey Ia:.!'n;i! aUii)'!JIflU roo. a~~ long ~liS~J;:'rms (]ol' ~ <ttIJ,~.ority

b~ raiood" i,. r'oto::getus b<m!~l;ldfll~Hliny (~iI.'l i:JRSed soleiy on ilS; s>!n1.rue. n the ot~~el:~the)! O de(:·hr-edthal a(!mdemic frcc:dom clmld be :>uPlP're$sed. i~'lI1ddml the lOLmcco i,:IIii.dm;;lry ~l;;td ilL hh;Lory of sucb :SUPIU~$si.on. The' seem i.ll,lgly OCUllt ra<.li~cWfy "'I'D rdj I:Ilg prov~d tOO ~nltd~ thi;;regi$nt$.t\l~m for .,

·e-I~CliO'n,,'i/hkh put. Dell1f10(!ra in ihe driv.e:r's, ; ts ~at. ll)~k pr·~)u I'll arr thll! <l'gen (y. Th~ ~l;lm:.1l in.~ der of the ooukl'l'be rover ruding ili'QI.J'ld(Over bud'geot :sllmrUa~1:5 in NASA'~ ,~~forlto deJilelop a ~alll!n(her to re!p!iloe ~he·SP2:(>e shuttle, ~ted ku ni!tiri@\iifl@n~ in ,2,010. Th€ !PW~osf!di(uts ar~ po!!rl or ~he <lg.en(y's 2009 ilnld'get req nl~~lto b@ ~nn(!un(l!dicm 5 Ir@ilru'ary.
-AND'RIEW IJI,iWIlElll

~,Sclel1cem,)g.org

SCltENCE

VOL J1S

26 JANIJAtR''f 2007

447'

N'EWS OF TI-IE WEEK

M I C~f!e ~ e 1iJi.~'1 g~. 11111 a~ (:~0 rt t y o n ~l.l~ slJ1lf)king issues \.\I1tno runs the Tobacco CO:Im,~ 1.r0.1 Pl'ogmm at the Roswell ..Pall'k ,C!lIlCCIi hurts aCaliCM:li,cfrectioll}" nOD ~'n,elps it," [nsrit t:I~ in :B uffalo, New YOrk, S<lysJ~e·5d is,in Ih,e end. the reg:e nts voted overwhelmpreeedcne w~~lcl],hreatens au culture of lngly to send the issue back [0 tile f~ilCI,.lIfy. An t appul ~lIlcdVi' ithil.hc delay, ·".11'"good [0 academ icmree:dio111.·' said Regenr jefferson. amendment charges the senate to d~lf:iJy deb,me 'IC~lde~lnuc frJl!!OOCl~,l1.'· he !>ii~Y:-;, but .'~ f \l;lh~'~[~er b]lat;'ftk~Dtebaeee fundi~l,g ban tTlJI:r ~~ you C~l:!m'~heose on 'th is, yO~L e a:1Iii C!~(l ese e 't Coembs at tl'emt~i.;o~~im!g, ·"It weuld etlI1IW~~ a b~fur~ the Or! ~tnYlhiilg." sig~tdlh!!l WoJ;;:do iH)!~ lr~I:;1 ~nir world-e~~!ss tIH'4;{'1:].l~!1=- ~]cad~~~icfl!~~dt'iii!l~ regents :m,ee~ag;lin lin May. -DA:'wI'm GR.IMM, fanuhy." Regent Rioehard Hlunl ceuntered U~January
!n.;~~jng~H IJC

Su fn~njciSiCO.

t~,m:'·!l nn~!!:~f us

Hl ke

ac ad emle f!"ee:d,olm~

thegoveruing oody appeared lIS div,uded as the senate. .. ~.believe 1], yes vote [on tbis b.llll] wouldestahl ish ~.very dtltlgeHHls

~ig htly, ". hu t [accepti ng tobacco money]

AVIIA.N IINF1JJEI~IZ.A

An

lIP~~]"~.'"

]!Il

H5N I bird f1~l

~~~c(j!1!lrtli ~Ilhn ..,

olllb~lI ks i.r.I pOllltn:y aemss Asi;J is dirjlvi.:~lg~,mne themessage tiln2lt

Early this week, rL%eJlm.wlliie •. experts, were porilng ov(:r the puzdetai ls or a recent

eveu c(iiUluliies

'~Iml have elinri-

II1.l'IIICJ.thevirus once !lhall Id n' t becume' eemplace nl. n~.e cent i111-

u ~~ghigh d eath to II iu hmu:!Im;. inel mJ i~~g n:;"{;~ntly de~:ii;:{,.;t 'l\\~) ..i1 cases Qf~n:We~~ionwith CL T<'llninu~ resistan lstf(!i!l in (gypt,~sai:ilg a gd]'l~.remiaderofhow devasiari~g the virus ndg.fuLE. be if it acqui res theabi li[y Hl spread ~;;u;i~y a l1r~oilg humans ..
Overthe I) <lSi 3 'weeks. T~li;1~laA1dl

worrisome cluster oW hUIl'mn H5N] cases ~I] Egyp'~. L,l~:e nc~'emb~r, a ~. 6-jIeiv·old ~ir[ and her 21..;.}'e<I~'"()kl ri.l1!1iC~, li,vijng iQllh:", s.mu:: house ill G~a;l'biya PI'OV,UIlC't:. both died of H5N I iufecrioa, Sequence infDlll1J~<lillon made "~lhJ ic on Gen]3~11l1<QJrn 23 J:mJ.1U<Ufy by I~e US. Nava1 Med'iJl:<'liReseereh Uni[ .3 (Nf\M RU-3 ~ in CairO' shews th~ll

zliDlS ,U1.d potemiaUy

and Vi ~tn1!mr~porlU;:d thelr n r~[ Higl~lai~eri!. Sd IliC~ H S Nl 'Iii rus W~$ (I~~f',,~ed iii! l!:;I.paJlil 1111:$ il'iIOrl~~l~ i It ,~ffi(~oIil:s ~a;!o'~ ~~il~ D~~1.ls !l1r:d~'mlA;lly re&i:>.111.t'l!1 I H::l!N'~ ,mltbl,;eaks all.~CiilJlg poult!)' ilill be-.en dle:c~in91 b~MJ!i ~<lffl'lpleslrom pou1:llty l1e~ghI1Clrill'9Iarm:!l. •he' antivi r.d mug T,m.l.u!1 a, at 6 months, Japan, ~.\~.lich seemed to Tami!lu resistance has. been ~101.Ve dodged dH'::b!lllet ~J11(,;;eit:;dlJl*,il:i' (If(~\!t- 6,.uks later by outhreaks in Jspan, "The reported in a r~'II" ther htUl!iI,llfl H5N.~ cases o breaks, if'l20Q,4, COirofirwm~dlhat the vi:r:m:;.llit one outbresks IIll J~pa!l and Somh Korea suggest fu'fh::'lI'pauicIHs. '!;VCIiC g.i!Vc'r.i1iil,C'drug, HrnN(!Vcl:. t farm, on :~I January and pn)b'lb.~y .1 second [~ me fl'ee~nyill1gbilrds <)5 the must .~ikel)"orid1e Eg}'pti"~1 st~.Ulp~e:-;shl}wirl,g g;~5:~st<l:nt."e were lil in."SliIYS S9 nmlS>, Bo~h !l:lQ[Ll1tries,1ren:liyil,lg to t r"'Ii.m, Oilm [he 23rd" Somh Kerea whklm, last ~<'Ikenj l!I~~:2 d;l)"s <lne:rrT<uni In~1 Il'eatl1r.U:!lll be~.<ln .m umJ~~I;('iI~I}ilih(]npei'ir1o(.~ in, '!I\11h:;lilIDr) d!e"wk~p NO\'e:~lIilb€fsHm~redjl.s I1r.ol(]'litIJ~~k sineecendetermi Ile hrfiV thevirus \"';fIIS rei i'lu(]oJju(:('ld,. taillimg the v9nls:in2IJM,. repm.1:ed lh~~rlhedlf~1'l "Tlse reasous .Ibr li~esef<li~lI[es need to be' res,istam:e, s"\Y'Sf<.I.A~'\l]RU·3(."QJlmnm<ll~di n~ 01'11~l~d l~!f'n~d lip on ,1 f:'IfI1l P.]u~IIfY [nnn. S(;)\t~m~ ex ..~~in~d<!nd !htll€:l'"m)~ ":~rl [4;~d;I~ec\.~:b~re:~ cer Brll~~ Bll)tm~[!l, ( '~fhf!'lt;more, ~1~virus ~n wsld h.~rdlsfomad de2lid i.m,Hong Kong tested ooIJillp.'lltiel,lts had <Ii rare resistam:e'-ool1lft:rr.ing says Sims, BUll ovel<.~liL.le :S<1Ys, ~le:'speed of posi [Jive [or H5NI. And Indoncsh~ Of'll the .respon~c, particularly ]!:J, Than~md~nd mumlion,. c~lli~ N294S., Ol1 O.1m in OI1C' 'p~~ Iy 20 J ,j 'm! U .nr)' reported :~lSf~1 hum~II'I d,e,ll:h i1111 Vi.e-lVl.un. "i~ <I pO:S;:Uli~re ign ,<uu] ~i'lo~'1'~ t'hat s viousH5NI p'<>llieillt inV~etmml. rmm thL;:virusjnj tl~[ .~() difty~. b.riln~U iL~ I~~ di.:iUi11 ~h~ survsillanee sysa.\3tn~ a~ '!iiV'J.:it1king:· 1hm 'j; ,,,,,'hy .Boy~Hon saysche evidence .IUl, mlmllesi.<J. fbur of du:: five: rec:ealt'liUJmliUl suggests "a more di.slm:birm,g" ~ tolm. to 6.2, b:Y far!he mas! of illIIY OO~lntl)f. llleory: lffit'l.l Th~ :i!lC~if~~ i~fl~itb~ks In til~ North~m deaths eeeurred in the J .. karta ~Ur~;,l+ :~n b~lll'i were li1feC't~d by 11 s ick bird that H etn i:o>phl;: follow~\'1ih,n ha~ bl:l~"tuu~ "In fe ~·I;!fpOll:;;e.~'l1ecity goverll!Tl'f;;lIll Oil! 17 li<ulmuy ~~~H.;<~d!y Il.u'btlr~d th~ liIL'mul:ted virUl!s.. Ifm()l~ esmb~[:sht::d p.luern. The l"C'aSOI;l the 'Scafor QJ:dered rt. kte]us vmo 'keep b1U:.kym~d I1cmh.ry .... :such bi.l.'ds C'x! s.~. doctors ma.y see more 0 somll~ty is St ili~I:L{~1\,,;e~i Umlet311)(ld s;:~y:)Le.s H5Ni ~%nieflh who d~,m'l: re5p(llll1ld wen l(~ 0 ru e<.lt will. ()r~Un the.ir bi!rds by~heellitll of the ~ if S hlns.. ~. \~eter.~mlriilllbflcscd in M~munlda, A UI:l;= ~nol1lh Olrillll\'Clhelm.~ <co:mlfillt."3tedarncl destrtl)"~d, Talliil,~,lrnn An(ll ir such fu r~si.sti:'lnt s:r:min Vo-eI'C ~ tm~i.], \,dlO advi'll;:l\ the lI.N"',,, Food <100Agr.iImt.e gtweI1LH~be']1l is w!kin,g of gmdmd Iy :weplaoro $pa\~1na pamle.1ll1ic, lhe \H:¥rl.d's V,JM Tm~linu ~ t! \!:! c~IJnH'e OI·.wrun~Z<llUOn(FA{n.~t ~s likely to !be ~lmgilThi.e ivenmill'keb lhm ClUJi[enl~y ~'('l:>Otnu fUll ~ stoc'kp:il,esrnlighl: be less. he.~p~,lli. i som\;':eamp~,cx imlterocliof'l am{)iml~ $l;;v~r.d f~l;':- 30'14 of pu~Ill'Y sales irl U'le:dry with ~~l~~ocr:. ref",s ar~curr~f'!IIYU!~llk:r :\l!,I'a)'10 dCfCf- ~ tors ..iJtcllud~ngl100ler ~mflel,~nlres en'<'lb~,ing ~.umses."h\\'Ou It! be' <I sea C;ill'<IJlge cU]llJm~,I:r," IlIiII:iine \!'!ihether Lhep<ltic'IllsiIn.lI.d the regjlstR1liIl ~_: ~he VI'fillS lit) survi'!:t; Itm,g~d!'!t~ltH;!'!W~rOiml1'l~e!'!t. so.ysJoln'l1!M~."ver. se]'Jinr adviser ~r;I FAO~n ~tr.;u~ bli.'lfurt: ~h~ytll(li<; ~b~ n dn~g.~ml \'ll1e~~.MrF! greater ~j{)U ~tr:ll~.e jll P.Wi!I~<lnniQi.~l. \1,1' ~nliel.' !for Jaka:n,a. If dOllepmperly.. he' smys. e~i11J11imlling vi:m:J; from ,. Ihi:N, sllsJlected ~.15N.~patient in ~,. fcstiv.als, :l!I1ld ~~m\'c,:melltsof ~vuldJ bUr(i:1L I:.oLckymd ]XJildiry C'ouFd~~~lOJ;:the a;pporiuni.iy ~ll.c ImUKcho.ld. "v.~'1.ollso d.i~d, :~Ili.S th~~ ! s,fIlnC ~.' The :recurrence of the v.~rus in S!;:Huh liortlle \linlSm ::!ulI'vi\'e ~J,ll~Jeemrironlllelll~" B,m Ilmlut<llicil. BoyntQl,1 says. Veter.~mlrY viroloi:S Korc,~ ~llld ~'aj:l;lJD'I particul;;lr'Xy I:lt)wbI.e. [n is hf,.; C!ll,ukn:uto; ~1~~iI1Sl .f1 !.>~ldd~~1 pro:~ibhjoll. g.UStS ;;trc iDhlU c.h~ldng ttl !\GG: \\l'h~lh,cl" the I,Hn~l.he 'i,\·imt:f ()f 2003-'n4· an:d ~hi~ ye:lr. t wllii(;h could lead s.m<1llhold!e.l'5~o hi,d)J;:their nmutau:d v~ms, c<ll'IlJe fbm.ui 11:1 ir'i[]::l,in Bgypt. ~ b olilbri;;aks illl South KOl"t;;O)W~1"e I·Q~I~J~li.~d 410 Ibw:~ refu~ ~ocoo~r.ne w:i1h~u)Jinl.d dis~nd -D'ENNI:S NOR:MmILE .liN D MAI1I;1I1IlN IENiSEIR.1N'l<. ~

both we re 911,1fecred

\iii

i~h ~, viru S

448

26 JANiU~RY2007

VOL315

SCIENCE W'Il'/vl,tSciencem!,g.org

~I II

, I I

AG,AI)EIMIG

RIESE.A.RCIH

New Cell Rules

The Wi500IlISill AtUI1iHli IFliestlilirch IFo'lmda'tion
school and an]~ iated ~ID$iP'ila Is. Oil"I:lte schools gf pu b~.e ~rn.l:m 1~ ~n,d ofell1 sin e'er i111;;and i It .• ::rpplii;)d !'lCi';;:llii.~;S. A:~lht)lIgh rome ~tr.livcn::itk~

Talkalbmu gre!u~1" 'Co~~!llbOrH~kl:~] aeaacross demic dep,arunems is c]u.e-.ap. Bm ~rnlakiml: ~.I:

[Ilap-pencan he-\/I;ry expcnslve, Last week, Harva rd lhri ve:r:sHyn~~~de $50 m m i~H"! ~i
down payment oa the coacept .• including seed mg~ley ~i~r first-ever, ~ll1ii""Crs~ ~y-widlo: a d.epO'l.rtrn,eJllt in dleve.loprnenl<l,1 <Inc! regelmerarive biol!Jgy ...The mo,re: i~ p.,n ofa nil<ljoi,' f':'!!p'IIl.s.ion of the university's science and enlg.inceml.lilg pr,ogrm:nls. ~l1dllding 1I planned 5:.:500> ~niU 1.0 u c ... pu s ac rossthe Cha:lI]e:s m Riwr~haT w<~~ U!l\l'dkd ~i1.r!i~r (his; mo!lI.fu,.
"Theres a lot of I~ felefl :i.nthe dilscip.~:inm.y

(1.'II'Mf) has .~(i!i.ed ha:dl.tIlUl'gh ~l!ce<llsilmgru.Le~ r restrdirll;J .:u:i1diitliT'lk ~.e:;;~ardlIii! i!5 ~roa.'(lly ( p'alt.ell1ltoo hllman 51em (e:lllill'le~. Pre'll'iollJ[s~y,a company 111~ed@dal.ic@1l s,e ~@f1 ~or IUl1i'!l'€,r~ilyboased research
III'S~llg lm'~ WAIR:FIjllle!>.

erased thg$'~ Hiile"S deeadesago, \'irnrking .,.cII"QIS!; d.ep<lll"~mI.l.e,r:tna] beundaries ~'11: ~:larvard is "n{l~],lrlli~ [workingJ'Ilj{i~.~~"vQ ~·niirul.)l' ::i~IMratei 11SI it (Lt ions," s<liys D ougla S Me II en, co-d i~c'ttlr o:hh\;i H"n'\fl'~T(~ C"~II hlM iU~II:.. SlieIH
I'rlym.ull

nat

remk~i(ifi, has been lined, allh~1i.I9ln .lI ~omP'~I1~ mlll)l~lm ha'!rell Il[{!eIWI1~D do il'~ O'r'l,lnt work Qr d'~\l'~~QPI)r();th,H::t~,The fJ!) I,l~ Oi;l tiQfI a 100

will chairtheuniversiry-wide

sciences, but at I:he &"IIIHC! rime ..there is a trend WWil1id COiL~C!lllL'1,tiF.lliol'l fresuu'U'l'e.IJ. andl"~J,';lro seale eollaberaeive seieuce," saysHarvard P rnv 0 sr S:~eveIlhl)' man, wh (1 la stwe ek anaoenced thl;; ,~!:'C<tli(]llm of!! Hi!rv~1rdthlivcr~ si~)f Sc i.~n'Cl;ia E.f!g~ nd !lll;l:r..i og Cum rn i rtee (HUSIC:C). TIu: cmrJimiuee"\\~l~C~l w.i:~~.repml hJ Harvard's presideru and its gov~rni!lg board wi 11:1. the $50 million I!) planjhe new use dep'':'ll1merli~~\'':~.kh\,\lm ]nchu.l~ ~:ttei'il] cell
Ilie!\.e'OUch-.i1lld seed ether imliti~~h'es., some of \vhidl (lou~dlead (n simi lar ~lCW dC'l)~U·ttlll1!rlt.K.

eomm ittee, wh icl~ ~.~hopes wll I keep the 366~)1eall.'"o~d ,~nstiUllion among the leaders in al] fields of sci 1..: rice:aml !;:ngin~erin~ t:luOlagliii.the 21 s~ century And HYlll31L l<lHHQlwd to he Oil! the shert :~iSl of Marva.I'di

presidential eand id '1[(;.t~.'IS$:mli1e;s

of rhe C o.l'l1m i nee .t-1, ,'ccom'mCl1
coM money.

that lI1.1;U1y

dllt iOIJ1L~wHI

··We e]!!'pe>Cl [I,he 'COlliiml:~'~ee]1;0

pmpoctiC a hud~et. and we know that ~ lot ~~lOre win beneeded." hI: 5'~y s, ,,, IU!t d~b; B shows llh.e [ilCLjI~yll;lat we <l.re serious," Ihl;1D11ew d~p~lnmemw.i llevearu a I]yh.~¥,e
25 to 30 Ilrhculty members. :lm:ysHyman-> about hulfmi,grming l"rMIiI C'~1\]:iil~.llg. programs and [[he rest [Ie\V ·"J)Poillil.tII.nent~<!n:liC~

dalri~ied it!> Iees ,~l1d hO','HlCademl:cs, {illl1l tr ensfer cell llnes, Jon Sode-rS'tr!Jril. 1lIl{!!1C1Qirtg dr~.e(~!Jr rVale's o mroc.e of Cl}~~eralivE.' ~5ear('h,. SBlYS old ihe ml@:s \'II@r'~wId usi rig, t@s~JiI:,~iv~. j r'!Cam~sand l>Bill.The (hall1lge is ~c,omil'l gait a \!Iery (trI.lci8ll lime f'Or llis...~ he says., a $I,h~ S(h(JIol is IflOi,:\i setlungl !lip a new 'stem (e~1 p r"K.]ram ..Ule S.tII1I.d ~~il!1r!(~ (ali:r(Jml!r:b"s~d !rIDU ilid'alion, for Ta~pil~,r .and COIl1i!1uRllerR~glM'>~IFFC)s.ays Ihe R move .aUMaJl.es its, C'(jIn<~fnS oul "il"Strie:lWe ~b polides that ~1.IolJld hiind~ifWQrt;; at th~ ~e\v C~lif(i1rtli~"1 n:slhu t~ fQU RJ:.generraave Medki'ffile. HOMlwr .• FireR. slill be1liev~ WMf'~ p'al~iI1~ areil1l\1.)lid, and I~e us. PaliNU and Tradei11!ark mfke hi mrlrentlll' re'h!i1ehll1lngl.madminisl[<ltive ~eq!!lesllfi!.ed by lhe 'grotllp to lrevi~ ~;llepale,llls.
-lEU liiUIN1rliSflliI

1~~.u\'".l'm pmfelliSo[;S have tr~u:.~.itrtollal]y't;m]~

dllj~ I~d~:hll;lir es.. ~h r~ CICkf][I

(Imld t ~i:Jming "Mith!!~ the nes .o:h ·p.articu I.u schoo>!" be it the FaeIJ Ity of A.lltsand Sciences ,~FAS,). I'h.emedical

report tobnth f}\S and medical scbrml deans, lr wjll serve :<IS>; ::1 focal :~10ilrn ~cnthe \..'Or~ fwa]"I.~1701[)people, ~!cluster rhat

or

II mg IBiucks

fOil

.ALS

Me Iton ca.llis "Ol~eoft-he hii\g,~.est eoneenna-

stem eell .~,d,el'llis.ts." co-director 1D", id ..... de'pi~ rlIlJll.ellt be a m the 'lcrivilie.s ofth~ II'I!H.unHe.. w:h,uch! 'wiU occupy spacein the department's Iliew building HI"i! the AI~~lO!1\ c~unpus•.iSchedukd for conrpl.etio.11i in2iO I O. A I~. hLl m"'l!llb~rs of 1:1\1;; ~ !fleW dl'lr;ar~lm:m ''liMI Ibe (lar.l of the ill.slillHe •. \"d'l~ch IHllVi.'h<'ls

tions in the world of Stem cell institute Scadd.~1m say~. the new ··cl·.illic'l~ C>lllt'illp:llemenf'

wm

Th e l.aleslsl{llfl or lineil1[Jf8iisinQI fmJllI' of dis.~ s,i,l. ad'l.l'(JGacy grol! ps on rtlse.ardil is a $ 36 mil~iol'l p~ed,gl' by two nonprofit grollp5lo idelflliry
n@w' Ull0tocula'Ft.ar9~1~ agar IiiIst amyQlrophrc 1;:11er!lt~ ~l~rosis (AlSo). The '!rerUm@cernes as OOe! of~ihe Ii) ro~p:sab~ndQlilI5eU!llirts III tadll>!! ALS via plJ h li~lhed drug 1.ar'9els • .AJ~l1!re5Ung 150 e-xisll

450 pri nc~p>11 fll.cuh)l membcrn, N.m.cy .1\ ndr-e,.~s.. de.1lal ofruliSic :s:d.,enoe:o;OIl ~h~ mt;dic,.ql s:d~)[)Ci~ ~I'ld~ O;Jo.,mnhu!:" (if ~rcp()rl. I...s~ S:~l:mllil,]e'r~ltl<Ll r-ecm.mnellded the :r(Jil.~mI,H~;hOnt ufH USIC ,~"'M~~~. d~~P'<1Jrull'l;:m"'~]l(:(Jhvio~i~ th~ firs!!: ·cho ice"&'(~r i ~H1:.rd i.::;;cpl ..innry ror;:!i.!;t;u·ch i be~.n~ Oii'lhe fiekrs l.lIpid growth. Il-IYI]WI~ '~~peCiS thll: ne\li,l' cmnn1.iut:tl to h,IW;: iii. di.:tillHcd plan in place for the new depa.rt~m:llit by I Apri~ .. ·'Theym;"e k!een~:!D begin I.'>ecm.itiing:· he ~.lo.te:s. J\n(lh.e CQJ,]lpares wliil,ll. FI~I"I,io:UN ~~doil!l~nl till.; rom;) ~mlp CffiOlll'ii-d~ r\:w ~'eRrsago by National. i.Dl.stil.~l.tesof l]ealLh D'~r""cf(llrEh~usZerhuulll for !lHlre hHt::td~s,cip~ inary program.s ~cms:s: tle ."gel~cy·:s 17 ill sli hLtcS [Ind C'c~H $. "Th~lt ~,V~]'[; h i~ t".r 1.'ii;:SpOiIrnNe to the cllm.1gi Ili,g W'O rM science.:' S<,tys .HYDl]afiJ, "AlrDd I.hi!; i~!>(u~r ~"pCi:l1ls.;;;" ""JliE,FFR:l1 MIEIR.'IIIIS Wlt.h reportUng ,by Custan (e Holden.

iJ'l9 drugs in 2;;!,OOO ,nnQrt, "~~·\vep.retty ~lnJ(h (l'i<m<lusted al~~he l~g,iGlll<i rgel5ror .1\15, ~sa)"5 Se,m Scott, I1res1denl of Ihe .AtS 1rhelri3iPY
I)~ve
[0

IPIIl1 enl IInsti lu I,~ 'AL.5-TD~) in (am-

blr:dg~, Massaa:tcIu3€U's,. A.l!.:HiDI is partMriflg \\0111Ul@w@a!llhi@r l'l'~scl!llaf DysIirGphy A:~oci.atiol'll 1M11A) in Tilcson, Ar;Z(il'!a. Scolfs, r>e~r~h s>t3'ff o;~.24 wltl add HI s.ciel1lti!>!:sprobing g:el1!;!' and l!llrot,ej n e.xp,re~· ~io!i!a:l:nm h~tLhya!fld di~a~d: !Il11ceand hut11l,al1lissLJe {OIlKt.ed (BHOU{j11ill MD.A's f1BtloIDrk ·of rlli~di(3l (~~lic:f.. The ~ool 1$to ijd~nmy geiltlS lh;:!l billJnaRiJllMer~nUy j nfllS j n h'Qil~)of Ii I'lain.91 OiJ i how l~Q5e diHereFlce5 aUoct tile df-se.alSe.
Jefrtrey IFiotl'i5teill, WhD
~:5 ~mpiXlr'tll1lg

work

on ~eI~A15 finO\>i s..e models allhe R/o;bert

Packard (en~er[or .ALS Res·ei:lrch througl1 John ~ Hopkins Uniw,rsity in IBallimo~e. ,Mary~[llIld. l1atS
hjgl1llo~~s. forthiit !'l~ parll1@Jsh ~p. Bil!th@' w(Urie~ Ulai~eocisling mlOU5e l'tmdeLs may !l.ot be r@lillbile eoou9lh to ~!!!rw as al guide" -~IENN IFEI, (!D Ull N

or

~,Sclel1cem,)g.org

SCIIENCE VOL J1S 26 JANIJAIR''f2007

449

A. de·cUn,e lin the Iwmbe'r III" ,c'hiil11lp'CIIIIiIZees;fJlViaUiable flllJ'FibiIllHm,edlic:a~ IresleialrlclllIiln the U.S,.liIias :SIHlldced alllfOwinQ dleibarle IIln the IIp'poRu'nines ,alRd eests of5;tllild~'eswiii~i our 'CI(UM:'i!si relliillti'"es
FIIIFlE.IEIN ¥IEA.IS .Alfiel', "HIE UNIITEID .ST:AJIE.5

~he~ine dml !l\~P'<!iI:<U~:) icul fwm unethical e-~'h

resea reh? \llt,~Ofn type:: of housi ng do d:l imps reG!u~,rre '!W~er(;!: wi I,~the mOUiley c{_ImeliQ.IilI:~ ~D :support~'h.e cam oh!;l,ima~s :rolltindy live )U ~i~.f~rs in c~plivity ,;'iIld C'<)Ul live IWicc: as ]Ollig·! I] as N U[ poopedy m~:Ul,,]ged ~"~,K'l~t i eai~~the ··clmi[llp'H'iIf!:.'leresousce"? AlI1d tvhy

~,.tI

was one uf" hO!~f~dm:(;ne(ltlin[ri~s U~.fI~h;:~d CII:p<t ive ch i1n;p21IlZCl!o!S <!V~I i~~Jble bio:n~erlica~ to rescerchcrs. Tij]d~y. it stamls alene, Every country except perhaps Gabon hasaband~nij,~d this ~.ype (')1" experhnentation fur a
brnmMiIl: of ethical,
i1IIJOM1C

also mandated uo hnagl;)i! needed for rescan:bi bemoved rro:~n pri 1mI<Lle cc:nT~'rg [0 retirement sl'm.cllJl.9r.ic~,"A sa. res ult, ~helfl~JPul,Hio;UI has. dl\o]J<ped from 1500 in ~9961D II)) ~n Oetober 100(,_ Nm!>( m~my

ln 20[10,. t~.e U.S. Congress

~hat(l]d~r "surplus"

chimps

do scientists need chimpanzees fOf biemed leal research 8'rmywary'! On one end, the Humane Socirely of the Ul'lil","d St!ll:i:!l:. and the N~v El1ghmd AmiVivisectiou Soc ~elyb:ve' ~iunched .•
C<L[Ln~

ia L sc ie'.n,l~ .. and fie

pohtk;,il
[n:Sliwtes

reasons,

N~lw ~hl;;U,s., Nalh)n1!~JJ

oH-I,e,lhh (Nlll)-lh~n~<lin supdellllllC over whether :il should

pa:rl>er 0 f ch lmpanaees maintai ned for biomedi,c,fl~ research-finds k\lelf r~lcil"lg an

incendiary

researchers who conduct hiomedical researeh O~:~ (':hi~~lp.fm;te~s tiro worried thOit f~~ 1"I!i;m(jb~,r ufbl"~cl~ng auimalsls d~di~ing .sornpidly lh<ll there w.illl roan :rmolbe ,eil:1Dugh ~inn is h,eadil;lg IOJ ill. duff:- .says Todd Preuss" a neurcscieruisr '1,~Yerkes N''lItjonllll?r,~m'~I:e Research Ce.~uer i:L'1. lU~;I'I,n;J. Goo:l::gia.which h,f~.s C(WI'!1:ry:" {lide$t coluny (If!"(~$eareh [ill: chirnptmzoee S" .. U we don "t sta rt breed iI~ g ~~est' cl~·imp.;;!I'!l', seen, ~:hiily're ,!:~oi,'Stu !;:g .. es e

paigns 10 SlOP <ill "invasive" ~;>\p1;l!'~m~n·ls. ~"ilh Ch~mp;)nt:~5 d~"l mi.ght harm dtem and
to .I1<J1'1fe !I11:1, the animals ,rm,~ovf:ld 0 saretuaries, 1
V!lC'~ ~

left te sast ..in t~.epopul .•uion, "The p()ptd~~

Some pfgm"U~;e researchers .. woo Ihn'e mis- !i

ghr~"rmgs <Ilm~u. conductlngme types of

phase out such srudies or breed more animals forlinme generations of il;h\1e~1igmors. The lssue has bceome i.;:,"r~(:'ia]ly ~H::UW over the past feu,' 'II:Jm,ollths.in the face of a
!1!¢\V

cine. dmg. Rl1Id pmhogem:sis

studies ll1m ,we ~

~
~

done in ~h:: United Stares 0[11c:~jmpal1lzee:l>.

..~ 1_·iO'll ~,f~1';(.() a r 1

Ilroj~~~h}!l mb[)l~~rhe f,He uf th~C(lp~i"'e

cbimps sel aside tCI:Ii ibu,Qmedici>l.lrese.'I[lch. Feu the p~~1 decade, N~l~ has i.m.pos!;:d ill. moratcrium en breeding any fedendly MIl'· ported C'h.im]l~. vihich ~TC housed ~u sil'<priIlill<ltecemers acmss the cmllH:ry. :h enforces the balm byllC~usjlllg to suppert .funy ~~~,(lWbfU:IlS"

aW<ljI,. and they're going 10 be garm.c fa.rrgood;' The pl~s;J~, breed more C'hil,1~.P<U1ZC~S is to

forcillg a reexansiaation of question s that hav~ lcng surroandsdrescarch wlrh O~j]· closest reI,ui>~e5. au end<1n,ger~clspecies thaf is ~Ipidlyd:is~.pPG~ring il~ \"'~kl. \l!,lhere is th€' VOL315

ol·l1:rim:;r~(~re~ilreh~ !'S.. ~1mey .1iI.all1omlibrlabl,e with ~l:' S<IY~ vi.ml.o~ 're ~is~: ,hlna~h<1~~AU,lQ:'!. whe eenducts AmS rese arch '\\'ilh ~no~,keys atthe Sourh west Nal.iomli Primll~'LlcR.el!:carchCenter ill SUIl ,A.mQliliOi. Te:xas. one of the :r~iCiiitue:s~m<ll does bioIlr],cd leal ell h~~r xperhncms. c "'You slumkhl '{be cOlm.~lbill1~,Me with it, YO~'I,s~.o:uld
~Ol

~ ~ :ll:
:;oi
;;t

i
r::l

~ ~

""
i"!i

26 JANiU~RY2007

SCIENCE w'INW_sclencem!,g.org

NEWSFOClJsl
bet'M~~l~he: I'€'~~rc:~ and th~ good 11il~~l('''{)nru;;s, orga~s for hUn11,f1!1 ll'llsp.lmm •• hnp]<lu~d m from it&lnd Ibe i:md lXlrl:".which is wll,l!t IUI~)~ elecrrodes into their bndn:s lO study sleep. pens to the !IJ.'a i~m}Is. [1'10 a d ilficl~h 'pli?l!ceto be. iillli]d used them 10 !;l<luge the cffcctl!i ofalee[f yO!U 'ne cl(lmfon<lh~ew i'th iland YOl! dOlft IhoI ,II nd mariju ,111<1. A~,d 011 Soviet sc ie~,l is 1 hm!c~my pmb~~.Il1I:] ith ~l. I.hM S a pmhlem," w ~MCI'lilJl'todtoiIlSC.U~li1une lhc~nwith human sperm to make :..."humaneee," Then'lga:~n.11 growing number of NIH l"urMiHIH:ycnrcred [h~ ehhnpanzee reseseehers, s.\le~ f~~Pn;~L$S ~l Yerke.",. cn~mduet so-e~dII;'d!'l(l~ injiil .. siv~ bi{1inl:H;:dk:~1 stud il;l$i" researeh busi ~l~SS, inI9C'ifl. \..;1\i(;'l~, CUi1~OO ~s eSI<lbl,~sheda ~.et\!'!iQrk ofm.egjDmd primate cenrhese chin11pmllzees that tlll!eysay cause them no ph)~~i,\;a h:;;ltlll~. I BI;!~avinr<l~ff:~~~'lrC'I~en;- t(;lr;.l for basic '~rldc~ill1 lealresearch, Snm~ (l f whoare allowed lOWQX1k at some zoos and these great apes were bred ~.ua. ca:pti.vit.y" but s ,HlC I:mt:deS-1:1 1:>0 slu;tdy lhi:i: popl~l,uion. ILllOniliY more \\c,..re!~~k!l:1li from Afrk11, until 19'71, "There's so mndl, \.\ie Cillm, do w]thom <I. destruc\"'~~"'iIl the UnliLoo States ;~ig;lIled the Convenlion ~iveapr:roach~o captiive chimp~:' 1>ays :~uscflll O'UD ~l1it~m;llti,O:Il~llIIrode ill En.d:fIJIl,gc:red Srede<s of Wild Fauna 3111,dF~ol.<l. w.i1I,~ch barred the G~}~n.elt""l prinuH'ologist ;n the' UnivelJS.iry .• Odirnmi~" San Di~go ([JCSD). wl~ooond~lcl~ i!npofhUi~.l~ wild c'bimp~~;I-'e~~.I~ 199b. of
use ehimp.fu!lz;ee:s 1J()l'ihldy \'l.lt'dllil.€:S. for h()'P~!!I~II S C m~.drespiratery syncy~i<1.1ivirus, as well as treatments for other diseases, NCRR. reasoned ll~~Ll demand eouhl be mel with existing (he ~Ilima~s. In ~,9'91. ~he N~ILQn~1 R,e:scardl OtmDlcit 'QNRC) e;ss.eru:i.~U)' comcurred i!'i.<;lI'iing a n;:porl. CI'i"~,.p(m~eesn R'<'.~'I..'(.~rt:h.Ihm !"("Ci (imm~tldL;d I.;;xt~ndi!:lg till; brol::dii)gm!.lfiu'o-, rium lInti I .200:~. NRC's, panel of e1>i::pe:li't:il d.led ··~m1Jrdlling"fl:;;\~ln~liJr lill<lin~inil~S" POOPI~I:a.t.lon of" about .~000 c:!llhnpa:rnzees, for research, IBm it coacluded Ihij~ihe e;ti.~ti~lg

populal:ioril. ' ....s "mnre i

11:1<111 .m:leqt:l<1te IiO meet

or

repaoductive brilo:~ogy research w:~~hchimp,<II1".:.ee~~lerm sainples; Gagne~u:. :rreLJ~s,. 3.rld ethers cmue~.ld lnm w ithaew body- imas;un,g loch~ul:.~ogie;f;f1ndthcrc~nit :l>C'qut1:1I1i.eing of the chimpanzee genorl,111!:,I:he epportusities <JI'e g rea te r t'~~n eve r h11~I se ~ or.liu'V<I!':ivl,: techniql.l!~~ tu lesm <Jib~'~il !;;'i,r'i;lrythingfrom ~U:!~MUl di:sease,~m:l ag;~llg W behavior il:lui.evohlltion,

f:lced, wi.lh, :il~,cl.'elD$~Jm.g demansls

for cbim=

Sti II otber
ml~kill]g
!!

researchers

cfludQa1

.ugilins~

b.1~:rnkel p:lf(x;~alll<llicm that :ul1Ivm,ive

experimerus

wi~ll C~i~lril~ecs

<Ire lmelhi(.. ... t l

"To dir.'J\M·'1 hYP'O'~,~eti!C:<Jlline Ihe ;J.i:L' dmt~ in [ I~ink docs j uni~ (.fI the sub[~.'L,!I:}1 ofth""\Ie ques~im,ls," S<lYSNO:f,li1I,12TI Lewin. an i:ITmmlllol(JIJ~g~
m!~~
:J::

Bii;ld~I:llf!l~1 Deat'PJ'i:~$$ M(1dk~'11 Gm~~r illi, Boston, whoh~lS dose Ams \1<l{;C~l1e'eXI)er~ '&.1 ~ ~ il1.~,el1il;S chimpanzees and monkeys. "These in ~flj kimliii!llfdi~c\!~:.;iollS need to befocused {In ~ very S])(:'(: if'lc quest iO:UI,$ <ibmu a, pa rt icui ar .~ s,tlQ,dy." let \' i::m.no Ion ge r eJipe:wi IlH:nls, ()I ill '~ chrLmp5 OIDl.d S<I)'s he c.m·1 see any cmn pell:ung til r''£L~mn ~od~yEi) U~1;l igurg~ i:lUJmb~rn Uflhc!n for ; biomed]cai rese.urch. 8tH ]iu: stlJ1es:ses., <IS do ~ m.my o[h~r~n'\!e.o;Hg.flt(lr~. UWIt th~s anim,;;!1 Alilhlltlgh ~~",rch~rs l1'l~l~~mted ~(] ':! model has :l1(!'d~o "eml,olfmous;~.y i,-. valua.hIe'· mcdtCliI1 ad\!Bl1c(s ]n the NlLlImber of (himp'ilInze,es AvaHab[e § past <lmd:nm)' \.~I'f'Uin dIe flitnre. fer lB,reedin.g or Res,ear{h in tilhe U, S.'"
Q '~

pansees IroJ~]esearehers sw(~]ng the emergr i!'lS: AIDS epidemic. NIH Mm;~ed;".br~diD"lS Iy.r:ogr.mlli. Nmnben;;; qlli.cld~ g;rew, 'Wilh lheinirial 315 I¥mn.e<lrld female breeders produciD:llg ~';l!Irly ..nm ofr.-.pri l1Jg by HXn 1B~! tthe c.~. IIIpm'!i';~e AIDS mod,", I ha d i pl\()bleL~.lsnlXllnt:~,egel~go. ,Rese<lr-dilc'fS hadl to p.n:)'a steep u5el~ fee, ~]ll.6~a:5t :5R(lOU~f ani$ Itml,llmrnd experiments of"te:n. used so few IIIITimels [11[11OJi1ly tile most pristine resulrs \,\iOUW! reach Si<L 1 i:Sl ic al s ig;L~ificanee. It <I~,:SO !1 [Jon became clear thm H]V 'l}'!pica~:~}' doesn't C$IS.~ d ~sease ill Chilln]rs .. 1I ndenn iai ng Il'lel1flode:I'::; re~~ Oni;:"~ seiernists discn'!.ie:rod 11m.~.987 ab:M.~l.~I. rhat Slv.: it sanian cousin ofHI\~ caused an AIOS-]~:ke disease ill rhesus, macaque:\,I:~ilLt became the IIlooel of clilQioe, :~[l1'995" 'With lew cJ~imps bc:iJ.lg used EO test A IDS'\{<lt.lc i11,11::>. Ii':i! N<!i~ I Cel~tcl: fOIr N] itHiia Rese,m:1ia Resoun;e:s (NCRR~ es;[aulisTm,ed;1 [mlor,fI[Oriu!1.1i!!nil br~~cliil,g Ihe: chiul,~pal1il~~sh s uppo.rted .. Wl.l:Il i] fu,rthe:r notice,"

research needs for atleast five }I'~~.rs."h ulsfl . expressly recommeaded t.~,u :IJUJ fm;:u'lities .. cu,th<ln~;.;~ (;hi!1~~17_(~~·.$ rnJ:"pt)pMli!l~)nrlcontrol. NCRR irnlpo.._o;ed the breeding <lind~ulh[lml= sia OOIlIS 211td,.OIS the NRC paneleeeemmeuded
set t~l'l< t~ hirnlJ;JIIiIZee M~llmli!,en1elll~ Pro£,rm~lI t Chi MP), to I1i'io.~,itOl"he :$t~.i.l,iIg of tile pt)pn.13t lion aad prallil:1iote ~1mepp:I.\Opr,~'lte use ef'these a ~Ini~~~!!ils. research, NC'RR. simulti;u1l00t~~.~y i~ ~st."b] i~1~l;i-d ~\%'"lJr.kiflg gW'U«J ~ U\lt(>[d~~'1pt;:rt:;;

~o advise ChliiVlP;TU,I~'lI'Ol'killg gwoup has rec\ {}mm~[lded ~x~~m:liliag lilt': bf0Iildi!llSmn!1!torhun three lime.'!,. IlIliOSt u'ecenl~y oIlM.iQ,y 2005. There '!.'V'IS "a l~ug~ number of chilill1:p's l]uul weren't bei~1g used" forresemQh. S ..IYS, veteri-

or

narian \Vi~I.~~$nMarlo!l. a !n~ili1lbl;:!iof rhe ChiMP '1vorkillg gro~lp\\,~.lOmlmS f~ prinrsre ~tl~_"l! Idng t"t~mpruliy.Paris N [~.P iql Edmonds.
Wm.l1ilrng:lon, '~They wer-e ju!>t sitti 111~ there," Wi! II thepubl ieat ion .0 r the ri.rst. drafl of

lK ch~lnp'lIl~~'"e ge~lQmein September 2rm5.

cansnm,oulli!cd I~l!:' CRR 1)0 lifll~;lC memtcN dllln, In.~ 'cmnn,ll.entary i~;l ~mne is,sue of tlm~ NdiWl!. the heads. of ~h.eU.s. pi~inlOJt.ece'm:~ers ~g<Liir'l' oe.I\[oll.;;,j 1~]~ beJm(.tfh ~.flfm~jnl:f~i~ling'bl~is, "'I.m,uque r-.;;:sourc:e'·<uld
\\i,o!fil~d

iliiilil

Ul,H if ~h~

\ve:re nOI lifted. 't~lepo~"ulilti.on

Illorlltori.um wouM s~~up~.y d\ei:lr~mte

.~ IUllri1f'ash~an3ib'le model ~ S·dl;l!li!tisiS, h ..v,~ cnndtlCi~d h~o-

..
ot

\.\:;ithi ij :5 yCil:l'K.
S im.ce then. one'

n~~co~3~ub()r];"J~)bll
V~LI!iU;h,~B~rg. r~Clllr di of S(m[~,lv..'eSl hI!!.t nona:ll

o:r

~ med,iclIl

,~A~
'Lil

~,'iiUl!Zce:l>

flil:-;emc h fmi H'llon:'I:lha:m,

pr'Op(Hltll.lt:'i

on ell im21

'cen;~lII''';I.

Ill,my

() f!'~~$

.~. played'l c~'~~,cia,~ N}~e in the ~.' dev. el?~mem of the ~':tccinefor '!I hepanm; B. Oil somll.;.lu!n~s knhOllI it! virus lb,t h<l.si nfeclc.d 1 b iU iOIll .~ n~':(_)iflle:" Bl!~ :>cit'rll i~f~.unumd 'r~h~ .~ I" I' ~\!!.!(lr~d!mve: .. Iso fUll: rQ3,:"I":i.~ ed stL1.dI! ies that ~tr,e now com;.idcr~d ~ o'i z.a r;re (lr brl~ ta~:~ T'he 1I.S 1-\ ir
I

'til Inall model. note. s;Hcll~ rcscilirch :::>

ani-

,Prim,ne
<I.

Rest:aD'ch
l.1J];lIl.y-' !i1.~nllSof

Cellter. h~" pc.rformod

mo:we det,d led

sis of Il~eagt" ;md heah'lm

,m!

lhl'l eh imps hUUSiCd ail ~Iisix fadthies. At OJ t;hinlp nH~'eting in Y~:D'k~sin Octobe,r 2Q06. Vand4;l:iJe,rg said lluu of l~e ~.133 animab the!l m'VaHa.b I~. j mi.t 100 remales '1\'eJiI:J potenl.i~1 bre~der.;.

,~ ~-(m;:(: $ C~lHl~,pOU;)l'I,n pr()g.~am

~~

'!

ill9'97

]r dIe

brc~"{lli Ilg!lrl()~Wrll!:Ulr1!WC~

hot tliilerLu i.llu) s,p.ace" Otber ,~ r~$ef!rch~ts h;j!"\I~5l~d their

~ :is

l'!Dr

Ii tled he addt:d Ih~re \\rilll be no

ft~;;trch cl.~il'l~ps~ftby 2;037.\v~en h

W'W'\IY,scien cemaq,!J rq

$IC111e: NC::~

VOt 3115 26 JAN U,AJRY2007

451

NEWSFm:US
~uil(lnbe~ chimp<l:~7~~S will have di~d {s.ee
gl,~<IIP~, " 45: I ), p V:'IlldeBcrg ccneedes rh:lll his medel is

NCRR pl1)mlS to OO!~\leml.£ h~.CWflrUj~g grOl,1;P ln .~vhr~<:I.l reassess themnramrium .. whi.ell is in 10

ernlecl ~lllih.l Deeember 1007.

'fur iIlISlmll;;e. i[ doeslltll~uke i11!~)(I ~:i,lCt har some l"aci~ lU!C!S, t their t i a have bred a f~w ell impanzees ShQl~ld NCRR lifi dl,e ~lr:loramriU!ln.debl.ue will d!l:'ilpilcth~ mO~i!lm:uli:U~)"Even sa" hisblc'fL'k ll ke 1)1 .imltel) sl fy over wlunD.)!, P C' 0 f ehi rn p prnj ~et im'l~~t~~r~ed m~~y ~!l ~nl;;l;:~~I1mg. rl;'l\!~1ireh is tihie,lIly ;lC~~jll;lltii~ ,Ind 'i!ivi:lnlhlhL:l ~ ~he i,l1tdllIl'iiingAj it V<u;ki. <II, g~ycOO:i,O~.oglSI,<'1,1 ues D expense=espeeially now thst almosreveryone else has g(ltl,!;;11 Q~~ Q~:the bmii m::~, \1i.'~Q hell'l,~d ie<!AJIt~~ drive h,) sequenee rhe The Nel:herl<lllds, the last EmopealJl counchhnpElnzee gem,Qil,UC:. "What S h"ff)~'!enli.ng now I::; re,,~~y,I di$<!ster;' SOly;.; V~l'ki,. O~1Jg~r::;,Bowd tTy toeenducr iuvasive research Oil! chiln~hal: the ;Iging ofdn~ popui:n-ioll is .dreOld:y j,impanzees. curlawed Ih,e practice as of 2004 iei Ilg b raillil. 11 ~.dbe h av 1.0 mJr~s~,H!rC h tful<ll and hils been 11lloving its colony ol'more than d!epeL1(1s, 011 ym.l.nl1!.er aJJim;II&, ~. 0 animals to zoos ami safari parks.Iu 0 y!;;':n.;G'." hea4 S~Ui;)Jt Z:ui.fl. \,vlm isa Jiil~(,';l~ub~:iOctober 2006. d~Jit ne Jflp.[lr:l~.~pharmaceuo ofrhe Ch,iM II? '\\'0 r'k:ing grou p and Lite ]:!l.illilder tical COnllpa:lmy st.iilJ conducting mvasive NCRR advisory council, enrp~~...sizes lh,1I resea reh in111;Ucountry decided to step: W~.t~.l m .. !iy ohhest: "potential breeders" may rI~~ :wppo rt ~:rOl~.lKyoto Un ivers il)i. l he COffin become preg:nil.nt 0.11" msry~.ot take C:!IrC' of olf~ p!H1y :pbu'ls, to e Pi! iii !:I ret ire.mc~.l sa nCDlla.rJl fo:ri 1:0> 301 ,1111 i111J,.I:;; i11 Apri J," The BeiPudt ls R~$;a ~'·dl Fbuf1,1~1lio~1 i n Poug~,keeps.ie., ":-;imp! istic" ~C"01J.nt lJ1C Q\1I'1l expense,

If,he U.s. g{l\~mm~lmt,h~5 d~kk~ 1'0inii{':Sl naore money iUll, im<luning the research ell imp ~,m~ p:lpll~.~tioJl, Imlll1lY invesl i~I.iO.rs comcml dun .it should re~l~ink whm l}fpe!l ()f e."'~1e[ilhelll....C'0!I!1l be done and how the animals arc housed" "Chimpanzees sliamdd not be [ISIL~ as furry l;e~ Il,i1b~s." )ys B~lU'ioc H~lm, a virol{lgi;.<>lI'll Ul{: ~ Univ~~~~ytlf AhibOl!~l1'., eirmj!~gh~H'i'i.\,~ll~ \ hunts. raJA IDS. viruses i~l collected from feces.

,;lIildc h~Imlp,ml~e~, .who Hahn

f:.'\pt;a:iml;.':lll.\dlm~

~"{)~d~ICl~ Il~)
(I

i:~,v,asiveresei.'loch. consends that "95%

with: d:u:rnl',u,:

r till.C

l~tUI!1~\"l'istlr}""

In ~he~r 20.05 Narnre com:lmlClll.HY. \mnJ(k:Bcrg and C~H)lutllm:n;al:];u~d chimthat p.1:lI.lzeesshould remajn <1\lajlable for tliise~lse rc~~G¥~reh fot '.~;;Ijlng drug."f!llld \lat~d!l(':~ a:~d
Van de IBer g not es t I~<II. SOI:J.~,e prop rie l<ll:jI experime nt s wirh monocloa al anti bod ies done for eommereial C<Hl.lIpanies .~~weled w illness or even de .• h of c.~.i mp~~pl:C'\lel1iting u IiJiilrmflLl1 d~:Il.ssITonl enH:rin,g human trials. "It ~su neth it'~.~ from 1'1 ~'Ult1lUIIll>lilmJpoint i:o !lOt
f"t\:)

lhi:>reseereh,"
In an opposing

he~4~)I;;;,

New YiJrn. \\ih~cl1 has

long eonducted binmtldic~IJ research ata ell ~mp<l:lfIz:ce co lony in Liberia, decided I<L~I:raU 10 release the last of it!> 74 .f1!1il'l1;;al~ ar ies, G,~bOIlI'!; Celltrel nte r:mL<lli©n.. l, de .sPJiil~gii'I:h:ey do. "Irs 3hmm too late un~.,efps, Recherehes Mc{Ji.cale.sin ff!lllccvmC has ~luHe's !'i!JLb5t,Ulli~~lbreed i fl,£, UI,l itiated done biomedictli researchen its small pepuq~1 ic.kly:· says .001.3. larinn of captive chimps :iDllhe past. but A~lhough Zukl is nOD (]il~oflhe~~l., 5~V~1';;11 rUl'~lltt'p~;;ms are U~lC lear, researchers fauh NCRRFor the WJy:it has Mm.'wn says he ~ Lil;a'~to see the Uuired S~fU'l;f;ph,,!~~ n~~ himl'le-dkai r~:$~;B;rch~\tilh m;lr!li!:g.~d ~h~ !.::'hi !lI~P'.lI\IZ",t;l resource, "1vln~1 of \¥h,u Ihey do ~.s.fund nms.s S"pec'I1:'Om~~el'Sa:nd, clillimp~IIU{!e:s. ~OO. ·'.I.t :scul.e~~ 0I11]~.e si.de of cl:iniel'l.1 r~$Ie1ln;:'h cCIliI.lers," :;':iI~SVlIIrkL who peol,I!C! 'whohiil\'C ethi~l~ argUlnCI1iI.s,but for 111[)~lncia1: r>e<l,S(ln:S:' S.l)1S, MOrlon ..... ~r)lmllift \ll(ou~d~ike the brt1edi~.lg. mOI1\I:or,iJluIH liftoo. "I
li1inkah,e chimpa'lLt;e~.;~rc Il. l~um in~hdr s,ide':· E''elil lV.lorton. "",ho SIIP]XII'ts the mo:r.1W· dtc ~m10nlMliiU~~1 •.thl:"rc's going to b~ hre~dil1g Im;;~-min,d,e(iif? lIsin.g MRI, E'me,ry Ull'li'ilm;il'\!"~ jalrTle5 R~Uin\i dm~~ {ompawillil,!\~ arnaLyse>5 or d1im~ add andll1ll!1marnl brains to 5tudy ag1irn9 and e,,"oh.lllion.
O~,1l0

i5:SlW;l.

UC.'SD·s V.L1li.

cOImnclll.tm:y
{:;.f!grrelUx.

and p'rilm~lol'-

il1l the same

og[st James 1. Mcoresrgue

dun chinnlll<lll1Zee>

:iislan d

S<LIII(::t ~.~

Sh(llLld beusel o:rl~y iWle,xperimeuts thm could <Ils~be dOl:ne elhic;ruHy imJ. imn1~lns. Thill 'lilr.'Oldd rule ourvaecine !>wdk:s ~h~l bu.;:nt~.omlill)l infect animals "vi.t]iI! potential]y ha II' m full viruses, for instauce, [v.e!1! with sud~ ,~unNI.mi'IDiIs", tlln,ey predict de'.I1.~.,nul for animals 1.0 rise, ',.~ SLI15'pectin 5 te 10 years from now, 1~,eTe'sgoingw, be IlHIIC:l\r reaterinteresr g in gtl.~.dyi~g ,chimps in 11OIldilngcro~ls. waY5," !;!I¥S \~lll:ki. Uchesechilll,lPl'> die off, he 5O,1,jl5, "ili::;t(u}' wiM look back and SOl)' .h.ID\'cmIld .1'(J~1.et that l I'ilftp]'l€:tlJ_" ln 'u;lmt'ircom m";~lm]"y. the)' a I~], ,C;)~0d 1 ["or TI.1.cn::;a,>ing nllldin_g t'O pl:ov~de cl~ impanzees \,vill~"f]ptim<11 living conditions," sucaas
'(,Ili~.np H.aven"s I~ci.hly .. V.~.H~G:e cUn.imps c<ln ~oC!i~.lize o~t:door~,fi)r ~mlC'h Oft~1C d1:lY. §ma bow Il.~:,nlyc:~imp.mlees tbe Cmqllltry rtOO~~R ma inlail1lQ "'iahl~bl'('CdiI1gpO.Pl~hu,ion to r-emai,:lu <I hol,-blilUoll! iss~~e" NilAe' fl:esness .. 1,{,\>t;Cluii\iL;; di~~~~lrlflh1;:I'li~\,;~~l~tiol1~1 ~ SNci.;.":.'O [nIOntl;M~(l1l!S'},iJl:l\lI ml{h~ m~~11h!;irofthi;: 1997 NRC' conllniuec'~lotcs th<llNorth Ameidc.:lll
L:~g 5· SlI}1

I~ke cr.ll:y.

<11111<1, once 'il;ga.in,,'\re' III h11\~e<l.~ I 'l1'nese

ri~n~J.(;!iit~dzc..; NCRR ~~1r i!;,,~~U~!re haId to r.egUllm·nll:\.'tll,gS o~~the \'I.'Orklllg gJPI.mp he ::;ilS,
~o IbilmN i[s <I.dv.~ccOlnd develop !) f~fIja~,~ cli.ll~ !lOO ,m.:i~m.gemeln P,~;'~lIbr ~he P'UPlJIl<!llion. ··Smll1cbody needs, to £m sp thfs tlh:illg !uld )Ohov,' So:Jine l!emlel'$hir ~:[NeIRa:' .5H),:!! Mono.lli. who
<Cl111,01'

chimp ...<lnd~\lh{l·II·be f<\lriP(m.i~g 'th~!llT'

liolll

d~.;.nwem

]al 11:'>1;<'11y~,.r 2~ Ins, NCR:R 5~m s:s..'7 mi iQiI1l,dlimpflIlz,ee cal\e.,. More dlirl1l. 201% of
Il~ Ch~.D1.tp

Haven

iil:1

Keithvj

pre'\,~onsly

f'dl1li1e

NCRR~flJ:LI1;ded priln;neceUll.~

tell"~n Ihl> U~h~rs.hy nf'li;'a~htl'lg(Qn. &:rltUe,

·'\~,11.~llU lhie issue S:.:II1~d hal are t~;le r.,et::l?"· .l:I1e w

Lou isiElml.l~lI.C cmmtn'y"s, Oldy publ ic 1)/ :fllnded S<l~llcl:~lml' fOJ the rel:iliieme~n o:r~tlrplm; duin'lp;lIllZees. and that anlOum is; cen;Jin lOrii,se OJS ~'Ol'(' almi~l.fd:;; ;fUll! roli!!"ed. PriU'\~ alolog,i:;'l Undrl
B.r,ent.
",,:1.10

ne..

h.w~feWl:.f ~hml 150 l41 ian pml.01:ees. dl"ll

he<ll.d'lS Ch,i~,llp 11.<Ivcn,.

\vh:uch

F~e~Il(::.~s and ,00:lmcrs should !)iIVJJp~ysupport ",,('Ore breedi:lm.ggr-ou:I) ol'ehimpa,11zees,ID:1l' ~ bkl!netl~c!!J ~CiLreh;;;$ 3!71 ]nsuf'<llnceroli~>lfO!l! ; Ifut u re e III erge :UlI.c::l,es.Be ~h Israe Le 1.\1i[I

IfuJ.cy sJ~]arc[0], blJiel!~ling,

the u.s,,,g(we:mmemrt

John ]-[m:"din~. whohi,;;~ld!> Ir'lf~!mHe re sou lI!.:es far N (' R:R .w,o'IIJd :~HH,ex pHc: ilJy addrc:s:s why ~H) I.ong-t ~Lmp lall ex is IJ.. B ~H H<lwuling :stresses ibtl:he linstinn~iODii ',iorh close!)' with the p.rhlH!.~C Ce~l¢[S o:Imld hflS ·',n:t~.ve policies, ami proce>;M:S to m:lI,l<lge OIlll,di oversee;;it~. ch~!1Il.p~r~z~~ r~sOllrce program." 452.

d~p~!ldl~ tm b~~:~ j:llll1'Uc .u-nd pritl<1I'10rl!ndilmg.. q uegt.iol1!s hiOWI:~,e fro ern II gove'rnmenl oem Id
poss,ib~y af~brd to hr'eCd I1U1ai'C chiml~il~l!Z~CS, The c:>I:iIn<!.ted lifet~,:lm.1Jecos~ c<ning for" of chi [l~.p"'il:lz.cc-c,;''1ptiv~rn~~(,'!s i\r,c fo~ )0 )"Calm ~ 01:13~~e:mge., lem,lies for 43-r.,.nges from $3,(IUJ~!UI tfl $S:U~liJOO•.sh~ no~~s" VOL31S SCI~NC:!

~~!'6'e.C: 'ii\"ilh

Ihis mfnim!.lli~ ~If<n~~. '·rf\'\.~\~

rs

:::I

lem:u;led .m.ythill~goVI':.r tl,l,e years" its l~l'U \'I.'C ,Iioll't know ""hHtt'hc next cpidem~~dc \vi~l.be ::JJnd ~,·1.Hll the m:.....IWl'iI<ljQf heah~,lc.ri:-l~s i=:;: g:oin; to l k:' ~ys l"1v~n .• '.It \..n~lld Ix: t~oo~hfl[dya()~ke t .my pOlienti" I an ullmll nmmk:1 the; tilble;· -]Otmll (f)lfl~

I ii
;ii
~
:::'l

{In'

~
~

2:6 JArN'UARY 2007

w!,iVVl,!'.sclen(;;'emaJg,org~

NEWSFOClJsl
INVASIVE SIIP'ECIES,
Oil ~~1~ [i;V.f1 r~!lk.~!lg-\j\io.'!t~r 'p:~~n ~~ar alse d ~$ drawwater from Lake Me ..d, Another big i C!llUl.oer;l'!..is the petennal effecr on sport. fisherie!'; ill lilt: 11J1.~,e. mussels filter huge f~.e

Feared OU81glga MUSSI,II Terns Up in W1e'stern IU1lnii'l'e,d States

S~:je!i!lt~s~s til')!] ng to assessthe pf:te~IMallfor elJol~o,h:;aland e(~n Gmi( da magie'a~telr are
fallrndingl a, Irle!Lllthfitil e<~ tl~le: '~jlllil~j1m!!ilUS zE!illra mllllrsiSel in U~e Co,Lotlrlldo IRhfll'r For .2 decades, ll\e zebra mussel has lormented the Great Lakes, A.lo.~.g ,v:L~b its close CQ1lJS]n" l~letlmlggllIlilU~s.sd" the [.nn<l~ liefl~!Y proliflcmusscls have clogM~d •. e h i,:lfuakep:ipes of power pl.a:l,lts. cuated the been on~:~]e ;al!ertaller several close calms,,, ln 2004 ..for e~.unple. ~h!;:y found dead zebra mussel s on. ., hfllf·,uozen hall sehoats arriv.~llg :U Lake M,e@d, UH1Ukllemercrs from the most westedy sisJn illllg of zebra oou!lSeb. NO''!.%li~·~ clear that the q~lagga mussel alread y beat l he zeb :U'3 'I'C':U'O S.S rt~e' G reat D.iv ~de. .0 1116 Jamu<II'Y, a diver dOi:ffiig ,I reurine illl~pecli on of 11 bre'lk\!,/;_~t.~T fULIl'1 da Cltu~ssa mussel at a Lake Me,ld lU!lrun~" a :WewI<ilo.
1I:i:il.eterl)upsl:realll

rc:nm.Q!vi!l£ lII1),WI11:;tJld"tDn ,lIlcl boost~~ mHwie;mns.This seem!' to h~ve ha.!"I1·Hxi !l'omn,c:rc:iR~ fhi~ stociks]1lI £on~cbut

volumes

or water,

rlll~all (~f~heGraat L~kxs. An emergency task force of NPS ailid ~~.~t~ '!.gif;lnc~lt:_.;n Ca ~iforn ~~I 'l!ld Nev~~d~l i hi
SCI:<I

mb.~~:~g preveru to H~mporm)' nl~'I,'H!re.

nm:~.C:J spread,

As

<I

Pmk Service

boats

!l:3.vebeen grollnded
h1!V{~beentold

and cencessianaires

not m transpon rellli~~ 'Dr:);][:5L T:~ey have r:m.lped up im;pecti.0l:1S ofp:rh'<Ile

h(l~~

hulls I)n~o'lts. l'I!dLt~r()w!l~O~)syMe:!I'l~S (If ~ out

whack" ,\Nestienl stares h:lve been so C{)I"J,~ cerned lh,.u in ~. 98, l:bey :!;rtmteJ..1 nnl$~itm' 9 clulilpllign. c~.11 d the J, flOth Merid i;Hl I Ilii uae t lve, d.e:s:igned toprevent beaters fronl accidenh'LUy trarrsporting the mU:-i:;Je]:-; uf ~llh~re:X~Jlk

D[II'I1" "I

lito III Hoover was heartbroken,'

species to their waters, N(}\\" 'the gf!thi'll]ve

h it. S. suffered iI.ilnlajor

defe..u. Earlier n:hi:s :IDluom:h.

[ll!I."g.sJ,'] mussels were f(»m~d i I:l

Lake Mc~~d. a SO.OOO-hcC't~r~

reservoir in Nel,'<lI.d<l" ndlast A

s at n~ lakeas well, Cali fornia iSlry ill!;l . ., ~ojncrease boat :ilmL~~)ec~:ions at irs rheee bOIW=del" :l>la~.i_ml:ll l~Thateceive traffic fro:lml L~kc r M~ad. Ami! M.lk:ol'Ii1I~.genera] eeurrsel for the stare's De~'l'l:'Llr[mCliM ofFi~iTh and Game •. says l hoeageneywi III ask the legi slarure l~~or the ~lIt h.ori~y to inspec!. anyposs ible ~!' rc ~nj~~ f ,.\"'Un: r {If! bum~ u e uf]d torce owners 10 drainthem. But the !lHJ!S5el.seems whave aheill.dy spread dow:lmsl.:L"'e'am. On ~1 J,mu'lry. divers \\,'il~l!the 1Vle~ropoiit<l1li Walr.:r D'i:Slricl ( M \l),' D I of S,OI,H hem C;;II~Jor~d~
found qlil,1Igg<lm:~lsseh ~~]l<!l.ke
H~NI~~U.~(;.fI:l· ]!lUlke-w lh4! the

\.\I>eek. they were eortflrmed dcwustreasn inthe Colorado ."" R:ive.r. There 's no sig'~' {'I f ze bra ~ mussels yet, bl~t observers fear .~ l:weir arrlval a.so11lly a matter of ; time. State off~,ci.<J~:-; iu Ne\.,.~(11<ll. ''ij: ~ ,1m! OJ Iifomill arc cOIn''f'l1lilllg sci.~ .... enti fie ...d,d liifY~"jf pane Is; tor. gun~
.

Colerade River Aqueduct, w~,:i:ch !!UPp~. ies \A~I_e]]" ml 8 III i.I:~ peoion p.~e, "We are goinglo be t~lk.ing <IIgg;l"c's;:s ive .<1(:1 ~on:' SJi)l'S D ebr a

M.IIl .. MWD's chief ope,-aril'lg officer; They planl to use ch,]or]ne or t'OPfx~r :~Hal!i,tl~akill ~~Ily us .. t m
sels or larvae .il'llhe ~'H~6~klll~:lol]g

~ mill "'th~]~. o ex pect ~Ull.dhow to t

Q

; ~~Ip~ wirh rhe qU01.ggi~. "Tili$; is an orga:~js.11fI l~~nwould be dev.• ~ as.mt.ingla,r >California's aquatie Ii ecosystems and for fl',eslli.w.ner :~nfr<]~ i t>1:n~ewr!l;::' ~il_yi> Rich3fd S()c~m.:'ilJ.water ,.. .~ policy <I(.hfise;IF rQ;nln:('alifQ~ni., Dep.utment
:~ flfW~i:~r RL:)sl.'lul'Ql;':s. ~. I3tlth zebra ( DJ;t:is.wmtl po(rmm1)/in) <lind ,~ q~l.mJgg.a (0. l)j/ge.m~is} mussels are native to E.. stern E!JIl'op~.They Rre tholJl.ghllo Imvt: .~ aI';rived in tJI;li: Great Lakes by I~~C b]tc 19Sfu

e';;~R~lll!clbly" (,l:mlgg<l mussels

saY:iiWen lB<lIklwi~l. presi!delliH of I:be L<lke Mead Bu~! Ow 111.:1"); A~SiOdatioil. alter l!:ire' diver sl.mwed ~,t~ohlim" Sincethen. divers
ARI.ZO:N.iI\pn':fer de·c'per. ccolee water with rclatlvely few ~mrucfllK so l~.1(~JI plub.I.bl)','!'on·t ~hrive jill ~he ea mi.." I!:',smysl!cl

with ~h~N@tio!)fI:1Park S~n/~c(: (NP;S J,whid~ nH'I~ the l ale Mead N,n io~,~.1 R~'crem ion
A:rea.h<li~"er{)ulld mussels illl fourl1I1l.ore I:OC3.timl ~,Based 0[1 dle :-;i~e of lh~ III lqs~dS :iW.:m
to ~ 1m11 kkl~l.i rk~IiII fll.r ..OIl" Robert McM:fIlmn t

Milk. ~Hii i?eolugillt at theCm'u~~H BiurrigUc.a1 F~eld S~at ion i:lll BF.udgeporl, Ne\"'¥O:Ii'k" I-[u'!iv~v~r. ir .<,;~br~~ mussels urr~'ii~~us\.~'GIL d~r;::~hiJ,!atitl!l could b,;; \\lol:... . Not oniy mig~l e zebra nUlsscls~]ml,:n.sl~ in the aqueduct. but
~:h\:!:y could

<'1I~o ause seriaus p:WQb~em:s c with

.~ via ~he b.11.lllSI \\'iU;etr of !!:onlainer s.hips .. In s-m,lle p I<lDe.~". lJ P 10 700.1)00 z:eihrJ m~l,sseJt·s .~cro\vd ~uge'~~~cr a single &{]jtH!ru: ~llGI(:l~. By hil. .~ c:.ioggin_g i:tm1a~e piqpes.. the mussels COSit till!e ." po-,vee.!" i~ldl~~t f'}' ma!~>'mlllil(Y~$ til' do~111!'S ~11 i

of Ihe Ulliive;~.i~y of -r:ex.ts. Ad inglom" esl iIlnm'e!> l:Ila~ they h.we been in the lake fo:r H1 ~~.1!$~.2: y~O)1rs.1l1~ .. (1(lfI~\..'!U~r~!!l(jlnl~'ky d.u~pl b{Jiuullm,offer <1.111 :i:de."II.laoil'ltri:J[I" the quagga ~illuss>l;ll ~,n~",s Ch,;!!t~ RiilJ'!iLch<![l;!!l, ~jmnol· So a ogislt <It L."m'el'ltu2I1l IJnuversily illm SlIdbm<y. Cflllail2l, ··L~k'e MClld is going to~~tvC <I hl1ge ill fl:'St,n~ioll:' hepll.oed:i:cts. >0 ~a~.:sarl)' \Vt)r rkdl abo ~Lt.~~~le~,11JIa.Ct ~ 011 ~ loc)\"Cr D"lm. '1\r~id~ 1111 year g:em:::lt<ued $1: 33 bm~(l!Jrok i~O\\:au-h(llln:; n$' ~h~crl!ichy. ;;undi

.~ ,uld~'d .Ul~.ainliemm.ce., I.:~lexo""dbly. the IlUlssel::c; ~ h~l\f!C spread dO\m 1:11..: Missi~li,irp.i Riv~l'and

rhe Immy t!lilches :find ptpcs. in l~]~ 1~1]per.iult [f[]g,uiol'l Distruct,. v.,111i.d~mike$. ]rs v.,'lJ.ter from l.1iit.e o~orado River. ":h's; cOml!Ceg\'ab:lle C ~~U~ e1l~aL~0ol~Ld r·~quirl;;' ~xteii)sivl;;' !H~~!1J~(:.' I"Jlauee':" smys Mi_d.l.,.e.~Mi:(unoto, lhe~ dis~ ~:iic~\;;b~'ClhJ,g.~~1.'I1 ~~mn'dn~t Sl~p~D'vi:mt e u Fm.' the .nm.omel1l. hmvever. sc::i,emis:~l> .md ~1r1!a;lllag:crs will hm~ enQugh~l;:~ldllche!:o;with

.~ I;hmugho~nm.w;h ot~~·lin.e !:<~lem U:mited St,lte::>" g, J:\.~aPOpll~fir dcsli ~iLlioll for bnal~s~mm ~ ;)(::00 88. the ,counlry" 1l~lke N.le<~d \-\'.u clearly iD'I .~ thl(; ['!lU~S\l]S.' p<lth. a~d oIJidfl~$; n:her.e h<1Jve

..

me

~he ~Ulfok'li ng
]l.r1!lIS.~~,

",ff;;:steHill dnllm,l~.or

lite q1!q~~sga

&1jI:o Rmnc'I~;;Jmll y'a.l~ U!\t h()I~g~].l ~•. ..~ frollt-5'e<ll t ickel:S tO~lle i11,];V.<liSJ:OUil.show:'·
-IERtIIKSnlKSIAID

W'W'\IY,sciencemaq,!J

rq

$IC 111e: NC::~

VOt 3115 26 JAN U.A!RY 2007

M EETI N'G 8,RillE FS,> >

2.mnH MEET~NG OF'THE AMERICAN ASTRONOMICA,ll SOC~ET,(

5-UI JAINIUARY

SIEAl'TLlE"WASIH~N(3,n)N

Three-Headed Q,U,B:S,Blr Premises to Shed t:iglh't nn Un'i'versle':'s P'i8S,t

Triple pkw~ are rare II'!baseball,
b~llll!(lt as

"ll~mSfllrs 11i1I~rgcd, the two b.lac:k boles would lbe';g,in orbi:ting each other si milmly to a

Dim!lry :!lHH sy£H:~!m"

rare as triplet fonmu.cons of '1Il.m.&lllfS. il~,~a;;. In n!c:t i~L~~ilg~5 d!::pict,~I,g m~l.hipl:e qml:-KLni in dose pro:.x imity have always been mLUlindy
inwrp retedas
IIIiraMCl>, d~n'l?rent

lm~i:>llillll,i;l~ll S'vii.:t:~ri~I~d. m;ing dU!{1;from thl;' Keck Telescope ill,l Ha.w3u.i. sponed evidence

for

t\

.hird

i..... qm~.s;;lr. m.'by ,<It ;t1lOul tire

f"Jiru~r iW.WIl .he o~'hwith the [urop~lln

ers but ,a~o

same rembi 11di:stance.

v iews or a
k!lffW!l

Further observatinns

singJe rea] qlJ;,I.'l~U, B~~!i!i!!I~ ,ahVf!Y!'. Ofm1,lgtllly I OO.O[}[I

quasars=-cosnuc beacons lIml be'~UlmI luuriI~DUS l:ad,i!lli.nn <I(;I.·OS:I> the m~~,vcrsc-aJbout ~00 h ave bet: n ,~d~m iJ~ as b e~I1S P,l irs. ed QU~LSl;l~.'1li, mcpom\:cl'e'd by mas;:!;;ivc black hoi.cs in the cotes of gaht1>i:.ies,. and when 1:\"'1:) such gabll>::i.(':!'icollide, ~hch eentral

Southern Ohser'V<lI:ory's VeryL::lIgeJ'elescope !!li! c]~iI~ ~(]nfirfn£ld[~liJK ..d findijrl~Effor~sl'o expla in. 1:Ii!Jleu~ p~e~ as <'I gravH'H.iolul.i-~c:l~s.ing effect h'l:\.'c f.. ilcd. Djm-govski said, m; .~".\\e efforts Nl find .IIlY iuterveui ng m3~,"S b mly t1'n.!I.t n Id h~ve ivc co acted <is OJ h:IIS,. Alld scrutiny ofthe c-olors of

E"''i;;mua~~y, stich()rbili.!~g N~ld hg:~~\'Vil:1 swirlcloser tegether and collapse .unto each o'~'l]er. cX1D: tli»!l h1Ipr~n;j.~ol;~i~\'!i;lr. B another co,.I:~kli.l1gga lax)' could 'brj,ng <'L th ira black hole ~o ~h~ dt!rl.c~.'I~· which peinr the :irnlllnll{;li!.:m:> gel moee l'l[)n~p~.icaled. R'Isio's si~Ii1I.Hllatiions ~hQ\v" '1"~clb{'t': black :~olcs ili'l!~cmc~ml'~(:r \/,uolemly <ludillnpn:di.ct<lbly." R;u,io said, "W~HI~ happens atthe end -.. is t~J;;~~ black h!O:~es b3.sic'~]lly kick tlhi,e:nuelves ant of the eemer ofth.u g.:'1I~iI.N:Y:'· Their 1r'~J:id xit occurs afteir the latecomer e black hole :pi:l.ilr~ up wh11 ene ,of li~.eo:rig.iml:1 1\"'0. gi\'ili~ the relhaill'ill'l!;lloser" gr;Wil<lliOliJ'lll Idclt~~ hurlsit out ohhe gfl~~'>;y. The !)~"\N pa:irrc~;:uil(jin the {'1ppn-'
she dire etion ,. :~ both case s the 11

hk!{.':k-h.()I~ ~wa:-;~~r~ retain their

separate idenrities, so p<li.:I,"Sc;au~ ~ulf\'ive<ll ~~I~l lor a ,,,,hi I~., COmSiml Or,,] third qU,1SiU with such a doubletwould he an exrremely roue event, but :,lSUOIHllnCrS :S;;L~i they h ave now witnessed the birth
q\!Ol!$i!r

black hales depan their p~!rem

galaxies at speeds relIIc'hillg thou" :mn:ds er even ll.:lI1:-i onhol!S<tmd:-.of

kilameters per second.

lfipT~l'<;,

or

"TTh~.;:~~ spcsds are high (:~nough

that the black liIol,e s recoil cerUlI!ll), etu intethe l~alosofllw galaxies ,111ld someti mes (.In even be k.icked all. the 'way O~Lt, ... meaning they beceme \\~.,n:deri~lg black ~~.oles dun go tJmnlghell:lplY
intc~'llaQt]c sp'Hce:' Rasio *,~i,d. rln the observedt ri plet quasar s}1f\;lem. the blllc.:k hnl~~\l ~w j tiSI h begun to ,eOUl'ILt': together 10 per-

"We: have tou nd the f:irst case of ~i physicaleriple-quasar system," sa id astronomer George Qjorgovsld 0 r the C filii~
~,I'I::;.~:i~HteTechnology of (('<lilted) ~n IPmHldeltl<l. leader
of au ilIiI.h;:rmui,on;)'~~ (~1l,~'i report-

foruia

ing the discoeery at the meeting ..TW{i oflhG quasa.1!'-~ the group .. dm.~g~ i~l nared QQQ 1432.-0 W6 .. were discovered be Fu rc .~9:3 9.A s 11:'0 IUl rnH:rs ,t) rig i[Hili Y believed that d~,epai.1i'was an :~Uu~iol:li ereated by gr'av itllliorn iii I h;",gi:!m~ (if a sing~.~ quasar ..Such lensing. <L:n e Ileci of Ei nstei ~'iI~~ g~!~cr~ll ~:~l~On,l )r r'I;lJfi~ ivi ty. Ot;;C~1 ~ rs \';~~m,ln intervemng inm;~·'sgri~vi.ty bends the light arrivjng :Iro.I1l'l, isu~m objects.,. ln d
~1il<Ul)i known cases,

light

~1'1~iW;;1dby

th~

~IU'-I~'I'S

revealed .~\,bdt::

form ~hei!fp<~rln~u'-S\,~''apring, daaee:

~hoG'Y'Ire

dlilrerellii,(\es .. I'l!irtjl~.er sU_~~S!.i~lg that the three tll~l~s~U's~ll\: dils~.im:::L"n usalmoiS~ Ciel'l~inly is
1l.Q'~<I
w'(:

aw 1:(:11witi:! ehe aiLl2'rlli111ivt;: cxpl~.i'i~uion

So ystem,"

glr'~.\'li,!lni:mmllens:· Djorgovslk.i said "so

~lfml it ~s ap'hys;i,cal9'ipl~~I<ls[JU'

Ahh(al~. l>~c:i~ systems should be rare,

~h1;ly .llf!;: Ii1etenr irely :ml!'rl'fii~~n~~ ~~l!1ld'Iheilf exis teuce COli lda ide tTcms [0 unde rstan d

$m;hlensi~ bends the the dy~;l.m~lics of £J~lmt)' mergers iIii the carl,y universe. L iglhl from Ihe u.ip:llels I:~ght firOlllll!l diSlfiJM !1i~liI:sar snmuch lh<llt it ap'r~ear)j 10 arrive .u f.uth fro,m ~llIll11"h:Lph:~ dep<1n:ed on ill'> journey to Eanh n'll.ore lh~ln ~ QI hi 1:11 yem,1; <]go •.'H :.:I [nme ,,!,11efll g<l~axj,es, ia~1 p,"",thw'IYs. creating the :~lliiIlpress.ion ofmon~
th;rUl ~f11> (Ihj ~CI"

:sti~1..beut II00.000 ~i,g;IIii.l-y~'~I".s, apart. The c:n,t:ilrc proc('s~ of .ni'tC'rg.ing~lIld evenmal 5>ll'lU[~ ~~i up 'iN' ~I :1, takea bourl 00 nlllill~iou years, ng RfL~i()'t>. s.i!llul,m~'I,)i1J~. itdiClllJ;:_ i "The process [liml Dr. R<l5ii,ohOI:!> modeled is \~ry, "~r:r'nr!ii!;jJ[h~1~I~v~" ~.~t~I;.)!lOfli'IJGr S1iid Virginia Tri nlbh: ()f the Un~\~rl}hy of c..... !lifuifala, lrvlne, "So in some sense, the ~)redtct'[on h:,~'ibeen ve:iifi~d by the 1J~~r\'M~ion.,amllh;e o'b5en'a~.uonll]lLS been ~"'rduim::d by the the- :it

e\'e r. M<t,IIii:y"sl.roncl:lrn~.e rs con~] wl.de'd linn it c:()ll[sit;t~d o.l·two d.b;iillii.ctllii~arhy GI~21~1Jr'Sai. ill. Il:ledshil"t Qf 2.1. i;orrespcH1Jdiil~gtQ <I li mee !i~~;len th~ unliVl$VSC W~1> 'll.m~nJ bi Ilion Y£;o:JQ:i> Qk~.

Later ObsC:',WltUCUl!S ofd~at qllmmr !»Is~em q~!~:.ahm~d t~.... !~mg~~mtltrrol~~li{l!t 11uwm

oceupied II $JiiI~.a~I~Ji' univers~' ,fl~HIc,;,r(l'!lIIding Ied to freGlUellH 00 iii sion:s:, "Th~ S,I; hili,;; n'!c ~iOli1!$·.!r~ i,~f~Cl ~I'lqit", emml.lo:IIIi .... said Frederk: Ras.uo of No.rth\¥es.lem UniVCf&iity in Evt'l,m.1cm, Winois. :~n
f<liCl.. :l~~8C~llllp\ner ::;jrm~da!jQ~,lS shQ'i/v th~l

my." 13m lOO m~n~ClIli ytr<trs is " ]~ng lillJ.ll!:: 1;(1 '" ~ \\"Ij[ t!) see \\11ethe:r the fm!illre It!el~,v:i,or ohb.e ~. ~~pl~roany 11~~toll~$lhl': dlc;;~lre~.k:~~1or,(;~a;.or. ~ r
Evidence
{iorX'llf

tlll,1l such, tr~.p~,et.illillemctior:l,s

:5
'''I

c[ll!h~c{~m~ ~t'ltl!\er. O\"''i;;~~r.A r,~'~~~ t:i h I~d p;<Iir'of b~.<J.ckhol.e..~ ~'Ould soon merge' OlI.nd ~

More '~!,;'Cntly .• 1Slmnomer.;f.l,um DII~e~h O1!lidthe Eco~e p(l.I~~ech,~li.que 1:';'CI€'r;)!J~ d~

g~.hl_ j:;;S '!."l'o~,'~doft-en ,",oUide a~d :m~rge: '[0 ... form Im.ge:I' galaxue:s. (]Jnd\1'Ihen t"vo g<l~ax:ue5 co!u;;lh~bilg c\:!HU'<l1bl,flck hflie-llm,>I¢l!ed
VOL31S SCI~NC:!

generalegr~vi~ta~.uofIJa.1 \wve:;;~h<lll).ught be I Mliec!able. Rasio ~miid. And there a~w e~ists d'nej:lfl&..;;:ubi.1 ity ofd~I~'t il1g ;;L w;;lndei.i.ng black hole speecl:ill1lg Ihlrollg~,l space. ide<lUy not h'C'!ldil'ng 100 clt~~ 10 Earth-

~.
~

2:6 JArN'UARY 2007

w'l,:VVl,!'.sclen(;;'emaJg,org~

NEWSFOClJsl

Mi dd Ile,..Earth Olein i,ze!os

Si(i~~11 blil r.;uh~r ~.I·~-llo;;1dngdw;;tr:f ''''!thin i~~

the .Loc'IIGrm~p of ~Ia;"ies. Zucker s;lkt Leo T corrtains both old andnew stars. g eaerat i~l,g'l ~(l ta ~ bri ght ness mm h out lya 50,0000 rimes thm of milesun, The presence oW new stars is :.iurpr:il,~in;.because IlIilOM. other d\\.'~rfi; ceased !>t~ :falrmalion~l:lng ag~"

Mo,b tbe MU- Way

Tile ml1y~~e:ry f o
1li1I~lY have

Ihelil.li.:->!l.in:J.~

dwarf ~J:Jxi.es
of la{lkin~li:]lr

bC'C11 ~()lvl.Xi,I~""~~{Id

dlwarf'K.

~u~rof!(]l.n~·~ ~ln\1

M~~'!\,'I;.'ll~~"n St,~,ii'e~,-

-'This ts b~!t>ic;O!lly ~he :>mO;lnl;:~~. fui,u~~l

star-forming ga]<Ixy known," Zucker said, "The L(ll:<'l] (~w~!p ,t;~H~id 1~ fU~l;1ld~",id~~uilill objet;~s like tlns," MO~O;l such d i1Tl. f ree -float ing gil h.•.x ies

illg furhobbits.

rW:;;~'ln';~(;1r.~ lrom d~e Slo<JIfiD'igit.!l Sky

SUf'I'ey repened d~.ed iscovery of seven p:revi~ IF;eculild"ho(~I~it/' Leo I (ce~u.i'llIi'i11.iM~· dlll'l!ter) is the omdi' ~u:lk newn dw'lrf god", 1\ ies :'i~~rro'~li1d:~ngsmalleM iin(mll1l s~ar-4()rll'lin9 gl'llal~. ~he Ml~ky!jMIY.plus <1[1 eig~lth .llt' .hec:dg.e of ~.he Mil,ky \~l3,.y 's,gravlrarlenal il1nu~ncc" A ]'loO'Ss.ible (:Xp~.fl~,;II:iOilm. for the disc~p,mlcy. Bec;lUse those small S'lla.xi.es were rtliLnl(l.i~~. sllppm1txllJ~f Ihe Sloan.ll1es'ults,. is 't~.'a'1lmany of a

could help e..... p~.<I]n.he kl:ck of dwarfs. in pasr

obs:ervl:11 ion!>" s,~](jllNitjfl!1 KaU~v~y;llJ~~ of the

$urv~y ·e) f o~.1yo~~..n fih of the "~k)l.dozen s

more 3Je likely 10 exist, team member Daniel Z ueker of the lhlivcn;ity of Camb.r:udge. U. K .,

[he d\,!~-ur:[ aro H)l) d~!ll~.ube; eflsUy del~t;[l;':d. s

"These new dwa:wfs me extlre:ml.l.ely faint," Zm:ker &llld. "\Vc 1.'C r~ndli~g lots of them, and they seem te be mm:hfu.ilfl'ter t~an anyolm.e suspected ~l[lxi.es.collk~ 'be bcfan::. So perhaps ra ther ~hanl dwarf gal 11 ies, we shcul d give x thenn ·the na,rnc "~]ohbit g~u~;fIxiC'!l," , ~

l1itlrVlnd-5milhmlli;m Celner for AS~I'()pJl,ysics in nl!nlbr]d~~. a~~K'hus.~fl", M

"This :mm~s,~ing=sa:rellile:; roblem is reaUjl p one of the greal.llrnlys.·~cl."ies:··she said, "The POin.l I!ri:<ll there could b e Ir(;.l!::-flu.uill g: satel-

said ,n 'the meet ilrag..

GJlnp~Lh~r' ~il1nJlIli:l:timii:,; ased en current b ~heor~e:sof g.d<~X)1 fOrnl,,~i()n sqs@esl ~~<'Iil; huge

~lh.,.;jC5i. s.ui~'nm;!11,c Min:.}" \V~'J'jsho~ld be sur..

r"llun.d~d.b}f d(l~_t"!l:-; (If'even :t~u;~drod:;, 1.I\~,url· of ..

sasellires. Until rocemly;. however, O:~llya few such :s<ue~ ires were kqlOWI"IJ. I

li.les h(mnd to the Leeal GOOllr,p m nOI. useesb sa:Ii'~lyto any :lIillajor 1l1emlber. of the Loc;lII; G roup is \.'("iI')I ~l'Itcr.ec<;tng,~lIld tl'S SOn:1~!Jljll~ i lhe ~ighth nrthl; !1:~W'~flla:\';;~~s,dil;:~a.gruU~~1 I~a!:ti:;; !lnl;nud'-'~ldish\'1fhe!"!i you lo~)k .u Ihl;: Leo 1:, is suanger .sti]~. At a dli:st<mc!I::' of 1.4 mi I~ si mu larions.' -1O'M S~EG flH:lllEID HOI] lig~n-ye<!rn.i~ is proLl<nIMym,H a M~Ik}' \V~, Tom Si!:~gfr:ioois a J«fiil.@rin L~ A.n!!i~li1s, (.a~ifomia,

Type ~~ .aslro~!i)mer S~e~h.e!'l~olds ·ot N~Clnh(afl)liIll3 Sti;ll~ tJn~VErsilY in !Raleigh reported at tile lIIleelimg ..The Nal~ rrevR.tIII. ~arge ilIlJ1l0!Jnt~ of irro:n..iii It'Ulale sign. oJ ~ ~Iumin.'ontldt'ilr b~i3t5l. Type III supernnvae rerna1'ml5 (1m lain mQ~ O)ii;ljg)e:n than ij~~I'l, "Il~re'~ ir,(I[1 e~lrywh~~e;~;l:.lf~oerl is h·ardt©~lrld" iillmund the Mas~ R€yn:o!d::; said. ~SI1~am Ule Si3t1llll e til1l1~, e'~ oonnmu!d.the w prest n,e Clf clr(lypnrotl:!llar ul1I.Ile-rial. "nl!al miillellioill, 1tt')'TIoias said, ~l!!gges~'5 ~hal KeIP~t'rr's~l,IpemQ'll'a lIIllighl beiong ~(D .i;'linew class ,oflifltle la 5tljmnovae h(lll1'l a ~lig~u~.y· -tmasshl'€·lhal'l-~rag:~ IGlrogefli'tOl"" mare
MaJppin.g U~~ d~ri<.niii!!i5. Usi!'19 aEr1ilo~t llOOQ I~Ol!lI5(!!J dbs~i:'!I1ll(l UnH':l wi'l!h I~eo HIlI[I:D[IJ!~.e Space rl!!t~!imp~. aMmmomlers have produced [he h~ghe5l· r~~OhMi(l1'llma P )fiel·of ~ ark IfIHIHer',> distri bu tiCl'nin lifu'! ~o\Sfill 05. H~f ~he ~ l.arges1 proJoc!. U'li 1Ii~ has eve!" b ~n '~!Iln!!' with tile spl'!oOO L>eiles(Op~," saitllfi4fck S{ovi~loe of the cali ~OI'nia IIn~litll~e ofji;ethlii @[O!lr:l!' ((a~l'e(h~ Pa5ar}en-i'lI, ~rin:iln Il:~pali nw~ligater' or!lh! NlASA project Kt1 OYl'flas COSMOS 'ifo r (osmil!: IE'OO-

~\\ascirr(l!lI1iIsteU1!lr mi!l~l!ri1!l morE! ,c;mJm:m~.Il~Y!i!!'1l1ll ij\\it~ Type- Ii !ilJlp8mOVal@', ~~lnkih OIl:(U r aneTa m 351live star ~u rli!S out ((I~IP p~e5.aflld expl.ode-s. X-ravs h'Om the ][6011 sU,fI~rnoll.a rremnant iIl!(;Dr.aec by (mandra. dindh the (i3!fiE1 [or

'C.au'!I'hlt. 5p'eedilil'gI. long [iJeL~ewd to be ~h~ Mm:.y Way's mlost IPm.muneFl~ SIllte~Ule !lillla:d~s" the M~,g,~Uallllh: nOl,lldl~ mI.;I,¥ lI,lrl!l eut to be 11Iler~ [!l.il(!;s~rs-by .. Rather In(l'l'! pur'Suilil9 a I!l~urr@iy othit, bot~!ii1@ ~(!rg@ .1lind $n)!l II ,",~a'gellt1ilnk e ~®ud:sappeiilr' ~!@ be Sipeedi ling by a l more thal1"l300 ",mollI1Ie~.ers ii~r secend, plJscsibl.y r.ilIil~ 1il!11Il1lu:gn fQr!ihem~o !ScapI! 0'1.11'ga~lIXY's 9ravilY" ~W'e do ~l~\i'e ~!ilQW (Qi1!~ider 'the '~S$ilbililY 'thall Mle dOlJdsSlfe ~~rha~ uobound to' lili~ M~~kyWi!V. ~s.ai'[j Nfrl~ ~lUlIo1!.~lL~ or th~ Hi!!'i!'i!rdSlIIllllhsoni:m C:emller~or i.!i6tlrop,hyslc5 il'l (ambr~dge, Massa('iluseUs, ~Ili\l'aydl amlllf!'O (It~,~ ~m(lIfiOmerS ,caku'laled: t~t1 cilOllds'v,e:loCilies by u:singl d.al<l [rom the IHilullbre 5p'8n T,ele~m~e lO UI1'lP,81fe' the clouds' lJIiIotiOl1 oIlUOS;S; thE 5ky to (fj!lU of wry dI~stoil h!l~(Jj h@l1c@ ~t @'~~@c:tij\l@lySi.1!tiOIlli!fY) qu~sinli. The Mil!ky Way (,[wid ,~m.l. eep the dsw:h in o:rbi~ k IK.aUiv;:aya~il !iat~, iF tl is l\~ ce als rnasslljle aswm~i'll E'S;rifll(l~es Slt!Jgesl or nils iI'Ivisi'ill.e '11.31.0' I)r 0 ark 1mil!IIIIH 1'I.i!I'~ odd s:hi!l pl!! .. an :Sup,ernellil'iI SOLlUlti·OI1lI. The !ipall[e·bas'~d Chnora X-Iray Obs'El'f'!latory ~as helped dealr lip ~ long-s~."llding :my~~~ry abou~ <l f~mClu~ sUp~mOfl;'il, :SjRlUed !r,.y Jolla nnt~ Kepl~f ~mI 1604.1 he s.~JIP~mwaha~ se~l1I,edti kelv tg b~ lr~ la~ a UlerrnQl'Il;Idear ('x:pl~osflj):n ddonated w11efl a wh i~e dWarf star ,acc.l,IIlI1Il!l~t~. tQCI nmJ,t:~ 1iI~,1li.li'!Q rtmJiilin~ta'ble. )Itt SlU FQunding l~e explm;iQII1I site1

lu~ir.u" $lJlrvey),~he prQjec~ traoed UH~~OCil~i!)-n,)11' i dilrkl'l1at~er ~(m:ss. il wilde ..-~, . '. pa."d1 r '!l~- s!\,y Iune , "~1fI~ . U.1i:area {O,Il'UlIol! "'1 ;·It, 1000", ~ t..·.anolls. d"IS"I) ,'I. .1. . ~ '.!'~s .I'll - 'I\i- 1... [an~e-5.O'lIl to, a de plh [OfTes.po[mdl~ngll:o <I tililile wihen the u f1ive;r5.€ WiiS..aoout half f~s :llr€Seri!ta9~. line dark m..alte!;; u!1lknuwi'! maooftial that 9i\i,~(In!'lll visible ~ig'h~, WiU dt'lleiteo !by the waV inJ]. glr.aviL:y Mmred tile' shapes. or Iaraway ~:i!Ia_:.:j@s.im@ new 1l'li:l!p:5, ll!!lbU,sh@~ !i:'!Sl ilii~kin trolu'e,. sliilJl'llj~h!i!t dad:: millt.le.r p[o~de~ a 1m1l3!siw s(an{l~diIfl9 i! mlmd whkh the U!.Ilive;rs!l·s moi f1ary (or '~oory(mic~) ilinaner (Q~g1orn.e(a.tes. in dlJs!er~ !DhJala:Kie-s" ~With(lLll this. da~1( Inatt!aif, •.. Ilhe u ni\lle'rG.Il!llI'oll'ldn'!. e:ld~l a~ it ~:!itooa,y, " '~il:l COSMOS: (011[1100' rator R:i(jharo M~ss~, atro o:~C:!~LKh.I'~~reare h~i1\t cr~'ii'er,~hat ordirnHY " h maU~r dot'S ut. allNdJy~ mim:ide mt~t~e dark~ma:t~er ske~!1to:n. ~1rhere aU! "iOO'le inteJ'I':~li ng di~repamlde~wilhil'l Ihe mi:llP, " s~id Ma~5-eY.PreviG)LJ~ darkmaU~r rll'appin9 bly Anthony il'ywJii !:.If th'l! 1JI!l~l,I'1ilf~iIV Q~ Catfrr(lrm~a,. Diilvl-s. and (Qlleillgl,.le:5sho'\i'JIW similalr df'S~r~p.am;ies.. "There' f'Ss.~melh Lng hmn.y 9)(1irtg o.I'l,.~s.a~dTyson, W~Q i~ inwl.wdilm thfl COSMOS, Ilroj,~~~ not "'r.s.
L .. -,,4 L~

W'W'W.sdencemaq.!J rq

$IC111e: NC::~ VOt 3115 26 JAN U.A!RY2007

455

M EETI N'G 8,RillE FS,> >

SOCIIE."f'i FOR INTEGRA.TIVIE AIN1~) COMPARATIVE IBIOLO(3,Y 3-7 JI.ANUARY 2(}07

PHOENIX,

ARIZONA

... N'ii:iir-p'E!rf'!I!ct ,eyes. Vampire s.qlli:d Ilens'~'$are .., dEl5ig nil!!il r(jr' ~~ciIPl® d~l3ils:. C'ifC'!'Iiii! '';Ii nt!.!l~l d~lrkn~s$_ .h.cl.p'_~re ve 11I. rhis by fu rli~~r ~~;;tbi~~,z~ the p I~g p.~\HdilJ.:S'lVGCI~i;;y ri1nt~_lim .1)dditin1lil.li~t:: igh h

positive da;:ar_ge he]ps keep the proteins apart. <l$!i1(.,~i J1ghow ~~ 1'!~lytmted tl~.esquid l~n~ is to do its job," says J clI]jatl~.an He Ill" y. a d\welopmcllw I b:iologiM"~ the Unii\'1::i;:-;i~y of m.i 110]5>. UIib<.l,liI;1u-Ch,,"np'1ug:I~. \J!.fhcll S\vc~ney scquelloodlh';u~S-el}'smmn g~~.esfromeaeious ceph<I~:opods. ,expec~ed she lO Hnd[bal <'III Th~s~I!I,;t'!f;pr:~)H_:i~s had descended from 01 c.:OHUnOI'l ce·p:J11.<Ilopod mmes.lo.r I:him had ('C!-op~:tIdt1'nc liver f:'~cZ)'me. lnsread, the sequeaees ind.~c,ued tLhn rhis adBpt.'1l~.O:I~. oflhc enzy.U~M~into~cns pmleim. oceurred laref;"ali:~.·tihe.1I['iceslro.llt I.""eptmtopod ~,ad hc!il~1I1 to duv(!:l¥e into. sq W1.~d, C!(:'iopil. and

"h'~

o'th~:rspeeies Ho\.\,m.u~y~I:rhahlpud :~L~Mages indlependently To ~wdylhi~ shnpler protein farnlly, S'Jh't."'e:ney divided the' ~lIid :~ell_Sinto foureoacentriclayers and tl!]iLfll)i;1:e;dits protein MJkenp tu;irlSllilI,m'iS specrrcsneuyend QI~,el' wchUil iques, 'Whh l~(,l' hal p of Du ke 's D1lvj d the details of apanel cf'ever-nasrewer b:l1ou::k Des MOIrai:s and Yih-En Alldre'\.v Ban. she I'ig;lIml wl~ire slliUp~-~_lhcl\ ~l Dl~~ Uf1iv~!'Nhy ~11 ~u\;d out I;l'v·oi~ltio!l3ry r~latiom:hips ,;Ij!nang_ DUJh@ln. North Carolin», ;i:1S <'I g,1.<Idtmtestu~lluJt.s;..quid's. tf:trst111lIi.:lms .Uld del·er,I,~ljllled hm\" den!: in Ihe' bb of Sonke Johnseu, she cemthe protems fold iI.IlG, inlenucL She and M~k~ai~ .nzQfdle University oflioridOl. M biued those resultswith biochemical and modeling data 011 l~U;;oprleal aad ehemica] Geinesville, .also sequenced 600 genesWol" PJ.\open~e:l of lens plooH:im; to n:oons:tm,cl:tll.le S acry~I<,I:~ns from .t dQle:~.other eephalepnd i 11.1 istory vis io:~..i n cep halcpods-vsq u id. species, ",Ifslile breadllHlr;Jppl'o.m:;~~ Ihat ] .11111 i11i!,()$~ h1~pl1'e~edwiUl." S,i1}IS I~QmJ~3$\Vo~cml. oeropl. <md their relatives. From just nne ancestrallens prote in~\~e rtebrates started <I p.lliiY':!!,uo~.ogic I ecologi s.] a~.North ~ Cm.·'OI.u.D1i1 wi~l1~\f~nUI-lh~S(l nl,l~!\Ie in'i,ierwbfll~l\, h.wf) SWte Uni\ie~il}'Un Ra~",ig~. R.e::searche.us i~l:e'd.d,yknew that S-cl'ystill~j.lls evolvtd lens-based eyes~.gtut. more 1l1all once. h~WCB!1 C;ltU"J ]oop .(."li)n!iilll~U(x!l W the ~_nC~gLrn:~ S"("C'~lCY :U:CP'l:m~d::tt I:bli; InreE.i.~g. The wn It ., eue of the f.~ ~i Illes I[ h a t is nn liver enzyme. E'lrl.)lon il1(, ev{}~xll:iou, ~hisleop people have di~~~Cl~cl the o.rigin.s efthe \~:,~shu". BUI~nn~w~rS~"'JB[I!.Ui~. \1)_ri!ll1iI$ •. it eornplexi tY~11l the ]e'm(' used by mallilY i:sk.1olil~er.'I'Imdllrn.e p.1U!:ei 115 m,c;e umre pos itiveiy \Wllt;;:!!'!!Ali son S\\"eeli1~ywanted ~Olem'Tl]I.'IbOlDl eycevolmiollil, s:he wel~[ to sea..\Vllri~e the ship solled beneath her. s~.edisScecood theeyes of squid freshly reteieved fD"l(lm .~ 000 meters b~ilmlJ ~Ild lest~d h.fl"!iV well each IC~lS !\.. o~v~d ...

rion li'(Wl'ilfli~15 u~dear. b(Ul·'~lrook:;Mke!~.ere'S

been :Im.ovelevohnion [oftlm.e enzyme]

came up with this SQl.u~

ill lcast

twlce," she :n:e~'l(]:lted. After her study, Sweeney is deeply imll(presscil: by ~Cllh!!.I~. \'h;ion.i:I'DlJ~d f;:&l{: n.o~ed\the sh.ipboard tests showe d dun the \~Impjoo ·S(lukr~ lens, wl~ <.!]nf~~n:dIt.lrly ~n ich d'ie evol[u!icmmy hi story of ~e:p.~a~()pods. "has a visual <Icuitybeue.L· than in a state-nf-the-arr Ze:~~ di~5el;~i1.~g microscope."

or

IMlus cll'8 F'iibelrs hiift :Inlo tHigh IG,ear

1

.s

c f,ef! tu res, :s"Y~";':VO h~i.o.l~fI!"Y I i blolugiM nldld Oal.Jr;lY lrf ihe Uni.\'~r ..i.~yof underwater

One look. ata bill~crnJ1'<1 <I.~ SJ1n~' pin:nLenes and P'1.111C~ (H,rh'{~ home the' Ir.cmm~kl'lbl(: ~b]~.i~y of mlr muscles ~:o...fia pl·todiverse b;~(lIm:ctmnOC:'ill c J'I.;'5j~ulliI(b;_ Mwmy Athd ~nd 1"~{l!lll~iL"Rob(:i~S,. biomec~.1<Innsts :It 1B~-m\.1Jj1Unliversity. hm,1C IlI1mV

ch<lrg~~l S\'JC~!~l;;y~~)fhxt The elder, f>hll!i.\:!" prol-ei t~:'i wen: e\'<t;:fl'~)i d istri bL!l~d lhRUJI.gh.oul
tlla.c t;cphill.opodi 1e:J,lS. But there W&'> iD.gr.JI.dit"l.lt for l~ern,~1r~recent c.r-yst'i:~!in!). wirhalmosr aoue h:nthe cc!1tcrimcl, relm-hldy .mare ncar the

~o und

th!l~

f1~ c~r~1:'Ii nmuseles ~le

Cfi!Hr<!(t, i~.~}t

CBU:tOn:l,la. S~nn. iBalrbal'3. See.ingcleady UlLlde~."'I,\1"al~eJ ires ;1 ,'jpec~.\1 Sjp~.e·.ri.c.a;l~em; requ with a hi~~ refracnve index illil.l~.Cccnler but

a lower iadex toward the ,edg;t: .. This Sradais; achieved\'!' ilhprogre ssi\'e Iy 100\'f'r COllc~m:ra~ ~t'lmlS"ill!1\! llle :~i;;ns·sCC!1Jle:!" au lf wdrd. of prolie~IlS C<Ined cr-ystaHimt~. Thl;; .t;~lle~.;~~rctysw UV!1$ i.;;\iQ]\I~d fnlm f th~ JI.lIp~,ic.ued gml,es of sm.all e~;lzynH~s ofi!cniflvfl<.lveci Qil] !:itl'es~. re'spoIH:;($" Vcrl:e.bra~e::l ha,"e nnll~ipl:!:. fam~J~es (l,r cry!'>ta]Ufls.b~.t ceph@ ~(lpC!ds. h~rv~jus.t o:n~ ~111lli]y.. ~liIeS~cryS:I.\niIlS. \,,'h~,ch des:Oil:nd from a Liver !;:l1iJZY:iIiIll;;_
t ion

edge, r ru.nOUllits dropped ,n° t11eec.lge]l~ll: ~ Sweeney"s <Imliys~'s suggested dw! [!:Jere h,,~d OO~rl pf~:>ili1,;'; seiil;;"C'dofi r],')r~ln(l fl~lg.~llfl(lps ]!l ... ~

amJ fame. :~ ffi~rlhes.c Iml scles have a bul,llrs ira".lwulli.101Ik mlm:imis)!ioml. s.nysAzil.i. "Thelr snLdy ... takes us one ~>tt1:p (,:~ose1l"o t ~m.d:en;t,md in!;!.110\\' skeletal muscle werks Jill
act!U';l·eaaimals,"

\r.l~)'their

shape tu oolOllll!,;e

nr.~dlor :j.p~-1.I

:says D<l.vi.d O.'lIT.Ue:I'.:1 C::Qm~

~.1la.e younger pmte!i.ns .. For ~!Ie-~:;, Itl 1.'11;; lll".'I!'!!sIMrem. cl)'~'~ullillS

U1J1IJLJ!>l

s11l',li' folded ane! ,evelrll)l dispersed

w cre~

alit: a gla.... y ~~[e, ThAt:~ irl!l~ u prob~em in ihG s (,.~~os,ed,;p<lc.ked pro'eilli~ orO) len5~ cr:mer ..iBm i:un ·Il'n.;:~ {1utcr ha I r Qf~:l\iel()'nn~. pro~dtns; caill~m'C' [he freedom to' fOHn ClmJ:l.l,ps. <I.lJid c,ulse Cf~h'lI,ftl!CtS_ It '~flllid b~ that th~ Iflng;errloop
VOL31S SCI~NC:!

]l~!";.u~vt:: ph};$iol~)~h,~ gil tl~1.;; University of Utall. Sa]'~Lake City. lIiille' d,na C,;onld even i.tas:pir~ be~~~r mbolih "TI1e~rr~ul~~ !Hay ~l!rt!-· \riti.e t'I .si.ml~.te me.ms for 'Ulwm<ttieol.lly varyi~.g th!c ge~rillg of~ robotic [m.m;cl~] w~.th changes :ill1 kmd..·' ::><1.ysWillli.UilTI Ki.er Qfthe Univ~rsij.ly of NO:li1.hraq-:n~hlo:L Dli.:lpd l~m, The body has seve rod k ill.lds of nm:scl es. each 'C~,;:mrfK'E:erh~~dy a r~[kl~~'~f b lT~li.gllme]H

!
~

~ ;i

iii ~ ~
~

2:6 JArN'UARY 2007

w'l,iVVl,!'.sclen(;;'emarg,org~

NEWSFOClJsl

fiv~ y~,alr.s agCil. r@:l~ardli@fS W@ii@ l~fij~ttill by.a d~rllpOSiirlg \~lall@ 'Calr'US5 Whey round 0111UIIE!!Haor of (aIHomia'[j. MOIll~'rey (anyml, 29[10 m,elers I!!m:ierrrrat![. Th~ 'tii:m:as!S WaS home I~ a 'thriving UlmnlU!lHy cf baC:lter;ari Ued tu'bE~\,!(DmlS, ~ail'oo Ose(}ax. erl]bedded in its dec,ayin.g ooHlles.Jhe v~s"

ible wow'S we fi'€!' all rle.ma1les, ead ,aueifu:l'«I by up to 1.00 WI;er()scClpic males
IDhlllt liwin
~~W.

the rellllille'5.gl!1~aI~.il'lmlsIJUlbll {Sdll'1J(e •. ~O~!IILy 200t'l, p, 668}. 'lli!I~, and '1011111£1 ~hilJt they have ad~:stilllcl~ edd but highiy largeled dlf'lle~ell1l1T1ent: ne~ falil lQ Imah! re, ~",(Epl w1'lh r@~fJE!!::t11J' t~e-h .c!1b1 to Ill!rodu ,~~perm" ~ ~lly S.ince 2002:, Ro'bert Vrijelfll1ioek .and ihf~,w~le,agues ,at th,e Monterey 18ay Aquarrnll,UI'I I'te$ean:h In.~titute in 1YI@~s UlIdilng, (illi~@lTUllia, havevi,~n~oosevU e,r1lil lITII>Orie \~h!i!h~hlln:!i, }Om!! of ~l~~h in@! I~esslihan 1000 1ITII!:!~llrS deep. A~. the me~'l~fljlg, ~lfI!Hill'lebjo~oghl Gr~g Rolyse (rr S;::rrigllP:S ~111:iUnn.io~r o Ocearilography illl.5flli'l lDi~go" (alUmni!'!,. ffililortm h.ding at lsast nilll'~ (}SWGX'MUlm species c:mUhese «l'rlC<I~5iE!5" ~1I1hey;;!!Ir,e finding W1~hI'i' spede-s e1Iii!JrylirJq.e' lihey go down. ~ 'XiysMark Maft11l'1da~, alB expe~iirl1le1l'1'I.ill limbllY-

biOI~9i$~s hi!"'~ t~k~l1i~ (llQs~~lo~ka~ t~!llS~ dwarl~ I

QI&gi'5{aUh~ IJllliver~ly of ~awairi, Hon.olclllu,

R.oIU~ii!' has ijllslJ brou!ll'l1t WDmm-(!I)w~l!l(i ''1h(!I!~ b(llll!s

~(k

1.0 the ~ab,

~u~~e he lJI~~d' elel:JrJ}1iI,~nd mflf~u:a ~laser ~.(al1lnl ng 1m iuosUiPY to d~wH iihe' l1na,~e-:;' ~tHI~()my, lin his ,rtarlier :itl.Jld'lte!!. ~QUSF? flOitked Ifl"~ If1:ille the OSMflX WO[m5, ~oQkLike !i!lnl',a~ of l!JIbewmlll~ lyph;aUy 5111e1fl deE1p"OH!an ill
~Er!lt!ii. Bmt~e

n€w wo~ks.hO'lvs lill€m~(I ~~

@'II~~

I~ss w~U-d€'oI'~lop!oothan

~ihese larvae, drwlat(lry

1)10 mOllith. ~IO alnus, al'i~ IFIOgut at alL l1h(lft"S I'll\)' 'systerll1, nar ,a1[1,Y 1r~e initemnal.stores Qr bac;tll!rialhM :~@tm::tm C depend on rQrr I101!lti!Shment, I~ repoHed, S·m.!!~~ew11Il(!lle~.heave tot~ o~ yol~ rlFOm ~ne !!g:g (rom whit:ih!hil'V werre Illo:rn• .tIIrW Ilarglli1.r lII1i3il~!il1av!! iill,rrm~~

Tit;€' dwau~ i!I'IJaileshav@

none. :!i!llgge!i~ing that m1:lb:ls,dep1i!!~e~hi.~ ~1~OUlrU! &11l~ey gfQW .• ~III :teadl ~'~ illU'! ma I '01] ai!Q~ SjperlilQ ,ill diHe;(l~rll :s~a!les I(lf dev'e'IOi~llVi!'~l s sMf the rIl.1!rn wo~m':~ boody Glvilly. Rowiie dis,cGwr,ea .111 ullliQu!l' sj:lElirm durt t hat openn'5 au l <lit [he~IP 0 r the male's head, Hi:s.~tU',dies indkale that , SpE!fif'l'I IPf@cl!!f!Sor eli ~I'~staJr[ (II!! l .ill ~hl! WOWnli'!l he~d, m~gr;ii'te toWoi!~dIthE! '~l:dIil.~ a IiIII rn nally Im®\!'e kuw,udi ajJIain IJe.r'ore beirn ill I~elea~ed. Raul;e ~d previilJu~ly 5U!:l'IJ~~tl?d tl1iM i'.I!;rt!miil~ ag~,!lhey a.ccuililiulate oil hillf1el'l1l (II male~. N'elo'lldillil i:la~ed O!il WIOm15 mUe{led d UlUil1lg r~(emlt ' ulOOeil'WiJl!ef ~~diliol'ls b(,l'itress Ihi!ll(iew, \vor:k sugge-:stslhat imfQa~ur'el'w>or:ms.hat.l.and en bene 'hlelmllfl,~ ~ellllal!!i1."",herll1i!1~llhm.!l thall ~eule t • on ~1iilsliJl!g~@!1na~€' t!Ulbe:s b~o!lle fI'lal~s.-iJ pan@'m kJ1ow!lin at le~~'t oo@ o~ller an nelid, RiOUJ5'~ lIDe; i~f!d eut, n 1f~~ ,aiilaJl!ys.is ~!'th IrnaL@wh€Jt~ worm is ~~.ith'!QI, ays lfIi'I~rilli~ !biow" s !;llist Ke!I1IIi1t'th Halany'Ch QfAubuflii Unijvl!rsity imlMil!lbarna:"jl remimi:l$ US ltia.;llt there a~e sHU fIIl!<llny il1!l~r~'M.if!g discaver~e~. ~hQ;l aw.d [us] in the Olea ns' d~p~hs;~ -IE. P.

ne

01" iudivtdual

fibers,

Biceps

have parallel

Sini1Jlllatiom; indicated

Ih'll pinnate rauseles

muscle 'C03;ltmc~edmore 1IIlcleU"I:~ghlererees, l

fibers nmJe..',;;ceias fast contracters, for~~"'OlI]1;"

p.le.. I~, conerast. ealf muscles are punmHe. width short fibers.arranged al an
angle UJ the direction cf'conuacrion. This e{'1imngtbu~lioiIiJl packs ii!'ll nH1~ fibers j)ersqlmre centimeter, nmlk.~,ng
~ dl1;i

with umgC'h,mging<lngl~ werethe slrongesi,

Az:izi dWflI looked ill whether real muscle s <lic~i::d thi:s \'\'"d.y.ilk: Imd expected lhal each pinnate musclewculd Imve just one gea:r ratie .. that is. lLltde~'gn a Chi"1!JC'I~[isLCic s;h~uped~Olng~. and thesefore be snrollg or contract faS! but acr ~~Wl;; b{ld~ ...uures, f:!.
Whe:U1I. ,~.eandRoberts evaluated

kiciking il:ltQ',I h!i,gh ge'I~,With ., htli:'OlVY lQ~d th,e angle ·o:fthc .flOOrs l"I;:1ilruliDilOdconstant, 3]].d dll.c muscle coarracted ol:l~YOil I;~ll~eand in:->teild became more oval, just as 1.11he SiIIHu~iltUom t •.
l!lll~bi

w~y. therauscle operated

f~l~ lower

gea r <'Irtld look fu U ~]dhf'i1llta£jee hhe

dense bit:ydc

I
~

muscle ~tr{)l'Ig(;r. bm

~'LC fine

es

puc'king of

pi~ii!I<'l!ll;; fib~r.;;, Ali;.:] ~010d. iI

~ dcm~mids~ A.<ilZ! IIUtl~Uy

con!.~"II::ti~e speed. ~. To bm lei ~ 'l'ompmcl:n1ilodc~ o:t linmlj,i' pilfHiI<nemll::;cl~~ ~~l,~\'e ·umI.der d~!:l!erelrnr
lI,i1W~,~J P'fl1~ to
tn;#:"

just as one cUn.:angcs gears on

re CIJ"~lJ'!.!,'1 up ~J11ever-steeper

:~~.mv drumstick muscle of mikcY:ll the shortened under differemit conditions, 't:I~~ rC'!iiUIts ~lwl"I~r~j;~d. izL ../\ ~ ing k:: Az
Imlse ~e lm,del,'goes not on.e sha.p,~

hilt ''"ll~(:dun~c-

~ gt',<'!Hlsro.r des.igl.1in.s video g<ilrnes. which

i ~ k~ th~1~NH~~~~~ ~LI~ 1::i:(I!Xl~Il~ Si(l!l,l~ i

!11~on. Hi~ VinL!~1 nHJI:;;cll: co~ldc(mtmct, .~ ,';~lorle'llillg .nl~nglh~ v{:'rtic1)1 dime~~~ i ~.J.on-lhul I~,. I~~·hel£~l-u.nd exp.md ~ altong OI:~.C'if."d hm:.~:S.iml'" 1.:0 11U'l.illla~n <I cO.n.= .~. stant vohml]C:!.for ead new m:uscle ~hape. .~ Az.izi obs.eIJ'\e'ed ~~lfther. <lnd to wh.u degree, .~ !~,eI1b~n;cl~a!~~i.; .. d~~4;li ang['~. r
.~
m::

.~ .sLllnil:)" less slrep<lllg_le. W.I'n.ea dun Imppel1!S, 5'i:lhe nm;!;dc'~ ovcn'LU height decreases more

i ,cl~:;.hflp~l"r!U~~C~]mruil~liD'l;pimll!l1;l fibers ti'J

,[I~.e :S.Daml~'::~lmns; s.hI:Nrl:ld Ii1M cer~m,1li nm.u!O-'

..

.~~bn it \vouid h,we had d~e fib~I."S n~:,)int<liiled

g,I~'a.dr Oin!!k.ln ol:i:t>cr \vord:;. l~'levi,ru'I:f11 El]usclC" ~ sJiaitl:ed ~lto du;: ·eql~I,iv;:)]ellli; f.t high gem,' lI:<1!io, o .~ hH~re'~~il~g ~p0ed of CO!Hntelio!1,+ O(~(;;lr the

chung~ b~lt~.ra.~,~~f,lf diJf~rm .. c: nt ~h.~pechmlg~~ ~'I'l.Jer ~lccurri~l.g .. to,I~.1 tic<1ny d~rl~mlillg {Ul th~ I.ood !i,pphed, d~JfereUlJt" circlH'IlISli;1,Il:Ce5:' he ::;,rid. Azil~propO:iieS I!han,.hhmvgh pinnme muscle fibers oft~n .rotate as th!::y contract, hem!!" Pan'lllcHng tbe '!,'~n1J.ml nmsde,. l~Jle~llrkey fibers lo'l.ds pull aga.inst the f~.bers Wtl l.nuch for lh~n 1.0do ~~at Thus, ;)illmugl.l pili1~l<Ile mw;~ IIti.emreh'es :S~,H:lJ.'re'ned ;,\bom the S31lill'e .amautl!l lU!lcd~r 11'1,1..: des ,H~ St1.pposedly !>p~c~fu~i,""l;;d f(H'(~-e• f()1ii" 1InJdler hglH dCUlm:<lIlii.l~.ey can <I~.SO v.~rk [f:;lst val"i 0 us' te:sl: condilitU!im$. 13m~~e~we!".d Th~ ~~Iudy I is··~~uttlt~ndi~~ ~rt itli!~P(ut'fiiml mld For' 'ls~es:siJlltg muscle' al1c:h~lecHI,re \II,".~l11 n,-el<t!Fan .aiJ'id $~riij;J5i •.MOOll)lf t ion to fll n:cli 011,'" ::m;:r.; AmJ n;w Biewe~ilell" ~] tile Il<lliJllslri1119 m lJ~de'5 bi.omech,mi::>~, 1)1 ~br\'alidUniven;:ilY. "It's have p,araHel liller:s sometbit'l.g l~l.<ll !ll!lll)f (ll"ii,lS ~lflV(; lho~~.ght ({Ii~!Jri!llIlI, l(!jl). Wth~,rc~IS :lbo\J t bUlncvelF ~~ lll,:ked or IIt(mg~,n thl'Q~lgi:1,." lhl)s~in the coil~fim~ -rm:~lllAEI!El!iHI"ENN~SI pi'nnate vigJrd. $IC111e: NC::~ VOt 3115 26 JAN U.A!RY2007

lion of'ehange inl the mllsc.~e~eOlJ',\l m'i~che~ [~(': mC~~!lnicii] dcma,~ld~ of eomraeelen," A z izi sa~d. M:oreoveli, the mus.de·s shiftilag o:r g,wn; j"~qtiirod !W !Wtr\l~HI~ t-;ystcm .i!lpu~.

W'W'W,sden cemaq,!J rq

457

IISIL.ANID'PRESS

LETTERS I BOOKS

~ POUCY FORUM

I IEDUCAT~ON FORUM I PE.RSPECTIVES

L,ETT'EIRS
doing dOOt,I,1, there i'u~, cOlmlput<ltuonal treJ,1(he:s. Stude~~t'l have a ham tile li:m.c;; ~iI!lQ18Ut~li!1g1l.;;~e I dic~u imo d~(;ir~;&JI,I'~slbr ~!>O!l~ .he WI!Ul!!ljl t'!i\t'f~, {1:falJl'Illolli'iuy (thC\;'dllUly jmLflJillsoC:~UIl)fing~llne v;U'ngl~1m!) and the i'lmher~ '? ,_11,8 ent be"uny Q;f~'liICn'lacrmnolocu~m· InOO,eiS lhm elncl'gc~ <1£ ifby m<lgic, from the user"'frielld~y c~y!'t!II~}O~I;r'lphic sofiw.~n: m':'l'umuha!ed {IVel" deca:des through the gellcroH~ labor anile field·~ I~~elm.lied l:oc,uproca~ nollE ~GREAT PEN1ARIEitFtACTlON~ CA SCIIENnSn; NIGHTMARE: SOH· w.1rt: problem leads to n.ve relnu::tic:ni1s:' space-cadets, 111 is C1I!'ie h~}ghli£!hts. 1:1ie d<mger.; of ,igIi.OT,~ ~ b iochemicaL results. o::om.'enntiOl:lfl~ but ~Q~icflily snlid, 3;,. was dJOIl~ . G, JVlincr, NC\vs ora.he Week, 2:2 Dec. ~~[T]h~se h:)V€~Y colored 2006, p, t856; "R,,;itl:<lctitu:'l," 'G. Chung in t~'!ti.i !!Jrigi~'IO;]I, p<lfJ'l:!"5 ~llld 01.;'> is !l~rwm~II~~J.'d tiwpnl i!~~)' 11:(i\;1.' h~ ribbons fsseoon iing cid :tleWS ~\el)()il"[ ubl is~.edalong \'!,;'it~,l du~1.'elr.lctiQi[!ls" SCilLll,ldilllers p er al' .• :2 2 Dec, 20O,6.p. ~;S'i')) iml.lll':!'."';'11«: g!,(,Ylfp fOirlnsrance. hOldS~l}\,!I'n in cm'l;;lid !;.wilie5 ~lUlJl]~l~d ]n .. (If one (If pH)lei~. c!'yt'~<li.logn~p;~y\ this cove rs 8111 01pa 9"9S ul~gI~morous 'INOrbdllY jourmds dun the 1~)t>ide~down~(JIpo~ogy gre.atest blunders. Vlidlnik an emMrr3S:Sof journals air's .iU·5t, asserted from the erreneeus $.~ruCtUt"II;S of the ml!~liiliug re~j~t-· ment to the innho~~. nevertheless provides the rest of the fgli;.':,ld \li'i[h some mlDd,ells, nat dartal ... " allce'p~llll1llpemrE c!j)u~dr:t(J~ po!')sib~y app iylo theprotein in. irs smsl ~measure of,t'.(l!n~.lbrl bCyQnl.dmi.O:IT: g \"'~r(: -MliJI~ler nnlCI"I~b:mnl(':~ul],n~l[e .. .., tl JJ. h.'s 1.001:md~~]~s~ [(,~U~IrS disIm.lus.sed as j lIS~,okl!.f'aiS:h iOJ.led bioclem ~stry.. CHR.IS M~llEJl1I !'ic~n1!d'enfi"ell(l.e.The ~,Hils~:ake l~~e;I'~.y se m~.sl"I~~c~ t'wl~ssflns ~~UII \V\~agi I~~ 1i'I'Il"",ardH:U!lh~s,Medicalllllsli!I!IJI!e. ID\~~rtment ~~ Bioch!ilm~~ry. Bramllels ~ml~rslty. baby boomer professors I':o"'tm down the UHmu so r O~I.D' :l'I'ro~eomic."i~)' Waitliia'rnl, MAc.24SQ, U)A. <!.rouse!!lJtg.ITi.dmrtegtl:llG.enl'O: [i Ithat dlO$e l{love.lyeolered ribbaas res.lom}·

B. 1'-h' '0'.'. 't···

P'reUy S,llruc:tllres" B:utWha:t Abo lilt

ing ihl:: QQH;.rr~,lilt! pages Ofj~lUfll,d~<lrejm,l ~liil!Jdel~.lliKlIid.u.l. am:lli:u) ~hatYflU i,~.vh;edL~astor ifytlll dO.u·t, kl1:O\\I'\~'I'Iill1your solb."iwre Ii; ocmf~Hy

:1,.

Reri~rnJl()~1 S. Nlimill) '[ 1E1li.az, , S:churdillel' ..,fElJ$lE.!t; S6i'l. 19~ 1~:2M4~. s

CrOllP Mod'e:ls IC02;r and CI~im,alle,Change

'f

lIN 'THEI~ RESEARCH ARTI(l~ "FOOD FOR tholll~~1:: ltJI.ve.U'·l]].iiLll-e..'I(pected QI\OP yield stimuI.aHon !;~Ii.thdsi~lg CO2 eonecutratioas' pO June 2006, p, HH S). S. r;Long et ~li. ~!1;l.\IG: lh11t crop models h,w~ ~u b~ ~ll'!p:i'tl"i.1~d

between i9~H and today. ]m,1.ail]~,Y because ~rud~ HH.;l1lm~{lSy deve~opment~ncludins: RS e fthe development flew '\,~<'I!r,ieties .im.,~ crop managenreut, Iand use d.lm~;g,e.and Idi~ and pr(f\il"i(;lm;)l!lo:t.gem~m (4. 5). Ch~~!,gtlS in Co,2: mate effects. which require a !'!'MH::b ~]Ilr~ coneerurarien aver [:linesame peried explain integrated appraaehrhan that oHen:d by less Ii'lOIn 4% ohbi:;; yiif;lk~ iaereaseassmning l{J!n~ (Hal.

or

to enableestiuuaion
tion nuder risinu

has <~ c~~n~id~l.~b~~ on p~di.c'~kms (If etTeel Their COIlC lusioo isoo5ed on <'L vel)' 5dt'iCli;rc £~oba~ food !ioupply (6). Long et (d. are also CJ'I:.tmpolill.timl,g rronn~ sample of crop mooch used in d:imate ch<1J]g;e stud ies, A ~,n:g:e:r gnnl,p ~ flJJroce5s~ the crep ],Jrolhu.;~:ivi~ypoer unit ;uca ofplo.:i'. ~.gnor.hl~ impomlllt reg lena I. and higher b,lsed erop models have already beenimleve] rel:niolllsh'i.p:s. Hn instance, emp mana rnlvl,;dandior S<lh~I'acWrny\'i.l~.l!ll dm~ agemeat, w:h,~chis I:<lirge~.ye~lermlilJll,e(]: I:l~ d from field e:..per.umcnl s, but they are iguored by LO:!lg (..'1 (Ii. ,( l-.n. :;ockl~l\eOml{lmic enndlrieas. isoften sub~,i:lIrthermore. the ~l1'ect:; 01:' r:ising CO2 un, Opti~l~,~l~ IL·educung potential JI ieldsand .• 501i!'l11;; {If the benefits {Iffi~ifig CO~, Esticrop 'i ields i]1. t}'nepast havebeen relatively matioas of regienal coop yi.eld's will a 1::;0 ~.L~,11lIl !~.ei('" fw::lm;J:I ~mclh(, tee IDi 111.0 I0SY develO mil depend on IX.':I.i'ldon~:~.ups b~t'\N~~1) CU:rI']I<lt~ fJpme:nl. have boer! more impOlftalmt ~11Idare I:ilk:cly to be ~o ilJll, hefi:L~L1II1'e j4~. fm i:lmM<lI1CC', and ~iflnd use- Ichange: 141. Cle<Qr~y,. future ~
C':ili[ll1Ue chanee,

e~, .<lind,

on~lt~n.: ftM,J<d produc-

fR)!\NI!: EWERJ•.•~~n~:frilNII,",IPOR1l',ER"ii 1108% increase per I'pm CO~ ~4). w,l~idl is MARK ,D.A. R!@UlNS;EVn!l.~ ~itn i:~arto th(,: d~t'lby Lcm,g d (:d. Thij~rc lalhr~ ~J"Isagl"li ficance oftb.e CO:! s ig.ud negates much liOl~rP"~~!lf!e!'!lt lIll!nt S~~e!i'l(j~~. of >Gr~p ~~rlt ~'O'd"'(:tlo<n ~1n,lm1gef fll.\; f.,:O~~il,,~!l!io~thAt t1ln~ ~'X.p~ri:!lI:I.~~!1·Sy.[i!~mg, 'Wageliing,f!mJ U!nlVi!fsily, P.O .. Be!!: 4310', 167!00 .Ak. Wilg ~,nilng~n,The ~lelilhell~Jilds,tFa(U~l~ o~ liffe~deJn()~s, tall technology lISIOO, to esrimme J~eld <eltmnges

Ull'lille'~~v ,or Coll!!ilh;lg~n, 10 2(3)(1 ri!01i~~rUIl, [)~~rn~,k. ~iD@iPiir~nn@'IIli~ (j@og r.a(ph.~. 1l.~lIIi\,@;rMt~, CalihornqlU!' de of 'LOOVOi;11\, fla~ p·l!s~~~r. " WIl!'!'.aiIflA~,·nillll''e. ~gi!!Jnn" ~

"Vo ~hom comr"Hgto!l!,d1encce :s,~~ukillJe ·!ldd'ffl~ed, E·;[11 .. 11:

;fr<lli~meLt@00IL~1

'~Sloo

1. S, ,~~ef1gl etai", fJ'eIdCft'ip~eo$. 8!l\ ,e~5 R i;20(l!4l'. ,2. IF. E.~ifl6 .c/., Agrilf. Ern~r, .6n~lOJ'!. 9~. ,2~9 {200'.2}. 3- IP,1).• l"l'I'!i~!iIl:1 ff ~L.~l\i' .. .f@~r:fnl'ir.:m'. 8~,H
{WO@.

ReJerences

iF. EwiliFI" M.. D..J!. ~oum5e'!'eI1. Ii. R·~jn'ller; M. J. Met~Bf. LN!ffi:I rn~,Agtil • .f(',rl.~~t filWttl~ 101', li[(Il (lmS~. $, I~DIXI ,Ag1i(~lhlrt Qr;amil<lli@n ~I ~~5t~ NlilooJII~ ancl tillll' ~.
IF(

y~G:lds afwitf1).l

in EJtlir'(lpt;; hfll,;'e ~le';;!liiy rripled

ffl0d

plrodl~IC'~ii.fl]'n dcrel~dso)ro s~v~1""4]ffICHns

(FAQ). ~ Sta.1ijilitaL D"'lab~es.. www·.liItI.or!t. ~,OOSL 6- IF, 1!,'-liJ;l~i~Uo ef .(li~ Eerll, . 1IgtQt'I_. im (prm.

W'W'W.scien cemaq.!J rq

$IC111e: NC::~ VOt 3115 26 JAN U.A!RY2007

459

lEITERS
IRlesp'o'llIls'e
OUR IRESIEARCH ARJICU Dill) NOT CIUflCIZIE
~1'~li~Llm('OJ P~anl Sd~mil!h HHI ZlJIrkln, 119'02'j!lurid"l, 5witze ~il<l d. S Cqim~~am.d ,e; lObil~ IDyIil~mi{s Oillisioo. m W,al,Moili:ll Center ror IIlmOs~'herh: R.eseardi, Bnlder. (0

as one ef the ~llP ] 0 economies ef theworld [Spain r~mk.oodg~lt in total GDP' in 2005. <lcoordi:I:1B! o !he Woxh:l Hank t The a buse S(1~r~n:;~d by Af11I;01l'nttlFerl"i~ ('·Spain~coJlsid~ ers lrs university referm lew," X, Bn~elt News ohheWeek. 10 Nov. 2006,p. 9~ I ), been achieved by ill1~,pm".edl harvest index ',"Quid btl itnIi,;QriC!~i\l.ilb:~en ~1sysle~nwher~ i O1!1Jd nHi"oge~l fe;;:ni1iz;ui.Oil. but J1;i:wms ,on merit is tilt:: pnnc ipa I ~:acto:rlhl' h[ru Ill,!; faculty. the se s ll,<'I.l.e:gi are d iIII in ism,:img. ~5, 6l. es Ald1Cll~gh in recenr time s, the wp uaiverA.:~l:lin~n.tgh we: ll~.o:pc l~'rlJUthe ilnpmv,clllcl:lts sities in Sp'lilll h,wc follo\,,'e{'j \,,'eU~e:M<d)~ of p..l:-'t, dec ad es ell n b e rLlI.~ nta :~IiIe:d.i~ l~j;b~~plroceduf'CS. in dh;l $el~-cdml offoruc1;.ilty. i wiUmuu hiiJppen whhom 1rmnov;ltivC' lil~'l,l\! thepl"Ocess u.s ril,dd]ed with iffiegll~arit1e3 ,It .,ppmac he::; .UiLd a :perce i\fed need for ,e:fiO'p ~1'I'lt;;!Ueil" tiliidtve['l1;ili(l~. \,d,~k:h iiiI'(; n19ii11.~g('1t1 by

[he d~s~~ of mndebM$.;;"dm p'ifCidic~ g,~obal d:l<li [I,gei mp~t c ts on I'm me food s l~IJP ~.y. Vile argl1llcd that ft:::'lb'Lble model projcctians IFeqIUrea ccurare modelparameeeri zat i 0 1'1. i D~wliJOI'tt'II~~ ful~,y UpC[HLir fldd 1!"C'~)[~n;cnlS with elevated 002 (1IFAC']1l'illlld,:rc.u:e;hatthe t ~t)~~Hnnn Iy ~t~ ~dpar,~1 i'!"llt;!h;!',il%a~i011 fur Ih~ CO::cfert iii ~]tio]il elfec t i~~Qv~rop~,~ mistic, \~ferecogntzedmat rising eo" ~s only one uf m.my fac'wr ... ~f[ec't~ng fm'IQre f~(ldl :->llpply. b[~t CO, has been shown to be pivoml , . ,-. I m pro J ec U!IiI g an I nerea se versu s a ,t ecrease ,~[1 liltme food :SUppIYl!LU~derglobal c])!fmg~ 'J~), E\~ienet at. suggest t:lillat rjl~sillg CO~ has ~,!(il_d ~!nd 'wi~~ h'IVC, llnle nnpact, <mrDb~On~ Qn~y4% of \\'liiu::'I.I.tyieJd im~fm'e]m,lenl over the IHIt;;! 30 )'~!US tori,dng COl. Tllit> 4% is cousisteet w~,th FACE, bUI :lIu]1 n,OI'l-:FACE results (extrap(ll::lt~ng from Fig. 2A}. WI/; 'Ig[-ee th.u the 50 ppminerease in CO::! Iml~,y have pl~~d ... :rolatively ,~lml~or role inl.hep'l~'! 31'1yr.::~ts, BUllt ls pnDj~(,;l-ed ro increase m :~ pp I1ilJJ ovc:rl'I!H! next 50 years <lnd thus, s:o ~las~Jig eP'O'[e~lti;:.ll{l be f~Jmore impo[1l'HlIt. Ewert Cf at note the rripting of wheal yieM siace ~,96n. due to technolcgy development. <lmit i,l~l plyth • th is wi 1m, C{lilt inue. u L~rg(' increases in cereal e.mp yi.dd~ haw

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lRerlerenoE!S
lJ. M, f\l~ C. l~nOl'l~g\ M. Il~\oeM!o~, ~i{~ TllIm, 1'1" ~,!@"jd_ :iii 8iDJ" ~i ~:6rlI. lUS (200~L 2. c, Rc~~ M" L IPllnJ!f, aMllf! 3:'1, 133: ~1:99';). N ~. H, Gi@y,. ~, !It(l'i',n, W. ~<rtt''''iflJ, ec J~!j,lIl!1,irn 'l~tlf~ Ch~tl.~~ ,2I:)flo1.: jlm,fMlIt A!dDpm!ron; ~i'tdItvIMMhJli!}" ]. l. ~a.l'lm~ 'fl a/" ,E.ds. ~!PCr:j(am'tiln!l;e Ullll1, Pl:tss. C,am~r!~, ,'OOllt Iil~ 2:~7-34" 4. M,.l., PI,lIIji,(~IlOl.l!~i~,A. ~l.e!iia~ M.I;h1~mllft. (i. Fi5l:~r, Gl'WQ,I €i:lIl'ir~ HIi'l'u;m Prilij~y
l,:a, )3 ',004)'_ 5, .M..p, Ei:~MMl. Pi. ~i!lleQril1t!d1l i~k StcJi,illumd\AiUt ~JII1i.!m~ ~'.r"

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ApplfJkn, :11.4,15,3" '12 OOSI. ], Sm~:!tl'I1Iall, It S~~~tt!~·Bradl~ it W;, ~5!:j)~. 1. ill FooL~e'~Crop Sci, 4S.1 i'S '(2005), .

1,'('10 blleim,s;

with U n~ver:siity' Hiring iinS,pail

SPAI~ HIAS COMIt: A ~ONG'W'A,,,{~NIHI [PAST 30

y....lrs" b!..!(~n f,~w~r~..blo :'H;:]~~~.~"I;;; l(1ct~;l.1olaad ogy. t:~.e{;(:ml.Hry :~ssu-rre:rit~g froml g,ro.ring pains on its \w:y to diJI.iming its proper p.13.~e

Un

ALnlQlUG~ DISCUSSION Of nu PROBLEM OF iIDlO\reedlirag :in S~]lliS~Wl[li\iel1l<ily hiri[bg: praetiees is V.1C~CQl'iI]C ('''Spain ~col1iSide']"s irs universi~ refarm law," X, Boseb, Ne'w:-; (l f the \Vcck" ~0 NO\t 200f~, :p. 91 I). it b. frustrating 10 see the del:mH~ ~om<trro\)dy centered .nound IIil.e ilre~ul'lrUOns. r-.e,I:m-edo d.et<1i~,s, t ofh:u ring lJiL1F cedures: The pru~1;;~H ,,}',"[~misfoeused on controllilllg t~,l!e pmceduD1es {~.lOl~I,llln.ke 01\.e:1I" ~l!l1~p~t1 ~y~~msJ ilfl~~~u.d the <)"~C(lml;.'i~ of [t is sadly true thllt ~f universities \!O(l;!I.'C to 1.~'iI.cir d!cvke;s, lOO ~~lal'ilyeelsions 'woulld {"!!wrm d be plai~lly ,nomlg. But legis~,<Iting to Ibrce (1,C<Id:cllli{':s~omake the ri!;lht eheiee is.a peorremedy for ;J deeper illness.Fhe real! q uesuons <II.'\! \\~lY nni versit ies <Ire nmk:ing poor hiirUr1gdeeisions, and hll\!J (0 reverse Ihi:.; tendency, The' ~ns.wer.s mUS;I.be based on the eurrem 11.'e.!I.I.ily I~.tl 'the se.mI.e:~j~iateresrs OfalUl'~Versity .~nd it~ drive lO\'vmxl. excellence f.ll"C <'Ill edds with ,J~~ i:~~~i:;I\lS!~ fi I.~ members, A SPO:!lilish eca~l diemi.coo''!''3.<tlays: benefits mach less rr"O"nmliiJ,ili~ i,m!.g \\;()dd-~:k.s;;,.. il!1d!;;pend~~:t !li~·w'I,::(!]E~;}.gl.!!.~ fl 1~<Infrom p!r(}b'l~.oti Iliga,~e~l'Nlm!LI il1i,ed :SllboiL'~ di nate ,f!~rem!y ~ll thJii ULl'l.~'er~i.ly. This .~f> rhe \ heert of the matter and wbere the deb-,ne :!>hm~ he fOClU!ll)d: nOlt 1]0\'" to ~bree aeadcmId ics to act :<Ig<]j"Jrnliil t~.l'I::.ir~:Jrnl,en~sts,bu~rn.,Q\'" 10 ii1Ugn these intc!l\:~t:S \\Iullin dlOSG: of tihdr uni-

~.en

versi t ies. A ell lnrre of excell eace si nks in very s,],ow],y. I.lUi1.d gl'Q'\!j,pi'~gnlUmbers of excelle~:~1ICiM.~el'!.~l~eS t hD"ivlil~g in: S p~~iilby are N\:"i.mlni:~gi!.g<1~.nstthe ti de. [tus time 10 turn it.lh;:mpnmry measures are necessarj, l~ey

slmouhl be C'.h::'<lfly idc:luU'ued

~IJli,'ersil:ies,

,Iii:>

such, d[~,tlf1~ filce the C'on-

~~islling durliDliYml ~hefiml,;tl~nodcL in 'which f

~(ln~m;:4;:" th~ir det:isions.. .~siil sector of :!lOCi,ely"

~,d;;\'r[~tiCJ:J!l" ~'f1I (H~rjudg.r:llcnl

.• ur~Cnl

~c'~ion

i!' [l ~~-d!.'dive;R t1!~£~nfgg rime:.... r~ql! irl;xl to g de:n::'lop :rm.e\v clJ~tiV'ars,adapted IOC~.l;Ulge' oUl!d .. ble to rel.llb:e ~!fl f~mH,r:-' fi,l;;;~d!' tbe 11~~.ghcl.· j'~niliz.,don CO,. dfcc! obs,el:ved in pwtHct:wd of;lllviromnel'l[s. Eqlilid~y. f~eldseal e 11m ni ~m~1JljOIlS () f (' O~. indudi Ilg. for ~~.;;t!mrl,~,tmpic~u~ln~a.~i~)!l~" i!li~!'llC'~i,o~H. \,\,.il~h ds~,II1l.gOZituu::, •• w,lll gene-Ik v<lri<lltli,oll.
.!J>e n~~d~d, VVi'rh~H.l!t: d~i~ nl'!J~lre :S(;;(:l~l~p,<ur!nletcrl z:at:i.on. proj e ct ions of 1'L.lt 1Jl1.'e!'g IQbal food 51Iecm~.ty m:rliY~i.lvc fed of elay. I r We

O)~y enh~ii tha.l p.lace a ~ow priori.ly I~MI uo, ,IlIRTACHO on n:::5carch.~m:::reaS!ed <ltu!onmny for 1i!I.IiI,iver·~ ~i,rie'.\iLq:1U cmne wUlh incre~ued respm~~~ s.~ bil- De:p<lrlm~t of Earth Sc'ience:~Ulmii"\u~1tyof (;lII'i,bri~ge,
~OC<l]go'Vern~lents (am'bridg~(a:2 3~Q, t!llt

or

and their

members

llY" NOI ~l~ri[,g lhe bcsa ~a~dudaEf;' ilvailablc ·~(ll,1Slinlle~ derel~e:ti(JIn, of duty by lmiversi,ly l'ilTidabt Th!,;.':y are .reSpt)~sJkil,,;'0 t'~t;: c:on1· 1 nmfi!i[)1 dMllm<li~t"jn:> th~ t~ni \'~r~i~y,.
The ne\v lInive:lI'si~.y:refonn
~."v.r

CORREa~iI)iNS, AN 1!lI(~A RIIIP ~'AnON.5

th~ Wrl,~~;; rob~ologili' ~ fights rOfi~ We· by A_ li!~Jer ,1[9 J~m,. p, 31S). The 'PiW,l~ (rredi~ '011 ~~320 we~ ~~ers.ed. T!he picture or i.ynJl1!FOOit~M:h~rl!l oour~e5Y 01 ~ ~erl,I!!;j\lii Ilorll,Lynl1ROI h,~hll~tNASA a,nd I.meCerm'~dl~ A!iJrooiSlGgi(.iI buildiing ~lIJJ Mildrid! i5 COUfW'Yj of I !hoi! {@lWr,
11I,~IU$O'f

repre~

f.\il :~n I;<]king lhese pnu::Jt'ic,11 measnres.lhe:f1 ~.ni.ght history ~ay. werc ~hc~ nlodding
w.lili~,e the '\'[)rk~"¥ ~mi~] SlUpp],y Imm':~n~'d'! STlEPI~IEIN~. LONG,l Ell~81ETH A.I/;IN5WQI'tBlI,~ AN DR"EW [ll. iiI. LEAKEy.,J I[l!ONAlm IR. OR1li;~: JOSEF: NOSliER6Elit '! IOmlD5!t:H IMIEL~
'~Di:!pa;lm~lIJJt~ai FiLarntl8:iologtV ~ na {;~p, 5(]en"~s, !LJlli· lj\~rS~I!I' ~llllirml~ a~. ~b<ililil"(hilrnp<i~'!lfl, U Urbaru. IL. 6'1801, USA. ~PIl1(jkl51'nU'le£~sRe:!<Elali\ch Un~l, WJ ,So De<p~ftJli'leifll of IIgr1c!liU~~~g ri~.!IItl~u~l R:e~e~ ~~fIii,~~. f1b~:ma, IL rdl ~ 6]l801" 'UiSA.]1:Mli~liIte~Ol'Genom~c Bio~Mf, Uni"'i!fSU:~ ~ minoi~ al Ur~na·(I/;Jam;paign. Url\lai'l<li..lL 6~8()1 US:AI

M:nt:.> <l su:p Ibrw'~fd ,i !'Illl,e prQIl;:if;l~S of ~U~ Ii,fi c<l~kl]"n of C 11 rm,didaH:s, hm ~herc iI.re no ~llIcelllti\'es~or l!Dlgver$il:ies [Q hire lh~ bes:r possible cand i.dates.lllbreooi Ilg and elh:ic ;11 vit'll~biom; ~1l\C]i.kdy H1 CO!ll~!l'nU~i)[ alll'!ilu 'the

be S!. u ni,vf'rs"itl ies in SpoOl i n

[Lnit i~

3.mOllo:lliny

~~d~~ponsibUi~)1

nlrl! 't:uf'!!curre!1dy. WANI J. iII.'AINFfUlilnll!

,~rf'~U!;bwl'91:!. Pinsbmgh, I

PIllQre$~Q~<!!"I!!Ch",lr, !1):~~r~i1"!el1fl;~ iIIIillhelll!@t1~$, ~m,f~f$i!y fiA 15200, USA.

lETTERS
:~~~~ fgC!!~i ~[l-~~tt preu, I~pthll!. piel ~~e.it!~t )'iIit" ,by .~. (olwzi A '(2.2 I)e( JOOI), p.18166). 'rilE (ol,ption o1rOOIll;j:I,a;ru)'·

~I'!jol'l o~tliie iIDilg(! TscDlihe lefrt,. i!nd~lj1;E! Iimldm~OIW 1'~ !h~right 0111

ing th~ imatge in ~e' 510111 mi5tNding. TI1i~'orH.iirnal '1'i\!I~

Need career insight?

Let th e experts IP you In th Ie pictu re.. ut VisIt VfWw.Sd'ence'Calr,eers,.org

nory natedliliat ollih~ th~e ijtOL!.ps 'th~l ha.v,e ~o~ikle<d h'"OO~h llih~ ml~~lng d~~ails 01 P,,~re~ma~'spOlper$, ~nl!l' (.)0 and Zhw claim to m\fe Jill~shed the rom;pMe proof or ~he Gooml?'lr~l<Ii:iolll c,onjBl;'t!J re .. In fad" Kleiner andl loU: al!1ll
iIIla~etli\;)t diJim.

Iprov.i!d~~.yD..I'l.!lildi::ellZil!

lhfeil!tlh r(illgh 'o~ ~he Year: ~lIhe'poin~,are' (onjedur'e!~

l22 Ik!{. 2:006, p. 1&48). 'TMe 1lI!1!1'i~

ICIMM eNT O.~," Evo~u1ii Oti'alry

Palf~s

IIJndlelrllyilFllmllFIOrwelr ICol'm' Valriat:~!CiIn

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Malrlk D. Iii! u!ihew

113, /!.ughl<1t ,2,006, P. 963) th~ I;1IU~f1~ of Mgh.r~fi'!i.~ridg;~sif1 ih~ A fl'l'ffhimJlm fll;irOlhd.Of· ~d<lf)tNe L1Ind'IS,.,pe, ~h~ir d~t;;!i a~ eQu~lly GCIfT!patjbl! With aclagJtj~ laF!clocapes havlf\~ ,a ~in~!t' Ipt'<h'l; !'1:d i'lQ rilll'1lEls. fhtiNt'l1lrnol1lrualio.n gf di\feiT'a 9 ~nlstiectiOlii aeross .Ii n~lirud.~~i1eargU:~5 o'Igi3i!lS1 U1~ IIHh.ouglhlWIi 1brey e,M/. (RI~~,
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IRieiS~P'ONiSIE lIlCCIMIMENT O:NI

,.EVlOllmrt ;~na IPaths 1llIIiIIIIlerllyiiIiIlg h ry IFI(!I'we'~C~hllrValliiatlion iln Aln,titlhill.lIni'

You II' eareerls

teo lmpnrtant

to leave to chan CEl. So to find

the right job or get career ,aJdlv~ce.turn te the experts .. A'~

$(;iE!f11(;eCan~ers.Qrg we know sclence. And weare commUted to hel!ping ta ke yom c.aJreelfforward. Our know~edlge is firrmly fOli.mcle~1 en '~he e,1li,perUse oW .science, 'tl1e plreMlli,er 5cilentillic

(~lnS'li~,pl1L@ AI1illill~i~,.AIiI,dll'@~¥ Hao rilla, I. Af!~rew B.aJ!'IJ!jll:ia1111.Chfi,S~(l!lIn~nl>&baUid.• E!i!lri~1;) CO!!;!!'!

Anl1litllbel. C. WtJmbtey. ~i~CiOld!S:

e,. L!iI[J9'~aide,

joumsll,. and the IOIr1,g xperienee of MAS e

ijfil

advan(hlJlg science
in

F,I:~!!I'~hl!r',~sertillll! th t di ~er'9JeIiU selection aerass ~IS a Ihybrid ,~iQn.e' i;, ~mQrnp'al1b~e<l'jilh OIdapHvt ridglt~, a ~liflot:!glll .. tu~tlveV~ rEMCln able, ls il'l[orrec.t Oil 5fV'ela!, ~mrlts .. fli~S aH~~m31Ii'lll!h~!IDth~5:i,5I[]1~ iI.snlngl~ m~~bl~ pea~. [I}redi:((S s!l~JP 1l'1l'o'immtullml:il!.lJan sltien "ness ih~ a h.yibrid rolW thlil~ h ~t Wfl,pt;lrt~c! by ai~l;g.fIS. Th~~, <l Mgh iitllie:s,~ po.1lh s:till ~rwid~ ~~allali611 0'1(pur data.
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EllrlO!Jlild the w,o:r~d.IPut'YollJi"sellf in lha piiictme with the experts. 5C lene e..Vi 5,i t www.scien e,eCare e IrS.O rg.
:M!SElil [JNSm!l, ... d 10101Ot:iI'rIItU. 01 Ttlo lI<IiI'''It'UIi<!ii!Si1!11~llI'ftj!.U:'iiI, ",011
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$IC III!NC::~

VOt 3115

26 JAN U.A!RY 2007

MUSIC

j aZ2" ~n~I~OS[respcc B, It was ~.perfoL;;,r;::tly ,~o:j"m~[,o!'!cen: a ~1~H!tk:1 led by the sax.oplla.oilm:i.s:t. R.udlresh Mahanli:l:appOi promotililg a n~w CD in the:;jm~~fhigIH;1!'1dj~ba blit I. ha I sea t s ah om ~,oopeop le, all of rh ern sq uinti ng in d ishel ief ..~ l~:n: I~ of d ~-ill~ P ce b. But there w... H~50 at unl<ll~zial£: secret :Wo:r as l~lfH;e imi,hekmi.ffi"" M ahn f~lh~llp{ll comp osed t ~he[JJiJ,llSi,c for his, llle:Sl albllml. Codebook.
~!s.i~l~!'IMmLJ~r ~h'l;oU)I---'~~J;: hrml!:.::l~M

!Hl~$k;;dbe~'ilavif!rs. "!h smmdl~ .!i.glumJo maner wh ;It ~ey~llle others <rife co m,p]ng i11." he
lol!d ~~!e,"] ui.ed .~~I.e:l!'nju~v,e sequences they dit!fll'~l:l<1.ve lh,nr:ro~~rT)C irmncdietely what lme.m~. (Lernnin; fiolPiMitrd 1:0 read the artists 'J.;)(.p1.~m{'l~.on'!;I:n:h.e pili.;.of

en

and

n~, I:'C..;.cI1lI.. fi.:I.:I.".g,r,id CVeml.i~:l,g New,. fork in. 1('.It)'. I\\'e'nl d~~"\fillW~"'1:I W hear some

t(HII~ibr,o·1lI~
Rij,r/((J$1i

~in cans

(I(rilE;:~d

~il p'1i~l}1 hnse

M<Jh,r.rfU""Pf/Oi

ami m.e.llled bubble gu nm.• you

li!'!liir/,etW

fair cnm~.gh"!~mve\'cr~ such subtleties d idn 'l~lmlil~edi .uely COInC a~I\1!;S IIlt'l'l!)!h?Sl {Iudh:;:nc(.'. "Th~ r~~th~m~lle1d~h~!tM'~iru ! d .ilrk: u h: to hear,' remarked

learn l~,Hil !1.~:oiii;lm~ =-ipl,!(~. time, eVQlut iOIili. ~.rtilk.iEL~. i'lilte~:I~gem,.'f;:,,~OC'.1

Pi A e C o.rdli 1'19S, N'9W 'io,rk,

2006. CD. $111.

Michwd Thaddeus
Un~.ver::;il)'1

{<1.g,l~!lg~-

br a iC geom eter <It C nl mlln b iU<I

mathe ..

]'0 IieSt ~~e ~RI ,~.ypothe::lis,. ~ II gaw Mk1~:afttlmpp~ (Iiilebest chance poos1i'.iJe to prove ~:t [UI.lg bY;ISSe]11,b~jmJg,m, ,~de.,.1audiW ~i1~,(;-ShUng'~~lund

with furrowed brow, "Welt il. is called Cm;leoooJiiaH.er (l11:~he q:~lip]'l{.l~~" for seme or Ami the pieces-vsuch .1 s "FIL\ont!JUlI'UltT ." .in which <I

Ihc 1;f~hlB \"riEi'! m~\'Ii:~!X:[\.vo

C fl]t:mne Ill~t[)dy has b~(lllil run ~b!:'tn.gh $(:\1.,

eral ~,diJier:s of e~u::ll'y'pl~JO:Im-no Imll.l'llI:l ear, lUI matter hmv lue;p21roc~ 'wo~I.IJ. 're able to de~t~l

ltiesrt.~)lcHi.mlg ifoUclI1 reviled, D!i Vinci COl'll') alid thetools ofcry))wgraphy. One could say lhat mixing :!'m!~lrla~d music I:;'nnlh~~,g [~~\~r<.A:;, th~ G~mHqn philo~vph(;lr (J<Otilfri,ed :le.ibll:itz put it. "Musk is the pleas-

musicians w.illhi 1'1:0form~1 Imthemmu.ical nainFi bonm:cn :SCqIJCflO(! (~he ia ~p irntuol1l fol!' CC'~~,- i~l!:ha :~mu:~'ncmm.ici,n:i.wu.lhno ~mls.uC'<I'~ <uining. u a cOluple of mataemat ieians wl~,o .LfC also urries of mil aUld mys.~~ci~m. including the maries thl: has born such frtuts as the
j!j.CCO~~1P

H ~Im.cd mu sieians.nndas a (.'out 1"0.1 gro~l) .• \ lawyer and a social wcrker, 1.p.<lssool ail'OUlnld a dl'n.cat S~]c::ct so lllllti everyone knew

~V:ha~t~)li~(,1!)or, lThe duy beihrt.' d'!4.'COI'!OI;':!'~.• J f

met with Mahmnhap:p<Q, andwrote
IIiii]tI.tic'l~

M'lh'MIl.lh!l~)I,J~.·~ in\'~~nton. F<lr~lm:rc I!PPfll'Cllil. li'llW1 !lny ofl'l1ic ~fll(liv~.dlmllli,llhelml<l;lica~ tropes \VOU 11111 nverall sense ef'lush ~omr.kx~l.y."ncrtJ:!1l~mer i:!1! particud
lar w.u:;. u~ingrhylh!'!1k classiea; musk in
iii

down the the

~ll;;~l.1;I(;lnbi fr~)m [f.!di~L!'! VIi/,aythat was. c~.e<L~ily

;:lr!,1iL<lleur."

are ihe humOlofflilld experiences from

buill to some extent 0111, 11i1I~tJ1~,~~u<ltic~1 Sl.ntct'LIIlC~. M<lhimnlil!l:lppa. takesthe r.e]gu[ hln ~:h.ip to ne:1'\J level: M,nll} i:ul'l j m;t the rnediu nm, his art, il is

CO~Lm-

(l:rd~r (lfQh~p.ieQ~!:! b~pl<'l)I'l>d.•'!nd tlltl mmhew

basis, of each.) The Iigl1its dhnmed

far beyond. t1~al of an

observed

ing w.u'~l\i.oul'being, i!lVI;,'i!I:re that it :is, Clmmt~ng:' B~n wl~ere<!s.dI music is

Mal~WI.~ ha rg~wa. ,I nu m ber theori st (and ~

band mounted the s:lage .• indt11e experiItWlll

rahl a performer)
Samir Chatterjee

at PdncewlJi Ulli ...ersity,

~ga!'l ..

(11iI.d£'edLhe dru mmer .. D'II'I Wei ss iS<I ~a t U· d ent <0 f l'~el'l:;~lfl'l,l\,'nc>dCo?Il.cuua t flhla pl{lycr
.j'

n~.c

M. aJ :1m <U~~. l h a ppa launched U!l: into the

deep end right .n\'<uy. beth musica Uy and

~1l:S0 his

..,~!bjiect. Take foil' example .his ~mrn.efUlr~;n:T and In 13'e"

sax.

or

Bu.! .f~;;:Ul~ ~hrnv \'l.Ii;~!:)!t. f.l1'l!oo.l1l~'1!lMio)1 f'IiHilrom; did begin to fin:'. Thaddeus deduced lhlll the imudodies ill "Further and In Bel\veen" gel

mathe mat i e aIl y.

:Befo~ l~.c rest
()f

the

in ~'tr1:Lmems joined:. he
broke the silence with
a S(tqUl:~tO~ nhiflUlo;S so f;l!>l and jirtery it was

tween . '·l1in .. meledies e

theifj~llllJ;Jy qJUl!lity fml[illl~';::hn:kofooIE01ltii;.ie dl,gils in the cyeU.C'2I1 nl.1JI.Vlmhcrn.lle Ml.d Uhargav<Ii were also wise to the math ,,,,it~ in alJ.O'~her~un.e. "i3uh,mced Perfoflllillloe:' which is bui~~on

b1Ul~d between

o~e:;,t~d

bass, aed piano are per-

set or a (:iCc:ienuing polyrhythms. f1\![iclll<lllilhap;lfkl dedieered dl.e pieeero steroid use CUI'i!-

mutm'ltl!1s"fa

~J;;ill,1o; bl~nl

like
Oil

fl

spider d.rlllcill~g: amf!~.g OlympJc aI.hI!;lle~.anclkllJ,owing (ills

on Ihe se:~ui.tonc intervals ( IA.2:.S,5.7).lt It~llpens lh,Ul 142.857 ,~s one of I;l~e ·c~ U~d c;:,1c.!tcal ~ nJumbers. ,\'hi.c:ll, have' 1.(l!lg f~~~ illi;i1:~~~1 !l~!mbl;r

~h~uri:m.,iHild
gists al.ikc
nmel~1;l
~'\,'{l).

lUlm~I'O~{l-

to (lOIili~ eye! jc'.I:I:Ill!lnbr::rs wi!l~ n d ig~15 have lite illte~.'e5Otln.g pmpc:l1y~lllfll.mllldp~yi~lg aile of~liDemby ,I ,~'hole nmnbe~'lJetween .~iIfId II gem:I.<Ii~:eS;) ne'i,';,'number :lrn,lVing ~~l.e 's.nne
(not

digi~ re~u:'ff!mtgOO ~. (~,~-, 4285 7 x 3 = 42&5 7l J. ~lS<iJi ('o!'lslrairu ITor i!'n~Jrovisalioll:

M~1h1]nlh<l~~IP~. l.hri:S'IIr.i~]~lm,le'tic<li1 curiosity 'U1'!!eS play tHilyIlmill.1g IQllig <II:!! i~ m; genemtes<Ii cyclical nmnber. But \""I'k"lll does the audie:l'nce ~flt out of Sill~'h IlilI~t~~em'<l:l.ic<1~ mi~~,ieill'! l'h~pe,..simi~t :ilm, me: $OlL}I'S th,~1[h~y g.et 1101il:ilfl,g 'lIt all. thil! it's j l.I:5t a. gilfl.nnLc~. Letls \''3,11'h<u~he II,lul:~ hy~X;Il~,ue5:15.,if' t
you

'w 11;6~fl, tfl' ~llod~!1!l[lilt ~nlil!\\l\~l'>.

~QU'n k.I:t!l.1I1'

bridgt.:. f~ ~~Q(':!1lg ~thu~I~t;':of VMe- l3li\1 &htlO.i,. d~tec~.ecllh<Jtj'ayerecl 'I'\,rilil,illl ernJ. of the se"... ~)!ltf,)th!;:~2";1~]~~e Wl~ISihUf~:';-~il ~"pl;ldri~[y "'I<!YI~J-\~~in S~m,"i~d~ ,~,'ll:-;c.~le. F:wm tlHli i.~<ill!::d~u :SmT!!U1~MOrM:-\'!il;;rel:~1;l pt..,.tQnn~1i· Ila~ne.s i.llmMo:r.;e code. ~ike :s.i.gnanLre's on initial simek. Ihe baml «nit'd the piece a!o['tvn wh,t!I~]llike ~]~rie!'i 5IOl!!nd paimll~, Soon this, h,FIIS,is,~.cBln alirCady mlc mil the of bOt1~~cr :lIltlC~C'~5 bC'~~orc :rea.c-:I1 tihc scem~ iIlg null hypOlhr::~js.Jhe nmlh is def]nileiy ntu.re il'lg'iy JTee..]1o.uing ,~bur ... :uLally 'Iuite !l,-um:t than jl1,St <'U g [1:]1mick. e'!;re'lJl the LUntl'J ianld e'<1I; W nJI.''t1d) s[p'<lce of limpl.iQvli Slilri:omll j<hzz. 'om Usi!lilg ~h~F~,bo~~(;di;lie:nce s~Bm$~lk~ BMI \\~l~llh~!l doot> dlCf.!~1di(;!1;(."~' g~~. or lh(l ~ gOl .m iub:illl"dVY C'.h.oice~ pedlOlpll> 'N!~.'gi.lii.g 011 iiII g,i~.1.1- eX]X'I'.~em:e"! This is: where~T~.e ,e1>l.peri,l.ne'.Im.f a lril~ k1k. \'Vol~id1'l"t a r!mdnm seo~lu~~l()~ do just 3$ im:eoo~li !'Is.. nle 'bm1d phiyed Q,';.rg ~llS., :I'lA Ihtl bre<l.ther bel\\'Ieen ll1i.elm~ like i'U ,r;o:ITee break .. "Iveni'! ACCQJ.dillg to. Mai.lanlhiFII(lll'l. tile' choice rises .;lbo'llC'gif1l~niC'kry lor hll'oI'C!I:5om:,. Fo~: bct.~~\'''CIllectures ~~l~ good !!:clC'!liinc ~"'I1l.ferOJ~~. !: says.. l<llkililg l~.e ~GI.lIIem;e ;1;"his i~;lSpih e:lml,,~.brimmedwi:~IIi.heated c[o.ss-di'scip~iIDI:ary nM~Of'll ;)Ind,,(mslr~i~t ,g~n~nu.cs VlHIl>1c h~ CQl"lld com'c~lil'>~tinn" Topks ranged fr(lm: hm:li' 'cunlnot Imve j)I'Qdlnced othcnvi1>c, FbI: <InolhelF. he str;li::nts. .'Iet <IS dY:Im.3'IIiI.lO~ fO:1Fcre"l!uvity {for ~:.'i!.'l!lilJr~c. th~ ~()flln(:t fOrfn ~n poony) <l!ld the: d~lms. dw ~n;)lpp(:d s.'(':{.llUeHC~ l];;IS uniqu~
l:<'ISi~

hi:s sax, The manic Indodics of the HaliDe. "The De,dder.'" ar,e a i1l~lppi~gonhe:: Fibo1l3.Ccl 5eq~l,el'lce t])

tributl.'!(Ji<lmIi exua sqLlir~ ofadrerm~igle'.) Nor di:d :rou.&rn.\r~to be iI. mml'ncmmJci9.ll (or ~ m~lI5l~daliDl' ~o Ob~f'I;fe m,my of the tricb. C:m)()li~.leTrtw'I,'-

b.

~
~ ~

.,

26 JANiU~RY2007

VOL315

SCIENCE w'INW_sclencem!,g.org

B·OOK$
biolo;gicil cnmai"!)h~,Tii.lh,.~PffU:!~I~~Fibo!lf~Cci ruunbers DIllrI(lture~D~1i1.e iml.d.epe.Uil.d.enl discnvcry (Fi!t[uI::'a rljet than in EllILmpe) and u SIC 0 f Fibormcdml~Hlben; in <linc.i~,r~t .Indu,u:l poetr),. Wjth om brains fi7.:Zing from s.m::~~. cross-tal k, tl1e second set was <l1~ more enj oyt~Me. the AnI.! thilil :>~upp!lrl:fl. y ~ Ik~i'n~livc hypotltL}m sts f()rwh<i~ an I!Htd~t;;HC~ c~~mt .gil;.':[rllil (i~~ <l. M ahanthappa perfennaece: a lin iq ue, j O!lZZ"

srAt.

I.S.

...... [Imlo:rri:'ll convicricn,

:.

uc!il i'I·~irrllli,g .\ ..~1;rd~~spekenwlrh $~Icb •..

IInlSI:

reviews

!1!ld

olr~rsrebunols

H}!.'IU l~(:

stan-

surely

CO:mll·e

I. .... [ronlI. all outraged liberalexasperated with the conservarive !l:lim.ut~ of America
wd~y. and one calm be .lorgiv(1!.UiI. :for lhill~ib'lg t hat ill a rev iew of T1~eGod O(fI!lsioJI these ~.il'CDle: \Vt1iutl1 uf Riclmrd Dawkim:.~ims~lf.
WII;jU k!10Wml ~~)rOt sijff~rh1.g n !'~Iig~mn:s;foo~,s g~l<ldly. Bul no. They were enten:"li;I~n~~lh~ 'Cllngr~~'stono! i?{!(.·(mi on .~ September 6: ~. 81 . by IWm.I;!: other th~ln S~n,l~gr 9 Barry Goldwater, the fountainhead ofth.e modern cons.cr\'alivc mevemeut. the ffia.mI. whose r.,H~dI964 run for rhepresldeucywas said to have been rtl.~.f:u lied in :~ [I by Ro[m.al d 'OS Reflg.u'l. am! the emIt! i,d,~~~wh!D~ ca~npai!!l~ Klog!l:[! wat; ''':~n'YO~LI~ Heal:D You K~'iOI\""He·s. Right:' .~fGoldw~l:e]" ~UH! been p.r.esidC;:lIi!t~or the ~).rlst six y~''lH'. 1 ~hHLbti:l~at' Dfl\\;[k~ns \\Could :havepenned such a lm'l'>'CI.iul. p olemic <lg~insr t h ei n fusion of'rel ig iom. ~~HO nearly €ivery neek and crn:lm:~y ofpHblicl~r:e .. :But herewe are .• md likeGoldw.l~el· ..Da\\lkin,S is sick and

t"'hoi~

fueled Q>I;.'cuii,i(~ftI h~e1'i;pl(j;N :i.de<ls. Tw", .. bJ:idg,e 1

ea lied .it "the secret knmvledlge ef1'e:C'f'-l:lm~u heighteni·m.g (lf~ljle senses fr(l\u:J.~. ~\\'IUH::ln;'~5, ,nil othi.dden ~Hye:rs ofmeauinY-ilm.d,~: ayree.ln addit ion. lille qm'l.nefs virtuosi D)' makes thei r phl.yi[l.lg a pleasureto be.lill.old no matter llOW OiH: fee 1~~ilb(1lM.lj~U. Bu E b~ for~ tflk h1Jgmy wortl for it. !I recommend repbc"Il~:IlI.g the experiJncm for YoUl.'.',e'lif-\\'.Ulh t11cCD .1")'0'[111.
(',l'~'~ C<l~d~. J O

dard m'gllments. for (ilO!:rs existence.He conccatratcs OEli dissectiag the 3Jlthmpi.c prirl;d~ Ille aad d~sm;Hnling .iln~lIigell'! design c reat ioniS'[;ll, r(A s pan ol·~:~].c ~alD..;rc Ifort&,&:tc redireers dle! erearioaists' argument flio:lim eomp lex it "/ to ~~nw [11m Gad must ~lave ·b!.ic:,n dl;';~igfled by ~1~i:~pef~ hgeiiU dl;':s,igmr!.;lr.) f.j~d ~II;': then bll~.ldl)i 3. case fur "why t hl;li'li; .<!Imn~t c\'rtn~nlly i:-: Il~) G o~J:" The remainder of the

beok eutl iIH~~p'Oss~.ble '\,'o~

ImiomrL"Y erigins of mun:~~.ily ~md religious bcli~r. <'I jm:rificilti!olJl for beull~ .~mrd on rellglon, childhood rc]~·· gious indoctrsnarien as
chi~d abuse, iI:DiId deg3.1IU an Cmlflllflenl<1iry alll Ihe ~):H)·· gress ivcil.}' d~m.1gi.n~ mor al ze itge lst. [)"~)j,'kns closes \\r~ l ribute 1:0 llir.e power j ~h,"

:~JoL-'il·I\ormmlre.

Notes
1. Tile ·ql!!aJUJelt(llllpris~ [;l~dlfe~ i'IIJahen1Jl1a!lp<!. liiji!Jy lyer, ~Fajj~pi~ rM~lnrl, an~ iO.a.f:l Wei'l!~_ 2.. Th~ sum or t.iI db parr d ,oon~eco:li\ie~lJrnll:u!f!:.'ill UI@ HlIona.cci $equel'l~e l@. 1, ~, 2, ;j, 5, S, l~. 21 .•• ,11 g~. tralH Ihi!' i'if!~t]h~,.ltlo~ 101 mM4!fIJ11\l~fil:!oi!la~l m ~rn1tben U.I', Z/3. 5ia, S/13J •.• ,]1 ~OIi1~rg~cnltie irr.a~ijo~3l rtgmbeor <~ro.6l00.;l3.~&', -J, ~ml:l~n ~the 911lclet1

and beauty of science, whk:h no I b,.'.illg \vf! I.!;.I: dces t~u~P'. Whell I received the bound .galleys fllr The wd DcI!I,'il'flU. J cringed ar t['[e [ide. wbbingk were more :~.eUlr<l~, w.~y not. say, The God ( QUC'!'ilimf?). At. I read dle book.l fc.m!'!dli'iI.'ty.<;e~f
Ij.~rinclin.g;<lt Q.,n'l,'kins·s ilTe[ere:rmoes to [re]n~i()m;

meal!.

l,

ThiJI'~~ag'~lI'i<lklor ·~WlIjl~ilJ!.yi·I'!g..~

tired ofbe·jlr:lg laid Ulm .u:heisuN .ne unmeral. seeead-elass, b~<C"-Of-tijH!-bus dli~~llIS" Ti~,: God Delusion is liljsw~dy of, like the lliow,m.·d BQ'a~1,;l ch~v~gtehH 11m.JilB 1976 fU:m, N(:mwl'k. t sli.ckiliig ~.li~ heud out tile \~Iilrn.dm'" ami shouting. "'1' III as. mad as hell. andl ·m. not go inglo tllke tbi~ "!liym(!~~." !BtU .TIi.f! Go(f Ddw;im.r is SJ() much more thana po I!i:.m ic .. h is ,Uill exercise to "raise

11tc:tt ii.~n:o POi>i·~iGrw.1II whidl pc'Opl£arc w

hn!llffii"dblc ils~h€'ir .uclillim.!); bel k:f.o;;. 1"h~re

i~ no· maJ:~ 1Jc!~lj,1:rlhl :lI1I.Y c:.m dl.aim ill :J on.c dchat~ ltum J1.~_'i!!!~ Chri$;n,m'Gad or AU<3h. or W.~2LIC\·~rr .:illl!· Cil~ I~ III i~ SUplll!'l:tliC' bdii'l:~.. The l1e'li~i'O U)S [<leI i{m~ ~h 1 :u:c~Hn\':i ng .. iIHoUl,!.lhm~[o:t.l1' 11IIDldlll:'l: mot hi~l.!lth!.:iii' ~,.:Ji'!lio·~ls do~~ whh wisdQ!n, W~1~yar~ Wl~i~~g () forciC gQ\'crnmclll h:ader:i!. into ~ [{IIh.:Jwing~~tl>ifpoN~iolii. :mo percent. .1fyrm.

di sogrce with
jpllnuoLubl[

I~C:SC !)cHgto liS goolll'S

Oil

consciousness liCl d1:e t~u;l~'l~<!It bean arhe~o i~[ is: a r~,;t! iSli:: .[lspir:;;UimL ;;Jl'ldfl In,fwt;: ;:IiJ1d sple:~d:ud one. You Ctl.ll be .om ;uhe~:Sl who is hi! ppy, Ib,d'-!I1c~~Lmor... .-!~~di t;Ull~l~! a~.~ l m y ftll fi.1Ied ." D'.J,!;"ki~H;WHI1 ts. a l~m.e.i lS 10 qlJ~.t S a~~,oh')g~2 i.n:g.for t h~ iIr n;Hg ic),~IJ,sk.c·WHieh;m • "O~, ~~.eC(J([iltr.l'fy., ili~.iOmen1.1 ~Jl,g~o bepro~l.d S of, l!:'ti.!.rid~ngrail tfi I~c~' the f~~r h[i!!'izo~. fO[i" athei.s:m ne.u~.Y'lhv1lYs indic<lteS1J he.,.hi:rij' i!ldl;':pl,,~!lIJI,;!iW ~ (}I'm i n d 'I!Hl :~ d ~~ d ,.1 n h~uhh)' mind," IID<Lwki 1Il!5a]so\.~.an;us to IIi\ isc Gonse.io1J~[n~!)S abmn the prn\re.r of' D~]rw.~n"!'j.d<lrmge~'I) lJIS
idc~'l of mlUlnll :se.iecliorIJ .. Hcocl~:cves. that

bci I~g lessinte I]lge:lmt,.poor <llli'e.,'s(mi~~. 01" even delmled. <lind 10 re]igiolls ~md~!l~I~S m, .. ;m~bit"r..::. of l~~nl;tis.n~.I .slu~c1(!ei·because [ llla;l,Ie l'e~igiou!> rr~.ellds 3~d colleagaeswho cI.onot flt these de,~cuU,p~ tars, mrnd 1 ~npmh~ll:'I~ the plJilJil !:.'llch por:jgr;!.II\-C ap;pclla:ai.oIl,5 call.lse~ll~Il1i .. hli.addit~onl •.1illll~. norconv i[L1 ced by Dawkias 's .<1 rgllme:Ln~·h;.t wi~~oul [l:idigionlhe're would be "no ~lli.c:tde hombers.jin 911 L IlUJ 717 .mifl Crusades, ,'W wi~:cTh~hlillll.lS,~1.oGunpOVI.'derPlol, .no l~.du;m partition. nnlsr~i(JlitPa~es~~[1Ig.ulwflr,.;;, I'm Se.rbfCI'IO<LVMusl ~m 1ll:;LS:S:!Icre"s. no persccu, li(H~ of JiCwsas 'O~risl~km~rs: no Norther.n .1 relorllild 'troubles ..... :' In my opun~JOI~.m,nty ,or [~c·s.(: CVlltfit~.--;;;lnd u.th~r~ ufl~~1 aluilw[cd poople
af;.

"faith -heads,"

J)lS

~old.y 10 rel,igiaim. by al:tIJe~sts-wereless [re~~.ghltU!>.ly mo~ivlH~ th<i~ po~itk;<I.~1.y d:rivC;::!i!.or ,f!~ lh~ Vier}' le"s~ iuvoh'~d rdig i(~ng t:I dle M:('\.'iclf:

IlilDrul. i,\;S~lC. 111I.t~y CfJl1.1plain.u·liI~y

DhrealCll

}'I!!}!Jl

'1I'ilh a 101>$. ohmuc:y

01 votes C:(lurH~'~i
:I

nlm:q)eopl~~even

or both. rn~ frunldys~'Ckuti tiiretl oft~:Ic

IPOI~lk'~II're<ichc~. ",cross tilis td Ilill'l.g n.'K:..•• n\~.~I w:.U'lt to ~c iJ'

morn].

~crl10n. ~. usl helic\'i,lj.n A, B. C .• u~d D. m Jm,U who do lI~t,"Y Ihili!k ~hey3rt:?

T~,e ll'I!iew« 1:5!ilt Skeptic !fr.!a9,nh~.2761 NOfdi! Ma:ren!J~

A.';I0nll!l1. A11.<1d~.I'i:1l' 9100"ll, CA

mmly sc~emrtusHs~do not fuUy undel'stand jU!il hm,,",,po~\~erfnl <Ill ~de<l ~.t is.. H~ ",nrlbu~I.$, ~hm ffdh!lre to ~h(,)' 1J1;~d o I,)~ ! ~ sl.eeped ".I.ui imme:rsed ill n<l~lII.r.d s.el·ecl.uon b~fll~ ylm l"f!iIlU'll!ly n.>co!:::nb:t'! hs p~)\wr.lr'I thi~ ,context. 11<'1I.ur.delocl.ion "s,il!au:ers 'th,e s iHlUS,UOr:iJ f' dCKigllwiihilnlll~ o dOllmill1 ofbio100~y.~1iI.d Iie<lches us to'be su~piC~!Ol~:iid·'IIlY kind of dcsi~fIJ hypothc:d:s in phy:;.i.cs ~nd
!;osmol.o,£-y .1S.\l\''(;:U:'

of pol ideal he!;,'l.'!mmyo I. <lIOOIW\ier ilJil.lIl!gimtt'd 01. book ,,,.j'th thi5 tide !,V,oulld ever laJnd m~ bCSlscUer :llhm. u[[m I.he Uniu::d Sw~es.Bu~1 \~I{l'" wmllilg. T~.e dmttl h .. ve
spokel1l.

rbe Gotl Ddusiotl :i.s a fLnll<LW<liyres.t~ S1,;:]ll;\:v.a !'!1!iTurb,:;tb;l$[ ~nmny Hi the hturl_g1,Or ~!'Iy
peop~.e-fa'r [[rnU:li.rr(;:·,I nm:'!.' hink. ti:lIOl.npo.us t
(Olf same![)~'t;h".a r{)~i.tiol~

rew~~I-h~vl;;

·of

Ine..8:~ige and pcw.'ea· to spe<'Ik for tllem, un s.~1.ch

~1Il Ii: Inqu,e Ilt Vfl'lCC •

[)Rwld

filS'S

I a.lc~l book

d§ltl:t.,mm

USlt E.-l1i1a/it: illil~her!!M~t@

de :H:l:lFYIi::imU Ilip Ie re'J.dill1l.g~. not jU:Sl <t~1) n itnPQJ:'I~m work of sci~l1oc. b~t ·as a gr'C'ill \'\,'Ork of IittlrraulI'e.

O~l~of obl.ig:~tDOIl•of cours~. DII!\\lkins . SCIII!!:.NCI!! VOL 315

26 JANUAIRY 2007

~,:soienc;ema9.ar9

A'lg'D,r'ithlm,-,Gu!ide,d lndlelduellzed Re,adiin'glnstruetkm

Ca!ro~1 i\lh:li!I(lrI~lcl C'I!IU!I!'UlI'r.~11I!Fredenk:k J Mm;,ri's(I!l,:l B:aulry.!l. I C~riS1lophell' :Sc:ha.~s,c'hlille:ide~.1IPh\lllliiS ILhlldelMool.iI'
s.

first grrai:lll!s;tud!lnls aC.hiieved stron'ger le.i3lrning ~!Ju(lm@s i!il ~df!,!g ~'J,ih~fIt~a(h!~rs i!1!INlJfl~d lastruetlen t{lir'l(l~vidual ''>'llJd~Emh under the gl!iJd~!l~~ o[ A21 S!!l~lW~re.

F~sIHntali!,li

e<l,J~ng mul \"'lI:ili,~g "lremmJrl!glhe: 1l.~.O.Sl· :~mponaJ,1t skiU::; i1:1 too~w iUlfor~ 's
I"IlillrUon.drivcn war:kt Y~I. according ;I~ Assessment [)f Educat ~ona.ll PuxJg~ss l /).mo~ thalli (lnC'-lhirupfd~Udr~n i:l], dlie Un it'f<il, St .. eslaek fu mi.d<Lmellt~,1 u re:<lrl:iing :s:k~~is,,, T,I~.is,rate is cvcl'IMghJcr for c~ikl.ren :

~,()~he Nmi.on

1,~'(~r1g ilugpCJ'i,·er~. IniM!!~ICY is ~. s{!cie~y-\"i. and {;'o~dy problem and is assceieted with increased r,nle~ of £,,~dere'telillioJ1. re:fi:mll to hugh Sd100] dropout, ~nd ju venil e d~1i~l(I~Ie-ru;'::)f ,~1). a:~y c'hi ldren fa i'~ M
.5J.'H)(:iBl~. educarioa,

d!i:~:~gn.l;,'id~"Q lilleip 5~:lU.d~Ill;'; <A.chieve prufi~i~lraJ phcU]ologic:al decoth,£ and \,\lord re'adi~lS skills t/,'n Meal'llil1,~-focli!lScd instruction is; des.ignedwte<~d.l cbildren ]10'li~'to exuaer ami. ~(jC~:"lru('1!ll~!l!'!iJmg fmutra text ( ] 3). The di mensian teacher- versus child"'lm.l<ln~.ged. insllm.c~ lion id!elmtilfic.<> 'who is rcspenstblc for Ibc'ln:!lil1g, the studems' ,1lUl'l11.iO:IiiI. on t he le.~rning .~c'ti\!·i·, lies ~t hi?lnd-1llc teac!m,c:r'or lite !!:tudclU (4.
16). Teacher-maneged
:iJllSlnIClio:1I1 :1111(1)' be

pf()fi~iennt levels in ~ei:1ding, onlly because ~he:y do n(l~ receivethe <1;!OOUU.H ~md Iype or[m"uuctkm they need (J:-J). Much, ohhe '(.X>:IiI:~ro'i.iefS;)" ~g<'lrding U~e[JoeM r \1'o1,J}1 to Leac1,1 >(;~.1 ildr~1li how ~o ~::I(1j has recused 0111 \,,1u~thcr ins~ruetion should be eede-based, such as phonies, or ~ed: O:l.l \~'~lOle 1<lI.gu<l~~ gJi!ul !TI;i;;~al~iJi!.~ (6-<~l'.bm [his d~hlM,,; InHymis.t': ~heP'OinL Ahh,ough moot children develop stn.·lNlger rea,ding skillswhen they [receive <I
te reaeh

higMyilltemCli~·,~. such as when the tc~c:~lcrs I le~din,g IJ~dL"Cu:;...,;jOf!. Cbild-.n\l,m~g~d i:ii!st!l.il,ctiou :um:ludes activities ~n which srudems ~re e.",pec~.l;)d work ind~pendJemly (see P!!~(H{)· eo
graph, :lc!1i;) or whh peers ..Change o'ver time 111t:e,ad~1'I9 ~1:I1S:tl'iliU:UG.III. Le~s~liI~ all"e! M U~ClrPri to ilild i~ 'II11diU;l1 :studenls.. ries, \Ve also fOIlll,nd...111 interactica between 1 iIlstnrC:l i(l:n _ ell iId ren wH~ abO'l.re"'<l.v-erdge \'oc:abu~<lry scores arthe S>t31.1 of the school year mude greaser gal.llls i.mi,\II,!Old ' we~dirig when they 5l)e'l~t:more t ime ~hrQlqg~om the year in chiM';IlfIi<l.raaged Incmril1g-~Tbc:uscd in~,um:~:itHII l5m;h (!IS independent r~ading)"
\l(l~[lb ul!;lJrY<l!:ld U'!f1ipaC[ (If c~o:I"ggl'lg the fOif.1I,lsor insnucnon time rill·>Otlgh Ibe sehool ye;J~rV6). These dimensiuns ng)c;:r~h~' .imultfln~OU!i:ly s Slo~lrn'IJ:<Iny ~angmlge am activity f~.II:s in one of fOll!!":S:1;'e~()r..; l tahle .s 1)- For ~~~)mp 1('.,~l1ildren read ung lo,g:elh.er in the Iibl::ilry are p,mioip<lti.ng in ~~dlHdl,nmua.ged .m.elln:~ng-.rocuse:d

capmres the

balance of ex pl i e i~ d'ecoo~n~ i msuuc lion

:i n

combinatiol1i with Illcallli,~lgflill rcad.ing aenvlties {7. 9-ln e'veu ,J, balanced approod~. theory assumes dun one .lpplimtch. ifil is~he right one •. wi II be €>llr~I,f~~ effective for ,}II elmi!dren I}l ,t H. 14). [nsl:emilhe en:inlcy ol'i:1IlrYP;<lfliou],lf i!!!:;;ll:lQCliuVl!.:i~ pn"ett~tl nl<.M.¥ dtlJ:,u;mllun the ~~iI] level of the sH~.del1l.t.lnsrrucrioaal stmtegies 11li2U hel~" ane stmkl1J! 1'Iil~1L)I be ~n:cff~)(':[ivcwhen appl:ied lO anether s~u:lent with diffe'l"em
'11"1'] j. S.Kh ,~. ~

;lC~i,,'i~y..The teacher imtml~clillilg thechildren ira, haw 10 segment or blend phonemes i~~

1:1]. ,~onJtr~s;t c1~i.ldr-enwho be.g'llm W.ilh below;li.verogl;; \locabtJ!:;iIry seorcs achieved ~I~ st:rOl'ligest growth ill word re<u:ling skiU:s with,) ooUJmbii-

natien ,of irt!.~HuCl~(lin panems: Their !'(ffldirlg sk~Us grew more wllii.en~he:y ini.liia Hys(pelu mere t~!1l:U:: O1;l cede-based !IC~ iv it.h;:fl. \vil.h

_, ?'n .l-- ~ '"

increased eh iId';II'I,~ml(!1gecl me'l:iIl,~ngHbc used

i~lS1:niNio:~ a~ the :>c1'It)o~l..~ar pl'>l'lgr~:Sc'VL.'d. y Algorithms used by Assessment to 1!Ii!~tnl:cti(l~ (1\2i) W~b·.~\'IN!,;1d sulh~'!:l~ tJ2) incQrrmnltf;it~~eM::cl1 iM-by-inSIHL\,'l( ion iateracrlons ;:u1(l forei1.C:~. t;hild.pr"{wuc!e llX;x:nl~,lllend.u.~on~upcl,~ledlm(lJ.llhly :~~g~ rdi~_g <lnfilmlll ~~ i!lndil1'peS o[inSlRlct ion (fig,5. S~.aud 82). ~ndu\ridmd izi IlIJgillstnDcIlOIli\\·id'n d1e use of

words is I,eadiug ,I ~e'<lc~,er~mam,ged. cedefocused activi~y. In OlLdd~ 10 reeom mt;':nd illg. ~mnI.!Hilr:;; and !lO]'iI types of ~[Islmcl~on. l.i algorithulSWl.se chilA dr~~\.r~adin,g: seores ro help teachers effeelively group the children for :uill..,.tmc~ion. Lesson plan~.h,g sof[~"'·a.n:I~.(!!lps teaehers ii mplentelll this ~m.dividmd ized im.s~'l?'lUif.'IIli,onO.L1a da i~y~si~·l,fLPP~lldix S A ~nd fig_ S2 J. The hui II.'V ~d~l<l~ instruetien izilrng il:ilteli'Venlio!~ ~·~k1,.ld~a.ch~ffi i

,Abi!l'Ulttih,e

.!lIi1lb~i!"l;I'>eli!ti~'11

tutcaeh r~ldl ng for {It ~~<)!>t

lna

p~.·'I;V~Omi

.:;'wdy.~hiM'n;:~:g who boe,gmlfir~

grade w.ithb.elo\v~ilvc'r age ~.e~te'r-'i"'Orul.re."1cdlung skillzj dlemtmslr<llttld gremer impm\'e[l~.efll,¥ilh greater illi1m,l)Untll f lime i~1exphch le acher~ o n1~umged ('Qde·fm:lls~d imslrucl:ioll (wMc S[ l A1 i does not :reJ)re~UJ;~<:I new re'<Ldil].g CU]fmiIClI't 16). For stmlellits who be,g'Jn with <lbove'" 1;!I\!~ra~eBf~di'l~ ski lis.. gmgttcr :!:l-'!m\vthIg'i!re~[cr~ hum lHUI nHh~ir.l!lGW '\.\1;)' (If i~lplem~llh!'lg f : \!I'OI"d I'e'.ui iug ski IIS\V"dS <Iclwla~.li)'relaled to I.ess Cllrre'Dlit. [-e.u:ling pr-ogu.uns. A~,~ ~teracy ac~.uvi~ li~'inei~ ~,"";ilch~r·marH!S~dclx!e·ftl{;l!!S~1 m::livi- ~.~es;~ind~~.ed ~~l dhm.~m:io!1!inf iJilSttlQ(1100: 3 c(),de- versus, mC<lining-fm:useci. te:<IcherVC['S;~~:J; ('hi~d.,;m!.lIlJ~!§l.ed, and ch~lnb<e ,overiiIf!H: llfl.oridill (i!nlff for R.I!'ad~1IlgRe5e:m:lhI, F~orid~ ~il<lte (J61 g,.ple .~H). he d!i:lml.e·liIs~oll de~r.ibr:::'S T ~h<lf 'lllllli\lle~5it)'.lrM~i'lh~s~lee.~1L 3:230 1. UiSll.~Oep<lrh'l'!ent of II\)ldl~~~, ~S~[ho~11 Ed~~i~, 0 U.nlVtl/SUyoJ I;tijdligOln, ood~- vcr"-;'~Is!UCilll ing-roocu sed .i n stirli,1 ctiol1i AnIllJ"troor, M~48Ji~, USA. C.apl~,I:eS the ccmt~m.l of the m;tivi~. ('ode~ "'/llJtlitor ror (~rtSPOi!l;-d~JlI(e_ E'l'II'Ia~ '((OilllliCl~@1·5i.U~dUt t 100l!St1d hlSlrltCI iOIfm ~ne:tud~s '!hnSlii act ivid GcS
1

90 miautes (~dJ,~ydi~l.rl!l,g d~ic'll~ .! block of ume, W plPovid!e inSlfll:cdoml, to ('h~.ldl\cll with ~i l~!i nlaclinJg sk ~~in smal] groWlp.~. In.d!o U I..~ b < fo]llow l~l.c.1\21 1Pi.:!t,:on.mtlendcdal1lmlUl[S~nul sped fic types :~nsntlction.("omro] teaclm.ers !;Mere ahu e.1>ipec:ted to h:we 11 deJ Ic<lilied daiJ:y re~ding 'Mock ~Qf 90 minml;1l'l u!l!d to 'J~ sm311 gmulls <1c{;ol.ding 1.0 schoo.1 po~.icie!>. A:io~s:;m!.lll'lu!l;'Sll,lts \~.t'~sh(~r~d i~h w '~e~l:ch·, e]".~(;append i.~e'~~ A. <lind SB). A~~.tei'lichers il~ S

or

.~n

1_.•

the i.rciliJimelill gOOMp rtx~dvtld the :'>!'Ime".lVlau:mtt ()ftn~im i~lg OU ho""rlQ phUli .lnd inlp:~ememf :undli,- 0 vid.J.lialized rcOil:d~]]~ instructkll'l ~Isirtlg !-\2i

i:

Control lie.u::l~ers. r-eoe:uved 1'10 Ilr-o:Wessiomll d~wlop'fTl;el]t from our pmgr[Ull during. this

I
~
~

464

EDUCATIONII'()RUM
t::xpefi,!l'It'nH~1 phase, Tl~lA$. tlKpl;":IC[f~d ens f(](r Alrhotagh h is !~Q[ possible HI fillUy dl.s.. amount and strucurre of I'eml~ng unstmc~:L(n~1 effil.la~lglc the .urn,p:ac~ or ilUlI,:~vid~13,iizua:lg were sim.i~<Ir <ICIlA'lS
gl.'OUpS,

received, I).hho~~gl:l 1]0 doubr both asp(eC~Si contribute to better I:eading results.
Thus, i ud ividual ized readi ng ~.~];!;tructon i

w:i~ere!Jli lhc

.i m>l.mC:lioll!

us i ng Ali

ami. the p.r.o:We ssional

development teachersreceived, i.t is ua- guided by the A 2 ia ~goritllm 'J~.d cuupled with Hkc.ly th~l the 1.·C'$dingmucomcs were due pm:f€;!';s~ona~ d.e\l'e~up.m.cn'l supports teachers :I mi. their effom~() provide optiiltm]~y eifC(I:ive tiO teacher tr:aii:L~ill1.galene. The treaunent ,M.dh,olld:!l teachers ,fill rceeived [he same- ·!i!.notmt of f¢.~dillg~ns.trUCEio~il f~)r{IIU c~ild!'Cn, l"hh, cll,;iswr·r~!!ldom~~ fi~id ui~ll :!~,k~d n'!l~i'lii~g:. et variedin theaumber 'of mi~l..1 i < y R!!;!f'Clil'l!'Hl~S a"dNoiW~ utes they used A2i. ,uld tle e.... ~eml,l to which whel.her fl:lroviding indiv.idllarni.zed re.h].]ng :ll. rilalj'ona'l.I!i~wn;e;l!lt 01 Eril!lCOI.lionit I"~rl~~, tmf' im.mqcl:~~m(as gt1ided by 1\.2i ~~gQridl!n:o;J they iadividualized ;~n::;tn!i;;ti"JrlT Thue •. we Ji,l'at;a~~~~ft ((1I(N~<!iJitnaL 'Geni~f,or EdI;!W!I~o~ would cause srronger SHldc'Dltt reading OU~:- could examine these two markers of >~ti&i~ 1HastJingRClI'\,D(,ZOO5-)_ comes, The sl;~I,dy was 'Ql1dwu;;:t~ illm] ~Ihrli~ fiy,dil:y 'J\~'I~;eyrelt,uit:,d Ilochildr'lilll.l':s ~;'~nd~ng 2. A,i. R~notil~ i.·IR, Cu, 'liNd, ·~f:.!th S"N. ~ :1;6, 1. 12(04),. C:<lI~y ami el'()alOm~:cOllly diverse school distric~ olHcomes.~iu:ddullig the amours o~· tr.alul~a:Jlg 3. If, it Wllmll1illN oJ., j, ~dt1(.~ydI, tIl. Mil U996}, in F~.ork!8 , in Wi schoo !:ll.wh h 4 7rnc~chcrs eenstarrt. \Ve ~Wound tlwtl the rtI1ore~i.mne il. r, J, Mcrri~n, H, J. Bad! ~n, e. M. C~nl1Qf; lmp_ffiwiff9 and 6 .16 :l"i;~1il~srn de :i>~1II de:rus.. School swere teachers spent u:!ii~njgAli. the stronget were :trlef!i\!Y ~ "'rT.\efr.r~: GrJ~1~e$ jF.!'}ffl~h ~e-I!.!I~i\l'. ~~. rtew M!l,1l'M, cr:. 2iJ{!&~. m~u~h~:daad pf!i~l.I(lnlo:..,y imJk,ator$. \"ljth their studcms' read i~g ~:[l;jU by lh~ end ef ~ 5. 8:, R. raa!1f!1<1il eI al ••1.fidU",. ~ydr. '(II, ].] U99,Jl}, one 1.'<lim.io\Ll1i,yssigaed to t~u;~' a ueaunent group the ye'm (OJ I =1llO:Ulll increase in IV seore for 6_ lil ~ri~h, iFll'1lii? Gte;;ll (~mC'Ullilm Odro~ fil'Q.1I" Sl.loatd ,t JI "" 22 teachers l.md the other tD the eomm I every ..ddhi.oml~ .50 m,[:~luCS te..iehers used Wtr~QctJ fl.Mdingt1rlfjM~t1l;, T. lQiII!'lM~ Ed~ ~I'i~im!:i~ rm;lrrlllifioo Prell>, Wa5l1i~'~t.l!n. ~C. 2\00:1.1" A2i. ~(201 = 2.97., P = 0.0'118: lab.~c Sf»). group 'til = 25 (em;~e~~. Allp"nicip;"J,un,g pp. ;2 :H1"~28, There W3:S a I s;o a I1i ~I:i,[erae DUOI1 W il~~ :>W= schools had students ]~.vifilg pOVC:lft.y (table ~n ". i!:, Raty,nue! t11. i't~Il, Sli. hI~li!l'rml'mt 2. n(~Oliln, . dents" initial vocabulary :scor~3.WiiJel1 S2 and appelld:illt SB). a to:, L Dilh~ R,A, ~~p~~" 8M!i~y~, (). 3i()~5-0'(1~951. St~dcnh"'~D1Ilwm.ge m"ld.I ite~y sk~U!; were tca.chcl"S used A2~.more fr~q~lemly. onaver9- .P~GJlI~M~e!' .aI" i, Mvt"_ ~ ~~, ~.~~s. (2(1(1iJl,).
'!il'~4;'~'il;IJil'!

m::tf;!'$." to A 2 ~ H~(UmlmeuHJilhort<; and profesI" s ional developmeat W[!S dl rrere~lt.

A!lgl!i!.sl

2M.5 and h~ J..~u~ry .~nd

1(.2'4),.. whic!~

age, ~h~ir ti,lud~!ill.~

conrsol classrooms,

dl;:i!1"!!(l!)sH~W~d sU'r~!iI.g,~iI

May 2006 with the use of the WOt)(lcock

.!f1lm_~TI/~ Tt:'}fl,~ ,ry'/khif!I'(!maru-lIi

outcomes campa.red wh~~.clm.:u~.dlren who had

similar initial \'oc~bul;;'!!y scores in the

Ad.dhiomiily. in higher

arepredicuve ('If ped'orlll<lnce OIJ state and. tederal aehievemeru tests (1) (~t'lb~e I. S3
R:eSlmrnJtsil!l)1(1I 1m pli,tatiion:!!

Our resu lts

ll'fl'\.'e .!led rhat iI.HJI~vidml ~ zed i i nSI[~leli{lJi'i!. ,~u~d~d by A2iiWd sup'port(\d

fide]ity classreoms, studems with belowa ver3:ge i[Ii rial voc.~btlJ!;n:y ski I.b 'I~ncludi ng ~UIIW ch iIdrc~~ from hi~ h-pover ly 'I'll ilies ) m achieved read ing scares that were ge[J:era~l)I
CO!ll~1i'ln:lble to students wiUl!high~r initial
VOC<lb~L.~<Iry
SCOlre"S

S3, and lhe figuee ShOW~l bel.ow). Taken with th,lt ufdu: ct~mml group. ctJimltJi;ooUing ft~r tog;1;':Iher, us i A2 i <n ndi mp I emem :ilmlgA 2i f.al~.read hag and vocidlo1i,llal)' scoresas \~;'C~J s, rccoIllID~.end;fllj,ous .in the ,dass.roL')mappcar a child .~m.l5'Clum:11 c~.trd,cu::d:-;~~:C51.we~idu1J~~leJi to cOlmibme W :-ltnmger ::>ULo,em mncome!l1 mecUIiI, diJference -- 2.63. 95% confid,e:m::e Olver alJ,d .Jbove 1~I.e tr<li:llJinlllg le<I:che:~s iilill:er\'~I] = nJ7 m 45)0: flpp~ndices SB ;)LjldS.C ~nd mbk: S5 t ~·lJMt:r ri!;~~c~buna~ 5~cre~ Ho·,:v.~',""r. i~dv\'id mil ~"jng!>nLdI;11l in~trucHig!le.1i!1l1l(lC~I:illlfui'YroJr6 L~O~,using Ali !;Onw.are. eVl.'!:I1Iw·iAh pa.·Dfesr :S lom.~. d.cve~O'pllrlC~n. p:rovct! c~:a lIe'.~lg~ m,g I('u S(}JIiI~e on~~ teiu::he~ il:li the ~tudy.\Ii,I'e (olptured fidc.m.y (if U"C,;itm~'ll imi [\,\10 \~y~, ThroU.gh classroom ob~el:''if,''liom; .md by ~eC\mtding ~jm~ $P~li i,lt';~ll!J.Al i'~~rJI"';~fld~ SC)" Bl!s.\ld '~Il X cl.n\c!'inJO'~~ldb..~!i~'fv.~lion~. FebnqUlry •. rline by Icachers, ,t 41 %) impl!emelmtIld l.beUlllerventiO:IilI wi ~hm{_Jder-,ue~ohigh fidd it)' (~<lM~.s4), T\\'O

by profess ie ~,a~ deve lop llml!e prcmoted i'lL stronger :stlld!el1i~ .1"e .. diflggl"O\llil~i~ compared

(~<lbl.es. S1 <'I.~~dI

ss. ~,;ig"

8;, Il'i. lilyl.lJJfill al~ E1Ml', SdI, J, l'(IIll. 121 ~20001. · ~,."~nr1ingh'lllh.!:L Hanlin '~J\iin.9 f~r:J:jil!'U.,eI)' D~Jl';wllftJ.ing ttl tJI~ Sch.o.oJ~ Mil G~.~MIru, I{,it Harris". S. (Jrillla,1D, 'I). ~~II'~.r;, £dl!. ~Broolllmi'! ~5., Cam1iri6gI'.1AA, t99;8}, p~, l~-16, :li2~.hi" rr.ti'!l~ Ittt1dlJl~' fmYlU:!iM .7ht111~W: lh>e r(1~f!Jf 80lai'Ked 1ieaffiir.JJI'~Gu:iLfWd\ t>!..:w'!o~.199m., 1 !3->-.~ A'O~~ il\t @,,' .~ydt. Scll.3~, 'll;ll M. ~a~aRilll,R~dlnSi P.al1el,. te.adri!11l rn~drelll till IlllJd: ~ 1 'e-Mii~!:m~~l!~~d ~,1,1!;:~~1fIl~"'~ Iih~ ~Cj~lmC Ire~~1(1fI g of, Iiitei'.!I1i~ On f!!!MIil9~i'id! lt~implit:.!tI~Mleit iU~ii1!i i'nmrdiom· OIlIH Pijb. Nn, OO...¢7M. tfaiumallll:!i~jjUUf' ,ml'Omilt!lH~I(h a~~ liIumO!lliOMelo;Pll'l!l!~~. llI'<I~lirng1tlii'l,

m.

'li]_

u.n:n.

'.g

IDe. 20!iX1J. 15. C. M. '(;Qrmcr a aI••1.Sdtre. P5y{h, 96, ,69, 1200.,), 'lih- C.i'it.'~nm()retul-. ~i.. SfI.td; ~din91'i11;I,{JS ~~n(l>\ll ]1. Cluel. C. Mimd·~rn-~IlPPIIi!~.~Rfs. (,), lS, 45a:'~2000l. ~{!. C,!it. 'G~flmgr~al,. XP. SfW!. ~~l~9'11, l4~ ·(;l007)·, 1l9. CM. (rCilf'lMr S i!I'l,. .~rJu~. $Jtt1L '}8,. M,S (2006). P l~. J.I(, 'I'org.tSl!<n, ,~m, OIJ(JfNI. f(fj. htln, 1~,55 ~21i1!iU)) .. 2:11. lit, 1_S~~~tibef9..ln HllOffl/lJ';)'Ab;\Ui~ lh~ lIff:J1u~ 11M !lWilW'.~M.I, a.,'DI@Mi~, [II, nil'liiJtn~e ~11I~ilm\ H!ll~at~" NJ. l:~96). PI'I.t6:1-1.9 1, 22:,.11 ·.~ther ve~~~ ,gl~;!i 'win be "~.!!il~hleo ,ai1Jer l,ll.ug!Jilt

2007. iI~lormaliMl doolil A2il ill. 3'iIiIj1aM~ 'Dm~i~e

~rnUpftlU1(i.,.,.,'$I;!~.l!.ImiC!i;,~~A2:~in.np~,

~11!i!!DSI~"

Alid~mo ~ii'ld:j:i;t!SWDrdIS'[I~ 1_ (lid:: ,fie, CI;"wornlVri!!lll bwll.oll tGw::aled ~Il>l1!r!r 1h~I$i II.Q!lliil'IQ· !;'1f!11i11211 lll@ soil!· war~ .~f1d IPf;D~ol. 01re 'ourfij:~jl)' ~~l;ib~ 1!il gjjlfl~i'
'r~Q!.!lrC:hll~~ u,pon r'llClll~t.

23. C. E. !if!C!W" R'eJMJing{Or UndmMlIcJifl(} ~ItI!iN~ Eldw(alioo ,~fld U1!1Sdeno~.!!n~ Teth~~ pg~iC)' tll'!>tilu'lll!, Sa!il~ Monk", CA, 20(11). ;;!oil. ~. M~lil~r. It 'III, WaO!!tad;, ,~a,kockJ~ fJ/ Al:M~£mMt fJofl1'intl~~aJ ~ (liMBic!!, Ilasca,It.

rJrret.~

Icm::hcl's, (9%1 did not Ul1lp~.clllcm ~'~.c .inlervelm~

oJiOO

tiOlllll mlxil't!')~\'lDrd the. ·end of the ye-~f" Tem.::.!rer:i!· mre of A2 ~£oillw"dre HI so vlIried. 0[1 <'i.'\'l.'!'r..Jge. !h~y us~d A2~ for 180 miml((1$ from I Septe.~.l.ber to 31 i\1.I<I:Y (I::<Ulge 15100 37.t:1·mi:mI.~ [!~~~"', tldhtl!"li = 19 I mi~lu 1A;;S, )," Si;.: .~a,dl~!'~ m m;e(]~he software fi)il' l.e.'!S tlmll,l I OOml~mll'Cs. whcr,e.u; II teachel's uti~d [~~esofh"'l.'Irc far mOle thall 200 mil,nate~. TeOlc~,lI:rs \\'l~o used Ali more frC~luendy !ibmle:.ndl~d to bcl.t~r implemC::'a:J.t imli\',ild~l<llized inS,tli,1.lctioll illihe d.a&'l[f1'lYul~ (cmj'ltrCIO\I imm. ~. == 0,34; P = (to l41.

eo

....

,Ci

ra-.
!:""!

'termillal.5COrl'S <Ifoe' ~h~n r~rdhildren hi ~he (mlltfO~ gin) up W n;:I@~) i.i!llili lit(l' If!l!lt!l1iC~ll grO!4!,p m !fl!!'s~; t.ro.a rumcml 9~OUIl fe'5Ut'~S llIaIJI depend img on tea<:h· er~' A2u us,e Hallle 58).

!.JJ~e~fI!'I'Ii~illLlte~ In1ieradilOIilI It!eitli'l!~eI1A,2i IIIseOlil1ll1l;1Q(..ltIlul.a'~y, :Sprillg ~.o~~ lor H~l >g;r;a(f,er:s wi:Ja :slar1eiJ ti:J~ y'~olJ with bel~w·.a'l,lerilge vQc;<I!Dulary (DI!lf~d ~2S.tl'l p~rc.el1lil~; Ws.rOrf - 474; a;~ ,eqllhral.enl:., 6 YE~rs}and a bQ"'IEh~yeta 9>e ~()c .. lbula ry be d) (75 till peroe,nliil('; W $C(!ir~ "" 486; ~g,~lIql!i·'o'aliml. a: y~ar!iL M~",!'!

1\2itotal

:3 '"

c ...,

'"

::;: ~

2001),
0

""" ""

~5- ~tm<Jfi'kA ~lJ,ergll!JilI!'~Ii'l\'ir:e~!7 ..rromerl.l)iIi~ ·mm~diSiritt"INldiJ~ .. iif!1i~8., ~tIldeifll$,.wI iIiI~ lSI' R.~emdl!I\r:s, urnt!n~ IfIIj5l1fllMided by'lb~ Iu..S.II)eIl'l,rtmM~ F 01 (t~!:io~, In5~'lJt~fQr [d~~aWn ·S:Ciemo~. (o~nijj~ .iIIld 5rud!i\fM ~'f'il[Jf1iiI19 (R~))!i~.(IJ1i; <lnd 'lilt UilIiQll'iil rnr.li1JJle of 'Chid! !'I~l'b Oil'lL!! lfhlrnan De<r,wI!!!1ief1liRO! 11I000fJIi World Om ~ts. ~Il~ Ja.~ 'Qltil!ibe~ amiCI )9), ~llIlCr\!t~ U.tC'~ f~J£, O'IM·B.l.f. 1M1, irll'I1t!"!iJhi~!, lh:t'i\~ ~fin!lDd~1 iilWJl'S!~~rn~A2i iIl~~!iI, !!!i!P\Pllrtiir!9 'Clntin~ M;at~lii~l
¥I"I!III'f.llril:!flDI!rn~!J.!IIWgi'tlJm!~nUlilUl3:!l~!ll1J.4i!lnxl

W'W'\IY,scien cemaq,!J rq

$IC111e: NC::~ VOt 3115 26 JAN U.A!RY2007

GENET~CS

,coI1sid~r@d

Single nudeJl)tf:de polYlmorlPl'Ii,$·i'IiI$ tl1at are "si ~!Il~"(;:[I 11 ~n~U ~romir'l f~ !di~1!19.·

are ~m.''''~.] en.etic. c.'lrn~.nges.1li.IO~.Jnd ~.~]..' g : illli l c,,~Jmg Rnd l1011cl(Ioi;hng r~qtJ{~~~ ol th~ gt;:llOllItlf.'. The genetic rode' is degel1l.eJ.7Itein thoU ml~'ii~ .!Jn~ng acids· ,tII: I:eu)l;~~ntl.>d by more UJI one triplet ofnm:leotide bases I~alXldol'l), Such

i!lgle m!{.~~~{iltid£ poly!l1;[)rph~Sj'us (SNfs)

MO·rffi.a! protein trenslatic n ki nNlc;
~, S'

1m

codon .. are COItsi.dG:red i>ynf:lrnnyliilmuii

MlI~lIy Sili:m~.SNP
(nnamlno !Idi:l dI<I n.g!J!i}

SNPl> me' "si lein" as they :re:s.ult i[I s}'1I.lO:I.U)'IOOLlS co~h1li1! ~ub~hl!l!tion~ B€~u~e ~h~~ ~ilelU :SNP'~ dora.ol clmnge: ~h.e amlTI ina .,U:UIc] composi ~iQU of"
the'prown Ilip~tldm;t,
they have ~fII.'gdjl bee,n,

AI'll!md prlll1:e~flI t rans lat Ion kimHL(~ , ~ .

).'

esample, a neuroaal ~ soform of CyU1P 1;!'srIiIl:i,c rmly2Ldcnyi~lion d~mi.;lf!il bindinM; lll:'oteiln in the sea, 51 ~I~ (A'JJJ,.t 'sier clIl{(m1iic{( ~exi sts in I:\JI,fO o,f"~]~-e g1.,'n~ aile' e:n,:tym.e"-~he:~~.-encG'of ~1j~~~!1~ rUfOUS: a S(lllul~:I~i~l,l£tiv,e f(lmml.1l~diUl the classic view o~'ge~,le structure .mltl I\IIK'~ [i.nsol~Jble active fO["IIU lhlll re_guhues ~)lU.1.<Ipl,rC dOI:lr-~tm.ted 10 break dJmvn al.m.os~ as, soon pll)tein syn;~:11esis~4')..The mclecular meeha'I~:~tlliHld bee iii, CQmpleldy nl.lrnH~ia~ed (2). n~::>m:;;~~<'II: can dri",-e Ih~lbldi~lg (if" PQ~ypepCel hnJar processes, ill cllildi ~~ga:~!c],r.l2lltivc' tide ehaln ]11 vlvo hUn <I~lCrnat1veoonlbnm'l~ RNA 5plid.l'lg and pesrtrau ~1'1Ji~JI~~l~. ill tionsare stiU mosdy obscure, protei medificatiens, cl:eme IHo:re than OI1:e proteinl The erowded oei]lI~ a~env~ro.~ul.le:r,litbid! illg I product rm~l Oi gi"'>ell sf,"(lue~.oomI! rhe g~~.o!1m.~. C<fllfl~y$~K OlC'OCs.snry pm~~.ins111 i and ~ C('iI!Ur~bU~1i: to ·p.m~e.in [ohhng: in the oeJm. (5). Could there Vet. iI is widely <I;ssluued that a s.ungJ:emessenfactors :JJ:1l''-'lC:~rig the flnal c{~nlljrm~ger RNA (!l!'IRNAlI;;IDIo11ding a pol~]J,;ip,tid~ be (]It~lt;:r Chfli~], ith <I defined am.ino ;u::ud sequence w tion of .. peotei n:! .~ 1937. Purv isa nd co.i~ 11 wm gi \PC fuse to, durh.g the l'lirooe.;;.,,(~n rn!L'DS~ leagncs proJ~osed.II'II:Il·"lhe \'\:1'\)1 in \"h~ch some hit ien, i.d.enlk:.~~ :~ro~r:.i p:~-OO.Ll.c1s ~nterms of 11 proteias Col.d I:> <lilected by~lrn.erares al: ·which thdr t>tr.lIcwr~ 1}lHJ ~~C[htily. :>S~lill:l~rJ,g tht'il ~ oc-g~Ob':i:l> I.hdr poIYPCI~[~di;,: dl~~int,;; are T~lnN'of these pMduc::ts ~m;d:ergothe s<lmeclO-<llld :P'OOt~ rated UI'I 'i,I~"O"{6). They hypolhes~zed thaI. cera h tn!!!~ hltiot'! ~ I !llod~fk:<.'lrio!i t[l~ ~~IL T1~ sin t~ H!i~ "g~n(,: ~";9qtta,;:~~c~s aw ~\lo~v!.;:d [(]! ccntrol .1s:t;jJ.mplion is based mol !hoewi.de:ly accepted the rate of tr.ln~l~u:ium.,d~lmlg"~lL!i)nuchthat s Allf~ nselm.':s ~}]ji~.H:le, \'.\111 ich states that the ip the synthesis of defiued portions o~' their in~orm.uiml necess ..ry to specufy thel1lmi-rc pO~~lpel')lride·cI\I<I~f1:> s.ep~.nu:t\d ~en1pm~11 y." i;; ~ The r.IJC or rJ.bosonlm,e~rnmc· m~ InRNi\. is th rce:d i~~m.e~l sioll!ll SlnLCIH rr: 0 r il!'lmte:IJn\\;.~i,ch l<ug~l)i diefines irs rll1Lilc~·ion-:ii$. COJIIIkn~\\fn to be JilI.o.nl.uuiform andL~,l.rinly modu· t;J~ILt:dsO~iely .i[11 it"!> amill.lO acid :se[llIelmce (3)1. :I:u'ed hy both the gem:.rr:<l~degenemcy of the A ~th~u[!hth.~ tl:l/(l prillCirl~$ (',01'1 ~ambe' ~ni;llk': code (!li:U~l~ly. l~!'l:ofs~(~inC' syn", the: iII,pplied ~.o<1. wealth gelJ1.es .~nd 1~1'()~e~.ns. onymous eodiolls tha.t is chm.,.,.c~eu.'.usl~C; for'l 3. jju~yp!;;plid~l:ha~!'! ~m,~,..isl i~ U~e c ,edt in Oil. ~uv~n~rg,lllilbm)~mdthe aV'Ji~flbm~y Of'l:{l£,'[1;1tunber of alteli,lJ1::tlive OOI1i~orm<lli.ol1Js. [revealnate' <lin i~1.o"''icy[I,-lr·drac..; RNAs (tRNAs I dun fer ing differ~ni fi~[I!C'liom.; <Indff:l(l' i!lclivities, l::ar ~tl,rr{lmld, t~u: 1:1 bO.'H,];u!e dl,J1.lii~l!ilt~·l!l1a~.lal~,f:Ji[l! .• poi~t:d to d:e~iver mni.m.oacids" ,"Vitllin, :1rn.1<111Y O~fl!.l~S!n!'; .11 slrongcodun IIia!>~ists (ii alld The <l!utihw qS in lil1e' Departrme!ll o~ B1QI~k~L Ge-obgkail the amQl~l1I;r ofcog.mde tRNA5IS. di[I,;ecl:ly prOa am!!![[jn'!'lrol'1lnellit~~S(ltlil(~~. (~el/!i!'~nil Stllf Uni"eiS[W, po.nion~ll [0 ~h€' frr:equeneyofc(ldoll iU&f1~e. IQ@'IIiI1Iend,OH 441l~ UJi5JI. e;'ifIIIi)]1: a.kDlIll11r@(!;lOOhio.fW.

e:..ert ao discemLM~ elf'ec[ (!Ill gene pheaotype .. I]ulm:c silent SNPs uliI.d.eed ;d\'!f<liYS $'iieI.1f! On, I'!<~se525 :in this :issue, Ki.m~hu.;S<l!':fuly w {Ii, (1)1 pmvidc evidence dl."t~ l!lo.1:hJ!r.!lly o'~~U!Hi;Lfi!g UI:!U SNP~ c,,~ a~lt'lCt i.i~ s vivo ~~J,u~dl] tbklillg <U1:d._ il:O:lliSequendy, funG~ tiou. The sndy sh()\~ thm su bS~Ii<I~e fk:ity s:I'le>ci of P~gl}'CoprOleilm. the' product O:l"neUl~lI ~ridrug :res~..ill1("t'. ] I~M I )1 gene, is a~ .t DR. ll"[(ld by :SNr~ pre:S~111med~Q S)':I.~O(I,~}'mOllS;md si lent How is, be l]n.iSrx1ss,lb~~I! The erigi Il~~1 Beadle ami T<I~'lJ1'lrnll~ypoli:l.eSlis
n~ll;etami tlil' i

assumedto

Pinillprot'l7!jn (diilliei!EiVI C!MIQrmatioo a rid ~ui'Kll:Gn) iIilF.~rljS!~ati(l" kilneti,::S o1l"d proH:ii11 f·~lplil'ig.UrlaHe'~dtr~p1:;;ta~i()n,~inc(iCS~sllJll'S ~na ((jrl"~(Hyroldedp'.o~!!in.A.bJl.o!m'~ llrantslaliclr! kinetics, esu sl\clliy thl! ribo~!Omc Im1Jving [asLQr.ofsloll'm th rO'llglh (@rt;ain !HRfMA fl!9ions, ea III pmdu(ea. diilr,e rent lillial. protl'il'l mnfo ~m<ll.hm.l!JJll.onna l ~netie~ m!d~ ilri~e 1m m a r.i le;llt :5in91le n lICIM~ld~ PO~~Ili'I(lrphi~m {SNIP) ln a 'g~i!le thall cmtE':s;i31 rod on syrl!Jn.~IiI'l,oIlSto the ~~ild'WlPelI!iI(lrL l1Ioweioter. ( this SYIllOllyt'lWUIS, ndon ~ub:5lill!ltijOfi may ~ad to ciilfer'QiI'Ii kil'Leti~ Qf m RNA ~pmte'in) traastatien, lh WI:; yi~kJ,,· e iln.g a pmlf'in with a (jiffeo~1fl'lil rinal.structure al['l.cIilUIRutiolli.

11ms. ilrnfrequC'IlI!:OOd:OIliS illl m RNA ~ippe<'lrto .bL:: slm.vht translated, whareas f~"qH~~t cedens
are :L<lp.~dJ;y mms1<l[etil.
1

Op~i!lli~3Hr.ln {lfin V~\'tl proti;livn Iioh.li!:'I~which is tlt{m~n 1:0 be cotmnsl31iomlll '(5)111lIigl1t~nc]m:le the .u:ilapiJllion of eeden seleciil.:H'I 'lkmg DNA land henee.along correspendhllg mRNA) ~o ill pm:dclIl,lli uanslaricn k inetics...Temponl ~ep'lraliGn ofroldi~.l.g events during protellD11 s,YI:I~hesis 011 ~~,e ribosome m]gh fhll]z.; <ffvoid "'L.llil:1.\\r;jInt~d"'im~nL!:,~[i.{)m. within ll~.e growing polypeptide .• ens,lw.illg high y~.l'ld of the C(lrrocaiy fnlded rr(n~in. Tbere for.~·.".~~:C1iet ranslat ien killl,e'~ s of <I d ic d!cf1ncd mRNA due to •.in I~rlii::t~i~r, ~y[l!QlIl:ymous codon substitutions wh~ch\li,'Oll~.d·1I1i.Ol a1L~ii11iI.t.~a!!liI1~nO~cid~qucncc ort~~~~nc(]od~d polypeptide) ~~luShi~drive thein v[i.v{Iolding f uffllii~ ~';Oll!~W rniW~ptijjdech<l.in in~o d~!reii~il~
I~

confUrn'iliM ioas (see lb;el'ig~fl:),

or

Tlils hypotlle.s.is i~very d.ifficull. 'to preve :in Y~;\,Q ~y5·~ems.because I]mllh~[OUS ql[I.8iI.UlY=COI1i~Jol mtx:l'nannsms CX.USI. ~.o get :rid 0'1' imlJ.::o:rrecl~y or "bmHm~II.~y f'oidecl <Inc] w'[isl'" ~;o.l!ded p:mlJeins (I~t o far. sUPPo:rli[l.1g e""pelli~ S f1i'!M~T'Ji dm~ hil~ blll;!1 ~ilrt.:'e. m.wkhr~g l~(l ifi.n.di:ng.. of Kimc]lli"Sarlaty et lit. j'l<'lrli!Cu]lar]y . impnrl~Lnt Thl:> "Iuth~)f~ sho\'I,' lhil'~l Qn'ilb i:~~c ~ions of the th~.'l"e :PI.'~v~on:dy kniIJWllrl SNP~ IFo]' the MJJR J geNi.c ,(C L2:36T~ 02677T. and '3415T~ a:~terr·glyl;Qpm~eun activity. P-glycoprold I1i iso III UI.I Ic-trn 1l~,mCI1'ilbnU1Crmlip leull p~lmp lh ..t tlI'lmlpm.ls. ,I,niom; drlll~ out of ~lIs. Its fULUilC'dO':~ Co:lU thus ~duoo the el1k.fl'Cy
usial!;

2:6 JArN'UARY 2007

VOL31S

SCI~NC:!

w'l,iVVl,!'.sclen(;;'emaJg.org~

IPERSPECTIVES
of dl~nrlilllhet';Tlpy lOO'll~mealU;_ K~f1"I~h~.·~rl&y et {d,[tnmd that r~g~.yc:opirole~nin,h,ihiwr:s CsA ami vcrnpmnn~'fCre less effeetiee !l.g<l~llstpro~ei[IS thatwere produc(;;ldffom !mplot}'pe~ rljl~~
('onsi:l!:tof the p{loiYl1uJl.lip~ie dOlJlblc (C JI:2 T~ 36 'G2fi11r. (' ~236T·C3435T.G2:617T;"(343)n or ni~.~'~(': lC 6T-<O:26 771 -C343 5T )\'0ri.aiH combiil~~lium .. $i.igg"~li llg t~:iil these prnk:l ij pl,udLI cts ll~ ave altered couformml: ions, Yei, C 1236,T1III;'IdCJ4J S1I r(l,lytl~orph~:~m~ do n~]~

~lfllim~. L imited pr.l\lt€-nlJf$i ~ and t~~.e ef [' U~

ClQl'llt(lIiInmuon~se'nsitn.vemmlOC: indeed revealed structural lonal <lillll'bOOiy
d~,frCI\CI1.c(;:,~ bet-

Uktil~ho[)d of thtl dl;.:\lekrp.I'Ii1~IU andprogres-

sioa ofll1,],my di.s eases sHell ..SAlzhe imer

ease. rJrTh)upi;ll(:lI diseaseleading
W

';5

di s~

a n::fimc~ ive

ween t~,le\"rild"~ypepro~~in
pj~ic hfliplotYPC1;.

and the p~)Iylhor-

defec~of~he·~ye). and others dmr are slrmlg:liy

Iinked 'to SNPs. Thus 'klfil.awledgc should also betaken .~ account in persorralized dims n:w lrC~lIl1'iielUand d~wlol~~n(~.iM ~~ro~"Hlml".
R!l!f'il!i!fmHl~~

tn

Anificial site ..di'rec:~ecl silen! Itml<~sell~il' of ii:yrtOlhj'I1nOi,J1$ cedens (ch~n,gil!.'ng fn:lm jlnfi~'_ t[u~m hI fru(,iu~'~t) i!l '~ei't~~i!'iJ gene~ ~I:h;,u f;~iPporlt]n.e~,ypolhe S!lS dnH alterednanslat ion

"d!1!"th;~ufmRNA.m~gh~ un~r;;t fiJfl<il~ pru'ein

cOl~lbrnulI[ ion {9'),.
[J ewever,

:ll.

chaage the amino

aci(~, equence of P"'glyc~ s

1I.l,l1It.i11 slliildy li:le:

j;iuii~i~l!l~dlmlili'l~ 211ll!!cemliet 2006, <CHI. U:i!~eiem~"

c. ti:il't'l&ii~$,a1'la1:)'.i!'! tll,. :5ciMC<!' a;jl'li. 5i2~ '(2007l;

andNOiW~

proi,ein. The

[2361" pO:IY~1I·1m:phiMn chm~.g~5' by KiL~~chl,-S,urOlJy(:1' ,al,., there 1I~,dbeen no

example demO:l'IIstta~ins that mnur,dly eccur\'an~~ions ilil _';YdlOi1}llnt11lS eodons in .11 d.efined gene can give rise toa prmein product \'V~lh the ~fI~~!J.,':fI~uirJo acid :r;;c;)qlQ~lI1!(I;'; but dH:' terent strucnual or :~hm;liOl131 feat ares, By
rill~ demm]~m:'lIi:~lg~'~l:lt I:hi~~ indeed the is
("<II~e..

:t
:11. -l. 5,. 61.

IHi3$S), E. L. 'ififl!lIm, Pau:,. NDrl Jl.~ Sd, LI\.:~;",t 27,499 (19'1111 C'.D. nfiM~ Sdif[j(f 1iiii, ,~23:~::n9?3). A ~,:5i fJ ~i~ i1t~" a9lI; (;200'~. (ell ]. flydmi:ln, A!:trn.!. !i:w.. 8Ii~d'wn.7'~. ,61!1.~ {.2:ooU. !l ]1. !?w!f!li~ i\I~@'.,j •.~ 8im.1.9;1. 4:1,3; ~!l~1)'. p, M.Sh,np. f. M. WltiiJ!1t J(~ R.MMlr~lU":Urli!"rAld';1s
·G. . !ydr~. W 5 izs ~1'l'll6Jo. C~1.12'51 443:

a GGCcodm.1 to GGT <II aminuacid p()i.s:~lion 4'~i of l!ile polypeptide (hot:l.l encode g1yditc I
and the C3,4I3 5T po~,ymorpb~sm changes

ATe to ATl .U peshion 1145 lb~llh em::ooG: i~O~ e'[LC~,nle). i~O\li,1,e\le'f. bath PQ~yrnorphi Sm1.l S re.!ul~ti,nl, ehal,lgcs Ih.lm freqtlent. 1i0 infrequeut cl)don::>mld[~lerefore ~ll<1.ylow down ~le ribos .~ome 'traHle at rthe corresponding mRN;-\
regions, These <I~.I;et.\u.io.lii;S ~11<l)! I mL~S alii:«:t the onlr~nsll!tiiornl~ j0oi.d~llg p21[~]\i!.rl.'lYf P-g.iyc:o. o p~nll;~n. re~l!I!lt !1gin" d iA"e!¥nl I·~f!~~~. ~ I;.'~ Inf~lI--

Rt.~ 14,

the

11:. 8., Butoo.. D, v.~IM.mJml A.lttolil'l'lth.

~:l(jM)" ('1999),.

slmly opel~:S up' a new u\'enm: iijheseOlXch "lid

sl,lggeg!s dun s:~~eDllSN Ps mi~l~t CQIJ,'tr.ibilJtc to de ve hlp ment ~mdpuJ'Sr~s:s io n of certa i11 diseases, If this. is the (!f~S.t;" then siknl SN~P:5 slw~Lld no~ be ~wgl~etl;~ ]f! d!$(~r~~liJ1ing th~

9. A,A" ~milJ. 1. Le!!1i~C, Rtii!i~JE8S:tWJ.462:.~:g1

Fu~lf,5fu edi ,J;gH!!~ 2'1 DIil~r!lboer 200« lO,U2.6,i\!;d~~ilIi',1138'2'39' l~cllOO~tn~,Iffformali!lln "'fltn ci1jmgllltlis pa.per.

Huid jeits ,o({lIrilli Ear~hi'~ .atnl.~,spheIN! and

OJ:@iHI!i, and on ~Ith!!r1J'!;[!II1@t>&.A new tiliror@t~(al, 'IIij e\V of jets; ~i3iS h@lpedl rew.ahlilhy

lhey rioUf[Jli3iild ihow th!!y il1t1HiI~'t~!{itl1~h@'sl.IllTo!Jl'ldlingl Uuid.

o:[fle:lm COIfil.t<Ii n most of the k i,l1:cth:: Jf:]Qcrgy ina nowi'~g ~lllUkl.TIl'f;Yare pervasive l:e.lt~l,res of Em:1i~'s m]IJ,];OSphenoeand oceans.where they lro~"I)[,_)r1 he.:'Lt C'h~!llk·';JJ~. !l~.deven bint<"lsuch ,IS krill. ami. they <lire a][5Jo ;i:lbund<IJlll on other p~m1JJ.;t. (s~J,Ci~he figure], . Jel'S, 3D'e ebserved to OCCIJII' spo~,HlIU!ml.sly' on n1m~il~g p~3IL1CI;ll whenever $I.ratificd ~tmo~ sphe res or ee e'ans :ui'elio:wc,ed i~.lO turbu I.el'lt mOI.knYcr therels a n'IY~I~fy [0 why jets a... ex,isl ,11 a]~.-'I"'hy lis, there this propensity 10 ~;:n!1{'~~tif<11.$ ~~';;1~}' ~I~d ~wm~~l!iIt~I~~f:f\ ~'Cond my:>\~:rious pr~~)eny cf jets is th ..;ri~y !;:.111 act as (1e.,xibl.elmn.tcil'.ial barrlen:s, :~Ilhibiling Inixing acfll!\..s !heir axes, T~.e ,';i~nge:';t eHs:t\\\ElnJ je~~ prmr[de ,expre~wiDys fall' the lr~nSpo.lI'l of dl.eJlI1UC~I~SmId bi.CMI "long their axes but severely DIllhibil mixiDil,g OIcrl()S;S lm.eil." axes. A .
~t No!'tlhi'!1est IRes~a~1iu5l0<~atK, r 1~I~evllill', WJl9'8~09, USA. E·I11I~il:IIlIl~~k;@'lI:W.I'al.Cl)m" P_8_ l~hY1!e~ij$ jnllie ~,p<!rLlMn~ QI OCNm9Qr<!~~~. up.~si'I!I'

et streams .. 01,· 'jjClls" rm;,bre~'ily.

are ron,

new theoretical

Ilaradugml ,~nexp~.;;)iM t:~.e

~llmll:}t'lty.mlllg~~

~Ilg ai']uhl

0111 rotating planet w.U11i.nM.'triably .1

formerty

cenrrated, intense, d,m,lg<l~ed now:;. tlm<lit

abundance (if jets, il!1.,my bulenn

p[(ldul;."eje'~mtd that lin: I,\~,'Q'mysteries, jet forumuil1n <lnd ~ht':h~hibidQIl ofmi;':il.lg.

phere fl:r OCCainl., inlfl 5hnpk ~inilu,er. The psmJiig~~. comhi nes fiddl1fl.eory 'i\'it~,ldmu~'ic l tJLJr~
il[i)l~raCl~£1i~S lh.~l fire Cli~td;;df:h:u rorn~,in!l muJ :S~<libi IiZ'i.llgj(!l61 ..h ShOWiS th;n m ix-

1I,;egardet~,l.'ltwo ~!:pm:'.;~~e phenomena, are iariBoth j et romn],1jlif1(ID ;)oJd the iiil~nb it ~Ol1i 0 r ~~l:uxilbg completely en~gmmic inrerms of <lire

11ui.d Im.Hlliul1.: i III doing- so. it ,cO!p!tmces Iln<!tel:yre]<lted LOeach, O~:~.ill.

long-n;;nge

Ill. P: Elaldwlii i£

Gf W~sUni"giton, Sea!lie. 'INA 98195, 'USA. E'ff1~II: rhin@s@)OlS~a'n.WfI,strli n!ll~n,,,;!dll. li't·P. 1l-lI.I.3ngl 1S iI:~ the UIITiOOHlo1ierl.)' Earlh !D1i~MI.or~. f.atiSiid!es" Nlf 1.09\64. Il!lS.A.E'rnlilil: mooi umbfill,edu, M. IE.M(lnlv.rI!' ~~illli the D,eP'<lrimemii ~~ Midtlhematks and: 1ihf~·eijcill

J et~ lfiel1lUaliidl far. '~il\Jei} Ma p o~ 'J:!aSHifE."~t CUIU~II~ spee.ds at 40Q.m depth, Silif'lllllaterl'tIy an eddy· re~j)hl'ing

Il'\h1!lS[c~. U:ni~erslil~ O'r (am'oridg ~,.C<lIft!Ii"ld':Q1!!' OWIi, (IB3

IUK. f·nNliil: m,e.m(in,yrti@dQmtp..callll ..;Iu~

(lC~~iii modQ-l, R~'d~ !1Idblu~ if!~ i c~U~ i2'iI Sltli'f~ ~lQd w~stw~ nl noli'll). IAd~ ptl'd h,om Rid! ~rdoS eJ 0.1'. (1n~ r(8 '~R.i,~M) Smilll :iJia~ 01 <! s;~mulation for ]uljJilI2IF. ,ith me! <lno MUll! iAd killl~l'Igi @i!,stwatrcil andl Wl')tlMil rdllil<I'Il's,. w [Mt'lplerliro m Heimpel. el ,(jl.. (16)] SClli!:,NCI!! VOL 315 26 JANUAIRY

~,:soienc;ema9.ar9

2007

PERSPECTIVES
~~~b~I'lem::e rheary, ]n [he $~~~.dafd from cfllubj,~dng d~c nrrbuleace-thenretlc paradigm, "I ~m:b[lit'lllt How pUIS Huia :pru1tdes. idea of mixi ng \!i:.idFn. rhe rield"t heorerlc id.e1. into tandem walks th~t ,CiUJI,se locsl ~llix~ng of.flV inversion arid. fml11givilfDg IlliLcl.WO every\I!";,~lIere. Mi~~i~,lgimplli~ tll,eIl onlYquam~ly ~,cleOl's .11'1 equal ~~OlIl1:LS. Phm.ctary 1:O~~il]:m1 prov!fics a vast reserveir carried with t~c now. suehas a ehemlcal tracer. wi]] e've:nU'l.:1ullly become ulliiform])' ofrv. wi~h OJ Smllh-~10n:1rn;rndieIllIDl'1'I.eumM:es d~$lrj,bu:tj~d,l1low~ve'". lhi~ ~ontn)d~lc~~ the W;;~'!"'~' p'lItlp!lgol!.ion, PV in\ICrS'i~l~n:;dklws us to d~dliCG ,be V~I~1Ch}' fit;;ld C'~)~5p(i~dirlg 10 ~lb~n.I~d inhibitkl!~ UlhV'lk;':ing ftt:ffJl'-;!l.je'~ 1J<~$. Far example, the: sl1.<II~ospher.icpolar n:ilghl jet disp~ooe~~,enD!!of tllre UIJ[D. (and .rv 'C:Qn~GiLllIfS),.
Slfl~ldf'~cl

:;;hm~.dpoh~Wllin::i by ilLb[)lL~ [CO Sii,lce 19 7\j, (~l. withc:orrt;:spolldi ~lgsh ifts incl imare zones.
i

New data sets and .high-reso~.utioll cO.lnp~uell" simulatioashave revealed .L previously unkumVll SC'I. f mid-ocea:nl jets (M-J I) (sec o

a2)\ The first

Ihe' ru;:me) <1~ we II .1,:> velee i1)1 ex treJIIlleS wirhin the A~mt..'ucti.c Ci~lilnpulBir Curtest

hi fI!~S {'If~~o\'ir Antarcue krill

jets

li~4;l

llin.e circumpolar

~re e:rn,c:rginlg. promising

has di!l~wl''I!t chemicel pW]'M:rtb

(1'11

either

side of'the je:'~ axis, [!tcri::'by con~<l.iI.lDng the 02::0(11;:ho~e: (2). Furthermore, ill the .'J,wndiJrd paradu£n~.momentlum is loeaUy mixed lHu!: ~ chel'li1lilCfIIltracer If D.hs we re fli1e ease, j ets i would ].D:Illllu::diald,y begin to dissipate, O'I.gOl.in eomradletlug llbs,;:~v.Ut(}!l, One reaso 1:1 fQ~:[i:I,e pe rsiste nee of jets iSI]marthere
0

are

W<lJW

mot ~mlS and ~OJii.g-nUlJge,~,10n~{lc~,1 Il3r~digm,.

imienl.ct~oI'lS! l~1;

the standaed

hmreJr <iI'e ,'wt considered ,un It is these I.ong~

are beSI, understood

Irnnge~nterm;:tio:n::SI:h<l,l

\lsi~g fldd

th~}Or}i,

i F~r e,.... nple, th~~'i;:~~cityf~dd cqr!r~ll~1I!1di~lg new irl:~dghb,nto the lif~ cych: aad f(l~1~1n toa :fi'I.rge"Sc:a~e east-west \Vrn,re pattern causes chain role of the :k:rd.~ (/.1. 14). the \""I\T~l\.~lIed" Rm,.'\,by\"'\rwe) 'to pmrmg,H~~ 111en: remain open quesrions with impoil\vest,,\,',nd. The PV eearours therefore behave nnil.l illilp~ications for cl imate p:reJ,:iclion. s(lmcwh~l likce Ills~ic:hands, ~!Ld it i.s~~~ snis I~or ~(l1'1l1pl~"n'n£ riJi(':ci:>c cil"C~umc~l:l1In:restlndelJ caM.edRos~by elasticiryefrhe PV g£<uliem:t dl;Jt which P'V stai K:<!SCS fmlllrellN"lim.s dushoe. QlJ<m~ !\'!'Irp(~rls d~(l ptn~M.<'Lth)n of RO:Scl'by \\,.1\/0,1;, lit<l1!\'<\;; predictions uf d~VOiI~t,lting ~Ofm!\. of Ihe~·mo.lliImll:i,ne .slmtdOV;~n.,UJd oj'd:ils1ocatiol1l,!; :illl The .key~D u nde rstan d~:I,l£l,ets iii l~,'U rv j not l~lQinl]~n;n.l'lm~. to behave li~e:EI tracer, tt!JlltiS eco..~y,stt~I1r11S; nd h.l Earl~'n.·:S ~ hem bal anee <I'b.o lt istherefore rOY d'mti sm,~,.i,ed when the require a mere d,e.'lOlil~lU'l.cll"ll1~indii.l:;ofjeUi, r1~~jd becomes wrbule nt. M ix.i ng W(!'2LkC:~1,,~ the sO!I;J.~'lm-aort h :rv gm,J ieu ts, redue ins the R!~rl!rl;'l1l{~, anol Noms RosshY-~Vi.{ve clasticil.)' i:,uldll;:nc;:o~lir~gi1m.g rm~ 1. ii'he ~t~di!iil'l!.'ffili!fglul :!t illn Amt;ril~n GitIOpllysi[.:l'1
1!,!I~li(In (~Pifflln (~n1lellnQ\, "]e!$ ,~ndi!mnull!i:r ~h'IJthj1e".!.:· he1Jdllm Saval'lllUliI .• Gili. 9 ~ '15 ,Iuuary ~OO6" The1W~~imiJ~ wiH b~ l'utlli~~f\II in ~ ~ptdaJ i~UI!' Illf

lh~r mil'!,i~lg,. This [lQS~lhr4;l fe-l;idmu;:k tends tu igm. k~!;;p:>the: idi;f! uf mixul1lg but combiues h \'I'il~, the lo.m.g~I,'illlgc imermake dmi;! nli~",jllg spatuall)" inhomogeneons, ,Jtc'l:k)J,~~. waves irlivoh.e ]l;;~'l(lrimg r{):~1;;5~ike TI1~!s.-ott(J!1!g mix~!lg createsa pr!i.)~·.il~ lariAU : in nude of,dlernating steep R~.d weak P'V gradit~.ose iIII all elastic Inedi'Uli1I~. iUJ.dllileygu\ 'if: rise w subtle long-rOlllg_e eIJ!.':ctY ....-m,!!I.sid,e the ~ms. "n(;X~a PV sl.ui[cil~. Tllil' s.HIU:case corc scope of In.e stJlld:J[itl p,n'<ildig;l~lI-ill:lcllld~ng
h'lllg-ran!,gl;; ~nlmL;lIU:l,]nllmn~po~[. by w~w("s (F'~di,align stresses I, A~!>o I;liIUO~~ the l1u~lhl(~~i (ln~ets 3:1\l' si!v:!pie f'i:elds sO.IIiI,1.e",vh.n Ihe electrosraeic field ~i~ ofa posltively c~.a:rged atomle nucleus, An .uom~rl its gw~md :m:w~coiu;i Sl:s onh~ nm::~el~$· ilind i,ts cl.ec;:tmstilil:tC ~'ichl ~ogcl.he'lI' with 3.
reS]KHiI.ds(tJ~ro\lg~ PV il1r\leJs,iOl~)to th:iin. fast ~$I.\',r.wdjl;':[s located ~l the latitudes of t)H;;:

The

!1.~"\'If ~1@ rad

quoumlm field des~ri,bed by Sc~lnlec:ling~i:'$. {!'.cunalioll. T,~.eqmmUlmfi.eld repn:~ems: the IiJecnnn n:rbiWih. \vill['(;h are c!.'!rri~d llklng\\,I'~I:h thc::' <ltmnuc ~U(reIJSIi1.o m~mer lmow tl:\i.eIUIC.k:lUS,
~~lf!V~.

a....kmg

$!eepe:strV grad1elll:S aml.strelIilgthened elas[j,d,l.y. F ~"m]it~ml{)re.tl~~l>t!~(!'lg~l~Bn.~J 'e~~ls'\~t·li,ty til.lere:, togelheD' with the jet shear. strtlli1ltdy [uhi bit In ~:xitllg a<:B'OSS the eastwsrd jets (1. j).. Tlm~. the IlII:\V I:)'<umligm e;o;;I1].~in::;: bothjel tQr~ Inali,cm auld ttl:l!::i.llihibi~,ton ~uu:.:.~.ng, of 1"h(;!: :l!u:n,v ptlnJ.digm d~mOIH:iU",I,tes 1-10\'" ,I rot<l~~uDl,go I;lIlel can '(;Ol],,(;en mile p energy ll~d nmm~'nwm il:'itfl J~(& with fit..l€,r rot;)litlJ'i yi:e~d~,:nlgot just (me jet but severoJl or m:IIlY n

Sd. i!;~O' ~"'i!ila~1:,I!;t '~ml:~JA'9i"I~",!IItO" 1lI·,0l{4Mrod't~!lfdlI C:!liI!!f!i!!'L~b.ICbi!,p!noilnG!lIl!~e!le~c:.e'2(106JeIS,Mm , ;;::. tPi'l. M-. tJIr;~ III. E. Mcln'tyre"JIla~l~ ~!!:8,. 51}!!I U9:;W, 1. 'flit ~·All1ii!l PV i~ '~MI'Ijloili'lr!i'il. ,Q'I :lHilu~~ Ul~ ~Oi'Iitlil!Ji' nl)lll'lil~ll(l tlfr~~1<!~ifiQ1~gnil!,lrli!~ ~..wd~o li)y'lI!I~ m~~ p~~ 'Lllillt.:I~·!.'a Mdlo~M ~,~~ Migh_lKI;rl'~'1I~wnl'~.
Wi1lllllh~ ~~rl<K~,mo!l'l!~I1!~ 3df.i!bali:,aLI!t,,'~on~!lI'!'il'LijCQ 'or'il~gl!li!{ !l!IL\Il!l~nl~1]l 'OIl'oli'JiI f;'!IW~Thi!h'ffiJl in~ '~~!11 'ilirtUai. swirl. or ~.rtJj~1hl!\llt ri!lall~ 1.0, '1hl!!plilm!l!t" ~ ~. PI!!' ~!lI\!!lifiiOfl! i~~imil\lrlD f;j~dililg ~!1!~'e!ll<~!l!; Ii~k!dire'tlD ~ ~h~r9~gf ,~~~~ra;:' fl!!l1Ittign qI iMi ~!<!m, ~~ ~

~~~i.· ~

minil'l"lillwtr.ri~L :~ Wlm',a1m.d:MiI!lC~ann,~~,u~M,pi.l'!' 9@n~~tf!olrll ],till U!i~, 6. ill. G, Ji!l!'HH~~LM, E. Md!lt}'re.j, Arlllou;, SQ" il1llpr~~~ ~l~~, Ol~ W!b sitt 1i'1 'J. T, J. D~mb:rliOO" It iK. SuItt.. J; A1rmw, Sc'i,. ~m j)r~~" .mo ill. ~b~ijleirim. e,~. Sh!!l~, )l, S. wy.Dllili:!:, 1'. s. Marro!., ~J!l~r J!l~·@I.I!nIM <!IEI~.;:Ptt!~ri~~!iI fh)oii@l :~!X'iety .mfl'Ui!~ 1[Il.']!I1g. g!~li]e IIJM~lD~I'Jld'Fluid D<fm~mi~, 1['Ho <11~DJji!mbili' 20~ '[~ii'tIP~ IF~M~. liJ.'.i,A, m,. 18<1'1'; 9. ('), fu e1111., Sri'ef,JU' ];!L,~ 117'9 f1'0061. 1D!1 h'i. r.I,~'k:!IlI~ H.IH'il~uml', J. pbY' ,o.MflI1'OBIt. ~~, 4io!l
!1I'IemII~p~tfC'!l'O'ep"""~mes,~f $. i?; 8" Rl1int!'S ilrigiM~d 1Jj1~ '~llrl:lrloolm1!l:r

un~·

~.n

a:o;~m{}\'~~~:sl(jl!i,v~}j i enn~lgh.

ch:;u:ge l'lma~ :110!dS the atom [QgciJ1.CI' is 8Il>cala:r neld ~il.II~d the potenl.i.d \'orti.eity ,~.rVl'. fV is a uUJ~fied ~~lc;)~~IiJI.rc (If 3el~l$i~ ]P(lli;:i~t ~!.I S\,.'~ r:~ or or ~IW~lr' ( illl,chld:i ns p.i:rmel:alJ'rolalion) <md~s coni ed
mld!E:<Ir

The nui.dI~i21k~

w the

{5-7), Thi$. ~c<'liin.§ H~i,;!{]iry ~i~ks ~.~.",. p\l.fI~'!~t':o;

l'<LdilUs,UlS

rOlal.i.oll :1:"<Lle•. m1i.Q,he turbulent t

Idn('l.ic'

t:'!lcrg)"

~c'\ld of

l:l~('t11..1hlul th(: <l.cccl'wd

<lInd sp\,lc,ung 0 filS j e I:S. [Ili E~rUl!i

(JClil~I~~ h,wG:m~ny

t~.~Oi'Y.• IlEai'Iit .~,ilU'i"iOt>p~.u~n; .fui.il~ j uto:.t Of:iJC ur t\,\"ojets peli hemi splia.e;e.\'\~u:.re3s §~I,piteT ,001ld

~al.c \vilh 111 i.s

wi th i'h~ ~:k}\'.v ii;;(: ~~ ch~imic1d twc-er (3). ~.

Sl:IilJ:L~g. clJ~npm;~P V ~~nQm<1I i,t::i.
them reoogmizilI:ble velocity.
~:)re5:\.Ulre 'C;J~,'lfy '!N,il

h,

(~~~ !h~fig~Ip;~, S~ver.d m:w p~.m~tm·y night missiun::;NASA"s Mars

ReC{lJ,m!lnSS<lIlOe

~200'~),

:n. K,]. ~~Il!I~ etal., .r:ie~~,!!Ies. ~r.33, ~(lJ6!!l~

10,1!.\D2mcOOS,G[0l1l6~$ (OlDo.6}

h~.1,o~'<Incy.<lntil
<15

incinding

field~

in

~he ~urlJ:oultdi~.gnuiLljU5t
m'l'litillls

OdJi't'er. the Elllll,-upeami.

Sp;il.Ce Agency's

Ve'mi,lIIs

ililomh:: uw.clci carry electron

(aga~~1

ptO\'.ided fh<ll l1ile D11Lolion lS s~:O\,\l enough). These :stm.ctures are t'lm.e famili!ar Il.mc:ake=like vart~~~$ of C)'CIOfJ'Q(.l$ and alui.c)idone~, from CIl:~in1l~Y cyc.loni.c depressi.ons 100.liurrica,ne K<;!trina~md j.i:!phe(~ Gre;x~R~dSr~~tKm.Ji;Ving the :rv <Illom'<!,lyis elil.ough lO dedut:e the cllJ[~re \Iv rliCX stn~chl.re" j list 8\; knmvi ng.lh(; II,nlC'lr:<rr c~mge i~ enou:yJl:1 w dedUloe the emire ;)IlOl1'lte ori?]I.;;-1I:struct1l,.n:~.1"hG; dcdUGtit~n of the \1Qrl:e:.xstrm::ture i{JrnJ the rVlleld is c<l]~,ed UglV~I:'S~on (4}, The !lew rOro;td~g~l~,CO!l1C'S,

E:,"3)J1e&'>,~ml NASA'S p~fmnedJlIno 11I:1!i&';~:on tD Jupiler-...r.('lre poised lopro·vide detaile"J CI:biSer~ \'<IlioJ,lS ofextmte.lFlJiestl'i<L~ ,;lunoS;,p~el."B. Tlme-se ob~rv.a~~I,)Jms ~r1;;!m.~t1d~d ll[) l~m!dt;]'lw;;t1md.H]·r e:'{amp:~e. §upi.tel"'s Gre.n Red Spot. "!"]li,dl In,l\,tl;1'~ 11$ i~~~'Si~M~t~ to [hI;; $uwrolmdili'n;jtl~::;
mue:! :RQS:;;b~ \:V'oives Its). Altho~lgh the l1ew ~mdi.gm helps, '~,,; under!it'<uni the mys~.ery of jet rO:nfiaiiO\~, al1d je l-roe~,nt:d phenmm:n'l

'l2.C: ,fill, H~~~, NI,/<, ~}!J!e~~oQ, pe~~~i W!R1!l!1~!l[><lIli~ :U. 1:1.. A. f~d1'.l.Ill, Klird. £I~>iSg.Il'~ .Aut IO<t@M9'. Rfl. tap. !i!~. 9187(2006), :Ji,~. B!, A. radiI ~. IE.. H(lIfma nfl" E.], Mw:pny. Df'fp-Sro fIA ~. JOC".etlm1gf"Jilet, PQp., 5~, un 1( '~20061

rv

persh;t~fiJce, the crm::rgi.l:lg detail:;; of m."ny in IHiture rem~n:1.l10 be nJlIiyeNp.~o~d, &I!:~h's 1U!;~lf) spht~ric j ~'l'S h~.'\l'~ VOL31S SCI~NC:!

U. P. 8;, :Rllin~, J. ftJJld Mrtll. 69'. 34S; ~1'7:S). U. 1M.~ei!p!1le~ I. ~D!A,,], WQdM. "'~r!~fe4]18".193 '(2005). :~7. ~'~iltJa nJ.i.,M.IIlI~.1i, y, H~~Sh~ B, liIams, 1': IiI.1"i\a,jMle$, 9:, J. 1iID~lkiru" \Ii, A ~:biMQII, n.W,!. fnMl:p!<1l!il1 G, VllIlil. ~nd:S.. Yoden!iI di~~~~rnn~ "Bill Ni$F aM ~fiQl' !umd:fng Ililll! cru,pmn ((J;1iI~!fl!,moe, 'Il,P.B, iMlS sUI'I~'lifE!d bf N~f~ ICLim.ale~nami(J; PrclQfilm (umdel ~lh~ Ill.S" 1(1], V:M Pooogr,~), ~OM'~ Olfi(lt'd Glo!1aI,~t.a!ll~, NlJI!5.,I\'~ ~IlPllOlilll1~ ~~iMr,h JIIld ll1dlnol.oglj' ~ram l[iDF 'riieo5,[p."e St;!E'fK;E'$. ~,~ Liirol!ll wilh a ~'ti!r ~rogr.i!m. ,<Inc H~5I!'$O~alns.. [~. and miill~~ plr~raiill,

2:6 JArN'UARY 2007

w'l,iVVl,!'.sclen(;;'emaJg,org~

IPERSPECTIVES

Pfi r"l.iltioni ng or ~!!lntr'Os.orIl,e~ du ri n ~ dh,ij S~OI1(lil

DfQs(lphiia glenll"!llinre stem tells mary !"Ie'l!), 'keep ,Dille daughter as ;a :s:lil1m (:en and rIl,01H!!lhe other l<lWa rd dliffe;r~l1IliaMo\l'I,

Each, time ~~ iit'~n~ll;,lell d,ividll:~il can d]vi,dc io.ldefinilel.y-il risks d.n.n age from errors i.nl till;: dllpl iC'L\~nlli.. .d .se~,·~gt'!:titmof g~I~~ic ;r~t m cenlll~r m<Ller~,,1 dun couM stent it.'! \';n~lity or propel it [mv<ucl a eanecrous state. Norm<l:U~_ em::ll d~\I':ision mUl>'l be asymmetric to ensure tl:!iIillOJ)ly (]~l€=d'LlIgll[~f t:~H difr~~!'1!~ial!""s. \-,!hile the' olm.er becomes a :slel,n ceM. ~fum:s renewing the stern cell popuhu ian. \el s.~.om. cells Sill rely ~'\,v 1.ImL.d divide F]:1.Ulr)! 'ture [irnr.:~ lhml other n
ee U type&. ~nc.~lId~ngIJ~.eUr' ,dl~lIghte['K, On OW.Im. [l."ge 518 of I.h,~s~s.slqe. Y<ll'nm;h~t~et (1.1. U)
C}HIIl'!l~"l'ne theilU!c ofo~c

·~,I;l

sues laee adiffieult

~~~~l C"".'.~.~~ ~l~~I.'iiil il'1iln~.~~m~ .th~J!r ds'.. .

challenge.

Y~i~~a."hil~!r.::f ()l. tn(ik f.dv~i!jlnO;lg~uf CG~Jt'u~ , semal ,Isy:unmltlrn:y to foUow the' fatesoflllollil.er ilil1dd.uug~h~r~~lTII:tro~mme:i d~u'ing germline stem cell d iv ision in the test is of .Dr(,)so[Jh~:/~~'" The~gt:rmlim: !ilten~r oli:~]~.\\41~cl~ givot;l rlse to sperm, have already d~\fided 12 {)["lllO,])e times \\.'~(:~n~.ey become c"mbli~lu:ld iml theirniche, I
;J(]jacelll

to SIIUIII.l,;1I,ce)r~iSkn,10wn as the limb qsee

d~~l1.g1i~l. n~~di~ slcm 0~nS Ctl!llpl(ll~ as many <liS3,0 tld,d,itiolU'll cell cycles over the of the auirn ..l.eaen rlme £m,tiilining theI1llh~C:I.\'es while procb::i~]l::!. ol1e IIlOl'H.1enlii ~.l1i:U Will~hter. thegon~ !I~b~ Each gOo:lmj~.lbl~t!ltd~.vudesj H~1. lI~t $;ilXt im!:s before dirfferellti]<I~illg,

me

onh~ mOiitflill.ld'lImcll~

tal ~~~I cu.!il'D.po!ll~~t:.>i ~U!.ppufti n,gf.~m ~IUlI<!i:n

eell nll:l(.'l~omL~du:.~Ili~rooorn,e. t Cenr.ro::,'Omes. ()rya~i~ ~heMic~)u.d'll]iil;\..;mlch mitotic spindle By indlH.::i1mg ~he b-XPOO5:5io:un. ~:~is ~llVtlre$C~In of (~al di~lect:; howchromescmes m:td other m3lit!protean lit different t1i.mI,le$.~Iln.eyeould selec:ri..~lsiJ!l!l:: d~)o;u:ibl!l!ed herween &!Ug[lt-e;H,"e~ hi t:L~ cell di\lisiOJ.1 (Mliwsis)l, The ,mthOl::S shew tl!o,,~~ male g~rl'l1~in~ stem 0~~U:S~!1 t~lu;:~ fruit 11)' DnJsopflUa ",~,rl{I'rmgl,.~'M!:rII UrreiL'emi,a~~,yP'Os i~ [Xl'l"',their m,t'I~he~ ~!ld d~~u~ht~!"I~1'ntrO~tl!'I1:!.~~ ( dmin.g Initosis. As.p.'1I11 ef'this sl:rateg_y. \,\;IJ1id~, ensures asymmetric d.uvi,!;,tonl, l~u,: stern (,e.11
[JeTm~l:Lemly retaiusthe ml(J~~ercentrosome across :nm2my cell d i\'isiollfl.., r.lI.is.i'~gtj;lC possibl I~

The ~lUt~~ol1,';gcnclk~Hy C!mgim~~rcd flies produce tL ce:l'~tnIM;unf•. p:D'l)tei~. k~w\~ ;IS 1 .PACT, lagged ,vith gre~J,llhlOll\e~celll~rote~~,
l[I

IhL;Imt~.!i im.:'I;~'fu~i:;.rnU~Ufud,Hl:I~l~d'ie [~R'll~~ii specificarien ofthei,r d3ugh~el:"S ,PllBecaus.e one C~'U!Fo;..;()m~l\'l~lint-:,i~hUlbpm.... ~m~~ '\,\;~.il.~ nile other 1H0'II't!S to' the Qppos~,I.e side Qifthe ce]I~, '!i..~!I;!nibll}'of.\ ~ULt.o~~C :l-p:indle J 5 ~lh"(IY~ perpea{ti;cular to Ihe hub. As <1. 'result ,dler eell dlvisian, one oll"lhe d~ughtcr cells remflim. a~$Ock'u~d ~~',~lh hub and receises :sigl1lais.ooll the hub ~R~ Ir) remaara st~n~cG']tMc,n~wMt;;.~h~ h luh. di,'>lal d3ughter ce I~11IlI,Q\~ 3W,,~y flail the niche mni in itlimes goni albb1!1td~.:nerenti3nio.n.. '])m.ashit a et a). show ~~'lAdU:~Ulb·pro:Unm~ mOdlil:l1i I..lclmtl·OSiOl,~~e ~s assoeisted ~vic:~, 1"Oh'U st mk:~'ombules dur~li:g imerph ..se (the IloIiK.lividir'i.!;l r !;i:a!k"C~ I'lml i.h:U 'J;om~ of these ~l~,~cro~ub'I~,t::S.
l:.l."Itl$D'J.di
t!.)

the ii!dh~~n:s;jU!lC~'~!1.)!,l~lhf~~I;.'Jl~l~t'Cl
the prnper o:L'il.ellt'llio~, t)f stem

Ibe stern cell liO hub cel~s".These mk.IfQl;~Llmlt"S m~'I"ifm~.ytedler d~e!ID'I_j~hrilr('~m~)!5l)n'!elth:tl l'll
hub.
t!ll.S'LlJt1Dlg

ThIs d~ugl-II~ r ~1em ,dl

l1!ni;;iim~imthe nkh ~

ily lha~; difrere'lU.li .. , oen:~~'Ostmm,~IJh~:I~n!lm;eis d i

stem, cell biology. Ulflli:ke mile, 1.:!m(lo\!!!!l ~nillll,111 eel I fl[.1¥~nelle..~_ the: two centrosoraes iml,heJJ".it,ed d<lILg[lIler by ceU:-; .u divi:.;i(ln.!ro not idl;l!'l~k~!I. AUimHrm~1 ee Us illfritli.a~~,y have one cenrroserne, eomprising, ~ mother ~nd d!aughwr eentrlelc I;'I.S w~11 .I.S peri'cer~lr~,ol<lrll'l~,Jiteri.l. The mother eeaU'iolc: OOI;ll.aimil!O S;lrI!:lCUJI'C'!i and I~roldn~.h~ll [ <Ire absellit frmn the d"1I:~tgl:t~er eentriole, and i~; ~~l~\;':~t~ w:),p~ !niCf'{1!t~jhul(;)$ than thl,;d~ugh~ ter, DlJ.ril~g eaeh cell di\'i.sim~ cyd~.lh~ !i,;~mm1;OIne ,rep~iC<IItcs,.The mother Ge(~,trosome' row ins~he or,ilg il~al ~:uo~hercentdo Ie.. In COIrI(1:<ISl.~l~.c i!flillgiller l1t!nnnsmm:: m:uh:::rgoes mat~ c rllll;aui il dur~llg III iwsi:ii ~.Ild d~!ring the G I O ph<l5e ol'~'henex.~ celli divisioll,l c~de. COll"~I.t= iu~ hi> inhcri'~J;ldd1l'llgih~~f cil;o'11~ritll~i~'[u a n~1.'V ll,nmi:l.e'r centriole j2), Wbethl::f this intriln:S.ic OlIs),rnl!'!'1it;;u')' l~cM.iwtes m;ym~'illeuic Sltl!n I;.ldl
d~\;'isiQl.l :Im.,.s. rell'l:<l~,noo,a myi>ll::'ry.

Ihis daug~!~r ~ipm 'QP il II;\;I\~~

[l;i!

esseutialm

nit.h~ to Il«omll!

~P~1I1l

GcmialJl)laSll

IC~!il1Jo:5@meillillhFl:ritill!i!CIil ~Hili d~lI! glli1lt~fl;ellfii!te. D !.!Iri!l'lll~fhe r:U'I,I'izi1;1111 ~! ol g!1Hm~inil s~m '~e'llitrllh(l' f)fQfj,(JpMla b~)tj:s, the mm.he-r oe!l1trm.QI.f'lt~ rrrernaim .adjaamt to thl' h!.ib ..lhe 9 reill~.r mkroli.lbl.i le Ii1l1Jxle.ating (ClII'la:dly or ~ttil' !fmILlicr (enli'OS(Ul\i~~rQwltil·f :SUi bi'lilie$ ·~IS o1lsstldolliolfi witt-. the ad t-.~'CnsjUridiO!fl,f~ ci li~tin9 sl~m (e~lllJI,u:ti'(iIn.

lively label ,c.~lherD].1.01hef or d,nlg.t1u:~r centro;l.n!rIJ~c~+ Motb~r ~nW:n~nmt,;;s. were el mt'l$t ,d~\"~}':ii 1(J(:<.\Wid near the hub ..\'\i~.i~h~~~:mr~d thllt aHer Initosl1; they wou Id be .unhedted b~ the d'lughwr du~t rem~im;inlhe niche ,Hlld l'emil.llm.S!I stem cd,~, D!i:lug~]t,crcnlrOSOme5i. 0 onth:e o.thcr haud, i!h¥,ays n:ui,grnt'ed to 'the
OPPosLle end oft~.e sle.l.H (.."CU'lnd\\'e!reullher~ ~!.(-dhy thl;'; do.'ll~ghl,,;r 'c(tll deslim::d to ooc",()nwfl gOlli.lI]b~ast Thus. g,e'rmline s~em ceUs. ~et;lill d~l;';]r mother ~nw~~oml~e rimn ~he t iiil~~they fI,r~l emer tThi,e.ur I':Iliche. \VlmL!ld~ll:lm!Bt: mi31rni sud) a ~tmtegy aml~ on the stem ceU"!TI~e ~,~lo~lli'k~'ly,ms\JL;eri:; LO [,Clp QOl'1II101 the orUcma.ln.QIlJ of,cdl divi~i:oo$. GCHulline stem oell~:s. un the Dro,s,()pltila tlesti:::;

cell d ~~dsilol'laswell as Ib.e stem cell-specific il)heU'~i~l~l,,~ d~~m{1!t~W:1: of ~~11[!l]'-~](~~.e" S I:~die(oi f ether ~.s)l~~.lm~l;ri.caUydi vidi I,g o (;'CII~ sug;gesl !)Qssibl.e additkm'Elli rQles fOF'!:lI]l)gr.mHlrl,ed c'emroS(Jf~lr;: inh~riwlnoe iUlr !'item (,'CII~. Asf:dcWlOll1I l~~d pmt~dpa~ iI~ ndlc r iflJl sill ~~ssemb:ly. centroson~e~, a:ssoci~te\'",i~lm. melllbl"dn.e=oomldi org~nJieI~essuch ~:s lhe Gol,gi i1Ind re(:"y~'1 l,;m!d.(J$om~s,C~Ill!nS4Ylli!~\~ 11~&Cl re~tging l<lite cJ'rtoki I.le£ is, by d.e'l:ilvering memhll<UleS ~l!,I:ymmi,;;uic~~lly the cl~<I\l;llSil;; :Fl.[;R)\!,' (4. j~. to
Ti:i~ls. differen:t

i<d cenl.ros.ome

ilnlffl.erit3.1lce

'The ,~u't~~or$. In the [l.!!I~Mm.e-,nt!!If ErmbrYOlog~ and are' !iowa rrd I'ii~gln'esMlfdkalinstiilll;te, (anneglie In 5.1.it~Jlioo, ,

re.·

might oo!t]tl'ib~lte In siC'," cdl mn.iilmtc:~~~ncc ::Ind d~iughte,r cel:~ p[QgmmL~]il:lg by p,Jni~~,o]ilil,lg
membrflllc-hot.!l,nd {I,l"ganclks f~l:Idsignal-

3'5i20 ~nMan1rn

Drilr,e, BatUlIlore, M[I 2121.:8, USA"

E·D'I1al~;,'prardllTIlg,@OW!l!;mi;l_ed::1I

PfK'Oi~f:onlh~f[' mi~olic sp:h~(.Ues. ,n ri!h1t

SCIIi!:.NCI!! VOl.. 315

(lll.gi.;;'\'l TO

ing ,~l,~o~.ecul!cs asymmel;ric<li~ly between the ~nllUI1:e 5'l(;)~1~, (JJld gonia.lb~flsl_~~deed ce~[
2007

~.:soienc;ema9.ar9

26 JANUAIRY

PERSPECTIVES
iI!.sy!l'lI~llelric l>egr(:go!!li.Q~. (:~ni:'''l~ dG'~t:rll'!iof nants t~mcmg~.lmcyd,irng: endoscrneslaas been imp~.i.a!u.edtn spec~~ring ce]1 fate .ilil the deve]aJ)~r1S Dm sophik~ nervous sy:.;ulrn {51. The mether eemreseme might ;,:I·IIs.oactas 3 b~:smi body t!h.n uucleates ~upr~nmry ciliu~:n, tH ~ea.<;~
in n~!~u~~mfulli(!.!ll tl;L'Clli11 cells (6). P.ri'~~.HIr.ydUa Cfu!1Sc;:rV~as "ig!11!~imi.gn~g.~I!lI;;~I$$ 1(7), ~nd II.night provide a meens for stem cells tOOOmJmHtl!ic.~~ \"d:(h~heir~,ii;;l.1e .mld ~nreceive .~

the u'l~other unilLerite~.by the bud (10). By i:~heriting mother centresomes, stem cells m"y ~.i~ewie use rl.1e mllust s lllicrol~lb~.lc array to. ropel molecules that pmmate rep IiC'<Il'i seneseesee. Hewever; in Vel
dH~]rn"~(l~hcr(:~nlr!)~mc.

part of

s«)in,dle pole body is selecdvety

It)' d';)I~~~i~ S!J~~t-eg}~ m ig~rt.;i,~!;~)prfum)l~ S~JI~~ ; (;"Clh·,:emam a ble s;nabil:i.ty amllongevily.

RHerel1l1[JBS.
1. Y. M. ¥arnasi'litol\ A.IP..MlIIOi'iald,~. It Pulim, Mi. T. 1~IIU~, 5deflU' :3;:1,5, 50UI {<lOCI!!'). 2_ M, DIl)la~tr~. ~ ~; J. (~J ~j:, 1.11, ~!61:9(2(D~.). IP, l- 'I: 1Y.i- ~~hit;!\ O. ~-_]~n~s,~_ T. fwll~~ ~r:i~e 3il1lt,

.W~~';1. QI~'n(:r~s t
aoi:CLllffilm~<I~e

01·da!li1J~g·;.;. such as e~rf)mp~ M)mL; ~~k~ that (:,n~~ lost>. (if h~h;!'t'i~y~{);;,ity.

tit<: in1[p(ll'l~nr;;t;;Q r matern ,II ~'li;lnwps·{~ne'~flh~t~J<II~oi:ein ~~Ji\Cllnmi ng

pn::~eIL'mli.J.lII.y

in the bud

un:so

maiutenanee sigJ~i:'I.l.

:~onge\lj,lYOOlll11lilirlS lHlC.ie1l[:

Could
contribme

w[el'lli(lH

of I~it: mother

l,.'1;;mnlti(Jil~1e

lo. 51e1.11 cell lo~~gev~lyas "!''eHas. to d,"vd.op.IL~.enl@~ pm~jt:iilnllniD1,g? most relcTime

di fre.n::11t.~.. i cel'l!tmS{ll~leinhl:'fi~ilUlI,{.'e the lOrlg\·s(Jl!I.y.h:~ .secl,'Ct of stem l'i;:.~I. flJlDlction'! It

] it>

vmn' iulOrumti.oJ.l comes ftUU11iI studies of asYJl1l~ met ric ceU d~\I~!i:i(ln in the huddi.!1Ig y1;~lsl !'jucdramnij ces a>nn ·isiae. IVI.othercells hjy~ <I. ftlwe:r rq}I~C'<III.iv.e1Xl1enl:uilIl than buds they pmduee .• md ~e.~li!ic:~.ivtdyret;"in. dm1l<igtld moleCll~CS(It 9). ~nh~IJt"Mling:~y. fling!} I ei)lJ~tcr~ the'

be PtlS;5i'ble maternal centresomesate shouMmlw

to d~[C]1niUlC \:vhG'llm~r

reta ined ll~'several

oth~r \",~]~·chllr,rl('i1:eri1.:6i1 tmapMla stsm (l~U D s,

~:I1l. male ge1111~i,ilestem cells. such lleh!Bi\fior seems lil.::c~y to !lIOnlr.~bulcto lite smblc!lsYUlllrIf!t:t1.l" prq~r"'.Jnt11lLllg,gh!em Coell,mJ d,mghter. A:mI.d,~l.is 1!:tLtisf),i mig to cOI1lC:m[pl'lliC the po&.,i.ld.~

l:)~1 f20~l\ ~. A. GlOO1I~ N al~ ('tll1j!:~, ?S, ~2'OO<t). 5. G. frne'l)'·f!~t. (~!'I':!!~z. 763 UOO~)'. 6. 1i.!Il'I. F'.reb~~.l "". l'iiidilil!~l5,:5. IK. ilu'cme'J; ,Cr.!ff; ,Dw. 8;ol_~/~, ~~ll~()OOl. :;;. v. SllmgLa_,1. f. ~~II.e~, titJ'j~ !i3, ~9' (;!OOJ'i). .. S 8. H.Il{J~i~ ~ilJ\.L. G~~ilf~~o~~M.. IRig c.Jlet 1. M!~tf.i5rn. Sdena! 299'. 17~1 GlOO3~; I~1!l~1leC 1 on'lim~,7 f~!bruOl!f)' 200:~ fllJU12~:5!:iet!!lJ~.lalB0l4liS}, 9- [(.]. Ui\tem;~D. M. (lIM;:r, !l. A>5.l~da.ir. M1{rWiQ1_ ~iW> Hii'Jl. ~~ 6'7", U6 QOO3). ]JO. ·Y. IIt!reifJ. 1. ur~r!.iIk~. If(. frldS!1l)'ttl. t. Sd'![tbe-L £MOO 1. 2;0\~m9 (~,D. 1!1. 101. . ~1J!Ji.)':D. A IE.GI!!~t5I;l1i1in!!, S6M!:l' lO:!!. 1008

r.

~'QO,~.

CHEIMISTR.V

A.~lngle

Woller

m ote{ule

COlIiiad

as a

Glt1lll~:tiln

a.

9J!'Is·phalse reaction 'b~r"Otrm~llg a (om p,ll1'J( \vithi[! r~J[tant tha.t reacts filriter than th-e "bare" Ire.a.i:t:ant

processes. FO.r e~.21mp.lle. the Haber pl,noess. discol'llcreJ about 1.00 y·ear:s.ago, Il[mke:s!~.seof :mpp~ln~d iron ,c,mlly~t5 l!!J speed up the oOI1,versi.on of·nlutrogcmiI and

o some, the\:l,liOfd~c.m~lysi!> COll!jm1es.

of

OZ.O:lTlf'

pl.eSieIll.l ill.1t~.le ~tnUlOsp'1rneil.'C.

up i.m~gB~~fim·~~-~:~Ih~i nd~~~~ .. d.~l.~

hy"'~rO'.;;.en t·o ;HL~Lntmi<~,u :mmdc~:<l~~lempeE<l~

ttures. ~\i'mJ dVl;',~de. this 'P1.'OOes'''l is. lliespom;.ib! e fil r :the ;;'18m manu f~t\1 re oi" nmf~~l~;)lnl I'm lLfd mi~ Ii 011lHf!tll"i.c t:O~:lS. 0 f ammonia (.I). llll cO.mi.![it!~.the paper by v.[i~~ri I'Igt'f- MOII1 i~~l')!f,N lli.
on p<l,g,e4(}7 of Ihis issue (2) 1l.cJ:d'rew::es III li.m.~ dam~'mtnd qucsl.iOl1J: Ol1lii s:~ngl~ molcci,lles serveas c:Jtll~Y sts:? Thati s, C.<IIli ind i\'i duall IlI'Il()~¢(:utc::> accd{:Qub.;: c;h~jmlleal reactions'~ TIl is {~nesl:iO:lrnis best answered bi' esperim~~t$ i~l g~srh~;!;iii. \V¥~~rothe: ovcr"U !h~

111 d~[nnetitl~ of O<'.Jil!lile;: ~!!~. be ~11:0ll,ght is: of O1S· Ill. ex.,mp.~e ofhomogelrn.eQu s c,nalysis <I (the catalyst is in the same p:hase <IS the reactmusJ. fRm,d~<!r in reactions iu :-Q](l'tiorl!. By coatrast, V(n~ri.l1.gcr~M.rurlill~zet ~~l, rovide p an e......mp]~c of gas-phase cm.tlysi5 more akin rcheterogeneous ct'lt.,.lysis. 'fhey rep0l.11'lm.u the reacrien lWI\\'4;lC!l h~idroK)'1 ro(liiclIh @iJ1d
(M.al~y~i~ ~]ng~El!w~jtili~ by

aeetalrlehyde, Ofl -+ CH~CHO ~ U;p -+ C~~)O. is aecelerared by lhl.'; p!.1rHdpa~io!'·pf si I.1g~.e melee ules .0 r water, TIley <u£ue eonvi.l1.dngly thmt this us because hydrogenbonded c!JInp.~e es of CH]CHO and lit, .... form aad dun these ('Q.11I1IplcKCS react raster with Girl: rat! ical s 'dmlil do i.:LU~i!{i.dmll ~1IiI.ol,ecules of ClilCHO. The b~lldillgrewater is IfuI.u~"ll.u~og()liii.Slol'li!ldi I"lg 10 ,-1 sou rface i~l

IHulI,lec:l!IILes. hen;jllj

P;("([I-

i~ ~eslor (A} r~a{lf

~Ji'lieoe,i'I hyd roxyl

01115be-

and ~bare" a:(e'~.al.d:Jlyde e mo l,~(ul~e:'i! ,(the "hi igher

r()~d'"

r:~ld iCi3Jl~

t flU
i.~

reaction
reactions,

C(lll:l.~.prbil:~ serie s of t:d~m~!H.~ry a ,eil.ell iInfol'\! [ng ~ snu I] I1itu~.lb:1::r

m(l~ecule~,. O~H::wen-krHlI\\fn eXll~ii.]ple is ~he caullyt Ie dc'5~rUI.clion of ozone inl.the uppe.r

or

~I!:~rllilill,UQ!1l:> !;uti.we:~n ftydr,o:l:.yl. ra:dicil~5: an.d

ace ~a!dehyde m~ letli le1!:

lr.acoo ilrl b1lu e) and

I
I"'"
I
~

aunusp:lmere by specieli: such mJ. h;d/og>e:~. HtmllS. llillL'ic o:l!ilil.e.,md :~;yd:roxy,~ r<Jt1li.c<l~s. ~3}.lhese t $peci~s 0'0 p!lnidp.,"IJ[1;) ill C!hairl ro;)lC'~~O~l~ mi!liim~1 .a[o!il!!D 'Ih retl~· (X + 0.3 ~);)O +O~:<IndXO+ 040X+02). ~~Qn P<l~:n Qf mi!1l~ mum \!i.'h~)~ m;~t::nilic! a s ~o lili;,l~d lUprh·~·C(l~1Vtll'$~nn eil E!f'9Y. vlillh e iile~gy ~m· of ·'odd oxygen" (0. awms ilnl.d0l molocu~el>l !I:!l"ei'l(e-s 91~n, 91'1 ~Jmo["I, ~o d io:-:yg(:n 0l' thtl.weby I()~\'(:.ring '~J'nc !In~lOul'Il Th~ ~rrow~ ,ar@ lIs,(!;d 'to t'! iindkilil,e that WI:I'Ipathway B all.l~he. r~OIclive llu~ ~hal f&:l (;h~5 Ine rni fIlim.!IIm .a~:rod<liwd wilh lhl! prerci&'! Clio,n (!Jmpl~'" ~!iS!l'~ Ihro:U911i llil@~nrq!l'r [rlil.l'ls<illi 0011 S!tal~ ~od b~oom~ pf(J~IU CIS. WiUH,Qll$ em IP~t~W.lY A S@!IJlC or til i,~ ~ lIhe authlllf i~.n Ut~ t!wive~ily (~emkal ~iiboril.tlllff, i ~ r rarnsiticm s~ale, rooud!ll!Q the Nl'<ntion ral!.lhe ca r~oo:ns show lihe SII1UI::h~lrns, at ~ len~fI eM IR~i!!dl,CJlrrd)~'i CB2 1.EW,UK. IE.~!Thin:li.w:m. ~~UXi;5nrlwJoo <Illhe in !'I~r .~ dg~
:~'inml@bh~ii!-ac~k.

~Ihal arenUJ{ iilted 'ml.h sinJ9!e mcdec:ules or via I.er ttheul()w~~ ro~d ~ !r ac~d ~ngreQtn~" ilil~ i:I,i)riz(ml~~ li!1l~~ ~h'IlW m.[[ocim!31 .and

tnt' maxima andl mini I'm,un mO~l (a~es 0,1hyilm,gerHbo.rtded ((lmpl~e~~. ore dell&ils CIre 9iven in {2L M

2:6 JArN'UARY 2007

VOL31S

SCI~NC:!

w!,iVVl,!'.sclen(;;'emaJg.org~

IPERSPECTIVES
he[~ro,geU'l.0~)l~$ C;,'!HI.I)'t;.u~.,

m •. u: use d gas. r1IJl~xtnlres c:oo~led to seveeel d.iITercnl ternperarures, <IS ]ow as 581<., by
eXPlln~i'on[hrollgl~ .• t eenvergenr-

or

Thelr e:"'.perimei:ill$, dive:rg;enl

po~sil,);ic to p~n a figt~m en the extent of the acce]el,'<Ilkm or~1~.e reactinacaused by rhe single water m,oh:cu]c illctil~lg as
:'I

CJt<l&ysL To

eonvert dle~;~,e<I~urud P~le!110fl1enok.gic<l.l tr.u,e

eoastsnrs for reaetlon i:mto rn.w con:st~lIlts foil" rem:a~J(I:L~ ufOI.i with CHFHO·,·&iiO. one

"Lmml"11O"2~;C ~4).lllfi'ormil'1llimlJ about the rate ef re llction is thelli obnl ined bY1l B~~md.nd niM~;[h~d:OU 11I(liic~ls~rc gl,;m;;rw,(:d hy p~tli>~d 1<l$Gf pholu]Y:5i:;; of hydr't~g~n pero.l,(id~. ~~mJ ~.Ill,e of the ir reactiveless is :m.e~lslUredwith rate

a('{'(t@ldebyd:'iil(6). dt1'Cnrlh~~-d by th0lrretlc[ton \\,'Dt~,OHmidicals,m.'C Si:lOl.il. so annospheric ~l.oeHII.dehyde is fOLIUl.d chiefly at low art iwdes wilecre rhe .~ven\ge ~empeli!u!ru ~s ;,u-mmd 288 K~E\"eUilm:1]{lOOAlrel~lhtC lumm.~d~[y.the (lqu~~ib-

'~~.~u~

(l,f~]

~~Qtllld

l~l~r!l~Ol,t ir!d!~c(;l:-;he OH r

radicals toIlnoresce.Tke decrease :il1 the fluorescence ~:ig~~1 ,l::i 'I~e tinllie: de;~ay between ~he two laser p ul ses is ,i ncreased provides
inf'o:rlllll,~t~01m(lOOU~the reaction nu~.,. dd~n,!il A ~l.I'atell'to. the ex Ilauded] g<ls increases the rate of]oss ofOH. ~$pc:cially at tht: hl'lt>'G.s~ ~I,,;m~r;.

amres of:the experimerss .. The imerl~IT:~:!lli on of ~'~.cex~~erime]llt~dun the reaction OO'l;""If!(;l1liL lr'<'I;di~']~)1md OI-l! .• hydrogcn.lbandcd eemplexes o:l'aee'tlll.ld.ell.lyd€ ,,,'i,~~ is f~i.SLedbn 'th.u bet!i,v,een OH~ldi"'41li!r cal to .l'UlJ "bsre' ~.ce,t!:l.1 dlYclc: n:]~')~!c:c d ules=-is

~~"ed~H) ~nnw!l~.c I;.lm1C€fH~l,tion of the COlnp~~~~ preseruin l~ ,exj:'l~rim~mts" Two dj,fficuhies arise. F~,~I. ii,l ,LS,U1.ol easy to measure these {,;~ll~'~n~mlioP.1l>. e;\"~lp]J;; byil1ft<!]'I;d ~Qr spectrosc opy,. inl'~.e envi ronrnent of l he jet" Se:cofl('_~ il: is n()1~"'.'ii:yto esrim .. te the cencen~nlllo[[s. especially asthese m~y depend O!~ the kinerics of~li1Ie CF~~C~n7'H~O assoeimlon, not just ~he~h:eu,m)(lyn~.l1m,lic:s of[his process, ]1!lK':oos~ing ~!Sthese roSli~H 'l~.an hnportam question is. whether they have <my releVOlI.I1CC0 mmosphc.ric 1 cl'lemismy., Aldehyde;!> urise inlk Iropo~phere cl~lririg Ile ,a!I,ill0spl1eric OIX iWilion of hyd:rOl'arib01ls.: CI.~ru,lC m!.i.dlize,,~ 'l~) al;el.I~.debyde. The I ife'lilnies of I,jolh, ethane and

ri!U~l fraction of ~lCel<d()l:ehydecomple •xedwith w~n(:ris:~ikdy TI) be vcry small. al'nd 'IJ'n("re~ili\: tl~.e,,(;IC~ler.!ti~lil~lf the 0.]-[ -'- C}f~CHOreaelion will. 1:l!IIObably be mi:mim31.
R:~eirelil(/Ii!'~

1L l;tU'OO~

2. ~. 'l.
5,6,

IE, l!D.tkll'i5kr., P:.l (Mill, lIlJ!tI'IN'U.il7, ~,2'1 ~2('ia~). E. '!iij.M~g~-IIt1r1ii'l~ f~ oJ•• StfML(! ~:t~., (~On 'l97 til, P.Vi\all'll~, OIemi;ltj}' fljAtfflrJ.Jphffa ,((};d~ Urn..,., Pr&. OJ:JG;rd, i!dL ~, 200II));. p, 164., UV. M. S!11i~~,B.1t. RI)I'!e. At{, Chem.!\le's, l3 •. 261 ~'(1!OO),. E:. E. G!~I1"'~l(l, s. '<II, M\lI1Urn, '1':, 'Ci~r'!liM"i~~_]. t('I.!,\!'Ietl~~in, Phyt, C~ ..~ :;!'O~'. ~Jl t2~OSl /. A, R. Ratii~hW:~lFa, 1E.lR_lm~oY,f_ ChMt 5Qc" fi:lfadOy rronJ. ~Ot, 21MI~1l9~~).

'p.ilthv...ty rrnn~~, 'e<~.ctiU~iIS 'p;'OO.~ . 1 10 isno (:orufiglliffilio,fil ~h,n the reactioa must pmss through th.u has <l hi~Jn;rw energy 1$lJjlIl th~lon:~lC .illi.daUys(:p~"";lI("d~"aC~OIIrn.tS. 8m there is <1 ··sub:lrn~.ersedharrier" ow "ro~C' A s f~:lram! ItC~~.!\~ ml!lllw!' of fas.\ ~~.~ i menl:HliY reaenons, l:iile kIeas ,or 11<11m.:Lt.Lon sl<lte tll.ear)' must be applied rntl~er.subtly (3). Tlsere ~u~, in dfi::.. . lWP mm~iii{lfb ~~lIi.e$: ml"oUlwr" i onelhal leads unto a sh<ll:i.mv ",'(;11t)]'Q [h~ energy pmh~"(!L)~ CGwrl:ll]xmdi Ilig to ~ preR1i_IClkm~,{Jm· :( p!c~ll. [ol!O\\;ed by an "inner" If'<I~s.~lDalli tate s w~.~Il1I,"1l1Ie~1t;'fgyfl\'Ii<e1fthrU'l, Ihnt nn~e S'ePfU11it0d I reactants but higher tlltl.f]Jtllilalof the u:d I ~see th~ fig.uru). Ihci!uncr mUlsi~i{ln Sl,ltc !'IOr'!'CS,IS the' true ·'bol.~lcn,cck" to rcaet ienbceausc the ifu.crmLI q"I.~lnhllm ~mleil ~l d.ri~ (:t1f~~:igur<'l.tiolfl ;u:t· more widely ~~<l~'Cdlh.m,lhO;ii:efor Ihe OlUlcrlrnn~i~i{m 81,1~L:: or. C'!i;]~lil\,l~IICr'itl!l" i~~Pt~~lltS Oil fr(,,"t, c-n.e'l!Ymmdm.~m aTml8lh~ ~1c~ion l)ath\'~y. barrieronthe
nets;

SUrpLJrl'!;-d. by I·:~~t-pr.undpll;s.q~.ultum -dl~mical (;1IIc1Jbd,O'm.~,E\{cn for reactiens with GEO PHIYS~CS u~,'!l"Qmp]exet~(;HjCHO. there il'> no ~ne~d'Y

d~O'L~s.there i

Mo,w,m~nt of Earlh's teclol1ic phtlil:) generra.tes tow-heql.lell.q seismic e\lel'l~. Tihe:se !illients ,i:lr~no,t Ib.H::kg round n ol~., Imt. r,~present s;lowewand '~e!llpere.alrth ~lUakes,
ogy ~md dem.sily of m::~\'\ro:rns used to :L~~oni.wr s.e:~~m:ilci,l;Y aad crustal defonnauon.Eanh S1(;:tel'ltLsts, are d i5{:OVeiri~ a rich ,~pt'Clntn'~1 nff'n,lphlr'1.': phe~lulTIIl;':n,f! d(;~p wutlrnill subduction zones, These zones. are

~tb

impn.wenli.e11tsill

the technol-

~f~tU~\\fi;l5~ Japan (2).. On! pt1,ge 503 of this issue (3J. Ito r;t ,rIJ" show llmal. ETS is also marked b)f the occurreace of very-lowftfqUf:'r.lCY {VLF} earthqaakes, T:~.is~.dicl'syet

~ifilotller pieeetn

lit"" l)luzzle of Cfl~lll;ll:iI1pOri!l)r

l;)"!wreEanh:~

l:ru~,l~

plates onmdto,r~l!~lirlg

Q~I~!~~1;1i1lil.~.che~llic.al CfI]c~~.I~tin!'l'~ ~lf!W~~~1

ill one plaee ~uWi'~g~l adernesth the otllll'!'r 3,1'11[1: C\'C.un.t~19~.iyi"Cf..cnl:cr.ing the lIPI'~r Imultlc_ A recemly disc()veredpheIlOa.Hel,1O\Iil C'1dlecl "~pi f.tl die 1J\~mfl1r ~nd ~Ulf' (El":S), iIliVu l\ftC!:> slrmU crustal :m.o.tli,ons.O\'CLW periods of d'IYS. 10 .."'(~k,,, di.rc;:cdnm!.I',)f!:'n~O~~()!~: i~,dIu.: S<.1i1i!.i[~ if'

deept:1" p]m:e dYIl<J:l,llUt:s. I\~. seen in rhe gri;;,;n Sum;:~lli<!-Afi!d,u'l~al1! e.:ll'Il1~.qll<'lke: of" 2004 (4). the mest <I\!':csm.m: teeIonic ,e~~raeteri~!!:tlc f IIii.c \\I'O:rld's ~~I'Mucli.m1J o zones istlile repeme{loocmrenceofgreal th'tnsl,
©i'Il1~~U:;;ike~_lthlluglii.i11(: A awmg~rG'Cul:'ir,~u;joe

HlP Ibran5 U 1;.'~HpLe.'1.wi~h CH ]:t:HO bm.1wd, by I.7'~6Idrn.ot-n., l.hiruLJgt~[\,\,'0 hydrogen bondis. [m~n~ctioJl rfdti~ cmnpie.'I:'with !In,OH mdil'<Ll o f:orms. a flll'lhcr llj'lU.l"\'lgen bond sOI:imal the enef'~y ofthilS teni.m:y p'ren:mctlUI'I c,ollllple.". ~s. 44.6 kJ mol-I lJe:I,ow f~l'lly sep~;llu::d CI-iJ:lCHO -,-H20 + OH ~u!ld Id mol-! k~\,,'e'rlMn ~h~' 27 energy orCi l)CI :~o· ,.1.17.° 'T' m 1. The p.fiiere;,{c~ ~hlnC{]!IiIlpll;lx can Ih{;;il con'lit~n to pr'oduct~.
::ll[;t0

"e:~~~ p,~ l\'l lKlfoiI\'~Y'ge'~C\f;i ,,1ml .~~ ...

nAil

:intervalls Jell; l~.es.e

dj,n."!:;:t~rGbtl;;d
~[l

cl.e\'m~t<llillig

l:emblors

<Ire

p.~!ltl;t1XI~wefg~n"CG'rou,;;:;.,

th~

uni;lnlh~e

mnl:i!~ orllLe Mn:~"'m~ildlJl.rI~~l 'C~.t~UIIl"'l'e5 i!1!

+ CI~I,p.lhmughl

an <lImosl energeli.t1!lIy

cquiVl!lileni trnllS;itkm 5.1alc(sec tl'le figure). Thl:! combined eXI)e.~ljment<l1J"md Iheoreli~

cal a~~H11;el1lsabo~ll the n:::llnt.~'!i'(;rca(:1i.vJlY

or

aloJ,lg the ooaslal ~mJ~ n , the:s.e llm.'gee :r~mIt :m'piru res is no! \"'~.II.~jnOI\\~IIiI" The sl!i,pepU!l<lllfe.'i re~'~.n im:en'<lis ol'm(lnmh::> Scien:riM5'lgtili:e ihtLl i~ the i~le.)!l:gmMe: <1seu::;j!j to yem:s, ~CC0lll111).allk~d by d~s,tinct :.eismic mk' pllllC mOlinn:s (.i.c .. m:otumlswi.~li\ioult ,e.lnhtTe:nr'(,or.! "... ose reoo['d~ .~{)ok h 1ll01-e ~ike b<tc.l~ qllil kes I ..~ de;p~liat[lal imcre.tse slres'''i on 'Dbe ymLulid. Iloise' thalli, ea:li'1~lqllakes. A.ll.lhCl~.gh the' ovel'lyinylocked pOi'lli.ou of the faulit thli1~t \v.u~1 prooeg;;.~~ ~~v(llv(\d iiilf(; nut nd.~ytn~dr.;]'S~eH)d. ullilil~a~~Iy00 ruli~'\,t.~dby a grefu co:\rlh~~l!{!ke. lrese.u-clli.en; Imre ~:deilltifiecl~'~ls; pl<lle-loou,lldi 'Um iI [~Cel~~:~)I. it V.f'dS <1ssmn,ed lh11l1h,is.deep aryl);;h,lt, 1t1r <!lIMy th~ C ~~S!,;<1J~lil!bdull~ ior'l ...se·ism~c' 51ip Of dU!ctil~ sh~arir'l;OOll"~M II Zone in Ihe lP<tc:ilfic Na:M'thu,'e!>l (1) ~'lI1ic:l Ille I'] !ues oocm,il·od m S H:nifoffi,1r11peed. "nn e obs N~llk~i Subdll:(:tinJl Zone ()~:H~lc~sIem COil.r.;.[ ~I'v31 ion of ~,11a'll «:5 f!Hl1l ~ySLCmill i~:sur~ 11:1ce J itlp:~lcement::> our omniiii Ultms 'F ~:Qb'llPosImHJiiQltilSprop~1g<l[e

:(',&i.3C1JO

,lUlU.!

C,~,~ll.C'HO H~O~~\:-"Jrd O[Cj ...

f,focj!Cflb; a.re OOlW.llnC!rl,g_ HO\~''0vcr. U IS ~.ol )'6'1:

The alllhQ r i~, trne G~o~kaI5~ Nell' of :(ililladai, lI'ild~[1[ at GeotllClence C~f!lre. Sidne!!", Br ilisl'! (!o~um'bla VSL4B2. C<!!n~d"L[-mail:.hdlra~elit@fIi.COln.gc.cO!

ilh)1iDil1lg Syl!itCD1111(.uPS), SI:fLd01l'~Q\~c:rfl pctilor;l of ~ever.tl \!i,'eeks. led W lile di:.'l~(}yel:Y of 11

"si~n( s~ip (1:\~imt"

elll

d~cCai'l('Udiia Subducti(l!]

~,:soienc;ema9.ar9

SClli!:.NCI!! VOL 315

26 JANUAIRY 2007

471

PERSPECTIVES
ZoI'lC;.'

{5 JI. l~l!isswdy

C;::S~;)ib~ ishc;:dl that

distinct episooes of aseismic :s]~:pof a ~ewcc:ntime~en. em] oeeur onthe

x ~~~,ok9~n~1iR9de-e,~

1~"W'~ueq.!.!1lnc~i!!feItiO~

Pilltch of \l'lf

l!ill1hq~D~

deeper pbH~ iaterface immediately dilW Iili(UP froml 11 complete Iy ~m::ked j'lilate boumJd~uy. lInl"eiolted to "my loeal
carthtll~l<,lk(: ::I,e:! ~vh:{

,!'IoS!ll~liI' 0-.1 rrne'!!!a1i!lJ1lJ$~e"lthq~F;)eo

I

~lIi:lVt S!!:gmeM I1Ifs!ow sillill e\Of,iU

don Of!1llids and tlmeil" P{)~Pli"~!'~llu.c. s, The reg ions of the su bduction zone where great I:~ust (i,e .• megalhmst) c1fl1hqru.aik,t:;Seceursre eharacterised by :1 pm!Ci.!y (~ffll!id:.; .mCMtlf In,,\,
pore~nHhl pressUI'(1!);., res.~IJ.tliflg: .in vel:y~m]:c.
~I'I :gre~~l esrth-

lJo~p lO1,i-lrBQUe;llQI' 1t'~i'I'Il!lr aDi! srO'l\I'~ljp migrdlioi'!

bri~~~em;.perilie:ij that mpture

A n~e:;,~mi~l"l~'OU .111 '~\l11~11Ib~t;': fif

Ili'>egi.on:al ()Ont~InLalJS GPS d,lta over

th!;;i IJ>IM d~,,,d~ ::::htn\;iI;:d dmt such :;,1p i events b;a,\'C oecurred repemed,ly akn~g the n(}fl1.~,~nu C~m:aJi il l~,lm,'gin mml with sm'u):r,15ing l!eg,lJllm:ily (6), Although modeled
t~,es;G

tll~Ia:k~ cwry h~lm~~d j'c:arn o.r $0. The ~kcp~.r ~gion& {If I~J;: '~(l'!Wl;trgt'!U ph~l~ im~rfat;(l., ~~~.le're'i:Ili£il~ temperntures (>550"C) and 3hul1l.dlmU fr1,;ie fluids render material ...plastic .• Ire chasacrerized by eoatinucus aseismic plate mO'[iom •. n.e~m~,l:-!g,liun regioJl betwille,m thew

s H P e ve f1Jts can be by :i>illllp:~e dis>~or::<Ilionsillll, ;)'1,1,

til..istie m~di~m.tb(3p'~)I~(.'.1!~ rmc~cs:

involv,ed in this dYG1lanm,ii,cbehavior em lh,e deepcrpl<lle i.n[Crnlce<ll'C not Wl!a;kf'~u.ll~s.. slow s!lip,.a!1d l~o:w·fn!otlllll!iI'I(1Jli:!<artluIiUlkl!s,. The W<!~m~.65 is trilleal:'ie~ (o·lIlve rg em Il~arl:eb4)1lI da rias is at n dynsmics of culfi""ergenl: pllllie bound- gJovemed hy tl<ie d ~~~ribut..on n!llid~ or Illig h p()re'iJ~E!s~tlJre~ aries \\,~S, c~~ai:m~hcd'by the cl~S-Cfwery ilnd by the me.<ID ~i~Gto! suo IIlger, nu~d1"ll'oor al5p~rilies. lihe shallM !portion or llile S!lIbdmlillllg p~,ate intfil"ila,ce is nund· poor orrl 'll{lnvllk:uok~deep ti!'~Ui!~~'" rduF!!g and h~$ la mge, ~tr()ngl aSlper'1ti estllll;'ll n,llphltf"t il1l 9 re-at earthtI1.eN<lIIkill,i S:~l,bdlJ(;dO:lllZone (iI, D'iI~,1 (jUij~es" ~he deeper [JI1<ll>eiilill\'rfaeo hcta !'ld~~ulid:·rh;h.alll~:5 c from tl.1eNmion~~~ Research h~Mhu~e (Ollliinu:CI!JI~,a!5ei~il~kp·laile mOlioJll" The lramitLon i!!onoebetiMl'en [:'ollfEiI:rt~. Sc ienceand D is!I!;H!I' r.retih,~sll! b'iQ reg itlli:S ma rks a nrroolf,J)g~!l{! QU:S, li!il'!Il'·d.i2'pti'rHJ~rU \'i;:nl~o.lfs(NIED) h~yh-sl;~lll'itivi[yeis- rnri~~.lJriGt fhlli!d"lic:h mater1al <1m!!sm<l~'·5(:aI.e b.rilJtle asperiilies s or IInOgr<IP~,l neI:work (l~ i-ner) enabled !il~al 9i'lle-'~ rise to @p,iS'Oil! of [TS aTh1:i1 ~~ 'VLF e<llrlliu~lIak~s . the id~ntil'tCoMio~ of :Srlulll-~mpi ilude ~J!,daJPld If'(JmIWe.l(lI. (J)] seislVllic uemees ~'h~ll: ~"i'itedfrom <I few mjmU~1-: t~ ~~Ik\v day~.\,vi.d'lJ pc~dnmin~I~'[ fro\l,:~SIJ~rn JaI1;t!~ "'I;) far: Th~ IJUdlOQ."S Ch;:H'IC'Dk'lri~~ queneies :mngiUilE!rom lID mHz a~.d scuree f these VLF etlll:"thqu<'Ikes by <I disl.im::t .~ack of depths <I.verngimtgabout ]Okrn,. F'l1Omlheroca~ h~gh~rreqLJ1.ency oompon~nl£ (i to 8!'[z) rela~iQll. ~!ml~.orl. and mobil it)' ~f]he~mm activd li ve to ~~ll!lIi,~.!r.;rl!!l ..gn~ltmJi::crd imit)' earthquekes, f~ml by :s,iglla~!.> 't]~ala:u:c p:redmni.nmlitly ity •. ~l \WS surmised th!lt the generarion of tre:mJO\IF~ ay be re~ilted tfl the mcve meJ'Ilt of m bodly \"~~,Ve5. Because the seismicity due to Il-u.:uds, oril,!;li nat ing fro:m dle debycllnl~ i'onl V'LI;" e;uth.G1,lIIi1kes accompanies and migrates \l\,1tl~1ndersteed u

A~lother key i,lili.:!lig:flt into real-tlme

or

1"''''0 is chanJc~:eriled, IJ,,;{ sp.n~,'l~and tenrporal in n uid 00nb::l1lt. ami pl'lre~fi~lidllrcs. :mre:s. ~e~ldiil.g~:o lhe phenamena ofepiwdic si;r)w slip ~v\;:~ltS aooo!np.;\r!i~d by VLf "~~D1hqj~l3kesami deepnemors. Can the exteesive d~'p[h disl.~:iblitiOl~ . of Irt1l1,1iU.'irS observe dill Ca SiC.ldi <l ~ N"} be aecOIlUII~od<1l10dl wil:hin this fI'IUlrlCwork? \:Vhat determines t~,le recurrence im:t~:I:V<tlsfOT sLaw ~I ip cvems, and 11m: ....are t'h:e}l d.cpendcml U!iI pl a l~ CO'~:VGfg1;;~C~ nu~s"pl~ul~ age. end
:IL peri[~,e.~ s

external stress a:rigge'rll'! B ..sic ...I]y. wfui .•rt <Ire episodic U:e'11IlIIOf. slow slip, and l:lm.ese' deep Vlf e<!r~~<ilm!1~1f! Perhilp:s.c~e most impor~OI~,l

'~e-neeessary

and sufficient c(lndid(J~lS ~:-~)If

question coacerns the relarionship of

Ih~'S!cphc~~!amf;:l:lia present-day n:g~o~~ll tOo seismicity, hils clear thal episndes of deep EI ip wil I modUli ;H~ th€l roue [If st ~gS aOc\nt'i!~ (~hUii,DIl ,1,Ul l~e' locked portion of the-plate inrerf a ice, thereby 'm.odu,:~"ling the eurmrlativeprebabi Iity Itmld cnnceg.\'i)b:~y ;"ciing as .1 ~.rUggcr) l"orgn::<11 sabduction t.hn,:u earth(,]~tak(;::-;(9),
I~e'

BIJI,l what

abcur earthquakes

occnl,~r:illgwilt~~.:Il1. tbe: subducting

slipr~l'~rwmt,;!llil innueru;~
m"ld tillfll.ilng ofregu,onall

of~mB'h,tbiisalts ii~the subdue:;li rill:!, ot;~i)nk t;nl[i;~, OJ: triggered 'by the' addiuen of j]lI~ds to a cril'i~ Ct,1~y~re$s"l.1ffitldhum. u;r ~lerh..p~lIrigg-erud lily Ille.urby eru1;~lCl[l~ke ,).cti.villY..~t was lfa.ot mnul the' ~tudy ofC:I:Soei'l(iia U'lcmm"}; I( l) that !Jlc int~~lUl1i:!' lrell~ulimillshi'p be'tv,teen I:remms ,nu~,,ep.i~K~ic pk"il!i::sl~p \wlc~G'!>I~lh~b':hoo.Tlh~lrcmoir:f:: WC;ri;: fmnui to be ide'm1I~:cal ~o~hose ob:-.erved iln. J~p..1f~. <'Ind fh.~y m.::C'urrrcd prodo~.~iil,mlly ;,'1.1:

~itW;:.I'i epjl${ld~:'i of slip.. or

.In, Sllbs.eq~len[ cxiliminlQ~iOJ.l fdh clHimiges, o [l(,'(.'orued h)f H i-n~r '1{'''J:.'~fertll,l,'illf'reP.i. ilt \"''<IN discovcl"ietll that tim prcykms,ly idcmir[ed lrenl1.or epizsode,s ]11 sOllithwe:iitem J~~pant\<efe .indeed

dJ;~"p '~ow"fuequencyr:t'iemms iilnd slip ,events. ~mdbeC<LUSelhe' specM.<IJ, (011t1:1III1 fu[ V LF ~\ 'e:11!~;:';~l~ggi.O~n' onrL~plurl;; dli!:m~ifm (J~ the order of I [I 5j•• t~,le SJln:~~O:rs, j)l,IOpo:re aJ ··(!o.~:tillmlln" 11i1lodd of:sl i~~, p:n:l.'KX:S;~S .inl.'h.c lr.l n."i I i(H' ~ol];e o;flhe ,plate ill,l[elr:F.~c::e tfl.le f~gtLret (:see ,~nlhi$ m[ld~.lt (h~ lr.\nl>ituonmlmt,; e(lnt>.i;,>~s prilmlr.~l.y of~",reakly CQuph:d fomlt <1Ire,[ISlllI'lt 1!ffl "l!bj~c'~to Ijph;(i~lk 1!S~h!1~k: $lir'l~at tl~cur ...o'r~r d'~~y.~ lO~''el;:'b dm'i I\g nntl epi,~ SJO(~C, ~:.Iowever.there are numerous ~I,~l,nn.

\V~lh the

slab aud o\'td~}lL~g JCI:il.I~[? DOI;~. [he '~I[~SS release ~;l1oun these more subtle and ben~gnn'p]me~
l[rn~

!ocn!Jnn

seisnml~ci.ly7 Und.er-

:>tilmli~lg th,escri(~~:mhm~h~.ps could provide sotid 1II1d'erpinnings: for e'lrlhqm~ke fo:recasoti :l1g .~i1 ~~lMuCl~e'ilm Z()n~g.
:L G" RIlge~, iii, I1ragM,Srie.flLi":~(IIID, U~Ol t~o.»; p~tJ· ~~hI~doo'~ni!' e, ilia)' 2001.~{iO.tiil6l1(~M!:~,1CieUe3)" ;::. t{, IQi!l;m, It Hil'G$e,. F. ~mami~w, Ii:, Ka~~~r<I,. Ge.jjJlI'Jyt, Rtl, Ult. ,~1, ']J).102W20t]o.4GI!LlI~-oe<loEl (20ii)1l~. :J. Y. 11.a. t:. ~r,iIJ, It ~iclll'j~ ~,se~l1I~ It Hiiros'e. Sdielm~
;!1~5, SOC») Wl!(!I1}; ~~!)Ij~h~ omlille ];0 r4()~mber lOO6 ~lO,l'iL2~clMO~ iU~~,s.::l)·. R!eiWetll()I!!i

S:li~.ghd)f :-ilmngeTp<1lches
event~
~Im, rcsJJ'Ollse

d},1t nu pu~re i!lil VL F

[0 im:n::.'IsICd :-;he<lr stress C<lllSied by S~[WI.'s]~,p in '~ll\iei'wlrrolUmtin~ IDIl~le'" <lcComp:ml~w, by crustall duspl<'l:cemellls;1~tlrib~ ri~l lim~led sire of lil.es.e palc::t~.es. aiollg The ~f~b~~ ~lld~etl p.~iC s]ip (1), Atthou4!h no ,cl~ar \!;Iulh~'h~pl"l;;-~~!lt'eofhighpore-f'luid pre!>$I'lrc~. II.nflch.misln!> 'fi){li Iremo.r genera~iml \lI,'t"l'e estab~ c:tulr<Jct;er~,ze the VUF e<lrl:hq~<lil!:.e' audils i:~(;,htld. he slmi~f.ri~y~lfIhe: (.IS phJ:;:n~(l:liI~,l(lmlrupwrl(l dturoli~Jn. n~:nl'~H'!>. t'!udbll!t~dI(} ~ ,H'e Qccllrr~ng :un bot11 l~U~Nal'l~ai .mt! c.) scatli.,. "um~ i.crocl'ilck i~lg" Wilh i11the U<lnSil1i.ClJil ne z.o
li~ghH1S poimlts to ~i!1 olligol.illg p~~te dy.f.I~lnic j.:l',J!(l!;ess !!;(lfmril~9n~Q young ~tl,Muct'~on zones" emlhGli~,Olkes, may m:: <Imn]~e:r by ":.'lpec~ or t'pf:s(l'Q.k p~flt.e sl ip a~ d€p(h.~u ~~a."1 $(Ju~h~ for over sou 1"Crc- S,iZ(!:ll,ind, !.'iI,lplu:re Ii met;. mudl f

~. if, ~~fl!i!I,.

~L t'l. or:a1]llerl!, I!:.II'I\\lm ~, K ~ie~~.e.10.ool'5J2l'.

S..Jame-s, Sc,/M(t' 2:';l2. :li525 ~2oal6;.pumll~mNl anUn@ l!JI~rll.MOl UO.ltW

Sa'ei1.e 3 ()$, 111121'(2Cum.

1

Th~ rht~~in:l.gsIta, ~'1 ,(If. rc"'eall.h@l VU~ 01"

~naneir [Ib.m thosefQr Vlf eilrt~lqm~kes, 'lhi;$ fllodd is gtcslh:e~kaUy p.l,ei'lsillilgin UIS s:impl icity: The IIIHture of the s~ip 011111.<1 :sln:~~ the f€'r~a."'~j,s ullu!Hfl~ely !kpc._~d(:nl on ihe dil'l~ribuVOL31S SCI~NC:!

6, M. M. millerl. t M:!I.bQ~W. 11).]"liltlmsM, W. 0, '~rnlm, S{ifMU' 29S ••M2.3: ~2!OO;n. 7. I( !Q!iJIJJa., . .scie~a' 1161' (lOO2't B. Ill. tiM.~ oJ., m~idfl!l'436,.$ill (~o,)}. ![l.~ • .I!\'UDlij,], Adii!ill~.8lf~l. Se'iml'o/,Sa., Am. ,~, 1'9:S.~ ~,OO~,

~'(j..

2:6 JAN'WARY 2007

w'l,iVVl,!'.sclen(;;'emaJg,org~

AAAS ,NIEWS & NOTES

ASSOCIAlllOIN A'fF,AIRS

EDITE.D BY IEDWARD W. LEMPINEN

~:11i p:lI':ticlIlar., the controvC'r~y over fedC'.I'3! fLmding lor embryonic stern cellro~<lil'oh has cast a ~Irudm,\' over Ar!1l!,::ric,~',labs, Balh!l!~,nre s ~"id,,~!Oli'ng th'" hi,~ ()I,',\r~gr.ulu,,!w ~~ll{I\."Il!:.~!<!ve I
OO\,ll1 Walry OIIbtll)U

fundull1l

\¥tl(;,~i [I

'ti1.(l.ir fUHliIr(;.

·'.Ifl>i Lmiqueirll my ~xpc.rie.nC(: in SCil~~.lC'C~l d the gon;t.illlK:l'Il .hs put :.hruckks 011 a cX:I'~;liu~.

iJ ~he-Nobel

E~i~c is

ing IICiJ.iC\'CIllC'Ill of a

SlUppOSOO. 1.(1,be: the crownsckmi 11;::career; SQ'lnOOrtDC

ill'l\H::I1vcd

with

[org(!il~() ten ])a\! id Ba~1lmore, Aller slmL"i nglhe [tH, Prize .IIl Mcd:ic.inc For ~11cdlscov el:Y of reverse uanscriptase sr age' 37. Bahum,ore hrus:
p~lb,~i~hed IllIJClf'C timan ,6,00 P,JIlCt;S including .. 'W.lni mil. re:;;~ii1.11ch the gCfICR.icZ'i f eanccr, the on o \..'or'k:i,IIl:!!;,:-; or~he HIV \'~Ili.l~ "lid AU)S\1lcc;ne
!::;Htd

eurrtculum understand

:1 loca1 school board' prm ~dil]~ !Hlhd!l:c and hclping teachers "thc' cxpetuncnral nature [lIT

science,"

B:."lilimm-e said,

kiml of resesreh, 'We've never had a richer SCI. of oQl)~)Onwlities p-m in rrom or i.my selenti SiS lho~l thc opporumltles put :in rro!l!a~ ofbiohlgists todrry. ye~ there is a IlC.r'Vm',ji'\IC sense orbci~lg
abandoned bj'lhc: Natinnal
]mH]~mCS of

his Ca.l"C'CI', Balumerehas ,daycd a key ro]e in ereating uauonal scienee Tliamugholll Im~ic)',m. ,I!l o'f,gOlnil:lCF onhc 1915 A~i~omar

]lcrlIlh."· he S<!idi.

Confnt;nce

id:HCS,

11J1d IIHl.tlhl·

I.hal. ~(~! ,..~111ld,H1:ls foil' :t(;"Cmnbinunt DNA h::;ehlto~ogy.

3S

B,llrumo:!'!;;' SWI][ru::d' do~~'n C,d~cch presido 111 11liSI. S~~~~t;rrnber. hut co nt iL'lI!I(;~ .'I~ ItObcn l\radr!:;ws !\II ill I il.;..n~ Pmfe~:ior 0 r B ~I(}I,ogy and hClld Or~l1C Bfuhi mere Lab ~II. the II n i,-

'I~'"

m11cmf~11 ebservarions i~ molecular imummo]ogy. I ,~e'WaS 3, f(:mndcl" of the

cochair ,ofllle 1936 NmJiona i A cadlcnny (I f

vcrsity.

lib clllrreurIIl'CSCaL'C~l" slipponed

lr.'\i' a

Wh~c.ilc:ad .[nslitutc fnr.

Biomcdic<l~ lll:.csc<II.'ch.

Sciences Committee on a, N m i01U11 SUtl.~c:gy for

t\U)S. and ilcad ot'tlllc N<lL~lon:l1 lnstiuues of .. I-Ilcahlm AIDS Vaccine
R(l.'li;-<IIr(: Bl ("O!luna ~tCCII]

'presj:(_bn of Rockcrc:llc:y Un1\'Cr1'l]l), and the C<llifnrni", hl:-ilinn~ or 'Iee hm1(}lo~y. ~llld,u i¢".. ding
ng~[C ~H narioual

~.99().

Hcr<C'ccivcd the

SI3_'t)c·m~~ grant from the Gates Foumh)~ ~,iol1i l.io:nL islook i~lg lor WfiyS 10 genet ic<llJ:ly boost I he i rn munc sys tern a,ga ifl::>~ i nfcci i(IUS pathogcus. p.miclL~al'llyl il\~ BiO'L~~imol'ea ~Ig:hcd \~ihC'.I'I asked irile ~md ever i '~V1\'ls,iQmxi Illm~e~f'!i5 an i~~u.'ifTh'tt~'llal advocate .,u~]d;ipo~p>cr~(}]l rmsd~~1C'Cin dll; ~,.~n1y of ! d,~ hri~, (:ur()n- "No, I W,lS j ~!;lI"~;lb r:m ~In,d iovl.n~ iL
] \·c son

Ilavid Baltnfltoiie
\I,'OI'I!.;

~.99t) t"bliol\l'I~

Mt'lda I .01:'

pol icy eommlss ions O.1fI UlX"Om bi Ilililil DNA end A II) $p;;scflrch, •.M ueh 0 f 1'111)1~jlfe slace the :Pri.le has been k"lKeIll l~P' \11.. i~ooing selencc," B~llh~.lOJ:C's~id. As ~l lac: ')SSl.mlCS 1~lC" pl'csklcn~}' orAAA,S ln f.cb1"U!ary, '~c: WJ.llI~IS ro l'eLU unCithe piublic: lhil~, "scleuee i:ll,lhe driving: fnru·' or •.!] moobm~~fc,. nOlj~I~H hi~O'I..'n,
I!"~ "In i~fl:". Ib.1:!:i,n~gll,i Iil:il1k~h ;!io~OOC~l IfI:5l in

Sciencein pan for his o.nL!\JIJS rescilrcUl poMey. I~."lt~ morelsas bC'C'111 eutspoke 11f11bom what he sees, <IS gO\lcnllm.cl1t ,efforts 10d iSI.ort and smJr press sciemi f:uc research, and hc thjnks that AAA S has 8 rolew play iu :fighling Ibls~liC'nd as \\ 'C II. AAA S":5 conga,c~im~;d~cl'iJtinUl!lJ' mld se ic!1:C1;i~MK.'"bric:.fin~ .. for !leW ml,~rnlx,;,] nfCo:n~~I;i'i(\ by Ill,.;: AAAS CCI1tI.;':I 1'01' Seience, R~~hIlPIUI.g,)'. <,IIlldCnllg:n;:;,.'i t;I.Il help rti:>tcm.~ :.l'i(;'Jlt11.;' S

or g!tn\'n;l~liItO

I.~~ i:;11S ~.corL"'-~U(,IfI(:C of C~UrCimi

nm)" ~m~reiil lI1,SL111.l1Iliollil." in

Billtiim.or(, will succeed ~he

presl-

clem!. JO~ln .~ Holdren, ai the close oflile .2007 Anm13~ Mec1jl~g O.rrI 19 February. I loldren \1; ill become eha:! r [I f t he A A A S bo,rrdi" as G ili:lC'fl S, Omenn steps down as ella,ir.Jam!:::'] J. McCar!~IY Im:->been electedm succeed 8ilhi~ 111Mm:. ,~::; prc~id~~n~.-'cl[cct. McC;.l!uhy i:> U!C
All;x,lnd~'~' Ag:;;uj:,;;iz .rrnli.,;~.'ijQ;r orBi{~,~o!!!;ill;;,~1 Oc1o,i',lIlugr .. phy.ti H"nr;.rd UIlJD\'l,':rl'i~ly,

]hc;: da~.~rmmil C.Qillg~:,~:>i'iill I[C'~·"'ml yi;'~I·l!,. .md .'~V~ Ih~~il;n~"\' .fu!o,~1ing il:\clhllndc r "1~;JC:kfrom nur:i.ol.liSsides, h ':s nc-o."C'r een ;l more impolfll'l:rn b tlm,e- for scientists h) take ~I,role in pablic llfe., .and dlCiPCis bardlyflll1Y t')!~bcrW'gill1liz,mioGl lhat C~1fI s«)c<lkasbroadly for scic~u"AAAS.:·' Ime nOlcd in 11rcCCl:lI,uluer"iC\.~·. Sd10(l~S are one place \viJ.CN" "·:ill.crc'i1:il,~llg .-:kClllkjsm <l1lJ.rn.U! m~~Qnil~ !hinking" has p.Jo1ccd ... 'C'k~,wcunder si~ ... "Cot.~n~in_g '~oB!illtir!lrlirc, J~
~l~:>c.fitldi(!lfl:cy ::n<lll.CmCi1I, he Sil~d. "Rllils.il1,!;; children ,~~I~o not kmjiN thm IiIfc Olr:lld our p~~U~'1 do cvo~vcd m CI" 4.5 bmJo(t') ycars of ehaIlgc i),lld

IlCtt.,,*,~ry P~!ICC],I]r.ltllioH;d

IloI)iity •. he- :;'Iid,

A"AAS

Becky

Hall1

CaU 'or No.m,in.ation ,(1'1 AAA.5 F@nl~ow,5.

AAA5IF€~I(IWS ~liho.are (tlrl'e;l1i IrIlo€fllbers.'a! th€A.ssodatioflJ ~ii€'fnvil'€'d I!O II'l()minat€ IflfI~ll~ber:i fol'_titm "5IFell.O'j\I~,.A F~~m'!,lisde:~irred (I) ",[II Melllllberli'I~li05oe~~lol"ls Olin ben.lit o.f thNtdvatncemernt. of ~d:· enON'Jf its apllDli.ca:rn(]lrn ~.iel1lliJi{:aUy or ~i.aUly rl'i,S!il!lgUik5hl:d." AIIl(]lmirn~~oll! IlllUllleo 5IJlonllorred ajN~ by Ulrl!1! NJA5 f,!:'UQlWS" tll'l'DoJ wtlClliIl muS'l. h~v@'no arlLllatiol1l ,\Ii lill Um ~Drnin~'s ~l1stitl.lliol'l. NDmll'latiOf:l'~ 1,1 F1de~go trel{f:eqv by ~jhe Sli~ril'lg GrQu~~ or lfle A51l0d~ ~QI1'S !leCUCU1S nh~(hair, (milrr-El~L Ri8'lir~ Fl!ll ClliliiIT, Seore<ti'lll)'. jj 111:1 fbur Me'Jl!.bl!~-ilHarge(]lr eitl:m s@cljQn~.EiK~ Sleer"l1Ig Group, ~EvioE'ws.QIllI.y llrlooe.rlomirnalio~~ de~!Jfl.i!Ited f~ril5~(lj(lil~. Names of fdb'lllllominl'!eswnO:HUl;! approved ~~ th~ Stei!riil!~ GiIOlI!PS pmsE!l'!:tedto th€ AM'S (lJilll'!.{ilfor eteltl'orni. are NmnlFla~iollM~ with cmn,plele dmUm'i21Fllci.tiolll nllL.I5[ bl!! recEived' by 11 MaJl' :2007. NG:llflnin8lIllolfl~ re,"i1~\I'!iIdalb::r thil dlii!'lle\lim ~~ held lor lh~~ollW!'iP'ngJ~FTll1re' JilOimill1a1i.QO f'~rm ::md' a,lis,lQr (u~rnnt IJAA5 Fellolil,>can be ~oulld om ihe AMS Web :sitea~.WIMY.aaills.or'£lJaoolli.aaa5l~elkIlI'i\S. To request ,a har~ cCIl'y ~rthe 1II1J'1'J~inol!licn form. pl~s!l: wnL'fllct IEU'IlriC!:Uj)e'Jll~fi:!t ~h'll AAAS &~L!'!i¥e O'~ffoCe. 1200 N'ew Yom Aw., N.W" Wasnilll~tlJl1, [lc, 20005, aft 2'02-3.26-6 63'5, ~:r at ellalr!J!nt@ai:talO.oJ]!,

devclQPIMcllI. h,~~nf>tl'i them fmnn par. ~ci~it~ I1lgs;

hfc.'" B ;l~ t imore '1""lI!U s: AA /\,$ to i;onA imile: it [0,1 st!"rm Ii!: S~Ipp-p'n of ~~i,;k'n'~~ ~~tUl;;'at ~ iQ~rrol1l1
iBg~IJ nlI'Cl(k:m

r.;inJ.~r~'ITt~n lo ~olh;:~~. ;~w~inl\

SCk:fiCC

lh;;!t~l

:JI.ifi."I~Y li1;.t1d ItllCd~, :iJ~lt1CCS", wi I,h C'LlYtiqllmn

h<H~ m~d

likmcyptogmll;l:> :iut~h ~s PnJjcci. i:06,J, ~cld o~[refLch (lllheAAAS A:~a~nl.~1VIC'Ciing, : AAAS :ihOlLld remind ii!> :rnembe:rl'> tll1U. ''ithC«l is :1111 l~port"mi1)1 forevcry sdclllisuo b.;;: o

413

HUM,AN

RIGHTS

AAA;S lEx,lpands, Hluman

Rii,gh~s,"~Eylein the' Sk'V'"

After 11 SlncCIO!s"s fu!1 YCfLr!cm,g pilot PM.!:lJl;l;lI11I, ,AAAS will dr3gnalk.d~y expaad its use of p(Jwcrrlll'~ S;JJtclllillC ill~cry and relllcd tcC'h~ r~ologies 10 h~li} ~'WC"'Ii,g~~w~n~c~·mlliom~1 ~POL1•.;ofh!lIl1.~nU"j~1~~vinb'rio!1.'" wi~hII new 3·y<:.'iR. S.760'.OOOgr.HlII fwon~tl~~J'oIUl D- uud
Cu(I:uJn Ill: T_ Ni:J~~AI1.hurF()~llldlltinn_

II...,

~:

Duringthe piJlot projcet, AAAS, it~ Beklr~ ilnd ilher :5i11ffiUiw ijmages or 'B~r K.4!dolIiIS, CJli!ldl,loo mEld d UrilllQI illlIaJHilckin Deoomber .-:l.i()OS Jill njaweed :; by rmi~Mln~ in Ihe 1)8rfu r (oflfih:t iii PfUI:Ctll~J.'ghipW.ill:! odtc.r orgm~i.GlliOl1s;. used geuspatial tccbaologles to document dk:vclop an olllucach p:,rogrnnm~hm!llgh \:\bb S~lCS, \Vi.lhiflllhcp'!'I..~1year. NGOs;Stli.:&DJsAmncsly human ri.ghls vlelations ~rolHn.d lh~ worlti. \~o rik:silops. nlcct Ings. amI ccrrespcadcnce wi Ih including in the Darfur region orS!:.HI;;m .• hrlcmation:.'lI. the U.S" Cernmittec ror .1luman ol'g;[llniz:Jlliol"]sfhm have expertise appJi.cabk to Bumm. and Zillllb.m.b1wc. Righ~:s .im N mill Ko.fe.1!. mldl~lc U. S" Campa ign human !':~§h!",Ilmjcc1ts. R'lll" Burma h,},;c worked \\I'i111Ai\A'S ~o nner~c \Vilhnbc l1ew .l,ll<tntlhc: Seience and [ lumau Righl~ pmgl'UlIl \\ ill C~llall:d upon i~s 20r:l5 2006 pllmlJlrc~(;'(:I. wUlh,u I~\'II' eltlpll'lIsi:;.nn lOuiioriulg it~ ~'(~liviliL:~ '[0 the: pdor.ul.il;_~:.i of inli'n:gOYCml'ltCIlHIi
org;)nb.:<lIli:on~ h~gh.IiL;'s;ohni(m :;;.,,~cl~ilC ilUllgcry \",'i~h gC\(I· !i!:I<lfI1Ihi,\': inroliltlmu:on. .';jygl.~I:n'lOl!nd lIUII'I1:i:11 unI(:l~ i·

AAAS
'cllt'~ 111l~]

IliOJS~'C11,1 pi(mOCF il'~ b~ingung; sci-

ilirnd COLlJI:;; Jn1(;~hc

]mcmIlIIOll!11

Cdim illufll Cour~ end the Afrk'JfI Ceurt on I,luman al!ld PcOJ.:dcs'R~ghts. "This gn~n~l.s extremely excitiUlg because il. i wHI allow us to developthe rnrst steps oh

L,H,}; Bromlcy. ~\llmi(llr pmgrou]~ .,~s..".m;;:umc In lh,,~ AAAS OL"rIC'~ 0 1'.1 !II!I;U-I'I<~E !l.ili'~liv~t;;. immll III hue J\I\i!ay 201.16, AAAS rdl;a:;cc:!uu; 1'i[;:;01
~lUJlMn

woddwide irafo:r:ILU!Ltkm ~")'Sl)c.m1lSil~g advaneed ~cdl'r~ologie$uh[u siDould help prevcntlnnu<lll L"i,g;IH~violmioss hetore ~hcy acecleraic," said

gene!: m raise puB1llii,(;aW[lrell(~s ali1d<~kn gov· ... ermnems, media ounicls, ami othersabnut hI! nnml ri!;hl. V.tobl~o.i:lS .. \Vh Iile the O\lC mill I) rice (I r new satell he iltllagc.ry and pG\\crJli1 compilltcrs to 3tlillyzc the I1ndimgs Ims 1:\."':CClblllya]~cl]. AAAS ims :~a"'CJvcd r other grants from a \'tll:icty oro:r,gJn~Zali<Dlls .. ~11.(;ludi ,g mll1W<lru or 550.000 Frm:n ~he OJ:KlB B
S~}C'.~~IY lnstiuna
rOlf

u;-dln]oloJi:Y Do bear 011 illt~rl'!;_uioIl<J1 hU.I1il';:1.I1glu s U151iiLI(:;S wh~~ l',egul!'lf rt::ptltU:-. 11!1.d ri \'o'ork.:i,hops; sponsored by its Sckuu~c rmd ] luman R~ghl5 pregrasn, The p'.mgrllilil also
SblPP'O[Qii

selcnusrs 'Whos~~mm!ln rights

arc

liJrC!ltc'lloo. Ibrollgl~ll~c

AAASi

1!IHmmRighl.s

mi(miwrillg

in [he Kar-cn

Action NCQ wo~k. ln J:aflltr Jry' :200(~.the new,'Orit rooroi rJalcdml ,;JJ~p(.'''llto the gQIVClTlll1:Cm or GumclnlI!J re proIC'Cl Fredy Pcceerell], executive dirc'C'lOW of
(j U.:1'rO'llU,aJ,~n i:lfl;H:jik· F

Sun~ ef Bunna. A £C'~ml·~ nUrn~'l wiU 'be on rdc,I!S~-d1,lh:tthih~ yc,~r.a~ ""ill I"CI]![l:t1:'i Oil Dr~l"rur
fin!! d&"C\vhen:,

An!h 11')PD~(lgy l'~t}.u d,~· n

~u IlrddhiCi.n m gm.herirlg and ilnn.~y.dUlg E~U; rights repon, ii1 pm1illC'.t:>ihilP,withAmJlCSLy I me I'mH lena I :l!1,mllhe Ziml.:mbwc Lawyers [01.' ilmages from COI11;POlll£sli""c lJigi~il]:Globe anti I :~mn.."Ln Rj~hls,co rely on Slll1t'~Hr:cima:gocry. Gool~ye. S.I :~~ rlm1s to !lISC the grant 111.0ncy 1;0 - B!f!rljl1mifl 5arne,s '"hc repon p.rcsclltcd evidence thai. the go~.'~ erlllneni 0 rZim'bal:l'we had destroyed cn~irc seulcmems, hJ:d~Jdlllglhelown ofPorl~, I~<ll'l"n. ANINlUAl MEE1f"~N'G fOfci.ng ~hoLI('mmh;,of residerus to i1.C>C' their li!{l~IlC~, Rcpon!;J (li~~ !Ill: Zi~llb;JI,bwi:.<L~! s .. l.e~I :il:1; SUl5tainabiil~i~.y i~IHI~~I:}' \VCTl; pu b]i.shcd iu si;\'C"wl ()lull~[~. int:J.lldiIl1Jglhc ~!h.';A~rlglml nrllt~, N;lII:iPIl ... •• 1 [he' AM5 tlinl'll!lat M.ee.ting slands i!liIlar~:lrsm olliire.r s.defi~; rk gaUl erring!!> in lts eml~hli!lsi!)on lihe Pdbl.icR'ldio, the (..J'I{{~f'(lulll.ml'd ~tcLLIC'rs. COiro IiI@C~10!'r b~twe@ III !lCl@iI'!(@allJd S(JCiEly. ;i! !:.rilic;CI1 cW:s.sflJ.a'i:ls (hat ~~@~pIJJr,e:din d~f]D._ih willh lhi~ .. \\'!I hi iLglooo:1 reach flll.drcpuhlllon for ~rI~lr year's 'Ihenlie of ~Sdeme am:n·'el!'.:hl'l,ol.ogy hn:r SI!I:5tai nabl.e Wl"U-iE!eil1l g"" pendence. hlllllHillii rlg~lt::;Ql'gllfli ~.t ions are Ol~I,c AI. the lBrtI AI'!nldall Me~~ing fill San Fram::isco ,i1(l!(~mo~lh, '5(ie;n~j:sts..i@ilIm~lfists" and Mle ~ub· HI sound the a:ll<llnl ;)111,", mobiHzc rail li(m:s: Ht: (an e)llp!icm!! 'the hlea O"~ '~uf>l;al abi ~ily a rmss al wide; l1Iumber of diKiplflllil!5~ lrom iJililliJte-dh.ange n qtlic,kly. ol'len bc.i'orc ,2;0'\ ernments or tile intersdi!~ C~ a ~d the rilltur~ ®~ lIiI.eocearus t(l~he p~~ili~s 011e 1lr!!lgYre1~ntfl'S, ~Iild ered IlIm i011<11 C()l1.nmmfty hm C' C'!i'(11 adnm:iucd I here is I.al'lg,l!IagF5~ iJn.~!mviiron l1ilell~;alll!1!itie:€'.. a problem," said Arie!", BHi~~cr.director OFlhc Iii! his. fnvHaUoli'! toO the ,colflJ(l!ri';!Jii,~,MA.S Preskle'F!! John risis VrcH;IHinn 8:. '~!i;:spo'nse C~l1u:r ,U IHotd rerrl ~aid., ~The (h.ilF.ilI~t~r 'lilf the clil.aUellg!!! ~ t,o the hI.!mu AnU1'csly Inlh::mtnIOfll;].1 USA,.t pilru1cf of ({llIlu:lhi(ll1-el'lergy. w,aler, heal~lh, dirmat.e., sewrHy. AAAS's SdcllccaraG .1I i1ma ill .!tigh"l:s: progr<llm. deVe~O!~lllent. (flllId IiIl(iml~lire tbelOl lackted by AAAS's 51 n.C!c ~ 97'8, the M a-cArlllllu" rom~dm IIm.s inll 1fIItef1di ~d ill! ~ry a Itll?tOa cb, pi p.m" ~dcd S~II}rO:rl hi <ldvallce hmmm r~ghls For Ilhe rirs[ time this year,. !it MM Antnlulll Mee'~in9 a~'(h'L!I'K! the \~imil~~ \"!~lirI a ~r~i<l~ i n1;CN'~t in innCl-'" 1J~(lg [lhUo.:Nne'ws,i!I<!BS.orQ ] wi~l. nwide e.:del'ls~e' mVl"r· p \';t!~iv~~1l'1:~(lr~~mn.l~ thm Oldvancc :i!!'i:grama a'(l~ horn S,;Jn IFf ':1!'!Cisw. Je-alli! ri IiIg lfe:ptms arid pJXICil'SitS ~l<Jk i~g r,;plll,;:,1UlIJ'fu~!l~b!i:D11:~ N(m;;mb~r ~f.M15 'ffrmH the sta~l o~ Sciet!CJe a II1ld Sciem-eNOW, AAAS'~ award· W!,ik~~ !fj:[~nl o~·S:[ :~ 0;000 to AA,f\S \I"·.H.~ to ~~irminlg S>c~elD'Updal~' radio pr~'illrC!ml.and MAS's wril@W5, lladp imcli!l;liiorl!)1 org1lUlil.~atU(m:;,. gO'licr.mnt:lli. (!~onglwillil tl1nlks to U.S" and! int!!mlil'ljollla\ IIr!11M(QlI!Hagl!! IJr ~gclt(' iC5, and 1fI0I1I!riO¥''';i1rlK:nt~110 rrgrunn i?.1l'U Ion::> ~he1rL'e'eli[jlg. (NOO ...) mpi!d.ly col k~c;!. \'il~ C Imtc, fllKI d~;s1ribbji.~ iuror IliWit i 0111 bout h r,Hmlll r.igh ts, vio 1<11,1 <L iorms -,kdyHoffl !:.Islnggrosp,u~iill] IcdlllW ~g~e.s.

II Oldttt~h'(id (k;,JI,thLi1JCOHl'I ;.l.g.tIi [lli'l .~~l 1. 11.' ~~Ild 1;,~I1'l1ilyfor his effons I.n ]OC;Ji.lC 1II.IiU:~ his ,e,.. humc m'ltlis.g,rtLl;I(;..~~,o;rn [he (lOlll1lry"s bJ.oody ,ci\'i~ Wflii ..
liarl.
\.'rI'~!O

'homl AllS,ides

"Im~

~"'L;~

26 JANiU~RY2007

VOL315

SCIENCE w'INW_sclencem!,g.org

lGI'!II,,~raJl O~'fims

S@(t]OI1'

,en IEd'u,~~ia'lIil

:!ieclion,

01'1

A~ed~QljL SCi@iIlOOS:

~llsident.£rKt: l1Jml2S:j. tl~cCi\lD'lhy

8(;~rd Q{ Dir:ecl,'Qrs: Chllr.r!(Mur!i~. lij!loo P.:B. ~'Ia-'hi CQmmlirl~' O~ Nom'~'()!I'Ij1Js: Rita R. (o,[w~li. JaM ~u1l)dleIl'lCO"

Ch'Oir"f{IKI,· GlWrg~ D. N~s:QI'! Mtmb€I.-Ql,t!1fi'le::GeroIG Wh~eler flec!'orot·e N()mla(J'irlrJ (I()mmin~e: (",tl'lirY!i!

~Io~ iE. Bloom, Mary L Good 1100(1,

It. ~ln!:hJ~,
Sifl!~H{l1il '®1Il

(afh~ Pearra'!i3 1Il(),

hglill!leerilll:£Ill

CfI!3ir.£fl'd.': Gail, H. (,assc4l M€mtlt',~Hojl'ge: It ~r! B. Emk;Q~U: Ei€oCl'C'r'CM~ NommtJti'tlg CommNlee: f~~r f. Mld\lallll~derl1ilan

weller,

Sedi~m

1I1111 A!lIlriCJu~~ll're.

anlll Rl!!,",~w<libteRe50l.Il'~s Cliafr.Ell!tt' Roger N. Beamv Mml!)il'l'-nI'MJ!1t: ChaHe~l. oi"iiml2'e1'll 8~c(()tf}!'e .IV:am!i'm:lIi:rJj {ommillee: Sllly A, M'J(l\"enzi g, If§dI!iJrd A., lDiii«ln

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Sedia n 01'1 Nielll m®t!>C:i,enc-'El a.1()'~-t.ft!d: John 1'1. ~yJflle Membli'ii'ltlHmye.::' !!::ha.Jii~e~ 'Gil~m I). Et~wNtJfe lIIomi'fla~j;ng 'o,m,lJIilt~,' Ali,5CIl Goal!!',

Thoodol'e IN- B~~,fll'

C~!JI'I,il D€ie9~te: P;a~<11;1 K. t;;;Ulhl
Se.!ti>OIll 01'1 Plnll'lr,I!I1I'iilUlinJ'lic,all Sdel'lal'!i

:Sedi~:n 01111 IiliUnopor:o.g!y A CliaiI'4:ter.t El.iI9!!nieC. Srn~t ,Mml"el'ii/'Mtgt: l~e~lie' AIe[lo C. [,1ecr,om,,!'!!' Nomilll'l,!'tJllSl' ,(,O,lul7irlte: (I. P l'Ii ~ll1P' ~gllUnire. Pol~on Sheel5 OrJncilD'eie:gale: Mr~hlilel t little' I
Sledi!1lr! (I!'W .AsU,I:)Ji!(I!1iII¥ CIl(Jil".:£tl!(./: lill ((lmelll~ rw ~i~~J-(II-~: Cijm~Midlil~l iJs.s@
fierliJ'f(1If! NmIi'~!lU.~!J ihmm itt(!'f;; Ala n mar<!:dl;L:lr, ~iI!i rt!rMloorg CorJtt.rU'D'rl6gme: Heidi It lHamm~t

(b()ir-t:fJ!a: 1I)1I11kilY ,D. s!IH~1I MemfNi'/.tlHerge.::' laln.A,. Stair Eleet orate III("'lIi~Ulli lfJ' Cammitttle: I

]0 icu'I D.

Scl1uelz",

Kel1llll'lh

W, Mille<r

s~a:iO!l'(!r! G~~~OogY ~rno~1G!(1'~~i!phy

Ctlwilcf(ecl:VicOOr R, Bak~rr
Membe("(1l".t~Jge"~ L~!fIfnleG, TliiompS()i'I floct,OID.!e {il.()mIDrJling Ct~mm'l're~ NeillCl. <liJ)dy:k€, S~Na.Y,il c. ~ril2. [mw,ril Dl!tf!9'lII,~' Dou.g'Lasj.5<hetmalil

:5ecli'@!l1 (111 Phlyslc; (1JaFr"E(~(;j': CIi1~'1'YI'lMaw Ah?m,~(-,~Hctgf:Sau~ [lawwl'! ~~~Cl.O-rau~ Ofif'Iril'llalifilg(@liI1lmiltee: B~,~b>lriliGO$$ N Le'lrJ, IEli!Ulbeth Rei!l~ Co:tJllciUle{egt:ues: Miiriallil1lPo Sarachik, W. (arl UlllebErg~T P:sycliro logry C.h!lPf-iilJd: Ul.l GIl'itman M€m~'~~hI.Qlge~ e:rWll Plir!nGemcSb~.oor M Ell@(tQ~alt'~' Nlomil!'l~U!flg (QIiI1I!llliU@l2': ]Iolhlrl ~, K~tnl.:sb'ilIm.Rijduilj,d L 1!kI1~

:S@(tllmJ:f'I01lJAtll'll@iSpoolt(. a n!ll HIV,dlrosp,tu!i1i,c

S(J]!'.lIIJlOOS

Section '~II1IHis;LQlnyallilil Pihil~m;tphy Choir-f{eel,: lha;mrus l'Ilickl.es.

'~fScii@lloo

SecHon Oil

Ooir'.:£led: All ne M. 'T11o:ITlpS~m .Membtf·~U.-lat9~;iP'tWr fI. GI@i(k EtWf)f(Ji<! UJOmiuaUng (ommrr,!e€;jtfHlif'~r A, ~rrand$, PillMa (}I.!I iliilil

M!!!.mber-'OHar~~,' Ki1Jen A. IRilider r !:lector'l'te Nomifi~li~9 (cmJ'mme!!!': t:;~thry!l 11'1, Ot~q" r:tidl~!'dl M. B\Jri~Q S>£t~tion '®111 Illnd!lls:trial Sd:en(')e andhdmology
OtlJir-f1tlJ," S:~ i'I 'Bull

:Secti~:n

01111 iol.o'glkal SdencI!'s B ChoPr~k,'c:r;' lIMa riian.::lJ W,01iflll'l'

MlrmIM'l'·ll'r"-I.al§ll1':ym.allda P.{mil Eh!c~OfiliU! NIOllilillJa:~litgl '®mmi.~!tn: KatiE~ Ehar.i!~H:iI, J(u~l HIubilmaJ11 Ci!iI~lJcilDldegotif!">: 9r'~Fld~ 8~s.s, N{!IIIl!!:YB@d,~ge, ](I"'~nmOlS(lllmi~t, (~lheillfle IKr'UI~l Miidl(1f.li (@.;(, Rolli! S'l'l'l'H-e.Ti)b~~lhl'9l~, O~i~ Otl~~.£r. Ofilme S!hil'k.es
01111 h:emlil.try C [/l(}Pr·.ftlect waYi'l~ t. (lllldfeller M'l'mbJ1't'"(]'t'-laf!ll'e: IDffillistJ.. DewJhen.y Hoct.orale NOO1iniilltil!l!) (olillfllliuee: Unrla: {. HsiehWilson. (ireyony Co IFu (!'Jundl De(egme~: C iBradl~y MooJ'e,. Jon ~ard¥. Nkholi!s Wino!!i~li!d

MP:llll.lelr""/·,~mg~' Thorn. Milrofl flecrDmrl? N![l.j~ill(ll.ir.lg f (lJnmiUqe: Avile", ]~~e A... l!4elltetcm

A.IfIBI

(11'1:S,®ci!iill~, ~1:.onIO:lfIlic..,iilnd IPriJlitff:[:a11 ScienCieJii live ~i!'l~e' [/uJir·1HI!€oe Itt!lls.~ell."arillilll Memf:;le"'tlH.G,tge.: Amclld Zl'llilleu E Ile<.tm!3Jte Nlo'm~l'Ii1~i(fig (i('1IIl1l1l iU!ee: Ga ry l.

S!e.(Uon

S~ct]O!1 '~:f! l~nr'@,Jfllil1in;iQIiIi~ (orn,ln!JtlITlQ. and

Allire!!:ht. Go!!y 'ltillig Seui'((I1il (l1'I5'i;u:metaU

Ili'!l,p{I(t!i

(ommu IWi'liatJ(!!il1
rh()it;.E'(~I: ,M.i(III~~l N~~on K

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MItmlt€f'11HGrge-.- ell rfi!Sliln!l' L

5edi~n

!'Mlrgmiilil flW,('JfC!'e Mtlil1lillllUfl9 (Clirlm.il't~;IDebOtB iii ts~rifll, G Ladrs A. (!~n~r

Sedion

'®iI'I

Un:gJlffis~.ks, L:lir~lgl!lli!.ge Sdel1lce: .Hld

Gngi liIeering Cllfl~~fCtl: ILems 8raln!;(uml) M, Mfm,~"(lHQfge;' R.llillit , Fristl:lbarn t E.~ert~rn~' fI4'Gf'I'Iil1c.atin,g (om miueec ~ober'il C®olHleegalii

Alil'n

Ela:iI,j'QIiII\

Ml'iWlbef·6l·.~&ye",'

(::h.ml'"~{ecl,' F~erleridkl.lNewl'llleyer We,rJ:ciy K. WlilkJil1'l5, EledorriUe N~Ii!'.rjfj6~il'i9'Commitlee: SaraI'! Grey Thof!l\)solli. B~rba~ LM!5l

Secnon Oil StatlSW(S Chair.£f.etl: WiiUiilll1I f.IEddy Membe',~Ha{qe.,· fr(l !lIoois~ s.~mq~r·M.oi~iwil'S(,h
E/~a'QfOr€ No:wi.M!'ffl9 Comm,t,ee; lludilh M. lIinlllF, Nomnam B~l()w

Sed!(II:['I (lnl I}eli'lt~.s:try aJl1!dl oarat Hea~h S(ffielllu:es (J~fr"fie(t Ma ry Mad),l)u!)!I1i Mi:>mlN?f"'aHal'9C': lP'ilIlillt IKreb!1lJxida IUeCMfGl',!:' fll'G',m~nati,rl:!l [ommiUfC': l.u~!). Aillill . mPletro, 1"ITiimM1~him!llfJ3i

5ed1on,

!I:)J'I MlI~he!1ill(l:lic; (llqit"Eiffl;' 'WiUia 11i'I1'~(~

mell1bitt"CH.fiIrg,f't

'Claudia fl4euill~l!Iser

,fiec.'lrJmte

N!!lIl1lllDI.Png [mnJJM!I&,' Firal1.d"i: SUlnt .. .aI'll,

De$~'(l/;p,' j!OOl

JUlin, Mfrn

Cmmr.rl

L. ~ebDMtl

~.:soienc;ema9.ar9

SCIII!!:,NCI!! VOL 315

26 JANUAIRY 2007

415

I]Qw di5lincu

mustclustcts l~ fQr~hit' wll

'b~m>l,.'T

IR.B.c.o.m.bnation andlhlB' INatulra i of Bacterial S'lpe!ciialiiio,n

G!!'IiI,etk S!lJli'l.!'eys 1'IS'\!'8'althe dlivE!ui:ly of bailt.'el'11a and le1lid t,o the ques.'l:ionfing o'f spsd!!'5 wnc:epts. i!!St!d 1:0 (at!91l)rize! bad'(!rilal. Orle i(iinic:~~ly dle'ff'ining !laderii'tl :spE!'(i(!() ari~~ h01lTl1the Ihi9~ H!'l@S or In r@OOmbiOi:!!li'C!nl ~lhilit n~$l!ilt$~nth~ ~r.allliSr@r of IDNA D@tw!@111 Ir!!'~C!rilfely dliiSt~r'lUJl' l'\~l,Jte,tibC![t~ri,~. Ba rfi~.r5 '00 I tli~ IPU()OO'~j i;'i!lli(~ (O,L!~d~e lIS!oo ~IQI de~1ne !i~IE!ci@s, IFlati!!r a Ily., oil n~ f!o:t '''ppa.r@!lJt. H~r~. Wi!! ~vi~~~ ((lIfI!:<f.!phl!'aI, !i!'\'Od ~~s Orb,,(:t~,F;~(iI~ SP~(~ a~~(lillculd cl~s:cn be ~ulr (I!iI'! p:UlbH simulatiOi!1r$ ,10 f $ped~ticu,,!1., '0 I)ll' fi n~h'li!!JiS~llgg,~t thQit the ralti;' l(!If rECOl'lflbiil'llat~(l1i! nd illS re ~a~ijOrI'lt~ !J~I'!,Ui( a lriilol'e<r,g'eln{~ have 0:11 tl~ol'l}giI'lUuel'l~@ 011'1OIll~c:rHr!e~,We p:rop[!sethat a di:>lil"llcth'll'l be made s i:!~twee,1fl dD:mll djJ,r,ergen.c:e amid s~xlll'al. ~pedat.l\:II1I. Hence" tll' make semlse oi bac:l:eriaL ,d~\i!en'it.YI we ne'ed da~.al mlO,tonlly(r1om g.HllIe1iic: su n,l',eys but a'lsofrlom Rxperimll'!IlIt.:1il detlumiuition ,~f!ie~e~Jtion ~\HES;~Ulr,es, and I\e-r;ombill(ltiol'll IrllU~S a~dfr(liil11 tlte~::lin~tka,l (iodl~ls, m

~(} 'l;n; ~n!.,"lI;;'t~\~~rlo~IJilh to m,li rli,~ !l· So\'H~:r,<L~~I! and lOr tbe ~~f(1h,II.I~'[~lW!.)' of~~~J;:hl;:~u:>[C'r be 1i~11; to !;iii~il],(:l'! ~s i~(;!\: II ~ns:ics;tt:11t 1I11{:ch~uni~ of s«J.t" ~n ... ci~ltion Ih,ul"pr.]~cs (1;1 OJ!~ll.'l!ac~I:.lr.~J,i!~1!1;;"clijve or !lie '~~!.r;g "Jld 1l1!(;'(:h"m i"!11~ oJ n.;oombill;)n!n,], which !In: biiL,'wn t"llIip;ticil~.liy to bi; 1..~";I~lirid)'

,jHr.i.. hJc:~'
h~(Idelillilg BacterlaJl DiVi@r5,Uy

poOplqliu.ion •;" u::;~m~y . I provide a :>Imk 1)~Clllre of the 'p:III'IC'I'ns- ~cno~)r I;)'l1'i,c cl.lI::>tcr~~~.!:l; cm1."\CguClHIi)'. C'.xpk~r:rn:g: Ihe d>'n..nues of poplllilaions. reClll ircs~hoo'~c~kal umxf.clls ~Iudicd usi~!g C'OI~lpll:tcr :'i~fIlU Imio!1!' ~w!c1 O1Ul.",lytical ;;t]~p3t'1l1iinl"l~·n$. Chl.~tL;'~Jng in !l!!!tur.d 'I'lOII]Wll~liml~.~'U~b th~'I~ bN;lOllliJJtLred \.'I'inh Illn'""i'II-!Hl'l
,G;:~lClic :.Lli'i,lC)lS of baetC'l'ial

froqllc:mly IElWLX!Jl DNA sC'qIJ(.'Jlclt'slli::H arc clmC·rel~l. than betwecnihosc [hill are simillar genorncs 10:1:' I'I1ICl1I1Ibcrs 01" Ihe s:!tl1bC, Oli" ~~iI'l1Iitlm'> (.!IJ-WJ. h~ c.1>lpcrilltclltaJl stadles ,oF n:i:ombilml··· defined role..s by uunatlon er rooombi~ll1ti()ll. We species i~diica~c the (!ymUllOC: nal!l!lre or geniC' tion in bacteria fmmwidcly diIT'crulIg ~C'nCira, a acql.lisi~Eoil!l. 105$,""radi!lItlns:rcr {l}. II iis l)ll.uba.lbly cOI1 .. sisient pmtC'm 0 F (fed i~~ci~~ ~~rxmJlbim~- abo showed dlilt genc~ic divC'~ity in ,"c\',cml the r05$,~blc.~hrmlgh a series or imcl1l11looiJl!lCll, [ion t<lOO as <! n~mctto[l'l ()il" :gcnt'tic dismncc 11<1:; b~C'l\''.rii,!I 'p"<Mlhngcns could be C1< rl'<llm1c1by tili s \'~'(;~or:o:. lrn~1 ~cn~ between any l \11'0. bac- been O~~'fVOO (.Fig.~A) U I). Thi!\ C·tr~L ha~. ~iITlpk- mot;k'iohcl~[rJ~ d.itl,! Un. W ..fer The usc of IlCUll'al mod'cI~ or Hlilltatioll nnd ~cri'l. lk.~id~$ the .i~h;g~lin!>n,", !'l;"(:omhimu~iD~ml b()Cl! I':hOW!l 10 ilc '1~""lCiu[~"(I \'I.'jth tiw v.lli'Ol~!i <in ft is not a denial of ~Iccliol] b lli. a ro::O£tllilioll f!'n1C~~f';lh!lt h,;adl;~o ~nc ,r~cqlli~itio!l fh::m] dism~iC'i~U!li:;;lll1S. ~l!;!l etcet the !'(l'qUtJ1C~' ~jmillarilkl\ t d dml f1uJ!ch observed 1]!Oll'ltll~miongcnea~c :5lrw;:~mc between (kI!lor,mc~ l)l'Cipicm DNA,Ill,rincillallly ~.m~b' rcl<llcd WM-CCS" Ih'C'!lC' is convincing evidence llm~hol1lo]ogo'lls. I'OC(llnnbin,ltiOH mOly M'lItS,.nlcdi n ~r:.'d IIU:is!l'I,.IA.'h rcptlir mld Rcc/\·· C<ln be '~'..'<.P~'li ~~~'Id tl s.ir!1fI'~'W' i h:n:n1_~t ITlillk.CS :'I;T!S~•• 1S.fl I:Iwl!l mUl(k~,~o C'xpioll(:' lite dy'mllnUC'll Ih;-qu~ml)rrt;'lJlla{:e ~lIm~l~11C'.\l,im~~ ~ht g('1JIDIMrL; med il'~El;-.dlJr;:"(:oJllibi inn ~11 J·n- R<o·cA i.s inn:f : umB on' Ileum'll d;w:p.!ificOlI;tioll urul I.hc eonditious of <111:"ICIC'I'imn wirh ihose ~rnnlm!lcr n~('n'lbc'n>I of volvod in ~I] ilirui !!gn;c(:oll'll'ifumniO'J'l bd ween doaer r~ni.d rc:C'i!JicllIt DNA arid is Ihlili 1I.'S,"\iC)m under which ~)(~rllihrlioll;:;; do. ,or lit." IiOI., separate ial ~lllc:s:tIntC s~etL';S 011"mull do:;,c~y re!.ii.~l~d species I l(lj" The linl;!orhomol~flous.iC()f}mbi~~miO;llvaries lot rcoombi~l[qltion, whereas MH~S ill~~I~bi,ISIiC- illm Jb.Qincl gcltmYPJlc dusters l~mll n~Dunk: d'l(! cmc~l;!ll~c of species. Eslimmcs For I~IC roles of g;ucilIlY.~i:I some species il flIPilCUl'S 10 be UilI'C' ::laM cOlllbinfLttoiiD betwccn mismatched scqlleuccs. ooc1liv.'itilabk !110m leads to ~he cvoMion of distiuct clnnal iillcagcs, One I11iOO~lmlisl1il tl~al has r(x'eh~d Ji'lflinicll.~!IIr 1iI111[mllion 3nd recoHbbilmliou empirkal :S1.lIC1ks it vmic,1y of i:t.ftctcria.lc,g.. auclltkm isjhe .Il'e'llliremera of ItceA for mibliwhereas in olhcrs these localized I\.'CO.l:llblmni{mru H'l!iIHy ,clUcicllll. procc:5S.lil,g, :'>Cgllllr:nt~ (ME!PS)" iUlpo:l1S. arise IllL1Ch mol'C fL'l!q.llcmly d1llm ~mJI[aa. Jim. a,~is ~hc Idaliomhill belwecll scqlllcncc dhrcr,gcnc:e rlirrld ree'ombin[ui.olll ml,\: sbowll in, \\'hkh ,aL'l'! :,hm1l reg iOIlS. ::lcq,~lcncc iacunaly ~10115 (j}. In "-'Cell! ye01lt'5. c1>ilOOl1:>ivchonnol:ogol!~ Fig.~ 1\, kM;"l~d ,U! eil ~~cr end ~hc O.DJ1[)fDN A ::i(1,1I1d ro(;(lmbi~~.!li[1n h, .." bcc!l:5h,(ll'wn lO be ~o wid'c~ Wc ,c)]Jdmate(Jpor~Jbdon mll~a~nOll rmc's 'I!~d h)'l11{llho.;.i~,oo 1,{l be n:"{juin.:'(lror rccombi n~,I'''rr~.;·d '~hUl 1M,.}, bo ~u,~1Ifdoo,I!,~III(~ !lOrm miller .. it HGn 1.0 (!Ccur I( I.:i~" 111is. mcchllnls:m c<ln sen(!I'<li~c (dCillot>oo 1I) in tl:lc range 1100 ~O. \\'hcfe.:!lS 1lI.."C'OJn~h,mlh'l: oC.."'CCJJlion. NOBC~IH:j,C£;S. :'i<urvcys of gCllclic dl,vC'rsil}' in ~h~ gcnt,~~1 rc~ali(ln;..hip ~c~lIlln: Fl:g. I.A and probl'!1<l1.iOll ml~ 'idcJUHcd p,' :;'IL'Cmore \"lrkliil!e. \' i,!(j~":'i <I CXl,rolxlro~i\"C' Cl'i~i~mrl;C ror '~hc Ic.r!g~h 0:1' ~l!r;;bm;:'~(''rialkiJw,do:l!l <J,~~1~\~OI~il)gthaI, flu' from r.m:gin,g Imm 0.1 to .~ ~19. no Th~~ V;I~lU(;'~ MEPS ofbt.'1\~'r;.1C'1:I 20 m~~~;O ~l;J~ P<I~m~ (bp) (16). ;,'I, QOmillYllll]l! Ql1l,ldJ"~~od by ~l!mrui~,u[J!.u~ ]il.l;llle!.,'1<1;. Oll1.: i:"'II!~'~~"r:"dl"Crg~nr:;l S(igll1l(;:lltp~'1:.\l.~'IIlt;lm ~Qn I 01110m{~~I!,~nwt'o' Ihe I,Illdeilyiu.g buologk~1~ ml~Wh:nl::v!"~r ~~Ri-C:II<lIni:;;m lm~lr:';di";''i.ti~~i I;k.iclinr..~ ~n clmngl~. bil!el.cri~ ~1,lJ:1III1~O ICIHn \.b:>·k~1N cd ~l.i'ilCI-· w dl.\'(;!'tmtolli"m:l r(it:cJnr~p.~nmi~mr.J~(:;l;(dc:rmt~1t1 III ,lIld r, ic;]~tyrelm(;d ~.tn~ill.~ 't~iK't:JCS}. CIt k~,k);1lor ~lnh~(: ~"I."C(n~lb:ilmniIOJ] ilh im;MI:;;ng. M1(ILll"IIIt;~} ri;spc(,lhl~!)I} by ii, COI~Slfun'l.kllt{)wn ·mi Lhe eJill~ g(:.lIc:nk :sl.~dim:l ~o Cm' (4-.r'i ~ "u::nl~ I:; l.hUb" IUIlC'Cr-· gcnc~, tli i:;.rdmiollship 1'(!;ii,U~l~ ~OJl!lihilil1b.on [\iXombiil1lmiO:DI I~m Opcrnl'C:> a~ the gclllornic ~cv- reclivc [IOpUlilliioJl size IV... sLJeh that iIj = JmVt/~ luilnty ~ruillll£ dlC r...~.tureOr~21~l:Criiill :>~ekniul~ ~nd p ~ 2rN~_ Our cS:li milites {I HlLa:rc oo::;c(1 Oil, d .1IJldlhc h]!lLlclnc¢ tllat h('lmologOill~' ~1.i'Co'~nbilflil~ d. pO;l:r;inl.iall}, altlo'l'\d ng sIJoC~iC'i>i~l Hilu;tru:ss lO gelles .appflo,\i nrmtcly .L =::;500 bp :Iong,mnd :uf \VC t i01n C'JlCn.,> ~Yr()n it ~ .. i') Clncl"';;C' <IS <I d)'llmnic coroUm)i 1.0 dive:I:S~:lical.ion mll:!~ a p~mbdble (';S.1km:li(;or the D'NAlnllli,1rCtull. mK~<lcl<l!p1,mi<m V~·. ). 9 Om; pml~·il,ioJl is~hm sp~lmiol1l (by which I"JlC tm)rul ;:; -.;[0-10 pCii' bas..: 1Pi)~ rpcrUl.':p:lic~'l:btoll. Ahh!.l~!ghllili~ pkt~!r'C' of $1l~i~.lion druvm \\,"crn~n llwg:~ncrnllon O'·pcfI1Ilml£illlly dif'~inCI {1'Jl.wc gei <llJmllpali::. e:slimaic rorth,,:: efT(Xl~\'C clu,~tcrs of cl:O'.:>CII)' relaled bac~Ci'j<Ii) cOI.l~.da.risc, by :l'Coombil'tal~!ilI\iB (Le .• sc;.:ua~) in.ool'lnpai ibii:lity ~ is appe.9.ling, cspccia.l:I.y for~hc p<.IrnUcb; ii, 0 11(1i I:lccm~,~ 01" fund."lrnental ecologic;]:1 oon~ popul'Mion sizc' NI: 0 r 10.7. lelf'S whh Ihe biological species conccp~ {l\r ~ilm:yr stlltlinls, or gcogrd.phksClxl.m~i(ln hut !<lIlla' as ,<I Til.;;! cfl~cth'C IXlllulalLo'H size is no1. direcl.ly {i7~ I.he dm:idli.lllo~ of [111: '1!Jalll~ i'latu\ol;~'deL,,! I of re:1!1II~cd w~hc cel~LIS rOpl.!~Ollion size· blH b rn~hC'r. a !UC1kSl!Ill: hownul!ch :!'ll,;'uLm~di'\!!;'\rl5~[Ydl~ r'\;"ombirm~ioP1:11 incomp~!ibilit\l i~,nly Me: ,~ o ... Depall1llilre"iIill af In~'WloIjJi~ Dlwasl! ~pjdl!mioll!l!JV,111!Ilp!!'Ii'iI~ cmrim'~IUC'~~l c:.,lrIy. tt~l~Y ~. !;lrIJl~ LdL;'!'<~hly c,~n r(lCI of Ihe ~Wiry (or ~t(l~U'~I (I r b>tCl~;Ti. J, spc. Cl!!le~ wndon. ~«ld!o!'l' 2 lPG, u i( W NH'i,uI k"t Ulan 't~lI; cCli~us poPul!alioJ'i ;l;i7C .. ~ ~I ~i1u~ion1_~\i1mL dri ... ~~~~ 'I" :;,tmiL1~ 10 t'JlU 't~~s.c t ... "To iCjhOfl'l oonespondem.(e' s~~u[dbe a ddlrtSSe.j, E·iIIlol'i t ":soli" g'L~.lCI.icjmi'it.Q; :.tI'iu foni1. It~W 5jlCcid!' j rcs.u~t of raeloifiii i>uch 8~ l'C£ul;it bo,1L~r:lk't:'ks, c ,ha51!'i@~IInMrilll..u:.u'k

B""";,
••••..•• cnvimrl'tm::rlffi i';t

!'<inmliittccliIl(l!r'l!h.. ~nll~ iflhC' b"CillOlYf~ of~~I<'lins 01re ,M!noo :ill the' ~n1.'Cw,~)r.rWIITle~wi~h~nbll<C~ t~\ri,d'popul~~io~~ O,l!Te (;o!HIHt'!ull)' 'L;'~'IOI~[L;'l~ZL.i'dl by
mile<ilkks :n ~H:'I'I hOI!l5~~I;;l.iJ;~'}il]lill~~ 'I nililmi 1[~IUS ~q~lcne(; lypiltg {M.LSTH •. wh~1tr eueh ~111C'lc OOffi.".lIm~d s 'Vt'lll d~IITcrcm slX1.lR:~tcC( 1."<I.Wc have d~vclopcJrrl modelin wl~kh stl'iJins Me dcfillCd ill. alte sante' WIly and in which~1 Ic-k$ dlr[~,we ill:

\v~dll an abumJ~l1ildi\l'~[~ S:llly of denor DNA. ,~nd siudies of nhe:

."p,",,,,;,,,,,,,",, n., 00<"

I;,"

(:om:ClWC-Hice of

rcconthinmieu

ffliling

11I'II0re

..am

or

or

or

Jm.

r-

or

REViEW
genome-wide selective :5-\\"~P:s.. or 'hi~ml'(h:il;<ll
T

suuctutc (J.n Consider, IQf cx"mp~c, an inlcc~ i[j'~~ ;:II:!il~1lt ,,1,IIJ;h, !.I:'l S'JW)M("I1~:(,:~~~ p~.r~f(/J.r(Jnif~tJ
At lL'olI~t~i~ro~ 1;1t'~ol'5h,,':S:(Jliil] m~ ~.ITi,:;t:lll" ~ IXJ'PIIIb~ io'.nl i>iz.~ m~m.y o:ltd\:~rs of n~agn:iiudc s,1:IU'lllc( ~hl'l~1Ih: "c~'I.!<I1mllrlbc-. nfb.1JCh,'fi a, Fi~1ilt Ihe i;.,;,1(:~t,"ri.d rorlJll ,~I ~lnil~ divided i111:0 d isl inl(;t p_":!p1I1<1u iomfi~~l,!~ Ihill i!id:~\i'idu ~~Ih~i~'i~tl:l~ ,md is ~:tm!!;l~ !'Il i~T{.-dby Sll'i1~11 iliOotlil!l. ~o I.hill. lilt; 11urnb~r of ii:lfcch..'tI pooph~ i'IUly be a ~na' ~nca:iiUreof I:IO~ Ili Iii!~tl si!tc~umt~le numhe-,ofbac'1criJa.&''''Ooiid. Irm~smiss,j:O;l' is ,se<lsomLI.wil~l ~)I,,"3 occ~nil'l,y k~ during the winiCl" monlhs. !l:rcalil1~ ~(}~IlC'~locks d~lring Ihe ~ow seasen, $10 tha,t the enoct~\'C I)OP1IIalr-o:~~size may r'Cllcc~the nU~l~r of p.."'()Ij):Ji;:o inL\.'tl'oo dWlri ng Ih'l."l'i,c bOUl'elliCCk:;.. n~a'd. ~heh~~

r~ll;l.bccanseaov el a 1 r.c-IC:Slhlll."nsc are unlilkclyw \)pro§I.~h~rjg~m\llly~hro~I,gJl!lhc I~ga~~" hr~i~lo]T!_A ft;,":~\In,' orr~~w:rol p",)~u:I~~i9m! ti,ir\11;11UIn;l in [he ,~I~m\'lJ, rq,):iorl is :.;'uml'lg gl,;'r1ot)'P,ic dI!IL<;.

U1ClIC QI(""ml I in

I hC,,",QOIl1I

P;:!!l,}'.~llg

S~!rPQning

Onli!N

Mmi,,"I:ijil~

(.27).

[)ivelr~ilJ·DriveD1 S:pecmatffiC:l!Iil in Siexu .. l~l:!.:I.cteridiJ

~.I~pol}(Jlllii t£m~w;d~ hi~111ml";.,.,,or ruoo.nll~.in:'1t inlJ. Ihe !lI.>dUCl;..,oOl!1.e Q~'~""om'bin1ll!i(ln bct\\'ccn ~wn r diQ~dy rebh..:-d ~r\,l'(;i~. co:rl1ip~~!\;'(I~o ~l~,tlw~~hin

fF~g.2. A ~IIH.I, B}. Thtl~t: d.~st"::Th arc unst .. ble, ,m1.>U mho Imlf,l-lll;!nn dymunics <Ire d~OJ!!t[,:,ring

,1iI;;l"rizcd by a cn'JlsI.;'U~I, prm;:c,\;j,lill which !ll!1~· jor cl u:.lcrs rcglll~arJ.y ~J11c[g.(l by C:h:i11!1:C, l\'I'j,li[, dri opm~I,,~~l1d ('VCI'ihll~~lIy IJCCmt~.c cxti I!.et

(Fig.2C). \;\ll1i!ul roc.cn b:buu.io:n :rm",-:; are hi'i1CI\.'9s~d1.0 values bcl\\iOOI1 m»:lour:thlIHlcllWi:cc the am1.iil· rien rate '~I)i.'ij· loeus), a tlu:cSllok] js~s.">Cd where chlslcfli, 1]0 Iml~r diverge ~UD ,:1/11'1:' ~oll~!1<lI1t:l)' feab!1orocd i:mc dw 1'lJllC'nt poaml a ~lml by the

~p.cck.~. 'l.'Q'If.~dl,;~ ~I ml,;'dkmi~'i of l\!:.'._"~ljtd, P ~~'ial 'ClII:I'i it1ltlilt'Llll:~n .lie lOC',pm'3linn of ~ecie:s.. bul it ~.. ulld~tuI" whellll~r tlllcreJmielllSchip
¢u;:h

i~fillliltio~i

n~lw(1r'k .~..;, hiea,~rchic,lnY~lruelul1L;'d i111J.'1oonjJln~n!iln(~. ~::m\~mUlli~~J;o:r l:Orl!U1Ullnl~~ Ltt".~

Hi!KLU! COI.1WU;t

cohesive rorc~ ,of roC'OIl'Ibinil!I.i.QH. AlI~ics (;"m .. :>lLCOoOO du:ough borizow!ial ~1~Old cvcu ~""'H~n
d~gl'l'.!;.' uf c'l.lJIslcr~l'Igis much rodm)~'!doomp'.rt!d with (he
~hi2' p.;tt~.!'lil.al I UIII!"~'lg.COi;,-;,; nt'll. The d

lOeIW";:cn, i.v!!~~i:lre ru~d roc:o~~1bimlti:oi'i,rnte b: J ::rufliici.cn~ to caase species to arise b:lol dri~L ,In, ()lhcrwol,d. s, is :ii Ilim.tlsi.b!c lhal. chaace vmimtol1, wo uM OOC'a::1,i(l:I~JII)i 0:('.'>111 h h~ S! u:J111:\, ,ti fie-rent Cll('iugh lirolulhe founder IU1Pll!'~rulion that ~hcy ~KI .tang<.:r ro(,'Ombin~ \\'i~hlhc 1l.'lW!!ldi.'1'S. oflen
t:lhllU.!:lh ~.

k\r;:(nl;l~

and so

O!Il. W

liml Ihe' enbc~h.c n.um\'lC1!'ofp:C{lI'~C

31gJJ mltril'lll ~v;li I'Jbl~ 11,)".and lQclIl ChJS~efh'lg or rDf'lll:lIiDlt" areund :'lrn:!l1 p<lfli(:l~\" of m,!!lril1ln:;;. ,t ':"n. 111 1;'C'lleralH1!lO$1 II<!u,U'<L1 'POPLI bti(ln.~ or blffLc~cria [IIYe ill :llJtructUl'W envl ro£nlllell.l~ with wcll-deflsed pa!e'tlcs of £oow~h. W~lCJ'l!serieus Igni~scxi=i.l" 01] the dispcu:>illn or nnvel typ-t.--:; b~"\\\i;'1.·np<uch~'ii;- ['"I;1Ihiisi]ing pbuibTr;; esti!1Il"h~l'i for N~ lor ,I diwrlIc rm~gt Orrni.Ct~'[i u, .h \\'ell <ls lidcllt:i Ijr.i 1IJj!i rhe 1:'1cH')n; I]ml a IrC'l'~ ~~. fOhcul'd bClI I'C.'Kifurrli pliorily_

inlech,'J;i i~~lw~r ~1,1!] Ihtl' a(~l!a]1"u~n~C'I~OrI'MJ(lI~l(; i!lltbcwd 12'.?", UW'l. JiI pop~d aiu!'l of !Ti Iliiom; .. P,lCll'riu C''ll.'! 11OI\lC :JI low (l.f~1,;~Ei\;L;':popul:niuu! !O i&~" Sin.'! il:1UJ' CtiUIl> idermions mill)' i1m nWlcncC' ,(;f· f(le~ in::" popbli I~D,iOi~sizes iiii rl\l!.rt~! e'li.\ ir,OJlunCI'iiLs.. such as the Il9l1iliOlliU1g or marine bacteria 2wDmidnlUiricnHkh coosm~ It:;gi.om. sCfISOIml re,[1,l!III;Uioncaused L'}I tJhc"b~oom-lulSl" cycle of

or

clomd ~"ilm~!on Wig, 2'. D OJ!ndE], <1mJdyn :uuic i '1n,lIy~ig\. (Fi!il- 2 f~ re\'cEd~ I~m mlwdu.~ lC'rlllJil thi1l1 dO(" OC,(:ur i1>u!1lllsi~~llt. ltis wOI'it.ioInmil!,g lhm. in bolh situ13rHtlJi&. 11:1(: dcgllCl: of d iversi I.)' at each lo eus is '~~U~ame sud s is go'!;' erred bJl uhc bal,mce bet ,veCI1I CX1incti on ud muuuion, The scxualpopuleuon contains mere d~::;~nncl gCIIDt}T~, based on (]irlIclICn:l comilill.<llions. of aslmilar mlml~G~or alleles,
R,ooombilm~ion

~f;:'rIClit;.dl!y iiiiOhu.ed '('0 ronl~,I ~~cw~lJ;Ceic.;:·! l.!f simulanons !'<lQ~C'~~ l!hal <II~" O ~h{lugh~iri slypc of ·d~i'ln"'!1ec"'!I(:"I~!(,Id rooomrbieJ!mi(:1I] ~<!H k'OJld ~p !he Clll.crg~Il.C~ of $~P<I".JLtc popl!ld~~~im1.";. 011!Y OCCli)':lO Ih~N lllldL;I'I,;OnU~luml:s, in
1\,l,.Imt~I,,"~II])l

!lIq,linlain gt.:'~J.,!~ic plO"~milY.

,~lld tll'U~

whichllll:

ffi(\Ollflbua'ilmion ruCe declines

Wll III

law of al~ck)l a~ one locu~ is

A.

i:,!,tl:cQ'LlC!H

1;,'1l{lugh~hal ~hc D10l tied to thei r

d~'VClrgeu£lc more ntpid~y than is; S1ug,_,i!).!smd by c>:pcrintc.'l1Imiol1l,~F~. 2. firmd ~). For V3111~'Sr o Dhi~ dcclh~!! cQ1t~istcm with f'ng. I A.. we did riLl" observe distill1.ci. POl)!;I~aliom, Cll~.c.T'gill!g in OUI: si nlll~lilth:m:ii, bccasse Ihe amOl!llll or ""<lriabiU!}' withi n :5illmblrOO POl:'lt~bliOll~ is roo low l:Or dlC rotiombi ~~;u'i.f.ln rnlc~1IP Va!')' 'll~r!\.'Cj:d)~yFig. 2. D l(

.. !
.:I
!III

c
1
~!

.,.

."

,.Iii

Tocxpku:'C :sp!.~if~.iol1" !,vc c'luC'Mclooour previous i1IIod.'c11 91~ to simulinc shnpHfiCdgcir U ~lOm;es i(,Fi,g. 1('). whh r~n cl1:eclavc popuh:niml
s izc N~ • ~ and a popu btf.on mUla'.ion ,nuteH O~ 1. i.U1d dC'lined cooh 5tu~ai~~by the aUdo's a4 ,.1 Im-gcr mIlllOCu of loci ~'0) III C1J1ll1lH.1' the C'~rCGI 111m o(Xasionmll'Ct",,\)I1il'hlnalion ,at <L Sllll;lIIC 1001ls iltils n II dist(lr~i ngucl m ionrshilYS bCI~re::n otherwise divergent or similar sirnins ,~1j'~.Thi~lnooel

'I
J
.~
I:!;.

a,m,
t!liWl 00001 D'% ., .~~IIl'Ii.I~ '" ~

:-: 'ObXWtl

$.gqg~'I1I~'Odl~t!!'rl'l;enS!i;!

ignores.
~I'lli~n

(i,'C\VCI,11 h~'!I.:oog.~n.eil tll~[1ll'1)' a!'ISC in I~ p{lp~I;lml,o:r1S «(!.g.. .• IUlm~:;;~, c~o~og_\'" and :L'tl(;{mlbi.
a

pmvidl.:' ,<I J'l'n;lijm~ 1]<11)' ~t:rif¥liQn 'Oor lh!,; gcnL)jJ.'a'[iollof di· "C:r,.;;ly by ,kiil ()tIJ.' dlQic~ of Ililr.unt1!Cr.-; .nu! n1110dd :;lfi.llCll11L~ i~. iin(:". iwbteoollilpa-Ol'lll i;';e b.::~ 81'1 1,,,eCfiI l)l"~I~i,biilil.}' and COml)lU~m,iomll, liun.i~il~ lions. achkved l)mindpillJ~y by redrucJlng Ihc cllectivc p()IYLl~mlQ:n si3e and! b)' us,nl:lg an IIP'p'roXil~.'ifUi 1011.a~.!!;jo:riihm notieling, nllmld.o.n of DNA SIiXl ucitcx."S t 15).
nlW'~ I'I1I'L), .lmm;~J.ll;.d(.~li

hut

::11.. (11.) R€:cll'lmbf!'!o1!tkli"! nnefClf a nmg~ of related dO!llo,~,. as al hl!!'!ctijolil !(jrft,~e IfJroportijoUQ 'of' seCluelliQe lhari: is dUrie;rerlt (~,~uJefi(e divergenl(e)" mot <lI1vari:et.y 'o~baderr~al rredp"ien,ts" Ci'rde, 8adllu$ sut.vlilb; ~~y~~e-, lJC(ilM moj~flsisi d~~u·"Q!'1d.$, pfle',gJm~n,ae;tri~f!.g!~. fs(herid1rao ,~Ii, The b~t f!~ tog-~ine,"r (JI.!Ne liSsiilOWIiI, Willhin,hH'(:epl: O.S%alliJ slllpe" 19.8. [D',ala a'j,~ rrom U2-H).] Stopeos ((lIF iincilhdidulil L l'Iam~dl sp!1N::iie:s rallge fl"C11111l 7 .'9 lor S.p.l1flnlnOllia(t to 2.5.'] iorr E. (01i..r(B) G,elnome o~ 1 5.. pnel:rl1:lOniae [hom (35H "l'i!dlloc{!l'jj~r1 air ~e'M15T g,~fI'~s.. () Sic~@ml1ti( r~,pr@~n!~ti!(jJl or~hE!! sim( lulated genCllme~ilil ill steclhaljlk l'Iell~lIl. l'IlIod~1. MI.S1 ge:n~ ,1Irt'lhig'h~ig'hted. ;)~t>~.JC i8Jtionw~

IFig.

Lhallckl\,'J'U

fJllhl"

kK~i.

~11

I'he

ror ~

The

ClCHih<lil·SexlII<lIll

'1Ii'11iI1~eslh(llld

<II. S~I'U'jl ll~ml!S[1 ~on

The most $a~icn~ fe.aluJiC on'll is S Imrie modd b in ('lOP'! 1",1 uon :'>~I,l]CttJIru W II h incfwsi ~g rO!lc~ (I !"homo.logo'us rcoor:nbi rmtial1.
\;V~1cn !'~'Cml:!,b~llalio!l ... L'~~a!"c 100w. 1.~1~ Pp\I· P

clonal :-;i1L.I~[ion.iu cnu't;m:>I, clU::;!I)Cl:,.,regllllf~~J'y bcwmc c:O;.tilll,c~ I f!'~!l' lC), fIIU!!:! (;;0:1iU.cti:OJ1 0:1." cloll,e.s is the ]lill'lindpal reglir~Uor dh~oC1:Snly as a \.~Il()~oC"(' h.!lslerlin,g CIlil, be: dellned 3S oYc:rdis~~IDn Ofl~lC gene!ic difi.<bllloCeS 'bctwC«l isolla~c.<; (r-~S. 2,. B. E. and 1.1), af.ld EI nlCllSlIl'C onhh;; islJlc in~de... (lr&~ociiliJj:onl (1(1). I ill C:Qj~Her . wOl'k.we showed how t\) calculate' this, index lor 1~'lII[m1 mode~s ,(willmlill I~'nc(_kpendencc .on !'cqm!llce: diw;)rgcl'lcc or Fig. I) IIQ~ and we

or

r Ii,. rll\L'i, <lll!Imlvgh Ihis COIiIC:CPtU1~1 :Irmx'td i!'i .1p'po.;t~lin!b .. iit ~:; not 1iuj}JJOr!l(x! b)! l~'i~ q~lilli.lita~i.,.e dCI~i~ OrDll~ irHCifilJtl!ny bctWOOIl g,'CfI~i:i~ divcmi'I:llto (":;ltL'01l<1m:! ~;:o,;.lltlll:kwl:lil~'}j~.

~,fl'Criimellla~~l!:dies o.f ttl!! rclmi,onsiJi.p be-!WQcn scqm':llcc d hC'l'~en'c <Jbm.l rlX'Qmb~na~['Qn h<l"'cfo,cu~oo fin i~l~rspceinc iml1.~f(:[ or [) A,
dml,is. lQ'I.\'ec!ll organism~,>that m:c uil W -20% !lHwrgent and <l:Ife<lllre<!rl}'['Ircsmm:-d to be ai, least oonlCwlm~ se1o;uIOlMyso~:I!oo. For lhe prrw;;~ss or i ~;:poolation n~ooeh.'d hCI'C. we :lP~ initial ~y inlcr~ 1.'8,,::,0, in the ~]roc~ i!llIa"P\""JCil~clrarl.\O.!ly, :;0

I;lJt~on.is ~:['I:1;:clnV't::~>',domd .ill ~~:!i,!lC'hll.rG" .~n~0I1fi~~

!;CI1~t',cllch el!l:l,.l1l~ ha~ ~l ~q:iml!ll!i: c\'otllt~o:mJl)

h.l\'~ ~h:(l!wn [hal the lhllC'~I1{llcl b;elw~l ,donal arl!d ~~1!>u~llro~i!1l1;:s; holds. for ,I \Vidcl'<~~~g",of rOlr.,lmCI¢r~ VO)" The tm!l~i'~i{j,!a htl;tW';:1L':ll clonm.l al~d :!IA,"X.UlII]'iclp!IIl!miOa:i sl~l!ICII!IIDCito;>'~~,di,._!dlill j i

!h~ n~QS~ irnport,l!]'t an10unl:\iUr!'i~U~'~OC

how \'CI)' !<l1lil,IU di\,.... ~HC(:, I!IIprn 5% ... ni"l;:l l't'OOllibi ni~Uljon.\Ve .It.iiiO'\''~,h3l bm::t(;(ia n~'i3;y" iiI)'
qUCSI~'Oll is.

or

4177

REViEW
1rn:1)'. mIDm;)jli~ QC~m such .. 8 the: Iq'll(lflcd I O~ I"C" !crl.. ,,~pm'!I~ooby ~,~n.!i!!!~l'C\:(Dmb~~mion c\r'C~IS ! in Iheir m~h"nisul" !Of rocombill;ili,iol],and ~\I;."!·~n~ 'l>c~w~"n !hc ,m· ~~~!N;~ihc 1)t1U~nl shm'!n i~.fig, !A mOl}' l1al 00 d~!Ciion if! woorribi 'mt~m~ !¥IJt~ ,(by ~~h~gQ"mcdi<ll.c'l and (:"~QI.d~~~h~ "~I I;;J;~~.·.ILI<'ImI~t .in."ot,;rh,:-d[LUd~ (.W), h~ ~llG~~ri,."'"~Wlp' ~1!tf'[w~n S(lJm('}JwN~ll.-'Jjlt:lir,. Q ~~mi'ilr I!H~iV~I~~"Ll,[u .. ),1;)1I",t. lor 'L;'''';:IIH1]pl~. ~ din~vl;'li!~ ~"Il~fod~I~!i.ml) po:rl.il]~ ~lllml;:il;:nl ~~q;lsof t;~qLlCnt"rl;) di'!."¢.lil\i~}', TYrh:illillt[lil~i1'tlmtd S. l!11,~(!r:iC(1 ti.l,:rtwmr Typhi ~l:ilt r=lalitul ~!.t'llCI,;"n gli;;":nI;li.~ l;!is'i;lnci; :mn~ '.·'~'~m~
bitllill[JiI] J.tttcs hUt;, 1k~~11ol):O;~rI C'J: (18), in whi,t;~! m:co;rI,~Y1J)o:ui

2.' ~ clivl;."mg,:irIl. (8.!9), Thus, ~r{)n::

~hcn.""JCOTnbW:1aJioll :m~e declines

~pt;ti"lion Q~tu~.·i~g by d iS~'UH;:C·!>iL;,dcdrc· 5 bf~,),bctal,l~ of ;ll'iii.... chaniNI1l1,~llktd ~o I!i~ Mul S COItI[)i:n,tt i:Oll. del;;l.ils or Ih~ dL;;PC.DtdJ.;!U<;~' ~h\.' !iI'iism1U~,eh-r~f~!~it ;Sy~k'I'iiI, ruacl 1],,;:10 Ilti1.:{'h~l'ii£l'i1 ['CCoi'fi'il'iiIlU'liCil~rul(;' O~l ~l'qll.eltee !:I1\t;trgclice ntilitiit

f;apidl),' lonh~ lin"! l:Cw qnii",m,ltll;:h~~ (~,5%n;du~lion IOf

"'1.'1']1

0"

con~~u~iof!s 't;'.m'be ro..c~~ <l!l)[JlIlllilc

fc,Ii'iihility

or

11'1~~~j'ill!f.;,ldl!i S. p~,Wlmw :lind .. m",}, In.-quc.nl~}"~h~ghl}' div'!.'f"I,!,c'nl. !hal ;!~ >5% This ,dL;'~Il[Jlh"~!1I'l;;;,i that, ..~ JGa;;;l 'vi1.lii~!i t>m~'iL;s~~i~, ~."Ir:I~t-;;i,\'t ~"tIUL;J:l~ di"l;."r~lIlIeh ~.:-:N{:i'lserl'(l

'I;t:k.',th~

dN:lii1l~l.hcre~1'lk. ro~·· l:ic knW""""11. (H). lews fi lot.~liul~MreliililJn:.ll.~p\,'ei)' ~ inni larm IJlm. Usinl_g.m.0thoo:; based {iil M,['ST (1). We can S~l in IXIl:;I,!.!d~ :;D~.,cli;oo1,0,clme (I:ig,_ .~A I, Sim.i~ itlcmii ry $Iro ilils from l1illli1~.IIXlo~bl~lions. of baeI:l'itiii
::;!U~.1I~1~c:il,:>O ~1:;1.'1 IJ~e

!!jr;;I~t1~iiii 110

r rcu)J,'Odllct:i",cfii;o~ l'Oll m

lmr

10 ~mlll'ibilllfiliOil.

MCt'h~liiism.'i:;l
d'ttli1

ether

sequence

t:I~Vl.~cn~e {:c.l'lai il:l~ exilOt. );jjd~ us 11Ibchcd increntlarien, ciifl"erell,ocs in DNA .. cxeluli1£:c by

'8

(['.Ui'll'

lliJOol

QM

o.Ot!iEI

(I (I

000,000

1:000':000

ID

!::.
,;::;,"

!!.
ca.,

3:5

il
W' :::I'

~' 'ill'

~ m,

(ic(n2'

,00CIOol,

!ll~

tlO(!e

0<

500.000

1:®oo,:ooo

IH

70

'~'

35

Ge.iilerallil:ol1i!s

soo.ooo

(I

1 ,000'.:000'

if'iig. ,2:. Simul.a~ed g,ene~k ~irud!llre CI donal POl[lulatinlill ~A tOl () and SE!'J(ui:ll of pO;Pidili~iomiW to n ,All p(lP:ui~ililjon5' are eJjIolving under iiileuual c1rii~talild a~ ~o rnO'£le1lIlOOlJI51y midllg, (A. [I" and G) 6em,~~~ 1l'111iPS, w'hkh a r>edel'Elrmllillll:a lby priltcip.d 'Qoominat.e ana~:!f.i~sUti):, ~res.ent the 'gen.etk ,cl~taii1ll!:e~ betwe~n ])000' ~@!1d~ml.~Qhooeo isma~rroITlth~ ~~t1'lat~~ p~p~latio~ ~foo'f 1.0ij '~!liIlllration$ haw Illap5m CJJordinaOOs;tl!rc eXlIlr.es~tiI ilil IIImts of :5etilUiell:ce ,d~verg~.~, •. (IB, IE, and H~ Alii a.lt~~l'Iartff¥elj.'ilayl'O n~pi'eSlei"!ll d!i1Sll~ng~:s '~e lci~s~riil:lllltiOIliillf S@t!Ullllt(1f!' di,llIeqJ.@l1oClf!'oollllll@1Ii IParil'5 of l!Salaoo) 'illi thi! Pop:lltaltioo. The thin ~ines ~h(M~he diSIi'l!llCi9' selweE!1il fiw rande:m :';1irairn ami all ~E! ather 'Sitr~~mJ!i'ill ti'l@ sg,mpf~, 1""Iri~n&M 'i!~t6iJi(ik.rtld ~n~sh'O'lr~!iUi!@ ~"s!riblJt!oil! of all till@ pi'lif\ljii~e[J1!5t1ll1l(;l'5. Where t~ere ]s,liiltle ,dmte:ringl {El. an poaif'l'lis.e dlis.t.uues are i1milar .amilhe dii5tributiol'l &aas.iii :sMtgle peak; \'lh:e:relhereis'>'lJony dmterml!jl W3~alnd HJ, ili~ d~slrib!JIlioi"!1 ha$ mi!.dlti~le p~ak~ oorrres.pO:lildki:!;I te Prainlitse 'mmpari5Vn~ ~v,1th~Dland beLwem cllil~t.41!r5.,U:, If, and 0 [)i.:!ilribulil:m of the paim!U~e mmparWQ~u eVi:l'l'li'ingi over 10' g@,liiieraltio~~" o norlmaljz~ the dis" T tnbliit!:Q!'!, [pairs

oil ~~ n1i!91. (F) ~h(ll'l!,iS iM:~~d im(lf--e st~bl..e pcilptllali~m! $~!llic.tldlfe, '~tl1i ,~ :Mable: diifr1l!$e' c~l.!S!t@rIll?frJg rTI(llfItain@di throl!!g'hC!1.!1 'Ihll' S'mllllati()l1. f'o\lram~llar~I.u@~ ~o.r I) and p., lhe popt.l~ail;:(m mu1li'litiiQIil i! IiId IrllW mbi'n anion ra~eb, ilr'~(oj< ~ 2:, p ~

dlirlE!rE!lill~ be:t'J.WE1'fl allld 7Q, ramh~;rtillill'l for ~e propo.mol1l of baSte paks as in O (1:1), (Et and '~IfU. he ~eig'l1t of the d·irSl.ributioiil ii$ r~p~ented T by color shade" r,mgiillg from b'lad!; (Q,O~ ttl red (::.O.1}, ~o that Ilel' k~ illl (IE!). (IE"" and (H) .. (:Qwrespafl:d i,o red $htltledareas. lAli (0, (flo" and -(Co and :$h1)':1li! cllJsiers; rno~f!g oilpart., vi~ib~~ as Flldl ~i!lks IDOV~li1guptnf©l,l:gh ti!'!"uol. Wh~.r! dJ:j~t@~s ll,ti!;. s iii new IJilMkap~rsll~ Itl! bottom,; cxtim::tiolils af!! apllli1lrllnil him; PI!1lJKS d~

m.

0.01 [00

'ro (OL

,~rtS()lla~s

,alf"e

'~~rnpa'foor\~r u~@ nil~b~ o:J aUaes~hal

o1!lr"@

~flt We also e.xpiw,ed IJ!l'Ider\llhkh o(iorufitijons diwsterilllQ cOIUIM OIi:il:u.r ]n the prre~n(je of tllgill recombination rates [[(G} to (0]" (h.tl!5rers \!lith high \lI1tmdi,l$~r reooill1bili!ali~!iiI '~I!'I be 'geli!.eral~dl mimkking~'ponttalf1eouS specioatn(lf1l [[(Q ~o (1)]1, bllJil ~eqlilir!'~ that rewmbin:illiolil Iralt> de~~liIes as 11 ~i.mCJtilon 'o~ ~equeiii(e ,d~versJen.CE!'ai~a ve.ry ra pid rai~ lInoehawaoteliis.tic ~f nliost lhig;(l,eri.al $lilj~l;edlt(ii dat~. s!!lch thllt p = ,2,Qi X !llOl-300i0',

= ~()i x mo-:Ji&", il'IIlli!@f@ ~s~'h~ x :SiNIu@nll~

dJji.I~fgin(,

~(I)O~ 'l\l

478

REViEW
ph<lgc-mcd~a~ed triln:lod(~Cli 911 Q\">'Ulg lei i~OOmi'r;1!i.ll iii l)' in ;5;lI:~. i;:1Jp~~bii!}' !!CIphll~ h~lt.~ or ! ~\lll
fC'-'lncriml-n~odifiql!~,o:n ;lys<h':[IiJ~ Or (I in~rt~Il.c;:"-".,,in INllk"foJ1tifJbillity ill ~po!"'W' 10 hormones U i . J j I, These 1'!iI!JC~,h~~i~I]Q_~li1Ia~'I; uo.\ YL.~~ ~lL-enil1\r;l'1 icah;~] i~'i Ih~ pn:l'~~;&" r o kt~,.c~!C'ri31 s:pce~1I1iOli•. bUi I. dtCir ~Ii.'i· Wt.acl tOo~d~d l\r'Orolmd. be
<t:IlQ
\\"C

h)ip(II~h~l~11~m

,<I

rnecha-

~Il sm

m11,~!"C~ 'to Sq'llIr.J!!,I;;l lh~ ~W{I

~1l!i;.'" t'illlimi etmlcl be con!!>idcr,,"tI iDmlt· ~I.~!: tlHCC' 111~ 1)!Clpll!0!~im~8 have cl i~'cfg~d'i.;rbcHlgh Ihrbl~1din!!l by !;'Y1T1lpalrit~'tl(mibim.:tiI.m to be lSI!.-"!lel iC~II:I)')Tllpo:.si.bl.c. i GtlJllll.dl!:~ia ns

oc aln~idi;I1tldllll;l~q;:~lil:;pf;."Cks_

pt!lp1,d[IJ~i.rnl& ,~m;l~lm~ thiJY OCM,I!d:hl,ls I

:51.ow Allopatrk
5!~:!i:u~[ B,!l!!(~riot

Sp@d.wtmon

if!

{,d IOP;llry JW u]1!!);.\dWla~~mi:!;vit,!;l and roi,.... nirnbin:,QJtioll1 bi.'t\\'L,,'('Jl bae~(ll;ja. and by !01naaH1 aoclmu!lla· ~inn of nlU!<u~nn.". ~\,~'Cl ~1'1.01U·.t!1,,;'d P C!P:1IIo:11tit}!L~ ,,~ iii g.i.~ll~1i(;,,0'l1l:f .cli· \liC:!W: ;11 1(\\ i("'C t~!C rmll1l!ionmt~ C1m)l As I.lrlis hUppt·ll"'.lh~ iliUilfl~ ~tC e~\Pm~ily fol' rooomb.inaiturl betw~n d~c populmi:ons is reo duecd, JIll.! queslion (hen. srises, at whm poim, li~IOU~d lhc}f be ~cnned SfJICt~CS'! !For wxt!JI populm i:ons (,11]('1\'0

ran'. we h",v,e COl1."iacred ,('ill~Y ~hlo! C:~"C or a ,~ills:l): PIlI)ulmion. Pw~lrlng:~'cl Ilh)l~ic ! .!1CPlllf.1l.tl:on n
So

Our medcl is an o\!crsimp.lili(ld caricauee of ~cllclic '0uvcl'Si:ficabton, and S~~Cd3Iioll 'btLl, nonetheless: ;1\'CS ~onlC h'lS~sht into ~hchllC~ f-ll<Ii)! oo~\\fc!l:ln mw;uion. n.'Cmnb~·

~m,iGIl.. :.!nldg~Ill.'~icdij\'~!l!,C'J!Ce. Fim'"

I~!I,' 'l0l~~' or

ii

•
~

~, ~ t.t;... ~'o:i.....

. IF~'I. 3. Gen~li[ m[lp-~'o~a poPlltation~9\m~ordiri~y • dli\l'id~dl Ib1 ill' strong ~arlf'ietr.Wi~h p:illrat1ll'~ler$ asin IFig. :2 'ror t~e :~'~}!U'ill OPlJ1lallif,m, i!!>pmi§: P lll'ltr,()dul::e~ <lIne'f 3.01),,000 gelleralionlO (A). Alter 300,000 generadcm; <lpart, h~ p,opulllltilJlnili, have driHed ililildl are del[lrly di~thut. U!:). At.uhi:~ po'irri~. tlil€' popw la~iolil$ a~e rel;ll1lited;a~t,~!" 10.000 g€1f'leatliolil$, Ultl~ distijl'l(ti~:ll ~e~ ~h~ eruical "''COmbinatim~ Ih!1l.""l'~!' ilfl.i!im (C), OiIntil @ fu,lh'~r 1;0.000 g;lmll~;l1iiQn~. IlQ !f1emli1.jjlnts-Q( tlil~ old). spcci;]l~ioll can be said l<D sepalral~o\ii1 are evndenlt (10

_,e_ "'_

di\fI@i1iil.y g~l(l"u[()dby nl~W!lmll drill. we have di!rivcd a si~npr~ pl]ler~~m~!lo:r.(l!gj' ~~~i~'$. of [r ~"!oX:o~l1lJ1i il)!~ is ~,L;.~:>(:onU1l'tlll rr!,u~
11i!~1!1nUIHi! it'lill•. [~bCt\:illl<l,tnon
Ji.Q,

i~

("S-

~n~r

i>cl'U.i:!llllly!rum] .::tnclDh~ p<I~)IlI~ [{m. C ~lt is ~h [~lillt'lcriZ.(_~by :l bigh degree of ('1u.~&el1n:g. tn litis, case, we C'Xpccl Ih;]!~ .flll.hough lliLllilrn.~ s-c:l(!Cilioil Hnd • gcogl.1lIphic $IirIllCIUlI"C' win act U'I un~l'!.lcl1.CCa l'IfOCCSS or c ~l~t<c'J:i lg lhal, ~ nm~1 be .illhl."I)Crll liO dona" pOJll~!<lllions. !;h~}" d'o n0.1 OlCllli;l.II)' cause Ihe (Ius~erill£, lr 1"CC000ubumiliOlil is

).

tlmvc !()clcl!fredwhcn~llc ~ions Illi.i! W blcin,d even

~hi;~l ]~,il'ib~'CIl

porln~a-· .~rDhc: barrier

i~olnting

ll.:mm'~'Icl_~r Ihi:' I''''~C' 111 whidl !WO pnpUl<llDiom ,t:"O'I'rI (,'xdllmtgJ:~l;:Ill,;'s ~llCu1d1'i {mille' ,gt:Jlt.'tic di~lfIIlc\'; ~~W{;Cr1 the MPIIIIJ',-ulions, 11i.(11~( ~hi~ diSltLnrc il;~low !1.IJiresli.old, rtx:olnbillmuol:1 C'flil cause dist~nlCl PDl:mhuI,(m::;, to c(mvcrgc ::md blend, :~[on, the ol.ber haad.this gcnctie dii:S'llint)~ is above ill IiH·cshoM. then rocomb'h.l1uiom:1I im:omnp.1tibili~ be~wcclllhc pOlllllali:ons is suc'h ~hm l~ne p01l1!! n!ions, can never bk~I~diand COlI ki ~ lc£itiln.Mc~y be consldcrcd diSlhtGI. spec lcs ,(l7),

Th.u:i!.,~bc degree ofdiivcrgcllcc ifld~{';ed 'by il~~ lopal ry 01: olhcr Im,"Clmni:m~s, sqla!l.·ation .t'CQu~~1 for spcciali,onlo 00;:1.11" :i::; :~10\l a CO],IS1.i:lJl~ but dcp~Jld:ii oll~h,;; I.IIC' of 1;«om'l1l,lal<uion be~wCt."'J~ ~ijmlll'i!·gl:.JnolYtl~. \Vhcn sCp'U<lI.ti:(an i~" !1.a~ -~umck:ll! ~D (;";;1~1~ :5,~)!t;;"C~lion •• m<B :i'i)I!Q1P;UI')" i.. ..

or

r:or So tJJiHIInO'lIt'ue;. the ~OOllil iJii,nal,ion :rmc has, been lo."Slhmltccl as r-<JllIghJiY thrcc~ inlC'S ~lilC ml!llationml.c (perlpcu~.) iJ'f) l,~~liilt iR" aoo\·\c the clrm'I~/~."'bII,,,1 !~g~hold, ~Q i~ $h9l~ld b;;;h;w'C' aM!
:;''';~WI,dpOiJ'1,IJ!o!li{l!!_The d~:>t;mc.; I)Clwf.)Cn sp~'· ~i(.,,, i!'; quill; '1'1l1abk. TIl'C di\'~rg(,':I1'~~ kr,W'I,'ot,,"lJl 5;_p,~clm~rnr;'(K·arldS. '~mli.~· is > IO'X';Oll dOL: bll:s:ii:"or Fi'g, lA, we pn.~~muelml t~lt'![\:;~O:fnbi· I mnio:rl iii:lC bCIWC'CI'I t~~m u:s ~mppi'CSSi.--d by Il

di\~C"rgc!1(eoot~~rcl1.:!1. isolates, ddt is exI;hl(k.>d bcC:~IIIIj1C is linke(llo il g,cu,(.'g dCWl'ilni!liug rt:l][.c il ~i~ln.;:..i!i:l;;illll;:(:,.which umj~·':!Jo unL1;'n'p()cil'it m~!liiif0t more ~rexllll:Cllltly I.lmn ,o,t111:1)I~hi~i!S an ( il1i[('R'SEilirg ";:"i:tI.l'I1pk of :;.cilc:t'li,QJ'i diTC'Cdy n.rrl\C'~ilig the g,cfID'lLc imcmlkll.CclilC~ of populatious, :rn.~bc:il III one :lOOtJS~, Named, ~'!lj)ecie arc cUl'I'Cfld,Y de, s filtc~] bya siriet ~'i'C.ries f 11~leuotYI)ic tests, lind o these hldcod cOJ.l\1.."Spondn c ~1Iir clusters, oi'reI latcd bacteria. ~:lowe ver, IiDesc ell!! SIC rs <l1.'C not unifnnn: for C?"I111~}k'. ,~~ pm.'1IJ1ltmfal;;.·us less dlvC!"J,'~li1l!h,nl!iJ.c O'IM~~\r mn~od species, g

q~!CJIce

more I'b:qllCnl thilIn lilis. VItilfCShold ,000llil eeembiaation SI:lIil~ 10 act as a r (oil(;';.;i\'C rollC'C rm the l'MlpUlhI!jon by b~1C;TIknng
is 'C~~ hl]k"Mcb(;I\""o~m ,il k~l~ and U't"Jt1t~cing ~"'I'IC~jc::: c'lu~[l,_"min~. Ul;hrn :sillLlnlron could in plinl;ilpk: S lC'<lcl ~Q dyliilm~e s:JX'Cum inn by r-h:rUlCC dri II, bllil only ~rh~ ;:m10ilHll of \'fllri:rukllli \¥i~hililthcpop~ I ilI~fLl.kmis SllflTIc~Cllt for It.'Combimlliof:l rates ~o

vary ilptlrrociabllybchvccil members ofthc POI)= utial,ion.. all the basis 0 r CIJr,renl e.sll mm.cs lor I.hc s~ics we have studied, thus does not occur. b~l il slt1,]\lk~ nm. Ix:: ruled OLlt. l'lU~.in .g~~1e]11~. bacteria c-anmadi do rOOl1 sexual SIl!CCic.~. aUld ul:;dmallsllils illvolvins ~dlop<Mn)' ill' nr~hc: ~pe-cisliznticn mwn be invoked in spcc~mi(ln" In. Ihis case, the SiUI,:uulon .~ Im"j~C~)'OImllogo!Js~o s~~l,;i!,uUo!l in highif.'1: (a:rglli~~i:~ns. wi'~ba.\til dlC chn:i.c~ U2), Illi Qur ;Jmll)':>is,,\'!i,'~ h,wc m'Jl dii$el!l~l\lJd Ih~ .rolc th<1.1 mlluml sclocluon may ploy iu dr.i;ving spccia~~O:I1. l:his~s l1olll!\.-'Cilll s": woe do~ollx;l i:cv~ l>CilxlhOfl 110' be inmpQ,r:'I:llll:qllhc the comrnry. R>lil'her. ia fJ:; imlslJ:l!iclh~e to I!Imko;rglill1d tile dy-Imrl'lics ('!·r nelnl:rnl diwrsificll'iionl Hnt! i>p(~i~I;ii()il~ to thelll ~lIldeLSt:!ll~ldhow di nCi\.'Jlt ty p.;:s ,) f ~dc,;:liofl migJ'n ~ullljjetl'c lhis IYN:~ess.Also. we mi~lltrl~!:I:Slbly ~lYllootlilcsk:l.e I,hal even ill ,11:slmc-IUI'Cd a~~t~\i'C llambcal)c. :ld apl.atlon lo d mClrCnl nkhc,<, may in~~oh'e :relec~ion ,"'~" $Il1<1i1l proIXl,r~ lion of hx:i. ;;Illd th~l~ tlli.!!lt~h~ gc''U:rOJ!~iQn of gt.".~omi(: b1u(iC-fl;, IL1 Il>OrHllhin,![ion
.:!lCCII,JliJilll~ lIl~iO]il01' };dL'(:1 CCll1pH~01tilm.s 3~:;Q(:1,1k:~

Wi!~l :\.~'?>;U<l~u,tUil1g u

blCllldir~g willi (KX,"ur mnn! nlpi(lly 'I.hma <lIJII&,lJ,nllc JiVCJ~KK~ (Fig. 3l~how~~'{:'r. gcncli.c
r('IOIOR.'-d.

divcusity .i~mmsiernly enhanccd m\'il~g 10 the IOll,g..~eiim f"C1'S ~tcnc~ of :[II:ncl('S ~ii:n'll ~(nh [IDP'!lIlmi@,~l::;.SCpilrnitioiil thrcsho!d:; <lndlhc dyw<lImics {'If bl(;l.ai ng aile C'xJ)lon::d nUl her in (J i'}. ihl sm1l1lnllruy, sill1rl?l.(;~1P.1iry \ilI ill !QnIJygcllcmJt~ di:;ii~lCi. b
dUf,ICI'S

of i>u'dins(we, \lory ~Oltg pt::r:iod~.

'Gompalr]SOIil Wi~h MI1IHH®(:lI,"l S'~qqJleJ!~ Alilla,I/I~~ The! nfcrlIil.'d g.cruc,lk:lfnapfor a ~'lmp:J!,l Orll\!."tcl~:" ~rom~'I(.';~11i~is !;1'OL1i?' .SIf'(lP'(jf,_'o~',:i f:~g.,,4 ~ (J'(J~
I~

r8.(;tor aboLiI. W O. thus, C"~l i Op.j)ORiJllitics. lor I\~ombi nm io,liI iL.'CtWOOl dlCSC \\'cr;; as IIt-cQ~!c:llt ~ lnt~poci lire Iit.~ombi~m io~.~'hcy wotl!1d !lOi. ~kncB ow~ng~Q gCllciic (1i\'cfgeIKlc, B;y con· trast. d'LC di \ CUS~I'l(;C' ooi \!I,'(;'Cn S. pm.'WJlmi.r.ae <Iud S. !J<St'lltluPJII',r{mf1J~ilje iSlI.bout J:%, ~o d1ial :inters(1'Ccilk '~lXombimJlion ::.hollid (lnlybe l'I;.'(]m:~d by a r:<lCiIlO'.r of 4 ~'l:!lat u¥'C'W Inl mSpo:.1ci lie ~,'CC{lIiH" lli~1I!l~iQn. III ~ynlpa~f)', Ih'i",i,s no~ 8ufncioC'm!y J~ ..
lI'(!r1l-I..'i!l~ '(op!'>C\'Cril two

Qr

ob(.;;!l~!ilcd from (hL,l ~"qUL;110(.~ of .. i~ of !hl; ~""J:1iI ~'!:!1!.:.~tll:!it dc~n~nh:b' S.t~pI.ol;:OccH~. M1ST ~h!C'ui\c mtd liiOI1iI Clll.rc\lI:II.~"g 'Hile n~'i~:tr,i:x of :ole-,
MI~

b~~n)d~l~g. iH~~rl;:',<;;lilTQgl}"I}u; Iyp~ Dr s.~r'L;pl~nI;G1CG a:pj:'iiru:v [0 shOllT1: ;1. i I:~I!.:IUi:11Nlnl1~!",..ophtl.1}'!~x.

;lriKC~. b)-'

Ihe

i\'dy

li!.':'1!aIJOII liI1ul,;~1 irloms.

slinni ~!Ir lilidill: wi'lhi!!

~lProc~;", gtJ\ ~r.IIU;d by :':~I'ifipll:: IIila k-s ...

l>i !~'i to ilall'

REViEW
~lml'>'Cdescribed h~rc.~n
~Nrr~~CIom resnhs to ~1!t; !<1IFJI,l,m- \',~h,.l~!I'(I f ~!)~ !';1l'(;J~1 !vq p,lr'~I!~lt:~()n sj:;{~. w I!¢'~ S{;"!q~~ iVI;: :lhrt1L~ ,~n;:<L~nl)1 [icd :!"'d:lli \'~~ kl i ()h-j,~.Smt1cndd il iouliI.1:lii~nlil.l~tion... mid (~i~.clll:;~ s io~1onhc e U~'lC,t i~Jc.[\~a.)ljn!;lNf! urc in (l'n· or The derjv ..~ioll n.r O!n~.Iy~r1;;.d ';;Lppn;:n,ill'll1lIiollt> to 11i~ 111rtll;; "'......L.";S, of ctm;'[C'f dj'1J'i;''l'i'iIi:c101 ~ i,~sp]liu.ing. " .. ex li:riL:'th:,l],l. blend; iii!!!,. ~tI1cl.It:.iml ilK drill), deseribed h~Wdll ht':~p ~Il expl.oriillg this. :>iJ~t':Ct ri.l(~hC',.

this :'iCI1.'>C,WC~ml)i l1ppl~c>lb!c tfll'mJ!It;'h

o~"\~ il~ fig. ! A ,,,,,,~"llNto b.; InQ ~!Ci]j;;llgl)' ~JllliI~~<lr ,~Ullrm~ bR{)kri~q. as din~TCn[ as StJq'ltKOL"CI. l:lat'l1mplr~'flll; anel
~og·!i.b'g~1J" n,;~lt!al]
gl,;lll,;"'I':JIi =-l~n.:

I~Wll'Cll'ly c::d~broJtedw~I1(!h.i}d"'!i!~ sets or experi1l1C!11<1! Qii:t:5er\,m~\(m~, For c)'I(lmpll;:, "'hh01.lgh ~hc

17. E. Map •.Sf!:r~~·C's,~tJd (hg Origj~ W.~J'e$ {(~lu~ia, lii(l"i~" i>re1~.IMI!<w' 'If.o~~ 1.9>l:a~. 18. M. C. ], Maiil~m "r ,~I., ror::.NIlI], ArM St,~ U.S.A. '9,!i.• P

16. ].. MajMk~

F, Ill. (oMIJf!,

6ene,t[~1;!i],

lOS::!.)(1:999t

3JAClUW.oo,
]:9. C. Fr.l~r, W. If', IoIanag~,. II. ,G. :)prnH,. Prnr:;,. N~IJ.,/kmJ. USA loa, 1Y6:B(:«JOS.l ~'()l 'i'lt :p. 1fI~a1~~, C, ~"~W', G, :S:wr~t.l ~Qf,~, ~;lII~', 2.10 troOO). 21. J. W, 11)r~~~, Gla,rL\e5'l\9llM, ~, Cn_f,!ID~,~, B" f" (row, GeI<!~.rict :14S, 1661 119'!lS)" 22. J •. Ma.~aro Sm]llil, Pmrr:. R. 5,oc:. ~il/ld~iII B. BlD.f. 5d.

Bodllu.'1 };pocies rnore I,llbrt is ncededio m.e,~· r~'"CO'rnbim~~:on ~4u~~bt;;I\'lfQl;'~1 do"c~:>, ~dOll.t"''d i
1.1",.oq1~ii1;ll!i!';and ,alliiOrn,mJics hm,'!.Ii;

so.

a,

jjuclct:i~Ib.l.icauM;"

A~~ !Ilten1i!ilivc perspecti ...c Oil bacl~r.ia1~pc .. dmion has bcc~lpro\lided bi' Cohml. \,dm iden!in~-.d ~lC eloaa If-se1(Ualllll'e;"~IOMfo,r tlClm"l~ dl"Ln

bt~clidOClllltl~lir~ld iii 50i.llC j;y~t~I'I'b~ Ui, }S,.l'!l'). 31~dalse '~O ~~Iimmk: gtiilc iloWwdlhb.'i m.ld be· tween Ilm.Lii'lll,POl)U Imio:i'l~"Ei'lli ~nimui'Ol~cd:,~ teeoeci {!Fig., 4) I'Cve<ll~ m. d iVC!f'iily 0 r pmlc:nls

.set,.

iii

"!'

~ ~ ~ ~ ~ ~

• -\o~ "

'l,.:":

'.

between rela.li"'i:~.)' elosely rc:IlLl~Io.x! species, as \.\'c'Has :l~~pm~m asy.rmnclrics hl gene now Ihat arc not t'O.'j1~ly ,expl~ill~d by ~ll1lnple ~nodcl~. ManiC' wOI:k .i~ "~8ii:1 :n.'lC]W1i'rudw c'."pl(l!~'C lh~ in·· ~cll~layh~rwtt'n~t1U~lbi~I<Q'tijnll and ,l(I!'~IJ~"'~~ On in nt{jj:r'L;fC<~1i~,ti~sL:,~le>e'L~vc 11,nld!'i~JlLpt~"'" i11C"!lI,Id· in!:l~n pm'llic~!h.lf the role cpi.~I<!.t~C inM'<llChon." 'that COlin promote div,,"ffi;ily and li:rnit~h(l SOOp!:.~for gIJ11Of!ll;·witk sd~\C'( \F~ ~"'WC!(.''jl~ ~ ln OWir ,pp:iHi'{.I:I'I.... utI(Ji:I'l\~1~ndirl!!i I.~!J.:n~~1U1(: <Iud

l45, 3, fl9!llJJ.. H. o. ]..l,'\\~tl'!J, 1". S, 11}r.d!;l'~. ,M. IE. l. r!.I~am,. :)att.!!'e ,2~~" II.~02{lOOn, 2~. 1. It 100.m11501"1 f'llill.:, Sfi'Ma 3'Oil', BH (2005). 2'$. 8Q!!l1y,lli~ iJIJ!i>!I!!1~~ d!!firtei!l as J(I!~. m:~. ~ 1II!!'ilItraL fi!fler·WIi!Jhl I!IlOO~~ ~bin I~ Tn if:lldil d1~'iNt SIMI! ~311Q~1 ~~. ~li:Ilites ~I'e samJllle:d rM1II re~mi!!ft lram 1!It! ~~ ~iI1erMion .,m~1'I e~~llprOl!)llbjl,~~. ~iIlI~~'au:' 11M II fij@i'lCItypI!d@llftf!dby "0 aIM~. ~flId ~di, aU~L'f!iI;
rnigfled coor,dflliltn

or

seq[lIemce: S!)i!!.1l. Mulialieoi'l .amdl recomb:inOltial'll oocllr ""'fin probam!li~li' bIN ~OO ql~ 9tl'i~, f~Spi!(lNet)!, Iiloflere'.t 'ill
U1e-~emg~h ~~1J;i1: ge/l~ j600 IIp}. I! !lJIlJ.!latiOfl! OOt!JfJ5., o~e g~i1ie .!h (ooptdlnat~ i~iiil[]\~i1ilieniBi:l ~y :1. At ij)e, ~iari 01 lin!!' ~lim~IilUillm, ~,~ comtlna1i1!1~ft M!I 1.0 1. iH!Mna!~

Mthin <II'! ill:i~!r,iiCl ~'Nlimelllii"~l

rn

o[!;laltiiZ::l~ of g':!'Icilie di'l'!..:'ll1llty CfIH t1'.~!~y be LOn :lchic"\lc'tI by lak~llg :1 Ul'iuli ifmx"lcd IIPJ)roa.ch to til\!' pmb~cm. Gel't0tic :SUf'I/C)'S can reveal I.h!! ex-

Ihe- il!lwI!r<l.t ~eqlrelll'if''lI'IKe ere rela,t~ 10 Ir~~ 'gfflelk d~IoI.liiI(_t.!,tby U!:ii'l~ ~ ~i\'Ib:l!mllil'lif il'IlXI~1. B.Ii~i!!:Il M
(orlsid~rin~lht IPFo~'dbililr of mut~,ijolfl,~ renrring Olllht h;c~Iii!lf! Q~ Je<!1 ~eque!!ces. ~Uel~~ ~!1Iafl!mi",~ d~~arnt!! i~ (!5lili'i:t!~dill~ Ul-(i'l!,U~,hiher(! l'J .~ mil' ~Dil!dI:. ,liIi!'I:~fI~t imll'le ilbllracl. Si!>llL:iffi« ~~" ~ iI1I~f~ of tm~ n1Q1i~" (il 1i'I~'OO;i:on$ ~1iI~t~ep!if~e5 'oo~ alW~~ ~~or.mbinatlon lL:ioo!i!d~rMl~prolilabJL1~ al(htl m IDI~t Jdilil~n ,ctlleM4 11l; ig. 1.J\ il1ame~ 110-,11"", F ~hl!l'e 1/.' i~'Ih"l! I!c@lwql!.el'loe 'l!li\oe't9ef1CI~, eli!~~'II(!'lJ;le
~1"!1~

tent and

ll.. ture

,o:flhe d~\ cn;ily~hm surrounds us.

n=

Carc'fi.lj~ c:-;«K"rim CflI.Jlioll cm]~'1i8hli&ln POW'Jl!im.! mcch;:m:i SIMS, fot" CI'Ca1 ~ng Ihe ob.'«!1I,red patterns. Thoop':!,t~c<!1 odels (;:m~hcll be asedto c'xp~(lre m

F'lig.• 4. Gieli!~tlc m!~1) (If the 51f~rO(Ouu:s:

'~'~r!Il!~"

Il(j~@d on (O'rKaJl~nal@dls.~l!~m:es
Q~@

,elI
ri® Uir

MILST gen;~~ (eoodUi d~n;g .ridl).

~aJ!1!1i!'d.ScpedlB.

higU,ghtoo, IRed, 5,.

:511 IIfIples

from

,PJl~ti'"

m,ol}i~e"; YIlUew, S. p'~elldOpnellm(}uAn!; purple, S, mi/is; Bllld II:mWllll. S;,om~"s,ihe tJ'Ir~' Il1gM b~ld e dCDts 'ep'n~'$ie!llt tr~iilsfQf ~\inil;:~ s n'~~ed speciessta.tus 'Ciou'ld no,! !:Ie asses~edl.

wilC'lhcl" ~h'C'lin k i)[}l\.\'C(:nl!Hcr:h'lI1iS!1n~ m~~~ ob, scrvatlon i!! pkLl1.'>illk. Because I~~Clcclmological car:J:cith:s ro'U' sc:qucm::iIlg Jlld sim~h.1Q:ing s(Xll~'em)C's arcbmh gfO\"j'n,l;l C';"lpo!1C'm:illlly. (he abl,~:i Iy~o ~illlktllcm iI,no a 1I;lD11~L,,~cn~ picture m;.I~' soon he li'lllil!;'d n~~)1hy Our im.. ~ill<nuO:Il.
RI!Jm:l!l!u:;m. ~nd: INot:c!§

Hammin~ di~tajK~ Kti'ima1.led at '!~e .a,~'I!1i1'd 9~m!! and 'ils

ffio, 1W1[~il1)i'J dl\iidoo Iii!l' ,Itll! tot;'!~, IMlgl!h [Dli'lJj'la~ 130..
~illl~

'~n~

11.. lit (I~hmiln.

bilil who hlils Cil1lphflSizcd th31 th.c threshold rOf sharill'lg' tlLda:l.lt~\i\Cpail>'lllO:t'J)hisUlS is, much higher i(Jh. ~eadins to d1C' l1o!ion Ilnal rOVlllmiOJ'l" nmy be ndaptivdy d~S"~il1'Ct l'1l~t ind~slin,guisl:mb~ b.y r!Col!uul H~afkC'rs. T!!iIo':\lC!'iitlKhc!' hi!VC ,etllphru;i~~ ~he role of adaIJ~i\ic umlitmiollS ill ~SlStlaling ''cc·mYI'JiI.\s·' a:ii p'l!t~li'!,'c SI)Ctics u'n. Otul' alJ<l~y• t'S :I'~Jgge;;;l.thm for I}opldmlon." \...i~h ~"CCClmhi~ ~ ' ~!OJ!tion',a'~ ,~l~c d\'C's~xu,d !.hl'!..: ...I:In;~'-~l;~tylJ'l.?'i ! q 1;;'(nlld .D<lpi~~I}' li~'nd ~lll}Wlld the old'Ip'!IVII;.' hn!~~a ;i;(".JP<:; clmnge ,IUd til\: bHm1;n; b~~wl.;ennl;~I;n:t-i 'Ix: rtuo .... d. ilnd '~h~tb<(;lil'W die :;t~:l!~IiII, t."' thrl:~hol(J. d iITcrentiJuli-on i IIlodi&incl. gt.~leli£ cJu.[,t(;r.'i arises. CVCllh'll ~h.e flib8cll,Cc of sck"C~~onl, Our mooel ll~gldigflh:;thC' hnpoMIlc\: Oil' <I ~ctil,i:1d qmm~i~m d~irllon c i~ oflhe r.ro~'S ~l~m drh'c ~o::~cialiQ'll" he sinn~]al,km"" u;~ h~ T ave b.[l~d on ~m:ric ~)lilU.;; parmm;iCl'S. IJI..II ~'blc li~t.1her lJi,'{)ikis, ooqu irod~op.rodlll,cc sc~ml.llalkms

2.
3.

~. 5.
6. 1. II..

9. W. IHI~nilge •. B-.. 'l1~pr'iUJt. !e M. IE, 1I11ffier, C. ~ril,5er, iWi/IM. fl. .5'~1C ltmfiM CSt!.,,9' 36i,~j.'9 (2006), 1!.O. 1\ 6. 1l<l~~IKA. !'h~,~I, gi'~ i6t iI~o? (.aOO:il), 11!.. I, l!.'I1!l~~, ffM5 ,NI~iM. It.tl, ~'9'}., 161 {;!OO'll. :!l;:t M. Vwlii~, F" Diool~~, F,T!ilddt'i, M" Rmilll.. PrIK. W~fl.

I, Go. !.aT.iftiilCI, ~. {l.. Go'I!m3ii1, NaMI! 40S, 299 (aooo.'. IE" ]. filii. tl !i'L, ;!'TQ(; lI,I'ad...4l:~~.So. U.s,A, 98. l&l 11,00,1). Ill. G, 5Fri!il~ 'W, p, !I!~ ge,E'. ]1,. "~11,{I.i\!'!'. Opin, .II1IC't!IJbIllr. 4, 6!Jl ~201Il!l. :5. G. Acin::u.~ Dr,. NDr,l)ft4~!W, 5:51. (~IXNII. W. P. iH'iI~llIg~" (. fr.a5ler, B. G. 5~r,;m, "Nil( 8ioi. 1 ,6 tW(5). I'. C, Venleor ,iN 0.1., Sd"f'I1'(-.e ::104, 66 (:!-oM), 10.. (IoNeo:r~fl' ~i,l'I,I'at~, ,i!.li~~w.r, ~. 'l33 j2005]' • II). FalUi$1I I1J,,1'f.JiJIM". ij'OJ1:;' iii. Sa(. ,tMdlm SC..r'. ,B 36ll, tit la~!i121006:1. '

i! r~ ~~IlO!~~~ ~~~~~.rI~ iiiI~I1!iii;i'l (iii 5dti'ict Onliifl~" 28. fi.. '~tlil. M. I'Ill1'1d'rii~,M. Li,p5i:tID,,5. ]inlc~·ao~ernoo, ~~ w~ji A,r,t;ld. Sr;i,US4 ?oIl, 'liS] (199'1'>. 29. 'I'. C. ,ZaI\r1!, s", ,Malo~, {Ja. NilitL Al'Dd .. .5'~i', U,S.Ai 94, ~86(1onn. J!'.I'. 1t'I1. ,Po li!a!'l;i~, C FcJ!il~J,. It ,y, :5,p~~ PllJIfJ$. 1irCl~:t lit ;S«; IMIll'lm S!!l'. 8 361 '(2(1'1:161 U.. I..!::" Hawmll1m, IR. lila renib!ldk,P. Gilustad, J. l!t:lllMM. 179, 6-589' ~U1nl ~2. 0. SdllJittr, '1r,mifs. [[m, jl!lS, 372(200[1. H. !" M. (ol'!a~. Am. Nat. 143, 96S 1199'4). H. F.. M. 'G~h~lil, Allml- S1E'v.Mr..rabiot %, 451 (.20(l".2J., ~5 ... t 'i'i!!ltl!li1l el'':'!:. Sd!!l'li'l!' ;i'iB, 498 '(:lOO~. .~6. rt IC!e\I~ollmffi1: ,Gilfe VI!i111l..III MJ'!,(lIltl:gl! ,~~d EnvlrollllllMJ frN SIlOriSlico1 (.ot/l,!Wtrng 01 fau.mlal~om rM 'ilati&kal Computing. ViA!nillla\ Au.~'IlrItl., 2GJQ~l, -J!·FcjecL,i'.I9, ]j\'. 0'. i5 runded b)' ill ~~~S<ldel)' l!I~jllErsiW !I:ew:nc~

,',I. I\Ietro,,~ ~ru!ilIl1l~!h.~

26, ).. Marln:ird Smilh. N, Iii, s:trIiln, HI, Om1Jr~ B.. G, SIlf21U, Fl'"ac: III'~IJ: ACit!Q, Scf.&S.A. 90, 4.~®Il U99:3~.

,~vo.'.

'raM,

W.I'.IfI. ~d (I,G.5.!h~n~!h~ w~~~ Vr\l~ tutwlil'ig, '[he atiitlot~ ~rki'lJ:!liLM~~ 1J&!'tlll. tlIl'I'U'IiI!!I'l1$ kom II:I'. FOIIu5ha!1~ O. ]. Sltiitli!. lhe i!1iI!h~ ooe 111) eDm~ntt 61' im~t.

r"r

~Uo~iitp.

kw. xi. U.S:A. 94, '97&:1 (1991:1, 113. )i. l!.'l!li~liii. P. ZawM~~ I? PIdl!!II'lI~ ~• .M" OIlliilll" C, G, Il)llll'lOOf\" ~Mf'fi"f. la2, lQ16 Cl:OOIU. i. 1\4. II', ZawM~ M. S. Raberl:!, Fc. ffl. Co~~ Gi'neNa ll\1'111.
911 (19'9)).

:S!l!!p~o~ti!l!!ll OIn'lf!'!E! M<!~eo~O!l

~,~~~rna9·!J~gilc~l.ent;'lillllB1SI5;Bnj<li6JOCl_ Mlilw!s 1IIliI1 ~!.\I!1lI'Iod~ tHg!o.- Si l1(I 5,7 If!.t!ft.rM~i!'!!

'iIS. IP. Sh~n" ~, V. !iloon~" G'tIIt'fiB" i:i2.. .j.jl (U'M).,

10,1:12(J1scil!m~.t.ti2'i)7 3

S:.BX.~rSIP8Cific UV aad Flu,oraSC'8InCI S:ign,al:s, in Ju:mlpin'g Spiders

of COUrl;;ilill ei.hell' rich or . 1J:;:Jl1\, ~{lICl. ,~UVI~\"I\'Ckll,glhj;. ~lnd nlJmCl'OOr.'l ~<lck.ing in UV \\·,wc:len\s.h.". By u!'ii~~g tml'lSp<l'" " lilr~lC:liol1!'i.for UV "cl1cc"Hl(l~ have <lb'Ci bCC'~~ (lill m~cr Ih,~~ :ia~oeifij,ca.ny Mocked l)V w a ve~t'lln,bl~ (h'hl.f!;,~SPO!ld~cI h,lt:OlJVI:~n,,\; UV+ H1<.l1l1;)-S showu ~nhl !ddilirm, "j~ib~~~ighlnuo~~rlo~ < k;i!lg!h~.r~ S I l, w'I;'P,.'Ill!iJVQd UVreIM"1<II!t~ (;U:l.~ !ill. ,! F~. 'lfV+), I'I'!O~D ,~~ 0 l' i!,!.l'ill\l~t ~.!C ~ltt>C rn~.lc Ih~ iuclln"ti b~r Uv w.oJv'OI~!~l1l~h'l!iiijx.'L;'ll in~pli~;SI~'\! in nl1~k'S( 'UV ~"UllclJ ~]n,;\'C1lil)l,;d I]UOi~i;.lt'Jt'"C" Ibm. I1lJW im~krt.>dlJV eues ( F ~" ~ UV I (F~. ill :lninl<lI.:;l~gll<lIi!1~ in certaill I11millC·~n\'crldbmti.,,:\ 01" IC!11<!k'l1.'~~ F l. \Vcr a:if;o p.1'!\.tr.ic~--d'l{mv.i~IOLI IF). (l·.,,(:n.lho~lgii tl~i.,~ !lIw,!.t,."S a I~o rolHi!l.ll!!()d&o <tHO in the pklrm!;.\,1~ of !WH'Ol'" {2, 31. '\Vc Fl'PO:lt <l cO!I1uH1l!licw ioo by ll!~~I~gl~ldi,.'idu<!1 gla<;.:,~ "rum'fl. :>ilow con~\;i~!.;'n~cy iI!~ !Coul1sl1 ~ ipr.WI:; C1oncll.lldc Byrn'U!l~lllUlonirl.g VV~brO(':ki!lg fiH~15 O\(lU' bOlh. C-OIM or !05ult.·;hip ~"'iMIIl(ll:~i~lM ira \'\;'hici:l oonll UV mil~ul ~xu'li c(,lor.1ti!!l:ll is ;1 (;nl;{:ial ·p~~q,u.~itc ren~"CWltC~ ;Bil1UU\.'.i~lIrll!ll~-d nU()It~'.'Il'OC:ll a~ 1,JS!;,'{i ci!lt(lr; (if nOJl'I;l nf th~ gl.:l&r..;~lfCili(iK. ,.~rt;l;Qb!~in~d · ~i::rr e{~b.iIls.h ir. r an. fl f;(:~-~'Pcdjk:basis, II! rhe ama1[Cj~liltlpin!:'l ~PJ- r{}1,~t ~eenl~lri(l."~"·UV~and :f-l; IUV-, F·: ·UV. F ~and -UV, r. d.et Cur;~1IJJ~Jlmsis nmbm~i('lJ, l.lil(;iliIlaJ:~~ im\'rl; JUtl1llJing ~M(n 113\1c c",!C.·dk~llt li'Y(!s~.!:!.fn andthe ret ~i1i9S r ~heprincipill eyes are Il:Il:OCWIl, (I U V~rcill'Cl~V~ patches of scales on tbe race 8ilW Under ru ll-speeuum .Ilghl. males readidy 10 hDi\iC IlhowreccptlYU"S. with n'lla:idma~ :scn.,,~bo<!y thaI arc dlr;;I)la.ycd duiiiil1g cOIi....pceific courted 1:C~m]~I!.~ 3doplil~g Ii co!llrlship pesby p'osllITing ,(.n, TilteSC' :lire Im;ikill£ ill ICI1lJics H). turc eornnrisinga Hc,xw-up abdomen, arched Ii "'~IiC'5 to UV and IlJ ~OOl lilght (6. 7). 11I1C·lJ V blU fC'1l1al:cs have I)a~r:! with a. UV-rCxdOC!d Ibril~h~ legs, and exicnded vibmtiug ~]<1~p.,>; fcm.<llc:5 ~ecpwJ:S have ::n1n."adJy been S.IIOW~l !o ~~JVC a ..:;. groC'n rh'!{I!I;1;!sQ.."'Jlce ~I~alti.... iJtb:iCllfIin meles (Fig. responded either with displays. comprising role inma 1:IJ"nmlc:inlcrnc:.iolls '~.'l!1. Our ~tl!d}r ~!~lllc'lICd legs, and bC~~~llbdomca by bri.dly 'Of !ll!ffD!lgly sugg.'l.'st" a l'nh.: fL" gl'L'l'11 waw'lcng~hs 1. l\ 100. as wen. The reneclancc spectra o.fe ilmhnwhl ['It!IllIlilngm.''\'l!LY l4). Will~ollll UV, rClll<lks. cidlcr lo.ejpilfA:m~m of Biologlr.al SOOfHl::5, W~tiOOi!t Urnoreml'y of made .1l0reSpl1)llS.C·01' simpiyll!t1Ilci.I a \\fay '~\\'i~h- mUm!Ic~ ha \,'C p,!1OJnincilt peaks in ~HII~1 U V and Si~g~jl«f. 'U Sciriliil!Ce OriN!;: 4. :li.l·7S43.~~po~e. ?O~jl<l~.1. (ll!.n f1l!nJl ing). S imnilarly,. ~JlOllc:ii.ci IhC'r ig~~orcd green w;lvc!cI1,gths. Zl:nd the UV"ill1dlJJ!ccd green msat 01 Bi~oglI dOd EIMID1lIlfI@mlal 5dente:, Urnl&~ity ~. C s'U~ BriightM BiNl 9QG, UK. ~(oIleg!l! ~f !j!e SdEll1i(fS, nonnufI"J.iI.'S{:ing fullfmk'fi bY'lH~lUli!g 11\\',.), ,or HU£lrf1;.;'.IlcoIlC!;; is n.'l\:l,~}C'~cdl Il\illlCl},liO' the r<1.lp::l or [,,';!ril'il£lru~d~ with a [(,(h,ICtd disl'l·I~y Im;:king sonte Ili;:m,~lc:>t 11:r,:.. S2).. HI~btl'i lImhie~ity. WWh.;ll1I<l300i62, Hiuk!~ (ibi rna" "To ~,hoi'll ~:or.~~~poo(!te!loe ~l!au!dl be ..ddlr,~~. ~E"mi~i~ IJd11l\ imal: Ck:ltbL,ll">LWC tounlLlll'I.lla 1tilnMc pmhere is WtdCSlll'e:ld CV:idcal>OC ror an ~Iml l)i~o.mcocplj}rs~h"l. aJ'C 9C'n~ •.lve 10 !.d-

.· T

\lfC c'.lIam ined Iht: COl'l.;;.i:..lCitCY belltl.vior in ~1]\iJOUI1-.;:ms n11<ll\\'ere

lug 111.,~llk:il Ihe: Or~~il~ !>c}l. \'\1!1~ ij n~~lfI.~r:11utllyJby UV-dC'l1eicnll liSIlt. Ai1l!lOI'~ the 2:0 UV-I "imk~:> 111011. aCli\lc~y (O!.H'l>OO F I R:mlll~.~nosi. ~61 ( fili~~d !o cmHt Ihc rcn\llllc when s.hc lacked lluercscenee] ~UV+, F ~.Fi~.mH,cv~nlllOl:I~l hrC'r. oohJvio'rni fC<lC.tOIl.o:. I"cm<linoo SUi;ll~:iiI!~C'III)' the same m;: WI!1rcl~\r !lm'1]l<l~ light And mnrmg ~.2F+
.[}]lC

or

in!h~W'R-~(lnoQ of UV l ·~UV... ~~+) ~a~~'~x!~9 "hq\\' ~n!J:,~:!l'~!!i;~m~ b~h~v i:o, ij II ij!S ;;LbS¢I:I~C { ~UV , - If HFi~. m)" To cn~~.u\J ilud i.h~ ("'(~1,1I1s1tilj Iil..r;spQTt'K'S of Ihe 5fIidCi!1:l.w~~ on cfl'(.'.C~of ~x.u<!!1 ookl~ iuid nn~ bdla\'illr.i~ '~~'lllgl';:.s in 'ttii~ (lrpO;'lil~;Wt;.lr:.. WG Q0111Pfi[(~d !he: li~h;.!.\'ijotal [,~'sp<l~S{.~ or iricli\':id~UI~~
!CTl'lC"IC'! ~.c;x 'lIIidcri"lIn ..... f!'L~Urd~ ~jl1l~lC'r or lite

en.

Ilbslidq@'lnlU:lcedilu'!D

portion

f.!f~illC

:'111111';:pruirs t~ u ...

:i.tlC'oc'Sft~>tallyil1-

JiJe're,r'leilllces Oii!ul ~t~s N

1. M. ~"'i:MI:, h~
r[19 \151€trQ.!.fl't!'.

~o, 455

2:. ( ..H'.Mill" .• f. W. Gromirn"It 'L. 'C<JIIII\lIllll . !!rI. ], Mai:ih<l~, $e~~ 30~,.SI ~~OA~t IPiJbllsh!d oolii'l~ 13 NQlj!.mMr ~Din ·(lO.11l6lKiel!1~e.108~a03~. 3. K. 'E,~ldI. I, ~. F. Owen:l;. IN. i. tI!!!~!!!n. 4. M. L II.io. LBm. Il. LL IJi~'.J. ,fifill'. s~. 8.9,. ;I'H t200cl). ~. M. r. i1i n~. '11'1 .m~j'(Jbir1~y a{.Nirlihtifrh; F'. G. Sarltl. rs.d. 'SjJlirlger"~~.r!il!:l, ( 8.mil'l,. 19:B>I• w. 5J.-7a. .

s,;~,t j!;~~,\lZ

'(200l}.

IF .~ ..
I~'~

6_ it. ItI.BI~~t III t liIardl~,. II". IW!lm~~te. n S" Wmiilms. J. ('omp. rnjr'~!.jDlAI'14S, 227 '(1~·1iI~_ 7. It. G,Pril~s~. G. Wl"r.fId, Camp. ~}'.IiDl.

!JV',*, 1'4. uv~ :FCou.ltifllp· paJl' oolD.u staU.lS

f+ F·· IF\emillisl ~@Ior s~arus

A lCl4. f:!~9 ~U.800. 8. ~. t, 1'1'<. Lfi!1\ Il. lU (Oiilip, Ph~ol .." 192..ell tl'OOQ. 9. Wt tboolc R.. t:. (o'l;,wI~ L P. ,K1]~. :1m!! t, I\t L~~ for ~m~Ii1!$IlCl, ~iIne;lm~~IP'" filll~ln(I!!!I.lI:!!ii~tI!nc! (!11'1~ 'ffom 'In@

R~-<nh

Nil!tiC!!l!ilL IlIni'l'f'fSil:.- of ~~,,~pore At;;,dem;c f~ tJ;i I).I!. ffi·J!S>1.(!!O(1>Uo-:ml

SUP\l!loOrI,'li1l!1l On:~ine Malti!fla~ Friig. ll. ~,A t® C) C fIDlbra,iCll sllmJiiing UlV al1d~LllImeS(em l1ilarlrdllg~. Mal!ers (lef~) <lind females (righurl1om WIWr.!:rum~~m~g'~rwgill:Jlnt(!nll!ulJlll!i~Sm fronM:l r(~op) an~ d'~:rs~~~botoo,m)ilil (A) hUlman-visllh\er (rOlorimag@s, (B) UV jmh~lg.es wilh whotl~r r~'giion'S5h()~"ing ~.aUJ)n, '~twng UV-re~lect~ve Iparl:s,. and (0 u'klrimage:s cd UV-imJiI!l(rOO RUQfe:5(rence hll ril?m~I.e:s {Materialsr and M3t~!ii!l~ <1M ffle'~ad~ MIl!~ho'M. (Ii,). to If} [)ec~easein the nml1lber gf -mmtsh~1l rll:5po:n~es ~'I.IheliiliiV \It<liwband1> \Jj1~re r'emO'!/·edi. '~i!J) Figs_ :Sl .. ~dl S2: R:~dr!.!c:l'io'rlif! '!!)!.!rtsh~j) illt'M,~tllQns ~P'= 0,.0041; Or!'!re"~!t~ signltest)whrl UVlI'l'as;;Iibs.e-rliI: 'rF!!lm ~Q~h S~);;e:s'. (IE) AtI~rMCe~ Mates \lle[e Ullilil1lt~rest.edwhen fellllllle5: ~lI(ked fh.J!OH!:5cl!m:eU' ~ (J',O'()03; '!Jll1e·ta]ledl f>l1gln •. olll1d '~I~)femalles tes-t) ']·~g~~l.ii!OO6;~Ooe~gJlt;!! nrO,.Wl!er il;OO6 we~e llIninteJre~l!1!d in male'!>~hat. lLad:etl U\f {O~Il)r:~ (P '" O.OO1;®fle~taiLM signl te:st). ~D.lll&l'~tle;fiIC~Jl .'ll5.'l U

Hiiistoc:olmp.atible IEmb'ryonic S:tlem Celilis by Partheneqenesls

Ki~iIIi IKim;l,M PilIl!Il L~HOIJl~1,",,'5; .A!IU "'I{.• II:II.1U1Chii,I,2,4 (la lI!,dii<3ilelllgerlk@,.l.tr.41 ko ~1;jJiSonW@:§~~,2~AI A!lliIdlr'@~~' 1 Kil[lllgy,~ M.a~k J. Daly,fi Gel!J:rge Q. Dill t~Vl.2;;!:,.:I ~ KHwtl Ng,I,2,1I
Gl!'n,eti~aUy l11i18ilt:iwa p tu rf,]lot~Jlt. emb r~(I!l1!ic'~t!!m (ES.) li:clils gl!al1!!I'!ltt.cd vii! IiIUld.c.ar 1M lI1s.h!wOF 1P.~rtl1efl0gel'l@s.is ~lpES (t!t~s) a F@ .i)j pot@ntll:il. SOli! ['Ce 'DJ hts!:oc:omPdlilb~i! ,(leUl~ and tis.5Ui:!~ r!O~ transp'~anMtl:on. ArU:::.r p.aUh@I'!Dglefl~tk i(:l'I\I'at~on ·of mli!filll@ QQ!tVte-~ aln~ ij!ll.t~rJI!JrJll]iln 'o~m~i(j~li$~or It WI!! i$lll!la~l~ (I~d !lJe!f!'luyp~d IPES ,celts aJild(harai!:~fh:~d 'mo:s!eothat ('grM@d(h~ full ((J,mlPt~rI\N~in (J:~ m~j(lr ~isto(omp~Ubilit.y cOimp1e:;: {MHO 3Il'Il'ii~eol"!$'o~' 'the (J(lcy~~ donor. mr'reri'!iIll'Datf1d~is;s,tJlesJ~Clm th~s:e pES (1~1Il$ el'lg! r aft.ed fJni If!m 1,111(:OImII:U;!~li!t WlUK '!ililauh;ed 111'I(ilIJI~e ~ Irec::1pi'Mt$, d eli'l'i~l"Is~r~~ij 11!1 th.a,t setil'df,d pES 'c~u.s. can ~rve as a source orr ~'i~tllwmpiQilible tisst!~r:llrlJi:!iflSplaJiu:arillfl. m:rlh~'J10.\l).;'~!C-;iScu~n."Lib. til.; d'e~'I(llIOprl'!L;'nl , .iU (:ornnbryo dil\1l':lly ~'mru~ an o{lC)IW Wili~(RUt r('I1.il~iil.miolll. MalllY ~u!~inl~] l1I]'1dplarn !>pceic:;.reprodLfCc V~f1 p3nl~~lio~eiK'S~<:, tJI~iti b ~~'ice"pal'llhe.nlGgcm.::Li:c cmb~o.~ devclop o.nl~l ~() !]~C cady limb bud Stll:£'C h<CCfltls.CtlllllJ.mnmliml ernhnyollit d:evclol~mcm I\Xl] likes gene' cXllres~ sion from Ihc p,ml.CTI'I~d ~CUrnll.C' I?~nhcnogc~lclk
('n'lh~IOIlic i1'iIllfi (pES)

(!l!1\1!10l1wmm. g)jl!~ ·11Ybr.~resistance" 111m is; p'Ir!im~<II:!}' n;1:cvw~~Wi bQIl~ l~l~\I ~~!b'5f1~Il~'l' liOTl (!OJ, A~ t"o:rnpHIl,,'t.I \·~[hll1il ...~m.dK\d, OIM:tlI~l.~, ~lfii[:d M~ ,m1lagOri '1lKIIlcliaialg CJlh,lIlti~ HUogalll.ll Ie 9.ur\'i~'al, but null MI']C-rnatdwd li5."l!!~ 1Un; tbe ~lOS~ Ilu..o~tblc Ibf tD"oln'l';pl:)U1I. Ill). 111.(: {l!nl)~ ,[)!,;'f~nil] ~mh ..g)' ~or1~void,u"gil']IiJ.l~l,IlioIQoflicCOC'llp~ic'tl.'

lion~ it.;; to IIr'dfto;p.iru'll !jC1icl~1.lIy jitl~'ilLtall li~u~" t.'ul. thi::; lii1~k:. m:1J]:;lilal1.ro~ ie!1"~Cl ~1lII1.01~u~ l.i1'il>tl~. IITUi~~f'1!<iam;. ix:IWI.len I1mo:nooy~1ie m .bu, fir cells ('located by oomati.C' cell IlllclC<lr lI~m~s:fcr.. UCI\!' we d'nrurnCIJIX.illl:: p1mil'l'ol.cnt ES ecll linc;'.] ~'1~C1'.1I,~dy p<lflhcnow:'ncsi~. in \\hich hOlh of b Ihe m"'~C'lnal IMUC loci hVIl l1>CCllm<linl:ah~od. [)ilTcn.'lt~i<a,wd ~i';S\I~ from ::-ltch MIK'.JIHlIlchoc1 ~ C'CIJ!s C<'lt! hie !ransp]~un~d ialWi [h~n.o<:)'w d(l~
no:nlll,lin willi.(l'l,lll(j,cttitm. St~ggl/,~fll,Ul~ Ih~'liC Ihll cells cOll!!ld be a Ihv(lmli.:t~c ,~Olm::c of h~<;[ooon1r

or

r~.;oooI111in'1Il11y

1'l'01~10?YMol~'" {7. ,"i~. B~C,1nlM;

'I~~!libk,; 1~~L!c!\' ror 1lI<l!l~Jllam.o'!lit'l!l. Rel':(I,!1IIilIilil,ard MHI('mllkjliu~d 1~IMIOES, ~U$.

i~ol:liiolli{If' p(MU ~ES tissues d('ri"l..Idfrfiln imrm:!Z)'goL.... pES c~~II~~ \11;.'0: rri<~rl~,jIh'll dl~rtrlg I.ljlL,,~ \\~(J.l,Jld ,~.prcss ml~Y une of lW0 Sl:1S.of f'fl!rul~tm.l cells f[(jlll! h)!orid I'll miec,. I!i:.~Ol'llliJin:lItim! ';:H:'U~t" hiSlOCOmJlpiil,lbililY fum.igt.:.n.s, lht'Y ('fiU 'be llx:Jre ot\C'urr:il1g bc~;wccu pflircd hQlrl.oI~~ru~ chmin .. I:rndilty matched. 1.0 !pal ierns and might pose :tess ~uoSt!.JlliIe~ InciQl;~ ! wuuld produce ("('IIL I.IHIl risk Q,f ti~ll:e rej:Cotiml ,(9). l~m ..'C\"Cr; nlll hctero1!a~1IrlI l'CSlOIL"t! h!.!tcmZ:C'l'gfJlsily m the M I K' loci zy~ou.s liI.'(l!plclus. nl.1Jm his.IJ.OC:OI~~~ibiliLy eem,(Fig. m~ R~combinlal~ou IreqIlCI1.C.ies Ill1lCC dlC plex {NI] I ) I~OllIDZy'gOUS ti~l~CS 1l101}' be rej~C'l.cd I1rulli~llC' U-2 MUC !OCliiS ;n -m.5 cC'l1limoJrg,al~s

,(.:. 31,. and pES ccHs iCQo.tcitnl!!le 'I'~'idc~y .j)Idul~ 'I)(} ~i:-,:'ii~lI:;'s chi~n:::ric mice (n Ahllllm.u! case of i!a f.!'1ri!h(;:nO'gt:l1.l,1t i:c Cil:i1llenlSlln hlliziib,,;~n ,c!(,':ii.(:ril)()j .~n~ whic:la !lrc ht::il1!:tl{lptDi~ic S.)"'![CTUand ~;'kil] W6"~ deri"od lrom ]pMllhen.ogt'lielL'C cells (·n Tn~ tocldil)o'l:! 1.0 phnipmetlt stem cells rrot'l'l Ib'lil !!led C"mbryos andembrvos acfL1W by oollilfrtk.cc1l mj~c:~m~ms,. fer ($). P8lruu:l1.ogcncs.is is am}~her melhod fa; CI,'C.1IAinS p:J~)li.I)lJlcnlstem cdlls that mig~'u SCIVC a... a SOOI['CC' n.isstlc ror lnusp~.mmti(Dl1. 0

cells have been i!lol"t~'<i by l'ilLl.1J!.lrn~ kiHcr ~NK~cdL~Ih"t ut!oogv~~;re Ihe ~inm ~~,ulhc!!.o~!ll.nic b~,uslncjl!l~ of ~licc and b::k of (ln~ .'1!"'\~ or h i~toco:l1)];l<J!!ibili~)' ,Imjg ..mll. a rri~nmcs U. 2}. Bo~h ImlOU~ rune] prill1J1tc pES cells um:kl'go cxtcn si\ e di rrcren!i:bl ion i~l vitro h:OG

'(eM) I'i'o~n !he ccn.tronlcre on mouse chm~m:t\QnlC 17 ~1..1)1. lhl!L" rn."(li.C'~iug t~m~.,ulf!l'Il;nximallJJ}'one

•
4n I(W~
hiCG

1;!1]h

en)
flE",$
:an
MH~lIi1at~edl

1!n{MI~) .......... ~::;:;:;t~~I1~'~

_O®l~; ~~~~6t11~~
Fi\~DO!TIb]l1I~tio"

ICV

1IJ!!I!!I1I!!!1iB~~h~ ~ ~ ~~~

I?t~~I!lIES
2n
MHC.iMUuelmedl

iHgM)1 cffic:it:JlI mClhod~ oJ ex peri L1UlC.Illrli which oocylC:> mJl\:'Sloo .m the second I1lCioDic ml;!lap,~m,'il:: (MH)
In~lj'illc1)""1l1ihC'Jlotg<:iI1\.':>L~ ~\'i.iSli[t;l

2nl :(MI~)

are oCht:l,nk~dly a't~,!;,1lit!..xi III I~'lc

pl~llC\:'

of

C)I-

tod~<ll~ln. ,;I dfll!g ~ll:!I. Ilrt\'t111L~c.',II't!6kllli orille secend Ixd,lf hoo}' (6), D'jploJdy is m'liun .. ined, and th'l;.! f'Cl\~dt!!l:!;l ~'iI:;;ClLtdO~Y gt1111; C ,U! d..:,vclop
i"rmn whieh IES cd]:o; ~m bl;: i:;.(J~!lk-dl wh ~h Wt~l.tnl pO\<H! ~ES ti!.:lIs ~7) J .. into

~~~~61:\1~,

~iiii!!ii!~~~""""",",~1""",'~h~.~

,1 bJI11i."ltOl;:)r::;;~

ICC!(O[::~B I
.(j·n ~MI~

~~~~

~,!!1~)ES

In some cil:>(-.",pES cells harbo.r;} d:u~JI~.cm of irm <li. hlllpllJid y_.cnOJ1'l'C ond arc thus bel levedto PC
lDivhiOQ !Of FI~dialJi( He.lnillfllOQ)liO,nwlll!!l.,.. children's HM:Dit!!~B,*IOO andl ~a!ia Farber c.anClE'J ImliHII!', BoMt'O!\

2n
...... .- .1n1!1~,O'!I(: C!lea'!!'~el

' .. 0: diMerel'ilt~h'lt>Omere., .

MHe:~

eh(!lI'Ilieal:pc1iva.tlolll,

IFig., ll .•.(lhw'om050AiH! dynarnks during, I'lOrlil1l<ll and ~rt.'ijfic~:al.lmlllriil'l!e 'oo{y~e m<lturat1:0ml. (A) Normal. ~!'rmizatl\on. ~liljHn.Ui.!rl! QOl!:yl'C's alrr'es~.ed all dilPlotem~ offth·!! ~irsllrIH!1io't'i( prepila,se ha'ribor .20 sets 'I:l~ ~airii1dl hlH'Imoogmlls c:n mmm:omes (lJi~alle,nts"'. A s.irn£II Ie bii'll'<liLIl !~8i~ 1lllmtlrat..e'llillt len; with Oliit. n'~ ris MA,0.21'1S,. USI"c '!l:N!lpa~ml2illl (If l:Iio'klgi(;a~Onernis~1)' 0100 r,E!'('01m Mnatio!ll. nU~1 ma~~M l ,~r piner\f'lalc:~~CD litio:s,omes; seg ~e!J.a~e int:o the '~ijrst ~(llalr b od~ (list PI!:) M.oll!'ru~r Philrm!ilool!J9!f. IHaward Mil'dkaL ScloociL Bmtoll, WI 02115. USA. ~DMs[lltm of tie!1'1~.ro~~ Brigham aoo •. dll!~~.ngMII. At Jl@fliU""ti,o!ll; ~alf of tl'i'~ (hromliJsom~ aF@ @xtJil!dl!ld \fiath@ s~((I!fld pol~f body (2!f!d Women's Hlosr;t1l<it ElQS~!lIIl.iMJ\,021!15. USA. ~ Hal'\l'ard 51@111 I?~~, and the 'incoming speum re-stnres the diploid c:ilromo"lor'l1I,e mmpl.ement IB!astn~!!; dM'IlIe(J1 bV (elt InMilu~e, (amm ri:d'ge. MA ((lllla. U5J!,. ~lDivi~iDn of f~FrHizati(l11l y-ne'l~ f€S: (el~. ~(G, hlllmgli! (li1i,orio~~1l: gll!li!adot~opifl1!. (iV, g;eormh~all iflilt.d~" nfl!i!eolfn~i~ NewOOm Metili d me, B!ig1n~JIi1and WOiinen's :tio$pilal and (hMrf!II'$, HDsp.it~~, H.!Irvar·d' 1!.Ii1li:li'llt Sdloo~ BiMton, 1IAA showfl at: top; h, hours. ~B) IlfIalrtn!!ml!g;cnet~( oocyte lIlIla~matioll.IVIU-arr,e-sted ,oocyl.es i! Fe activCl.t~d in poilalr body" Dl,ll~oidy i:5,maiinta~li'Ied am::! the r~5l!IUiin~ 0'2115" U.s.A. &(fililWf I'~r HUIINIII G~neli( 1Ri!~~an(Jh, cyooc:hillatllinl B, blo<.klng ~xl!rlllf>ioni ,of tItIe-~(ond ~~s~u~e~~ Gen~.<!llhl~p'il~t Bo>!O!:J. MP! '~znS"USA. I;Ilasooq!~t:s y~dd ptMIII)IE) Il:~~!s, IRI;lI:~!'!1!biil1i1ib1on ~f!ts· prooli! w MHC:-lmat(:noo ptMIIO'E!i (dts. (() 'PilIrtnen ogen;~t.klll(ibi'v:atio:n 'of il'IuiIlilllure oOOfles Uti). Inhibiting, e'KtrluSllQI1 (If the lirsl po'larr bodV "To ~!bom ~:o~~e~ppden~e 1!IlIG~dlbe add~ess:ed. ,E'Plait ~reVte;n~s.egJe.g"lfcm Illf 'tlh!1'hom.olog(!lu:!i (I'Irn:m!l)~om~ p<llrs ...E:Ktmsicm of a pill;u oo~y 1"E1'''lIor;e-s diploid~'. ,!l~~·d!~lI!K@'~ni,ldrl!ns.h1iI'!fit'J'id.ed.u

482

RESEARCH ARTICLE
<l r~~C'nom~ml:n kllO\\'Il w O~iJ.1IC~Il ti;:.~m.'!'C'ES ~I~ U5), MI!fIC:~iI1i1!i~~cliu~d (M~)~ !\1~~Its..]~l l;I !OI,,'(:OL1t;1: ~ d~vd['l(lm~1tt via ~I. p:t'Otrn;{l~ ~h,.il ~\~] l~ ~1\1lI'~1.hod ~.iml;:d!ll~l,;Ill,'li1'u~i!l~ ~~'lI1Clic'llJ.yma!dKId: ~n,L.;ion of 11lc-~'t,l)!ld p!}J~lrl't!:xiy. PltlltliullC 74% pES edlL ftuJ1fiI ('S7BU6 !Ii CBA F'I mtx, We in· .. duced ll;if~11C!1:(l!gt;nC'~ic dC'~ot;~rmc!1lt of :limn.• a of .u;ri\alA~ (Ii[l!C}'lt$ ~h;li~ c_k!vclb~ llo·b!kk>i!:QC}'!:1Itl., conloo!ioouc ClJ~~Irr:l 1]t.il!.c1l ~b:r om add~!kmlll ~4 !llfe nocyb;:~whi~ !~!"rI(;1J1ing wilh ~lwsc.gII'Cwtlign, 'Wl;'; !i.dift~'j[112 p( M II ~ES (:cll lil¥.1' (T;;t.Mc I), dOl}'!' in olid.t"r ~p l;,~nilun'l; Mi 1[· '11l~H£CU·It,;1'(P1l.;',". .. I'iOPUlimJti(m ~I]ilh;:~'ial. uf or Ih~r.iHUUl;d .lrbol1tnki~ou..~ ditCilnos!1Ii~~";!Odl,lri~g \"k ~~~Id}m:~yJiii;:t~I¥;~ eh!lilln.\~Ltlimi (Pf'R~ simI ([IIi :II clin'cliCfll~~Lr~ iiiJitpli [tal,i,OIl ftd lowed by ;I~I:dt~P('C.bfi.e llCSur.i~- .Iike (1clb (14). nim.·!t:JU~~~ld p(J\llU)ES re.~ls ~'i~1 ~h~ !irs~. meiotic 11'I.clnpI1al:\t: (MI; .fi!!j,. 1[,), llli:;l hacltlOI. lL'eomblik'tl 31.dte Mile .1(i!Cii bypolyjWOlocol h~i...~e~1 rr(\pot.!~ 10 )Ik'id pilrtheilO!k!l~ cl~~JlI11I..! digCSl.iOJ1 OJ' J i!l..'CI. :ii.\,"t'I~K'!iici!i!!\ .. geilcl~!e Ciil1lbfY'iM I.hfll. fire !i~il!elie clo:UI.'::S 0 f ~lc lIiul~l~~uclet)li;clc :jXllYlllO~lti~tl'i.~ (SNPs,) withitl l1'LOtUlinb.m Iilcll!~YSi.~C:iia)b'C!>sc(1 unllY one of [lilC ~he :~I,J: ro!:)ion, or chmrnOSOl1llC 17 to (fc~cr.mill~ Imrental [\,II ~ 1C"p!.'ol!.:ins ~,i I.2K.b rO:l' (;.) 7Bu(~ mice: oocyte douor b~i pl:c\'eninil~ sc~,'e~"lio:ll 0 r Ihc hQmolo~'!.JU~ matcnllm.1 and ~~"IlCf.~~I!i~hromoc wilC'4.h.CI' n:cCiJlthiu~:JI!km had remored hctCM'LY~ Fi~ 2B" ~v·~~ere,a." [VIU(, -rnat(:h!:di pt1\i~ I hES H2Kb gos:fty llI,t t~le ~mcloel 03),,24 mn orn p(.MIII~ CCU~,dm~ Imel lli;loombinoo expre:S$od ~l :iOllles ~J;;;~" .From.~hc 56Wn or aCI,i\ a~oo ()tOClrlC:': Ilmi. ,~k.:\dCij"'Cd U~1IiO b:1<l~wc~s, we isolml.'iI 23 and .1i2K.~ on <l~1 cells '~Fi~ .2('). 11l~ d~~ool1ES cells. ~3,:'Wo)h'lIool'od the hoter'O\l:y~OLI$ M'I Ie ,gCll{llypf:! Qr liw- (lil)CY~l'!donn!' ,mg,. S IA. block lim1 111(:helozylY,go:u .. gIOl10~}'PC by MltC Olm~g,~~l !ES cell linl.es, \\lhi:Ch \,\'C term p(MnES cells ~ bk d~", I,)xpn...~"'>1i{ln. Amnj~,il}' of p(MIUES ccl!lsimo ,~ n.. I). \l\i'~·d.eu;~u~!~lt;;·dg,,"llO~)'p¢$ of !he cvrok~; mno 11.\l.. S L B .md C). cl(>ut'£l!.y;pi~!gor (1lmking m~U:K~L'S(m dl:l.tIUnMOUIIll...~ ~,. COUfil1!l'~'\! ~10nntllk"l)lol::,1p>l;;' Uig" S11\; II = .16 U11;nTn~.1 M.~Ie fCgim'l or lite NM~.)ES ccU li!!~ by ff'R fic ~hC'MI [(' hC!~\rol!:~igosily in III $\ibsCI of eight .$C'" ~Ulry(l[ypC;; (Jut 19~inc:;.~cl'i~cd),A ~nil1.ority of il!!'uphlk~tion f[lII(l\'\!o~1 by <JI ~!l;:lll!-lq'Ioci resuiclion cn~y!1fK'l di.f!..... 'i!i{lv~or Cii!'l."Ct s~lIcnc~ng or ICCK"eJ 0011 ~;inC':\mg. SIC). Wcc<d~ ES~"CIL"l 3:l' !l(MU IES oolb Sht1WCld~!!~~~o.,,"cac X ·~hr~ SN.r lC!(;i ('Ind~rC!l'IlIo;;.tulI!~~ 1'7. and \Wln~CI'\'(;d MHC h~wral)'gosil)1 in 2~ cdl. li~b~ (fig .. liIlllb l~ ll. Pa ~thei1{i~e'm!'~t~c oocyte ,~d~'I!3:ti~j'iali'id (eU d'erij~3i1.i~i'ifroiii ai (5 7'IBLJ·'6, x (IB,I!/; f:l ili':j.~ijSil'. S1.A•. white cirek:5: ,n:) rug. S3 h Gr:UIOlypil1lg NlIIl'mbe-rs indf,c,at~ '~,mbryoiS. r<e.fithiI19 ;j! gffi\i'@ rFI $'~i:I~,eiit.(I~:aI illOcyt!l'S" of ~laul.I':iIlg Ilflru~'kcn;; on chrrot:l~OSC!'!I'~ 1.7 een.fJlir:ilgfl(y ,r;/ p(M'm€S ,~rl d~t;:"t;Iti(J1:l ulrmcd the ila(!tcro.zygO'llS MIJl:' gcnC!l~ypc and 2(!el~:i 4 '()~II~ Morul"l L\iUIe id~l1.til)' wi!.h lhe ,oocyte donor hJ .~ nllcm· 0 'bcrs oJ a selectedset of ~:2 p( M I)iES cell lines 1_S:Ofl'5,a UlJiJ.'5i(l I.ZSjil.!H~ U7!il!SiO 1,l'5Jl'50 721Ul, Rate of 'U1ig. {8.3;~M t;8o/~) (n%.~ (~4:~M (10·0%) dlllwl.epl!mMlt ~65o/ll MID -m<utd~oo pi MnES eellswerc Ie diffcof.@ir;Jency r;j p lMI}ES cell deriV(litfOil """1Iiah .. lor 1~ om.y... i~1 (:(1 ~ture Ol" dCI1C!'ihed ~ ,63!i~Q Rl.lte ,,,f 231,6.3 BffU1i: SlJU2 75/11, U2fl12 abt'IVC' and cx ..'u:nill.cd b~i \ cvtomCh"\I.. 1l1C {3:f~/~} (100%) fla%) {!2%} (6fO/~) (S6%) de~l~lj;lm~!1t di ITcfClllimc:d dcU'iv.lJti"'~ e1i:llre$;5~dlDotJ~·I [21{ib

in nv~ meioses \:vi~1yi-cld acressover I;WI1~.\iVc ~QI!Io:'A;lQdI!lfg"W:rc O('iCylCi:l from CS:7Bl/6 -.: C'BA r~m~oQ .<'I!1gJif! !!i;~",,-d (l'irl!hJt;!~~~fti.=1!:C·l,;'n1'b:ryo(l

~i;'!;.edin!ll f5 m<lnllcr

ro;:oHlbiinOltll,

M I ~'C'-mmciloo

nl~QnlC',

r(M!I)1;$ ~~,"" R",,'1;:!D:I'!llryjn,1I1t M~1(, .o\!Jl"'I,c;:h~1 p~ n~~s. ! ~l\. M ~ w~r;;: d~fr~IU]IIi!L'(! mo ~~nbT)'otd hodil".'.s, f~B""J ~ rot 14 days, lon{)\..~d by eulDlI~ 0:1'1 ~kltin.

or

or

or

~s

or.n

san

no,~~

mld

U2KII

0!1

~H ,cells.

oonrif!1J[i!1Jil

h~crof()'~

1il1lbl<e :2,.I@r~l'I:omQlro'l1i1ation aHtetr ff1~e~tionl ,of rlriffe~e,i'I'liat~d:fES, P'lflMOIES•. and p(Mln:s oelli: ~in~~ MIHC~a t,c:h,jj::!lf!d "f!riSlmatc:h i!:dl redpie !'Us., o!Il~i!r\l'~d 'ror] mo~(hs, IRggZ.-,I'-,,(iC+ m~(.e a ~@ onrh~ (57 B,Il/iS bad::glrIDUIlld. N l!ltrlber~ relprll:se:nt ter03Jtornas formed\llllJ1lt! ~ mi,(j!!ifl~~dW, (P <: 'Od)Ol[ortne! ~Yipoth,e-siI5, t.lhil'l hilploldel'ltity orfu:W MI"IIC mat.:lh prredicl5 t.eralom~ Imm.aition; s;ee me'till,od~ iin !illl~·

p orti I'! 9 0 mlillJ@ FJIIm@r~i! ~ ((IT

il!5:5I11.!'ilipti:O nsin

!lti:!tl~il(.:! l

(!!It!

~~is..)

!l:~{ijip~~~.t
1iI11(ltli'$~'

flES

~~~ls
(C51Blll6)
+ (41:4) ~ (0/4) + (5t5) .,..(5l5)

It.alJl;:!-I-lyc-'-

(M

(S7,BlJ~
(57B1J6 x: [BA Fl

+('sm
+ (3/5)

+ (~/J) ~ (ru3:~' - (OilS)

+ (6JS)

+, (.4'4)
~ ~(13)

+ {3/3) .~ (llif4.)

,+
-

(313)

~ ~014)

- (OfS)
+ (3{$o)

+ ~8J12)
.+ {ll2Jil2l

[(15)

~OJ511

+ (415)

.+ (SIS)

IJ'C'Uecl ii~gthe tCI'i(kli~.CY or fliimleES cell S [0 looc ruJ X clrrolnOOOI1rne l7:. /5). Nine ]'inDcs S1ilowed variable dbmmosCY.Il1.COOlltCil.U, A n1:1.)UCootai~od genomic 3lliJl~Y::1is, tllal (ti::!ltLl1gl!is~le5 these: dip10k] and .~ooHp!oid classes of p~M[)ES cells is prcsenllcd below, p\(1ri(:efltrQllru~fii:c, genotype: Gililld p~Uems or r1.loo!li1lbin:arUoi!l in p{MII) arid! p(M!lE:S cel!~s. To ilI~'l;WSS he !~,I~Cm o:f ChliOmo.~(I!na I I segfC~,II,iQ!I; ()b:'lcrvcd U!~cl,C'rth~ d iff1;~"Cnl. ooc:y~c iv'.lion

,of'IH! MnC~oc;llL) ~lY ~11r:t1ICCIlmI.Cin (..~':"'P~ ~i.!i'Ill !Fig .. 20.). ln seven r{MHES~'n l!il1~, we doclm:lli)JiDcd a. nnmliil] diploid chrm:no.sonlf: ecaWIlU by d~fC("l. ("liroJllloool1me oOUlII~n~ (I~g., 52C). SccVC.i1 11'(1\;111)85·ccllll,i~t<:shad 39 dlmmO$..{lll:l~"
it}'

IUl'i!ldlirte'mlilll1Ia~Bd' p(MU)ESoo!lIs

a Di!le:rentiilited

C 'MHC"m,a1iclillled
IP(MII~'E:S; Uilisi O

11UI'

MHC:'IIW,iI.k:l!'led

I1rotQC'o.l:;"~llcl d~wml iTU; ~h~ dl.,~Ml,lC 0 r gli;;.~ m:tic idcnL~lY of lh~ pE.S ~cn:;; wi~h [he {l{J~yt~ dul1tJf, we dCt>;;Tlnil1e(iI:h..: IfX:ri~le~u.n:)lli1ICrOC: gCllol.nlCfmd dishll.1 ~:ac()fIlbinllliml of iJ:Ir:p(MII)E.")

,0.

'.C'~

p~MII~E$ eel ~s

p;~MI)eS;oelms.

;[~ildIl(MIjES oell lint:.>;;I.IIS,I!~lg SNPs (flM di:>tin.. glilisilihe paU\::nln~ m.oWiSC :>tanil1l::i (C".S7BV6arrld C6A) {l7,.\i¥C hy]l'cnhcs:iz.cdtha~ p(MmES cdls '!htJtlM IEpredmllinantlly .I,IOJno0"gou~, wil.. .... h ':romlJinfltion rdlecrcti by (l tl:lmm:ric 1);i:OOomi,. I~Jllice O'~ heh::mzygOU:> SNfs. Co:tr"c~d1y.,~,,'C 1t!~(lIlled lilal. ~hc'P(M~IES cc]ll:;. ~'(lu!d be P",,"'domilJan[~)'
~nflct!.>d by
~gOLIS 01"

Hi21!(11i'I ~~~~~~~~~~~~~~~~~~~~~~~~~

u:l'

·it

,..'

...

hetcl,'Q:tySOLl5. \\'iDh l\."'COmb['l.la~ km IdmnC'I~C: redominance oJ hOl1!lfF p

IF!I!}. {!:. MHIC IP~oteilfl~xpr~s~i~11l p:ES, ('~l~s, ~ow (yU)m~h~C;d!e"te(;~io'l1 ,of If2K PfQt~~1fl exprl,lsslQIi! is (lifl F ~rnol;'llli! IJiIl (A) L1ndiU,e-rel'ltiama IP'~MIUE5 (,~ll!\.'(IB) rd~iife'f,e-nil:iiltl1!d MHC.-homo;zyg'oLl!i p(MIUES, (f) ,M11H(~lell'e:roZY£!l'OlJtS p(M IIO:E'S,':Iin,~'(D) MIIH(~llel>e'J'lolY£!l OIJi!> p(IWU)[S ,C!el~:'l.

SNfs. M.o~'{l"''Cr. ES oclis~~i1b~oo rmm d~~ ~"&ull.od l'mln ~m~lI['alrl;Ll1.iliZOll~1l t"".:nt~ b.i;lWt"l;:g'l~IDlil)s. or i~~bJl-d ~niLX~II S ~~ I~ ~:i:m~1-~ mm:ing:>~' sliou.ki Solinw 1l1.:kmqgM.i:ty :'1. r
(;f:llbry~

RESEARCH ARTICLE
:;;m::i:lt!scnll. ~:n~tiU!rlC Tcdu~oIClgyI\Vhi.lchC';jJ.d 9(" .-JII loei •. because the ~mctCi). derive ~~o:m hemomap, ""'~li(;h p,;'port~d a ~(lm~ ;jMI,IOO~,~ml! 11110Jp ~}'~QIIM' p..r~m~~n \\ih~·h m~~nl~c reeomhina.~ i ~rg'\;,! i~II!)' ~!'l\i!liih!i,;,WI; !iW~~:IYI~ ~lro~ 'e;;ngolh (;""ti]I~'.I!'!Qr 133·:>: ~M (I 9)_ im~ s .Ana,ly:;;l~ortli!~ R;"II;:ombin!:ln.io:n pitl[('TIlss, of 10 SN.P:s.OI~ chlro~~m$mlfllc n in T4rtMU~ES, 1:3 r(MI~ES C(l~I(:.. Slb!::l;WlC1d IWl'! disiinci ~iJibg.rol,lp!)" PtM .. IES. lind 20 rES cell lines. lind II tlingk: I pcC'.ri:ccn~ri 1Qli.;ri1;: SN P on c,!ch~ihwl!l1f1i,(ml1: i~~ Twelve ij!r~hc r(j\11 ):ES wn~showed a rred(l1m1. n~~ p(MmES <JDld r.\I~ p(M!jES ~111~il!~s lilg. Pll!lUll p'1'I~m of 11t'1.cn:m.}'~osiWI.l!t;gi!1l1iing OIl the CClilmli'i!;:'!l;, lonli!\l,/~-d 0]1 ~oml;; ditmI:lo.>..ilIl1Cll by S4}. ~~-d 01] .he l't.".Siuh.:. of Uli~ low-n:;mkltlotl ~df.l.lfil:tiC' re~og~ ofhO:i,1~!t'i!!llliil~' ,~.r-ig,.31" IiliJ ~\.":IlOlyrilig, w..,;: ~ck~[(;.J :J :':(:'1. of n p(M.U)EE and ::m j](M~.~IS Q~IL~ ,.n.d pcu'bimn'l.'tI h~!:1:h:oJ:- Jig, ~6BI,W'B~ell the gl:iimlype d2it.\'L of il:l~ t~: ~j_f1CS; were ~im~lllr!y c\,f1llilllt!!d ~ii100t the: asreSOhJiEi(}i~ ~1C~IQ~ypiill~ Wlil,g !II ~t:ilml,urd panel of i(l~.~no!JIsc nn!ll'lu;:r:> ,e\!Cil11_y )<lc(\d aeross ~ ~hc iCnml~e~ c.x~~illns~onf a 1~Vt!do!JI~~lydean. o ~~CI'il'!!Cc.I SNJ? $Cl (Nr,"~i. A t(1lta~ of 5 ~4 mm'ikcrs ~ll::l11illiIlS the 2.25 ~ipIl38'cs (fill) aCOO$j;, Ihe 1tJ, m.r!.O)lOllttll. wl.,,'rei~![orvn:t:[i''''c:Th!.')' were p.ol}'~ SlIn~i:i;Q:11 mdinn P'lUll:fIl. ~1K!CoCd. the Il!l<lP ~el,glh is "lI!P~ becausc jhcreeombi ~m!llc,hl'fl'm<l!f'QmC8 \ iII i~~ .... =>Gm~ C"f.JO~.. co-~grog:!lj\l,! by d~I](..'L; {or lJ..y nOI.1gi!:;j~ID!dimll'~1.11p!ll~ :I."'IIT!R,;' ~111, Iht:l1l'by i Jpf'b""iCI,.i:ng 11:ctcI1Mygo~; iy~i:1 ilte gl;nutYiJing ~1;S;([Wl.'S:',,;d ~!}' (Jig.

S!ll.

~lol'lPhic h!.'l~""e,~~! C57111J(} and CBA ,~ndhOld <I

((mlroIFi omn;:e!Jy id~l1tiJi!"-d
;t;;

ICilgl.h was ~i:Ili!kantl)l !5l1PIUC'lSOO (~) 10 ch~]t DCl'rjtc Ihe: high c!cn"ily ofIKllj'l11nD1~hic mafkm cX.<lmincrl. 94 ef ~hli!il~ 'lu,t~~om~. iUI Ih~ 12 hl~ Wr<f~ ,o~(:lrv()d w· be eon~plc~dyhCI(,"rOZ}~· !;l0llll!'l. (ng. S7 1,1l:l~~ abl'oCnJ,l!ion,~ (U!'ij\'C~h", . ~
;I~. !{'j_lnlPlU't)d

or lin

Fil; intC'p:ru;;:~, Ihe ebserved map

Or'i.)lghit tl](MHES cells ,ccnmrL\llnLliLllil, Clr.!li:!plctc hi;:'l\;)JD:lJ'gC!!>ity 'K"I~ .111 looti. SI)Jl:g1. !!!(,~~iD'i!5 .~':ll~tie .itk~lilY {If ·riit: ~u'lht~li.ogl;_"rk1~c Ci'11bryt; ~I;'ilh Ih~ ~lIle i!IDm.u il11 eITt:\e'~,II prurIh,:,j'lO~itC1te d(m.c. 9::: ori~kllllll'y k.'JXifb."tI ((6). I Io~vc~"'i:.aii'! or I'heei~~n I)(N.I:[IES eell lines Lhal:
~lb!l~1!

~row ooltllpkle ilclcro'0I.~~ity af:e hI facl.al1.CHI"'Joid. wiuh a Imls.e or dhln:unMOmc coums beI\"~n

3 fl. snd

n, Thus

w~ ~1J!CCIII<!!tc lal. Ihc~c ~

~'~ll~I'!i;jt·C'm"'}i~w; I)E.") C~U~(_kcJ'i~'c Ir'OlTi MI!(M

d 1i~~IJ;;"mzygolls ~bn';lllCd nl!;lp ~<t,;ng~hllWIl

to ·~Il!l:t f u

I( m<;;llnlling in

an .'I\'~'m~C· In!ctnmfk!.:f disemee or

the !~O\ll~~·ES cel~ and

<l ...mndmd

Fz n:;comhi-

ra plrl'id h~'l~U)C:t.'l$~l.rtw ..~nir~ inMdi~:>' of chrnn!t!stlnw] ~~Dt;g;:J~~ll cll,ll'wlg ,tl()ri\'alitm ;rna, cl!I!'I~Lnt'~·Indoo~~" ~"'h!,.'n oocY[C'£ .. re 'Jii.;~i\'m",,'tI,<;;0011

4!.J~ Mb}. SNP !;leno'YpiP1!il, t"{1!11fimlcd I~I~ hYllO~:.hc$i~oo f}'l'L(i,}m.." nfre(,1(lll'ibulfI~~ mll bl,l~ ruve:d!.,~ ~I };lIT~ ~ pri~in~y ]~~gh dl~grt'\C OnK'ltlo:ffizygosity fur the ptMI01ES (l(;lls. l3ec~l.use",i51£( c.l1ro~n:lInd ~c.g·

re.!1,llliol'l

nrl MH, 9~1 chronnosmm!!!. u~ear ~h~~1" ene !.I'mne~e."for om: of (I,IC two Il<IJiCnla!sl m~ns (Fi£,
lis;

preverucd

well: Sllh."mnli!ZiUy

IlO.L'DlD'l;YgoUS

3.1\ am.I figs. SS ,m~d Sl~).3:ccauiSC orlTJ.'Cmui:lin<1] ~i[Jn,lhe hC'~,,)ozygn~il.y of SNP nmr-M'\J'S in(i!i~JSI.>d in ~l~qlllC!u::}'vn lll·aorol1i.o.n to I h~ f,!.!:;'ncli·C' distanec rrom the CCnlron:1Cl"C ~f~, 31\" I:1Is1n pli!lllc~:m~(~ng, 861\). n p'l1C dl.C' 1)I:\:'S;].m\Plim~ es of Ilrodomi nant hom_ozygmily in I1(M I] IE.~·ccn~, we found hO!11{11ZJI~~i:!ym O!l~Y 3(i._~~ orloci illn ;JI. S'l.ll'vcy of 17 p(M~]JF-S (id~ linC'S. wiLlil .l r.'L11W; OnmTnO'£,Ygrn;~~y bDt\!i't"Cn :!J._ 9 mid <Ii,., _5% ~lig.

A,,1'r
f=o.oot:lil!~.0,3;6 ~'!)!l3et_

S 71. These dfllfl

dCI1ll0Jl:> .......te

Il..u,

ctmlrm:y

ro

C'X~J!...'Cl1llDo.n. :r'ocomb~lbm:[on renders lite mojorhy

of loel h~ p(MIIiES cells Imtcmzygotls. Some p(.MmES cells htn.tborcd gcn,otyrk'S

Ihm :ililo\Vcd Itmuozy.ga~il.y of SNPs .ll~\JI~n one rarel1tai $1rn ~I~near the a:ntroml.'1C. IbllOllI\'Cd by ,a, .rcgocm of h~l:cro.zy.gosh.y, <1m:!dK:il a I.CJon1.(:i~'C rcgi.o:tlofholfQl~gosily of SNII!l:.Irr-olnlhc oiher ~Ilt a ~ s'tmin ,~dll.'01:110i.o;omCS, H lind ~.O ~Il .F:ig, 3.1\). ·nl,is panem i:i;, consistem w~lh mciolk L't!~

!Dim 2!j)SUP Irnil~r

40

1001

00

1001

·t,2!i1

(1!!~~_!1r>!)m

cen'!lO!f'!~PO ~P!I)p:~.

CQmibl!1.lil~~{ln OOIWf;IC1l ootll sister dumnmidp;)ir:;. or ~'K)!I~olog(nl~chmn!{lwm~, whic'll occurs dlll'im~~'v~ ,ibno\\'\;~ ~"':'iga.·''g,,!.ion i'. he bi;vall.i!llz.. ~ by il'lio ~qllmH"': i:.\;! L~~p'£m .;xnru~:i,(~n l!he l:illOl ~'11m-body •• nld lIl'lC<n oeU"""l,'gn.:g."I:lit;!Jl pf Iht~ R&.onm~ Dillfntl s.L"'t~r ciUlJl]la!l.as.inTo the :;illlll(; ,c:x:Udae willthibiliio.11 of cxtru:;iOl1 of~hc second po.i'ar body (lFi,g. .~:B "Vh~bo:tlll s lster ehro~lat ids 1J1iI~ ~ ~C'T.go.~f1;g fCcOI'I1Ibinaluo:n $cp~rnlciy. we [jOSI,!JI(I

or

I(ll~dthm p(MU)ES cells, 3.hhtJLlgh

cOll. .imil1oo .

i.e be 1t00nl;,~go:uS at Ihe eentrorueres, wouki ~nanillCst recOlnbil1!<ltion <It ,I mle I:h:m would be eqn i.... 1.0 an f:l: in.!crcl'Oss betweea two f:1 alenl ~1lice. \llte ca l.I::ul!<!I~cd the gCII1C'l~c linkage map lro~n Ihe ~~tj\lm)ES g~'\rm~yp!."I). llsing lile· pa:~ .gmm. MArMi\Ii{ERJE:"'{P \'crsion .3 lUldl<'f lh~
II!"I al;ms.s 1111~Iu.lo.k:ngtl1 of~329 EM .. ni~.~ i~ bromlly Con ..i:.[('lu \...iiJ.1Dhc Mm;.
iu!~~l\;'m""". ~(m1\;':>wc ~~~linia[l'd II IDlal Di'iap

!ilIO· DI:ICBA SNIP I.:IHIETSNP'

sw>

,,,,.Qi' C!!1!'l!1iI~~1~!li'I1 ~ntlll!lllr<e

go

!Ii!!

!IJW

eo~iOW~~Il·

~"'!!I!I'

ml10dd

or ~11lf:!!

tFi'!lI•. 3i• ·~n(lm~'l1de :SNP gtUl()lyping of ,ClI ~epiit:l$~nt~lW€ dofie· ~f p(MlOtES. ptMO E!), and ptlnh~e,ro\1;f!lL'UJiS' t~(M®ES H'iJls. i'Aj, p(M~II~E.S't:elis. (() tll{MI)ES (eli!s, ~·(l!Xlr1hel-euQ,zygouso tptMliE5 relJl),~rt pa ne!1s 5iT101'1 t ge:nelYIPes fo[ each (hmliOO~oFl1e..h10m .celillm:IlIlIlNl ken" top~ to tfl~mer'e (tet bottom), me!i3i~ing b!(!oI;ks~Ilf th~pIQl)'pet· ~r m!l!t[lt~r5-. G~!.ln, htHIOO;zyg~t!s CS7~U6 ~B6) SNIP; Qr~nl1l~. h~!IIl~;QL!$ eM StMP; bliJc!~. 1l~l!r'tI&)!!I.Qm.(Him SNIP. PJig~t p.anw. ~hO'w tilll! h.eterozll'g~i~' of Sf!lJP rna rkerli plottedt ilgal]liI~ SiNIl' NI'k't!r dht.:inCi~ tmm ·~fle [t!I1I[l'om.ere.

484

RESEARCH ARTICLE
arbor cy:LI.)c~<l!<liSinIfClbtunenl to blO!l;k M I,
K'11';l~

"IR~ ectedenn
!"kinl
~1]~1 wi; Wa:'i

r!oi:d b!a,_·.wij;;YS~~ r"''X:ltl,f:!l!t!}' P:5ll! II Ul.f}'l, SJi' ruN S9}, S'I~h I,lll,bryg,s, \,\I'il~ )'~(;~~!l,;'I"lp~oiii;! l)t1\.iU)E.:o;; 1;1,11~. mId it iL~pn';.\;:ibl~ dKli im::lllkRliirm '!.~'iih

lF~.g. "'),

IlmJin. '~m:lmH)C)'~c (iris), lind

l);:5.ing pn;,;j.(jot!~~}'

ho~ t[}f in \!~!1Q d iff'~n;rlli~~tion observed d~yi.hmk~~~o:ntn~II;:~illilyimE~s that

c(mKi~IClliWuU~ ~rudi:ml~.y{jcyi:C: .dc\oIdtlll.lI-

p1!b!!r.:h~d 0 !,~S~Il:'l,

~;s lines.we ~rIQ~d~raI15\1ll;IJUllmklfl excell pcrim~Jflt~ !P1 imm~m'Od~J!c~crll miN amJi imrl,;~d~li~I~!.o;; !hrl~ wc!'~ ci!h~r I\IU IC";ll1:1il;:!j~~d or .mi~m~u)I;;lh;:ll\lVh~n. WIJ i~lj(1::h:rl 1.£1' lDldi.U1i. ... r\,~lllifllC;~drES OJ rES~:.cIl~sab~~.l~!lOCQI~P-~~I~1
CI!I!!'~!lCQUi5:1iJ

cliIlIk:ilhyhntn:irltlp1,lrim;

n1:l)'

tli1~TL~.Il'L"tpn:~li(S~~~

:.:;kCplic<~I~h,n~,·lL'cp;lnht.:llog(..'i!lC'lit clones

~h~~l I(:! 'lhnomm~ dlmmrlfoQI11;li! .'iH;gf(l:g<J!~im~ r o c'hmJ~l[l(,;l,1illl<1~ I,['j._~s_ SUI;:~ i~ 100"Cli, 111,1 inwi~1 i;mlilJil¢l.c Ii.~CDt~zygOli,i~}' wlu::!'1 gJ1:li.otypot1di iii:' <~ f1t1pullHliolfi bUI. do tint [';('p~1:n ,a l.aiLl!: t'blfiL~ cc:Ul'l WiIh gcl,.cl~t" itleJ:lt~l)' 'l~lhe (i(]oL'ytc OOliOii,. Theeraical ~y,.1ll1rr111K~K$Cit\.~1i.e cllfiilCflWOlJ]cl ru'i~e (I!~Jy if the roool1j1jbin[llljl1~~ events j\ilj are SlQl),l~It'S.'lCd (It," ir dlC'J~ is i.li~1i:~1I:tc cO-S!.'gr!i:g.niml of .he rooolll~blnl.'lnt C'ilrnnmoo.<lmc$ ~llt:D the smnc daughter ,~n. ]3:1I!."'Od our c"'!fl'CL·i~I1Cl~. rule' on ~~'C

meillun pM]. pMII, ;1I~d IES cc!!s" ,,1:110 'COP11· r~f.d)I'i; 1~lmh(lrs oO~' hcm'l1.o,I'l1.'1i~!ic 1 c~nu;:n~ ,,~,
m~o.("~:-;l,Jroclb>, nic nhyk:d iU,~M~.baliccl (;oll::nl'i}". IOtu~ling ~L~I ~I~y'" ~iJ'icl.I1o,Y ,t')',ttlliJU(:,lric ml,i;1.~Yllb. lOt d1>C h~I'i'I~I.oJlIO,k~li~.: Illmkt.Th c-K~[, CD41. and

illlO il111mUP1CiOOl'l1llll".'!L',:nl mice, we

~iB~I!,Id:~.
~!ldk:.~l-i'

ClI~C~

dflC,iI,;m

'(i;r.lti:l;l1111

rGrrm[~i{:ln.

or

or

C,H~ be

iso~'I!!..Icl.. lI!)'iUe'rE!l1tia'til(l1'l1 ~oh~:III'~i<l1 p(M[IIOI~:S a rnd of p{Mm)ES(~Il !]ne.~,Ttl ,ao;..'IC."S (II"" plur~pOh.mcy ~hC'M[ !C-!lm~dh~'{i pES Cil.,'n~... "'~C !I.:\'al~I"h..;d lhcijr dirlb1;;l1Ii~timl PO[~~!li[~1by un t'ilIP ..ndiu~ \'i:\Hl :)I&..;;a:}'1>_ \,Ve injt!cl~',(]. ce~l~ £W)b!';Ul<JnColl"ly i,lil\O'

or

i'fmmJnwdl!~ick,:mltiJic~ flill!t11 ohscJ'\'cd nIDUS!

l("ru~

,(MI ):EScclis. ~'Iiqoi~ogyor tcr.:1I0milS. made [rom rtL'liil)E.' cells rerC'.dcd li"Slilc clemens of all ~h!'oo '~:,'fldll'}'tlnic Wo'!nl !;;ty.;;:ll': I1lCMlCicllm ,!bOl.lI.\ bone mmro"t.. muscle. <IIfIOcrutil~'C )', endederm ,(~pj.I'J]{lf)' 1.'P~~lclilllnJJ and c%OCnnc p<l.nclU!J»'. W·ilg. 4.. [HimpaIDology ~~rat~rna:s '~m MHC~ rn~OCh~ pfM!).eS C~U~

Wfill<l ronnmDOIll rrollin mLlhip~(j po(MI[).ES "'.~ = II ),P(M ~lES(il = 4), .and~E ~II = 3)relJll'ill.(:S, The' di~cretllimion ]OOtemia~ of p(MIUES cells tlo1lSbeen well rl'oom11Ciln~xi (n but not dlJlt of p

n:.'cipi('li.'t lin1diIT~i'Ci~tiaktl~1OtI~ ES ccllki do !:len tNprc.<:;:; COO llO1 11'1,:,:. 10, A and B I,We i,\~~11i.:l"flli:d M.I Ie ;fl1:'lIl~'C~t~, \l!Ihich V!'C :;pcewlhu.cl'(,'iJcld~ffi l~leil' S chi~~lc:riemice by illjo:til~[!; p(':"i~U}ES (Ij - 4) ~mJ ~h.')(CI'tili:tlc10 U!::;iocl~OIJl 'by N!K edL~ {5 ~ MI ~:(' <ll"'~ ial.io~~ 0 f ,'fMI):ffi '(I,' - 3), ,cc~b into ~'Ci:piem 1l!111.~OC'J."I~. iilnt'~&!\,i!n$ expressed al~cr tli f[c'i"C n~l Exm~:q)Ie:; (If I}(M[ljEo~ and I~~M~)ES C~H:5 ".1ch I'IK'lli!SC ES 'C'(U~ in, 'Vitro ,(.Fi,s. 2}. To dielcrdC<llmn."i.tL'3tl!d rem!1 lh'C'L·chin-crism ru,d hisJ}·kvcl ~nhlc \,\1 ~ICli~CI'di ncrenl~a~cdl issues dClf'ivcd sk i~~ ICh~ullurism (If ncluh mice: ~F~g. 41.), No ~}Clm ES cells 'wouW be "eccp'lodil:;;, !~~~ue li,,'l,.".rm!iu1,,' 'tr.llIl:Sr~li~~i~n ~1I!~1~1"'''' I1lm ~hc p 1 W'tl fn8Ul !\;'Ci p:ul;'!l!. rn ir;~, wt~lIw!l.:-di nl.\r~~ll~ iill~d (M II IES er Pi MI)ES (lL'1 ~W,ll;; !l[l..,;,d ill 'L;'iglJ:l ~ r::s ten:!: i!~~iO' EB:~ ra]" 2 w~~,ks.. ~h,,~ninj~l~d Ell cells ~\!Ix:~nOlrK"!i,,}~s~}' bnvmmooC'lk:i~nl ~mo nliil~i~~~'i, r~m"lcC'hij~ru:!Th";;; ~cnCII'<l!oo Il1CIf'C of Ihilt ~ni\:IC or in!n< .vmnumo()OnlJM,,'l'l..'J!I MIIC'-mall:hod thall 700 ~iOJ;lICHY. MC'I!'I;,'4'I\l(,.'I!'.. ~~j~"'!ior~OrO\l4.'ir:50 mld -1l1ii";l1IlUoc:b:,\d l\:'(;i~l!s,_\'\1.:l ()~lV~,,'d~I.,,'JU!o, H,'!<'IfI~oicl l.;'l~b:[yO~\""i!h pri'vll)ES od~ (~1ik'(l 10 ~'~lU~[ in ~.~~O\!; hinh ... il;(Jn5L~I~lll wi~.h Ilh~ Iknown , mlitl5 in HJlluniru wl~n we ~*"Ctl,i'dI'lJt'>4in1'1\.,n.u.s(I.Jd, IES tldls i:Juo ~l1'ImurltltkficuL~1I1 Rag2-'-t)'(-I dl~\r~lopl1i1lcru.a1 li:JCn u1:llaon Dfp,mn.hcnogctlclic ~ni:ec(rnbJe: l).lnl kmI'1I1UI(X'OD~'ipClC.IiU rocipi:elli. mouse l!mbryos lH A~[&~Ollgh nOl !iIi.Uy compcrent 1.0 stll,DrIill ot'W1nisl'md dC\IIclopm.cmbc. unlioc.. we Oh..."'CM'tI teratoma l.iYml."ltiODltrotH prec.msc Qr 3, lack ofpalcn1.(l1 iimprin1S.p(M.lnES di flclfClllialoo fE'S illUlp.ES cells of1lly i~ therewas uo ~';ill!C~llisma[ch \-\~illh Ihe re:cip:ient (Table 1). i ami p MI):ES cells ajllilC1lf !O ~~hallCa rornpal:1lllc' degtee or nllllllj,~inc,Jgc lis\'iolliCd iU"crem itltiml 3$ All ES cells d1Jlll c:mriedlll<ol.hCS7BU(! :m(] Cllil.. rES «II~ Nli I' :gCOll(1'lYi!!C:!i.r;m:ce:f.l.l.Il!UyCIfI,W<I ncd in h"~cr()(f.),g(11Jj1 s M I W ..~l;a'td!OO C 57 BU(l )( CBAF! !iI~st,(lliXllli!lpii!~i[~iUty' o~ dijf$ere!ilJt]at~d [I!I:U~ !'f,. elillY QfMH(~miilti( he(!J[ Il(M~n ;1,IlI,dpl~'M I)IESo(.el~. .n::cll)ien1:.<;. f(li~cd 1.0 fO:rllilleralmna." illl rni~but r1.1ClldlOO hOlllOZY!:l{)US C57BUtl or ('SA l\,'Cjll~To ~C'.st tire hnnHme c{ln1Jnllibi~hy or selected

of I~~~gJ.-"nolyp of Ih~ ES 'LX"II1;;; .. -s o:r I'iliitt:. Li'kcCl:IJJ). en'ib.ryoJ'iic t issues,

or

'Ci!f

dlnd ~Il:inlddfilleri:;m. (A) (amlaig!!l',.(B) Borii.l al'li1il iJ01iI1! marr,CiW. ~OMlI!li~l@'. (D) [Brain. uE) Mmli1loc:yt~ (]~is). (f) 5kiil't fG) [R@~ pir.artl)~ ep~~heliIJmi. ~H[) P.alnC:~~i, O~ EI~ad!:: 11:0('I~. ,oolor clhimerism re"5tJUingk,olilil p(MI~ES (:ell:s iiiijew~clii1loo bl:;l!lIr!(.ljSl5

(r,om 3 ~lbGJl Fl moose (whil~ co,.;t ootsr).Scaloe ilatr, moo 1JI,fr!,

RESEARCH ARTICLE
en!!:'. {T<IIll!~ 2:). I:~mnozygm! ... rss {IT IJ{MU) ES eells ~,gm !lQd ~Il M! N':'""~n<lR:hc(! hom(}7:ygml~ ~~Id h~~.~m,';Y~o~1iSi .~c;!pi~~ll\.'"a",i:il~ ~J, IJcc.:llI[I~"~
ES ot;:l,;::lL~f the ['SA o strain !hn1lll lerllt{ll~ltl'"i~~

A oo1nil1!:, hi,ghly ,el1icie'l1l mejhod

ror

C'.x~

~b~lR.a¢/y~-I- n'li:cc. \\hic-l,l an; m~th~,C57BU6 ~kg_mulld., \'11";; calIC'lude tllm~~ho i!11Imnll!~C re=-ponS,1;.

;)Ind

11O~ hO!'.l :s:t[<lin fitC'I(li~.

i~;;;1'I.~[Wlh .. ib!l:L)'
im'i1l1lUltJ-

foit the 11111uit: or~emlmJll:1 fontl111ioJI ill

COli:i~I£llt

MI :~C~ll~ism]1[teI1Gcln:~t"iJ)icil"'~. TIk~",", dfli11 ~tl(lii['iini!l1,.;e.hc hi~loco[upatibilol:r of diUl~I·· CFllnmoo, lis:~e;, from rES rutd~ES cells n1lfIL ,:.hiii1(; g,el~clic kbllily III. lhe MUC IQC~\I:,ith u'IIn:!{lk'ml.

reciflit:lllK
1[l'is;(uSS]0i1. '!;Vi:' ,fcl1cribc two SD1,lIicgics rOT i:'>lObling phllri~tcnt mil!ri~le ES cdl}:; tllillt rule' ,g~'ll~liC'<Q~]~ ~!~Jlthcd ro oocyte donor 1tIDhc M HC' loci. By "rllly~nllgL;'!lol)ipjl)!:l OII~.a.iy.~ijs ~1 pES. ~~c!lsr.ollll~d fitMflh)'b:riJd CS1BU6 " CBA i F ~micc" \~"C~m'!c~lc!Cled ~:ines "hilLt 1~!;BIitl Ihe

.h,,~

and ,~ !;ItJa;:.ll· iIl1IPlol)~~, :J!laill \'!Iilll ID n:11unOnf\hiplE'lwt:Oil Ik£l"'l"~K'Y h~t{''n:l\lY5'1;)",iiy ofm;;irl;cn;; an:;~ di5~o:.; linm I(l:!c C'C!100mCfC'. g ions ~h;il1~;IW reverted ~. ~lCil,i..'ifZY.r.}'£'(:I~ Iy beC;:'U,I!.~ ofl'1i.ll,;O'mtl ~!1<1~ il(ll~ CV~I1JI:.~ dLIr.~!'!g Ml U~. Beyond dG'mo!is~r;:ILI:n.,g .he 1:l1,'i1.o:lti.~ hiSfi:lr I~Ol]tl~E~~ih~] ~[rOr pm't~i('liog~'1I!...ic.. ny d.;;ri",·od .e OU r oal,u d.;;mo"''litr.n~qll,;~1; 1U1:J:~.\P;;C'~L-d~ ~1,1:t y th~ mllljlfiliily of .!tilti inplMmE..o;;rell:;, L1lirlC~iI li~uo.. our 'exp"'~rnlti~ljt'i ~'iiruvi(l' ,Hn ~1Ti~llyh. of ... n.'eo~~lb:i'l!lni:o;rl, Ih~re'by geil..:ruLillg II ~~rodo[u. gCiiclnc p.."Co.lulb:i iiallOil dill~~ rf~r:!hc~w&-~lt,,'ltbC r'ig :.leli\'i1li(i!~~!ind d1iSlinJ!liJb,hllt~ !'tm~em::; Qf reeon» bl'llmll~ 1t~ti:ro~!~Ili;:; ~~mme, Sj,m:i.b"I,.. ace hm· tii:m or ir.mmli~!lII1'C: Of;}eyle.~mld :iIl~li~il.i()n of die blntlilion o'h ••el"'l',oo when k.lllyo~il'tCs,i.~ k:; jJl~l!runcin~ic diviQ:lem essuee Ilml.:-lI.h::1.t:lllll~iml rilp:tcd dillrill~ l1!IiCj{l~is~.~L II. ivlo~"O",cr. we nole Ibm i~(lkuioll ()f,~(MIIIES ~cll$lb!JlkDw!.ldl h)1 SNP hcwllO"l:y;~iI.)i'i::1. p.ro$cl'\'odru:;m~lhe geueme, Cl'I,CCpI ror thase reSiol1.,,!hm com:~\fllW homo ... gelID')'!}! ~,g 1)I"O\'~d~ 01. l1C<U1.';'of l:,'C'H~~k m:l!.rlliUlg 1 zyg(n,~ity because of !\.'lC'Ombial(1~io~t oth I~Mm of loc ~f~,~hlJ!1{1I}'I~Ih..~ COI!l be dl~fi!lcdlIY'ct!ilh B
Sfll,:'lfJll

r(!rimli;fll"~lfI'iiWiIllI;J!lQg;~'lC:.l~!'o. iumiec intern pts ... j\IU! 1mdf!l."JM1~t~ijl~<'II~ ";!~1bl)·!I;Inh"l C{r!l1'i~5a ~tLI· r!H;:tll~.,d h!lp:rQ;dg1t~II.mne ~l:,'S 11>1,,);11 tIi!:I!l ~~L"ubt;:d

()f ~nct!c n~ljt .n Sl"Cale:r dl~JlOe5 lIDnl~hc OO!liU:!;)mI;W.Simi!rn'~ •• lnES ccll llnc derived ~to!n :;'I. li.11!i~iZl,;~~ h!~"'~QI;y5Jslloll].I;J b; "'OOHibinl1l:~ioJl of

m,.

r1fL"l.Io:rnirl.m:u:1 y

liaClIIilO'.tY30us..

,~~C(:pi

~{Dl;-re-

or

n~t

~IKHCl1m~

[\.Ii U

,g,CI10~}'po::!

by .~i~lIllC'

of

~t)o;;i llc

~n.'i(uii; !1.\t'Q.Jinfuiil.l!il10n(l'1.'Il,'H!:S ,lIld i'11hll.~ri(.MC~ ~h(: mi..n·''llil :si~l'~l' h:r.t:!m;lI;id~ \\'i,ihL;~1 1.l1l~ g~" ~ ~1.\."Lic:'(l.~j;y dC~llL,d pES eclls :JJ\: pre ..dm·~K:'lrnli[ulcd i;IilI.o EOs boef~)1C t]Ciil,!l b\l;ccl!..'iJ bun immunt:!>C"OH'IIpc[(;nlL

or

(1 r I)ES cC'n!':~e! a iln the mi~oC'hon{!l:iO'lJ genome (If Ih{!' PI1C)'IC d~.m(Kf. lmli~~ g~'leli~;<di~l~muC'h(,;'d .

..~~dP(M[) ,c~H~'!:<Bn s~k~(;"!('dto nl<li!U.~iL1lhc be l\1~H'CI;lCllmypc ofl~u; Om;"j'LL~tmor; A~Iff'i.HTf~~ d

~tC

f,ll1(lyol" E.'i cclk

l1!e~er,e:n;oeo5· ,a.liId N1~tes

ES oclhl

1" 1\1, 2.
3. ~_

dl!.ll"u!1;

Cf(,\;utl.i'l;;t~))'

111.1d~p'

'tr"dmfi.'f ilUo

t~lCYlll.":;.f!l:"n~ an u~blChm;:d

donor. ..nd; 'eht~K~furo

m.ay ~K~irl immu!iologie .ucjL'CLi:tm.dll!lc 1.0 :ll11tiigcu!i c~t~dJ!..'li y rhe rl1.utoclmrldril~1 gcliJOIi'I(~ (12). b

1:\.'C1PiCillimlee. the tl'i:SMeS cngmJt. as long as theM I[C genotype 01" ihe donor eel ls is ~m[chcd lo~he I)IX "p!e111 IlIDtl S;C'.
/I.ltlmllgh unableto
sll'llain

Usi~lg a protoeol lh~E. ~f~d bcClil report~)(1 W

CfClll€: il!milhclrog,cl1.l!~iClILllyc~mlcd I!Il:lhl.}lQl'i.( I til. we £,c..neimlled J sc~ oir cir£lhl~Ml1ES cell's dUlt l!e!lEliucd ~omp!c1C' hc~elfrn!.yy.osily :II a ~loci, Ulll... l ,]iyi~lg l!.C!1CI.ic ~dc,mi~y wilhlhc OQcylC donor. !lowcver; Ut{.~, eel] ...\"'~'1iC' ,Il~ ~iuher~cl!l·<lP~¢lidr o uflct.lpl()id. su,ggo>ll!t:l,g th!ll~ tliIC)' arosc[l'onn em~l}'QS thm hac] fll]JedtCl CO!1\ICIi! to diploidy

5.

S. C I!;t!"t~ K, H1H~ Ill!. ~rnil:, SGr~ n~ g!NtI~t Bill.. 1413 ~1 'llM~, ].. (I" Ci~em S .01•• :>C:limce2'9S, 819' (2002~. H. lin !L'! aL, Sfem (t!l'~ 21 15:2: ~O~., L Strrain" ]. P, 'lIi'.arn~; T, .~~~Olil. III r, BQnU'I1VJt ~ql, ~. :U. 16<1 f199!Sl. 1i'II. M. Ridel~t 31t11,~_IH'OC~,et;ilil19)!if,'". ~, G•. Q_ 1l~1~.
{I., 1I(1en... M. A.

ltiallmism..

Ctl110~'. 1.1 (,2'0!J2).

I"u~II01"£'111~sm<ll

dcvclopm.cnl because of li~c 111.'C1\ Il,Ilicm:1I of ifnIWil1t~, ES eells derived l!;t'lnl p;~rlh.cnof,l(l,," JI.~'1i.c ,cll'Lbryas aplPcaJ" 10. 'ilJ,~ iP'h.1r:~po1!C1lu. \VhcLller tissue dmcrentu;uion and CIll:gmtlmcnl into ;;)IJ] ] im';:;,1gC$arc ,l'OhlD.'5l and '\,-hc!'Jl1.~ lh'C'~o..,,'" of

1£. H. l!aQl(if'f; A K, Tar~li:i.l, f.m~f. f.xp. ~~li.1'l. !15. :2'~ U976). t. E.l Ro~~rtsoo, M_~_ E.>Bi'1!;, III. lot Ka'l!Ifm1!l~',I. ~ fxp. MQ.rpl'!'''t. '74, ,9~'{193:3~, a, M. A. $I!IF~i, s. (.Bartt:n, ,PI\,. L ~rril.~Jwg .;!;Clll,5oi16
(l9l!~),

dl!.uing the ~)Mllh'C'!lO,g,CIlC~'C.:lC;li\'l:!Ji()1!of iun· [I;.;I<l!UI~es~!ctc'~oZY:!;!Jlsi~)' f critiC'.1I~ o !ilCIlQll1k: lfC,gi:ml:sn~ ~gll~ n1i.l!lUil; O!i)C}(~l':jl .. i\.~lhou§h t~l(;:-li,O cd ~~ t.,III}, all 0 frht;; nmh,,1I1l!l111 chmmtlso:m(:'S.. Ih~}' d i:;'[f1,llpt c'~11 ~lurll;:l~nn t,:<llgm 1.led ti :w;l!lUC;~ rt .. M.~n~ in
llikc~)' to he 3. lot nrnlL"'plill.lIm irm, TI:l~ I>llllus ,of mmcnml· or pm(~nlfll.$pcci:l:ic blllJlrim genC'SCfllfl ~ .I,mmr~vo:rcdo icicmi fy pIES t cel~s, t\mrlysis of t~lelintpri~~le;dstmll5 of th.e 11:Il11~ph:lIJlnmkm. .Diffcp!n!imtcd li~ucs lirmn IlES ccl~s th;]J1 RasElrfl. Ilocus inll, (1 prM I)E'S cells, cOJlllr:mcd t!:le e,,'Ij,pre.~, onl)' olle or d'lC tw'O SCI::;,of Ilarcntal [\.'11Ie m,![lileu'IrnJI~]I."ttteun or Ulrotily.llru!ion Oi,g. S II ,: hO'.\,eve •• the <l~'ljl1Cm of ilnprim loci in ,ES ~Hs ~ml'JilmYPCSf\Ppe<ll' 10 (.':llgl':flit in hClemz:,vgous .~,. dpi.cll"''' ~f(Jr CXflI'l'nllle. C)7.B:U6, imo C51BLt6 >; U'I!a.y Ili;- mnc:1 i,lbk:- givcll l~~cir tcndencytOVi'<rr:d . ('13,..\ ,FI l\,"'Cipiell's~ Tabl\::' l). TIl.is, ::;ugg.;:s,tslhie cpigcllC'Lic in ... "4nblll~y V3~.'['he 00,13 IJU'C:l!cntcd hetiC dCJnmJ.....U3te~hal disc-emitl:!;; IDle distim:l. pm ... l ~hC'Orett'C;d iXI:i>5i'bl'lty (~lo':llt Ol. moo~"SHiz;ccl bank i rl1['J.ilil o.:.:mm.l qLl'"-~lion.'i 1.2''1)_ T~(':;\-e ,tm'il"('u~ J:ml.~ wjll~"l"mflnlding. Out dam (rom I.he IiI'iIU[lItC SyS!,('fjl C$limlblii>h fII prtmr of ~li~LI1eiple liIIsing pflirth;enrJlilr genesis m ~'CJ:l~rIllC MlIIC'\m{lILChcdll')S~i!s for
are 110t Ul:lr Cl0~bCli ~Jlldfire !ill[

ml~L\\o'Onhy 00 llOCt of

I.h\:.i,u(",

9. (.]. TOI)lDr fl' (1'1.. ~.~n(;fl:~6>6.• 2:019 «lIDOS}, ~IJ p, !'IggL!.rM ei 1lIl., !m.tmmLl"" ~. i1.!5is.:n 991.1. u. G, tlJlpOO'~ J, Wl!ljoi,d:, I~. Ii)r:iIi~r. S. Sch~!lr., )... M:IImn~. iI!w.. lm.trojn"'!I.~Rff .: ~ 3:3~ '1999t u- M;iU(il'l~ ,CI!ti!;jjrrI!J,r Eliot~lii'lol'~ inIOr~iiii~" B. $, H. ti!~b~ M.. 'IIi. HOr~rrBl1~, p, It mnllilKt.. iI. IBi!I'~; 11'.1. MMdlall" }.lm.tmtI'lDl, ~Je~ ,n6" 191' M~~~, ~olI. e. y. f!i!~ P,IH,Z;,nd~~r~ S~ (fIll: ;!;~, 133~ 12005~. 15. i.. Z'Ioe1lkO'!'a fl,a1•• NrJf,Gerilt'l; 37. 1174 (200:5}" 16. ]. ~~!lk. I<. ll4i,ld'i., m. IN,i:ibe'f, B.. i!.I<!!'o, Oe.vef"prmml

n.

consist~ng of IlES

C"(;!~liiil, hom~y,gol<l!s,

for tile ma~

tcms
ES

jt1f j\ln-IC l1<lpkH)'JlC[ii COll,~d .~(;,rI·C: ~ .,1 ~llH'C'4: of" H'Ul!'ipl.m!,ul~l,r;; '~~"""'WI"'Mttn \,"1,)]] n,'[l~'~C\.'1Illi;:B~~ thl~~:~

,'~]:y.

[10\';'(;\'1;.1',

blinn;

~ll,IfI:OW ,I nO:

~1II;rhlq):;, Ulhl;if

~h~ !!l'CnQll~!;i pn:)vid~ ;(!l ~ninl: Whl,llh!"'Tl!i~l'I;,,, <LLt: tlK~ 1'l~llll

ccn Ili!]C'.'1lhr4lugh SNr

oT homo;r.yga:sily

<ludllctCNI0"g<lsit)'

~n

g\',\!l~lyping ,lc!n~:5, dI;I1HVli,,'~ m!"'~ns~nd.cl~r·

(l,r

1:!<lrdl.t;lUl-

~L"l'i~IC.~ Iiubj(~~l 1'0 .rcjootioll "n"C

by NK

!t"t~·n;:;..

whid~ can d~stiUlgUII::ih 1:i8l>LIIL~ thm. CXI1L'Ci:i:;;' the ~.illl complerncnl 0 r M ~K: l'l1IolC'ClIlcs .~romn1110'.'>c c.;"pre..~jng QnlyOl r~~' (JO'ltI,~~OCC(" mhror hj~ 'Cli. tocol'I'npa~~tli lily antigetl~ SK:atlero:i OICros.~tl1e gc!Iomc ~.nigh~. CQlilP!] iC'21i!:: i:li:mlGliP~.tmi~u.i(mIh... [llfl l\Illi ~JllJgell.cic 0:::111bmlk. F'u II Nt 11,[, m"lchil'lg, (ar indeed. ,~iun !:lcnetK: identity I, ~s mos~ r.1vornb~!c for. predk!hag dlC "'tll"v:i.\,..~1al'ld! rlllltcliO.1:I or s.olk~~ orp~n allklgrnlls ~ni, More di9!:rilllin:JJ!ins tmnspl<l~wl iDn eXpC!I'imcnls <Ire l1~edecl~od~$thlgJ;! ~sh ~hcrc~~!li'10 mCfi!~ or!i:Stl\I~'8 ~Ic<!i 'C'~from hOL~lO\ \ 4}':g0!.15 MI~('·~I1>L'tdl'\id rES 'cclL~. IluUl' MI~C· m1'i~u~chl;d pES c-eJiIa,.or ~Mlgt'm1 it:' [i~~I1t~~ d..;rH",~-d
lC!i:i lTIil1C~("ili1l' 'i:JlJllb.K·r.

!:lt~IK",i:;.luuel~au·ir.:n:toOrcr.. or n;)~L1r.11 f('"1I1.m;Qjlion. T.hc ~li~.gml"'lic 1),lI[lCU:nOr;) pt M:~[ES oell Hm;ois )

I)crkx:ntro.rll!:'f.i:c

It1Ju;WtYI;O,siry

flm:! inc;J:'e;;"W>illg

hct"'ro.Z}'g~il)' as r!e~ysical alld gCnlCli~ Jis~oo.lC(,' lrom tile IXIllJ1O:rl);;L');' r!l'lCllC'a.~. 11Jc d~gllO"lic I);(I~ ... lan of a p{Mll'E5 oc-li. lill!: .~ ilCrlOCrUKl'll'L"n:C hC'im1il)1S"Psil~' am~rl :m ~ncli'll..'asing [rt"ll'ICH,-:i' of :S~p'j;lD:rtin!il1 Crnllfn~ Mate'lfial. " ••• wM.I!~l!flfema9"d @iigilEOI'ItMli'fllliltll3:3-5<42mCl ~lOfllo.zy!!usi,ty,1Lt l.1l~i'S dist:!~to lit:: Ci::nlfOJn~~ n I&ltrfl!l$ ,00000d, ~lII!llil,ilIii:!; A cCl~ lin.e d:;:ri'\[cd Ion ill~ mlhuyo plll"ldnced by :FiIIs- $1 ~~ !in m.llclciLr ~rn:rt5rCf fmun:Ji so-malic cdlwill. rO:I" the Jablll.~~ most 'I~.lfl.'bCOI oolnp!.lel.e gCI1~l[t: nllch or Ihe ii~e.FM~~
m,IICIc<'lf dOllOI;

~O, H. K Mi!ll:ld~ .• v, Fujiw.lfij\ T. ,;t. 5mllae@llil;. ID. Tra'o\"lr. S, H. OrtiD,iS'{wd" ~rl. 500 {OOtU}. ~:L L HI!o~n~l, :$" ~~I:!D"',G. IPli~~~, l. $!I!miltl:, (. fI'f!X. Nr.Ui!:Jit1'ld, .!i:L'I. ,U,S.A 100..].3:344· W)OO), Z2. ].. \!l~ d'e'Wolle!~ Qt, No. fnrJ., .1" 1\1'N'. lZo. 1377 U;9a9l'. H. O. H!Ii'!lp'he~ fl' ~L. S{~'(t ~~~, \\15~;lOO~2<1. milt;; iru~ "''a'> ~Prxfled ~ £Ir.iln~ fl"Olll' INtH ,ilOO lIre NUl O!J,~_»!;'~ F'Ir!m~r J,w<!rd gflll!: N;.J R~;Jcl~p lor ~~t ~rmtL (i(}.IiJ., i~ re~iQl.:il!nl01 1Iii~ I!l!iff~:tr;IU~lboon.e ~ F<tIIiI:II ai1ir.a~'S<i.@nIi~1 rd ~ iI 'm:.mIliitlcm~ Rf'!@~rdJ.. ttl(. 'l'!'aS5l.FjlO~edI by I he- Cad!e1s ArHmi~ ~I!!btii;m alHl ... 01 S~~l SQrntJ!;ir ,!!If ~tm~1lfliJ;~nia ~n~ l.)m~ So[j~P,l "1iI~~ilfl!y :!iUJll'Qlled Illy N!HIritlr!l~g gfilfit P<!'h«i~!l9i!',O'f r!.!~born ,!n~ ~'~rnel1l;!' !P.:i~iili {B.<l: H[lIO]\~66~. CJL. W'M. suP.poi1E\'1Il)1· ~, Ml'd.red ScIh~ ~f:tij:lllil fllilf ~~~~Li~!1 M~,

i'. mti@ ]~d;!lol1'IL:lOO1jjIQi)'. _.iilfom:>~~,iM.af9l' ~:2006l. 113. ].. L 6 oJ,,, Rfi.:!l!6., 4316 (:100;6). 19, W, r. !)ietrkh ~ ,~L.W~~ ~i!IlO!. 1!,0\1~' U 'lI'lI~J.
Milirilll! (il1'lNlIIfle

11:ll., 763; (19'11

~'llif,COIl'llv ~ OC01.Ul!lC!lC'eS of mi~Oltc ~mJ~:lbirllll~o~! WLl'!dcl ,Ii!~: Ihe ~PQct,,-d raHL~.nrSNP id:trll~~}( FlIrliIiOl1IJon'e. rhLtt sl10uld
~ lio

A!!9~' ZOO6; ~ 00i:~ >6 1!;h;!~~b~r~O~6 Pu~II~H1M llI'i'I_',f! 1M IJt!(jmth~ moo;

di.'!.l~TmmM!; 1t'li~biey

Jill' Ih~ ~'t1Ii1i1biINlll)[1I1

10, l1:26i''i{iBIiI(~, ~M2 IH Imdllil~ 'ihi~irnll!lim;1~'OOIWheti dtil'lg ihi~ pa;pe~.

486

\l\h; <!ntill;;llmt~ th.]~I~~Ceombinatinn

wiCll,IM off1;:!"sill!hs.~.11ti;L1 d}'lll,lInil;:

Qr

I~!,';

l\)j,'('I

K~i'lOI~~-"

b;l!,'1i:;id,

IR'fJ'VBlfSible Swilchiilng of HY'd r'o'ge I-A"c'tUl!8ted Na Inost:ru ct:u res into C,omlpllax IMicr'Dlpl,atlBlrns
R!il'~polII~i\l'! b@i1,iJ!vior" which ]S intrilll~i( to, lII~tli!r,,,l ~ysUjm~, ls b@[,C!milF!Q i! ~~y l"e<j'l.!in!'nl€'llit 'f,~r ~d:~a !'!ooiII aJrl'irijdld li!1~t~r iaJlsar1t1 d'e'!fion pJ\e~'nli!ll~ SubS:t(1!'!ti~l KliE!:[!tlfi~ .:IIli'~ @ng]fif.l~d Iligl dlal!ell g@, We dle's~gl!'!f.ld YIiI~mic aC:~lJatil)!11sysL.f.l'MiS by l!'!teg f3Jt! ~g hig ~~~p(l(,Nath'J,~m(~'!i1 d ~.alnlil:)C(lilumn5-, e'ith'~r hee'Mali1dil'l!!J or Slil bsratl'!-a'l!taci:l>ed. \'II!t,h ,pI hydrogell.a)IE{J'. 'The 1'1(1 1ilI(Ii(iIli[lI!1i1Ii!S were put in I'I'IQliQmI bytl1re ",rn~~d~" ~d' ~he hlydrog,el, ~\lhkh SM.reUS or' 'COIi!,ml~as, depemiin!!) ,ol1~he MHllmldflly ~e'lleL liiris acluatliDJI resulted in .. fa,s;! ~~Iml]bre reoriefllM~ijtlln of the n:aliilocolmnlls hDm mted to, pElJrpeliHikuLarl'O the ,sliIrf.l,u::ll. IBy hlrther ,mfltmltin~ the £tf,e1;S fielld il'll!i1e lilydm£le1,lh~ foniftall(llil ofa variE!t)/(!If e'!artlorat~ reve~si'bl~ a(W!1il:l!~d mi(ropa~t.e,rns l!la5; d~IiI1i(li1~tral~d.Tlrle iiTlet:ltaJllli(:'l ,~t' th~a(~lJatiiJl"I ~m(:e'!OsilIave Ibee,rla5:~~5~d. i[)!ftnamlic 'mntml oilier l~e Irlm~l'Ilent and '(lli'ienttatiol"l of smfa(~ nanjolf'&'1Ihlue'~ at tlile m~tml'l .[Ifldl.~ubtmkmlll KCiII!esma,V haVE!!e'Jicitilil'g <ilpptica tie I'Isil1l ilctua.tors, lmiw:dhJIidlks, ,or re'''>Jl(lIll:5~lO!\e mliilt.erialr...
~I

on ,g,.l(m~~lril;:lil\:,'1lr.ml1J:l~1n!.c;i11t.oi'thc h}'tlru~cll... ~rUm1Lilllmw!>im('ltH:"-~ To ,u;;hievc 'li'!~ ocsig!l$.!1.hc h}'dmgdw,L'i; ~orm'l,,>dinl~h!; CPr!nm,.'!H(."iIll bet \,\'C(;r!J AIR:S ;1I1d! ,I SOtti!iidllI)'
Sl!llhs.t~ c ~ .... fig...

I q l2J~" .rol )r:gI~t::id}'1

ml'i~Umer)'I!llc (PGMi\) P'u!t1a.~]y .1I'I'it:.idi ll:I!ry~it.aeld, which fOntt s 3n1 3it-ehon1n1g

ii~~ \\'i.lh iul!~rfiLCc

Ihal. i~ii,e:~ ~il i1!..'"1lCIJ,\C cpoxY!;Iroulbi I(Uk Wa!!: chctnis:orboo on one oFd1,e sllru~rn~C!!>.A ~;\\YC:I: of pol:ya.ci)'lmnidc £cil {.Pl\l''\.G) £m'l:kd to llie sub~.r:m:tc via PGMA w,:!.") ror~nC'd by 111101.0- or. Ihcnuo--il1 ~li.. lCcI in :<.ilIi.! t<ld~c<!l cOlX'i~>~ncrizautoll of ,Jl!:'!)I'lrunidc ,U1d N.N' "mi;,'lh>'.I'c!W"biii.aCl)'~<Im~de a~ 1Icm.....,,·lil1 kil'i,fJ"tg!;1ll (fig. S I). The Am;S ~l'liJd ill. our t'X~r.~I:Ilf,;iJ'In.~co!,1!;i~llCd~f ~1l:U1iC !lm.ys !pI' \iel1icaJI,y ork~ll[ccJ. Jimiroml ~Irulocc~llll'l!n:'l willi d~"mCII.1'fl5. d = IOf' to 300 nm, hei gl~~:;: = 5 w II Ii! J,lIn, fI\\I](;~ ~u~in!' MJ = ~ 10 SO. ,IILd ,l!!)'~il!)5 d~cili(;;i; P = 2 10 " ~~m ffi~,. ~B-1Wi! de v cIQil~d 1,\'\10 il}rb:rid :l«,hu[I.'Ctur,c"S
USHi'lg

dn,m,gc proper! kG and rLlnc~ im~ ill I"CSIXHl::<C to extemal ,~linUl~u are tile focus orresc ..nch in fields ranging ~mm1'll~\dir:i~C'.um bio~!1lgy!od~\;'mi~~!ry. physvcs, mil~i:'ril)1,..eieucc, .. .t!ld~ngiu~\lJrijll';::. A wi[l'c r'lJI!,l:,", o~ Ollr!il'iej,ul resp:!nf>:~vc1]'lJ~tL'erbls. nm",Uy ili~vu~\'.in.1:l ll'oIYI~tlrs.. ilillJl. been !'C'P{)1'!Cd(J 9~.1 ~}'dl!ogcls :rrrc 1l!TI!ra~
iHC'~I~'~\imn\pk"!i o:r ~~~I'CIi! nl,"iC'!~,lh,,- Tilei If t'II'~"'I~C' nl!.:''1:\\'m::r.:::o: sw\,~ lin \\'rn~r. !I~ I()win~g (Hie 10.](::hi~\',C' ~h~ dlt~~iIcd (k;"'grlx...':-i (I f rm~}','l1i:t l1y,r:1rnli:on m'l'C! W:ilUlHll.C c.~mrl~e _ A 'l"ilriely of gd .. ehm are re.. s sponsu-vc to hlHnidi~y, I'll I, n:mp~r.ltul"e, electric ndd,~~ght, and ion slucnglh have been ~:ks;igi'bOO 7~ oml pauerned by means, of ~ithogrnphy 1(,1( 9), These il:'CSpol'L~hC goc:~s ave been S~lOWII 00 h iltll\'C po.ttn~kll aOlplk.alic:ms, hl micronliliclitc~lissue cl~g:i~'l\,,'Cll and ,13 $llmpC'"'IHC'U1.'OfY nmtcriab. ng" ,lxlHkial musdes, ,,'''~ vcs.and "!;~l!l!;UorsU 9~ SYll!thCiLk: rOlJll.!_~ an!cl &'lini\".uiOll~ ~twt~giC'1' lead i11g! m :~lcw-gcm::r.!I,th:m. dymlmic.ll1y tml1lib~,e: Ilm~cl'i>'!J", <l1lC OilC'll il1iSpg~d by il:lQlogTc<l1 ~yNi(;,lll;;'" ~h.rl ":how OJ! wWc r.mile (If ao,lpti\'~ !'I;;'Spg!l;;'i;\.'$, RA.-CI;;'Ul~Y, !lllIn~bcrol·~~i~; ...h,I\'1; d''lll''Il5'tmt~ .. tl~I,~~ V~ni~I[lt~ph)"'ic{)!;;b~nlli.«I! ut~~~rI 0 f~i{llogl ~ iC11~'mn,.;~iuh;lillm al\; ~"IJlL:.r~Llly~lUlj lli'r Mmiwil I v

'

dript!, \ c materials mu! dcviccs

jhat

ScV<!ral cflhns ~nfi\lcbccn made to ftr:!:mcktilly

pmduce naIIDSll1rCUt~C'd SlLlf~'l{CSln pa:1'.YlHCriC m'lIcr.iai,s IhJI mimicli:lioioglta.'i suueuaes and rtmcli('ln~ U.f. Ut The: imrin~ic nl,,?;,ibli~il)l or
ludY!1!IICfl' JroqllcnlL'I" k~td~ lD unr.i~iftld desjgn i:!:! Jlun(lriOl~!i, ;lilt pom.Ly cm~lmlk;cl in ,gt~n~~d.l and ,1rJ;l OfOC'll
O~I!C(l!m.U.'S •. 1'!>I;"(;,I!l!:-l~ 1~1~fi.1<a1Ufi!;;~

sor~

Slk<;'(C()1

iMclo irreversible

eonO!flsc

(J.n

We ro·

or auaehed ,[ImAIRS-2) f1m1ocolunnn.~. The key cl~n'lcn~.or jhe HA_]RS··~ dc:sigl1l i,~ use or hydrogl!~. tbe embedded siloccm Ililll.ocolmnm;; ~Il!in detach from Ihe original AiRS s~li,oon wafer (F.Lg. t D~ 11:'0 achieve Ihe H/\JRSr~ s,u1Jlcwre {ng,. S2klhc anOhOfil,g la.}'X.''f orohcm~oJhod IPGMA and <!c!')'.i'ic acid WM ;[l~pli.cd to the sccend surface whereas !llte 1\1RS :~cmaimxi umlllodi fled. The prcpO:lynliI.'l':iza'llt:S(lhuiol] "'1:))1 inuodneed bet ween /\ IRS
fn:ce.sn:lifl(hg ([lAIRS. I),

either

~~o~!"''(1 ,dWIHIlJ!i,,"c tll1l ;;'Irp~n,lch ~o]h~ Ihhric,llion of 1I.'1I1C)\;lrll("Ul,l~(~ urlil~1'i" ill whith ~W!III,lr a [Jrt~y~, f wl;.~II-dIt~ri I'ImlOMJ:tI.cI.Illrr." with li~l· o trn~:d HHte ~iJ;loC;,l; .... (1) 1.0 300 Hill and a5Irx"l:~ nmios of 1 [,~a,d~~ng were fbltnc(~ 100 ~nsili(o!J by mesas of

rul!(,l~h~ l'llodi:fi:cd (."On'Jiu~n~ ~lr(~cc.A PA.'T ~'iO]}'l1~ S Il!~1;' 1\\'0 slud[l(;'C':ii W"-,,rJ;l ~el!;II"Ol~~,d by aiPplyung shear ~llr'~ U)lhl; ··~Jld\'liL(:h" t]m[ rc... SWlliod in '11'1(' OOll.lpllcu:: bl'(~"Lkin~,g of I he COIUHlilll~
l;.Ii,7_."L'UiVI1,

the Hosch proccss l161. these suueturcs are steble flInd their gcol'Unct1')' is; high~y contm~lcd. This rigidil)" h01J!'C'vcr. makes Ihell'll lil'oSui!ablc f:or lLscin arnplivcmlltlLCriah'" and devices. A mat\: p~omr5ingapllL'OOlch ISliO creaie nmu::r~~::1, ba!:K'd, on Imf(lroo'~l cOlnbimH kl!l~hm Cllph<!U m em the pJ'Qpel1iCl! Ofb'llh, eil;'ml~~t'i. SI1liM~.'ill!'ICr· hyd rophobi,c Sllr~acc,s liUi.\'C l1(l'C1IlIbb,k,uc';) by ul)ing pollyclcctroly1iC' l!lll!l~lilaycn;. ovcrcomed w i~:nsil iea mlllgl'~:I'1 icl~ U i},~n!iC'!\."\'O~i~!g -\;I!111" C

,I

i.dc~ oj,' ChJ::Dl1!ic:;.l:~I}', rllIil.gali,;lk' .. lb'. ,1I~d li~~l~'~"TI!I)11"h.'CphO'ton'lk: cl)'::;,~,~I:>. whic:h w,~n; ~n'ld.,;

~l. Ihc base, The dctruc]ll.'tI OOll1ll1Inil.lCIIlmill~(1 emIm(Mcd inlhl! hy.drogd Il~in !ironed via PGMA 10 the cmltlning ~jr~ficc. 11l.is ~lrocooure was ~.ollowcd by rinsing and dryillg anile hydrogel !OI'C'IHQ'VC ullreacled 11iI0>~lmlle and [he solIrS ,r~ln. C:flu$h~g the comracrion of the min and .he. ~s",,"}ciim,!"-d slomuin~g ofll~c. ~llmuachcd, mmocol LlIllil:i!, ,(Fig. 2. A allld :13)"'[Ile ~urfil.cc [lC\,ca:lod domail15 Wilh nOIll-lllllilorm tilt dil:cc~iom,. '011cel\';lbly lll.d'Uccd by i'llhomogcm:h,Y or ~hc h}fdrogd mm 'll'Kl {or 1!(Ii!;,d&~lu~~t';J,c>c c'&,:r!.-"(:I.,'l, The
lill "1l~1A; a of I h~ C{llU!HllS nh.~(,'I"vcdill ;;(,:\'1,;1'"•.1 1.,'Xpr:,~r.~Ill~nt~, \V"ol:>(lOa(j 75 , ;Ind lh~ 1.;"',N:H;i'!.iil, lenl.~I,h I \V'<JS ij, 1~}:2.S~~Ilfl. EXp!}N~m.; 'co hmn 1di ey kx!IQ :;wcllillg ofthc hydro;g,.:l and re~m"'1Lliolli f o lhe drying S:!I"C:\SCi:l, As iL re."1iI ~I. the Iilted co II·

.tL'is~ ~r·'011'l Il:l,,; ~.:lrl'liI.']OC of highl:y dc .... -c!iJpc"!i1 lilIJ."·' laC(; nlallo.;.roughl1C:SiS ,and CJoi!qul b;,jloC' Ili.IIilp· lind IDnic:oorc.~t!Ji~S {IO 1.1). NmilO-mn~ nliC"iiO::1rrHC~ 11l~dID~ ,a.cv,elop on thc ;9I.Irf<l.ces, f ~ko o lbel. IOI~~:5 ~e<li"C's,m:nd dC<lci!:l <lnd btllucrill(y willg~; for , ",."i.;j]1I13~le. IIU~WiCl..e~hl,,"S'C O~Jll i5m11S\\,'itil eXQl;;p-Iion,,1 a.dhc::ivc. sclf~c.l.call ing., wi]lto;;~'-li'!i::pdIHng. and Pitol OIl i:cpropCr! k-i;,
l~ll l..ii'b6l.!l~lll'ill!slAk~I~l.I_rtlI(jl!lll. Murr~~ !Hill. N~ iD7'9'74, USA, ~Mail: fI.<lirH:k ImtitTUI.e ,or (Ollmds alld Ime'd~,{e!. 0·1442:11 Pm'Sd"IifI. GIilImi:!I'l,Y.

by d:~fq'lningrig;d l:o<lloidnllJUnt it.:I!~ID1tO h,ydro.. gc:l's. h~lV'!.: 'ltX"i.:ri n::poiftod (lfJ, J9~lloo~ike tlllild· in!;! b:J!QC~:;c0l15i:r;tilllg of gold illlcl pi)ilym~r

domains ha''of'C ~il

~hown to scllf·as£cmblc hnlO

!JllnflS movw10

Il'lcii" original o,riClltaliOll per-

a vmict,y Oof" :ruPOCj:stn.KIHI"C,~ OW,!, \'!Ill; l~mllO;S:Cherew u.~lhe c'ol'l'lb:inmi()'11 son and lmrx!c~mcmsw ell!lbic- :r;c\'Cirsible actual.ion of rilgid sUil:ll~,cc nanQ~;-~nd~niCl'>o· sh'!1actl.lI:cslha~arc' SIC .in Illotion by Ih~ ro]ymcr I.
laycB',

~')r

"To ~!bom ~:o~~e~ppden~e!IlIG~dlbe add~ess:ed. ,E'Plait 1

jailn!nli!!!r'9i@EiIGitel.tll({!nl-com

hycliro~'C1(Fl,g, M,) willt high 3SII!1!CN<llin dgid ~1I.'I!9CI.ll~'V\IRS'J (.Fi,g, lEI) uilito h)'lkogcl·AtRS ilSi~nfib],iCl< '~I.IA.IRSHfig_ I, C~n E). Th~ATRS rmviw: ri.gu~~ily, nICII!llIr.. l!Iluil. pJ.'i."(:t5iom., \vl1elt:· M
inlcgrnlcilll

W~

Ihcoo~.lilml1S returned to I.heir i.i!!i..'li. ,sUl't(;, By ru:ljui'>i:iin..g lhe humlidit)! levc~1lilsinglhc: hu~n 1d~iy c1 mnl~}c.r (ami Ihus. regulil.tblg d'Le d(."g;~c or hydral.ioll, and~hc ~,'C'Il'Ltcd volume' change of dlC hydrogd), we c<l!nronlroi lile:1 ilt angle G r Ihe Jl~III(lOO~lUnl\;;'l.

wend;i,c\ll~ffrIO~hc :surface (F-iS.2C). tJp::m drying.

all arm)'

of isok!il~d.

:it; t~k':

hydmgd

IMDVioc-;:,.

Ii:'S:POIl~U'l,lt:' bchm/:ior;

OAClll<lllon ~imr:, '~hC' l~lDO~ with O!I hi~~·!S[~'of;."d ~id(;"O C~Hi,1~n;m I(li}. Whll.'li'l w,lIcr dml~kl i::.pl,rttd dri1U dry Sarii!Jlc. I:i:lt:' s.'!\'~,.rChiHig W dt~ :f'h::tJ1t:'lliI iCLi,~m·
W;I!'i !\;"Co.r,ol.li;l

Too ~n~a~ull: Ihc

REPORTS
,\~ilh~h~ (f:i.g, 4A);. A~'arlCI)' of'more (Fig. 3M. The aCIm!igio.ll~ycle is :;.i!:llil;uwto ~mt 'IXMHC111Cc! (11i;l~wi~d IQf ]I A ~RS~I: .1;,""i"ffi'1Ifl: 10 !nnlll~iggl}' t;;omp~c/!ipmacm~ can t~ ~hilJ\ cd boy ~Iii'il:!,g ;In ~~h~trn!~~ '~I~l'~W!l:d !I~g .. tOpo!;lgrr.aC,IWI~~ thJ: hjrdmMgl to S\""\!)~!; ,u~~~ ~I~::o:,.~h~!>, I,IIKlI,;"[~:yj.llj;l ~'~(l!IIr¥lnll!!l: '~!l~I!,llloorlJurmll'1 In 11ii\.~irp;::rpl,;"'I]td;I~1~!lr phy. Ollt; :>w,':h ~.... :m~.Ilk is .gi\,"~il ill fi!& 4B, "':IItl~)~t':il:;Lke~ ~4 s. Ml:I'I.'ic S2 dt;piCI;s the reo:ti.... niuilml" To~'i~lllll~~fI."~ LIte ruuri("II:n;tiiall rnx:e~:i, WhL"'iJ:1t.:'i honGycO:l:nblilk~p!JiJ"_'m an ~J:IC >t::ollfinirlg \'C'l.;SC In\n~'" ~.~o\,~1 dbwn h~:,!! n~C'tor 'of 2, {ll]d !he d(lpc-ilW!.';'~~O: or!JlliC' Ii hllnglc 011 'tI'!C'dc!lf~ :.W!dlloo oonU'ro~~lhe' rO!i!1~~inn of a ~1l~P~c;,: The !mU!;.1Froorill;n!,l'uiOlra if;;,m 'D.,; .u;:C'OI ~~I't~d,i r (I l'It}'~t~uiou,Lndtor hum~di~y.\1-1; pb~~d .. tll.npicl :.lrm;'1ur<:irl] 'INiIl!:(:h"Irm}'~ or r.d'l~ o~jl;lnh;;d Iy co!l,Illi!ml.~Orm ·~'ilil,;nlU1o~~'\.mf·.iml l Ihe rI~'r:,k:d. by US.il'ig ,Ii~n(",,'!i' ,in Ih~~ :'j'$,h:;ufi'i !IIi.dlor (I r w;J~l,,"r til~lit(:" HA~RS;·2 ~W1diu,)~ mid 'took iUl ~Iy iltc [(~!L',i i~ 't~u.~ l k:~'ii,pCrnIUI'e, TI11? 3.ct~i1U~ uoti !!Oi:'Mndry of ~~ ~eri1p:llfllili!!:sltru'LCC. Alll 3ITdY of oplne'lll: Itirtcoo~TI~ph ~i.c ,d~)lillg, cJ~,C' ffi,~,38~. The \!,I!ll.~r ~OltIC1~1 31td the !>wt'!U il'ig of the Ity. c)'dc wa,.. repeated on '[h~ MJillil~ ~!ill~~h~' mere . ~nic:i'rnra(k"'" il1. \\!h:~eh cvcl)'!¥OOP of four atmathlddl nmioco[lilimns ,of 1i11c I ]AIRS-2 dCi>~~i1b: 11191'1.20 UitnC::-. j nfUfI,eJ imel:v !liner th.\: ~reW!<ita] io.~~ dro~1 gritd ~mn,:.' de@rel'l~ aemss Ihis. region. mitd mol"C' dum 25 tiiil1!!S 6 mond~~ la~er,. No held lO&e~hC'r b,~ iii. h)'dro~l.ils dl.!pktoo in Fi~. iltdi.lcill~ ~ I'ro~$i\'c i~lCrc1li.~ inthe t.ih allgi.::s <te. Sud! 'Clli~npJC':;: l~mtI:!1'n~d~1l('IVCrn(;,ilIS would deierieraiien or d~an£e h~ the: uC,;<;:jX)llse lilfl~C' n rlhc nmlocolumi1.'i:, was ebscrved. c1C'mO!~lS"~m~~ OJ ,remm:knb~,C' 11,~ We rurtll'Cil' CLllli'tilll!Zoo on ~lese new aClll<l!- li'iC im~X'i$,siblc h~ II~c ~~p0l100 <lniltc:i<lii-nmsdc ~yslll:!m.s., ill \?,illkh Ure po ~)i CI:3 aJ;e aeltmlOO by m ~~tlbHhy .mcll:of.!lI!SillC:;:S of'the dC:i:i,sncd nc:~ua tio:n .~)'i*~rn~ cxp<ll1d.~n~Iheir C,lpabmlic'S to bJi Iflnn rc\'c.rsib~y ,lcUlm~d micm,I~~~liCrn.~ wilh an ~ioc!u'ic flf m"~!lelic field, E'IIC'n !l1!tWC imrion ~)'~~cm. \-Vhcn~h~iJJ~~d!o{lri !I!!: lo1y~!' of Ch1'Jn]J~rllil.;'d ~XW1.;)1nt Ihr;,:S!.,;[]i()M.Cm8 o1!i'i',; ro\'c~};.iblc. ill con~:r<l81 Ci:1nJall~x Ii,fI.;'O!TI.ttrlt"& B~ilWIj,i~ ah~!ill diro~!ion 'H101l!ll,) C~I&,.~ r ~lnl:eH.1~~1 (I [~'UIT'n~m(lm ;;l.l1d <111>.. PGMA ;mdl <Lt;l:)'lie ;JC'id i~ 1lppl,i~dtt'l I.1;Ii~AIIR.S 0" nh~~ n~:II(liC(1ihll'l1ri& lh~ HIIJRS-t dL~igll d~· in ,,,~bstrnl!(,': ,met Ihe c{lnHllil~g surfaac ,~ Inmnocl~ !]cnd!'l (In I ilctoclll dcfeelS on Ihe cm~t~ning .slll" ~'C'111bl'y reI'lot:~(!d in (llJ, :n~Slfl!C:~~lres~hl)\~'11in Fig" 4. 13 and (', QIJC~~ do.w d~9xnt!i!lg QIl ,md il~!Jd, 'the hy.amgclJ ~Um ~~ grOlfkd to ~:h~ ~~,~!1O\\,C ml!i~~Pl'!cd !11Il!lthis {I~-ij1,;'lI"'liOI1 {:o'l.dd be ,"IH1UC'l~lD1)d Mufat(l' I.fig. S2~.'!'! IhbIIAIRS,·2 cOllftmlk\d by ddill('1rmcly ~Jl~!'Oduci!!!!l'1 WJllo~ 1,l!J';lphie pauern nlll.o t~! MUIi"~ICiL;"In:(k\l.'il, ;l h~~ll. i.~ . df;"ign (.f'~!'l'.lEI. 11ll."lijydr(~I,-cmllllddl\dV!1mo. Iy i!mi foml I i~l.dill\.""CliOI'l of n.hc nanocQlmnrk.~ \Vlll> ca!Llf'I'UliS rul1rmin :J[l8t"h ... w' I.hc origillal $aJIb~;mh: ~, andbend upon llI~cdrying of (he p;Olym.cr film :lchicvc,j b}, lh."'i~lgU !Confi 11iiJ1!lt>Llrfrn)c IJ~::U~was
oriemmiQll I:;; c;.;!tI\:1l1cllyr1!5~ (->flO ~ns), Mo';,' ie S~, d~~!!;~ Ihe precess slowed dQ\\'H Iby i1bo~l! ,~ f'~U;:~flrl[lr "1. lh~ :~V~f,\i~ m~I]_~f(l!"rl1,,~iml'(l !h~ ~illcd .$,t!li~ q1~lV~~ by III m~is.h;:d ~nyin!!l or lh~

or

or

rouow

race.

'Oon1raC'ted 5tate

I~ me d
m I~ l~

~i

SWoDoo
state Fiig. 1. (M' Scinemlat1:( ~n~serntatTIQinof the :strudLn~.and (;oIil1t'~sitlion of the PAAG glrafi:ed t.o the Siil.5i0:2 subS/mit.!! vial Ull! I~GMA all'lchorirg,~ liJ,yer. (8) St;~nnili!gl ell!droll1 Irnil:rogr,;lIphr tS.EM) 'elf a s.am.ll~e AlilSstf!lu::bml' ((lmpoSM 6.'1 al1iiura'y ,(ill :!iiLkom 1i13lrmmllLlliFlfiS, Alili 'iClwt on, 'lhe right re11{E!.a1s a handing! ~!i;lfl!tem bCQlllopi f'I~) llil~t '~~,cha ra,r;l;oe,ri:;.tJ1c , of~ne 1!l,05thrabrtCQJ'ti~1ii p.~o(e;;.s, Th.1!' bdiii'ld ~'i!idtiiii$ ~200 nml, (() IfIV,~rog@l S:lJiFlt~@S"isi!1'~!i! (,Qn~~I'!@M~rit U11~AlI.RS "nill a s'e((Ill1d.ary s;lIiil)suat~ (.ar g!ass slid'@' Q~a of si!J1mrn warer, e<ither na~ or topog~aphk.arly pl(lH:em,ed~~iParated by a space. or the deiired lhkknreMi (hat ~'!JlII'laJt,es th~ 'snv.l' ~'~: '~h~po~fi'l1lerfUm. ,aD!) H~brid BiAIR5-1,~'!Bigin. 1I'h:~ rililno(oh,dITII1S;, ~niIbedd:ed into the hrydrog,el, I.a:y>ef ~l'aIf~.ed ~olhe PGMA-modifie~ wnfilillng :surface, break fwm the ,onglin.all,w:a~er !lipan tih~5.ep'a ri!ltion g~ thll C(ln~nlll n!l,,;ub5tratl'l:!! and Ilec:omf' f!ll U.y lrr,m5<flllrr,ed ornt'!J tile ~emnrda ny s~rr~ce, ThE!' sYrr(!Qe ~@pogr.!:!!phy Cl1illrlg~ hom @JiI .cJJnoil¥of highly tmed to 'I,I'~rti(@llv ~ni~nlxldl lII.lnOC:O~LllIIlns. (E~ ~Ili',briidHAUIS·.2 de'~91il1. 'The hydrogeliihn i~ at1i:ached diredly to Ute' tIll'RS S.1liCIllIl \YOIIr~r. IrlJancmOWllIlfl:'l rernllin O'I'Marneci m ~h.e :!;l:Irfacetilull1eond lJipo:n ,drying.,

I So !ii' . 'S

Ie a

mm

fliig!.2:. Mkm~t1op¥ S:~lIdy of l~e H!AIRS-'1l de~.iglii. (A) SEM imag~ D,i a di'}' ~FiI!pl'e \lliewei!lr ii:il~~jJ;i.ifi~ dim ~ar to In~ sliuf.a,rre reveah lilted ~Qrlul'll1ti'l~ organl!ZiW in ,dl)~amrS w~h dmn~re'"t tllt~ dir~(:"' I:i(lns;, Iha top inset shl:lWS ;1! :boi!J:rI1·ilil:vi!!W ~@r· pel1ldiicular lQ~he flLlrra.Cll!. Iii !limber or 2:00'~mlll bClJild'iSflO (lJiI lh~ em~r~irl9 p!liIrtior! of '!he ~"!1'~ ,milYmns and the hmg,lh OJ the wlumnr prajeclicm, III ~ve~eIlfOCIl1IitDreao dre<toermine the' Length ~ Qlf thE! t

ne

,e<llpio:;oo D1!i!'ioGlllljllillll'lS r

,II", 2:{lOJi',

ImeafiL!r,edinr

IllI1TU,[I1'I;r:i the-lr titt angrl! (sin Ci =(111). Tine SEM '(ilf lh e ~O$$ senl~ii'Ii ~bol:llOm il1i~et) d~alfl\y :sI'!rli\lfSth~ IiLt.@~ iilarnO()(l ~!JIrmrr!!;Ip..adarlly @'m'b@d'd@diiI1!OO til@ liiI~drog l'~!.aye rr.$(0'1 re ilaJ5 ilninr~ls., .2 ~ll1Y;L '~B and 'C)oOptilGP~.mkrogrilpl1!S, ~~g~l1g tme s:.gme f\~gt{ll'l ,oftne HAIIIRS-l. :sys;l:enl In ~ ,dry (In .and 1'1 wel (C) 5tdte,. sllow llll,~nmriel'ltaUQI[I of the nalnO!l'i(il h.nn) ('FrtI:fI!! @ titt@d tQ ·!!'I,I'~rti(j~l ~rtI:sili(l1ili l!P'~l!1the el'!:patm;ion of~he hyd'r'(ilglel. Also see mQ",i~s51 and :5.2 a nm! fig. S.l.

4SS

REPORTS
CI]Q.lmm!S 11mC'JIU.i>l1 in i!~:chit~~m~i I;<!!I):stnJ!CliQll ~hc Immidi~y leve], as illustrared in Fi,g. 4. [l'mw coined !!h'C tenn E. Their movements '~nb·w i1~eR\'liJO~~"i\'~ ~}' U~IC!kmin~~~r-ft!n~!",~'II1Q hoqh;1\' iOfl'lr ~anl.i\(ln.l~S. ~.~ !:lUlt~, r IlCdi:(:c;:~ o 1<L~i;l "'~~~:o1~gai" (lIS) •• m>t;ll h~ CO~OOiIlt was l>'lt~ pm•. y '(mu~tUm.\ft,.. d
On~

micremeter SCillc (,Fig. 4. A <l11.Q B)' wh ieh might l~ ~~lmic."ll~ CAI)~oi!.W" such as in l~e context Oh~lg(:irnll!lli~~~c" 11ho~oni~" (J;'il or
llIIAIRS·.~ sy~~~I'I'~t> ~o~yo~~d IJI~ I;:{lacept
[~nl:';I,"~.I.y.

nlktofbrceps-!ikc

~Dro~U,m".'!'>kl\1I1-d

~'ChinO!l;k."jJms~.\\'hOl>t~I~.lf}VOlllt."IJl~ IX:';'-Wt1L11, '~hc Op!<'U and eloseel :sn;;lt~ rill Ilc!;n!lsin .tcc.Jirlg and dd:bn-<lC' ;I..,d 1L" ,<LPl'loB i:l1]lI~in,gnl00h~.!.1i~!l1,; Ilrt,;· \1~'I1 cing I:h!"~ !'ot;~tl\;:ni(l11~ gnYli\.\clh of 11iIirn~.~ ~lIld n'lil'llll. {2'nll~t:'""t'bir)lo,giC'1i:~snmccul\.';s~~ik,ni<ll.l,I'y . C...:.iSl. as jlt'l<ii.y:; (i4f ('lIl\' irul\t~1IL'<tIcallyi\.,,~;:,;prnk<i\!e !\.C. tuators, a<::do Cllifm'tiiCi:cia:i diyn,1lltltic PflI~I;~rrL".The P'OI~i11k'\l or thc..'~c lmklllC' snucmres ['Or various
is ~~owbei'l1~ ~"<.pklli'Cd (15). From <L mOCll1lihl~dl v L'e'I~1'!!Oll1,tdl~'C is a~n'll(_'ll' d iI:1Ct'e~lce OOlwo(;.'el!lhe I [AIRS-I I and '~I,I\ I RS~2 ,~)'S1lt:mi'i,ill l.he 'way ~h"l Ilh~ mbnnoo,hullm~ moe d~lh~'1f11I.1~I, tB!"" 'C"dt;C of!lllCl HAllt"'l·l :!''')'~CIln. ln 111L:'11:;0 11,1111111." (:iiJ.JTL~~ld (~...enui01lH)'~o~t~ Il" i,;ll'" 3~plica~ion.~

as :I. meehanieal mudd for [h~ (jL,~1I cyBJoz.:!kckuJn 'PI). S'~(,'n i~'lti~gri'j)y S:iuut:iu[t':;. consisu n;g of ~tirf rods eomrected by so•. ~ringl5, ~ UnOl.':fI!::;!"''iin~ a~~ 'C~~!lll},bt;:i!}£ i~\ ~~u~'O'Itocl ~n tilL" tm1'[\txl. of n'k,ltlial.'ikl!>. Ihr t~~urillhL:'f¢li'l ::;;11~]~ 11'iit:tij£ii:t)' j:ricl. oc~:tcd pol£ljlifl~ 3:' :nCk~illot.~(};~ i. The .II.AIRS-I :>ysteinii:. !i 1:c~liG''''1Ii.iim 'lC~~g· or rill' ,m. the- ruieroseale, WhcilC the lli.ififiOO l!.iitmS, llit;."dil\,~l,nll.e tensile romes from [hcg,el. (whi.c~~ 3. 1):11,0:[1 ould jll~~ Ilcllid ~\'I ;J l\.-duClj('m w l,'he gelhycr Ihicklw$S1 ~tHO ,I 'l;Helml :tC~!J.mlj'(}ll Fis,. II»). { Thoe mO\iCmCm s is cG~nro~lcd by 1l1C ...oll.!mc dmn.g~~of the gel a ndthe h::ng~hnfl~l('! nanoc{111l~n!1'''>1I' ~' 'lI~d 1."!! m[lr1l,.'t;,tn~ 1.!1~· w \,fOr~rnQo uf
posed

of

in C.hb~ :>t:11i:;c d1tl[~hc b~lHb~ 00 no, :.Iuy ~"!<!.!gill.,~~d C~U'lf)' compp.'fu",h'C' IG.w~, bU'1 i!l~'W<1d !hey ~nd tFi.g.. IE). Typ:ic"llIy. t~.;;: ~tinhl,;'s", of
'Ihirl bL;mtiiS iis ~t~l,Ich k'£S ntii lX"iJidUI'ig thum ill '11is.~lIImia'ig 't!b.m:t ,niO hLIGkJillirg {H:Qjj DO J".Jl .a OOlL'iCt'lueilcc., tL ~!litC .rm~~im'i I. Ihe deft)rtlbii,HOn~ ,"~'iCir!1V is ::tui\.'t! il~ '~hc bi:nt COh11n.~,
COliiliprttoo1oi.nM

or

or

~h(; gd~1]

Bhe \'l'CI.tlltt!

~l~~lt'lI'awoo under ,ron~pl'CS,\1ioll. whereas Ih,e Shfi'lki:ng i!}'d!llgd esens IJ.'I!!:-<ilC' for~ This ~~1'p(lor dc~i~n~ W01:'''trJ:.>,ooglli~d lorl~ <I~O Rwil:;o

tiNel)'. Ihe!] 'cos ~hm

.f

= II

f~ -

~h~ dry s.!"nl;.\, Ii(.... pt"C· v~l" ~nd sin a: - xlh. )10 ....
lilc

'~Jdl~>w~1. Witll
.in~his

dhlll .. "Jl"

sIm!:> l.YI)i:c..~I)' used

.~lud)'.uhL; 1.:]~~'r;)LI

:liC'Il)I~~tiQn! on I.he order of_1f = 'I co:-:jJ.I1IL A ..11I1Iarc ..is;

cemplcx
Illl.cmoti.\i:

IOpc;gro~:D~cl~! uf

I.he cou~~il'iing surface,

~fo.~· 'L::xmnl)lc,11c!I1.qlJji.'Ld:rmk:J~Mnlgc. meuts of lhcmT·.;lY of i1m~ocO:ltlmll~. the use of . Ilm1.OS11'll'Ctlm~ of [L diffCiI'Cf:U sh<l;l:lC (~m:h as b Iades), or a oomb~l11ilt!ion thcl'COf ·;J~low rol' a varie'ly o[)frew.:~iblc p<HtI:!fucd I]lJOVC1l11.Cn~!> ~Itthe

whkh t>i:lttI to go bacli.: to lit..::.ir Sl:mi,l:.hiL sh>!!I}!! when the 1!C'nlsi suesses fi'Oi1!~~he ~cJ,are released I!e Ypoll I"Cwct[iny. TI~C' am.omu. by which each C{l~!1!nmbends is a measure zyr the forte exened by !h~ ~C'I. This i~in C'{m!,,~~1 H)lia~ I ]AIR_S~ I ~)""'" '!~~n.wll~~',\:~h~ CilfCCdcvdopOd by tl!1{; gd h,ll1 nn dir'l."Cl innul, .. ~~ nil. ~h1J Mn,I(:,tur~u fu,;1' dl)'m in.g. U~~irlg sil1ilp'k~hc..m theory, we c,..;;!hml!tod ~hc rOfCC f'u whicl'! the bond ing o.r[h~ :l.i:II~cml n1mp.colwm1,~~1] Ih~ I.U\IRS-2 d~~igll~oL:~'U~S (fij!}'>- S4 ~Incl S5}.ni:-; rP'J<:(~ ,d!.;"pcnd .. \/\!'11' SCffiltgly 011 ,llc' :1."3JCC:'L [miio" f,or alfl iii: .!Jlm.h igll roluunu wil}) fn~9sPCcl.~mio ordt~a.c" 10:0,40. 21[}, or 1.0, the bending occurs at ~50 :nN. ,·-:500fiN. -8itN. OJ' WO ~IN. respocclh'c~y. Morcm,cl'. du:

stmim al. thebase

CilWUlot

exceed the fracture: Sllrnill oilly lmdii;!r,go~lmivcly

of '''i.~icOJl.The COl'L"C<lU encc is I.hal, co~mm~s, i!h w

a:?'ll~'~~ ll!:;'Pt.'(~li<l'~[1 'CIlI_ll

Ii"iig. 3. Mkllos{oOpll' "'~l,Idy the HIAI'IltS"2. ~1J1iI'C:· w.re'. (11.)0 UJ~~"'fTIil~gnlfi((Ith'ul SEM (I~'ro ~r!.d h~g~· lIIiIagnlifi((!tioJl SIEM hight) shew the hyd1r(Jg;~l ~'@r L thil t 'folFlll1dicl1ara c~~ri'5tk onion like ~I" - ((I1ili(all r~~~lJn~s tlfl~ bilUiJ'ffi ,of the !l'H~!Jcol,~mlfQs, ~IB)' (It r.lpth:al .rnwJgf<:lph of ~he drying ,edge of the IiIAIIR:S,;,2 stndu ~E!' taken IperpEl'lIIdicurt<lr to, the ~urJllil:ll. iI1he d.arUicO'!:iion off~hii! .1dlliati~1l Iti,edll~ antiisrrnis :'l!ho~~a1~tChrelil1ati{.8llly beQO!N Hue mk~o·' ,!:!~aph,.A dashed a]ii1,~inl~ne~d't~ma,lk (i(ll1'e$p.DIii& 00 '~h~ fl)(al plain€! 'inl the 'im~g@. Thii! ,(j'~re@ or ~ydraJtiJl)lIl or ~!k'i~Hi'fl.g Jl),ftihE! po,lVlmer layer a:l'e~ 'ClFfN!.s~ glr.a:dLlaU¥@)c:~()SS t.h~ sample hom t~ril:oo r~9nt CJJn'~5~olldilllgrly. the ~an(lc:()kUTln~gradl!~ ,ilIlly cnalnge tilleiroriellltatBo,n from perpiIlitdktlllar I;Q tilted. irlw i!\Iho~ !"-Ilng~ or 'l~~ ,ori~l1t@tiorl~ liS ,J:i'E1,f}idoo •. thu~ proviidi ~gl ,11~tiUi liTl!i!gl! or~h~ 'e!n.tnIII

or

reo:rieolll'l:atilln pM~eo~S;.

IFi!!!.. 4. SiEMaJl'II(I(lp'ilkalim~~~s sh!nwilil'g hlighly C!nntr(lU~ lPaUerlilformalii(lli!Iil11 lne HAIIRS de~igmt$. All th~ stWlttU!ir@.s !,,~@ rsill~y SllI~U:ii 1'(1Ih~ or~ gfll'lal 'Y@rlical ori,Et!'!t,nlill!'l Qf~'h~ na f!o(()hJ,m Iii!~ lIpci!l I Ir,e~'IIeUimig.A) U:niiro:r9'!llllfie-nt:ali(lnloU~e tiUed ~anocolumns ( in t.llIe tIIAIRS·]~wlI!.1!lrtlE!' templated bV the ~WMfl@~ (O~irrimlingsllll'fa~e thglt l'~ ~l1)lriillg Ilines. Tne d~re(ti~1l 'o~ th€ UI1€5,s ijndf:~I~d by ani i .:!rPOIO'II.{II} R~llI~.aT ,ana::r of IilIkr\(lf~.or~t~ ]nlJhe HAIIIRS-lil 's;trutI:IIlW'~ t'l!lfIiIplate'd by the sILlb,S!trate 1:i!!i1.=1rinlgr hroney(omb p.11tem .. (Ii:) An eo:llilmpte o~ <II o)mp~.eox HA,IR5~2 P<ltl'll'!1l, £rnowiililg a~ anay' ,of a mkIFQtr~p.;!:•. in wMch every 9lr~Ulp ~f four .aJtt~~h~d lI1afl~(;ol~nlImi!; held tQg@lth~, by th~ hyd'r'@f1el. (Ii), 1I11l!l E) Optical mkm91r.lIphsilil1agil'lgllli1iHCI'I!r'1Ip!i,!;ho~~llin. (Qirltl dry (D) ,1Iml OJ! \Ii\i:[ (:E} '5:l:aJle'.l1h~ sm ~c:hin:g ,of thE!' nliillnoc:~dl!lIII n$filiom ile:nt:rou J'ftl~d dl!l~tt!r:'l to a ve rti~alL ,orieonrtallio.n i:o; d.@ar.

REPORTS
3. V. OI,aIb, A. Ma.1~ub., Nt/IIlH! 3'76, 219 tM1951. and Ihel~ 11l~!jJJt !~e;ilJ \,':hcr~M b .." 4_ I!l l- IEleelie ~f ~, l\I'cmllf 411)11, S,8~ ~200(})_ ~Qhl!wm~\,,'vth <In <1~[1>t;'(:l r;nIjo 'Or gO csu fb!Iy~N S. f. 1+'. 1~1I!.~el~ Sirrm6! 297" ~. tt.O~, ~o !O~I~~~ ~hA;; l'iIl!IrI~OO\''''~Ililol]!hlV<lI.;;iu~, All ;:I~p~C~ 6. I~IlAIi!ilnnl1(1.1., .stif,l'l~ 29~'. 171 !l2003.~. rntio of !Ii k~!l~.t ~o i~,n~,", .. ,d",-d IfU s!~1 ffici~Jq~ 7. A. ~el1dll:i!l" I-!,'f.lig~g', 10. ]U!1lIl~f, R. ~l1g~r. ffii'.r,[1.~ 4a!4. 87'9 (~OO>:l. Ibv:l1(lin~, of L~.d:l bc:mll iiJl] i:Crl]da i'fu;[~(:igh!xJrs tID &. IF.ICh~~]~i el 1l1.. M(ill'ta~ ~rd' (omqf]l" ,'2, 12134, nbl,lin ~l(;, S~!1Itctll~ ~hnll,"\'n ill F]g, 4, C ,lnd D',. {,200llt Thil<. nbi,,;'Ch~I~!c;_d 'l\l:.l~~men!~:uI1~'kc"" 'po!;sibk:: ~, i~ 9. M. Ula~r. l- ~, i~ G~IJjI'iI. [I... GOO:ll!, G.1\Ii. 1miIif!;~ H, dl;.~gn~u hydmgdrmlf!oct).llIIiiQl~~ oomblil1,t~ ton dm~ iPro!erJ1lffi 4, 2366 {;!lJ 04l, C'.I'iubli.;::. 3 dc;s,in.!d dc!!!,,,,","C (I r Jjilt.~o(."tI ~1tltIHl;im'i. 110. T. ~~" l I'en!il. X. Gil'!. IL ~ial1;9', .'ke. Oilm. ~ ;!:g, 6,.o!,iI! {21001;lJlImdl rilfftfo!Hli(_~~llu!r~n , W~~havedevelcped hy;brhl 3ochilc';:I(II~ .1111 U. lIt AYU!f!1 foI' .0'" P{OC,~ll, AflNf. Sf!. U.s.A. '9~, llil:~2 which :9Jt'mysor hi!jj~1 fll.'>p«'D.-ru.t lo ~il.i~OJl imllo(,2'iXI,;n ~ndl rtfel'lllii~~S'Mr~n. co 1!.Bm~~. llltcl' mLl,\'LChcd or 1h.~~~nuJi I~~, are c 112.Ii'. \!'d:ilI!ll~, ], lit Sn~I~~, '{{jr~rt·~,Ol!li.EI~l «N)tI'~ ~l'Idi ; Jdtr~I!1~e!, ~iIIBfejn. embedded imo a~l)idI.ICi£C~mm and ~ aCIlL8it~'<i 113" A L G!iml til' al, Nat Afattl, :2', -16.[ I(2;OO3i~_ hno hi~hl)' contm]led, complex, luicl,,(!:,!,tllllCll!Ircs :ll~.s, 8, lI!i~, ~O~5t~ [l" a:. S!I:OIll'. H. E. W~rlrilltr, IR. ~poIH~o,mmclion <lnd/or s.\\~dlingGruhe rob ~ne" A, J, Ru~~en, A.rr;i, ,Ct.iem, S'oc, 1216, H~"O' ,~,ooO, J. The aClu,~li,m is f.1i."1. rcw~~~!C'. rCil~mdll~i~)~~. 1!5- I~ ~~mal'(;heo. litS~l!IiJi~. lB. .Mtd1I~, 1iI_ 1!!~~!J~k,.MI':, <lind rolillll~l\l!,r~bo.;!i~YI; t~l;,IItlhL,.~ OJE:'hill'twr"1;'1'o /&tm .. 9" ,{oil (199i.1., :l!6. T. H. Kliupmtin, .1. A. TalflLor. t iii. SMr!ll'id'fr, S. ,!,j!JlJ~, Il101:!' kacl te a \'~n.i.~~y U.' o .,uWlut'.oti{~IlJl, ~nd ud:ing ~aftgll1ult ~iCj ~i!i,Z~ ~j!OOIl~. O'Ltl~l<l~('I;n;:.cnn~mll>d 1;;;\ ~fSlhi~p'mem ~Qnl1<l~ 1l7. IL ,:!i1'ai.., F, (. C:~Ii!l1'd, R.E. (Ollim. M, F.. Itulimer;, .Mwi ~iCl'n, n~icml]luidic1i..fI;,;'\,~I'lI.~!c; ~"rifdhi~~g: or !hC' If,[t~\ 1349 U(lOO. W'~.!~D iug ~d1!lvi~[. um ..bl~ phCllonic :.;1:n,U(;UUlrt)5. ~" 5, It ~e., in N~MidB: Bnlij~ B~$ for NclllilUO,Mn.oJO!ID'. lJ. M. iRot!'l1.O" [d, ~tilM!r'. ~~. '!!lank. :)Ir!if'iej!111 Il'iu$clc> •• md ~k~~ ~ys~l)ITl!'i t}:5)2004). !!'P, 145-.1.701.
~1l9.M, K. M:M~.I,.IK. l~dni!1i; $. Pc

s m1lU ~1K!ing

22. K. 'S. ~.!ftr.IE, :Z·d~~o.: H. M.ail. ], PiQfi!et:1';. I, lwiml!'" MOtr.Q.md~wfes.l6, (15019(:!!O~J)_ 2J. it ii. GatJ!'.! .,1' 01, An.gtw., Chtm', In: ll1..43., 2700 (:;!OO\1~. Vi. e, E'. Rwwen. IIU" F6l:"R.lB, &ll'll~ lfJlIMmtf:lWlrJfJ)' ~~ji (ill'!: Thl;Qlil:l!\ !6!:~JiI~ (lit, ~. ? 2004}, ~ 5. Illi jillIlt1ClJllllr. eur 1j:f~Uriim~rl!' daM ~ Hlial: ih~~ ~l!Ict~M m dlrll!l:1 the UOW' !f!d ~\WI!de-I!~!'!~POIl1: ~ ~n~g5ll)~~ t~ !h~f~n~it;om I)f ~~~, im ~'9~ln~~) ~nclthLJ~ liin<f! f,iIlIMt~~ in Imi~lrofitildr($, {Ii~ rl!l'milhDy d'i0!1'I!lI.I! ihe'i1 pfiQlooi: proptrlill'li and thus can lit amsideroild Ihr hm~Ii!~' p~ownfc;, h"u r'~mib!Jt 5I!ri!1:1i1 IMir 1j\'~1i~!I lMJlll:liiitiffrom~Lip~)!:IropMbi{ inlliill! h'li:mj~ ~r~~!ll; t~ ~y.;;!r;""'iU,' fg U!O!:dry en'!1!'Q!'!me:m" ~ f'!~ (a,i)hilP ~nc I1lI~s~ ~I!rl!id~~ IE!~~I'$. [II <l6. It. Btidiclm1I'1'i:1.tr·flll[(:r. twtfgffD Art lIftws AIln[o[, <11, U2 (1~6:U" All!lDawi101bLe ~t _,!l!l!gfa'lJlf~Momr 1bfi'lG1t!~p.aP~IiYlM$I!9ri1¥1I!ll!!>!SO!Ul1im I. n. 0. ~, IIlI~b~, ,Amilu.. illY. "'f.'lQ;L 59. srs U.99",. :2 13. Mi. lDoi;fD5lI~, ~d!. !i!le£ (amm. 3(1, 3U, !:2:00J). ~9" W~U\lank P. KI:ll{l~n~, 1.1Il'l;:~r" I!_ lurilil~1Ii ilrI(j1 I. )Q'kolQ'IIJ:f k~[1M dill[ili~kiinrs:. W~ iMnk It. 'Simi!h
a,~!'i M. UPPil!lorttdLni~~ aMiIMI1I~', 'ThPi; ~IH 'W.iI~ ~lIl\PIl'rI!d n~ pari: by th~ OrEicellil ~OlJlI:lI'l.litIl:Si!'aa:In, <w0lliiI MOOO1HILS-HI"09.

5l1l1pIPortill!ll1OIll'lfllli!

Nlat@ili'iial

A!lMr;,.Aid"v. ~ft

,';SIlI,~.

_.j,(~Mrema!J,aiW'~!1li!cam'.@n~wLll3:1~;eW437IOCl Mi'lle_rloilll; ·ilOO Mi!ltl1ad~, FIgli- 51 ~ ~5<

TableS],

R~fel'elil(es <Ifill:! N~IIJeS 1,S. Wri'!:o, l!~we,P~ ~r.rnaWI': ~if1j Qfll1 App1i~!~ ~el~eI~ .Ami!'l,1lI\ 2006)·:lIliidlf~k!fell(l~. !I1lerl~i~.
't_ O$Ol~a, H_ O1LJ~~ !fI, H~ J\I't:ltiJIlg ~~~, .~.~;;: U:\!I9~L

15. 11101 (;wQIr.l.. 20. ;5, ~a~ J..!!l. ~i!1!\ S.;I!'!I, ~1;1~. 3'''B {z,004t
211. ~1"ft~S 'iilmd' Illi~ild~

(JJ!l~g, C. A Mirtfn,.

5ciMt:~

~dereooe-.s
,;!006

1IIOOi!$,51 ~nd Q
,2;!I Sfpi!1!I'I]b!lf 2000.; <I octplflll ''lII)KM'i~tir

,Ill!! <IVlI111llIlt~,!So sup,pat1in!!l

nia1leri~~ lili S~nCf' OnHn.e_

lQ,n;l~ielfle~_lum,Q

IEI'Bclromsc·h,8Ini'ca I Re:S,'OlnBlo Irs Iro'Ol Gr'B.phene ,Sh,sels,

l

~i~"" oJ'

~hc~~ !ilriulflilICJ]!;P."l\Ol]'IIO~:;>l.

il~C'rLldil1g

~O!~auC\;' fnt!!WlJincy. l!pring Ctll!c..;i::,UI!., buih-in !!:~IS~on. and CJ~!<!~~tyl:tC'tOf. SUSlp(;!wcd grap'hc~]J:,s.hiX'Isarc filIhricated willi, a pLi\:'lin.g 11]1{lC'.l.~'i~i.Ii~ilar ltD Iha~. ropO:I1eeJ, p:n:.:v.i:.
oll~ly ~5-i), hi m,lIrCffSC:. lilt; gJ:ruplwne :s!](;"(~I~ a:1:C modimi.i.cfd~y ·];_~JOJ,ilii~cd (D"(Tprodc'~ini:t1 Ircraelic$ ddmd ~l'iItofi stU:! surfecc (Fig, !) (9'),n~ rcsLI~l is a jn kron·SCille dO'uhly cl!ampoo ocfum 0., catuil:CVt'f Clamped to' the SiOl :rurif"m.ic by van cler \\'aa~s, ;[IIHmCliml. Some devices, hm. e l'rtlx'Iitterned gold clectredes bctwees ~~lelfl:mches~o IH,lke dcct1'ic<l1 oomoca (fig, 1. 1\ and D), t\ !1OJ!CoJlI .. C'~nlOdc "'1.O!~dc rg~C ~llkro.~c ()\fI~)

N~Mele(tr~i'ii'Ie(h(ll iIlic~il SY!l~~sll'ii~~fa bficalt~dlhoiliii slFigl!e- (II!lId li'i'Iwlti~~yer !:1 rrap~iHi>e sill 001.$ by IIfi~Mnli(a~jly ,~x~o'l;;a.lilngthiln shee,tsrlr",mgralpl1nite Olt~~ '~ren;:cl't~~ sjllk",n oxide.Vibfa,ti~n·s lmi \lIIth flJnd~fli'Ieflt~l re!lll.i'la.i'iit fre~lJ!en des, If! tne III~e.gaherrtz Irange .areilllc~liIi!lted e1itliier olPlica Uy er leoledrkllllly ,[In;di d @~e'c:tedl ®'pti e.. Uy by il'ltelri,e,romet ry. w~ ,d,emon"otro'!te roo rrHE mperalure, ella rg,1li' ~ensiLi\il~tne'5 down tea x:W~ 1E1'~ectrmlis ~r r®at hertz. Tlhe~hfllilli1lJ!lst rl3'S:Olllaltel" mmiro15 of iii s~n9~e suspended IlilY'er o,f atoms ,alld re\present£ I~he ultimtlt.e limil cd tJ.~o·d~lIIiIerni(lll'li!jl li1~nIlD@lectl'''liIle'(halnk.aJl sy:s:t~IiI1S-.

IhTtknC'.~ of the

w,~ uiWd

'!A;l.

qu,UIlII:W':!!:vdy ~UC'iJlNl.U~ Ihe

'~1~.$

(lin tl~e S~lbSIr.t,~

:nC'..... , W dlC

. tILl; 1;0:1 nh~g dL"4;a~l~:s ;I~ [lliI; mill illDWIri;a.!~iO:If~ of c-f.L"CI.ruH~C dci\' [.cc,~ '>'1'.1;;, in dt~ l)rk",l.ol!J~ on{;;:>,.

he milli ..lli,mri ~i{l!1or d!,l~lroml;iCh,ulic.d .....

r"l~l.lis~~

'\ io.""

l()

be

,l\::~

,l,~ reV{~!~111 LomLry iu~

D,,,::,'icc£nrn~giu..g

fn:m,

fl31l.0&Cfllc .rt'J;;OllillOr:s.,

:>w~chCi:l, ;[lftd valves 1\[I''o'C ilpl'lIiciJ!lorrls h~ til::;:,k:> 21S divcl'$C <!s hl~Ol'lln;'Ltion, p~ .. illg, moJ!cclllilr IDIKlonliplllllLtion. m1.d sensing, Tine Ii'rol()WpicGlI nume-clcclml~hilnk:a:1 :1;y~l.cm (NE~IIS) is OJ IlmJlo;,~O!!IC' ~m~;]!~or; ~ !J,,;rnn mal.cn<ll Ihm "'~rni~ in
re~llseto <lI,l1 applied
\\(()liI]d

or

Llli,lmlllc I:imil.

extC'mal foA."C:(i. 1~, The be .. resonllilor one algm

togethcr by weak ,'an clJ:r \,VilaL~rOuteS ("n hhlls c.:>;ul:mcly higb ::it~l,gi.h. slm'i'lCSJ;" <lil1ddtCn:llffi I c<mdm;:l~"i~y akmg the b.~~~ P~<lillr:" 11'1 addilion. glCuph i~cC<il11 be exfoliated! QI1~('I Vl!ll r 11 Iming ;;lli :1;llhMmte. rrodtlC.~llig mi'cron"~lzcd gmrhcIlc sheci::> withl~licknC':>.~':> dO,,"""Dl10 a silllgh~ alo:lnic ~ay,el" \,r .'1), Thus: far. 1.'C&:an:h 01:1. thC3C Ibiin gu.·a;pherae )l.ilee!!'i 11<1$. ~.bc~)!cdl l~r.inlOlril:y on I.hd r
cleocll:onic
G 511~X:~1di JIg z;,llCC~" P\!~f Can

~hic'.k. lhk~ l]lIIl1! <I1c\'Crc ~1rn1!."~r:"~ on Iittl !m~· gnl~· ~...:illl. As .. ,\lil~!ll~ I..y~[ Q:~'<,!!\tlI11!ii, i~ ~!l{llJtd be ~'IJl:)U~I,:;,~ii'f" Olndt;l,ibJ!i;. (irllplli~i: t:o:n:;,i~I.~of ;-;tack,lXI IfIY"::I:i; of gn.'uphcrbc :sh.clL'ffi l>'Cpm1iII!;-d by OJ mil ;;1I:ad heM

mu~

IIl.'Hch. a",~h{n"11 in Ibe insciln .f,g, ID .. lh:I\\"C'\!cr.

~<lr~h(lCI!1nhinnC'~.·~~l'lll.
~Url.;'llilC;:lll:;; ,Irl;l

I;Inn;.!linblc (10

2 00 ~ nnl. ""'I:d~!1Il~'''' J'2l f<)f rh:;,....~ w~

u~d SiJ]3t i~II!ly !1"'i'5rltv,,,,"d,R:;Jlll1>.1n :\l}l,"4;III.o~opy ·to Il:kunillc the ll.uIDllbo;;r () rlay..:r:,; U~ig.I CJI UO-i 2). lb,; .gHip.t~~m~ sheet ill ,F'ig. 113 h~~ un AF'M,· d~t\:,'m' incd h<:ught (I r O. 9n1fftl. By cm~:lparison w~lh previoHs r'C:'L!IDS O') J 2). IJl:e shal)C of the r~alfnan [)!::O),I,;, neal 17GOcm-~ sugsesls ihe s~lecl i~I,WO ~aycr.;ll~kk, ()'v,cr the area, lying:, on ~hc

SiOl! Iflyer.

S,\!b:l.tllittc
O\'oC'li"

{Fig, ~. ).\'\,~leN'<!S C

sl!Ispcndcd

Ih(: trench h... sitJlglc

.he ~"cCI.iQll ga:<fIjl:i1<;m:

lC:orlne~ (l?.mror 'ro~ Ma'~ri~l$ 1ti!S~1!rdii"(oMiell Uni~"'sity, HN<I., NI'J" 1485'.3. USA, '~PomQn~1(o(ifge, IDeparlmllllil ~f

1ilf'(!ilX!'rI'~CS.\'kc dcmol,. .. tmlol: <!mclhod

5,inl';ll"" and

mll

II ibycrgu:~I~hi:ne

Plly$ia:, (It!~(Ii!gI, Gli! 7:1.711. uSA. "To ~!bom ~:o~~e~ppden~e 1!IlIG~dlbe add~ess:ed. ,E"P1ait

'~!~ndi~;\1 and $how '~h,L't ~u~h,\1'11001:5, ~

A~~ n.'OOllalm" m~~l'C'nlc~ls, <lire ~rfO:I.mcd ai, romn ~'cll1Ipc~,llu 1:C Oiind ,j;! I'U"C511!l1! {D f< W-6wn:' rt: unless Olhet"l:vlsc indt'<:a'!!;;'d, The :rI.~a~O~1~re OlC'uJ~lh~d h}' lII~,i!m ~.jlhc!" c~oc~riC':1 (Fig. 1/1.1' (II," ..
Qptjeul Itt{ld!lliI11Ii{Jl].
!lIlodiJ~aljon. !i

m~~III!illi@«J'1U .oo11l~ltl!oo

Illillh~:,,~. ddaircd l>IUltt~

1x' ~i~odb~unio;;:a~ly.a'!;'tuah;;d. TI.lis\vol'k. ;d~o orlh~ Illtt,'tbmiit1~11'ir.upc~.

I!~[he

IL;".:II~

of

,..:lccll'it'lll
frO

Ili~lit:-\"dg:YHtlg Ii!lu,iio

flt.iLl,lbi.::~'icy

REPORTS
F'!!g.1. tA}s.c~~,liJt" it ef oill smpl!!llIdled '!J ~aphene IIrtS(lf'!a~ r, ~B)!An ~ptiica!lim~'\J1l!
nfa deubla-layar Ii) rraptienf slii ~t 'that be,colme:5 a s'~l'Igle SU!\P~NU:!ed ~1IiYerll!'!Ier th~'ill~ll(h. S(,a,le boa r, 2: "IIiIl. Ea~h m~0:r,ed diid~cOf,r~p~j'jd$ Ul! ,aJ Pl!lIiflt WiliEl'r1! a ~mn spE!(;:t.rum MI; ilT!rt~Sli~d, (C) Raw!I!.a.r!
:Siigll~tft'r(l,m [!II.': h
iii

SUfi' (lJI

the glrapheD1f

piece. ~dl(lie d scanl ns l

Id<ltl.atalti~n alt. e.<Idl '01 'Lhe ma.tr;hilll'g m~err,ed

drd~s. The' tllP S(,a,j'i is used all, a re<rf,rn~e

,;Ilndl !L')(!r~e~PQ.l'ld~oo

th~ ~mansltilft. 'OJ bill~k grilll p hit!!. [[II) A~ !IJ,pticC!il imag;e: ,of rel'll-l~IH (-·4)graJphel'le suspended
IIlV>er

,contacting ~E)A
'$.'(Q

a~remlr:h ~md a 'g,o~d

I1m~,g~!~(trt)1l

I!.'le~tmde. Scale bar, 1 ~lm. (lnl~t) A tine $Canf'Mm

t~ I'lIl1n!!l molle AFM C:Qm~:\lpondifl:g tl'lthe '~a:!lhed lii!i1i@inl~ne optkal mic ro~ot}ll lPnagt' (If,~ fi!'\v-Layer (-2~) ,graph,e;nil! 11~(I~aroJ.:5((!~~ Ibi3Jf,1 J.lJt1"

ima'gl'.

lit sIinQ\Y$<I S~E.'pheig'ht ,of Jl ,.5 mill.

IB

(]craphene

n;l~l9,tw

la·.. 'U!

['actor Q 2Bl Agll~c 2B shows suni1ar res!.llts [,orthc: ;5;il1,gll::·I'\jlcr grnpht!1C 1li.~lmIJ}l' fro~!l F~g. 183::)0 = j"O.5 )'\An~z ;;mcl Q = j'ri. rig~m:;;~ 3 sl1JJ\,\'s
'Ih~ rL~~~ilh!\ 1tIC{1L'Olll'l;rm.'ru:ll of

of 33~onl:~!(J1Si

\~~III.

,'myil1M Ir(lllil iI ~urbgk:rHomnc lil}4:r IQ sheet:; 75 mnllhic:k. Uti;: fucqLll'n~cy_r.i or [he fili.t· dan~ellm~ medes varies 11'01111Mliz to 17ij1.M I ~:z. l

Ihicklt~

with qmildlY factor Qof.20 to K50. For l1~cdaallica ~ 1)!SOllaIJlr:l lInder!cmas.i on T.

Ihe f!.luadal'lltClltall I\."SOlmllCCmodc

/0 is

giVCil

by

Fr;eq;!!!E!'1ficy(Mtu),

10 Fliie.t1yeilltY OilHiZ;)

15

jo.

UA

l£11)1!!',l: ilL ~]~-I

J,i2057 T/JPL

2nL'illt.!

(2 !

F'iig. 2. (A) .Ampl]~ude ve r~U:o;rireq I;lencyfl!l,r ,~ m~5·flm·thlck un!lIhiiaye r gW;:lip hene re$o:nat(lr'~ kemwilh IClp,tkal d'riv,eo. (11f!;s~dAn Qpri!Cid ~Ima'!le or the reS~PiIal(l~.5(ate oor, !S ,~m. (EI;) A!i!IIpl]~\!!dewr$;U:; freqlH~,nc:y taken wiil:h optkall lIriVie for tn,e fUl'Illla me nta I mode of t he ~in9le·la,ye r gr<1lpille nE.' re:S~Ii1aOOr~h@~\iili ~!'iI Ff9. 19, A Lorei'iuia'j'j f"i~ ~t' the dig~ ~>sh:Oiii'i'i in red, vtl1'U1g': 3 j' ~~I. rru~luc",Ky(iil:;:L1p.;;ril1rlpf;l~1d lOp on of il OOJlSK~nll VOh,:'lgc <tl.ld,lPII~ liedto the g.rilpht.:.m:: tht~ b'y"~' 'Ih;;l,t k=lItlf;. [0 moti:o.TI, hI oolh t'<ilM:i>, ~h(~~nal ion i;:.; (h~lec[oo lil)! :lllo.n~i.lt1ring, the· :UX:. fleeted :1:~,gJnl1!cn,'l;.llty :~g.om.;u i SCOOIl.dlaser with Il Ia.'i~ plw'todi,oCi.'c (91. Pig.Ull.': .2A S:l~~J\iII~" Ihe nJc~~"wJ[('iIl;u.npli,.u'(ic
vCr5li~ i~tlltcitr,y for [I ~5 .;nm~.~~Ii.d:. :ih(JCD :;'lL~ •

\,,~hC'rc It is~l1,c: YOI!U'I~'s mooL~I~iS; is the 11l<1L..<;;;O:; p d>;;!!~!ty; f. 11', 'illl'il. ,lUiC ~h~ dimci!lsion~ or ihe
:.-\1~P01~~~ g!O.ipil~Dl.c ~ho."I;.ml,~~hl,'l .;:~mlllln~g

:>b(\e~.The fesuJ:t is an ~I~c~W1$pcndcd ~ph~lC

I'd
. !:' :;:: j.

~1~Glro:;~I!I.uc~'oft~· ~IVI'OOl

sheet and ihc Sl1biMI<Ltc

pended over II 5-!J1,m,re.i~dl. MIlI~tip~crcSOiJ1:mllCCS I are observed, tllCliil0i»1 I)romim:nl m~c~tthc 10\\1lC'iilision. :1)0'1" both eastilevers and ooa~ns, ,~'~~ero \\'heu,'C elI.' h:;~hc dcrivalnve Gfl ~u:,': capacil;mce gate values for bulk grnphi:tc C:S~. rl1C'(jlll.en.c:y. We associate' Ihis dl:n:ni mu:U, 1~"'31k we have l!Isecllhcknowll with respect to t1i~cdisl1!!~JO:;:ru ~;Icg;_lte; and r,~;!1~ with Ihe ~lJ.mdmncnlal \'~bral.i:onal mode its p 22QO kglanJ ,md II I .c TPa ~JI, n is i~a O'iJlOEo 1'1> <JR. respccl.jvc:ly,1 he de and I. ~f1"!C-':~nyin~ detected lnlcl'J.i.i!y is. l<ifgC~lwhcn ~ht: n10IL{l',n is 'II<llid compilldson cOII.'l.ldering lil~ extensive rl' vlJl!:<Ig~"" \ppll~>d ~. dlC g.1Le U3).For op!ic.~1 in11ha.w'1ClWos Ihe r:n~i!1::MDsllCnded SC1;:tiGPl.Wc < ~horn''I1!ic.:d and ex pcrimcn~'1I~"'olk ~hOJ!I:J10"'NS !he h;!s;.~pbm~ gmphi1.c '0 have .,L :J,li!1r~larVOIluc ;.u;IU.'~1 io~ ,~h!: .~~1'tcn''ii~}' d i(J>cl~ h,bcr fbC~~'OI;;1;! wi U I illli~ Ollt diseu !'.SioPlpri!l11lri~y lQ thi.. fun. CD f,~ ~(U ~~ gUlwplltltC1nld ,~mh:';Ju Ilmio~'lI~"~ H). E. {J. damJtnll:n] IllOOC. A .fi~to ~LoOC'm:ci.J1l yield.. ;1. n~ [he ~1l(iG:[ i~ 1'I1.odL~h~h at Ffl;J!.1l!H';11U;:y _I: .."d, ' c iiiilll~imig a p'l.."Iiod~c Otlli.lrJel Lg,.m.fexl]mlt'i,UOnof ~mll~L rlX."tjiJ>;;:licy fi!j, = 42 Mi i.z and ~ qi.lil.ilily Tom::tOLilil rOt~sibl~>: lnDl'll ~1'iE, ".... plCl11. e

. ~ '-lit·.r;·'

,..-'

.'~·I' + CII.'~' 1.!¢ "'.Ii IJ(,~ ",.r"_ .. "Ii

u~mei~I~I.A. i~1,0) tbr dtJ\IMy d~mp!.,:"d ix,,~!ll~ ~lIJ.d0.162 lo[ cantilevers (l4'). ~Il uhl::~imla of Mnilill tCIir:>ion,. Eq. J pR:di:Cls tlmll .he 1'C~(}mlnC~ .[i:\1qLlcncy}o sceles as Ill}. f'ii!;lure 3A shows Ihc ;I'C::lOililill Ilirc'lu'Cr~v of Ihe rLm(bun~m<l11mode ~o,r, It:'SO~lator,s with t > 7nuln as <I liIlldiOIl pi10itw..'tIas G~loo :sqU1HIC£. Al~op.b~cJ h: Ihe ~le~iic.'ll ~pro:-diC'ii()n, Eq . .2, ill, the limit or zero

or Me

or

4191

REPORTS
dashedl ~n~'i!, correspond ~(I '\'<1 lues (Iof E: - (I,S thi!l,t TlP~ <lm.1~. TP", The dm<! r(l~lQw~h~r~Clictin~l:) fla"LwmlMy ~(,l~~r;I~~!y.i:m;Ji~ntil~~ ~Im' ~hi~"tJr

The sing ~1<lY~f grnp.h~llc !\,'OOI1<!110f :1hQII,¥],l lfl F~g,. ~B inl!:;.tr.nc~ 11w: rmpQ!141gl!:C of IC!~iQtn if! ~11'1; ,fll.!llllrl,,;;l.~~O!fl\l!~QI:'l. h h",'" !I 'lu~~~I~~1J11"1! f!'Q. fC~OJl:JItoms. !In,~ in [he lx;n!~ing-oon~ilmlb;d li~11i~ qll;J1.!ll~')'1m = ",10., j'vmz, much 11~!l);ht'fimn ~h~ l :5_4 w.uih a ~~ ..ncllUhl" E Chant(11~i~~i.c or ~lil· 'bulk l\,H Il.z ,rrl,*t1,.~lt:y~:>irMJCk'tlrt]r L h;ul~~onl- biL'IIIU ~ h"C rml~cri,iL lhi~ is ,1!nO:!lg!h~ hig!'lCl'i~ m(:u;J~lhl!'o wi ~ht = r03 min. L = L ~ ~un ...mi u' = !,93 Jm~.
170 Gr,~L U~ From 13q_ 1, '~hiSl irnpli(p-;. thai the gr.;l.p'11L;.Ule ~\,,~1$111l[, ~l~ 11 'bll,l:u]t-in ~Clil~;(lnfi (:If r = 13 1:~1·\t n Flt:llli1 fllir.: c;.;pn.'C,'i5UOr'i ..J!~ = T £!n l~m~~eerreM spoll(1l; to a :-tilli.l~ !:If ::/',2 >< lOll'%,. An ilnpo~aitt ~1'i.CflSlLr,~ :Iili'ly i\.'WfiiltOJ' it, [lilC of nOnfm1.lI!;rodwi:dll~ 00[' lite rcsmmnce peak ehnraetcrised by the qlllillJity raclor Q }iJ/8/ . .I\ h i!1;ll Q is c5llromi.\'L1 ror 1'I1m;~<lrp~icmiolt<: ~'UI~it In-

.m)1Qn<l.1.(l~ ll], cl;;l~L;. gI'L.'a~,;;:r t~m!l 53 ~o

12· to 300';lTUtiJ .~~ i!::'k Si C'lIJlli k'l'L::'~ and sin~~~,lr ~1 l;i.nn!!,tt:"\\';I~J.Gdcatli(Ul n'i<1~101~ll'I~~ l'll~d di runl'iOJ'i'li EMS {H 16. 11}. I,n) Clji'llrJ::-;\ [0 'llll.l'dd~in! Si

ealu~levcftl:. 11l.i.:: ,~i}P~IC~I~ reS(mil.ror~, shtJl'l't,I 1\'1) dc,grn.dm·ioll in Y:oUJ'Ig'~l1'l'Odu~us wfrtin decrcasin,y ~hickflc$~t 17), nco 1\."l\O11:Inl rrcqlJC'm:k':l; \'CI'lll!! ~ IlL ~ ~brlhc reso:nat:ole.i wi~il I <: ;' mn <l1lC' Sil.GWHO!." OJ;l!C'VI ,yqlIOlP.-S i~~ fi~ 31\. The lh;quC'nciC'l' !h!.;'."~ !llim'!er .f!;.·i'Om11,{l,~ ;:.:hm'l.' nl(~,!l;)~'tCO,l~1!t.'f; wi,.~!h~ !)'1,~tlrityIwvi!.lg l1I.';:.;tlI!,IIl't &1."tJI)I~m::i,.;.';.>l hig~~r 1,L~m d p~dicLCd b)' bend ill!!!:(!kll~C', A~ikiC'l)1ex r:1~U1• ~kllfn • Ibflhi~ is nhlny or~hC'fl(."Ml!1'aWI1'l· om;: u ~~dl::r ~c'rl~iPJ1. "",hi(;h .~n{:TJ;la~.td,~~\,!~lppo.nlinMO!'llil1~ I ( ~ru.;:~~- he t~!1~ion likd.y 1¢;11;11ru. 1hI!'11 till; rabricOiT ~irmprot'(~ when: dlt~ ~i]C'lion Ol'1:Wtlr:!!nhc grull!h ilr.:allldtJ-~;c oxide :s~rr30C dlllrir~g m(lehauic~1 ex rol ~[iOil !l1r~[ehC'Sthe W1'P.~lCI~C sheets across

Clr)l,'lg:ru~ M)!] gmph~J1~ [,~';Sr;)mlloU); ~hQI[II he IIJa'i.M ~I,bl:c 1~i IO\vh.:nlJ:>i,..'1'lIlUR:.:>_ E.Vl!n wh~n "I 111.."5011 .. is not bei!!!;; dl;V~"I.'IIl" ~l wi!I~lil~ ~ilhl'lC'dUiC'to ~hmT!",lo;:d~,ll!{lnby !)'k,~~]~llilll\~ (R!\ltS I o.lrn1[Jolm:l, -~U~ = Ikl';lfiKrll'Ila. ",,'he~~ Kdr = m.:tr w.,} = f1735Lu1tP~:! l.. ~bc L'J.nixth,e :OPl'Dltg !.'(ill~fulill of lh~ It1fiitfc (2~ All. CNm11flj~le 1.., :S1'1oWul in Pig.. ..j.;\. ¥.here ri .5.;mfi~liek

mlii-thiJ;k l\.'iVOfI.<ltor r~!fIld;;l, '~~mn;IUic increase in Q \'\l~tl'~[~~<I!"ifllg ~e!]ilI)f;."'l.~U\; Q = .~ m 300 K ( 00 ~(}Q = ISOfI ,I! jO ~J- ll~i:;1;'i\UJ.iiW-S~'ii !h.. t~i~~ Q

or

'0[

'I.~~.

l'l'SOOiIlor with
ha~

to -

35.S MUznnd

'tC,df-

0.7 Ntm
n'l{)'~i()i1r

it lioom~l.oH~pCl'm,tlU''C1 hcr.l1tml

RM S

ar

or.'I('!I~ 76 prn For resonaiors ~f{lrwhl'Ch the creases the ~sil~vv[>, ofthe ru."Oll<lIOI'W 'C'.x~C'J;nal IhC'I1'111<l~ v ~bl~>!i~ions can be measured, "'t'e use I il~s !h:emd RMS mOliC'lB lO scale ~l(: Illc<I!!i\lrod j'lcr1urlmlio.n. A p;l.otnrli1c Q '~C!N)!.~hcthick.l1co'l!'i. t ,d~!li~gr.:1p,t~L,'~!.,;" .~m~'OiJ.·$ (fi!i:,' )]3:) ,'i'hOW~. IilhClltldi.,,!Oi'."~or\'('lIWl;lC '\\' itll f4,;':\O:I!n:!O~' di~p'hn~Ji,.'n

rm

~h!ll. Ilm!.~~ li~'!I1n (;'k~Lr cl!.1rc-Jl~h;llfiCC nf QOl1

~~w~n ~(: :;;l;Ippol"li~l~ {U)]~noj ~~xq"Fi.gurc 4B ~

nm~

!]~Ctrench.

:\how,~such ,I re."1C'Ollfcd I'llo~('IrthC' odii$plil!c~mcnl II};.on~h ieker a!11p~ilUd~ \'l.'lfl.lU!'i.rf·cl'rhre v",l:mg.c,'lilc !'t'~O:!l<UfI'r NEMS !i(."lillm.~orsrahriC<!led R\l!l1 ~i~ie(ln (lB.), The ql"'~~i~'kC!;Qn;; fU~ !OOvn t~~mp(:r<.1~ure~ru~nw!.,;"r i); !i!w,;;w Will ~od~~«lhu."r(;nl!!,"nl$i 3 um, or 'on Ihe than d1i~lIti1Ol~d :EM S (2500 to 3000) of lllirni~ar I;lfdr:.1r of i&snh.iekm:-:;&. whl;'t(; 11O'JlI~1lr.~~rilLIC:-;m;;l\!ilN volnnlb~ and signi fit'~IUi.~y im~t{.'1I"Ihl'lJl h~g]l-Sl.re~:; ci:lllLcdw~[h ,rl<!d.uli:ormllr:Ii:i>uo:n ruJrukll'lOWn to SCI. in r(2j.. l1i:ii~ IlO.ui~iltt"~uri.ty 1& characterized DS"u de.... S~lN4IUlJl:(lsl.rin.g.s (200',OOO}r, y~! :sin~ililruo Ihose ViilliLlil rrom I) lirrlear ill.el'c:as~in 'Ulllp~ikidc widl reportedin Silflg~C!.Wailed carbon u~~nmlubestSO w 100) UJ, 16, 19), P.n:Uminru:y slmltcs OIl a ::!O· driving laroc and! aC(Ol]lPdinli~dbYH decrease in. Q(Fig.41Jl. Two ,<J;pplic<!l!iol1 .. of I1runolnccilanical resona", thickm.'l';'."" This eneuasts wi~h
~'eSli!

or

,r
of

tors are l!tt~~I~O\\"mass ·dclcc~iou (see ,..up;rnmli~lg Qulim'wxt), and ulrr~~<jc!~~~~V\C' fnl!CC d~liI,,'amn. The tl~l~mate litnll. en tlilC ,lOO::eSCI1~:ljv n')' :is SCI.bj' !.Itt ~hcrJn:iilinllClUiltti.o~s in the I1CS()lm~01:
!!II (1il

1.1•
."

..
;oo,~

__~..Ari
",,~r-

.400 '
!!II !!II

III

1!l1

scl1si'tivily 01"00"9 fN/~-h:~. from, Ell" I, this; {'o,r~ IJ'C:SPQI1,d.~1.0 a charge :>etIS~li\iHy cf (/Qf = rJP'

For d'i'Cr:~nl,1auor J11Fi~ 4A •. ~hiiSI'ClIuh5 in a .eOute

so
F'ilg. :3:. (A) A plot sllowin:g th~rl!'~i,l!i!ncy 'of ·th>e·hliiiidla'mell'lti:il Imode ·of .aU th~ dOiUbt~ ci[.allifllp>ed (illnU·~E!'!i,!\~,r~ ''I!i~rsu~, C,j!liItii~.evl'rs, tfl<l.IlI{1les,; d'OIubi.y d.al'llped Ibe.ilmtS wi~h t:> 7 nm, rWeirll5qlli1lln1!.r;; dl()l!1 dalmp>ed bei3im~ with, I <7 nm.. lOp ~n sq Illarre~. AU thkk !lIess.~~ a re d~t@'~m~ned bly lb·Y:Alr::~' 1fllesoUd ~i!llrt ll!i the th~~~~l.(al predilCti!!lflJ IO\Ilt~lhfWOOM](IJI'! ~!lidl f= 1 lP'.a. 'fhe ,ci,asi1f.ldliru~~ .• 'corN!1'poml ~o of:: 0.15 TP.a .. nd ,2 TPil. (B~ rh.e qualilyffa.ao:r of the ]UlI1Jdiam.fllllall Inode Wf.!>lIS tl1ij(,1<ness: aU 1~:Si(!n-alO ~s; mleQl:iUi~ed.
~H~,jlllm,me

tie.

'r(l~

d is the distance betweenthe £rn«)hc~~e s~lcel and thoc gate electrodes, '111S is, a high ,'geIb."'i~ivu~dcm()11~j'm.oo .m room ~~mpcr:amre; ~L'. krw teIlnPCI'1lI1llVCS. \\,'i~!~hc OIlSC'I of !lithel· q~aliity r.:1~()ffi,~l could dval Ihose of rf sb'l8!'e~t-eClmJl mm:iliSIClr cl~-tromctcrn rt ~ :.. W- s ~!Ul:~H ],(1. :ih lhc hiah YQll!1og~~ mool!l~L~. C)i;u!C'llllebrln~'1 !fI..J.M;, ,and :Ia~c ""u~1l« <1«... 'vmik" ,hl.'l'!I: P'\\Q!lat~t"l\, W\e~.~lySUiICCI,
t{;{tg.lJli! x W~3C'J)~

10--4 cmz'l.._

400' [Ai'

,.......".---=,--~;;;;ulo.oos,
.. ..
'

;'C

'"
_., I,'.

~ I: ji

'f' s

.ro II'
I
1

I I

~~~

~-..~~

~' 5V

It 2(11

I I I

~henna:1
Q.4, 3.\I~(,mV), 0,6

32
~~lm]i'lCV

S6
(Mtib:)

1100 8Vlll(~V)o

:200

F!lg.4. (A) ~~OiiS~!X»WJ d~!l<~itf~f:S~SrrEiq,!.!e-r!~'f t~~~fI(jt .:ll ~~O~l!t~(lfI1 1!)!:I!ndl¥licith,of m kH~. (!r;(:iltt) An @p'~(1!I,irna9~ Q'r th~ ~·sool~lQr. 'il'h~' ftSQIi1~l;Qr h!l!~ rdimemill,m;; t ~ 5 nm, l·~2.7 ~U1n,andl w~ 630 11m. S(a~!1' ar" 2: p.m" ~B) Aimpmude ~f r~Cln::aI'l~e! and qILiarityril~t~nll!rSlllSOV~lor V~~C~ 2 'Ii, ~(] IExpl'!rldedlvieo.'1'o~ b (8) for "Ima~l. ,iflV!!,l"

REPORTS
toruse <II}'!. mass, I~. ;:md 'Chl'!T~ sensors {:J ;]."/)1, The ;j1r~~!!C<!i!onof groplWHC N EMS, extends hJ;)!9:I~d j~~~!11~,:,,;h'l!liJ;~~~Om.:h'O"'S, 'Jilli:;;; !O!:I~I~ tm]~hl~~illlil nu::n]lpr:rurn~ c!m ,Ict as :lII 11;Il1n.~HI\i
"'!Ilppoltilll,g
!;in.K11l1D"l:

o:r1ulomicllt~y

'uh:in ~1lf;Jflhr..'lII~

Itl!. A. G'~j)t.a, G" Chell.. p, ]~~ s:. Tad1llladiir;t<ll P. C, ':kwoo. Il'@nO ~etr: 6,~6'67 (~OO6). :iL2. 10.(;tar ,~0.1", h11p1.Mrntl~!rf!J:!i'S, cadi;si~I'lI'Ii.'I·m'!'imdff 060iIOOOi;lS 62.pd~ 1(2!JQ6.I. :lJ3. 'II!',S<ilO!l~W "~I aI" M'lr;rJff: 43::1" 2'811UOM. :ll4. 5. 'liil1allil~ml:.c<. D.. H, 'I"OtIllg" 1I'lI, 1i'i\1!I'Ii;t1. ",jiJmiIM

NctJmi 39Z; 1M f19~6l, 27. It G. ta'!mm~, It 'it CII.e1amtd.. ",H.IJlit 4;;;ilii,,291 {l{HJ~. 28. ]'. 1', BurgI So. lit Milf1~i!:.Appl. PItyJi, tllt, 8,~. 2,698,

Z6. f4. H. 'Cilelll!D!l, ,M. llRoilkB,

taOOlU"
29. Wi!olha nk (. Iliml.\ill![m O!OO t B!!!llal1llll!}f ~tlp cl'fllh Itl~ III ~pe'jtfOliQCp~: It. Reic_~e!lbi!~h f l1e!p ~hlJJe mer fL' ~~lJil' ~ ~1it.~~h'~~1.1~ ~ill11iiE:5!rp~r1i~~ CI!i'!~a1~ M.!t~~~ and P. lim','!'. 2ll!an;g. It Ilif.. af'ld t::. S~'Ii !fir 1I'!t,fllll dimmlom., Uli~ ~~ was ~~POI1~ ill)! Ihe' US F IIr!r~Ihe' C.e'.n~r .fo!. ~moiO'e :Sijs[e~~!'d th~ ConIeU C.emlfr r~r ~LMiills ~~rcli aJldi bi 111@Nil.':root~cm~ Mr~(b~d F!elwdl (;~~ti~ foolJ!ii:dl ~oo CtflU;:J QflI

s'C'Pw.Jt'~~lgwo disp;lI:ut'C' cm~j~mmr~I~, l

R:e:fe'Jielilces ollll,d Ni!lI~e:5 1 ~,G, C!lai~!t!~Ii!,Scie'!'Ke ~9(1, '1532 (2:0000, 2'., f!:.~. E1iinci, ~1.. I. Roo!lie'~, Iff.." Sci" ,1nl"1IrW'll.16" 061101
~, iii. L KeU,)I'" J'bJl~

Probtem;; ffl.~fJwmfflW¥J" W/filey, ~''I\Ilf'"

!!,!ii, .oiI;::5-l'~1.

til. 4,19N~,

4,
5(,

6,
T,

8, ~,

!i!1Ii!~~a' en S"w.e Onqi!1!~. lIli. A. C fefif~lril~r oL ~ R~: :te1t '17, lS,J'4Ql ~:i10G6}.

f!/ ~illtll' ~1'I~i~ s~nc~" lnr.d1l11; ~gl~mxld.. NLli~'a:l1I. M. Wibo~\ PlJjIS, il'ad'a!' $19, 21 '~2006~. K. S, t.r(Ml~b ('lot, Pmc.~atl.. Armt ,set !USA j[O~S il, '12000,1. K. S, N'(p,I'("'i!ll~e,1 ,,1, NfJMIl: ,4i381 191 ('005~. 'i:. a, ,£h~ng, "I, 'III, i~n. IHI. t, St~lTIIer. r. KAnt N(JI,~ ,~:f!a,,~D[ ~2a.OS~. l 's.8!I~,1I. 'fr. Y~illl, M. Ilmm~ K. Sciolio, F. IL.Md;'lI~'f\" l\I'tllila l,M,S, 2i!1;(l~~, 1l\a,1fliia'l1 and mtl~odi ~r'f!' JM:lila'blt ;;!(!: ~l!pO!llijil1gl

~oo:st

~l!S. 10.Qi!n, G, ll,IMa~I1~r. w. t::. UlI, M. f. "tltit 5:, li:u!',oIf, ~. MIX!!, 1i1tv. 55,. '195 ~'ZOO:2l'. 11.6. IL :5eh!'i~ fl' a'L, Appl. !i'JI~ ~.tfI, :6l!..,~~'~~ (200il). n. l(, X. U" T.tO,mD, y, L W<Iflg.. M" fj.a~hll, App}. PIlys. l.t.tt :63. ~iBl (2iOO~. 1l&. IltL Y,~~mlllra ~r' a!:. J.. ,mlCt;o,!1'l~~i!'t!i', S~: 9, H1
(21JJOOt).

~ri~lt St!wUI~
p!!rfOfncl

<!1rTi;i ~.

Samp~ fahr'Q~i:!11~

1l~1.S.

s. 'lttmnd'g:e,.

t, M, Ii!~m\

]" 11\. IParpiil" !It, B. lleichffiOOd" H. (j_ 'Gfai!1!~i!:!iI, J, JWil, I'flys, 9\1'. 'Ilo!l301l

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20. R .1,5;:i!l!Drii'kQpl. ~. Wililtttfffi, Ii. A. ~MJev'll!i~Dl,I'. IP. Il~ng. D, L I~r •. S~C't 2:60. 1,]6 {X'.l'9~)o, 2lI. M. D, ~iIIi,)'I!:, 0, I'llill,B. 'Cam~ilOI:~ K. C'" Siiliwab, SaM(J! ,30"1. 741 U.oo·4). 22. Ill. aic .t(,QI., J, ~. l'lIlys. 95\. 3f1,'., t,2;1)Oq), 23. 1101. V.lI.ilm. 'p, G.IlIill!ki!!, App£ PlIys,.,fff1: 82. ,iltm ~200.3), 2~. Ill. l llI;i!K~ X. M, H, H'l.!an~ M. t, RoIDie5,App.!. ~y$. ~e,l~ 8l4. ,U6\l! ~2(1(1.4~. 2!i.T. tfutlt'l.l' •. llEE SMOM}. 1 I'll flOlm.

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mokclIll!M kvdl o.r tile surtacc under s[lIIdy.ill Ihus way. 'we can d(;.'~t'lTl1j.nc whether tbe 0.) l.;i~bCtics of !he ORR • S!!lJC:~~I~~!li!\;in.i'ltc,(iv) .u:c ,
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Tille slnw r.a~~ Ilif t~e OiXlfgefl f~d'U[l~Cm ~eal.tio:n (O,RR~ in llil,~pCllym~f ,~~e~tlrolj!'te membr~nn,e :~ulel.oeU (IP(M~() iis ~h:e rll!i!in lillllitltiol'lrm diU tornoli¥~ a ppll~!:ati(lu" We de l1iIoflS~r[lted thatltle li?l:;!NI](n]) :~urra(e is lO",r,ol.dl rn.n,etldi'lll~ ro.r~lhe ORR. tlilan the ((lrIr,esp,on{Hng P't.(lll) :!im~aHla nd '90,.;f.ol.dl mOI1e!adiv,e! flail! the CUlrn!f1t. s:~le·(I[4hh1utPtlC ,cat,a;ly!;1Ji [,or PEMIFC.The Pt"Ni(mil.1) suriaoo has dirl UliUJ)SILli1!Il @L@(~rllmii!::s!rlLld!L!r'e W·bandi [I!!nl@rr posiition) and tm~ln9@m@nit of lSlIlrfaoe ,a~JJlllrlS;ill1 the 1ilI,e"u",5urraml rregion,. Ulld~r olllHatim,Si mlildiUol'Is uelevanlliOrllJll!!l ![:e~!s. H5I1ear·nnfiilc:e laye, 'E!':dil~brii~s lili~llly s,~mc:tull'ied 'GOmlpos1.tional. o~dll:e'l:i'on in Itl,eot~tennmt ,11 :end thkod l.a.yer£, which are Pt~ridl. aad 111the secend a t.Omlc, laye.r, whkh is Nii'ri[h, "!ltEl' ~wak il'ltewtdion betw@el1' thePt :~urraC:@ atemsi1tll1d Mlilul:adive O:tY9@'~tli!dl~lJ@de$,il1ill:~S<es. tii!!! number (lir aai!,llE!' si~@s lor
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~nc kin~.."ic Ilin~~mtiQI1:-i tht;.' o;>;ygcn 1.'!..JlIcl,ion 1'!.:"{I(~ti,Qll (.oRR), whiC']l ~i~I,VC kdl to t1U:'CI: .~ulldall1.cnlll1. I1mbk;:ll'i1!> (I 5)" Furst, t~c ,td'l.~DfUIDifll ovcrpO'lCmil1II lOr lhe Ol~R "6 nn all practitJI o~ro.t'i[.g ~urrellt dCI\,.iaic:; redtKO::s dlC lhem~lIl cm&:J.cm::y wen b.c:ll1IwmiJ C
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sarface chcmi;".~~on :ldngk"ct),sml urlhres, ai, ck.-mrochcmkall iflterr;1ces IS rc~mill,leJly mlC':lI.'I'l Ol..... These .agueous imc.faces a1\C more comed, p'lex, in. lhm,uh!.:), n~iiLrH~ cO:n!afn oolvcn[ ,<lind CIOC~I,'D!1 ie/ionic cll<!!rgc. ,mC! lC'xp!l:rimcnt>llly )~l al 0.7 V I. ver.ms lile Ihcorctical thcrm,'lj I 'C'fl'iCICJl" C)' (Of ~3% ,n l~~crc\'~~!'l>ib~ pO!C'I:ni:d for the is vcry dl;".Hcngi~.l~b:\bl !\dn!r.lc~'.lht[,J M ~I.S~ in ORR U.13 'VI~. Si.;;ormd.llwppw);;,il1ll<Lt(;ly liv~'· si'~WI )\u:dk"l,;'-'.'lIi,;n~ili\'l; I]'!t,;ilhnd" to clmrO,l~~leriz(.; ,
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lUJ). [lill~,~.Ijy,IJ1Cdi"whltIro ,Jfidfor lo~ or fl. Strr,l:'aoc <lJiCa.ill d'ae c~,lhoo" I1nl~ be gl'\.~l~yrcdLI.lICcd. llR'N:: lifiiilj.iai~m,... ~u,ul\Jbe cilimiumlh::diJ:>~Jljlc c3n~odcca!lllly~b>,~d,dl i:l:nl orrlcr Oft;ll.~.glJj,illtie ill~'U,'CflSC in ihe sped ric <lei u\'ily over Illai. or~talc· (I r:dlc-mt PllC oe<lilllllys1:s. com be deve~oped ~101. In IJac h!IJpelh>JI1 :1 cO:Il1hhmiioo ordilTcrelJltnetill~$ woulkl have im~~ro\!oo C'I.,.~yl~C aClivily <Inc!~""a" bili!)' m1mivc ·ro .J!05C or<Jp.1.nrc .~m:·[.;:!I, Ihe ORR hOlt'! kt:u zo;~ILlciLiodi !Il!~ ~Ulni;~.·aU5 hi· Of nndliI1I11~· [!I~:I e alloy;:..; It 6. ,Ji}. 1! I '}~. u Th~ sl!ll:d:iJ;~ ~mi,lCIt;d
to

JlOlI:nli.u[!..indlll~1l,'d d~,Ul!lJ!,~nlll~ ~turl~LI;,"i;fJ'!uPij cJ:iir;s ...nd, ro"o:lcthilly. We Imv{; tll.'ilCd 11(;{illiilbjn;'llion~ of t'X :oil u find in. Si,illl :;urfa.c0~~L~ilivc ~~rohC$Ilnd dcnsily fUJ1Clio:rml dtco.iiY (1):f'J"mIcLlI~.tioliil\ to stLid}.'aho;;:ORR

()U P~lNithlt'll' sillgle-cryskll sum1.~cs, id:ellliJy which SiiJrfaceproperti(;$ gOV~iiil th(:\'~nim,i:oni> ill ~rea'Cii'Vily of PIN iC'alnlyslL,>, altd dC:1!C'mminc 11.0\\,' smr.1oc :siru.cllurcs. ~~t"r,~LCC scgl<:gation. and il1.tCl'rt);;:ialilk bondillg arrC(;t~he ORR 'kiltcll);:$. \VC]~dmr.llCIer.~d PINi s~l~glb-crYl'i~"II, d~l:rooc $\lflaces ~"'~ fmm."'.d anti ch!!!l3C~rizccl with UHV gJtc~ods ~QrSl.uf~ pJ)CfI'Ir.;(~i(lo~l<LIU;1slI!u'nhocc my.~i~. llwiiic
sW!rdhtt~,'!\'(:n:; lit'll'i,alfid

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dj~ :>1;11)1 uf~hc lily

HIIV.prep>urtxl

REPORTS
Previous st(!dies or.PtNi ::;iog~C"~.ry~'l,d surfaces in U~ IV h;K!~!'!d!c"'tt,.>dUw:ttlh~~ surtace enrichment j]" p~ if! th~ f~r$!I~yqis.~~I!}'~"!t~:r1lin~x! by itl'i Clbt!~n~ il~ rt'l,t!1 i!~lC_ !;kp'\;1:im~ inu&u::- Itc'X~ two 10 l.h~'4; ~ltonili.(:' l"lY(i'ffi, lh~ oc?;!ull s uf d]j;pK1Pllro~!t1I~ mId Ch~LLI":!IC" :>.0 1.lmi.tlla: OO.ll~ciltnnioJlpro.r.k: ~x;cillflk"'" 1IImumi [he bulk \1!llIC {14. }5). 'We ,1cH1.oa.~lmt(ld ~11!Lt ~c-;ri"'-<!tL\(lI~Gr'~3Nilhk/~ a.II[ll}'surlh'C'~ ilil WIV l.!!.rcsumrmui7cCd ill Fig- I.Th~ NlLrJ:';IiI;:C'i5Cn5ili'l~ sr;.'g~!il>"!,t!;{}:!1-drjv""n nt.;'M~...~rl1!~i;;c cO~llpo!;lilif.m",1 ~~c:.l1l1i,t;J ~¢:!\ IJial w~1'¢ ~I~~\j] il~t 1~Ilf:!~dlow -cltergy ~hHngc.:"'~""lIiITtUn d i!':l.im;:'-v~,ckK"I:rn~1t::'Ijlro'f~n:il;l'i ~ i orlhie rINt(hkl):'IlIoJ's.. C'!{-Clmll di!rt.ll::l:iD!~ ~LEED). Augt"r ek'C:tl'On. :;peeThe ::'lJrftiee clC'el.rol1ie :;,Ei'!LlC'rui\~,Wcre oil.. ~rOl>OOjJY lAES~r,{]j\V.t.'ilK"J.~ lio:~!>ClIntrill1g (LEISh taillcd rroi~~ il'LCk.,ll"O!liitd..oo.rll!..'Cled UPS~P::(}. 1I.h~ ~.ittl ::,Yiteh roi~"Oicl-b!i.',ed It~!:l11l-rcsoltitiu~1 ultraua (J6). }\$. :>lIItmmri:t:ed in F~ Ie. ~ht;: d~band vielet 1)&~o.!QC.~niS.'>ion SlloCClWSOOjl)l {Uf)S} (sec dcnsill.y or ~~.nes (DOS) is stI:UCIUI"Cc...:K"11Si(hi'C', ~nc~hoo.~ ~hc ilfl ~'iilllfi'porlin~ol~'lrflCn1Jll.l.CI'h1n. E.1dl of lhc;;,.'C mel hods: has ceria ill IJU,h'atlt,!i,g.C:':;, oubd aucllilcp'rudlion ofthe~H~aJld cemer shitlL'"limm. 2.70 CV(l1l f'~iNillIO) 10 J, ~OcV(U~P~Ni{ Ill, ~h~y yield COnlpkll1cntrn'ylI11olr.ln:M'i(Ul. The SUT~ ~hcc!'>Y!ll!llcny 'l:'Ib!01i~1I.'tI rom LEI!]) "ll'I~.)'~i:;. U) 3.14 cV Gin ,rl~NifBm). FtI!F~h!'cIr.IHOre, hc f DOS (I f Ill!: ,~~Uo)' ~!UfraCCli ~~q'uih'; d if1i.'f>f..'nIfwmn _~how$ Ihu!.. ~""htl'~""~ P~,'!Nit I ~.I) i':l!lrtr;"'LC¢~N1]1(;: th(IJ\ (lfl;(MR'spollding p UI1l.' p[ sillgJ:e CI)lll;l,I~f;;; ~ht[r il ibilS: ~I,l~ I) p~IUCI!'11 (F.i~ .. ~Dr) (i.c .• lIH~~ [I r Ih~ x bulkII:'FI1~~nmim~), the atomic,lIly IC$)1 d'cn$'~ is, on the l un. 11.00 I, and (~.I ~.l' aUoy ~lIrrhccs" the d~bmld OL;'~~liC'I- dl)wl~hifwd by ilOOlU! O.l{~, is Il~~Nl(I00 ~'SUIJ"(l~ !\hrl\,\~a (I x 5ll'!1.\oonS1:nOC:liolll p.:.n~!.I.nl (!h~ .'ii~>C;;IHed"hex'" p]l."Lw) in botlr! mh~ 0.14., and 0_34 (,l,\/. Il.."SIP'\;OClUvdy. An iJll1pol1!!UI yiJil ~O~..~J i1lld 10 1- ~ 1~ i",'Cl'ion!:s (Fig. ] E). AI!lll.yto>~ IU~(lI)'lIIllt;x~'I('Jfiiil~~c illl :;;lIIIIJ3{l~! i t;]tel~hl"lInIhc .1Pt:iNi{~.W) LEI:!.!) dam (rig. n~ l1.dic::m.:,.,tt!~1 is'l:!)' on :II.U()Iyf: i& u]]tk:~n;l~,lrld~.ug hrYl.'!.'chcnm i50rp.. ~i ~llis !>Llrf2ICC mil)1 t~1\:hibil. r~mi}"lurc [I ff',t ~ x ~I' :mii tien r:::I11~n];ie5 comJ![lcc with tlte iL\'cr2lgc em:rg_V oujhe surface 3tQlinS to \\' ~'I chjhe i 't I pcri:ooidtie>,lItc 13~1crlJcing, kflo\\ n. as I.he: or I,he {1~S~3h! ildSorhlL1c b~n.ds (i,c.• Ill!::' I~~illl clTed.l. Todo~his, 't I x 2) n~issi'rD,g,;mw S[mclllll! (22t The co:mpos ilio~1 of the (n.l!Cl1ll.G.'l;l ,}Ilomk: wc comp~rod IheUl!::sul~ on ihe (1111 ~Irrno::s wieh (he same compositiDil ::nnl alTrulgcmCnl of laycf was o:bl:lI.i~ncdwitl'l LEIS, a,'>pre\ ioull.l.li sur~1;ccatoms ~1!i11\vidl ill di !lfcrceol t.i~b'~1U1 center ,~hOW~l 101" Pl:JNipo~}'cay)1l:lma~ iilllQyll. 15. 2'.n\ IH~ili{m, A.~lhrlugh~\'c ,u'e l:Iigh~ig~ni~!ghow aii.\;'1' n li~'I<Ii ,m:nn~.,,~I. !\u rlllcc O'l'!Oinlic bll'cr of.flil the three .P•.l:N:i,V~kl') crysta'is is pure fl ~Fig. lB.} [i.e',., the dirrcrcn(ic between 111C' ckcrronic surface ! suuciuecs (I r P~(] I ~ );me Pi)! Nil( Il·~ ~can a WOCl .1! I:~ Ibrlllllhc so-called PH,lkin S!.1'l.I!ClltroCS ( f 7~,:;'0)1,
Mlrr<l"';:c WiJ!i'l,deiennincd

wilh a C'CI'JIlbiIKll!ti()Jl Qf ~i,lt~ !lillrt~gL}'~Ill;;;iti\,'c ~ro~)1 \)j,~lh ck"A;In:l!t;hCl]l~ il;".J!~~n~hw;l>1 ~ t~~\1~;~~.i\'.rty !l~kLl_irnl.'ihi[1~ 1,':0\11(,1, ~~ in

!he <lJdoorptiQn <If f1«~eCIti!WI' speeies md lile ki~clic)1 of the ORR. Ihe I~ndi!lg:~ lIre gcrl\.'J:~ly O!IppJ~c<LlJ~cl(]olh~r ,~~n~Ii.:~_.."~1<lll'oy ~~~rr~~fL o
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iC'.;.II~ (,mvironml;lnl. Surface ;o;,-!~)':;'~~~W<Jiug r.;X:Sl ""lOll!> lIS';;,o, IQ ch~r4~lcriz¢ 'b{ljt~ the pLJ'lr.;;llliOiT mli3:lc of j,1~l.b,.i.~ily !IS \1{t.:'.i11 ~.. lille J;le'1r~iJ!lIfm=c MIl.'iposilioJ'i oCllle allor ki l>ilLi. Oiily restillli,~ ~b:r PcJN i( III i fn't': ~~~11Irj._'tI because the.), ~j!l1),'iidc Ihe most useful ~1]rormHl:ion !lOOul. dl!;!pol.i::mili~' iildu.cM changes ill tine :;unlbcc s.tructurcalTld ~VCg..'ltioll~romc:n brier Jcscrirtiml ef re:;ull:iJ ~br (I Hll,mui l!relO) c.an 1'lCfnund in {lh The

fC'nnill<LtiCin ofdlC PtJNt~ Ill) laUOCi:::11 the smfacc in 11.'I'nl;;"'; ~k'HI~I~t1Il cmll~itir'knm!d of :ollurthcc
rol~x.nio~ w,~ di.~!..·!T!lin~o,cl by n.11.'O,lI,,"1If(l!V1~'I!1~ ,:;t!f!d ~m~Lly~i of dt~ ciI)'i>r.l UlUIt'Uliol1 rodll (CrR,» ... .
I(]:/f).

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or

ft3Ni( ~I~ II~~"d'lC' filCiC'~I~I'Cd albic I(roc) l,mi!~'C\\'ilh :mnd'om m."Clil p;uion ,01" si~c;" by PIOl.!1d lNi.,u!l~~~hi,~ !!li.vl;.'l' ~iw SX.S I;ll~io!..~ 1)0 ~~i!l!ibr ~l:I nll!!1 (lbl~lil1J;_\d 1mI'll! [I mL'!llocl)!1.\n.ilnil1~· fee hnticc'.
Sc.rt,~ili\l.i~y 10 ~UQllIIit'!3ycr tlOllilpi:l$ilillll

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k'C~uakljjCS in 'l.vhach lite illeidem x-my eitcrg~ is I.. etl 1.0 ~Il 3111mi:e m:fui.ortJlion edge' of ~l(; .l1nlam tcri~d < lQ}. lj~lII's. .titUIl1rSXs,,~'\.,c o'bl[~ned in'lilrIHaI.lon abm.lll the lXl.sitilcms ,mld c{)ml~i'i itioas, 11)0111n~hc :surli!oc and :'!ti!bsmf1!;c(.' IHJ!;:f:i,. The i ("rR ;]:n~I}':'i)1 )li~{l~\,!,ll>l!.. ~1 0.05 V. l!hl:' lil1';l lill}o\t;.r l is compescd cll.irciy or PI, ~md. \\,llCutaSlhc socond .uornic ~ii1ycr is N i-l.'iCJ~ (52% (I r N~ ,>!is 1IJ0lll:-

Wiig. 1. SUlrif,Ke dllar<l!:teriz:ati:onl of the IP~ Ms<li'i~1f crystals ~i'iU &-IV' (!!i'id '@1t<e(!trocn@rrnic.al 1~111;1'~'QJ1ffi1~nlS. !(A~ AES s~(iI1l"a. ~B) lEiS5{Pect~a. ElEo. wll@r~ E is @liHl'rg'l D'f ~!!:.l!U.E!f@ld
l@;te'CtrmnS<,ilInd

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~he ~ndderlli®n be<ilm. ~O UPS, :Spe~tril. ~D lQ If' I.iEED pat:l:enu w'it~ ,oone5pond~ing Ibi3iU. IiftOOeil'5. {ill to U C~dk V<lltamlrn~rv (CV) if'! 1:1004 {(U. M) o1!iS 'OOffilj)a:fl~ to the voUam,~ im eli!y o~ Ute umre'Sp®ndirn mI fit '511ilgle, (Iry~.I. {gray Ci!ifV~~, IilHIE.
r@lOI'@rsil)~!!IrIljCbeg,(!ifl @lmrod!!.

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REPORTS
lXl!~,1r.'\I~o 2:5% orN iinjhe lmlk r~~hc Ih ir~ ~<!~'Cl" i~ this surlace QV\,'J: corre::;.ponclillj,ll w O1.g<1lnPhmr~'c'hcd l~')'%) (fig, 2A~. J!.winI,l which \\';:18. "d('!il~9111J~ly nlhmcd ~1I,.~~llil].l;;d p,1.!fC

P~('~I), I

~~y p!U.~m1~~d

d!~ ll1;l'~f"c..urC..~ ~~I·n,J(;'U,.m;<110_0,$ V, . ~h~poti;llni':'L1 qrcloJ wh:ih~the' ~<ItiCf1;c-dx-my \W~,

i]~h:,~~iIL)' Wflci IUK-'::Ic.,;urt~d II [1~R IliD~iiont~mi.is at r; to

cy~I.~llf,lr!1 ~w ~k",i,g}fl"'t(i'\cl, ~h;I~~i,,~r.!n~c. TIl!; .n;:lmiC!lh'" b~I\\'I;;I,~I~ ~h~~lrf:J¢c ·~k~tmnil;: r:rupt:il'i:u(:'~ ;lnd i.l.1l~rok:l1~l~d.-d¢r;:I~dl'lll :ii~.IIfm."C
CO\'(l"~'"

!he rm,CliQ:rml covern~s by Uy~1lnd O~~!(H! (~H~!md (~'o:I-!:,w~\\'I;lN' dl".m!lU~c...IJy reduced ~ ,0% r<~!{lii\'~!.o I), \vhid~i~ i,~ ;I~'I(,.'_ M1J,."~I~wiIJ] th~~!U~c down~~l:~n ({tJ4 (N) nfth~

r!u~

by n~ltWlrbingspecies (Ii~c .tdSOUplion i~(I"uli..:m;_. inFi_g, 20 Wl,"rC I;;"'t..hli~hl;',d. bjl (;01111. rannglll!; II;xpcrurm;nla.!ly dCICIlI1iJijll,-.dpusiliOl'i of tlt~ d~bi:JlultK'nUcr:;.W ~I(! iT·d.Ni,O'I'i~1.1oovcn:tg,~";S,of 3t1~'.Lb."'tI ~~)'dJn!:jC1l1(II.~ e: = ~lll'P"l' w~~CI'C'III~ -I b.L.1C:flwi:;"; the e:~a~l,Jg.cs Ii11, l.hiC' ll:.-r.iY ~cauerill.!:! :Ii iguuil an: lill~y reversible, II nd We r.~ thlil. the id rer~i::> to the uOKk'TfXiI.C!I'Ili~.ny dcPosit.,;;d h.ydro. 0;05 decrease ~u in~nsj,l,y arposhi \'c pot~lui [II is COl~~ £Ji:n) bC1}~~J~e~1 < {; <" [I,o.l V. \'Ihere /; ls tI.~c
:\'C'm;ili\

:l'lI!r1aoc !C~ni\alion,IJmd i:l\ufacc' rough.~ !1~" I~r-~g" Al'i,'/c lind II~aoouh t~~c 2 .rhN~( 1i l) surIl1I;(': ~ !11lI~1~1~ as wd ~;J~ UIC 5cgrt,;~1uli,tm fi ~c 3Fe tloJ'itp!L'i.C~'y i:;llIhlc 0\ er lliri:. po[e-.ifIl.i[~1 ~'ttifi~JL~

cl-h.mK! C~II.IL~tp~it:ion an dll': :P'i~kun :iitrnduuc" As ~hOWll iua Fig. l , H lIud l, 5~mil,~r d~lIigr:;'.s
()<;'~I~I~'T~d

rm~

ra'r

Ihe

other

.~ WI)

![oiiln.gJ~~·~II}'~~;JT

an. UllWiliOO reillixation ~OOnlr.li:Clion) atmnk laye;' nil fu;. ~\CSlLh ~s (01)11nlloo by :;Ii mi I'.u ralCa~il!reilll.cnt~ al 01 incr wei rroc:l~ "mice IK)~i.ionsJ. The ~'illHlJrac!ionor p~ surlile~ O"IlOll!"", i~ind~Oi.;'d by I:I!!L,! .. dt'(l!l'P~ ii.1!~ or t1X)'!ilt'J10Jn.(iti i.p<;ei_';i;. Whith ~t'i d"'h,;runil1!..~ by mhL,! N~"indLJC'Cd modvfk,mion Of1lilc P~~kinle~cc~rm~iJc '''~L'U(;'t~lro.A I"li.r~'"{:l con,~I~~"I1C~ nr COI1!mC~Ll(nl
\\I~l.h $llYrmCC

~~i. ..!Cilll of the

apf)licil m Il!il

j)Ol.enliall. ml111~)ildroill.yl '''>fiOcics (21 i10 .... .1i,.:l<O~ I c-, \,,~llcre 0:1 10!1I! dlC adsonloCd is

hyd.ro",),1 ~:OI!Y'C'~l :IOOW IO.fi V (Fig, 20. {' a reli!11 in,<;pt.'!Clion or Ill'" \'OJ~ml'!)(lgl<!Xn.<: i~~ Fig. 2B ~~\I~O!J-cd~hmCUI .Ptll i! )!-!J;in. \l hid! emiSi~ uf
tht' 1\:wn~ Cill!llrr..Qt.; dealti.ily t.lfVl 0,I1I,t.ln1!'l ;'I~ P~( III ~ 01.

tlie PI.( 100 N-klri, fJ110pI W~b'" It.\l Wlp(id 'by ::::15"1 .ebniv.c 1.'1:1'.PU 1001. ~:rid 8otl.1dwa,'l .. 1'Cdi.Jt\."'i! jj,~1 ~2 5'%.. Ol1P1tJNiH iO~, Sltl1111 bllil d~t'.II)' ,di:>cC"n~,ub"Jc J()!i'C<lSCS in EI!.!I"f'II (~m'Y~ l ru]a tl~1! .rraC~kii1ill, C(i\"Cffi.~ by OU{(I. (Gl01,u) "~O'%l'\vcllC' ol~5:c,,:oo. re!a~jlvc·W ~'t~ l. 101, To dcl1C1'ihc th~c CWCCIS q lIIan~ln:ati.. C'~I.WC I'iClfol"lll.ctl DFI c<!'Ic'L!la~iol1~.u.."~Il£ilif(!Lociopol,m!i,~I(ol ,I l~~.ml;!'-\v<'!li'cb;!8i~i-lct (H1 ~h!.'!ndf1OJ1}~ ~ij!d ion
SlU~"ritl;:~< .QUI

d~:lilm,uic neg<uhe shill (:;:0.15 v lin U "iiKI rbrlm!'!iol1 m~dpo..... i'~!,.~ (=::j)J V)~~! 01 !/lid :s~l~n of ~h~ ~~p:mo:s.l]~)'\,lr of Pm~$killm!hL,': I)Om;I,11!ni,~b ronr~ln'[inn !J0CIi1 mId roJ~lli\'C'~o P~{!J 11.1 ~~..gre!')· nliC~l[ ~\,jlliJthe (llfl";~~ of ;Jd::wrIP·lion. 01'1 P'1'lNi~ I ~I I, ~IigJ1l~[ Illu UI 0,8 V is lht~ il]~rC':ibed :>Lub~.~ of s il)' Fiig. 2. In~~tLJ (mradeF~

nltl(lel~d f'~~~ ~~lItf~~':J;. .. I) ~ wrt~l(;j~'t\ond,~tL11t! ie: bw,!;:!'1:' contilini~~g, 0 or $0% Ni <limns. ln acid ~Iw ion,",. f]ol i;[j!J \\!(D1!I!1r:!w-.ca~:;,t \\'i!hl I+",v~d Ibm1 llio CHl the '~'O!)w~Y;"'t "Wlr~bC(·. dmu~ U!fIIhe ~ He 11,i~f('J1iiuh~ }xlItenti>dAU" oruw.,~Po\'CfJJ.1I(;'~irln un Ptlll i Itrcs;I,.I~~.ingWH:iI Siubl:ayer N~ .~[OJtlsi!s. f

oi O~! ;lilt!

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izal.iicm ill IIU!' PI1N«1J:1) ml HCl(\ 00,.1 M) <lil333 It (4)1 SlS d\ata
5t1rfaae "md (A') mlllc:entratijmll

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AU = r CrJd! (0,1 ~.l'i't

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tamfl'H!~ry ~lJt{jJ~lIled Iro'!1il

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(C)

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and £..;,1.,(1 [~Ol!!b"'~ the arl!;.o~)li)cYdl Cltcf'gues or arc 01] and ~liO Of] P~! I ~I }wilh Of \,\fill~o1.!~subliltyoc.r ~i .noms, rc....f!OC~iv,cly. O!no F is~hC' F3.mday <;onSIO'lnl, The DFT C<Lh::;lUb!iomll\how,~ pr~ilivc
s!]i,~~ {1( 6U = ,ell {I V when ~hc sllbltlycu' hil~ 50% Ni lfi.lOlllS, The '.:xp'C~:i,I1i1IC:I:n .nuld Iht"oD)llhwl~ IJ'Cllch nui c;;((l~lllcnl tLitti 'CJ;mllttul.mi vcngffiC."11lC.I1t in

Here. E~(OH

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Ie<rage (dt(~lat~ ironn ell"' dk voLt"llI1lrno~Jr:arn5 oi' P~Nli(11'iU (fled I!:l!I we) ~rid Pl(n,l) (bAil!e (lLW@i: p,o'l.alril~tiol'i (l,llr'l!les abo

Ihis ease :ill1d clc{'Irly establish fiU ..::icNmllaC effect of siLd)sUlifncc: Ni on the Pl-O! I.ehemical
lxmdn~lg.

l~in@(j hom rQt~~Tlg ~irlg

d~. ,eI!oc:trode (RlFlIDl) III'l!l1O:IF

5lI~emeints. (':"1b:ild!!.5IJ1fa~, i~Ql{e'faJg~ by i'!,,;jsorbed
~qJ

c.> ~IPt(11~~'-\1
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The ORR is a multielecuon rea.cll.on (~bOJ

2~ 1+ ~ 1£,clcm~fmU)'

(ies,
lla'l'S

O\(.~ernatoed :;pe(m) i(Jiree.fl $late re-

te inydr~,iE!n peroxfuJ'e'

pr:m:llji('.ticlIn 'in de-.s:iDnatBcl ~()tent~~l r~ion .aJF1d00 ORR wrr~mltS m!!tl!sllf!!d l()Inl PillMUH} (r,m ()lrJ¥eJ', paU.:m {bi.LJ~(ll!f\I\~, (IF!d po'lycrr.ofilillilii e P~~9r<lIY (lIi'\i'e) :5ud.a(oeS. The <I rr!IJ~v,s aln~ sh,om ng '~'h~' p osiHV!i1' lj)otenliaL '~hiirt lJ)f 100 ImV in el.eii:uod'e<

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I

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leliill~diat!;:t!, 0'. 3m, We I)m:poso(] lite ORR c,m be ,e..'Ij,Il\j'~~ .fl;.~

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tJ I) I ; c01i~

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'l\~enJ ORR po~81rizat~mll

i'liI€i;ls~red l()Inl Ptpoly (I fIId Pl3 Ni{U, lll) $uff~.(_m. t II, and IIIII~epr€~!lll Ip~oo~tl(!1 ~egiQIIiI re~pellil~Y·

i
0.2
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CI1c-rgy of adoorp~ rolll or rea,d,iv,e hlicn11£:(l'i.lles, ou~dTis Wll1J~em~lII~, n.he ~~vatto:n orEq.l. it ~ is ~l~med Ih'lIt {i)i ~lC OR~l:;iIkes ~dacc 0111 dc>c~ Imd~"SII'liLl~ ,Ill; mooi:~cd by OUIl!!. ;miG~I5,~tc" ;;u~dI( iil lhc J(t."iU:Ii:\I'C i~~i;,"P'l1l>Cdi;,!1~'{02 ~"lid ~h~ I
~ iid ~or!)Gcl
~o ~!JwC'CI'I'cr:agc

tlrl~ rtrt'!UlI ~Uirfact: tl[JVCit.J~gc by :1(£:>orbed! SllU'li:3Int ~JCci(".~ [hydroxyl ii.ll'id ilnicms: ~.or ,,;;1oi:fIImplc, OU~dl {0.0l~l'fllad $pccjJica~J,y ad~oi~JC<lanions (eA!!dN,~ A,GD~ is I h~ CH'bbs
Illlioll1, 'f:')Dd ~s

,(lIf 1~1.a1Kl1 "n:J50rp~il:lIi11d!l!:!ior"ptien pro;Qesses, do~b'le--I.i!yer

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REPORTS
:B~<;lOO(In these
a$~\IUIl~)lion:ii, !Ill:!

kinetics

(}l:fCclttc~g(ln ;:I~ ,o';;:!1o,.'l1lill ned ~'Y ~!J(l~umhcr of ih:,¥ p~ ~i~g;;Jwl~~<IM~ t~!" I1:1w .. g!\orp,~~ nf'~ {the ~ '800 rcrsn il~ Eq~4)' <Illdhy~hl;,i!l.{;;;!i~ clH~ ~Jld!l~flttit~1l iuit::m~(~di;li\]", ~Ilu::tl..fi::.:! i~'I1t~ ~n Eq"2} 01] l1lCtll~H1~Cl!~ preeovercd by m!aj}- ~ 1I~ ~hi~ ~L'Ct~O!1 p1l1Ih\'\my and r.11"'L)!xp~_"~i{lll fil:N~ ~tl ~m.d.>'7.c [he ·cnt,"U\l of d.L'l;;lumic IlmpL;rtks
o!~[hc kh~('1it".'i. rthe {)RR. on Pl3Ni('~ 1~.)mKi o f)llt ~,~ I.) !~I'bL!.lh":il, by e;QI,~pafjill!!; aeu ...hies utI

or or ~Iw
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:;;lt~~'C' 1{'IIJl'"'t

enhanced by lacter ofl U on .PI{ III ~1>,kiln '(.'1;:1en P1i(~..11). 'G~vc!1llhm'~r.lgncl~d P1

;} 1.0 ~0 tijIH~{'ith~
!if\:

S\lr!'~.c~-:ii h\IJ\'~
O:ltfJly ...~" 111'Iat

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(E'::> 0_8 V),nh~ (ilt'ii~If\'I,x1~.T.(!I.i}fuk ;;lcti\ ill' for th~ ORR Olfl IPijNi( ~ I ~ I is tile hug.h!..':S:[ 111m~1:.1S t"r ~" been ooifiC'tlrod O~~cathode ,c;u~. ]yst;.;;, ineh,lId il1g the . Pl1NilHMH ,md fl~Niq ~.I0) ~\lfra'Cts~
Sy lic'!tY b~UWc¢li ~\Ip1l;Lel; g,l-.o,n1~rry ,~nd ~I.IF· I~LCC clb:::ltoni:c ~11I'iLCtur.:: Cot lIlt" ORR i.~ illhl.['.1.k:d

~iI(l~p~ ,~i,{lll~~ th:'111 the tiili.~c.o.f:.th,;j,.""''lrl PilC

cU[['i!.'nIi1:y

o:r(f1.1I: f n111:gU1iIUd~l. S:tn.tcWI~ ~{l;,"il.iv~l.y or~hc o Ilm:r,;,lnlll:rn ORR ou the r~kl~:'i.'-ind~.x s.iH!gIO-C[}'N"'~ ~~lr~'i,(;I.~, '''oIW!jll:, W~Dilc ,1~i50 IlIq,lijr!!air!ing!hc in ~rr:~ilo:riC'HCid h,i:' been w~n 1,,';S;t:ilhli~h~~ ,~3,. '11O\\\:'I" di;~I'Nil)} I \1\l/(:n1.'), \'!HH,.dd l':li;,.~I.~I~~~gil1{:..;r.inlg in at n,.os V Olncl ~mHg~ng ~li::' clecaodc p;o~c~~~i111 "\'i~h ,tc'!i\'il~~,'i:~alcr>C'"s.~I~g theo:rdcdP[~ 100)« of P~.3Ni( H Il-sk~n iike ~1'lnoCal.aI)l~ls., p08iti\l~!y, \\'d·dcri!1~ddiflfuion··!im.~ling 'CUD"" Pit ~.II,~.:::1'1"( ~ lOl, ~Fi.\}3), These dill""'I~!l£e... have IlIlill,~'er,Iiln:I)~§; ,1I.liId N1(lte<£, rcnl~~~[om 0.2 to 0. 7 V urc fo~kmrQicl, by mij:'(1.,,'d bl.il.·ll,uHribll~OO 1'0 tliq~ ~1!r1(l!!i;:U~~fl,'liti\'~ OJd'iO'lI)' 1. S, GDnu~~d. T. A. l;jW(]dljn~ki. ~rnMl"rln(fS,m hem nf ()Ol !.::id em ~~~t/Jkll~~n!clI li(\ il'lhib~!ing {:;,i~c kincaic-diillih ...itm control ~guoll b~~aw~"!Cn 0':'1< afftl~!remiw1S(W!.e flfNi fn9~~I1~. R, (, ~. H~ lieri5(l\.er, I[)_!IIL 1':OI:b.. (_ \C'l" TOO~:5!. Edil. ~lj';\i~ey.1;[( Ii, IJroclk~ Ilgj cffoct CIt] O2 ild;;on-plion. kllhc pmlt..~ui~J E < U]I V. Fmthl'1I' iTI:S:P('CI.io.1I1 orlrig. 2:D [,c\'C8iI:> Wi)inn~~ G@r~my. i99~~, \'at 5:. " thaI d'Ii::'(}Rll.koit<:.t1c L~acrelc;rJwd {llll1iiePll'~ ~.~). [legiol'i orO! I.[Lt:istl.tp1.h:m, [he' Slruclure sc~l&i[i it}! 2. Iii, IBe'511f1lmher ~t i!lt.. ScieRu' 2:79, 1913 (199,&'1. the ~llNi;lhA}I=~in surfoOL"S lIDS I.hl! same ~ki r.cimi VI! 1.0 r~ll'~1 ealls.1 Ilg the poshiw shi'fl n ~ )\ s, N, il1. Mil Ii'. ~, IRQI.:!, SrJtj;, S6,ifep.4!i. 117't200:lli. of I(JO mV in lite' htl~r.\Vfi"'l! potcn~iml Gi"''CH thm. oilignnl. ~. W. lIie:lslitJl, A U!l!l!l1. !t !l<, 6-aote~ge,r; ~.fmdbaf)~; ff/ f4m eet&: MMtl~~ul~ r~cboolorrn aM Appli(orion$ ~:nerder to fC\ c.Jllilc role of Ihc clcclroUlic ,eOHil.;lii'>ml~ll".mled on.lh C' PI,-skill stt\lctU!re.~hC' (WII.~ Wf!~ )1!I~b:. lit. ed .. 1, 2003). key rilrJ~]lC'~cr thatdcllcrnnim'S the ~uuJ~!.mlly SIJ'!lJC111I'C oato 0 R R k inC'1lc, \'IIC cOfnpJred e lee5. ]. 1'0:.:, ~ll1rs~m. , H, ~~ri1;tim1~cn, .~U>JIl~ 3:11,2', .13 22 ~2C!061. c c;'I,~ytic &:1.ivi~}1 ofPI~ Ni~ ~.I)is lilt, low enver~. nodes wi~h~hesame Sill rtace mOfl]lmlo,g}'. The 6. A. JI. l\P,plfiby, {orll'i, Rfv. 4,. 2,;U (i'97C} , t, 1'0\1, lit Tar~'II, Jl ;s.,,;!})D~'!J~ ~ 'l'1~a9:e~. !iI ~ mo~tt rmlmll!lC~\d c'f~oC't was o'b\.;cr\'w on ~l!c a_\lC b}i OU.~ [ie., dlC' {.~ '0,0!""" ~ WJmin!I~C' 'o.th4m~r..e' li'ft;lll\!e in E1t(ff~miHi'J',], 0.. M. kine •Jc cqaation (0:1' tile ORR]" A.dditimml OOIl.- ( ] l I ) su rfaces .. Fo:r~hc same NI >< I I .f:u:rangc8adlr'~, a E. 'GQfflOO,!" E. '{iNig:ll~.S. U. ~. ~H\3m,. I1nm1l11011l h.at n~c ~h.c~iomdoovcu~;gC'S 'by Sj}cclau mcm ufjhc l.opn1l0:i!'tlayer, the sante surface It. E. W~it~. Ed~. (!-l~~ll!m. Uew Yor!t,ll9831. d~p, 6. tor SPCt~,~'S<1J1: im_Ic'C'd 001' !fQI! il~g the k~!U::liQ'i; r alOI1l~ic density. ..lid! I hC' 2>illl!le surJacc ,ooi~n· (I 8- K. Kill~ltl!'" fl~irq(hg.mi~oJ' ~~t;i' l'edi..rQ~' !)l1II1ey, ~hcORR W.QSfn~nnd "m.~y.zin!,llil~ l'Cl\.ll~'~~ilm ~m!\;il iml 'i 1(l(]fI/", PO but 1i00';;I d i1l'L!l'!;;'!!I1~h.'(:~nm ic h~' U!!W 'f0ik, 1t99·2~. 9. It R.,Ii,dlt~. in ftmll!}~D1'1l1¥li$, ~. lfi(;l~~ F. N, II:~, ~hc pm"'l,nilOlII [I~~gion W~Kj)L;' Ibe "ld80'lIJtion o( :;;11IllIC:Lll~ (rl!![~m] = 0034 rN, w h~m;:~d [IW] iii Ed~, (WIIJ!'I"I.I(H, fl:t!';';'I'Ofk:, ]:99,00.. lh(, tt·hm.nrl (:("I.'III)T :;,~~;11). u.ht" f}RR is being ilyrlrogpllS I,nki'llgpki.c,l' {E < R2 VI. Bl~t~f~IlI~~ aU 11t1:1i,;~ Sl.tirlhC\.~ I~lt.!lfIO!ari"'~lijm!cWlI'\r(~ j.t\hihi~ 1\\'0 di;,lilliDlgui~h;lbk Ilitlt('l1Ili,d I('Wiill1L~.:1:1.)' s,[':'111ing

dilTercn,t P~JNHhk/l s!..In1Lecs. tiC] e:.lflblbb struc~un:: senshiv iily. i\, dtal<J.ctc~:isl;i;c~e~ Ofi'iOlN:riZ:11ioll ClIf\ICS nhc relml'O'.~~of i \'Cl'SlJ:> £i fo:l' tlrlC ORR 0:11 PI,po!]y. [l'1I'~ ~~ and P~s N t( I '~I j )1\1rul~~n ~LI(,IO~ I I( 0.1 M} m 333 K is Sll!lln~!;~d in Fig. 2D. For

hl'i,lPi.~ 3. A~ !Il1.mmnrDz ...tI..he d~tlCD'Cl'lI.:lOW~iiidt'X 't ~r61cc:> have it1mtN~d~ydi n~i"Cnt oeli v.hy ro.i:'"lhb:, llie'lclion r(hat is. P~,~Ni,( I U[};k\ki ~~<:: 1.'t,lNi.(H [I);. ikh~ «< f't$Nt(lll t-~ill. whh the ,dUl!nl,!;!,c ~n
aClivII.)' bctwecnthckastnctlve (WO) and (iIe' m{ll~1 active ( I]~:s\lrfoccs being gl1C'd!I.Cr tIi~ru] n ) a

V) UfJ, 14), .. rpl,~1 g,Q.~olcl i;:'fl h,I!]C~~n~UDI ofPtl ~ 1 ~)-sk~ 1'1 \' ~!Xl,I!'l Ih~l~ of PVC hll]; been .m::hiL"\~11 ('mi:ilj:dcl~l'iI!:llhHI. '[t~c rl3Ni~ 1i I,.akin tiliil1iLcc C!(hihii~ rhe b i.~h(;":>t ealtilJlytic :'i:ct~vi~yu~m hiii'!: COCi; 1k~.111ktOCI~. Ihe ~hilU.i!jjl.&C ( ;,v,Quld 'be to crelLt~a ~illloo..'a1a1y.>1wi,t~1 ~b.::lronic !lrliid Itl1.Qiphoklg;i.c!l!1
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3'li' {2(1f)2h SlamenkooflciN Df.. ~ 21. {.1. (ilmpbelL itlmCl"i\trn ~2- P. Ji.,Thie~ 1', J. ,E:~p,iil 2!l

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Friig. 3.•.Ilrlf~lI!l!'f1I(.ef th,e sl!lrra<!!l!' mo~pnolAlgy anaj electmnli( sm~ace prope.rlie~ on the kinetks oiF o 'ORR. RR!JrE 11I1:(N:I'S~H~mrel1ltsr(lr O:Rljit ~n H(104 {OJ! MJ' ilI1B K ~\ii~n U(IIO r~!.i'~'ll:.lli:(li'i~er m~Fli.ne 11)1'1 p P~:Niil!bknsmWac;C') a~ 'QliIlipoarledl to' the (Qrrr,~,p®llld~ng Fl'dhHiSil!riaces ('" horiwllta!, d,aiShed gray ~.if[Nl milrkt) fipedfi"iU.U'II'ily Oir pollycJr)!,j];l<l111 ne' PO are ~horri'n. Specirri:!:: <ld.llvlly i15,'g~vllm ,ili5, ,al kin~,ti~ ic~n~!1t: 'd!e!liiitJ1 'k' ,n~~:StI!fed .at: 0,9 "V ver,S\!iS RIU, V~liJ!'~ '!;If d"ib;:!",~ 'OO!1~er p~iSitio", Qbt~inJ;!dlfrQm UPS ~pedrll il re llisted! flo r 'LM,ch~lIIrf i!~,e rTHnlpiholGlg:; i'! nd Ii:llmpa red be<blleen COrfe'5PO mdlillgPt3;lfi4i(hkf) arnldl Pt(bk!')s.urra,C!E!'s.

~5.. y, G:i,wllli~I. srJtf, ~Il, il'!t. :f!, 1;5~ (19~6). 26. 8. ~ il,'Mn, C. tee. V. Slamenk:r:tric,N. Ml.M.iI~i" p, I'll, IllQ~ NryL 1!InI, B 71. n5~2(1 ~200S~, ,27. Foi'IIlt!, IIIhtr 'twollngclMf)''lI.II ..1~~Iil&J.M ~ml@ ~I!!a, fDl.mtillbill. U.e' Pit~i;n ~~JINt rl!lPlled in 'I!!HVi~ slilblti, i!1 .!!Ii '~~rl)~h~lt:;!4 ~~tL l!In!j};~ th~ lfl'tlMttl.QO'~~~lbIm irntl!fh~~.OWl!¥~r, 'ill!! i!t«ti!ll~~ inrN!lcra!:1!! iNM Jitl~['uilllD n bt UII1 ec~fl~lru:ted ,M .alL IJ(Itffitiah IDe<aU!!lIl' ;lmiilO' F o~ intluoM MlMp~tljjj. 1M S.XS m,'il1l~ ·tne i'hNl~Yl!m fer

REPORTS
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H. M. M.a~,. HI, .~ Galroeiger, p, It 1tC'.i~,.I\. Fli,ys, (htrn. ~?< lIi11l:!l n:~~)}, ti Eq. 2,. jj i; ~ Im~1:!tt (liP If.eruc~ Ks ,tIJ~ ~1f!' ('Mlfai'li; .Iii!, ,~tlil(l(ljar IlII .miN PI ~ila; ,;my ~ tl'lll !;)flOI~ ~ f.mor5 {~,n ~~ <r;1iUI1~ '10 !lol:' 1&~;~~ R iii ·tIl~~r~l
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am! VJLS .. O'!dincV!'led,\J;e'Ifl>e' ~'!\PJ*il fro!11 tile 9:ln1Ir!!l[1 {EI~,,,J!;tQNi!fiCIfI:lllJ;5 n ~!W~1lb 'ij)!!l !lJnl~ll$i'l)o Ill' Clhif.!i~o:Ai1l~1'ID!!'>IJ.( ~ 1Ih! us, ne~iliI'lntf!nt o. IEnerg(p'. W~ o1(kncm!le!I~~ 11lCl' :mp~!iI iDl Gtlfte~~~ MinOOl, ,!!'id 1'1~!il1'ul diloWl~iDIi!; 'l'iilb fl. ,\ 'Gai!eii~ef iml!l IF. T. Wa.glill!f. CA.L. 3nd U • .ildimtM'1Iii:I~~ihl!~~ppcrl
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~!iO('lJ:Ikr 2iJV6~ :!f~~~ 1 ~u!m.'b!!r ;lOll!; ,Pu~Ii5~ed oo'lWJ ~1l'1 ]a nUilrlf 2007,; lO,lll61~ie~t~, ~'~1 ns Ilindrn:l~!hi~'lfi:iOlm~~oo 'WhCfl iting Ihl~ lI~p~r. c

nil ls ilItud)1 wac. to im'C'SDigme thC'ro~c (I rWaI.cr ns real c,nOlIY$I. inchen~icm~ sa:::-ph<lsC reactions rotcntia~~)' rel,I.."\'<U~~ a~nU.lSflhcric chCm~)1,II)' for and. in pm1.icWlio.a:r. to ~ll)ovid'c a ck"f' examnle ~hr'l w,u.cr.c'<n<ll~~ed nldk~<J~·'l1mk·eulL;' n:ii.'tIiCIkHl. The ~h.<;I,III:-atom .ab"'l r:nct i,on react i:on of 011 with accUllklchydc preved jo !be' .J good lih~li~ ~,~S been CO,i1sidil!lfabl~S:~'Cldlat~oili! ab@~Gt. ro~~,of wat~r ~l'Idl \.,,;uI~r((lmlple.>i.~s 'inl 'r:::b~!1fIk:,a!l :<Y~lcmrO:r ~tlld)'inglhc (lflcC'[ of \\'''~!.'I''a!l:!ilr~ t~,~ s ,1,l,1ltion 011 lilCl k iueties, 'lhr.,;reO'!{:ti-Of~ do~ u{n ,~al~~'has.e reactions, fnd~d~nf.llh~ ,!::oli!.j@~'wre t~alt ~\I.ate,rmay ad ,~ 11mot'e(!J'larcatlllys'~ 'thrOlJg~ illvc,h:c IOJlg-~iH.1.I .... ,rollgly 1x~1!I!!d illlL'\!nl{ldli31,,',ll it!;aibiiily Lo f®nYl llIydmg,f!'ni b(md~,. IHerl~, we present kil1ll'lii( :stlild'ies jnwihi('n [tie em~ct ,(l'r 'w,8lter oilier cm1l1!"lk;>i.•. Orll11liiTIg ~ 'ClIiiIh,e ralte of the lreadion bel\lIBen hyd'N'X~~ radic:alis allIdact!ta~d,~hyde !has been lIIl,eastired d]re('~y 9,Lt:h :15 those r(H.m'ld i!~ t I)JmOIcClll~f~r reactions. fot ,,;:xj[rnll~~, i'l'l Ui,.;: rein Laval moule ex', ans:ioM 11'1: ~ LO'Iy~elTllipoe;<lltlllN~S. il'lu!e<l'~il'lg: en h <lft(!emli! nil of~he reani 011 rate by An oct i:{m or O~I widl CO pl'OClfooillg vis! 1[OeO adldiad W.il/~er \'lia~, fOlJlnd ~'IIH~h ilJew!.1Isi'ng t!llrnpelra~ur,es bel'lle,El'n! 300 aad 6.0' kelviilil. QUi3,ntum dhemic,a l (iIItl!:uti!ti'orn~ dirldl :staltiS'tfi1!:1Il rate ttll!O:rySl!lJlIPOr~ '0ar !l:OndllSioHS till;M:th~!5 o:bcSlEllPi!'.1Iltion ]~ ron1IJl'c'..'«$ (J 4}. due to the redu:ct~olm (lir anill1tr~nllk fetU:tiiOIl b~nieil' (,lI!!1:s@dby :spec:iilii!: w.ali2'r ~g;grll9atiion, 'Time Itc<ccn~ c.x[)CrimCm3~fLn:d lhoorelicmJ worn r@:SIiIIU~li!gg;~~t thail @i;I",llIrI~1In91~1i!' water' mol,@(!ul~~s 'Call €!C~ ss calaly:stsi IfI Irlldlical-mo'~e(1i! I@ we1!~lijol'l!i. ~~r.lSIll.'OVided cvidc m:::c fo:r the mSSlJ mill~! 0m!thai, 1 !hc rc~lc~ion proceeds pl1Cdollillirmn~iy IIIwl!gh e mdena coli ision pil!n;n~c:rto ooli.isioJrmUy s!ahil ire ahSjrnclbon of~hc aldch,~dic hjldrogcn atom ~er ialtl:'OOllCeS ITlOny lUll1Stml Jea(l9~2). ~dcHlion the on rodkJl to the .il1tc.t1~lcdi~!K"5 or such reaciinns, Un~b,ti!lHm!ely, ures into ~~ae kinetics and cnergclics aHI'hcS1::' cffb:;:lS ,thc'J:caJ\cr IlCj1::rrec~~o .<1<S (lie deuble bosd docs nol t<Iikc place hcCllI/I~UI,S . . of ehemical and biologicalrtill:l:im~ sysl("JH$. which are di mel! It to rauonalize on 01 Cm<ll~i:;;L'" r.ltdil::,)~·cG~nplc:x,m" the C~!(!rgy· aetivation '!.;'~crg)' i)1 much hJr,gcr lh,m lh<!l! for the , h}'(i!ng!.;'rI <dlS~,"'CliQoJ1 (13, 2.1). !n",dditinn. ~hc IfIm~oCculhu~c\!d (I ,2l B!"'~Jolld~he'w~~U·kn!()'io¥1rn lr,,1'llsl:Cr nnl.'Ch.un~ is!1il:ll) 111,<1;"1,1 :i nllllcncc ovc-r,,111 <)I.llkhydit;. ~Iydm!ilcnalmllil has.u rdmi\,'~~}' ~'imr!lll pm;pt::'ni(:'s or bulk \\';Jlcr, th'l'.';R:is; e",'idll.'n(;(;'ltim! ['1;;'<I.(;li,oJl yi,,~ldi'i"illd nlh.'cS in binlOJ~(:L1llar ~Lt!ir:ms (UI anti. ,ICC ofi('1] diirrictlll~o dU!\lGn~lilsh. ID'iltd ill]!!; C:llc.~y compared wioh !.1m! 'C)fSlyd roeven K~nl~~ oO:l1lll1lcd \'II'm(~r mo~Ct:llllCl> ~n meleeIIIlnr 'Igg[cgal(",., may have fi dec isive i[!l~1:u(~l1ce For cxmnph:, 'llt!:1iC' is cxpcriuncnt;ll V5~,ind. l gen ilton.l~inlhc r:J10lhyl grnLlp',. A ptL"'IJ'cfL'tdve tI'lDOretUC-1LI q.~), evidcace tl~,%l II~c hydro~)Cr.Q.x.y romp.k:x 'CU:ln·IO",HO has been p(HlDullatml, O.!~ g.1Spl'lilsce'hcm:i~lr:Y tJ). 1\.1] imriiguil1g quesby 1)'Allmi e~Ill: 1[1..1) a m:l A~vfilrcz~~.dalboyef at ~IDnis whether wmcli molc.cuks can even act as rndicflll II ]0:2) IO~Ul~ fL~~y~Jrog'l!iJI.hOil.dedCOil1ll· e !ruecamlysls .~n ci~C:ll1icli~eacuons, for CXfll~l\'P!e. p ~ex w.illh a water moOJcu le, \'\Ih:hrh hn... Ihe [)o- o!1'3), and its bindi1il,g en.c.rgy was esii mated to be r Ilrt:!'Cacli'!'c tClnilll 10 alleci I.he klnelks of .~ reacI.iom •. ~J,;.l kJ mo(~. A~thotlghlmnsicnt 10:! Ihro!liSh ~:hcirahility 10 rorm hydrogen bonnls. A ~umber ()rmolccu~.'Q,-wru~cr C()tnp]cJ~cs UUlW eviA 100lglhc!'>e I rues., the r.uc CO'lms.,a~n fur I.hc sclfoompl!c\"re1 'Ila\'e notboCCtl dil'C(t ~yc~mll3.clcl5It;od (;.1!;peril,l'tCHmUy.~leir p~cm;;~ has '~~n k1emalkd denced p1ul'lochcmisu), dis;~i]lCi iixml1l l~l.\'tt orihC' readiO'ul 01'.1 2 ]0 yIeld m 12102lind O;i!. wh1ch illdil:ecl~y in severn I mdicaJ-l1w'l~cc~l:e :~cacLjiO.I'Is proceedS \'ia a h;mg-livcd in~[enl1lcdi.au:, j",s0I1fIe· ioo~atoonnoJe<;lJlcs COII~P'l.e:CS orw~m:r with and a~,,:>;ociate<l wllh a ,tlCgs,l,ive tc,mpcrntilll rc OO!JUlC (.1), sulfuric <leld (6), Il~lrk acid P'), l1i\\':~lflil~'aster iIJlhe I)I1CSCIlCC (I r waOC'r than ill its H'(YL!,s mddc (;"(1, ,<!i~~dl!lhninc ,elioxidc {$I 11) ilIcl1C c dcpem;k1il~' I he 1'Cat:I,ion ,L'3~C ~';.'i,.?4l. Nil:~scn, am;ellcc (4, M. 17), A similaJl' e~rcc~ llas, b.~n c."i..lnlU.l:k!sf wmcr~T!C~;~tl.~ ~I}'J[ ~m~y rm~ obsCil'vcd rOif the n."<lCI ion (I r hyd11OjlC!i()1';)' u<)d· ~nld co-'wClJ'kcll" C',aloulll!~ oor:ricJ'.~ {1lC1.u.i\'c to o be ~h!:' ~m;wrn cnc!.'gy) to ,l'Ildch}'~hC' H... 11~tr.I.~ic,~I.. wilh r:!~lI'Og.l.ln dioxi!l~ in ~hl:'pT1.~'nCC of ~lom~,H "1J'lul~phcric rhn[oc'h~m riS~)'. in li{lir~.]'!~~hyl '1b:~H'H;:l~~m, OH ;to.di.l.itl.!I1ln I H and B~,yo!1d ils ~.hili~y I)(~il1.r'l~1il;11IQC :p.hotOl;;h~'nn.~ W<lJ\,," U/!,'}. lio\,\'i;\I'cr. thi.;l PJ:iWI! 01' lhe (.i{lCU1; . I 0 I ,.'.....Il~{l ~-I I ~.. I kJl '14'" kI· J ;J(jU)'.. 1{jn~, ' " , , i:·m.y. \Wlh,T!iS. klllUw;n(o :tbllll1l ('/QlliIpt;';.~\1 ,,,,itlil ~n~~IlI,,;d, i'!np·'K:l C!r WOIiIl;£ Oil ~htk il~1.'lkt\ i:1:J, gl,'i:n. DI1,1,' c'n rll <1I.clu"',.;: ::;peek"::; Klich 1"" Illdlcld::; ,lIld ptll~u' Il~o:lt... cro,~ rc~nl,lill ~ L1Hcl~I."flr. AI.ihmlgh . ~ncn:~'IC!DO:!:l" . iJnd 30.9 U unQr~. "!J~'tliV\.~ly(14}, A R nlCg<!Ltiyck:fl'lpC!rm!U[,I.! d"''P'~n~llc'l.: tl n~ g'C;l,'I)C'L1 in~uacrm,11 l.'(~LOlioH.·' l2. 11), .<IIlt:! :sudl ~~ (Ij!, comuJ1C'-'"',,-'l' or will!Cr in ehtmlicr~.~ refrctiollK have comp~(;}Lal,ioUlmayfl!t\)J' im:l'ia1i:>ic N<lctLGI1. btu:· bc(;1'1 ::>ugg.cs~cd ilfllhe past !(M)" fl cOJlclu::;.i,'c lio'rrl fmc 31.high{:;rICilTlpcrnl~res has also been. rier.o ~f1 bhl'LO~C(elilaf .rca~liorL'>. lin itdditlol1.~hc mcc~lill,~sm and rl dcmUccl Idncl,ie ilflal)lsis 0:1.' :It:jXlrl.cd {22) Willlo~u ;j], dlcm.'C:ti:cn'l.al1llli!,'sis or nh(: dleci.. w~rc noogivt':ll" The g\."il,"~['.dprob. !1tdic:.LI"(}ollilpiex. m!..:ohatli:s~~.'10( M~i", w~111b~'J>l.V.UI the ~'y~b,.,(lns to hil\i('!.~ dmtm;i~'rii,51:it'~I1!~paICi rooomh~lrulio.ni 011 ~etll rOr 3.n IiiDicl.cr,l>IHndi:D~!J. 0.1'~lGehkiiltclj,e :.y.~. We; l,ilf.. ~tnrcd rnt~~s role con."~3ni:.. lor Ihe ... lind: . OIJ ~ ['~,I::lCUO re.[tci~{!n il1lDhe Il'rel>el~c.etmd p<rocessc'<;, whidl could '!be cnhlllHcoo by wale. tc~n:> iii nlmi th.cy dcll!i::lld upon i.OJ1llx:mwiI'C. a,k~(.'~lCC orwa~er vapor by using.<I pul'secl :lava'l, ~nok"ClIlcs. lasL ,,~JtoCrt.l'l<l.y ac~. s a PJltkli]~ rl'y pressul'C, and speci Ik ll'llO~C'!Cl.llar pammelc .....and a .. W reqJuil:e OJ, clcl,ailcd killclic ml![1II)~sis Jsillig l:eL uo:r;z~e c....... pm~sioll {J5, 27" ~FUg, Il\.,.. The I'e:!:C~li1I!!I!!lUt fill PliIySi ~1:~{BIe (hemle dot/' !llII1lWEJ5it§1:GEill:ll'ig,m.,. Dion~\'a.;;;:rolioWocd'by J\aser-'~ltdml~d Uum:esccncc act uon mtl,:' theory and hi,gh-te,,'d q mmtulll i~rnnlatqns.raMe 6, );70n GffitUngen. 6t.l'flflan~.4De~rLme!i!t (Un to 1l1nliUlor the d~ay of.ol i lad icaI COIIF chemis'I;I:Y ~!.,~ of (hMri51ry al'ld Cile'pill'1I081!11 or Ealtill aoo A~illai!gJII!lI'1' CC!~rn.lion. The r.;;Jtc cm!:S~.mt<; WeN dC'tml1~l1od FoIIO"\'~~g up on.;;l iong-!lW!iding c'onjcclllfe ·il:ii!n~!!'t, PUirdlJ~ 11lIml!iers~W~~t.'6Jf~te. IINI47'907:"2OB4, 1jr4\. . of 5f:lto 300 K. The UII;,I~ 1;VI."!Il\ling Ii; 'm few \\"'''~~TmnkICld,!:'';:IO ~mw motr a 'l~,!T!J,IX"r.lllu:"r.nl,!',l,C . ;i1kt CIu,;~~1 ic::. ~Ii M,L"II1.,;tal auld 1t:'11Ict~(}J'iimnrieffi o('lL'OI·Y rmti,fiilb~carl lx~ fi~tI.,."d wdl by 0'1:11 ~xJlKlI1~'n· "To ~!bom ~:o~~e~ppcten~e 1!IlIG~dlbe add~ess:ed.,E'mait li!lll~l1itll:w ~f~.g. 1:1) • .P!ot;,~or~I!I:dO.~iffit.o(].Dikt I 1 il'i P~lrt~c!JJlm"or faVOrl"t~Lctiofl ~.iitl""'.11ii~.!5Q,:I:l' of Ib\;l~l~
0lI.

Wate!lr Cataillys:i:s, Oil a RadiiIC:sl-Mole,cul,e 'Gas~PlhBSe R,e!8ctiio,:n

nc

or

l·n.

or

~nor"

or

4197

REPORTS
rotc C\lIIl Sl,iiU~~ versus ac~.~,~chydc cnncemra~!(l!1_'" (i,~~ii;l'ogono~~rrot;f ~) m .. ~Q~Sl_;lnt rom! ~ dashed ~illc::l o(ll1l'Wcggng q_he d;!Jt<)P9,~I,lt;;. m;:lJt~, CXI'l!t,,--rimenls 5\!g~::1t ~hm ~m;J1I <lcemld("':hydc\.... m~Tch!Mel"s. ,~I'"C~JCWlrl~ are '~;r:.'$po!l~illk: bv I the di1To:ron~ b<.~I.\"''Q~Jg thc~wo C!m~ sets <ll1d i]~~I~'!,r.~h ~Iw cJr~l,;~ of ,u;kl~. w,II!,,'!", !S;o,;p~'1:~. nhL,l O!b~Ii'I!!.;:d (,ifl'!J'~~T 'a· .. \'1k 11,O,to.;: WI,; did not ,nqlsl!ll~ "pure" rnt~ that ~n(,':l~i:s; !J!~n;::ilt;:~ion.w~th.o~I[ <B~dc~ W:llt~'Tin or tho;: nUigc IJcl.,!""1,.~n SCU] mll~ ]:00 Knn(\~~l> u redby cO:IL~i~mi&, lirnsi!:ud. t&~cobser v cd l~mp'6rntmli.:.'" Crow~oCY ftr (II. {2J) ,,,,~~d Midmi:! 1.'1(~/, 09~ are d~rc!u;l(;nl m~J.;: OOn:5,!11111.k( n ill lhepT"C!5CI1ICC of 'i\lal~r is t~~~~um or'~wo con~r,ibu[ionto. fmm the shm'l'!m .m!:! cornr"u-cd wi[h ',"",If d .. ~~~point ott 300 K (Fig. 2.ill~~i!;!i;).~ll Ol,.lf :;;l~ldy .• we :.ignif.. unel USh;:iI'ot:..1;!:lind du::;:[~rC(1 s:PCciL;~,fi.";!!;i1L;C'1 i\'dy. iClimly c:){l.ci1dcd lh~ ~el"llpcru'llIIn:: f<lIlg..: and HOI h ililtt:lum'is 10 mid .~g@J !tllt:: r~&..;,oc:i'lIh,.'dwilli. confirli~OO thenegative lIeileili(k~'iae i!J.i"fi iC[n· Ihci:r' rare COl'i~.~~ii:U and k"iir Thi~ s.itumioll. ~ pcrnlure or Lhe-reaetien rate dj}wl~ w 58 K. ~n om lX':~yl1t1ioo:li.etl]l~y~x~re;,scd b~ Il~dilion. small ~1J}'tOmrt.~f ;[lI.Jd~d1 wster aco celerated the react t{'l~1 (F i£. 2). A Iti~o,u~hr~hc c~rcci. ls small a~ 30U K. it becomes rromh~\.'l1t allo"','cr ~C'~n''iiCrnUlirC'iS, T!~c c~.ec~OlI:<o .ooCJ)eml)1, If! oroer .~{l! 'm~i,[lf1<llijtx' lhe enCel or W,~lcr~u!c1 on the- w<I!tcr eoml,,'n'~ of the bllil1cr ga._..., I r~hc Ct1n~n!['<f!ion or \\I,~rer~~ ~ignif'icm1t~y b(111::lW '\\'.ul!r ,el~L"l1.I"l'n:em thc· ri':<l!Cl~OTl nne, \\'c.; p('rfo~l"11!L..t!, ]. ~h" rio ~ign! ifie,I~H ·1~(:celi;r;1litm ~~ s.c'l.~n_ On qwmw.lu Ch1,11Jik-"Ll ~',~I(;uhui!Dn [(;tll!l.p.I'!,_\1;l·ot:;[u;:o;;['\,!'.!t .... the- o~h~r h,md. a ~V;UCl'ro!~lloQlltof rnore I1mal wu.h jo;inlgl~mltH doublll ~Lifld ~)c.nLl:r~nli,,'(~ l ..iIJ.~~ ~"l%increases agg~"CgoJ~itl\!l~n sueh aw,~y liUlIJ cseuanons, C··s.D(n~ ~o dllcicl~\lc ~1C' ener~li~·.s 1~ucl $!l1lC'lilll re 0 rth(l i!lh;''I!-rlllil)du1~~.c z>llOC'i(;s ~h('l in~"'"~i~}'·of dl(l ,n~1 .'i\i:!,lu,<l1 ~~ig!liJk'~!lUy i~ ~CdUC'L!(!. ,mdi Illie fC'aocllonD,,"(:!t1'm("$d~fl'icu.~1 to ,~rtilt.' rt1:11~~im~I.FioM- 3 I, ]3;~!C,lue~hc aeeell ernl iUrI cf[!,?,Cl11;:15 b(\("11s.cc~ un~y fur !;mall w.3l" mt~'~S!IJ[,(: because of an 0'\ erall q'lJclld'li~lg of LIE fruml the lk:p('nld(:l1o~ of the ref~Cli(1~] nile WI' ItlOnttllll1'lli:(1I1JS,. we pos~l~d .~~ sign~.fmmi'lrole en water ,~uftdm::ctaldc{hyd~ ro·nlcenurn~ionl$ and or srrngk'. \'Io'3N!iI' molecules h~ ccmplexes with For comparison. Calcll~::ltiol1s the dens i,l.y mnge ~Il~'" hich we fmdln :Ii ilC.lI' hy acetaldehyde., or the r.l!.C versus UC<lC'I.I!L1.U and waier 1::011-- were' also carried OIJ~ lor the rcactien ''o',iI hOUI, ccnu:atuoll.<i.. a_" \\IC II a<:lhc- cxperhne ntil~ ob\\1 ate r: We refer 10 two kiads Orc{mlpkx: the servauon Ihm larger clusrers {11C"C'IJ.k'Jchydc ~wdro~C'nobondcd om: between m::cI.:lldcOl}'llc and waR\f, wh~ch ,11I;;·tl':, as ;a nmg:cnl'(w~lh '[)~n and u~~~_I.ldcllyd~·'\l\!<Ilu.'!f) much less reacare tive, lQtu.li~g to decrease .·bc ilC'IIlIm_1l'C'dCliillltl ill [h1;l~xpcr~n~I:'V1~~ ~vi~h ~!:;!O pro.;.~!l!. ,u~d th(~ COl]oC'nlu·,ill.tlO',l:1 (and in~um~]le ml.c'oonS!Jml. ~]lle ~rm~~ ..il1m p11l'r(~,C'u.io~1compk'\IL* CorlJico 0.1 [

cr)
;,!I:rio

~1I;1]si~yyi~!d ~h~ ~(,;~ol1d-pr~fm!A; gom,!;mls, Two I1D'("M;ri~11~ml~, ~ti~m.u ror a h;~In~q":~II,I[\Ll arc of K wi'th ,UlrJ W~~h()~li[~dtk-d Wft~cr \'O'!{po:r (fig. !n,Frmll Ihe ~~Oll~~~ [he pIG'~. or

'n

f<1't~ CQ!l~t<L~'~:;' of4.,fj

x !f,n

'~ufl"I~l['i']il,I~::ll~e-I,:s,-:1

~,W "

10-11 criF'.ni;(lllI,.'ClI!c-jl~-1 w~n; dt'I.\.'I(~

m'ii nK"a, SI!.""''-'r.'iJ. ~"":.iX'l'inrnclit~ were l),\,CiJ1!I!!-od IL lict'.r WtMllirullar L~~n~~nlii\: (2'7). AVCG!I:!:,>eiISC'tCtrld, oruer :tii1I.e co:ni>l:lii]JS ofl.h..:: i'C1lcliol~ !l)fOrni Wil~1

~,OI..~!lldch~c alone rulld~nlhC'p'~scft~CC' of water \!~l)or III ~em11~cml.lIrcs el~.'Ci:'tIJOn K. andM.I :K b arc ::I'i.o"'\/ni nFi.;. :!. [)Cvia~ion 0 ruh~~~ libll'll1 m~n Fig, ,2 110m lot a Imrnml Arrhi..'.!l il~"'·l)'1l(: p:I'ot f~fl1l k v c ,,!~~ns lin is :>mllil,.~vh.u arbilnU)'. ,mod. ~h,,~r~ XIO ':I;!"!>.O'U~ iiil W (::lip~·A;.'t 11 li!w,~ri".:.."I:~i!Dn ol"t~lil' d'IW. Nev(:nl"i(:~ less, the cho~cn r~OI is ofien used in c..ses
\";'here bi mo~C(;ul!,~rale 'oom';~ilm~.~m:re,li,~c i(h [ w dj,,;crc,~ i~!M[~~IT!I)CrI.QJWw.Th.e ori!i!:~n of dl(;i>O' .. c <Il1l;1d n~g.3:~ HW h;!'Ilp~r.nllll1U dl'pr;.n!ucrlO~ de-

~x:Jlds;$1.roJlg~y 0,11 the 11lIo.ll~uli.lr $y[:;~C:-IIn1. As. iOhown below, olic reason fotlhl! Cil'i!cl mil)! be: a "rcc'''-lYllCbfllrrkr below I.he ~n!l:I'gYor the rcll.cla,tll:S~il!he C.1lI mll!cc ehsnncl on:l~c' rcactien,
as oppOS\!dI
~(l!

a mueh hu'gcr b:J!lTier

lite: ly!]

~Il ether

cases

I h a~

would yield

[cal

Al'fI!C'~1'hIS

liloeh.wior: Ih~ \;v:ilcrcru~K'm or~hc beam WO!\l, ll."I!)1tll!'l.'Xi ~ with o:a:·1inc inJrnred sf!ecl'ros.copy and t1ctel:~nincdw be ~%. Tlsc shaded area OC:lWeCH!lhe

,A

II 1.0

II
I~

:2·48 IlflniI

EX"Oilmer~taS!fJr

1!!!!l!!!!!!!!I1

II... tJ~~~.~.~~ M F
M

!,

II II I~

Q.O
-20
:201

'H)' 60' ~ ,( '~I,Q?E!is)

"ilaser

r ~,
e

3.5

nm

~ 3.. 1

2.£!iI L...-_ ....... _-i. __ "'"--_--l.. _ ___~ ,tU) Od;; 1!{JI 1..5 2,02 ..5 liCtina CiH:ClLo, (~·Ol~·lmmem.d·e !t~),

F~g.l. (,0\) Lav,flil mJizzle expalli~:Vc!ll'lapparatus L!s.ed:her~. Po r'i?:preSlents; the ~rres.slirein, 'rf(ll1t off the b1ial now~; Vli1lii!clIJ2:, valve$rediilg the' !iUilZ:l:le re:s~rwir: F, fw;(n~~~n(:e: l,. [em ~yS'~ml J1(!f tl1:~' (Q~!.~ct!~O ,ofi~h,J:or'e:~~nce; M, mflnr~lFS; amll PMT,.phol"Om.ulltip~l~r. '~B) liIIF p.refile·5 ~'i Oli iin the reedion bil'l~\!~@f!' ilC~Mld)ii'liil'd~ art!~ (]I". radkals (r.';" n K~ Pllli =:8,'S :x lO~jj,

mO!f\CllIle cm-J, fl.a~d~!1)l;1~ 1.6 x lO1!t!IfIII(lI~e(lL!~e{rIII~~. Pol!! ':0;: 1.5 x lOU. "" mo~ec~le (im'~'~}, (C~ Ps;~~d:Q"fir5~-c;rderr ra~~ ~!!).ns.lal'Wb ,~s i:I 'r!,.litc~ioi"!1'of Ik Oi!o~~ldeh~~ 'CW!(~fltJ.l!.tlr~:rI~IbIJ.d·fef N2• T = g~, K, ~ml = 1M x lOU; mo~ec!lll!f1! {1iTlI-~). C~l"des., rei!~ti~11 WiUIO!llt wati11r; Sq,UfU1!S, uliilCoon with

~~at@r ~sif!g:~~ ~):i_p,ii'nm@nt$)<.

REPORTS
plus ('n~(l!OI,w.c\~QJ'I and For thCNiJC~b\)1rn h~I.\WCn OJ! ami (' I ~lCF]O Iii> .I"!ydr(l~cn· IrH~!~;;1,qd gO'~~Jil,h:x~:;;, Op~.i~niz~d :';~!lI!;:Wn;l~~hr con~,p.k:xG~ m~l~ ~r,HlshioTI "t!l~e;:;; wiH~ and wiihml.l. wMt'r !U\_~ ~h(]\"'II~,il~. FU!!l_.:L[':JI.~cuh~~oo structures Forl'hc prcrc.tcliml complex (Fig,. 4!A} '~IC lr.m~·~lin~ sWlc (F ~g.,4B ),in ,I;~~
~C,~! r I~

fC""1i,0IJ,.
mte

<!i:i.

charac terieed in t r~~. with

as

Qr the

<1'~ o\"~F,l1I

~1181;O,!li! CXPN88Cd

,mo

!;.'PI!:<;~Qll Ik'q~~~I~cii,;s; ;IICr..;!;11\;!~~~~yd'c·N~ i for ! l;;o~I~· SI!;:II!iS. tlll1gmhig;~I~~~!Y ~IiO\,v~"d .IUll ,.~I~ ill~!,.'rm~l~..d;~ljc. II,.W.i:1 up to, pn;:..'iun::... of Il»lr. . ,c-mlmJ;l oooolli s~!lrmly ~<;I,~b:ized ,,~!dIh;n ~hC''icaclUOIl is i ~i l in i~~ !M'i"-pr~""""In;: Ilimit. lltL;:i~ I1lldi:(1g~ ... ll,~in (_~ p
..... I'~o liI!(::O:slIJ'It f.k:~lidcnc(; 1i!I!llix;o.;li fOWiliid for hy o Dhii~ 1'C'~clioJ'i (22). Olle crlllM 3~llIc Lhfll.ih~ :.i'l~m:liOr.i ,e-~!ll~gC:;
W,hrn\V8~ei' i:'Cil~!~.c~:.nil:i'oo!;j!!i11lIS II. co,liii:>botl pai1L~~er.A II h(iu~h~he ·00.1 ~isien ~roClHCi1,cy fol:'water

shal rlJ""" RKllcml:'!1 \1i,"C'U.A 'mnr;Lm9~ 01" !he sem {If Kri;)1'W kb>d! \"jlll LCIUl a rd-Jenes tlnQ

ahsence oJ

wall~r

:LL'l;:'ill

~gw!;,'lIncn' wjth

ric~

k~. klbirvr

vious. work ll·n

l\.,cco:rd iil~ 'ltD (lLIj}Jtlun~' d~("iniC'!t~e~lclilal~o:i'iS I( :Fi~.lA ~.t~IC~;1~fi'b::'1rnelionirl':$lctionl i~,:.J.ei\'~moo . 0)1[i il:ullkw'o' well end ~, sm>'lJI~ brun:icr s('p;lralin~ and Cl b:l'O}.,

~U1{! kbolGk .. rc ;llp.t~ifie I:'lll~t;o[m:;;!ant~ (17) lilid ,L1Cikild Ui"JOl'i •.he t:i1cl'ID' and d:oc:~LI'ig~1 ~![r mo~ncnl.lIi'b11 hit\::' ilJU'~'::Iil'i'llcdi]!~h~ 0 l;.'Oi1lDP;leJoi: .• sl)(."Ci:r:

~he p!I~uca('li(ii'l cemplex and the products U ~\~o mmnt!ntmn. A ~~hnpllifi!.'.d :ii,talh1:i.COII adia~)[iJic nc barrier wnslidcred here ~ml~1 ChallllC'l model UJ~ \\'<1$ <J!j}rlic:dki)' l!sil1£ be dcscdbcd as a ~lQbrm.'if,W-ld b:JIJTL«'or reef Ol~ cn~~'C':> and cakW!bll'tl \ ihmti:OmlJ 1h.'quenciC$ of,thc rea',Hl!1Ht ~lC' rT'(.;'Coln~l'!e.",. lnd the transiO ~h~patil rirmini.llwlnc;n!.\rg)' Oflhc re<lclion. Gil,e tien $tn~ liu~!l our ql!~n~lWII di.L;'!IUi:;.t.ry ealeu\"" lime ~1I~~[gyn'!l!I:\imll:!~l H1,,~~~{lIli,V vhe ~:s;ynr~p-, 1!l!iom.;, Tlse ,del:Cliluilli,,'(I [filii,)' con~..I!.(U1'~ .~n Iliu:.: ~wi(;' 1'C11,!l!i.;'rrl'~ '~ntrgyi'H.~l ~lhfW";; lb~ cu r!uiI.'r(;I]~ C'r~y '~ni~liIHlq~n associawd! \\. i!h uhc PI'C<f>C<l'C~ b'DW~ cm.;'~,~ rekVlllll For the lCmper.:ulIl'CS Oflhi!1, C'fllnlllb~. Th i~, vul!. ~iolli!Cf.l!:rre~lmnd~to u lc ~J; ~. tJ)I,pcl'ill1enn si1o~~rcd .hal ~ifclilmcs of the COIlH" c-nlnmcc rig id C;.,jl (;OH1.pk'iX-.fo!l1fD irlg biujorlf,)'x~ <HiLt.; ~n 1Il.CIrnw",- pkjos~'Oond r-.qngC' OOC,1!llM;

be rate eenstants k!i;]1"'o1 and k!J;!d<~ wer,e e.."il0ulat...::d first ~~ 8i ~;um:l.ion o,r cnel'gy and 1!lle m~tl~!l.r

~(!~ICiI:SY

1:-: !Iboll~ I{) t~l.nC'l\~hrL~ forn i1ro~c~~ '~2l!')\.lhc !l\'crtransfe ~'i:l in a eollisian L~abou~ S lin~C's 1000J'gCil: 12;Mllor w..~crC'onisi(l!IL~. Theretore, \\'all.;'f di hJ!~1.;1dI aJ1.1>01iU 3% in ni11f'(lgen sh[Ju~d n{l~ w

IX; I'l'I.Ci3'J..'i dficient

ill!:: ~h,.; re~icl;u:n

~1~ll ni'~rof;ll~l!l a]mw ill :<n.'1bi!i.x~

("otllwpl!(!lI.o.;lt

The I:":g~'JIt<;rgy I

Iml1."i.i~fdHcic'ne:y (If water l'oC'hu~",c 0 the nilITF 1 ~Il bu,ff1,,'1!',g<l!." is con~!'JCn:;,mt~d b~·'t'> '-'ltlll!! con... i C~I1J[~ !iol1~n .. [he SY.'l1,'i;m] a!ldlllWl~ C'.. ~ql~!Ol be 1t"'lj)Q!i$ ib!~ rOor I~I~ ok ... ~...'l;,o; enh;Hllt~~'m~'1!11 or Ih~ ~
1Il:-11t'D !In.'" ion

We; have 3~SO used SH'LtL.,lk:d iliil'li.liwk"OLILir:r. Ihoory {2il' 10 calculate the CivC:Ii'3~JI th()j'm~:1 r3ll!' i.lQJ'iSUIlIH [~s a lUi1I~lli on tC'lfIlper..Ml1~ 1/4). Re.'>MIL'> rrol1l1 Ih~ CJI:CuhLti.Gll .lor the o\,cm:ll] 1\,':lcrioll \1!~~hmn added wmtcl' cx.p,~id~~yH}Dkimo
ImlC

or

Ihe CI'lI.'1!:g)I ~nd li!lC .m,g.tl1br mOJllC~mq m ~~po;;'!l:de~!~ 0 r 't~!C' r.,w con SI;ml>1 {or~bc IbrVH,l;lion .of t~~c ')Ul::rc0.c~ID.1l complex, H." we~JI !IS dlC compethion between rorwwd reaction over Ihe b~ricr and backward di...;..'WCirHli(m .acro!di_rlg~o
~tC(:{ll!l'l1

the Ihm:J,I~il'i:11l giv,,;:nUI.1 041 {Fig" 1). lllcoixlcf\'od nl,~;t1iv(; tempe..nu IX; dl,',pe~ldI:n(:~ w~s ~~II;IIroo pKKllICL'iI. ool'lfimrnin,g ~hL:'p~12'J1C1; 01' '~10 proJ;'C1L'D-

tion eompkx,

li1l!i.': (lllil'C'IIVOO 11~~tl~ivc

tC:l'i1JpeLTll~Llre

[JcpenoorK:c CfI!rrnbe mti(:nifl~~j5ooil1l tenus of d1(! d~IYeil't'nl Ctie~ and fnigular' ntomcuanu depen-

dences of II~c speeille rste 1.00

Om'l5!fmls.

kifi:,1!!.\!

and

,. (11\)
with

riig_ ,2:. Rate 'C)Oi1S'tll~ts k'ror [he re-;;Ictijoll -011'1 OI~(l1IO (1"11:20) 2 HjIJ[]+ CI':hC(l!· measured ~'~re -I... (s,()Ud c!1n:i.eS) ,i1f!d \li1th!)YJ~ ~w,Hd diam'(U1ds) il!i:l'd'@,(i wat~" (Il!f!:iI;Ht~ Datal fif(J;i'li! (22') ~(joP'~fI s,~ulafe\J.)o., U {o,p1el'l ~nlr'l:loestartri this ~'IoI\CIrtk selld sq:(Jin!e~folr Ule- reactl! en ~'I!i~hollt.,vater. $o[icl ~i ( ne i!'ldkalEs, silrnplJfted SAleM (atwliltiQIlJ,

~n

k'ocd oflhc ial~'I'lcdialC eemplex anrlDhc rest!lth~g v,il~rn[lmml.2lJ1d rO!ru!ional. channel swuiLchillgs I J4).·.Il1I:is i"CS'l111 shm\'s lilm a IX"Cr~l)'pc n.'1mcl' am:1 the OOi~po:ndillgl rnl1si~imllstate may be fi.'iJ~delcnnil1iing dc;~piic their IlOtel1l~al cllergy lying below Ihe cnCrg)i of Ihe rcag~nts. Ol~r qlmmmn chemical cakulatiens .~hcd

light

01l

!.he rn:igiuof

W,~I1,,>iJ'~~ ,1IcC\!'ICi!:~n cf1'~'tt ~I!.i:

Fiig. 3. R.elatil;!@'~f1(!f'9~~f(!)r (he re'ali1ijonin Ute amel1l.l!E1 (A) and the pR5ei1le • ,sf

(ulaii;oo at the ((50m revel '(!If I.hoo!)!,wl1Jldile oo~latiQI1I

,QtllliSIStefiJt ~O:lalrize.(i 'lJijp!~i!-

water. 'The ,e:nergjes

,Ell'll

cal-

,.~~(l o((:c-pV1l'Z)

ba~s

se~.

REPORTS
o;~ihis U~~om "b~I!I,;<l);.tig~rwcl,ioll,
~,,!cl[J1!;r~cd energies

:si£nitkaH~. imp<u:l oj' "il;lC~"lclchyd~d~l'lt!.'n'cml be ~-.o1J;; 0, k~~ c"drLId~cl.~'UtvC IhiR .nl1ulir~~'l il:;;~~ ~~'i ptOUIC!UllC(:d~Wlr'H~i!.IIIT: 0 f plo:1N lik~nilltRiC ra I; I in, Fi~, I B_ Ol1la" ~~1I:Il!I.~'ti'cljCC .is. Ihmlatg~~t clllISr~D'iti!5 "'0'0 IIhcft'i.CCHilblS. dbviotL~I)' d~rc~~~ lh~ aelive ilcarl,)! i~(.'i1cr~lic Str:ut~hi[cs'. Wilhwllle:i~ '~o~ l:ic e lese to kilg; illlli there fore Ol~~.~ Wi!ak 1)1 1O~1tiI11i1 ill {late ~F'~. ~qand :5i(I,;:-OOi.iitdilillhc ti!impen1Itlro-t1cpcndcnl. 1he Inflrkcil! i~'¢liIfLT~\,(,~ n.'OCI.3i1t~Oj~l1rolf'o11,. The ~!In~c ~liiltlmwttoi1 hoMs, fOl"nc..:n."Iil1chi),de--WII,lcr el:lIS!C'I~S:iarg~r. oliler O':]I~, 4D), 1rlt(liOWi'"botu'KI eluster reacts tcmpernunre dependence Ail) inllhe presenec d LI:st~'S wm alse dccre<l:1C' the ~ITec~i\icI"l'aClm~l, n r water ils ~lCi'Cfol"C ml,'il~u~oo 10 Ihm of I~r wilh Olm v:i<lialrau ..,itim~ state . simHaf in C'llcrgy to thrnl cakul<ucd for 'ti'lelrlakcd aceta I!d\:l- a Ilcl~I{lI, k~'UJ" F (lr~callJX"L~<I!Il!lros lose 1.0300 K. rom:cl1trn~io'lI. ~ngcw::rni. 'wt;)ikinS. in cono.'llIl<ic ilydc. with ll:os~ulinc,<ll1lt impael, m~ thc 1iC"H::~inl~ ~h'C'dbgl'i.'eor ctu::ti!l'ing J~ i:s: ~nl>il~Land [he lion and d\::!ni~ily IJCgjOllS. "~'h~re ,11 irl'C'ar depen1Il.1UT~~r. inro,~U!'m1t 'the ,~~,k.tlmulcl pC¢ll.....cm~ reaeuonis Hlll !'Iu,l!chcnhaneoo. With (_it;cro".l$ing deuee 0 r Ilh~roue COll1flm.~1l ,"' fillllCliol1 of S ,1!~ fOflll ,.~ c)'c.llic 1)1'(..'$ooe~i[H! OOIl~pJL,,!}; wu[h Oll-I ,It <l [(:lliQP('r.auufL:,'. the dl.'gp,;,'Ci {1<f dll)';l~.ring in1Xil~~!1trmion i~~ ob,'1('!'1:(,;d, Ol"\'oi:dl>L~!\'~ cnmplic,a' CrL"~'$.•""hidll il1tI'C'<l~'-'$ 'nhl~ W[t,:;. TOg).~!J1il."t with !iDll~_ It iii ;s:bot~knt)\val 'mh,!! :ibnn<; ~;mtl(1k"ItB _~'L1b-"u:'!Il1ti!~illyImlll\L;T C'!u:1rgy ,~fig. 4 E~. ]'C<.lrl ing the energcties ofdu: n.'aC!illH. thlJ~.J~of C(¥JlI:L'O~l1 ~~,hh Ihe' h)'dm.'<.)11 r<l!dical OiI.:nCl ~~!:Qro\~ <lK'f C lusl~~"'. alse te a !l!l!bSt:illnll<l1 ~;cd'l!C~iion orille cn.cr,gy of ~'!llh:;: pn.~'U!11 experi ~~Xl!lt how('\'(.'~:,"nO!WCIiC ~lnl l'hCl TS IFig. 4F). ,uld~l!u~ the h"lffiC'f of IhC' the e.rw'l:Aic cffoct ~mli.~li ..'~~o ,h~ lYIJI.';S i;:o~llr.!l;;i'i !i'olil. lu,'C,ul!II!lc IDr u Frorll~hJ;,' aI:!lUmL.'!11);)11 !tm .1Ibfl\'£), an ~nlJ}('H'!"lll fC<J¢li{ill" QUOlIl~lun ch"'ll1 ic-,d ,C'.:!k~'U~~l~OI1-" !h~ .n qru.''lJllilyi,suh~ t.~mocllu!".l,t:iorlerci I:!!,("I~O .~ l;il] nile mil':; OOn, ... mll.~ w~~re rl."COIljfl,;d ~lC!lL"WLI1l,![}II itlT CCSDrn I~v(;j,o,f 1.l1(;'tJI}' Ih!ll iulcludcz!"'"ro~ ~'II]dw~~c.rt'l[li]1lC(lItlt'.II.[art'i., ~{l;irn ~11,\:,Tgy curreetion y~,cld 911 :;: kJ 1il10,1-! hydrog,(;u!,~ontilld CrnllHJIl!'xc:;; in the cxperiimt~m!; wllcrc -]% uf r~I::!,ois IJres;(;nu in rhe gas mi;.;:The cma~}'lit' ctfc{'i[ 'htlfLt i'i'W!ifeslS iiSdJ in [~11 barrier SJ~~M,r.uirlg,th~prcre3e~ion complex tures. Unlon.Ullcllldy,. I.his qli!~m.il.y '(;,'MU10i1 be iilon::.~ Qftll~ ·Cfvcm~JI bi,l,ol'ccular 1'C21C[tOIl rru:c is fmull dt~ l'S ln jhe ahsenee of water" btH,only a OJ') kl m('lOI-~b3rrierni'lvohing lhe side-beund due to c~usteri:ng of ~le t\'m::lai1l 31 spcciric ~hes s 1ne;TIIS!lIlrcdnor cillsi:ly calculated, L1oCC;;Rt~C i! UYly depend ulxm the k ill,c~ks ofthc :('1,'I,tlCI If) ! .~ I:l!O' ruJd in tum a n.--duc~i()nof anl ill!lrillsic It'OClOCm aocta):ck'lilydc-wawl" corsplex, Thas, ~lC celeul~\1fricu"1lrolJgb [he ~omm~ioJl of ~lOCilkhydrogl;.'n ( COml)h:.~.in8 proccs:ii,Esti:uu.timls 01' kwr [11'01., lauons s!UI\vtllal CV'C~Ia ~in'il:C' W:J!IG: molceulc ("<Ill acccleratcjhc rcaetien by ~iglJi,I'ic.1!~H, b<!iD"- cC~LVi~ibrjmn CO~1"~:m~s CQtlilibrillm 'OOI101."H= 1')(1:110$. Thcwmer mnlecules :11~ n~ Olclivdy inand \,Ol\iC(~ in Ihl,;'l't\,t!:rion (C',g" (tn~ugh bo:nd break[rOlllim~s f'or the I: t con~Il·lbc·... (w~~ ml;!' an idl fi~r ~o\'\lcring. ,~~' lC water cllhmlccm~'an of the reaetien ,L'lllC upper limitw their aCW!ll~ cOl'I'Cclnmtionl L1sing ingl other than iomling hydJ,'(!!,l:lcrl. bonds. \Vc' are ~ aWUI\e tliml. :JIjJ}' gCillCL<llooloc]Hsl,ons JOOUI radical our eb in i~ dm.l s how I hal, the lCmpcrm.mc: ro is due 10 a kYi,,\'CI,;,I1,Q even a removal oflhe or dependence of the 1:~]j[C' cons~an~ (Iuli'!.<l!.ti\ ely i'l !lfil'~:ii:ic !'t1l.~tiol1 b<!fuicr 0 r ~lC p1'1Cre.i,ction \\'3flCr I1IlQ!CCUIC' l1C:lc'lions !!1\'oh ing !~tC'S nnil,.V be 1;l!'II."IlI1I!l11iIII nlhf.,'l rcl.r!1,;x! ::l~lldics of rw:~~CI!Ui!,~1 r~. (:ow::Jmcs wi.th J~ fo!" ~: ~ (:ompk'XC$,UMd th.1t compkxe:;;. Eq- 2hcjllS i!1llUf!dcn<;.t;;)Indil1gIhm, uhis m<lly appR'N,lCh Lmil), ~~)f k~llIp(;r;MlI~ weill dlt":;l iD'l Iurn nrl¢f!;ijl::il:.~ Ihcll{~[C e(ms~:m!" k:iIlff'lioio Ihr \... ~'t1;:n:;'II"~}rM~,, it \V<I:<i~h~IJllB Illml. '1I1h()ughrnllTj~T:':; IJdrmr 100 K._ ~OC ImR~"tI ll;[X'Ill hydrmiorl. Ihe: d}lrlrulrnil:.:ii :31~d ~he ImWll[~ rcht.l.ioll li\'~l;tc It'I~:.C'fing ~100rkdue: 00 Ul'Sidu~~ lJf~uricil;Itl~y ~Im ull~avoro:b~e,I~r !!C'be A~[!mHgh Ihe kErK.:[ic:dlTl.ill.mld Our qUfLl:ilunl ~1IC'kUl!i~1t !~Llunbclt of nmk"Culnli dc:g;recs of chcm[ca~ cll~ei.!i:l!a~J{}.ni> Slrrmgly suggr~uhm [lie ac[ion I[29), [rIOWI!J\'CIi, we do, ,1Jl1 k:ipfLll! Iha[ ttl~I> ~i'c:i.'(IQm o:flhc eon~pkr ...(fillS, 4E.~, lllD '~hc cXIil'crnc. Ibis c ~r~c;l could hlCU:t'lSC the o'llcml ~Im~~~O:nSillIl~ mllC (~lhmloeul'Cil[ is dlle 10 ~~itf1.c Wimer CllliiSKTS effcct ;is IlOl I'CSlj';j.C'I.OO to the :IIO-CI ~lClIIO sY$· lor C]ltlSDCrod react3ms 10 lile Ilssoc.iotiml=llimiloo aitd thm even single \m~er mok.-cuics may ~o'Wer lenll bti~a ,gCllcrn~110\\,'=tcnilpCm1UII'C' reaturc ora

~ir~!~in!he ~h~~'I~L';:OrW<J~i;;r(f~" 3 ;Ilid 4, A ;In.g B~ ,~U nr~(ir~N:liri!,;'lr, i.e., dl~~ difl;_~fl..'!locin ~JI,,~rgyb..:u,\it(;n i~.: prereuct iou ('{m~ pId!. t);iI1IC~ ~ure 4A) ..ndthe Ir.U15it ion !5;~tclSlnli{;ruulli: 4B). is

,I"I~"

;)1!Il~CI,~I\,.'l" or f

firs~. \\,'e ~~ ~I;';M;~

V;j):I\~. k :;:; flw,s. c(l1'I\,':'j,TX"Illclinglfl k

Ihc Inm"pI'm,1

~HOJl
!1l~CT

~0lf~1m,1lC' ql!l,;SlgOn .d:.l!t;mt ~herole 1

01" of

ot1."~v'\'abk c<!Ialylic dfoot

As. no~~d ~~~rn'~ ~ld ~~"p!l1;'S~~"d j~lfcq. l, ilw
~n~'.!I!.,,~m~], [C'11fIp~rnl.I!lln;-"'~'~f1l1,;:n(iI,;'l1'. V1lr~~Ol1:;.hll1:t

clusiers arises,

Al

the

1000ll"QS~ tCi!nJx:mtli!~
tl\~h ~~ .;;'tL.~!';;1N

n~

il~ pri¥~~ip!\;,. !1l~T

dU:5I,,;:p'

COn\; i"~~!1l\'",ilh prC'vif!u~ f'L;'S1ll1i ,~24~ Nc",l, u~ oplinll,iz,r,'l;! till~ g~TI!~'h;tr:i(_'li.~:I' pTmp¢Cli\'~ ( WcL[I.:'NlectlCl,ltk:h)lcl.: cOlmplc1l:i"~ m'i!d fmnu:l

\'i;\L;:

~(r, .~nthe pn~cl]t",C of \V'.il!"."r.i~, i.h..:..,um of I.wo i,;()nlri'b'ul.ioll~., mm!lCly. t'rom !he d~l"~e<!~'d and u~lI;:hl!iiIli;!I\c-.cl :.p.r;:1;;JL:S, ko, ij~~ gi'lli;.'!l b}~qh~~d ~inC' ~n Fig_ 2. w ~cr'L"a" ka,ggl' ~C'~1'U~ n.f Iii,.; ~1lr alJ<...cllcc Or~l:i lnLri n~;iiicb:iirrU,el', I:. exp~cted to

:.1c~~tm,Mt;:hydc::l!lJ~11~r Wal)CT-ai;:CI:tll:Ml;hyd~ romplcKe~; nl!;~y ~llso ,~1{~:;,t ,to ~mrn~ cxk~nt,. A

or

n"S~

or

or

r~

"~"~,I."

r~

Fig. 4. C;1!lr:ulal@~S<trll!C:hl~E1i(A to f) of r'Mdi!nt ,cmlp'Ie'~e'li, prereection 'CiQmpe\l:.~s;,andlHUlislllWi ~tal'eS. Sir!JIdures (bo~cI ~elilg.ths. lii1 A) ~~e ,chara,t am~;$tic poir1It~ ,awlI!~ the ~~ijNl \'Joominates
(F~g..3) ~[s~i'1g '~~~MP2 ~!!1!'~1 f theory with Uli'! o i3il!l9'«(-pVOZ ba5ns ~t W <lild (B) are ~(!' pre= r!lla.~tiotrl ''ilmpl~~ and [he tralmsiHonstate wi~h-

'e

larger class of :~eaclioIIS of polC'Jltiilllie.icvancc ilt <lLtlrLosphcrUcchcmistl,)" Lastly. ,"'C note dUll llL C:l!Hllj!~ic ~'l)leot wm.erin (e ondcnscd Ilhoscl orgJEUlic rca.tl,ions il<li.'> been reponed bC'fb.re. 1.,~lwc\'Cr. the ,eI1'e!ctdescribed here should be dC;lIi)' dr~lhlg.\li:;lH.'dfrom e'i:l:ecffi of m~cllOooll\alion (CllI!lS;~I1~ ionie ,J,i&~'OCimion), cl<lSsk",1 a..ei(~= L1'<~~c;"<I!!)'si~, of O:I.g.Jg}il;n;~.C'~in!l"''' ,~nd 'C,;3~, in \"ih ic'h w;u~r i~1~1;;tiwly i:!mvo:~v\i"d<II:' 'IP;''''I.'~ ~Qn
rmrl!lu.'T l~ .g_"

"~'iml'~m p

drNmr OJ <'Lt:c~pIOil

,t
3.

:1-

tRe:rt~e;i1JCle5, ,Ii! iIId No.te.5

Il
I\t M, i(.

water" r~s\IEClivelY. ((~ and ~[)) ar,El prm~et:lwe ",aI~r-~(eta~.dehy.de dllls<t@!i'S fr~!If! \'jhich (Do)
'(IIU]

\Y<lsirI!enliilieri t:o lbe ret'etlant f~r tli1ere~C~(lrrn.

{lEI is In,(? prE'I1(?action

rearnolm in~hi!!' preM!n(ie

,mmptw:. ilnd ~ij is [he ~r@fis!tioJll ~~t~ for th~

,Cllf '~'I\ilIter.

~hfl' Rl!;;_ lNt.l.:I, 1.1'''H' U'l9'bl 8. ~, iamlb. Ill. 1N8g~mlm~,M. T~kl!)'lloo~~ H. l. ~m. t ~~nill:ikj\l, fb~. Clie,m" ~ lIiQil. 5101 '(191m. !l_ 'iI. V~~!!d!.1J.~, ~,·A\lf$. C~m, ?(i•.. '!lln~9<1)'. " ~O,It C. Dum.~.[ls,;rrm,j\, Am. Clum 5«; Uot. ~as6 (1~92). 1:1- K. iOlll~JJ, IJ<. ~n9.~IiI~.R 'O~i~,EI,.oI\, ,~~i!tIder;.J. Fl'l~, r~tm. 9JS., 5S')'13 {I99'2}.

4, $, A.lo1iD" I. $. IFrancis(n, ACG (Ii'k'm. Nt!, ~~, e~s:~:200!j). 5. itl. !\roll. V. V~idiJ,}, &\Qp~~1lf~. 1001 1930] '(:!L99S:'6- 'iI_ 'Vili~~ It, G- Kj~ef9iI:aatl, p_ (_ llfimi]~, I), ].. OOm~.~on. SOffl~ 299.lI!i6ei (200~), '1. ~,~. r<!!!. IK. f!l!\l~i'IIi, W, !{lemJNl~Ilf. [l, V.', ~'el'5.;m\

s;oM~

G;!I'~~~" 5t:ieI1{,e ~Q~, ll~!i {:!l'lM. ~faiil. It'. Frram:~NOr~.rt ~3G, 'ill. U:1l11},. 'l. 'Ch~!'1~. '5, Il. (jr~~~ C, 1(, R:~~. JI, L ~f~~m<!!t
'Zl?

J!&1I2 ml\ n

REPORTS
n, w, Ill, S;m;!b, A R. Ra'li~~am:~~J. PIlys.· Cl!fllll', A 1,06" 419'6 l2JlXI2t Ii, ~C. H_~', J. ~ IFi'antdi!l:J:\i CMmJiIl}1Ott-m 3, :li3~ {2.oo~. 14, l irre.}. ClreJ'!l. SM, FafJday !'il,I1~~, ,0, ,BO~ ~199<it 9 rs;, K. S\i:1I\i!', '1'. :Su~ili1i. 'to EfI~~ SciIW(e' 1I::U.U7i8 f200.6l. 16, L jI. lftamElIIM... J CilMl', f!hYff" &~. 3692 !I~'1S~. 1; .. IN. II. llolmore.. 1,·Phy!. eliI'PI. a!3. ::I!l13 (]979L l$, '$. ~- :~~~ Ill, L 1'!i:~~,J, ~ ,(ii\~ e{l. 15~6 ~1!9$4j, 19., ~ ~. Uli~n~~ D.,G" ~i~ It lil. fi\r!,mm.. J, Jlhy:t Cll,t,m, 8~.•
]2,

21l!, ~.R l~)!'Wf, Mi. s"lb'hlmaf1, i!JlI, Nif, B. Od~!I!!e1".IP,~at. hoc ('i'!d. mit. .26, 497 U.996~, n R. A'iJ!;i1"!!I!fI" J. /nip. Chfm. Rq. ~iq :t 61 ~19'94t

16~O fl!9a.n

22'. IL iN. CW'i'ltey. V. Sitll~wrnar<l!l, P~ys. ,Q'jffl;l, (/iIem. AA.p. !5, 1(16 GlOO:H_ 2~, Ie lit Al:,,_,!oid\!bo.~, til, MQra·lDi~ it ).ilalfdl, A. ~1!11' 18t!n!J~,,}. Am, Chi!Ill. .SK. 12:~. 20U '(200[1. 2~. IB, D'A!iI~ el oCl~, p .. ~m, '{J~ PtJ~ 5, ],'1'90 2S. T. SipaI!l!li!l1bl!r,;,. ~S.tt6I'Iter, S, lI{!1I~muilll1J, 11iJ., W~~m1il'h. IB, :Aliie~ J. AAys. ~"I'. AI 'lOa:. 752':1 (2:000 .. ;1(>" l!:l ,=5il'lglA:~n, R,. ~, CINi:!~~, t. Am, '~;e, bEl2 '119:)',61 :27'. Mah~'jfiilb ,IJrnd MiI!,IJtI1x1~ere <1'11i1ila~~ en S!:if!00l OriiijtJf" 23. i'\ F. 1!'I'.a ~!,!e'{. Pro" (omo. f~~t :;';9, l1!'j"3: f200.2t :19. Y. M1~_@f.R, lB. Cit1ll@r, J. Am, ,a\iMJ., Soc. 12:8,9594

:KI, ~por1

~oon.

rromlh.e Dellll5dl ~ 1\D1f5~l'!un~5gMlei~&d\~II, ID~m 'rn,lIrin ~iru~ 5011d~rklrKlvufl~'betl!l'ioifl351 <11M I~ I[lFG· GradwifflI!m~II~1 '.l3Z, .a.~ \\lI!!U ,M, 'uU F!ltttk d~r (htJll;KIl~,mEa!lw!ie:" i~~ralll'rul1l' ,1KI!loo!I~ed'ged.

'~m,~,

5!1!1PiPo.rtingl Orn'I~:rm Mat@\n'ial mm ~ lIaIi!.fji"iW'liilll3:1~:eHi4·t7tocl

M.a!~n~ls '~igl;. S;i ~lilIlI 5:2. d~

Jiibl~~ Slla S3
,~reretJ~e!l

,:1'9AIlglm 200~; ilOCt;ptM 4!

10,1 U6!~iemte, lU44~4

II)~plllb~

1f.106

!2006t

LU1.CerlaLIlbecaase 01"1h(! absence o~'lhc sp;oco:rafi IXItcmiml COlltrol). TI~c lotll '.l~C'~pcates ,.-cprntcrJ here 3!I\'l b,Ii,~c1 r on '[!~~' Il~"W2\un:!l1ent~ b)' 'the ion TIl!.'l:s-, alyzcr ... 0lI1 OMA) o.f ASPERA-3 v\mll)'zer of Sp<lICC Ma rs 'I,II\il1'" ane!:! \'!Ii~t bUi~.i5 III OW dry, (!IliIcl tlhefate of Hs and eillt earben dlmid'E.'EitmD",phefei~ on e olf the iJiggest p'L1z11I!9'lI,il'lrD.a,rtiiClI'I p'~ane~o~ogy. We have rnu3;a5,liIu3cl '~I1e-mrr,el'll bn!> Irate' due m tille ~ot ..r \'I'imlli IIIte'CiI!:tj,O 1'1 ter diHer11l1'lilspe©ie5: Q~O") ~:t b. Hli~ seeend ::: 4 gram:; llerse'(!(!Illid ~eil' (gl s:-!I!), Q(O~" ~:Ls. _102:3 s-:!I ~ 8 9 £-1, ,ami IO~CiO~' ~ 8, 1022 S-I.~ 6, QIs:-1 1111itle en,er'9'!{ Iri! ng!! t 1(1113'0to 30.000 ele!'(trtll1l '!!lollS p1ar (Iharg:~. ThieSi@Irates can b,~ prolPa'~ilted backward O'll@lr iii ~@'riod of 3.5 blliltoilil y@:~lirs,,, o!su~tif!g In tiflf.!wlal, r!!m~'va~l of (1L2w 4 lfIlIillioor ,(If !l:.t!Fbomi!l:iiDwf:d@ and ,~ fe'l7f ICE!nti!T!@lte~~ water. llf1~I~(a'p@ rat~ i~low, ~nd ths OM has t:~ olllnti!mllil@ sel!tn:hiir'lglrQr \Vi!'l@f ~r ~e:S~f1Joirs ~!'Id (a~boJ1! d~oddil'~~ores ©f! 'lH b~iII,~athrhe 1!)~~f1i~talrySMlrfai:~ and ~l'Iril@sttg1:!te \\lt~e,r I~SI(ape dli~li!ifil>e~S.

Pb''''ma.<;: <lml EJ]cI"JC~ic

Ato'ln'l}. :111 in.<;:lfl!n1cnl

on1,}{l.;lrd th" M4m' E. .. !il."'"'~ n1i~Km 1/2. l3,. The .p ~MA dfli(;l\.nC!~ ~~I,v~ di!l;'t;11y 0+. 00+ .• md C ~~. 'mi't tli!L;" :l:n<L~ ~nWlpLl$i~iol]cml 00 Qbmil1~ld b)f lin:i.lIg uf the wide 1fI!l~f; IKi.lkby G:)lLI£f'illn pei"Lk."eom::spondi I~~ Cll tl~~ inti hili tiu:d IliIfI!1l11k'S. The p08~lillill orthc peaks as 3. IUJlC:tiOIl oftu~®,

at.

ow i:;.~hc~'lpbr wind C'!'P~ion(l\r~hc mar~i'lI] !IIU1~h(;f'~ rcl~rtIJ .. d~orh(' ~~.i~I·£lfY r or this pi.[llJlct? The "WOO'Dl and \,\\C~ c<I__fly M~U'S"lIQlodd S~!ggCS'lS lhat f1o~:vil'!,g liqy~d wrocr
qll!jI:iiodlhc surface l;~m.iol1 lbiu! ures, SlI<: as h ouulmv chmnlds_ The \'I,ra~t'f \'1',,,,"" ML"I:;,iIill~(:d UDl ~h~HqLlid ;;;1~lC b(iC;:'dU~C ora lrluL'Ch 1.1igi~~r aC.rm)o s~lhcri:c' 1(1npL:mltllll(:' ill the r!l~1caused ~:Y a

When ~hc .~Ql1l!.:r w~nd

lH'lgIlll.'bz(;d

crg~'O!JnlCl1' ,~ !1On'

'IHU~iJlhll,.'lic pl<!!ml. u void

solar wind IS [:'on:no:i. Tire void CD'C:llcd by tire mil,gJ!'l-ctic fick! 1.'Cli.ldqi ng: lI;om 'I'IIC' currems in· ductd irl the ionollphc:ru j;,. CO'IIl\;:(1 ,mnl vllduocd
15). It is s~p...'IJ"nL,;-d tmln 11:1~~t}~ar WilllLl by 01I.n il'ldutt"d J'll1!1:gm:to.:.;;phtnt; NlUlH,:11Iry_ Pre" iOlLls iawcM~gm:i.tJiJl::-; (6) bfliSil-d on Ih~ Iwag· netic f~cJltI 1ll'lef~~tlre.11liCil'lDS revealed in [!~ii!same region 3.00'llndfL~ ~cml~(] nlii! nlflgnctk>pilcol:lij ilou.md!ary !(MPrn, The iderrlli~~! of these tM'O bO!Jm(llilrics is still to be cOlllir~lled, 'fbc void i::1
IIH,lgIllOl(l .... hr:rc p

or the

and ~mI$:S. 3S, ""''eM a!>lhci rwidl,h, were obtained in. la'bornlory and in-ni~~n calibratlnns (9). The !~x:h~~i!qu'C'v.'Orks re~iID~Y exccpl. lor (,00+, \,'!'t!k:h C'!Jll1o.t be $eparn;~cd lrom .0:£1-. Ihl\IIIC\lcr. dllll iOJ1<lspltcricmCidcls (/;,J) prodkllhi.! CO~ detlSily to be ill least a rae tor or tOO lower thanthat or 0:1+ at 300:Ockm ltd gDn, whercilse iOl~(I;;~p'i~cre smflSHli internet w i~h II~ S(ll~xWiIl.Q (.nWe~hu.'" Ols.."lIIl!lC ~h'l~ the whol~ !lux !lssoci[u~cd \vith Ihe
O}:*/("O''' poi,'rlk US.1!Ur:ibll'[(;"d IDunoll;"(:lI~l!lr'nxYMP]_ The obsclVlUio:n ut po:~O(~ iUI qUC:;;Li:orb is fmlll I Mmy :l:OOItlto 3.0 May JOO6. \.vliaich eorresponds [0 l1p;proxh''I;lI~dy eee mal'lllill] jl'o:ar; The Sp.8l.tilid coverage is rcgllr:doo as a.C1Jl:.'>C and sl!lttitOti'Ca~ I~ :>lIIlli'ident '~:mpportin£ nnline I("XI),

strong gl'Cenhousc cn'CCL The

nlcx:dcd urproducc

m1mOtI,IU

of wmcr

the ObM.'l'VCd

giClUlbr1Ii1l!l<,..

logic fe.'ltuu'c'Sis about &00 m to I km orlhc cqn ivalent waleI:' layer (I). The arneum of atmospheric (l're5..s UI"C' needed to l1imili1l.ah~lhe reg 1IIIJ!OO groC'nhO'l.1.~e c!Teet wou~d be ,[I rcw b:;_.~ of CO:! (11. The hm;:ntory or ~he Im;:'j!,cnl

The 1.~a1c.."'C"IX" ne is caklilimcd by~napn ll lied by csca«)i-ng or accelcraied ions, wh [:ell are piing the r~m~el:<!l:}'ion film. dislriblll,ionin dlC 100-o the $nf.lplm~Cl U b1C. ,Fill'Sl. t the [:ons Irol1,] .he m,l!rliilll ,mn]()S~lCI'I:' W]lh (.'11" p\1ene pcrpendlea CQl.y c;\jjc;eedi n,gIltt' e:ocap!Ccmcr~y (2.·~ cV IiiI' 0;>'1. lite di:i>tiill.mdon f~lncllon mc.a.'>urcd ~Oili"thrce icms. aCO::%:''1<l!tion af!;! (}".. ~*" iULd CO2'''. <lkmg dK." sp<!:C(;i::rnfl tmmm01m.s, Gif w,mer an<i CiID'I.'Ii01l.d:i'O;",i.dc II~lC m.QSI At IC<l!;~IIWO l,roccs,'>CS oflilc cs,!;jb\lishcc:J. 'fhru:-c rnl\Clhc Ilick-lip ~m:l>Ccss~ f'i j~wry l~imcgrn:[cd ovcr C]lC'~gy ,[lUld :i!oHdallgl!c. rccc-m "t~ondTM:\lvoJ:y" of c:aIDoI];nes by Mars amI acctlcmtion l-e:;uiling from cll,u;gc ~C'PiiL~l- 111coill;aincd int~mll ilui'i. I(c:m-:J :>-JI) f5.~igncd Ei\pn::~"i ,UH cannol: }fl,;:I; klclU1I~y;l!UjiapPl:«l'1!ib!h;: ~.mounl~ of~hC';iiCn1<u~ria~l'i, HfJ"_lf'..hlc wilh!hc ~ taoth~ o~bV!i~t",w,d cO~~"11Q~di!!~~o 19:1:·~,~MA don cl!~c~f1~hlc:clcCI ric I~dd )',. O~h"'r ,'!1l(,'C i~,~· (1'5' ;.u:nQllll:l pr\.'.,l.llllt (m ml~i~"iIl~ M<II~.·_"', ~I hlln1.I~~! !h~ ;,lA;;CliImu ~llioll ~imlC i,l] thi; fr'..un~ 'l'irn;.d~~!:l~ n:i~!ll:oi,I~ r:rw,dl~Y 011111;0 ,Ictivt. C'lri:i.'"ml O ~~ dl~11,1!i:d In du; i~kTP.l~lll.'tmy ~r.t'CI.riC;iidd_ rlll.h~R J'l'n;:'i~~m.1!; Ihe \\',Ik;r in liln; p~L~~is ~nl(:Lmbi!:lI!jl.- (9) Imv~ t'l;I<'LML~hl.-C I~.l!L"':>'~"lIliI.~'itlilm (~f[l11; {If 1:hL: Cflrl:l.":.'iiflnlCOt)H1ulTI<)ill~ j;y~w·m. MSIE {M~Irl\, $l,Irrlnu~1:y C.:tUI till1T1,I,:.\1 by M,u:-; ENploml.i~)lll Ro\'(:r;:.; c:.t"'JIli:ng ~)~fi:'inl'l. ,]h.:y roulld 'th.n a'h.,: i]1I"- :r.uiOi:\ ot/{)1- .md cot/O+ arc 0.9 ant! 0,.2,. :u-c. ,[.J,. \,\1 hm h!lpjl!C.ilC(1 loahe m~a:aiallWa](:II' Ill'bd 1E1(l('&I':LC lidd~. CiCIII:Cc...'d fIE M~n:l>.lhe xr~ i~ fll... IXii IOW~lro the S~I:l,;;;conli!;,lllS \\,:illh I.he intcf[planec:il.rbon di.oo.idc'J Apart rro~n caUil::;,tropllk; 1\,,--" :;'~I.i'\!c~y. 1\.1)311:1 rrornlll I:hc c..~apc d ~c 10 the ml)' elcc!rk Ucld plane, ~,n.d )' Cornl)l.ct>e:> Ihc ~lilova~ the ancienl almosp:~C:fC by .m ~mpilCI ac,c~.cral.cd iOlls. d1.t' luanian almoophc.L't ,crodes. rig~l.htnld. sY.iit.:)]1lJ, ll~c d,UfC:cI:l.Oll 0 r the dt:~iric d~le 100 IbC1"lIl111liI,pl;MlfI~ n;:iIlWY<l:I,. spunc.n1ng, am;! at the elld ofl~le NQa.chiall. and S-t~,~ I"w-lhcMars Glooo:l, Io.~s of IMI ~!WJ:n,~ '(~nosll)' oi'>.ygcrn [~C'$1iI:lI:illg ~idd •. E; :is cakul:ltod rruFnih~ di:'JOO,'erod hidld~n ~rvoirs. Ihe ~~<JII: whK! rrmn~hc c;:.;.ogC'tl dis.'>()Cial~\'C Il,'ll)Qmbin."lt iml. ic Sliif\oeyor me;'~1 ~mcllis, OJ), At; :.'>IKWI.'il tI'KJSi. C1:osiml is aplatis~blC' idea, to (;Xp~ai" lhe absence This stll.Jy rqmrts the CSC:J~lC n~l~ or inns n;:cci~~I~,'.by II5). lite ion,~"'SCa~ HuxC'ii.are o!rof noLolblc amOLltlls or WOller .md carbon ·dio..xicLe accck·mi.oo tQ c-n(!rgies in ihl.! r.mgc or ::10 to g.1nizcd b}, lh-c li-I:1ickld~e to spocciflc i.()poiogy Oil Mm':> tocl8ij'" 3,(1,000 cV IJ'L~'cil:!!_rgc,lhc carHcr mca~lIrclfl>l:l1.t$, oHlu:: magnetic ftdd ~nlhe w .... region ,('wilcre ke ~mlil1 'v:ll:i<laim~s. or the magn~tic IIC~,d He in dlC or Ih~ csc"'pc: fMc ill ~he s~milar ~n:cr~}' .... mgc lS~Sh h~tuIE d 5p;!«! f1111fSics, IlID:Sl2, '9S128 ~ifiWna, 'WC'!'f! "~ltcmll[ed byth~ Phob~2 mis.....Gn. The i I1hlllC 11IU'J~!1d~~!I:rn' to E j and dli.l~ W I~nit~ S~ ~'(fn1l:li!d'1l1!ildeSpatDte des Ra}llllllil!l'ums. BP~j:46, o~~n.'iJirll:d 00.1-,.;sC'"q~ "~~Ol, WO'L;' 3 'Mrs !,!,-I lnJ) mld ID"Dllr.ldi~~ the '1Jl'Ia~c~"1)I imi~ CQ!1Ilp!II.d;.l~ F-3:102S 'o~~se. f,~~e. . 5" ~,02;1 ~-l (./I ~ lllll;; r;1~L;i~l (HJ) i~in. the L;lliClID' with ~hL::'plml('~OlI) t'li~¢_liIlli; ipll'lll~I~Ury ilJm m:1 "To ~!bom ~:o~~e~ppcten~e 1!IlIG~dlbe add~ess:e-d_,E'P1ait ~m@lrLs.@ nUfigc of 0.5 10 24,000 eV U.h..:lo'O\it:r I i:llli1 u:>;. Ihe whL1h~den,\' lrnwi ~rc.gio.n in:l!idl:=; Ihe MPS lip to

or

or

or

REPORTS
4 (fig. .~ ),Ue !]Y1'\ d'iruilll,u ion CQI'w!~'(iCS "vcll \\'idl !Il~cE·~i:;k~ ,O!!l.'e!!OI1f.ig,~ ~<J!ld1),. ( Th~ ~,,~ !:.;H~ pl~r; Im,1~Io~~lill,,~~4y !J~I;PWl~h ~h~ pla~n!l S:1:!91,."Irirld ;J~jl,L...i",~ IT"!lij(}J1 im;.u~nh~ ~hc )I(~~.! -

M ~)Bwilh

'lhl..: 1~11L::.;ilT~~nn ~'SCflflc

r"..I~lJ

dcqlin .... ide

instance fl11X. '<1f1 reach S" 10; cm_'2~-I, W~~dl the ee lips!;:, Ihe C"M:<1ro:: <UJ:;1!! is ron_her Hm ~wd and i:'. "'1)PJUI~\!'¥lib!~~'YRM 0.: 2RM -4R1<,I'!2. where RM oorR','ilQnds~o <l p!~lettlryilJ~'mi!ln!J;;r {fu;!],,,.ily 01.' 2 i!';, ~l~ !TI~t~(J~ af M,I!N (~g.. l ~~t i", WOrl~l I~q!i 11~ ~o <C!lt~', h~t~!~!1I~~11gn1;:JIN :;;hown !'~ ~Iu; fig.J (l'\'(l!'~h~ ,urqu. o'lJ'<Lil~ tlw Hl'l:ld ~I\'~m]!¥=d.,.":1;~INlJ.l!I~11.1J.']1~I~h. m~I'" b:n~I~~.l.:JI\'q~I~OO illll:>!; ijn (h~ [hi; r:rcllS.ll~ Slll~CI rq:li~!~ is alaout l-uf ¢I~-::l ::;-1. tl:lc C:Tr~C n~U~'S i.~rcc :>pct~i~~ rtJot

',Il;

,ar s-11

QtO+)

= = =

1.6, 1Q,l3 :;;-1 = <I g [35 Will y..;.';llr:-I

:0.,-1,

!\-I

E
2'

Q(o;t) = ~5·W~
250
<l
~)J-

=:;:
If. I!').g

s-~

toOl )'C'l]II',.1
::...

flilC'F) ~\'; \..~+\• = I"z:! . -I = UI ~ = I. Sc'l, TOIli~eall':-1

-I

mB
1

Uc
01re

l]J.to~ OnilC 1iOti:I ~~'.Q'l!C ml!.')S.Icli1!:~\ C 1.0o,..;.yg.cl1 Qln~-liQ(O·~= 0,9 and Q( iot~"Q~O*1 =

a; iii
N

05. The ,muil'! (:tli'~$\il.Uet1~ ofth() ~"I)in,g Irlms.· ni,~:l1'C 0/ ,imd O-j. wiLh :1,!llJhrn:i.\~'lI~,tlfCO;! ", .Altlilr.lugh the mlio Q~02 +\1{)to+ , j:;. :in :Jgrcell~~nl.
~\'iillh~h~ m~in ruPO!1lC,a,hy (9), QK02 +lfoQ(O+) i~a fhcmr or 25 hi.ght'!:.n i~i:(01 :~:I~H!i'd OI!I'L;'~ 1lQ.
llIt::C'I,Ir.JlJL;<1TIi;i~l';sl~~r~~aion 1 in

01

thi~ sludYD~LW(;(m

-~
105 -,2:

O~ + and CoOl+ perrofr'l1cd ~LlItQI~.lMk:ll.Hy. \"hL~ri('lasin (9), il. WaS made Iri'ui~.Iii,:.i1 y.TlH:: numocrs ~ oot."Iilftcd 3,rc,~~mosl..a 13.ctor of I 00 lower rhan allOse from ,Plmoos.J;, nlC! Cili[D'(;ill ,ese3(lC' is ~:l!:,lrem.cly weak. The OCCI:UTCncc per $p~'lIi"lbill is hi.gb [ai, Ica!lJtfive ~ampl~ ij'i!L"r bin (fig, 2)]. mill dltl!s Z)' ~he sUI~ii!OC;a~LH1C'C<l.tllli'u!)I rel<!!~cd w SIl;Ui:1i samIr)Jin,!;lis. ~,m!."jdC'l'Gd low_ The 1II1c(,!r'wiI1W fac:[~

ll:lll("J;i 1.0the i!~~l1!ll!nt\ml, l!:Moun. ~uld ,1L",,~lJ,Il~flp ,io!'!.-",'C (i) IK'I.\ IiI ~lr'!vcri~.e OVCiI' solid '1I1gh~. c -:2-'1 -4 Hi) IUD nll,I!_"~ !:i.C(l'lm.ti,{D!lhe!w~C!l nt :um.1 ('0+, I[iii) lUlU. fulll UijJ&"i Zii(;~:rru~irmhcl~.\r(-en 0.2 +tlll!d :x.irooe. Flm (_l:)t. ~iv) d(:\ illl:uOll ,of the "IIlIX {h;:,c:~i{lI]![.Olll Fiig.ll. int.~lgralnl.V:es ((:m-ll S-'l)o fer aU thre~ rolll~ (0*', O'l ....' ((hi-) togc,ther C!~r(!g,~ili Oil/elFthe r.l nge -0. a~M<:: Yo::: O. aRM in the ZX pta ne oIl' th e [-feILatedl coo rdimldite :'l~sle<m (M:5E). for Ire~e.r!!IiliCie, Ihe -X.'IX is, and (v ~ IJnrCr~ai:rllics in~l.1ii"; gconllt:!iDul foctol"' rorf,ow lenergiCis t <:300 ,.;;V), 111(: oililso snol.lilii'l a~@ ,Marrs (r@d drd,eo)..~ne @(:l~pSie reglio!ll, aJnt! thl@ MPiE! (,fM da'!:h~d ~il1i@) as '!JiV~FI by lne lII1odlel.o~ (l9), ifh:e' white doilililhedl Liines rohow ilpprox.ililil1lile :!ltn!o'lm:lines usedl fOFlIV!!lra-gil1l'g te ,~et overa ~;II u neenai nties h:l the measured ,e.'i'C'aPC l'tues do nm exceed 5\l% (SlIPI:IOrtfu:lg, online: mapsinr the ZY plane, Tille :sl,op~ or ~fuie :stn'",mJ1'ne:l1 115aJ silrrlple lif1ie<llr il1lt'~rlfl1ol~liolil ben\le@1iIHie s~~p~ of ~1iI!ii: JI~IIflBand the h(l~wmlta l e'I)jg;~(If th Ii;: ecili,p~~. lrhe 'Ii~~tiCQIlred ,dQlsn!(ld ~inle diiSI1llays t n~ texu, whiCh is (ionsf:dercd low .inlhc' context or

n~

~o~m!!Jrl ,of the mPPlng p~IiI~,

sueh studlcs,

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.~ ~2:

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11

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av~'r~ged over :;·treaJml!ijnles '(!!lJ!t:ijdlil' the I~dip:~eo olln~ a~Qlng th~ XMSI; dlk~·~l'lQ!l l~S<1:dl~ the ~(:lijps~
{!Fig" l).r~1! lI\1'err.lge~lw: dis:mbl!tiol1
~nd ii:()<l~!(,t€di~ '~M (t08R,M -2R'M

If·iig. 2.. ,l'( rnaps

hnF.dd~e), aJiiildCO~ 'i" (ri!!lihlU.lI'h,erhjl)',)~safle'

of till eo imtegwal flu:l[e~ (~m~O!

,of 0+ ~left},. 'Oz"

is mapped ,ontG' ~h!i! p'lane .A'M1I.l1 ~ .x. rCMJB:RM, l'ii")l~l alf\l'~y. FDF '~~i(! fIned

distrn IYuiliolil, on ty ph;:e r3 that have been sampled at lea Sot five times were ta~e,]'l illllO al((Qulnt. IWl bin$ \Viitl ~~~\'er '~rO$:~ng$ were dbr~g;a,~d,~.,for rl:t~f~!lIC:~. w~ ~b,(!! :sh~'!'!i t~e Ma~ Ilrmb.th~ MPB;r;:rQS<s :s,~~ttio,!'!~~ th~ , di~lim(e x ~. -i2R',", The dirllc;~!lon of tJiJe iirllt:errpLa,netail eledricr-ie!d is sl1l'!wI'I by tit'€' 1100 'I.I'~Ctorr .•

REPORTS
Pr"Qpm;mi n§lllc mC'~~'IJroC! r.I~csooc:'k"'f<lm in ~iIHC, {I]W C<lI] ,c-.:nim!iJ:~(,llmc ~(Imll mfl~ln~ ofcOl.riJon ~ ~io,,,,i~J,;: !mdw<ll~r, t!M{C01:' "nd t!.\lIlJhOl,·,
lo\Si~ ~hmlLlgh th:~~ p~~I:in~lJhlr .r;'.i'!aall!d
(l ....

!,.~f ~.~

gig;:lycm":; ~G}') {~lupporiiillg olTllunek'XlI.

L\MKO~ l'- 0.2 4 1l1lb,')f over 3.S. G:y AM~H;lO) < I.l:w crb'i o,\erJ ..5 Gy :
Usi lig II. h)'brid eede, M{!(k;!o ~4 al. p 6) ealrlUb!i.~t1 the eseape role:; for and d iJ:lc~'Cm. ti:o~r OOJ'L:dilio.l'~. 1ilIe eabuliincd loml escape i:r~teIb:r the heavy iOJ'Ii:1I~O"'" Ei;~~d tl is O S .7, m2;l, s,-!. COllfflfk;ll'Od 1.0 3.1".1 ~ S-I in Ihis ~"Iudy:i,c" the c:lIcubloo and measured rmes alle' in '1.'0(.'f}1 good Olgro"Cm~nl. AL<;;o, Ihe mc.'l"u!lI.'<i escallc !'lIIC,j; ,m; i~l g(l(ld,~g!';~ml!n~ 'Vi,.'1Y ,,~ri~hvRlllD I Ill1l~!pl~Ltlh)rdr(Jd>'!\;.lI.111ic;.l sin~lllhll:itlll!1' ~.>' ,'J n They rro,,'~db jhe rn'l~ Q~O+-l'- 25HI"'~ :il-~ .. Q(O'tl = BH(D~ ~-I. Q!((i![)'tl = 3.Hg,lJ ~-,I for !hi:,;:;.o1<!1f n~iHP~m,ml Jlditir;ms. i'\.~C~1l1 ~~1. ~ be

~h~ "WI,;-:t <Hni;lw ...n~ I..~LrlyMars'" model. Th~'n;:· . Ion.:• ..:;dltlr o~h¢r~!lrK~ chumllc!k;, wen: or arc O~:M.'i,m1ui(lm~Le.g., i~~ll1tC"~ a~mosllh~f1.cre!No\',,], rho~O(:h~·~nk 1 or col:d :pl<L~lll"lbulk (;"C'~p¢. Of .. 'w~l!ra'~Lndcarbon dim,ide fin;: !';IDIft-d iIi nm'li(;kJ'i'

The e:;:linmw~ l{l~al, losses Qh:<!!rbon (li,oxide ;:me! \\ <tIer are n~uch ~{lW'Cl"than tl~fCw Inll1~ ~d ~l](.'!q!1'~n hD <Bn~ Iw' Ilt.[ orco~ g1,']I!!Iin;,~roT

9. e. ~son ~ ,rJl • In'lll:!.~182. ~,:ro(200'M. • lO'. It Ll!]K!i!)~,,'•• ~~. ~,. hrt. :ll:i!';. &13 U 9W!, H. '"" ~~rlijiil '!.Il!l.. P1Qlllft Spau: Sri. 39, 131 tl9m, :ll:ol, S, Itn"bll~!1.e1 01.. ij!,! ~ .~~(fI~,' !I'1l~ .SrUm!lfr<
~~, (!L 1,ll'ii~n"

7, IlIIbmlm d ai,,, r~ 182:,. JU mOOI)}. ().. E. t'!\Il)imm, ,It. lJ;Jndil1\ Q .. r:loiberrg,. M.. Pi!;O~lfBrukg" J.. ~plt%- ~~ 98"j991 (l9~j).

e.

W, I.Wi.

S~I!I!

~~~iii\!!rign

~il K";liie:rvoirs" ~1!..'tI

\;llc (,irn'p'h~~b.:c; dtlu lhi:-. SlU!J~i 'C(1111~;d~rl>ll~~Y (

cr

otraa-

eae ch~ml.cI for the esespetc

space, T.he JC~Clmi:rmtto,~l of the escape rates iUI t~~C'rOill~ of cold I)Ia,::madouds ~bu ~k esct1~)C)or v it"i S~)IJ~" tcr~~~ and p.h(JI~ocilcln ic-a~n!<ict toll s is beym la tile' iOCOpC' f lite ASP' E RA ..J C'.\i]"iCrlment c<1pabi ~i~ie:il. o

S~·12.>l:Il\.20!.l1), 'pp, 121-·.139. U. A ~~0fI: ~~l'11IIIill)I' of '11i1~DIM. '~Bfl~or5 jj'liilla'bl~ 01.. i w~j1IO!1i~g !'!)ii!l;:I1i!l/l, 00 x~,e O!1llin~. li.'iI. ], L fo..;:. GtopII1I. ~J. .~!I. ;24,.~~1 (1997). l5. A. FeWfoY ,,' ~, '«m!'s lS~ 329 ~OO)·, :li6.. It ~O'I), G.. lIt (~i!n.~ur.E. 1~b1lml1\~, IP. ~thew.s:.
Ar'II'I. Gil"J'l/;iJrlikol! ;0l3,~3:3: {~IID'~.

ev .... n.'ll~ e..m~po~.lilo:!'1 i~ ~l qui~c 'Ij,\,\~U R~:l~.'OI;i~I(l!;!d ..

:17. y, 1!.!I.il, A. f,. lr.til9i!', I. V. SiJ(ko~!}'I. C. I-I:arl~en.!..INophjfi, K. i!Ir!." 1[1~'•.AIl,n :U. ~200o!Jl" :1I.8.H. i.amllltf elll'l,." kJlrosUt!i. 9 12003~" li:9. E, ~iliD,}. Ge.Qph~ Rer;, ~j)rl. 11.B:!: U:99,6}. To C'O!1fIPil!l; 'he T!lC'<l!':,wnoo n1JCl'>~vi!h ~hc ~{)l li1ii~ AS F'f;lt!!·] ~r;rrren; .om 'he ESA.~~ ~1lres1i~ ~ ml'ICf ~~;IP('! Ch;Utllds.\\'~ dL..'tIJ!11ll i!ll1JdlICIIW~riTh!lll' jiltmt ~IIon: G~ :ltS, IlIitlat~wrii!':! Iiom '10 cruJllriK.. !lll wpr1led by ~I!i~r "iJiioml. lIge:mii5. w~tlNW:. ~n ;i.... ~I".).lil(;d ,OX)!g.1..~all I.':i>~,~p<; I'<II~ uVI.'t ~JI ~L\I',~i illhk h iMi.'o'I!dl agem(!H .amd' 'Ilie ,llIe~arlmeli'll a-lltlinstilAl'les models g~\'cn h~ the p::vtc\'~ I.U'J'l. The (1'i1Qw" hwtillg tl'i~ enert~, W'eoIMlticJ!llaill' Ilunli: m~$'i\jedi!il"l ch~m~!C<l10"" esc .. pe LCOI~~'iS ~"'~l~ln(l!.t~ld-tn 'be m'c NliIli(!Cl<l1 5~' ~rdl for 'il.!IiIP'l1Ril1gl tJ;ie- maim F'l'ioop!!e' 2- U)~ :>-:1 .on ....r~TOl~C'~dw s,PUUL)'.rijll,f,!,. ;'- ~0:2:1~-I ~ I~igalb:lt im~nM~, .amd(t,16!~ ~~ppooinrigibe' IMA

~c,~r~

BI..~I,J~ I.he e2Lk;olJlm.cd escepe rules are .in good ag-l'CcnJic~n with dre 1lt<:as"'lIn:11 fmc'S ;)il. nhc: ~O\Irur Ill]inhnu ~n llo:nU iLi,QJ'I$, we rum lillie; r:cly ~l~l !]~eealeulaicd :ii!.clmofthe escape iflCU'CliSC :fonhc ~o\lrur m .. ,~jmmm com:lilioHs;, Mooolo >1.4 06) su~elllClllS of the cold plasma, IJhom.chcmic3~. r of. put!dicte(]~h~ fad.o:r m be 4.5 tl 101'111.C0"'" escapeal1[~ $JlI.Iucring C'sCJIl'C rates m!JJ;i:,jOlw:lh l"uture uuss I,OJ1:;", rolle increase, and ~vl" eN al. (!71 P'I'Cdictoo!IH~ ,,",''lJnC'racwr~.ol"l\.~, 5~. Sca.l.vng t~l'C'mcasuredemes 4 OIl. this f:J.(:;to', hmin:gs I ~IC K'tUll CS.c:.lI'led amount or Rll'rIl!U!I!IC@!i '1Ino INot!!!; 1. (, P. Mcl(a~, C. It. ~\:O!k!'l, ~'1,. ~ <1:7,1119 {19.1ll9}, carbon d:io:l!,k_k:w O.R 10 43 ]lloor. :lIm:ili!lJ.e upper 2. I, if. ~rti!lg. f.ONS 94,1 U99jJ, IiuniL~ for WmC-f to 4 roo 70 em..Tli!ld5,\'!,"e (.m!1'C~lldC' 3~ jl·!P. l!l~1!9 9 ,1:11ScienC""!l' •• ~i1!7. 15'76 1'(Ii!l!iI. ~hmlhe :SOI'lU".~~.i'ltliIY,;;LfU,II:iol"l) C<lm~nl inCI'C,~~C' V
~h", ~':ollil1mUid Wl:(l~ ~'IpC" nil! " r~r u~· .~ 00 !1i."tYled 1011 c\',ohll iOJl!tl)' n::!cvmn :rI~nl.tn::r:.;" 4. ~ W. SqU~5 .~ oJ"" :5ci&l~ 306,. 11M {.2004~, 5. tR. ~l.m~i.!l f~' ai.. ScielKe 30$1 un ~2004t 6.. M, III A(.1JWrl (1J,. fio!?mi!0l79, 1676 (19'18)"

aud the bulk/rok! 'p,lflS~~im. "-,,,~.;i~; H026 s,-I [I P11(l1J.p".~.2: d$l:.IbiL!.~C: U.'!'H. At> c~m be SC~fI. 2Lt IClIS:tlllll si ~m:Lllmions, csrimstcd phl!ilocilcl'l'ilie."I.1 er c:olldl pt~ma escape niles arc SIlI.bSliltll.iallly higher l.h.2i:11 rcp(}r~.cd numbcrs.Prceiscmeslhie

cilll~~om~ <!16!'nrtll!,!il'IE.tiili! 5ipnn:l.l~ d~ ~l)Ilntmgn.~, T~~~e.. l1',a,n~. W,;: <l1I\l ~no~edI to:) i'i'I!!iilml!t!; ~, lhi!! tlJi\'i~lim Sp.li~~ ,~~~!!:lI ~~.PiI 'for Ih~u

10 ~'IID~1ik 011il mizaoo lite iIIIill"li e;pr5~ ~1lI! 1II.r r~pW;!~ devet~piog iI ~pj(Je' Wob~ 10 IIii! rs" ~~ ~r~; IE,S!!I. If:i~~nrn!om lI~d ",laML '1iUii!'inlte~pl:dn~;a1i)' ,e1~:1ri~ lief!! iUfii~ ilI'w:n~, Mil: Il.il~~ onille fflm G'LalJal wn..ll)'OI' ~iii (M~n~Iiie-~et'/Ueth'OiI ~Ileeh,nie~g~l meo~eti11i (p,rflKlp~1 ~m~!ili!il~ll'.Ir .M•. ~jjmQI ~ASA, GCI!:I'd!!rd S~(@ flIl!Jhl {ei'llef. ~JIff'Hb~lil; MO, ~.
(Olfl~~

511!Jppodingl rn'I~'I'Ii! Niat@,n'ial O

1fflNr.~iMlJ!i'i\il!l,ar~gill:.oi'i Il!aleriiill5iMldi ~olJi1'llrj~, •. lnW'(jlll3:1SfSallliSOL\OCl f

F~ 51 ~:rilll S;, ,R-!!fer~oM ;!In::! 1ilG1~

,2S; ,!I;ugll!~ .2006, at!:~edl4 1O.112~il!m[~" lU4~)8 IOKW!! bel" 2006

S:ll,ow Earthquakes Coinciidanl wi'th IEpi:s,Dldii,c Trlemor,s B,nd S:lllolW S:~iIP Evants
We rell,orl
011

is (Ill I.he oltlcr of l, s. 'whereas, ~.Iow SJiIP C\"~lll's . (,J -4 ) do 11.0~ mdi<ll.e .m~)'seisl'Il [c Wllves and ro~::lC crusl,l~ dcfiOlTl1mions rllat c0I11i~~lLC' for:::! lQ. S drnys. Drep ~O\A\.frcqtJienty [(;'CI'I'iIOrs aJrld shn,' slip CVCIl!l.,\ ill .Jiallml hffi"\:' ~ delcclcd by 11'l(; d:lim~,:.; s miICtwmk of tile Nati(ln:a~ itCSCJI'I:h Illstilutc for
E,'l.1lh
Scicl:l~:!.'uild

I)i~~r

~~'CIl!.i:QIl

(MED)"

the

very·i.Gw·if[@,quJlIlI1'C:Y

@arUIQ'LJllIk,@s i(1!!:(lln~l1g

~11 lh'e ~anllitiQ~

z,ol'l@

,of Uil@

:~ubdu(l]1fI9 p'l.:Jt@, illl:@'ri~Q~ .1!le:r.g the Nilllk.:'!l sUID~h,l!("tiliJnil:cn@ ]11 sllll!~hw@st J;1!P'e!fI. Se:llsrnh: W~\I'~ '~'~,F!@'raJltl!dI V!l'rf·loW ..JI"~ll'i!'ncy @l'Irlh'qYI{!k@s \yit~ ~li'ijsrni'c mOI1'!@IfI~ !Tll"g~~t~dl@~ of 3.1 to, 3.5 by lP'r@~l))l'!I'!i nandy s;ho\1!i aJ l(lflj~ p'~ri()d !Il'ff ab(iut2Q1 SIe'ii:~Ii'!d~.1I'he se~ Simiciry (If l,i'~ry41l)w4i1'~(Il!tl;n(~ ,!!'arthqlui!!t<e~ (!()(omp1mi.'lls, i'lJrld: mliglrat~:5lMth the aJ(ti!o'il)! 'of dlll!,p lJ:lw4r'@(ljl!'llIU:Y tr,@mQf5 i!l1d skm :~ii !!\I@'Jlts. TiM! (01 flti:i,ciI@ r!"~OIr th@s;~ thrft' ph~nQm@JI iI ]mprOO'@) th,e>d~toctio!'l o'l ndl (ho'mlCOO-rl!l:o'ltil!lr! P '(iif ~l()w @(!Irtnqurc:@s, wI!JlI:har,~i!(lI!!'Qht to imJHe(!S~~h@' str@ss· (11111Ulpdip" Im,~athrust~rth!!l,l!C!k~ rl!lpWII'~i!(Jne s..

\,,~hiC:hC0I11pri:st'S<lllctwmk of -750 hi,glHlC'11Sfrlhdl.y ::I1:i",!lKlgWph !lC'~wod" (l!-li~ncO .md 73, hrQ;!'(LI. h~td ~d~Jl~!1lflh I]~wnrk (.f-lt~'!!) ~mions 17). The 111:i·Il.Cl:;1;u~iIHl!:\ pHn'id~; ,:I :high~lc'\'\.i ~nclu;;'t;.~IJi.i,litylor hn~._r:n;:;1[lu.,:nlto/nrollt\m; widi ~mlll~· iln1p,~j~~I.clcs,~gliill\:> n ~ nn~] li-li£lam:l F-I:IC!.S1mio.l:Is ill:m d~ocl. vCl:}'-lm¥.liil~lLcf~'y (VlF) Mlq;u."ike.'!i wili1nn lhc aC(;rel~o~<LI-Ypri~l a~,d"tc Updi:PPO:itton. ()flhc N,mkai :>l1!xlw:tron zo:m:'ll? ,Nf)}.
(ec..~)

A tYlio-eompo~l~iJl h~I;~(;!ilf;iiliviiyhori!i".JIrlia:l

N
~rom

31, Ii-net Skl~ton. m~\'O'C<lfli.c deer ~-o,w-~i;:eqm:~ICY 1fI::m V" .. ~r~m periodically em -1i()O-~"ill" dme Sca~C1,,~rl~ ,(I:IIl"I~C'l lH.I') is in,'\"nan~d ea,h :~ or.:: and sliow slip iC\ e~n:> jJ 41 occlilir ThCfCis 01100 ,11 Img;e gll;P in dIe sOllICClim.e '(om. rile il!C(:cI!cl'omt:!e~ 1l!~cti.Gn <1:-; li~tunelcffi: ~lfn'l.~In.'nlrousl}'in -1lI1'C tL',If;1silocm ;rone j5) ,~ro,l~r1iesol'l\\~('1 !1I.'t".(.'Ill~y d~i!io.wep.>d OOmni5lbl:l1, m locked lo a~i8n~ic slips. ,It the ~I'o\,,'~~,~ipof ~1~l~'41!~i~lOrOOCWUllulalcd S!Jll.~~ll the 1m.!15,~~inll N:tliDMl Ri?5'!MrI:h 11l!i1li~l!.Itefcr Eillllh5d elll(l!' am D1sa~@~ p{:IJ:iI;iG~ 01' ~11~ ..d"'I:I !;iuW~lcfi{m N :W~~!;.'i in ~utha ~Q:r!i;; 1lo!;.·pkw..-.ln.-q~IC!lC)It~'!!li:1rll 11~n.'dnl1li. ' ~fe!l11l~io!l,3;-1 re!'IIf!O!!la!~, Tf!iI~lJba3@s-0006, 1~~!I. wt5t J!q'iOBn1.. M\..'g!llImbl ,"'<ll1hqllJak'L'1'. willi ~!:;'i(S;l'niC' 11:1.111)' show Ih::-ql;DeJ]eil::~' Ik~1((05 R. ~1~l."Suirig II~l '1'0' mlOll'l (orl'espomOOf'l()e slllo l.ild be ,a~oi!ressed'. E-I'II'Iaili: iJ:b~ ,e-lmf'JCktii:;~ic li~tl~~ selllC' or Ihli:: Iilm)t~iIT dUr.iUiOJl ~@hIlQi~oo.i1l mJ10tiirill. 11li'lg!bn~ud~~ t::~.o::,,;-diJ]:g fl 'f.i.CCuJ."iJi lhi!S.

lk.-reraocm:i~ witill a wide r",~u,,;rrICY"""Sfl(]iilSe r.Ul!£'C I:r.om S I :~zw diirect...cLIl'If'Cnl,co.mlmncnl,

REPORTS
~hc}' arc ~~~.~d In ~n.tlyzing $I1(1]N!:lm~ $10\.\ illip ~~i>I;;ml!l 'I).]! i,:u~cITIjLl m!d f"llel <1J)C {J' ,1~:'lL1tl~l~d ill q1k;llDln!b.l.g !h\:l n~~If~Wfl~ telloor s{l~~uti(}11 [~~! of L'tlljhqI:l1&~(l!}~ 1'0 LlJld~I:>i!lIld the :slfi.~.I:da',(fliimi protx"::\.'" .;U~ ~1~w.~n~i~iOl.l mIle, i~ is .iH!flGr!'l!I~. 10 :id!Oll~ify .d! :-;r.'is.mi~,~~dl~i.;I;il;: l~h~l£)n~CiIl;l!, l\! Il""""iiI.'fU. <I la~c gtip Oi.i~ OCI~irU;Jl lill!: di!lRtC:liLTi!Slacli~'ie !rcrI1e;s. fot Ih~pl'C\liou~~y jdC'i:ua~lC~d .ocqI J.ow~fcqiJe.I'lt.'Yarerl'l(ltl; <"I:i'kl Lltos~ fo~ .~Im.. ::llir C"'~lIS .. !VlohQftwjCk<C ,di3OOll~i!1I:J1j,\y (1!1.li; !I;l:"W'I-I1IIi:1!1t!.; botJ~.:.~· ~<'lfI.~lqj~ua"'iQ. wh~ch w~ ~I~I~ n Vlf' 1.,.';,l1·!'lqtl"~~, ~" "~'llc;:tl.l~;noo n,'.i;:~ii\"G'.!.- 'U!!~iCIIl:Th~~lbrs;!~.{n}. by E:iI.rthqllml.;",';S, wi Ih :!lin~llhlr ot;J:i~r¢IJl,m[:k~ h.,;L \1(.'L.'tn 111l:i1hi~1k!l.:5Sof It~.;;: .x;x'<~l.iict"Qllsi un Ihi"n;l!linn i:;; 3Illp.~ibud,,'li:n~lISW-"_~. ~tt1m...aile! ~i!:lh rlft~qu~tlc.i:e:; ~'l"Mmli;'dI in b.: -) km ~I:(ulfdi:ug 'to :IIfI HCID.v~J~}lut:c i arc ob.'\'i;r;."ed em lfaul."~h~n fiu.l!~bl mel are iden.• ~~Inic CNI)CriI11i;~ (/.~J.1lK." lbe .. ! d!.;'I:!lI'!s of VLf t~IiOO'b<;~"l~mv;1l.-lhqu!i!.k~ (J6~ c O;!!'fihqu;lkc5"mdi~~ribu~l,.'Jd 'Kam"l 51~gl:~.Dy wid~ The (]Ib~"'lx! \\u ... fOtil'iiL'.of VLF ~·ar;t1.1fIuak~ mUl~ (fig. S4.~;~hc;.t\'"~cd t1~~p!h~ ~Ialld!llfd ~ ;lIla ~vimlil1l'i:' rnft'; -10 :t ~ b~'imi.d 15 ~.9 kil'iiJ in tlit:: .Ki.i::ir.r tNrI1~1ic.d by iJitLl~lliluhil'i~(Fig, .2 3rid fig, S3~ T~iIi!q)i;.."Ctu.cr:. (if VLP ~~j.qll~.cs ..t~re OOi~iiltoo Ie Ttik1~Hild Shi!.:ukbi. ~i()~~ ['l~.vc.~. nil-;, wide lliJlliliYOW !wId Mmd bey Ih::;Si.ir.l:ilicc·p.rojr..'C~ioJ1of dcl~ti! ITIl~e {';;,iiI be aoribtirtoo. to the rl..1Olh \."]fii'ilion 3'01. to Js,,}on dr..'fit~1 COlltOUI'S oflllC 1)I~iIC~l~rl'~re wilhr II b.l:oOO~'ak o'·wmnoe ml.ctiou cru~uh!C'd (1:.1£. 2~ Molt()\'cr;.nilcir Cjj\i!Ccn~ffi cl~rJJ' O\'Ci~<ll) by ('j\iHI (~;l.. SS), whlich. SI.~L" lhilli. ~:IC focrul d.:(fLi~ j:;; 11m wen cons~rnillcd I kl\vc\"er. theN:: foc!lil li.vilhlhc ~if"'Il r:d~yof l.~cp 1:m"'-fl'CCll'C'~lIt'ytll:ilml'S. oolltill:l. are JIt'It!gNy cmlsis'lcnt ~~~~l ~lC ~\!IiJd~cl~ng Ti!IC depths orth~ il~~mc imar",tc are hilL~"<l'C\lllhC' h igh~fre'1l(I~I1cywlnrClllCl1i$ illlhc 1;"l'Igc or ~ lQ g I ].,;.This r\.~~![ ~Iggl,.~!f:. <I'lOlh(,'if wpe flf slow
~\!It

of the depth

\ .;!:n;m('Ill

,or the

ocean ie

\-VI;!cx.aruiued the I ~1~·it~'iI: P..tlet. di9t1L wi~hlhc ~~Id lIS~ or VOO@!ilS bruiuilll~:l'J !a:ltoCf':" wilhiu ml O'!.ccraU range or I to 0.005 I rz lU1 IlIn eHbn 10 dctocl l!~~i,de~Hi~od ~11S.-pc.I.·iod se ism lc s i!£imlb rndi! ated n'O~n~hc lrn'1.<:.il~(lnzoae, \~"Iith .J b.andp;l!s:': n~~~rofO.mlo(D.(»s U~ w~ Sl~CCOro in,d~t1.'Cain.f; d l>1!i!1Cl ~Ulo!ml.'1lolli': V Lf 5igmd$ r..di ..li.Id Fronl
IllL:' !.nln'h....;i! iL1n ~l:lJl\;'rrKOIt lhe ~tlun:;~ IiL;"gio.I.1

Im\'~'lh.lqucnc)' In.'!T!O~. These slgmiis.. obtcf'l.'>I.>d al several ! ii"!1C! TILT ~Ind F.iJl!;'! :>liO!li(;Hl.s. ,O!ro L~ idl;'lllifiGiJon nlK~ mdi<lill and wI1i~,d,L;Ull]rmJlG'!lL", TIll;' VLIF l';ign .. l~ I( Fig. ~l an.; ~l:eOUilp~lniL.'lI by \\'a\~' IlI1liJ't~ortk."I[~p :k!~1i!-f[l·q,LI~.iflt')' IrlJllJO!S. The lIi'pI'fIlocm lo'!JJoc.it)1 cSii 1IiI'E}Ii.iJ rtIicir [Ij'iri,~'alliuncs

or dL,..'\;I,)'

~oo
i' ~
~

21.Tstations \~.ildlaI[I ,cpkcmm~ disl.9fOCC eX>CC1.:ltling

50 km is -6 ,kn~'., \\'hieh o'\lcrk~pr; Wb~l P",~I\J3V'C

~romt'

vdocilies hllhis region .. This \irulue su.m:.~t'" thaI ~hc ro~rr'\'ed \lL F ,.;;igllah. 'OO'Ulslitlltelhody waves, To (_!ctt~ VlF seismic sism .•~s. <Iud cmimiltC' ~hcirh}1bOC.Cnll'!f!\. !l:_Y~1Cl11]1;l~QII)'. \'I.'C adopl.Od ilhl,;'
grid mmw:.'Ilt tersor invcrl1ioll (G~iI1i1) mpp.rodih US .f 5~ in which \Ire :Ji.."Smnc ~Imttl~e !~~~CI'CJU'lhGllI<!kc JJway.-. occurs at a .grid roi_lll dl<:ll is ilIJ:l'illlgOO witbi ~ro oonh s:ll~ and time, US~llg~hli!'.
~,ppID<'l(::h ,;cLime!, C'fllliLillunut>
s(:il-i.I.1l!1griUtliS. oomin{;d SOLJrcclll1!"!1tl{'lIl~

from

F-Il(,I.

wcetl.k,l[.Ihll(;'(i dlcpour.n

0_1 horilorn:til J km dc~Jt. flltd l; s ~~lt>b,....a! rid.point SpocJ~l~ (llg. 51" f'kst, we g ~illm'ed ~iI~:I!' pcu1oos!hm 3llCpolCllljn] ~yOOI:llfYmU~ ~m!oo \\'.IJLin scis:illioc \w.\'C> fmm teleseismic fll'td omiinary regrkmH.1 caftltqlmkeo rromlll1 t~ll! GMT~ ,~o;l~nions.Nc.."i.1. using the f-'Illct and I :H-nctlIl T dmrll. we ~Wlied ..he tCl'ITmki IllOI1ICmce.[loor ilil~ VemiQll (eM-!,]) ,apPlm.ch Ul.. 15) In ",1lidh ~lyij:xJOCl11til'S anldftl!Jh 111leciumisms of ILhcna:rgcl cal1hqlll_akc:; 'l!1:e COIIc:tuI'a!ro :'iiimt!~tllIJ:lC()1.L~l.y 10 tile r':nl n i~~ing (lMTI oolutton;;,., W~ d~cl""~Vn:L!1Y VLf !ici.'>nnicCW!l!.~ !;oin~ \:'ki~Ul wilh HII:.l dl;;>':p l~w ..riIl.i.q~l("ilJt;;y !1'l;!rnlO~ !ElK! iilm'l' :Jip ,-,,'cnbi'liml (K,:l;Um,.d fmlllJ.mU:L!IY ~oMuy
[CIfiI$t}1" w~ulirJl] at

IFi g a. :1.. Sam:p~!!Sllj,SIll1Q~ramliof til VLF !i!'ilrlhqlll1!kie at. £Et;[e(;l-ed sil€'.5. Ro'looll mlllilpom~HL'S.a t :!jew;) I~ii~nt't 1lL T :!ll:atnon15 re shmw. The ab~Pf!led ~vav·e'forms .aIJe arranged in epiC!enlr.al dl~t:an(p cm:ler from ~nl estimated a
S(lUIi'(e' toCQi~OI'l, ne

liiz t! 11d'2 to 8 1Hz.. IrtSipl:(ti~~:f. me verli~ ~bers 011 the

1"00 and I~~~k tl'il:IOO$ re;pre:;~nt the ,~bs.e1loiOO ~lla-~r(lrIlli'lS fill~~d

ngh~illldi(all!:~lle

~~W

baflild~(If (U)2
amplitudes.

ro

a.os,

dis~(ellrl~fl[

.2006 (udl~; S1)1 TIl\::£ VLf (i\'I,~n" widl ~,"i:;,rni(:' ~llomClm.llnfLJ:lllrtLlck.;:.; orJJ to 3·,5 ~Iaclocated OJllth..:

bch·~i:kc diSiribuliofl of deep ~oW-rutqLlc'lCY IfCll1I~ along ~clC slrikc of the .st.ibd'Llctillg. P~~~llirpl!rti: ~<ll~kl •.c {fig. 2). Wxlll omnnaryC<ll"ll:iquakes with ilK:S1lHl: sd!SL!nicmOll.li::ni mUWl~tu:l~ flSihcsc \fLIP C'vc~1isareulI1t li5'lcJ in~my !1iei~nic Cfrullog, ,Ve COltlJi'arorrJ<I'Wa.Vi:fOiim or ~lC VLJ even! w itfl 11l.1~of ill'I oro:in.·u.y cmihqtlake ,M tile F-oo IllIong.i~r~) S!ru!ICII'J UMJ ~lig. 52). 80th cvc~n~. hmoC t~les.'l!~n: ~lOll1Clll nliilgnilmS:: of 3.4 and al'mos!. idcn~iCi.'l1 IFi!J. 2. VLJ ,earth"l,uClk'es '~,Stars);~eep lJ:)w~heqlllcl1(Y tremors (drdes.), sta'tiiOll diistl11budoP1,. and Cijlioorll_!!1d cli~timcc Olnd dq)!lt~.BOtll cWUlls hl'!'·~ lmOlif'lelil·t ll€'n~(lr ~y;lut~ ~I'IS '~~VlF ea rrth ~~a k,es;.. The IPlu~. alf'ld dI~~ m.~l'Iid$'Ylifllbol~ ro.pre~€'Jlt. the N~ED !i:'I\V-f1M:.1q~Ito'lC}'OOITi;pont~,..,,, w inti ~i<~!1lik~f,miplij~ Hi-n~t ",nd F"fililt. s:t.l!ti~!'IilI•. r~l)e(tj;v1::ty, Tille dll!pth CO'f!tQ~rlP1Ic:i~(~U~S; the l!ppers.urr~li:~ Oif th~ nucl(,"~ lUI dlt: ~I~C of(lI"02 I.n {I,05 ~1z.1 ~mifC\l1.:.'t. 'Plhilli ppilne Sea plate. nH~ r,eetilllil'91e iindicateslhellNi!iI ~holWn ill Fi:g. ,3. The yeUow lrnl1lll.gle ill>tii ~lItes 'ihe ~i!!\l11L1 Vlf \,"oc~l. does. uiol cl'mjllnWt~ ally ofa tl'ie @p'ir1:elnter of <lin orlriil'lary l1!:uthqlllake \lIi1~ha moment. rm.a!lll'lii!m!e- of 3:.4..
O~

REPORTS
p'Jme i!!t'C'I;J;~nc dip <lIlg!CS of thl.i: '~~;'lIP:Iarn,'l' Q1llping, ~CI\;~',,!1d!, ;jJN i;Q~is~:u \Il'illh ~lC S~9P;:; ~hi;) ~hililP~~~rw ph.llC: t~~ .a\~~(;"d ~l:ipan.\l:!~ ~. ~II1i;1~'il;lnld~Il'd ~c\"imimllS [Un; 14, t- S in th,,: Kiij~ T(lJ.mi n.:gL1;11l.~1I]d 1:1 ± 9 .un lI]c Shikoku [\'gHm, These '\I.,I~I~ Wl1il'~}'dl<1!1 vu;:· ~thql!OlI'kC8 n1<l)i Ot,.'"!C1.!IW n ~ht tlullil;lWl~~!lIg ~~tL,; in1,(;lfi.~'t;, o The :.'1ill]ull'~l1:::tlU!l [IO;:lU~I1.t"C of I;i!:x;,r Jow~ fn..'(WCi1ey I:I'Clmffi 31W ~,kmf s~ip't:\{C~ll~ l'IlI.,> lx.'Cirb LM~'t! krlOOildh\\'cs\ J!lpHI'i ,t.?). These ~tiv.itil':S are roi~~cider.l. oonh :::pa1~fl~ltrRUcl 'ten~~i<i,II,y. ruKi fl1i::i' c1o!hi~jt !I ci~ar ;1 igm~km jja:~tenl 'witCIl Ithe n tlLCI:ivily or deer !ow-Ii1..'l.1uency 1iI~I1!iIOlS high '~~f} is, ,1\ clear Ilol'llha"-l,wam :Inl~joll of ~le )jC'l\mf:cil)i of doep .I{lw-Ih..--quency Ircm,ors. ,1l~d ,gcOdClk d~~jbml<!lion!i b~ a :;.Io\\~' :;.Iip t'i,,"I.;'nl wort O~f\rOO 111 Ihe ]{ui-- Toklli !1.1.l:io:!l in J;nul1lury 1006 U9h

or

C(l:U~c\Hr-alt~i, ,m

m!t;i~yor Vlf
~I~li;

l1I;J]liou

OC!~~II'iorn'lmigrnliK)fI (lrl~'l'C' seisCW1hql!.. keswas (n~CI'\'Cd IU the (r-iJi,;- J J, Tll~~niW'tt~lI)n ~U~;!nlorV!!.F

IIt(:

~ugg~~a1ml

Ihc \.rtF
.i'!fe

C\'Ctn
~\Hl!

and

1!11;(~CP

b\:v-

~mi1hq~I,IkJ;: ~itillili.(:ity is C(mliis~J,:'n~l \~-jll1 bOQ]1

~ko:p low-[rt:(JuoJ1cyi:ocmor. OV!5Cf',,..aLU!l or Ihe £'!.:Y(_rl.:ailJ;:

o~o!:\';lai\'ln intdic.'1!~ <l dQii~~n;:hrtiQI~";hip bt;,'1\VQ!;.~1 ~bI... Bliji! m'CIll"'~ and thea'l;li "'i'ty o[lxIth V1LF ~~~II'IIJitJ~II;IIk mid d",('(i' .I'Qw-r~~(~LIO:IliO;_ljI ..~ IftliliulL'l>, th~hyitC~«riil!:l rhe s~~OOCi.Jifl~il:'l:!iQti lind
jt~a;o:mioii, IJM.~ in [h~Drnru;:itim~ ;ocmc. The ~xei!!lli;oj't~ of \\i'1IAcirnills,C!litjscd bj V1P carnllqll;J:.!.'S appellr hJ a 1)l.\'<IY:; ov~i,laj) wi,tlI~ the l1!t'tdt ITll'JJJlil!~c ofw<l,\ec u<tin.i1. catl~"'(1 by d~l' Iklwlh;qltcnC)llretnm-:::: '~Fig.I), I.~ow(.;.'\~';h should be Ilnt~d 'th,1l dC'(,;1Il.1'o\v ~.rI\.;'qlJt;lC}' 1!iI.'nlm1' OCCW!f\,'I'ji!ll" out the "-~1':dl):)llion VLI' ~'ll'.hqlll'lk...... hi~ !\.'>Sl!If;t of T

;1I!::ti'i!i'iy1Ul~d (I:IC (!J;rorm<J~i~~, 11i~i~

di:ii~~~lct r~NmJnN:!m, C'cm!Si~~~~n~ prJ:,'l;tQll~il.U~~g:; l"Vlf oompfIt~~ o mlI;;"Ilit-. of -:10 s (.F~- 11<Ill!] lig, Slt Lhc-ir Ji!,};'poh'iIC 10 dE .illil)\UOC p«~l> .rl!uy CUfR';sponld 10 1U s,f,mvO'LllhqL1akc ~...'i!h :u d1<JmcrC'ri..~ic '~rne~Ic of n~j)llI('(; dLl!OlIJt!ot1 O-~l Ihe mdi:,T or 10. s, Tlhi- I'uplun.: du:mluon l;; king,.,- di<!1'1 Ig1i1~ lifooji!li,Ifj' ~nt~lqu1lkt.~ i)lm ;JJk>o
L"1CiLu):'

~b ."qIi!lC!l~1tll.'H~ .

1.11(: J"Ir":l~iol1 liffii~-d

lOt dCLiP

k;,\v-

~h'q~~'I<!Y w:n~ts, ~~.s Uti ~OW~'o"l~:; Ihd~ 10..1> dLimt~n is. ,ob\!~oo;:,.~y slwnl.:r tllliiililte .1- lC! 5-dHy dum~ton (Ifregi:Omilly observed SICM slil' events {1 -4),. S~rlIiar v LF cmihqumkes~Jmt occur wi.lull, l
~1>e :nocl.'dKlI1<I1,,), Itri:..~n <Ill.:: liloL!gl.nto 1'I,;f~t'C:1Cin :>'100\\1 cru'lh'llmkcs with low ~I\..~ drops, ~ow n~l]llll:C

'I!C'locili~.!lIld low slip \ docilico,'l on d~~j~u~~ 'P~m)C 11n.."-"ltifC urn 1l!i.~ olIl!'.CJV"· ni:tm~ <'i\1ggt..'NI't~I~11UI~'1: ~~'P'f.-'l>. ofslO!i~' ~,;;Jrthqu;d,~

\\'ilh high l}tl!l;'"llllid

dc""llm ...~rIl:lqU"_'Il1e)'~rolIlOil'S. Vn. F ,!,mt:hqtmkoCl'i, aJild ' :-ilow .\l~i~1'Ic~ CT!'! eceur sirm ..~'mnllo!~l}' in~hc ... s
IrO'll!~iliOT! .zone of !hc _>;'utxlllqc1inl,l. pbl;L,l il11X:dll:ct;-

TOllllOgFoulilhy :;;!lllO:i(os dl,1I irfEug~ ~iis;uniC' \'0loc.ury $1J1Ij1t'WUR; slimw timl 11 J~i~h 1B00i&'OD]'I'\;IImio

flI'!i)unti tiiJol.: OOU!ooc It':!l~(m of~hc dCC;p' tC1W~:Ll:q~~'l~fDre:nmr ~'?O)_, A. roCCl1t PI\);!L'>'C suiidy in, nhc NHUlkilb s.u'bdocltiolil <!Ditcpi:'CipOSi'S II linear di~lribtI~ion of :l.IDv-rreqlllC'l'OC:}' CII:l'~h~~'lakcs.sil1;~Har. to Ih~ di!i;"smcalto'l~ or deep IO\'\,'-rrcql~c~~CY lromQf'S wilh sli.~hl'~Y oiwi{ms POInd :"1 phases, <It 3S~rI
c.>::iSffi

40 km ~llml!ld

dilC' ~.lb~1,;' i~UCrf3C!1.:!u~~d " cO!J1I.~~ld·

inoS~oa zone of high .PoJss()I~'$ rnljo illlloc: \'icinil}' ofn.hc IJmlSilim~ ZQIl.e. This S~S!AlL" tlIac rQlk~wil1g
[\\{'I Uli£Il"J:1..::lmions: ~i) gC!l~r.nion or !o\V-.[rJ.'qL!1;11CY caL~]~qUillb~:o;lIl¥ ~l~"!f slip, ,:;lilt! ~ ii) rf¥mmrir]!] or U n,...won nr high j'Ml3'!;,'JoI1:Llidl:ll;-';S_'i!I1'C due 10 nhe' ITuid p IIClta..;u:l futml Ihe d.ch:",drm.itm n:f db[: ~Llbdut[!J'(l, OCC".illilC crust (!l~\f:nrl'iid:::nJig IhM thn:::t typoc"s f o slITW c!il1hqlmk£5 im!llLdiug low.froQLl!.llCy cfL11h..

liIm;e,ir~mJlan. 5,,30Q!6, '(days) IPilg. 11., Migratiolii ()!f :se1iSm~dl.yof Ib~th VlIF egr~hClI!Ia~es ill'lidl dE!ep Ilow·freqJI,I~I'I;(Y Ilfflm~r$, illiidl ~l'~ dlan.9@ at JOlillf Hlr-n.~t ~.lt[:oflr-OJKZ'H, HlAZHl URSlt arld MGWH--be:lw@E!fI ~s].lmlJar1l' C!~dl25 )arlllJ!(!f1)!' 200.6. lThe t.emlportlll varialtion o,~\illF ea!rtrn(jud kes bed stain;;) a IiIdld 9'ep to~~·mre.f1uell1l'CY tremco I'li. (dn:Je;5) is :s.hlJ,wnaw!1!!J t~€! s~lUitih~-es.tclil~flhlN!st liflei~m~iniS@.t. r€!pr'~'~iltili'ig U\i'il:5trik~ (Jlrn!il: !llJbdlu(l'i~9 pLu~ .. in~e.rr:ai:e" iI1hE!' c:olo'N?d Hne s are reeards oir~he- E!asH'!Jesll!:ompollent fom of the till cha nge;s '(looen/,oo at Iii l-net stal1i(ms, The ~Inet ~hmllS the d~slriibution ef the< VLI~earthqml~e.5 afll:! dl'~p ID\'1'4re,qmmcy tremo:rs d ~rimJ91the' $~iIl1,e p ,~s '!,!fella$ them-'~'nu 1II11[]' SlS!lll@1'I 1[.(I(atio!ii$ (phiS :symboh).
J

~ilioo

qaakes oceurin th~IRLifl'O~tiQJl LOil.C ~Fig, 41.Ihc cxj~lCllloo elf a ZOfIC orhigh IlOllCo-llliid rres~lre ~]lflj' ptaY lin i1t1Ixnrt\'\'illl mle in Ihe ~ilfC~CC of t~acsc sl'ow Ol!l'thquakc5. On:: ("!Qo~jblc ~Qlli'IJ:io or~hc :>U:'C-ss..rela:uu.itm IJ~:> in thc U'alllsllOC:.ilr m~ is based on on aSIJoCli~1 alOO;:l (12, J'J), inwhkh S100J:.gcr oollp!cd Intdlcs of Vl!" eaIIIJ~'lU!akesII,'!; < :i4Uimm(.1t:d hy a..'>1.;sndc slow :ill~I)reg~ns. "TIle ~lIll~l s:ha!I~5U\.'UJSt~of the sIO\\' slip ~gilllC:l~l is \">'C'<J_k'Cl; !I!!.in ,h,_nrJir]i~i,) ;':~rl!;,'riwQf,i l:m.. 1!,~!h!\I'5~ ~'",niKI~;lkc bI.."C,~~ u.hr..: hiJilh :pol"(,~,n~idPL\,$UIlC C"ll~,""
n:;x!ll,t:II:G1lill QhClrnQJlmi~ ~1n_;'&'i lh~; OJI

pl;.u,,: lI1t~11illP~~_

,I

~fEll!'; sh.t1,~::;tl'\.,'i:>,i; l~tdli.J,~y dit~ ~li!Ibd'Lctingplfrt~ .R."'<1'chesI.h!.: yu~'I!:1 :ml.';:;s Qr~lC bulk fo.r~h~ s!ow ~Iip :5CgJlncTll. the :::lIow slip C"VCl'it s[m'io in d'K: tmf ....,]~.OC-)Jl, mll.e.WI";;iiI~hc sh,~:If :;~~ irac-ICiII:lI.."i> oil~he ]mldlCS of VlF e<li1hq!lakcs, dtllc WIlle (ioC'liIJrIiCIKC
or~h~ »10\\1 :;Iip C\i(;fll, i.!\l.-~ rliliC:I,I~" ~\Ab"n!h~-ln1' I'tlpitll'C f(fi.(;'I'[i,...: iCm~'l.ul.'lJti\.: S.tucSl;re.ada~ 0 iI.Il~ yicid sIre..."S n.;dQJIceclby Iho::; high lloi)re.:fill~icl pilCSSure. This iIl!.lp!Ul'C resll.llis in a :low sln.'SS drop ;mdpo:!l~ihly \Mao crack: generaUl'Ig1 deS'p~ow.;IJ~,LJe~nmifl~'
,d'e>ep Ilow,4reql!lem:y ]!I! s;o IJtIiW/E$t J!~pa!'!.

induc~ dlt: bclla.\.olmora VLF carlill'lu<!k.c. M.my ~IC'\,\, InkIOCfOlCk;. with shear :;.I~'!;ng.tBI "Cakcr Umn u
~l<!~ orofdl~!arymLcrooJI1hqlll<lkctOo andsflLI~c ~,~ Sll1l1lkt '!:i~,UI1 of Vlf' ~ow!hqLla~~. [il,1l rm~)' !l1,P'lU'¢; ill Llh:c 'lrdINllIOli, :.iDlbl.' 1~ 11 n:;...L1~lof UlI;' i!ooll sI[~';);_ll .

Patdlru 0.1'V1LF earlhqllll.ake

lFiig. 4. Sc:nem:tk ~f the sensornic: SOI!HUl di~~ribu~lClIl for VU ~~ttHliUlilkes, tf'@m<ml, a!'!cl $~()1,1i s;!Ji.p '~\I'~!lit:s at~he dow!F!dip' poOnih:rn ~f '~h€ :SlJbdll!('tn~rn <W!1:~

chmigc

!truLllbti

Lhe pR:n1l!: ~ likrutet;

IJtJUt"(,..J b:.'

Ui..:

REPORTS
IDlligmliQn ::;:10\"51ip,. The :iit~I:>t.~iliol1 (If a seismic

~igq1;l1 llem each ~~n<ll! li:lit~lre;rii,I,,~tlk the- Q~"'OO ~~L'l;~~l'lr dl.;~"p.!:m... ·ff\;-q~IIi,'lIlt;)' !!!.1i,'mOp;, Low~ dLiFII.1~ di~Lril~!ii.~O:ll rrul11il I.n tID 4(l1 koHli. \."IJ1C'J\!:uh~~ .IIC ~[r{m_gseism ic refm"1,Qrn,,
~1~lVC1UM.C'xt""''l1:oci vc

fnlflrl,li,;'I'!(''Y t~ln~N imnllc C~di,L

!lubdu;;i~!I)1'l :mI1\i

!If!'rli!n!nu;e~ ~nd! INotl!~ lL IK. Ob.llr~, Sd\e1l'a'2!l~., 1679 (2002). <l. IK, O~!!IF~He, "'h~, ~. '(~m~rni!.llJ\ K, 1!:<!~~~r~1 ,Get1,!11l;ir$. ~.~ Mjt~l. [o[)'.m9:lZ1JOoIiGI!llIW9>111:! (2004), 3. lit HiOli\, IK. Ob~lra, fonlil Fl'afitU Sp~(t' $·7,961 Utillll~~,
~. It::..

Obar~, If!. Hii'o5e" re,c-lO~op;!1pit5 4,[7 ,33 ,(~OO'6.l.

5- R 0.\ Hjln!!lition, ,!,w.amg', 1ll,r.;.Ih:!fIIB,

I- G!rop~

R~S.

~~~gg""~ling ...... !hci., i!ii~IU,:(1) of Hukl.04llii(.\'l~ !!ItCHQ(lI';s, :.L'i<O 11iiI11YI'M;: 't;'3~L'1I;,'!dl1)' II.hc ~t~J>~ t"alllng,l': nut);idc rh,.::I!I]lIj:;.i~im'i ",[}ltC du~ LO iii, ~,I:ow );I~p C'!,iCll[. bre~ll!liCmJ lliatliiW mig':fl~IO'i1,orl.lt~trel1'iloli~i'llnic· II)! and ~Io,w:l:~ip o~.I:'Y~ im lit..:: DI:>Cildiill :iub-6 dUCliol1 :wnc I 25l, ']'he n't);lliu:u;n£, ofnN <mb! deer I~ -.frequency ~re,l1lOJ'S, ut ,Oilso VlF cafflKJum:;; mllly be ullCfi.llto b a..... l!he ~t!ss O~ tile: n!pt~u'c ~lC' ora 11I!ICgmhrulll . ~)t C'.mIJlqlU1l_k~. ni~i'ol. because du:: :th!.;;lilf !'iu\""SS ,nn ttJ!lO o:!~ri!..'i" of't:Ir~~ n~g<ll~ru~. ~;;mlh{IU,IM intu'~t"~ p'na.}' O)IS~ [\,'"i1uh of ~,f{~lI' e'Ll.1ilq~I;l'k\'';.'o; {!ra!l~ si~;fi (l!C(,~umng

61. 8. 9.

:!JOG,IU13 U99~}, I,!I, J!ltJdo, I'tc.'kiil~pia zs, 69 U97!5h Y. O1ad\!! Ill' rrrl.. ft:rllh I'fMrm: ~II'OO' S6, .'1111 (~iJM, :It OIriira, '1(. lao, ['Gmt pr"JIll".r.1~.f! 57, ~.n. '(200M. It'. !~o, t);, Obi!JF~, 'CiI\"p.~p. iIII~kt(; ~3.
'11'.

lQ.lli:i1'NlO~~)$I:t'J: :il'll. ,It O:b~t.!JNd. ,~

(200J),

ro.

lQ,11.();!9'/ZII05Gl..QZ5!27@ (~[)1!!6). Uo, !t, Obi!,~, ,f;eapnys. 1IIe'£,tett ~3.

(~()(ilf,),

.~ [J'l:YIIIi'fi.'6.. 'iI111M3r1.];~~~1.9~

19. S, Sd:me, lit Obarill ~ :Him5e, IP~'!le'f ~eDtred ,;It 11i1e: )<i.p~n ~enc:e II.IrnOCIn MW~ng <!!OO6.,(hltl~ J~1iI, ~il~~' 2006. 20\. K. ~SIlioml~tt Gb.lfil\ lit ~Ol rffim~ 420. ,M5~OOtil'. 21. O. !lS'!lI;:U;,-, G. c. 8!;:1WiI. 50, kI~, S. ~b:ml!l", Ngwre 441 lL8.S t;l!lI000), ,22. v.~, H. 1(a!1~mori.. iI ,flm\f)j'[]I!7~ ~k1~. n.~fl!1~1 P; Fiidl~r~~ W~ ~1i!:~[1 ~~~~~ !.rn~,. 1N.~~inimg'(II1. DC. iI:.9S:t), ~, S,'i9-J;;92. 23. f!. b 'WtilQm, W. IL. EIls'nol1i1, J, ~~ ~, 78.9·527 U9f~U, 2~. H. ~o ~r(1[,. Nalwt! 436, 841 {200~J, 25. (ii, IW!I~~ Ii{ 0!r~~!1!;" ,K~~30(!,1I.~..2 t20!U); pv1l4lille!l 1)1i11~ (j ~f ,(109 (1!!(),lU6lMien<e.l0:llll1an ,26, ",~.~ nlilli'. ~~~IiIi\ A. HUlko, and ]1, IRo1O!~' Iri!ilrlM" m>e!l,iiI<~ cMim~nl~ a:rnd diliC~5I;i;o.n,

at Ill:.' ,~lm"'~ldlp :pmtocm ef tile $l!ixlllction .ronc, VLF ('af!:i~q~IOi!k~ o.alRO y.~fid imlictulH'S for ;I!'C
Uf;,111

lIZ. '11'. Uo ~. ~I" ~hr" j,. 1111. U7.J.I]17 ~:Z006~. 1!3. lit K~IkM~ 8~1I, .(<iJffflq. 1M M.l1I. n., 2'1." (1'9'J'8]o, ]4. IF. T~jil11", C. M!lcgmin, [l.$. IDifrejJet, BI. ~~<rI~i!;;i, ~Jt ~~smg~~~,Am, '?2:, l3~ ~.2~2)'" lIS. Se!! :l-up,prA'fimgl i!I'I~l!mll.011 .SO'!I!l'Itl! Onl:im~ ll6. IE. A. 'Ol!:.a~ L 1'1\. SIe:willil:, ,f:mil PW1iIft. sa: 'en"; S1" 7li
:lil_ lit

5!1;1plPo~~i!1g1 OI1!'ll!'!e M.neo~O!I

¥MW,~,il!iOCem:l9,Qrgrt:9Ltcomiei'1M'ilIllI'113'''',5~mCl. f.!I:Il~Jf!llI~ood ~,\i!1lI'Iod~, Fi!Jl;. 51 t~ ss T~bl.e!il!. iRBlereo~e:!i :2.8 Au!lli~t :i!OO~;.!!IOC~p'ti!dlJ IN_m~~ ~OO~ Pu~Ii~liIed 01'1'~,~ 30 N:wtm.bI!r 2006; lQ,l1261~il:ll]te, ln4~54 IIi1~tllll~lliislntrilXlt1~~OOI 'When tiling U1i~ Il\!,p~r,

~"S!jmwiU:ril !!~~!(,,-..;CtlIliJiLitm or th~ ~p~l;.I!'C .;mIIL;' ,1IJ1Ilir'iW!UI!,;,d~~1r,":!Nn1:lrU;>1. (,JJ1iIlqll!lk~-

USIa;!). Shium~

HI,SOltrI, tt

Ob.ara,

Me OMakt,l_ GMPhfl,

~s,

'JIOI9,UU0ll9~21)0'JJ600nI'4 ~;o(l4), J!R S, ~ira ,gj {Jf" 5~~ 304, 12~'5'fl0041

S:tr,01nlg Rel,B'tiollnlSlhilp Bletwe,en DMS and the S,ollar R,a,ldia,tion ID'o,s.e over 'Ihle' Global. Surface Oce,an
,Marilile bi)ogel1ic d]m~lhytS!JlUid:~' (tDMS) ]$ ~he main iIlaU,Jra~ sO~rI(!e of nl!)rp~sph,eriiC $lJllflJlr, ~\lhikh 1m ay p'~ay ,[I It~y ~ole ~111aud fa'.!:rmiatnm!i iii n~ allbed ®, (!'lJer '~he Irel'llotl~ ,orre"IJni. n rroug ill <31 1J):ila~.dalta d '9 illl1iallysi:s, '~li! fio I!Im:ll ~hi!lt ,Ii)MS Will Cl'lil'~r,atn a r1l' M ghlly' po~itivel'ymrlre ~i!lted \llith~he solon radiation am dose lin tine L1pp€.rr mixed lay,n o~ tine epan ocealill, lrrespe(tillli!' or !atll!!u:le. plall1ikton biomass, orr tempe'ratllJre. Thl~5 i;5. oil liI'ec:EH~SiHy'(iondiitioll foil' ~he' fe;iil!!ibmly or a neg'l'~live fee<dhad~: 11111 '~hkh ~.ighl;· <IIth!l'lu<'ltilllg D,MS rmI1s.:s:10IlS areilll turn dliivell ~Y' the. tlight diolS!:! n:!'(!!'iv@d' bl~ the j)eli!gk ,em5ysL-em.

ObSC'I\ralo:L,)" loo:lIed ,(andUl5 associated

l'lt.lOynm

:1t

41 )fN.

2 43E

ln~ile

OOflsmlln.arth'1o'C>IMooDtcrI<lflcrul. \Vc flOloo.Dhmlhe li~ht e,,'';!'U>tlfC an idealized ~m.cr 11tlltick:

eeanie biot<l :influCI'I,o;:cllirrmne im ~lC Ilfltl,g, \1I./l; \'>'runed to c,"'rr,op;: ,,,rhClhcr D1v~SOOlKerIImn by ~mrillg Ihe biogoxhemkal cycles tr.u:ocms m:e ~i~lkcdl 'IiO 'eli'~I1Clagjc ecolo/stcm ex(if clcmcllIls, C,'i8(,'l11! iaI roJ' E."!!nh-$y~tcI''ll ['l1..~'1.~Iit!!:O;;o.llin':~diia~io,n. A 11X'lmhl~ sm]1lp~:in,g(If 1"\!!iI;'o; PM S OO!1-C1I,'!1~r.Mjgn..... ,;;1..'" \'I..-dl <1!~ hinlggic<l' n.mcli.1f~i~lg'~<o\~h§Lo;; c. 0, N. :r, S~. Wld s.J' U j) ,
;,!!:nO~ll~11,; :S;l~O!1 \I.;'fl'!l by I.i~i~!p.n,\;in~ ,clIil11~[~~lI;:lv\'c ;;111(1 ri~y~c<L1 v<:iri1lbk,,,,\\~~ QPl1d~IQlW ~l)ri!!1il2.00.3 ~l'iiQ;\,,'illil th(~ ,ullll.:1;;.~)h~r~ (_grIi'Clllm~I~(,'!.~5~"£, mtd ran £!W 2mw ,ul 1l1~!l1,1Iu;:..;;. B1Ly Mk!(nllJ;i~1 oxOC:iflllt 1U1~d~:ight :"a;IVI."llgCJ);, alld rro..""1f~IK:al !lJKi ~.I.e't'll.'K.~ p:rt:"CLtl1\IQr.;} (4 i'll Oll(~ of i.h,.'SC' !!i-I~i!S d imclhy r~ll~ (DM'S), \'\ih~hrcpreli:(:m~~he ~]Jc

disselved ~ub.~!m~ces H:~ld u~ot cmi.Y an I.he :illrfhoo irn:mtl,lJ,KCand its l!11{_1c'J:'\,\Ol!~cr attcmll<ll[Uu, bn also Oil the d~l14h of~hc mi"'t...o .i@ycI" ",.jaihin \'ihich tlm~ P;lrlk:!~ i~(1oI1'iim,;,d. 1111):'0. we C-lSlilillO'ltoct d.;;; dsi~·Y.~~\,(,'i'I~d :ilIk9r t.!~ii<Jrri[}llDett,~voJ in~h~,; uppc:r mix.c.,o .i{J.)fJl:1' (lJM Ll. or UML to.olnr radiflltOn OO$'(; ~SlRt:rj, fmm I1lC'Jt\LJf\.1d dlall.'l. o:f lh:c dfai~yavc;ragcd SJllE1oc~ kf:ad~'Lltc~~h(; undcrwmcr Hi~ll C1I:!nnlJlion QOC~l'idem., aJlld TIilc mixed ~aycr dqJdl, {MLIJ) fln A :lli~K"u;Ii\:g~-;f>LUIl ,lIml),:;';" rc~\~~kd: than, dl,lliDng Iht' period t~)!ul],'1iIK1d, 'l.11c S.RD fiCOOllJll00 ror 94% of d'tC variance of mollthly SUln.1OC DMS COllcclltrnl·im1s t,FJ:g. I ~ Ap;pIi.'1:'a!ion of~hi~ aml~Ys.i~10 atnimUllia'1 (~9fJ2:w F)!:]"!} lime series DMS concenua!ion~ at 1~'lydm~m~ion S injhe Sm:gmsso SC~ (I,n {3211J''N. M ~ij'\V) ~~~nd~I~~'~ :5<i~lil"f rt..~\d!~. ~.~:l}' ..~~ ~rr.Hdi;L~1~~~ ~'l\lrr V'l4I~\I!lJdi,~B~I'!'~l~ld!l 3S, w(;'!] ,~ M!".P, ,U'J~ ~xtin~[Lo:FlI l,;oGlf1l,~i~~'II'-' ..
organism,,">J depends

or

or

Ia~'-S'i:

.!lialur.d

~)I,In,):

m'i3jO[ prut'liIIr.;;(jr hygro:ito,p~ {i.e", dOl!ld~~rn~'1ilj® ~r1k'tcs, ill. ck"flil. ani' oVIX Llt~ ['(1i'1,o[1;; OOl:mlS j 4, 91. l!tciw1 ~tln~ [0 rcdnec lite ait:liJliiI11. or s~m"md;im~'m tina" cm;:~lhc am~O;l1filtCi\:' andiiS <lIA<so~'If.,'d the ocean. A 20cyear-ald hWO~h~i::< by UrJ) posl!~<lIoci ~Iml.mru-bm!' phmkWt1!. ck1i.ld albo:in. .l11d ~laa' rndfn~i('1Jl am bl;' 'c:m~nlc(:l'i.'i.l~hrouSh OM S~lrodIJC1iol1. \'c!ll.i IJl~~!l, :md o,,~ clmio:n i~~ a rM~h;u:;k iunl;,'i!:~1.Clion: who!1hc:r thi~ riX\dbll~k
Wt:M.I!!~~ be p-rnli'tiv..;

or

of ;H!hlll1lliil~lC'lic

SlI.~f~lr mocJ ~I

e,

I
;2.

I~g l. ~ilil,e-ar 1fI~~@lSsimli (1:1 :: 15. .. (2", O.94} of s~rr3l(1'l OMS oonc;enctreJtion~ WI'SIl!:!i SIRIDin IBtart~· Btly ba$lalll!l(lifUiln~s~ Med'iiterr.i!II1:ea n). [)(J'iS ,a,f\l mo~tMy da~ Clh.!lrin!lJh~ t IP~nQ~[!ror1il ]al'llJ,uy ,2003 1'0April. :~ClQ.4.IEmu bar.s repreSient S~fIi' darri!il dleviatiorn, of tw~ ((lrFls.E(Ulli!lle' f.ilImlP~irn'~ d<l)fS e\i!dl m,onth,. ,Pl.. Spe-artmalil ,C)orrrelati®:n a~~Y$i5. rof the ~m~ ,dataJ g~¥E!S afii~nilikilflt 1~0:5ili'lll' (!oeHicielmt p =: 11.75 {P «

or
~r

11J~g,,~i\'~

W,11lO1,I11!i;:(;lll;lijll-

0,011.

.Bi~l'Celoo~Ii' 3J-4·'9, ®BOO.) BaKe[on<r, CmiAJ..mi[l., )ll18in. Eftlfl,1I11: sergfo,valll!i!1i1@icnil.cic.es. CkM.VJ; fSirno@1~,ilR, (sit:.~ 01.5"

Ip~l~!l-tde Clilil~es d~

«(510. P..~~l!!l ~,~t1f!'1 de ~

20' 4(1' 00' 00 .~ ~20114(1 1 001100200 :220240 2001260'500 00

SFliD, (W 1'Il~)

REPORTS
measured at the IkliHU,ldiB_AI:lrollic Tlme-Scries SMl,y z;;~mi(ln rrellt~b}' t II ~ w(,,'11>l used ~o c;~ljnm~c ~hc'VNU,. sob~'!'<I!O:i,l~~on ~ID~ tm !h~S;:U!l11~1Iy!l,~1'. " OMS ""[Ie.. m~,l'ii~IITO,Tti~L~ ;lrl,Ultr l1 i~~ S.IR[) V the
l

cm~ ii~;j:J\'<li~<1bh; ~III}' ~cn~:'! di

fmm tile d~~al)J.~c.

C"Xu)lf~inb~ S:!% or 'UI~L' wlri;,ul!::c' of :I:nun~ihly .. urs

~hcc D,MS cm!ool~niO:!lS

Wig. 2). ni~~eon-

BO}'4X:..\\II.nn~~,gtlf ftI al. ,(Nt} !\:-';;~!Hly oon.."'1¥l,ltl~da i;:('In~.p~h~'lsil\r~ i;::li'.l~;,IIttololi1Y org~ol),,:d MLD' bll:;;cd 01] 11J1~ thlllll.:j,.OOO,OW ~~~~lp"~mtll~ pmli~i,.';Sbilm.illL'tI bi;UWtWal 1.941 and .20o.:t \v!..~made o

c.;m:ii;istcJ1~ Wilfldll~ IOC<l1 time ~'11eS,a s~gni !l-!;'.~n~PQSil.i\'COOI'f{.lI~i~n\\r~ fQ1,lnd between

O!l'!,'q;I~d

";~lr1~J;~IJ[V1S i;:91~'I,;e;l!li;!";l,.io.ns;!md

sl1ld9t~ i:mltliilit~ ~.I[1) MI1si.derii'iga lrilIuh,r~,i:.sien CocmCkil~ oro.s thziI. is, :run ~Ui1rlil.'i:SI)I~Cfic lhe Of~iflal p.ll!lnktO'JHulrUI'.cllimflte hypGlh~ reduetien by II half ~liJ~ MO'IIII!~l.)' globa~ n",!tii n rSJ:tDwcrc obDa:il1,cd room I hi!~~'firentcl'lliml.Cd esis ,pOShlfll!fI~OO Ihl<f!l are3t11,.r1OfY ~nc~mh'e) feedooICk.\\,'ollid OCCIllf irpos~li ve ch:lnrg,cs :~n slmHI~h~ variables: in ~l~e:l.ID:l1C' WHI)' as rbr the locJI s~ll!dics r a:ndlor ICIn'IX'rnllwc caused increases in lu~rmll)i un. For chl-a cOllC"Cnllf'IIIOmt :Illd SST: \li!C' used ~rodlll(:lion anit! a"'Wlcimod c:im!!gcs in DMS, piQtr~ :><.l~J.1!c~~'fi,~~'d e~.i!l!l<!loloWes ~II X Mnnnh'l.y lim" ~~i .~ic u I;,u'ly u~ low Ii!lti~udl.;"~ }.i-iiJ'i,\I',wc!". mJlll1Hh 1)1 ~Ii!.lclil1,ddi~!riluJlio~i': nr OM'S. chkl. SST. ,mti dm;c DMS 1mo. el'liort)phyll .. ((H-,uj '''''lIlecj1W'''l~nl:l~ SRD ~Iww ltiml .lJMS Ihllm"':;, ~ohl't~·.;;rcli'lli,o-n 'toI1coon:rmoni)' l!SCC~mensure 01' blom.,,~·;.) \/,I1CJ1C much nIDI'!!' c ~O;;w~y lihoul ~l (b!lo,w.~ phllk~on
:iiE.LldY!1.il~

si~"lrnl with ,I reeeat 'W~lrk U3) KI~~:lwi~,gIhlll ~hLl rli;'t biti!t'lgiCiI~ ]:imcl~UCI~OI1 ;Jlld e.rnlt~nml~ KlI1I o.F DMS ill lila: UM lWlL~ ~lighllY COOCkLl,"1dwitl'i Ih~ lilwdViolli:1 r~.di~tkJ;rrIqUVR~ dese !U Ihis. i>m~le

use of!h!ts.di~ll!<1!Jtl;ll'Qg}' \,vi'th a modir!calioll of h;: ~~dTIli[io~ cri lC-,t]on(il). The d<'liIy <II:v~"lglf.;'d
z>olar irmdilar.r~ a~. UIlC' 1l:1f)-(ilf-lht,1-alll'io~phcrt;was c!llcrml:l[~d 1)9) fll:id cml\'C'tlru il'ilt) eeean-

SIU) lIJ~n~g thcf!ld!~niolrl tlo.wrnrl~'1; tfi~."I). Ncl~,hly, t~I~:K~ :J :.llt.ln!!j aiilllilL.;[it}'ooh:V~"4;:n is ih..:: ~l(lpc ,md the .~I]!X."IjO\;p~orr ~hc glob;uUy d~"i'"ed lintmr cqu ..lirm. and UllJ!.)lC [lb!;!i ~~OO tl1L; S'I!1;l.:;)Iz..."IO ill St.,. (Fige,.'~nd1{l'I;I~l tM~ dalll ~t:!llln Ihr the

lh~

=n

va

rllll

1'l(Y.lli!i v(:~)' om-rolouco bU~r.lIJhcl

~~'10\.'od .

o~

bi~m,r..":ii orh,'ml}I,;'U<llul'C Wig. ) h

W¢ d~vit!cdr~c ~l;Ir[-'l-C(;orlh", .g~nb~into J:N bOKC:;; or HI hlliLi!II(~ [)y 2.U 1'u!'1giIH!ule_ A\':lil:ll.:!ie D MS 1l:l~~z.ill!ucmC:lIll:S ~Ir:rdcfIIlcul{llC(iI SRD\ :due:>

A,~·J

J A ~ '0 ~HI J F ~ SSJ~«'O)

pal'jt~ p<1U.~rlll,~ill bo~;hlhl';l~!,<LI Mooil~!T"Jnl;,;'';;LU~ i(p = 0_5~.) .1Llld the S:I![g .. SSQ :S(;~I ~p = 0_(1) I( .I j I ~fll1~lPtilng sacs, 1I1]i& iSlllcu p~lU'lLCl,Il'nr ~(: c of Ihe;".,c1wO wlllionls. J\W[U)I room. l:fle eflUtltnrial

,I

.oMS OOJ'lCCIHrnlaOI1S, Iii:!>lmlly p~k~n sllmmner.~n ubtmpi~<il and ~ow to.'n~pcil'~ ate UL'lgLolli", Ihi!> m..ximum DMS coil1.C"i:d'cs. \:\'it~1 a mmnimurn (}1''P,h}'IUplar:lklm~ bi(l~ll>l'J.\'" This fca~'!.!re lX"IC11 called the "OMS ~mmn;:r parahas dox" ( I $~ lH high~,i'iU.ldbs; and OVcrnlrr.o;'\ of!h!,;'

region .. surface

wc~:e u\'emg-cd Yo-t cseh nH:mlh and eeehbe«. Me'xi. we slIbdivided the mnge of SRD \ alucs
tfrom 0 l.o.2W "!,V un-2:1 inlO spaced imc;\'~ls of 15 ''VI' m-:l:. and mean J._ st~m:lmd dc"ia~.ion was

cakl.lilllac.a for Ihe' bo.x-il.\,cragoo DMS concen[rm,tim~s cOm;')1pondi,l1g to coach of 'th~ iutervals, nolll., fmm d ~W!brcm h.nniH[.(k.1l and n~omh,~ were S(!i~Hhc.li'Il ecsn, ma I~IC other hand, dlC summer O a\rcrngeclllOge~hcr 1L~ long as. lh~y Imtl a Sil1fD'i~!!!.r suaace DMS ~ml"hlll.!lWcn-necurs wifh n chll,[l solarl1iLdimtion dose, The hll~h,"Sl 5% orthe' DI'\~1S ~laxilnuull, and bOlh 'II'~11~b]CS Imlk :>l!.l,()l1gl~ box means \VCL'C: pl1!!f('Ml~ly no;~ liken ,~nlO OlC" COITCI,ltoo 5'). Sud! .a bC'CL'ng.t.~nCO~I:;; b(:1'Ilol~ i(l:r (:flllnl in OrdL;'f 10 ex-dude hrigllDMS values.asrt,...."II'L'l in VO!'}' \-\'I.'<.I1k glob!ll (;O:In;lm i(m, ... b<:lwo;;u~ sOCi,~~~id i~h IWllr-.ogl!lic (;otl~H~1 :S;Y'''U.t~llz.i and w DMS 1md chi,..;). 115, .i~ local b lo<]il~.l~ (Jf <iI~!!lm:::llml pl:l'Ioduct"'oI'¢I), high To fi1i},(".-uwhCihcr gl(~IHll IJMS ~~~ ~ribuL~o.rls llI11flullilS of dil'lllelh{yl~llll ro:rliopropUOllm.c (DMSP). g encr IO:!I!ow those of solar radimion or sea surwhich are we.lll-d.oclllll~Cilltl\d short-rean SOLlI"reS, f~cc tCl]lll!Cr.1L111UC C:;ST) thlilill ~~IC)SC orr:! m~k.tml of DMS I.hat WOlliM havea di$rro~)on~omllcl)' high weight en jhe 1:1\ crngcs,. This e!I.U-on:' eribiOl~m,'is. we Oo:rnllik:d nwmhly :globlll m"i1s or m:vai,~aMc I) MS cencem r.ltimllS [rom Ihe CHoh<l1 tC.tion ooughl,Y corresponded 10 an lIfl'r-c, limit Smr~ce Seawarer tOSS) DMS c..lataimse '(17)" of 10 ~M Uig, S I), The rimr.:1m~mberof'boxThis dmoI.H.lS\: ililclilides iLbom J(M)O() ~mllvkl~ momh oon~bimlliom" used W~ 54-5-. and the lItlI,:1 dam points collected from 1972 liO 201]3" mml numher ofGSS DMS dara fXJ,hllS indlL'(h,:::d No in[onmtiml abrnn the' 'OI'fC':SpOHi(U~g in sltu was "bOll! ~6,~OO i',c., ric-ad)! ~)g%of du;: orig11\1111 ,elm.a. MIn. surf.LtC irrna-iml.Cr:s. o:r: ~i~u extim'lioll .00--

AM,"

J p"S'CnHl

oil f'M

$R[J,~~)

Ii"iig•. 2. ILii1~r r-egres.'\li~1iI "" 33, (a a.Slli) ~f $1l,jrJiI(e~MS ((N'IOO!1l~ratil!Jl1f$wr:sUiS ~(! SR.Difl Hydro,. 'iltatio.n S(Sar'Y!iI!»o)e\":!). 005 are ~oru:hJlyda;t:a d~riiffl!:l'lh~'pemd fr~~1 ]o1fl'lllllmJj' 199'2 1:Q N'ctve:moor 1.9'94. Error he rs repre:lie\nl sia nderd d'evl;;alti~~!> Qf !l'iillttipte $iiiiililpting ~'ilY5 I~a(h rnon~'h. AQ~etflmla III CIl'rIN~lail"icm .anallysis: Qf the' ~f'Iii,~ d al:.a, gi~~ aJ s.~gll'!ifi((!nt. ~o5itiw (o~f~~(llNltp := 0.89 (p« {)I,on DMS dllta i'! re
fZ

5
A,~·J " A S ,~;n.1[I .II!' M

rr~m

U.7.l.

f'i!gl. ,~. IMcmlh (AprilltrWLJgJh ,Manll) by i:.1tiituoo Pl-oOt.:s ",feli rna~ologlGCil. g;I.oI:tal dfi!illln1bllftill,ns Qlr XI:te'lli~-eoorived chl~ I!lClIlC!r'I'ITaoons rCIH~, :5eaNieiMn~ Wide fi!!!lfli.cd ..\il'i.elll'SE!IiISOr, 22002: to 20(4), ~jtBmffildem,!!'!d ~e& $lIlrfaCE~peraUlr-e (SSiIIg1j, Clirna,t~ Dial;J[l0!5tk5 Ol~~r. 1971 tD 2(00)j~uria(@ OMS (OI'I~Bnlr.lli~]otU (65S, DMS databa~), an.d Une SiRD (crrkl!l~tedl) .. #iiIl '!igr'.a~les< oiIr,~IiI1IOn~ll~ avera(J€dl !by

OL_~~L-~~~~~--~~~~~~~~
(I

2(1 110 00130

100 120 '1<\lO,1&1 160 200 .221il1 S!RDrW'Iill~

~o

10 latiitude baJmis- Spe.;!fTll31lf c:oeflfidelifns (P)b tille ometJiliom oolli\l~ ~E! lilli~mJ:rrti:r d~§mtbL'llijol'lS .0{ VMS il!nd Ih~ 1000er W!mbllls ~~. [l1MS '~~S\!iS (Hlw~, 0.08'; DIMS \!Ie!lSUIi '5:511',* 0.161; DMS wr5l5

:5RD, 0.56 (rJ ~ 1!)5~

REPORTS
glob~.~f'C'I!J!.ti9I1Ship i~ql!itc I;:IJgt (shaded ;;If1,,"<!$ in F~f}4). ~!w ~Ipp.;:rll~d~ow~r ~Q!1IfI~11ffi ofille ~n.t~;r =-II!1 <ihm,\, cl(,;',uf pl·UJ1!t;:In!i9!1~1.~tybi,11\'!I·i,N~~
OMS 1Uld tlrnc: SB,n Ahho~~] ~J1pCr-(:!C'e.,un DM.S (]Yllmn~cs. h!l\'~

emission Ihl..'" eovaried w~l~lthe ~C<1l;;:M"ny avera goo ~Qi!r~~oc SO!;lJf urad ~<lllOC ott d i fr~'l;;:nl hr~ihl~, Mor~ !\1Q~!lI.!y.tI~ dq)~h rI!,~!1!;UML w..s
sJ.;I:.m m hm ~ II. 'i.:gtL~~li,pry ~f]ntl,l'!~~">I; DM5 on rrod~ctioll1 and t10ItcL~I!lmliml ~!l :!.glob:d seale, imd i~ \ViIS hypo:th(I~~~-d .i11l1 sueh :rcg,ulalion WOftW partl), Q(;c~r ~h~Dllgb 't~~ dlr.~i;;'~S the of
MLD Ijjbllk~on (.'xl'JOi>UI,\:' 10' :>.ol,Ul'mdi!llion ( 14, 15~,Us HlI1g11l~ 11'IQ~~ ~litipfC'h~~i .. i,"~ dutl!! ~t 9vfiilah~.~ lod:i.)" we show I~un!ollrffLcc D.MS COitCc:tllro!~ml::; respcnd fiOsi~ ~vclj)' to dil(; UMlL
on

<i!C:~Th~:I

!:i,gJn :>'Call~inJil, <Ind. nm.ing ,depths il1~h~ Qt;;'C<I!1.,

I~lOO~

hl1fl'G~m.

bee n [he (lIbjctl of extcn ...\' research, dd~nij Ii \'C iC

C'O'lI(:lll!'i~'On~
,1~XlW!~t~c

1~@~'@I,@iI1J~!I!",jl N1~tl!<~. ..a.~d 1. P,. IS,. f.IJ~~i. NaMI(!! 3SI. 2i2~U9~l, 2. fl', G, f.al~~ R. T. Barber, '1/, mJeniK!!lk. S{,renre 0l:9<1.

rmuun fllcl,ot.'lffioonlmUing

DM S cm'iCt nl"'I~()m h;l\ crtn,1:11.~I1l;;d di!illi'I1:,

iiiJiil '[3Ii:-; ~ba."

:t

200 U!l9'ai,
l. ~ Il!!liIlliil,. ~ ]. W~UM,. G~J~€hl!llt

(yt1~5

pre\'~liI~-d gi" iii,!; !flii lilll&:(ll1i,vocm.1 for :my 17c~dback~iubk. bel W~~~~. el iltlfi~C'.fLibli DMS. EN:I)crimcl1lrul work (Q. 12) nas UI'ivcill,oo selar rndial:i.o:n dose, !lnd this response :f"ol!ows !he i~tIOOl(:lioll oJ:' ImJ~li~k bi,olicml.d abk"tk over the ~l,oba~ OIl'Cl1 ~Iaycn:; tc.g, .pla)IIiO~llm~ !.:ton c0l11~msi I ion alb!:i the s:all~e l~rop{)l'iIrm~aJlily nceaa, irre's~'('~,ivc onallitlLdc and [he brge "':11'ph)'si,oki£;icml ~~.u". zoopJ JI~ k~Cln SI:m:.i11!i£. bac~f,.~:iall n'C'l~vily mul diwrsil)i. and p;holtdlyslsl,. iab~l~lyo r:. ror e.. alnrl(.', K"lnl~l'a:t~ re and !U'~hic ~ and soIJ 11.1' rilld~11iCl!l(1'!~ld~'Sp~\i;i,d)' l[VR, 1;.'X(.'Il'~:;. slalW!8. I
:>.iglll
a Sll'b!iiI~Wb'~ but U('Il i;!.~.

p'":1- (fiJJl:!ellJ, Sdenbf' 21'~. lQ~2 U99:l'). 5. H. Si!li~ e.lcl •• NrJtule: 4:lJO\lMa ~:2C)orlJ. 6. A, L. ellu,'" :5. M. l[J~rA,er. ~. 5<. U~,5'Ci~f ~!f!!', 1151 f,too:;U. IQrl')O'ffllI"'1 01., ~rure ,4,11;;1',6,:12 {;\W{I;;H. 8. A. L Hir$dli fr aL, ,(OI'tl!!al~.:l!'h~ ~k$ 2;1])', 681ooi8. :1D.111~9l2(,H)~GBOO~44·3 (200ii). 9. It Simti. r~ Ec.w' llVDL :16.~8:7 '(2001.1. ~I;I', It..~ (~~n, J. l ~Q~~~ ~ 0, /IIld*~',. 'G. w~~~, $. M:!II~~ ~l6. ,6S!i ~U!l1}.

;11. MLO. AMj\~~,

14.• ~.i!5(2:M~. ,

t. C

~ulfni~i~ ~ !ol"w;lrd

i!l n~l~n{;'~

(lni;

of t~!J..l Oh,lllt!li~~'l'

m~ r:l'Hu~~)I Ul~m (21 23 I. A p'~~!1icuh1:d)' !;lor 1t;1~ ("\'<lilt. flJCC'IlI hypt.rylhc.'lis $Ugg<.'$l~!'I1.q:! DMS 1c<!k1Ig0rmm Ihe "llg<l1 ooH is lh~&ly~pmducl (If

\ystclU ;!ojt:iL;flC~ : !.O dll:{:id ..1J.'! how lite b~~ph~.:rt.~ i!' li~!~ld.~ !oclimOlIC in~\la>,f; dlia[ in unn ill"

nr

'!Dd..)":;, [E,I!1Ih-

,U!Hio",id,~n~ 1l:I~'{:h'lUlii'l!!l (2.1 h Gi\r!.~I~ liH11 high. liglll,Qhigil UVR) ,~Io~~'$ D:ldH(:'ll ~ o...idmivc stress {i,c~ DMS release) ~Ild iU'lhibil. bac celri~~ DMS OOIlSUlliipl ~Qi'I as w~1I !(1J), DM S ~Ilay aCCUll'biIll.llmlW in seawater, Phyto;plilllktml SIJC()~iOhlIO llig:IIIi;!I'~DjI,il's.r Ilrodw::c:rs in sumOiL

,';Wllfur:-bm:¢d

llueuee e~iJI]I:.*.':26,. dC!cm~~!1l.:~hcifOfi'C1.<uion ( time M;;'I~J..l. ,1111.1 elari I}' \\'h~lhQ'r 111,1;(';"': ro;;~'H:II,,""-'1\ c~mlt;r smh.illity to d1t' dilirl1Uh) i>:%[~m ill frolla of
pcm.ilrmll.itJJti~ ~UC'~Ias; anth'1I1Jptngm1l1e IL)~1 eng '" urorn~lr;!~blt~dl3lilgCl. TJ1C l~g~1I eo~pling (I nJiM~S
"lAC

oxidat ivc :i>! ress on these pmduc>cl"S, and o:llid(lt~';.'c dallHlgc an .oMS CO'J.1!'llmC1-Smay be coucunem n:ca~O!l=, Whji DM S (:o!Hx'mf<liioll.\1 ;!~ h ighl:'r~n hi.f,lh·lil,lh~
~lCT WJI.Ct'S.

strntilicd

c:onditimns,
A~ocent mlaly,~is of the [DiI'II]S linne series in ~hc SOIrgass(J! Se:l [i(,'w;.·tI!lod !hm~h(,' ~C'milOl"l DMS \/,o:riation cniIC'rg,il1:[,l f'rom such .. C!iJF.llllk\x cycl(ll'!;,';."<t;rllhk'<'i Ihal (If I tile: I!ot;a~ UVR. ,md.lhL;'
laU(ir \\'''~ .~IlI~~Cl\terJ,lII:;,~h,: 1lr11~(jr,dridng foro~

~Dll~e solsr rnd imion dose ~12lt condill.i.Cnl lor I,he ocCmflC."HCC oJ ,:I l'tCgati\'c leedbad:. bet WOO11 plankWI~ aI!~d climate t~rotJIg'hlhe iducnce of (he (olUler (Ill Ihe rndiatuv'C' c:'nef'~ bud,g!i!~(.I!'h No: [all]y, it ;d:oo provid~sa ch~ on ~hc t~l!lC''C<ll~ of l such fC\i--dlmck.TIIlc wklr I:adi(!llicm dose of the surface ocean varies sh'O.n,gly over Ihe scasona I c},c:ll!c as :iJ: OODJscquenoc or the coupllcd vafi""tion Q I'::ill rthcc imld imu;~ aud~hI.;'MLD. OUf .0111,1 inobserved is a l1e~WIIY d LC<It~~ il i'im Ihi... ~Cffiolml . ..:::.alcIh~11 the that cpilllCfuigie uMsy::iICHh IC:qlOrbcH [0 ICIi:DPOl1a~ and 1!1I,iutdilrml dmngt""" in soJllr~1idil}liDIlIl)' chtlrlging Ihdr pror:lu.{"~ion. igthl~HCmL![l!~rlgol:l.~ilc v SJLllfLlr: E.xl}loro~k),[I of re~J(ms;~:,\ fr~ ti:UbC ocate::> shol1cr tlama 3, mOl1~h sllml,ki be carried (IUD widl

concemr3!lions

~~"Ill. ~ ru~ li1Iel!lOOl. ~ r,e ~~liO!~te ,~s $~~r!1iDIJ iiilail!iiia'l 1m Sdel'll;~ OnliM, U. 1. W" IrliIcl)'. F. A. Hoil'J1~.A, IF" ~lcha@lJ;" S" Go 'IN'.ilt.m:m'll IJtreW5f!Cl 1m, 145, 2005 (1m), 13. o. 14 T~I~. o. iii" )i~ell, G.e~~. ~ Ult .31, 1!Q?3(D8, ']fi.1029'200~G'L01 ~5111 (200!l). 14. R. :511116, C. l!!\edrds.,(!j_'i!!. M1iu.e4'OF,!~ sss n.999~. 15" s, ,Me ~ilin3,. It SiJrJd, s. 'G~riI fiJuaor Biog£l!Ilili!lll! Cycll!'!; 20., 'G811:Qi14, 10.ilCl29J200:lmeOO2S97 (~olll6), 16. f!.. J,. I~'tlle .~.,a1•• G1iQba' 8jc9~'~ C)d'~ ll" 399 {1m}.

n.

11. GlCtbil~ Suilf.aoe'

~eG'Watef

Dimel.~)4sul!li'de' ~:ni!b.ii~

(llup:l~.~!J.prne~,q~~,go!WJdrnsQ_ 113, G d! ~arl!l' I~i!gilttj 01", t G~y,t,

Ki'$,l.oo. '],0.102:9/.20041(.002:3,18, (2(04), 19. 1f. I). 16roct,. f~ Mudf!. :1!4, :I! 11l9S11. ~[]l ~. T" !KiErnl, E,Vm;:oo.eIiU1>,8ulJ.Am.1M'~ieoflt~~ St:i(. 78, 19';1'

f199n,
~1. W, Sumolll, O. ].It\idJer, Ii:. f. ~liI:n~, s:, ti![Jl'nlll1~l1". N@r!!ife 4lS, 3 fl {21100,. 22. ft. 'Sim6, Crl1l',.l, ii~h ;i!~r.rillt Sci. 61 673: {lOI!!4~, •. ;a. 0. A. V~~, dol," DefP-SrNJ~ I 503, [36 ~OO6L l~. IT. S, llaie~, III:. ,I. ~d~n, It. H. ,(jalmmrn, NtlMe 32:'1l',

i( I J )",;1,1]Ilcllllcr I.hi:;, \ICry :;amc qLmm jm~iyc DMS.,

U \til relm.ionship wm.nh:1 be ap;r1 ~eflhl~ 1.0 m,031 or [ill! global oreml was mtknlOWul hilll ~mnkel}i. 'b C,'(:m.ls,c o •.hcr ~ca~ rRctOO'S such: as pJ f111k.mn ~btmdalliCC and ccmmunity 5lntcllU1C \wuld iJ,c e,,'Ij,~K'Clcd h "'\~C a ~a:rsc cOIl1JllCilucntary ifllllllto CI:lrCC. NQjlClhc~,(Ss. Oli, pioneering wort.; by BmC'il t'l {ll" P4} shm'lcd, w~I h the few dllm Il,vlliilabk: m ~he tinll::, Ihal,lhe
S'I."<I1'OI];[1I:1y

on

319 (l'967), ;!5. It. :5im6, ~. OOdili, 'GIa~d'l8i~fJ(bf:m. C,rd~ 1~ WI8 'nal'o291200lCi'OOIlll8:Z'J (200~}. ,26. p, i'iIeir; p, Co~ I. !i.iriICe. rremt~rc)o1. mL 21l~~
(,2006}, ~7. weoUilank I\t llIila,'~·l;a <lll(l f. unlieim for ~l'ing $GHI~ ~1.U'~r I~I (lata.; C, d: 8a~r-Jllo!1lI6g1lf1 fllf IprtJljdil1~ ami O1Ie' PIUI[I cl,matiill~\I'; J. I'i!J. Ga~taOO i!li<e- 1Il~1'I'~B~ Mlaob~L olMM':aII)I}I',~t~difINlm !dl'!~i'IlIp[irig ~!IIf,sMn~1'! imdm!:fdiniIlijoo; I. I),a~~, A. Mifhllet~ .imd S. WJ~ham r~r the 'U~ ,gf tht: ~~II1!!d\;! ~5 daflii! ~'; ]1 11!!i1~iO!I1.rQr Uile ifiialrii.ti11fIo~~(~ 1IIIhil'~,~: MiS, dilllab:i~eo; ~iid U~ Se\i!,'!'MIj'F~ IfIf@j~ UI~1i1Iatl!l~ (i~d!;lrd Sp.ilOe'I1lig"'l ,(J~I1!l!:r Il~~,g~e IMalnclfli!! ~ mr~;D!i Alim~:ph~ii;

h igh-utso

~lIliml

'In~lIrcment::l,

of [lMS and so~!br

Ilveraged

D MS

,',adialiem in coherent W,<!I~crmasses. \,Vhctllcr Ihb;. r~l'lilck wi:lll iltoo, o;peliil~ cmlicic.nI~y III tl~e long{;'I" ~illte sca'lc o~.·rIinlhm~~ic . £llotm:~ '~131111i:11g 'wHII dc.pcncl onimltoced changes In gD~b>l1 dou,cl j'1l(.'%Ss.

F~g.4. Lin,e.1!.Fr~g f,e$:siriHi {n = 14. r~ ~ O.9~H o~ surface IDMS ,()ol'tcen;tratili'J~s V(;l"s~ th~ SilO ~fIthe Iglob~l OIP~f1 !!:u::eiilfl. lOots oil ~e ;l!1f@'rtl!gle~ '(Ill 10 'by .20 (l.l~itlll:hi!' by ~ongl~ tude~ b\tl:li: m:e~!i OMS· (,QIfl(IM!~ t ... tij o:ns gro ull~d' by ~nter'!l'<llJi Ol~ 1.5 W m.. z(!l~SFrm. The :s,inaaed ,1l!realrcPN!':5!!1il1S I th~s,t1lIild!ard d'evi,. <IItioaii Qirfhe aver.ag:e1!. The' l'Ium~ belfS bV the data; pClinlSif)ldkat>e ~h~ ,i1!!lTtJOUr!t Q'f DM5 boo: n,aJiS
IJlsediffor ead avera:ge'~up,pet' liIum;Itu';lr)amd! 1l1~ ·amoo FU of crllflru!ilal ditltaJ f,rK~ud'ed (nUmDerilli pareliJllih~ 5.!l~. A :S~ilinllil[1n (ionEllaticm all'aly-

ru:J'mJ'r.istraliIJn" Ilh~ 8:ufrludl ,!i,~lallltil1iT1l1@·SHig Sluil.lI'. !he'18ell1ll.U!lal Bio,OllliC'5 f~4l, oil m~ ~!tI~ &i~i!~n~ :Sulfi!ce
ItadliatlOfq IM~~(W.RIl'IOfor iIIi~ing dala ij!.f<!il!!i~~~ ool.,e,'II1j~ ~Ofk. m;5~lIpp(fI~ii1 ~'l't~i! Spani~ Minim\!,

.;If 'Edw'~~t!!l
rrna.l,1!eI'~~

~!11~ ~imal

~qn

tl!1rW~h th~ ~~ "iWJdl:lil'1i1! 'til1~ $Ur,ra~e o.::~1iI ,and !file'

<!t~phl!l'l.'· (tafl'lirilli:'t CJlM~OO5"1)6.~B tD I1:.'S,}~nd a Ph.D. 511:Wt!:(rt;lilip 1D S.M.. \(!., b)'1~~ (alal;m 9IM!fnm.tl!lt (fuH!l!!gll r~nt 20a5)G~(JOOiH (gO I!.~,~n~ ~. ~h~N5,. g !hl'O!1lgli grant O:filPi"ggil;ro1m (to, P•. M:l,"iil,i~. ·1Iihli~ i~~ Cg!1I!ii~~nia!l IaI!Lt' o!:!je-~~~ pmgf,~mSuf~~~~ ODe'ailllil~

~l '!:!t4: 'il!te!i]j!~lfitina'l
A1mo~j:lt~ $!l!Id~ ~S06l!i9 !l1'~gr~l]].

..nil

tn~ Nl!wori

ollf.mlll!OO!' £,UR·OCEAtM,~ ~h~

WFOJil~~ l!I~jD~'~ SL'dh fl,am~r~

S!!l!PiPO~Ul1gl On'll!i1E! M.i!~~,~O!I 1MWf"S!Z.ieoo!ma9Jl19!!;gircCl'llentflLIIIJB1_S!Sllll.ltS~1 MaIMials ·iIi!!CI iJ,I,ieU! OO~·

:s;lisofltJe

545 box

ffiN!liS gffV~
[oe'Iffi(~(fHl

(j

.ilO

6080

1'0012(11140 SRO(W!;'f(~

H3D180

200

Fig. 51 Ri!I@rfflo!S.
9 _51 :1;001); ~ ({epteil 6 IDecI1'IrIDilr2C006

5TIgnIiirlll:amtp~iti\'e 0.47 (P'« 0.01),

p-

lett ~261~i~m~i!.lH ~~BII

REPORTS

A, plll,a:s,milnDlg,e'n'~,Acti'v'a,tinlg P',fot!e'8ISe Spe,ci'fiic,allilly IC'O'lltrlolls the Dleve;llol:pmenl of P'rilmllBlry PneUlmo,rni"c: Plalgule,

Primary p,mmmonic pl.a'gl!leis~ramnnmEl!d: ei!s~ly, ~roglresses. rapi'dly, ,lind eauses hriiglh mortaliily, but tille meC:~i1Illiisrlils by ~hid'l lffl'f!!inl'O pestis o1;lEn'lliielms the- IlulIIgsar,e la,rgely 1IIi1kIllIlWI1."We ~~h t Iil::tt~h~ pia ~mili'l'llg@1iI il!iclwal:or Pt:.is ess@ll!tiall lor Y.pe'5tis t~, C:alllS@ jJ rimary pli1@llnflelllk illW P'~ii!91l!~ is ~tS;S; f m IPO:rtallllt ~o'r dliSrS@lIlIlli1a&ioirl IliIluring P'Ii'I!H.!lTllo:rli:C lPi.agll!!!tna:n Itlut ,ei urilil9 ~l!b~:nk IP~a!gIU~ E;I;p~r~m:e!lt~ !iIiIil!rnp~l"t~:ngJ ~t~ t!n1IPQl!'a,l ,~;(pr@ssi(J.r1!lhow!d t h::!it Ph~ ~,uo~\iS Y. pe!it'~ to •. ~eplk"te rap,jdly iJl! th@ ,C1f~ys. (alls~Flg g l~~halrUILFIi!i~3JI't 1Pii'i'~Lllm~l1j\1!,~ ll!il~Xpr~ss,~, ~'r infla m malti~i'I is a bonEd, ,aJl"I(i l!,Ingl repai~ is a(ljva~edl. lii,hibiU(!f! (If' IINa eXPrr~$$iClI:!iIPlr'QIoIiI,g'ooU'ile $uM1Jal ~d ali!illlla,L$wi~h the di~~5;~,. ,on.~riUiig .!'I' I!.tierape!llti't: ,[lpti~n to ex~.,dl 'the perio,d dl!lri~!g which alntl,l)]ol1:cs are ,e(iF,edivl'.

j}J;tgtlC (l2~ sucenmbed to the g~lrcclion ill <l hi~dy ~yn~hITIJ~OU~Il~OOTIiICI', In (:Qni;!;"f>~, (l!nly $00,4, or ,Iw m~~~i~ifc,,!.qd~'\..'i'h t~l~t-YJ/fI ~m~il!
~~" dop~',t;1.,ctm~lIl~p~!I~~I~ ~~Il~T 7~i!}'~, !md Ih~ ra~,,; lIi.wll.Ueh 1:~bL~ miee di~d Wfi~ less :lY'l1ot:hrun:O'U:;l Ihalllhc'~IIIJC (1hi'!o{5C ink'lC'tedwidl~hi; \,,'iki4}'pc

:.~r.linl.r~~, A)" ("or.npk:rn~n!'II~!r.mof~h~ p]IWII,lIlll

w(tll Ihe codiP1g :«;ql,lt~lct lot P~11iillny Ii;:i1nu~ud ru~!!iJ1~. l1'lI~, I.he ~~~tk of Pifl t.lnl~I>~ai'nill.~~y 110('bJicd lh!l::'Ii~~l~~10 deall~II\':$U~Li rro~n I\.'),;p,.irnllll), ng inJection, A~IhOLJth lite kil1:Ct~.e:; e f LHLC~riIiJI li:~'(iWdl, dlll'h~~ ,il1,r~'Clkli'lwi'dl COryl p:1)ooecd!::d as C~~.~'C'~ccn'IIJ t bacteria ~ mUl~IlCI\\'~~ in 11~liC' Ajdcl-

II

'I~

100" ~o 1000.1:01<1 fCM''I.'f Ijm'C't~riOl! ~ru 1\;"'C!tI\'O!'I;:"C! ftQn1 th()llIlI~ (1rilf'liaw iu.fu'C,~'Jcl niliCL' COI~lP!il]'iL,o'd wi~h T!~~~I,I~'!}" ilb w ~~'.illd"lYI'lC'inR'JC·til:m.Ove, Ihe n'~'.X1 :2 d<l~iS. dl'C' ~~lUnLlC'fS tVlialiJ,.u;:tcri .. in the luu~g~ did U>l.1l f ~~ 'hn:,~i;l ~poc iC'oi1 CI,f .l~I·shj'it7 p~~hQ- u1I!dly ill the hmW' ~illd 'rl(;'~l1~ h~l!"'~ PJ.'WlII).10niOl :>W!b1;:~•.ulitii!dly c-hOlngi;,wlu.~,,~ \'\,'ikl-tYIH;h<iClil,1r~ i]l<1 il1crcW>l~ by ;.I~fl!m!\l6 log_'i, In mUlr..!'ill. \~'C dL,.... ge'~ie ,h~~t~m1J1!"~:.r. f.H!8l;'~~s ~u~1!~b1ml~ b~IDIlllOlt fibr.in ,d",pallili.OH \\'ol!lll(1Li',;i'ilJiicl [he :lhil~lY i owmg w us alJl.l,lly m (:<LUi.>~, ri,~guc,. [l"!Cl(,dlJnC'~ri(l lui the l>pklCiflS Q r~\~~ rl~iC'Clg;y 72 Q n!llt1L'fim.~o (",~{ifll~ the :uc~fr!i['J!l.ol)' $~lSl(:~rll. Ahlmll_gh iLlSllaU}II.mlJiirrnh~d1 by 11111rurtltroptxi "\Ie infecwd C5713Ull m tc~ iliu:ruua:lL.;;nUyw ith lilOm'S" iu.dic:1I.1iug tlun escape rhe llqAu Sitlrf~il1, ~.i:mn I.hc lung. s to' dis,lal orgmls !>Iill occurred vector, it is during eases of sL"COndiuy lPl\eM~ wild.type .Y. fw,wi,,; CO\l'2 •. 911 ~:;:O',gcnic I: IJHWi;l; U":ig. I C},11l deed, til :ilil!Ct·~ hn~.MC five mice ~110nk p:lligtic' '!hat ~: IJ{~'lis 'CI1I1I~ Sl~rc3d from Slrn,i~l ~llcking PIli. 't C092 tJ;~/tJ t or the i¥pla Slrnhl C{}mpiCiincll~,oo wid] the- ~"Odillg sCI1jllleb1!CC had a l'lOlcl.criJI burden approachi:r,g Ilf 10 I r{J p'C"rson!o jOC'r.sOJl !~lrolll.sh I he illilill!al iOll of resrolony-rorming units {CFIU} ill .he s~)h:.'Cn, pir.UIOl.'Y drol,lcts 'CllLrryinglhc bacteria I 0. This Rw P:Ia. Mke given wi~d-~yre)~fWSJb~ C092.:1. mlcllIDd of [transmissio:n may i!nili:lIc an epicom.:-spondcd wilh Increased numbcl''!' of bacslmi~~ isolaied I~n :1 J~!I"~ case ofrnc~mDlIic dem ic of p:rim.. ry pneumou ic pl>'!:guC'. \'\,'hich is and

in Icr~oo C \VOlS signi moc

n, Afl~f 14 hours. or

Ik'an~ I}I

a I~crod t ri~. I. B

or

or

n~s

nl!"l.l~r

nm ~!'J;;'a1lid ~'.<JJi)l,. Pn~~um{)ll~c P~1ll;llll;' i~

~o r('Jml I. in~ "'h~ C'vcnt (l:( 'L bimv<Jrl:m:: ,m.u.::k ':'i"i~haemsclized J: {Ni.. (,n <ifis }~'f~il!il'j viruk:m:c in 111I.,urnmlllls.equires the ... r y~c 'ty~,,",ln sccreticn _~ys~L;'liIIl (T3SS)1. which is ,~~'I1LrCtl mllilong 1111 three pmil'IOg.L,."nic~pl_'{"i,(';S U)I, Uri.~Uker jJe.~.~j:,~, flrlleroty)Ufi!_,u and }~ p,~l..~l~ }:
dQ'fldJ[<I'(:ulwii,> are IOQdbomcptllilogens
.fLlbCi

~lw JmK<;t Iikdy ibnn

IB
,II.,

,
'

1~

t;r

.•

'->
.~

--_ii,
g

ilI:i!Jj,allly result :In sclf.liimlilillg gastroinh;:MilnaJl infections (4 6), Thlls, tllC' presence oflhe Ysc nss; [llo'l!Il! lis IK\t sutflciem to cause ,[I I:f\pidly pmg~cs..~ing.respiratory in rcct~oll" r. t.!(;',~/i)" ,~I50 eames I)PCPI. OJ 9. S-k b plasmlid ~h.n ,evacod.'c~1h~plasm UllOlgCn .a\:'l~i\ aior Pla, ,,1
'~Wlf!300 ~]!:otc,~~'tlilm .It-> ti!i(lll.J;l~l~O l?fmllqowplm~ ..

::14 4iII

D
i! Do4!
~I

"
,~
r.t

~
1,1:

!lJ!i

"' ~ ' 'ill'

~

uni n ,(J,cgr.L<im ['On of'm ..dil1lclots 'II. ;

ofln!OOntc

IlI<lg~lC'.Pia

Pl'(llllot\."il.

~he im,~5io'l

n III models

of'

IU
:~I Il~

V~, !,Ill

\Vh..11 in~n:i>dll:Cl.1d by a(",I.u.,,'t~l, t Pi..'.';;I,rs lackur:ag_ fku wil:S ~.t'!P<Ia1~d:.0 liIC ~'qUlli'\'ilkHt Of IlL:ur 1 cqIUh'ilh:~n :in ViIIHI~fl.CC to wild.lype by m-cdia.ul

)~ J4';','#~ r.mm ~l!.!bcwlt"m.IO!.Ii:l ~it~:;; of i!HlCllr"4im~ into dw IY~llph,uaijc :->y~<H;Il:!. ,IUd dl.;'Cp(.'l!' ~iss~liI;..o~ IJU~ is di.'!pcll,R;IMJli Ib~' gmMl'1 , .. th~ ~;t,,; of ill(X;:ll:~~iO:If~

:::;i -

1..........

~'I,

1I!::::i=..Q--=='" ... -1l,~

lii:jiii

"
iii

D,o+a-""9'-'_"':ii---',---li

Ictlull dn.'>I: (L.D!<O~flflil~)'si» (/0, Xl). bUI,~hc

fI,mg!\''SSion lUllS and S)'st~mic diSCilf.'>'C'h<ls nc'v\:, ~Cll CValll>lI00 in, a mod: Ii of pril1lmry p'ne~m(lnicp'I<l,gll:e. On l!he l~.sis ofl~c:;;c :SMI~, Ume$ C~IU ~er orgaiil il'l the Iltllllg~ Onafld 5:p~~emi (0 were dletenmiiiled. Ear::h IP(DifiltrI~pN~$~n,gth~ WCp:i\."<IicIM .hal rcspimtobY inl~~t~ml wi~h <I IilIL!mbr~r~ r~!I:'(l;;ilnd 'from aJ :~l,ngt€' IfIiI(lU,S~. U@ t]mit ~f d~t@ctiofl ~iS ~ndi(,a,too by .al Ida:s~@d lIne. :slrnin gf }:lJ!tslis ~ackhlS Pila.would IINJCccd 11.01'- S,ymonlh ~elm~~he' timli~ ,of d,eote!Cliolm l~ePlre5lel'ltt mke-~'hat s~HVii'!led ~I!lt diid mrt haJ\!',~ dnenaliJ.1eIlll.lImb~r~ (If oor::lerial; (If'll "x" ~lldkat:lBa mo,I.IS@ lhat Sl.u::(~mb~dU)1 tli!,~inrl~(tiC!l'II. :A soHd llrt:e> 10epaptilTleni !l!f Mtdlecu!ar Ml(robr(l['Og~, wa~h~l1;gllon. ]ltclicOites the ,FJiI,edt~nJ of CFU reciOver-ed. ~'P "" .c1.037 •.... p = (1..00'2. and ~"'".'p <: (tOOl. (lilllIlJi!ired , iJl~~r!iity Sd1>1!!Ol 1 ,tIWi6ne, 0 St .. lWis, MO 631:10,. USA. te~tt (It)) Gm~s, \'IIll~iglhl Vir lurtgls fWIII1I mice ~l1fe(tlild \!I~~h tI<, pf!§tir; CO~t;! (blaci!:) or CO:iii,~ tjpla lDl!iJl~liUlllel1t of lr'~di<ltrk'5, Wa,IDlrngl.oo ~nhrer54ty ~hoo~
,!!lola lIIall@l'rusll~oo

or

Ifi !iI..1. Pta i~ reqlu~r,ed [or if. p.1!5Ms 'ttl !!:<lIIJ£!il <I wlJltminant. ~n~ect'ion 'Df~hel' llJlng~. (AI Sillrvilo"'a.l,,of (!)tHl:IJ6 1l'l1.ce-inJe!.te-a ililtranalsal.~.y ~!oIIil~l( pestis (092: (bl,K~ :!iqll,~r€'S), (0,9,2 tJ,pJa <whit.e $q'LIar€5). Oir (092 t¥$fj c,ornpll!llliHmtca w]lh pIa ('whi:te '~~.lIlif1illild~·h (IB il!liIdi IQ Kilill!tks elf il'd~li:tion~vi~hY. pe.st,is (.092 (blac k) Olf (092 l1pfa (~'IJ~i~e). 1E!,i3iderii 11~w.re i.ntmd:lIced iin~raflll,>[lUy,. andl at v<lriiom

of liMedicine, SI. I~o~,i~> 6Jn~. 'USA. il'iO "To ~!bom ~:o~~e~ppcten~e 1!IlIG~dlbe add~ess:ed.,E'Plait

(f,j\h]~~. CS7B!J6· lill'l(e \"er~iA(;e<;too <!liS @!;IQVl:i. ~nd!l!t,\I'~ri,(!~,~ l~m'~, 'l~ngl::i Mlre ,~~(j,~d ~nd Wle-iiglhed.. "P ~ j]<,,01.<ll'Idl '''''Po:: 0',,001. Ead] ,expenlli111ent mali rep~atedl hrice. Emir biU!> ~epresel'lt. S'!n IIIdalr,d de'!lnaitiCiIil.

REPORTS
teria in rbe Il!n~;
U,,-ri11 in

however, ~li~ ~s m,;,e!y dW!1<: W tlw .mlmq OngnlHl!I~ .<liCil\l<!i'I ion ~Il~hclungs by nl p('l~ym(,'1::~1~cf\,'II;ir;al!~ ion of :5i)i.Slx::m ol];;;mi~~n.~oo~.k mi.!~lgq1!;l!fll.i!,ri ivereve ..~c-trn'lllc!iplj:on ic hno ~Iw~ttlrlil~U3} r.ullu;r thml O"~ltgrm..'!h (]If b;,JJ;;· ..~ ~h,ti!l !'f,,~!ijOIl! (qRT::'CR) tD ~~nl~~I1!~ Qlo1l11gl";:'S

·by~hc foll!flwing
~~~!!]l~f)'i,

ilm is (lrgnuL

in lr.lm;~ril)I q~![iI.Iltil.ie:_..;Of!ll~.!'lipk~ll"nan~ma.

Ihc;s.c ~Jj;:ushm'l'
C'l.·i1~ Lu:n of

H111l

PII!i(;'onllvls

i.lfl~ prol
~:5

if·

es:'o;It"dHu l~Cllilri;J !o di~mri~Olt(,,"~ Thill is <I clislili(,,"Cr~ie~lotYrlC 100lil Ih;lI'I!trjbu~.cd to fU~1 ill m'iodeb.of IJlbll1(moC p.!liigWlC: \\lI;("liJ Pl3-i1C;g1lIi\"L: }~ pestis Ii5 iill~rod!jcc"£1. :;libc'IJ~mlooili:illy •. tli~S~rl.,i· i}~liru~ b d't'fLI1ruLliC'a.llly reducoo, bUI batl.,;:rbll;ii:ll~

)~ f~~':!I.lb: !.!he l~I~'lg.~ bW!l in

net

tol}' 111(:dil,U.(lrs" ['ml~isIC]:Ii wiah olJrrn;vio~~:;. G~~cli"ii'<!,t!:ons l J J}, I'l"l!icc infCc,1,cd with ('n~2 re-

m~.~n~d Urlr~pr!nl,iH: C'lr~y iu !hl:.l inri;"C~jml lillil z;J1ClI'!IVl;:d ~,iglii[Dit~mt CY~Qkil"iC UI'i4",;g.u~a,i(l1li i}}' 48 hourn ~Fig" J.A). Si~'iiri.!1n~~y.n'iC cyl'oki'liC' Il'fni.·

ill:r;:: ,down·n;;~1d:j!.tiDrl pI'" ~Iw i~~ U;Jll~l.1~~I}' Wot,;'Sf}l]n~c: to iJ.pl~_ 1Il11,l~,.,he 'I;hll~ ~v,;,'<II~Il,u .• 0 u Ih~ ~ik';Cll~ of .PI!iI, ml ~U"ui.illn3mntHd.o!}' ShilC o#> ~J.u nod iIII (l,lC I.UHY. and !Jl~ illlti.>c~~onis IlI!n' IJt..~ .rib!,,;: ~o PTOgn..~~ ~o~lic p!'I':J-il1!l~.~ml!l.[try phmiC ... uhal we ~~~'ribcdpre\.~it~\'l ...y ~.I'3), l TIi~[~ eyto:kiric i.mm,e:ripl 3"i~,Omll'" 3g)rcrurod

r~n most

Q<l}' I(;)'W~~

or i~~rC~lion

11'WI11'l,cri.p;t

nes QJl,'l;g'C",""W{I, :'<llI!ggCi'i~~

g,row1h

III ~lC

i:ocal site of

itl (celina

i~ ml.~lrOCIOO

,[01:,9), :~Ildccd. we show th'~1 rates ,[IFdi~scmii1>1~ion rml:l~~hc ~Ili!i:ll ~iI.c of C,O:lo:nlza1iml IOlile ~.~IC\C'n WC'lt! s!.!I.<~lmn~'II~y . increO'!!;.;;)oo\ hen I~Ja\ I!.r;,gm;~i\.'c bitt leU·i" were ~nll;Jl:ld~ocd i !1!rO.1!!l>!~I!bl (:(limp~~!\;d\,\Ivrh Ihe $.l!J1bcl~~0lI1COU~ l;Oll~" ,twblc- S2'h
Tilit-omny hi,:: clllt; Ir.Ilh~ hi,\lWy V'd~C!ll~ril;\:d IlOLW.1:'iU orthe lung. ,!I~{Iwil1g ~COl,lX' 0 a Pla-.~lC'g.ati\. e

ser.upl 1\1.i1Ub..~rs.n the 11[l:1'i.~~ i ofnBi.{.'C ~uf\)el~d \\'iI11 At'Jia were also rell.llivel)! iliilc.hr~il~"'iIl!iirl:r in d,c: in [ootilm.. A Ilcr "1M hours., however. cy.. reki ~lC 11I:a11:'!l!:rir'l mn1l1l'I!C1'S :in response to flp/"a \.\cC'1i"C ~oh"i:I,~cdr on~y ~li~hd)'illCI'Cascd. m o <lnd

rOo slahi~:i,L"'; ~irni!'lllh,,;tl.loo'C2ISC t

~liec' ~Li~gcstedlh~

i~~the

~;/;a.iljl~li.\d

ilf!l1fJn~Ulilluory k's,ul1I'lS WCtl.'C f\.~ol'vil'llg. \Vc irl'i'iim.Ul'lo:M!lincd i!nII'coclcd~lIin~s; For lillc Pl'OlircUJ:li~lg cell Imcl~at' Ollnti;g!cn lPC"N.I\}, 3 nmrkcir I'm' 111O~l ec~lu~:Elx

pnllilloiHl:ty

r lew

l'l.l!lt:!~riOlithm'LI~l ,m ~dvl;lol:lr c"r~lli<lfY <!ltd Ih~ls h~ili,~li!18!: }I~II,.\mi{:' ~ i'fI~t-,(;liOJ1. i\. il~IJhll.1rk ,of fllm~ pac[C'riull prli.:lltilllO!ill[L iz.;.

Ihc di.!..'C'klpn;,(:~n of cdellm~il]i:li~ hlllgs, whid! ca Ii be ItlCasuro:lby jJ, dUllll.gc b I gross ~Illaig, weight as fluid and cc:1ls ,oonlribL1tC '~o ineressed mass ol.'the ol].m. A.itho~gh i])ou''>'C' ~U!ngsin fectoo \\'jllil\'o'illd-4,YiPC }: /JI!!I!/i,f wcrt£hi!d SiShli !Iknntly
mlOre Ihm 'Lminfcx:wdl:l.llJgs,lhC' I~mgs, rn ~oC' inth:l'~'.d with dlC fi,Jn.a ~t~"in nof ): f'l.',~/f.'i;i~lO">\ilf.x! JIIf'l' clmnI,lC' ill ~\·C'.~,!;lI1!,even oallC!f 7 c!;Iy:;, (Fig. I D}. ,';;MSgcsthlg thMlhc dc.atli!.or mice inrcc~cd \v.i.tIi~ ~his snin ~:5.. uet clU:c'w ImCl!.mnoni@. but rathe,'!: is
by S:'lfS'lCmjc inteetien, Our ut$uJl"i.d'l~rc'" I.QI:iC, nnay c~!)bill1 the' :-;:~lllIi~JI'" lD:so \'<llll~ w.il~d·lyp(' lUl(! PI<l-Il1;llmi\"it.' ~I.r,~int\ \,vh,;milllh1tk'tl. C\'Vl1 dlo~lghr pt.'s';J; Dl.'ql!ll in:':S f~!Iw C1~I!I!iCtL :;t;-.. \,"CtePllCtnilllDli:iil"

or

caused

ro;

\Yc fib>!) cxa~nniJlcdhrlg h~stlJ)logy of wildIypcmu] nunam [ Pt'.~"fis-inl~Cetcd~lli{:c to resolve 311. ex lisl i~lg controversy r.,cg~ming paIhoiogy mld .ilnf1ooHnati.on m the S,~IC of:inrcctiulfa U~.9" !}), n infTIlIJI,or .fnflmlmmlOry cells was A detceted in lh~ .llungs ln both wild.type OIl!:.!i'Jp!a infeclions, 36 hOlJrsfl~lcr ihlocuJm,ioll~ in b01111 cases.lllii: rn;:dmnim!J1l1 int11[m~ins cd lYP'C '.~<1S P'O:I}'lnOJ'l)lroll,lIcllC'~n' I P'i,g , :!), A.lth(l\~ghlhi;! $:~ze of Ihe ~HdmlO:1!!o1U.·Y !t."Sim~s iiJl thc wi ~dLI.~;}C ial!;..'C~ ln l !lCN;I~!:''tI over ~imt"'!'&.·.:'l!II1:!jllg in~ i.~"'J:c k d~trtl:l;:~lo!l! tlnd hC;:IlilI'tilih'IY,.;', the ~bci{~I"inn,lnn~ ~mni{lin!Jnmk"l;" il1'li;t:I~~ wi~h C0"92. tJ.{n(,* ~m.l.iincd 1."c~f11vcly CO:I'I~WlJlt ml~~~.lri¢tad~ i bl 21.ddi~ioll"",,;~ c1'>mnil1.oo il~ltx:cted 1'LI1l:!:;seerions by ~lsil1gimmUiH)n.ilIOre..~1'CC '~"nth <~1l1 antibody 2lga.i m.t ~:!J(.. '":J:ilo;, NtmlCmLJS exuacel 1u.!arbaC'lCii a were associated willl .inllamflilJlliol'l in Ihe ~!i.llngs of w ild.type inrt'd~dl,nicc.. hut .n::Jla~h'dy r~w k~.ili.c~riut were dc1cci.OO il~ ~j.rrl iu[fuctim~ 111M dJC wererestrictcd in the much :;;;m.ll~lcr iflll.mmnJJtol)i
Icsr~.tls (Fig, 2). Tlms.both b<lc~cria~o Lligfl'lwth and $\lbseqiLIICllt illnm.lnnlal~o.n~n the h.lll1G;i'il WC'J~C' dCJ'l!CJld~vll on.

4111

7:2h

4B trJ

r:. ,fiW.'s'lfs

SlliiIiti ...

lh~

I.hePb~ surface p.l'otc~"C, 1\."1'i~db, sliilggl!~1 ~h l h~,1 ill~!llI\lnit}' ..

wi'~hm~! dtvJj.
111.';=s!''lI

c'QnUm~ t~lC'p~lln1omu}' in~I~li{ln

~(j

CI'P']ng mlJ il]VL"r. .... ·~lchniI1g .~l]fII,1Lrnlfllatm}"rL'1.Climl ...

l)l~.•!i~tI:tiri,lc' ,

bl1lt'lL"r.~111. TlkF'cfOrt\\Vc'

IFig. 2:., iii ~sWlQ~W .andl Illlr~S!em:~'of b<'!d:er~(!irn tile lulngs (If mrii!:f d !Jril'l9 'mIle pnlQw,essiiJIlII ,of jlmlUlffionlic. pLili!1I1U~,Mk!~M'i'.e1N~ iDlf~(!t!id lillltJraln.a 1;[lUy;' atv.arri'OUili, tJ me5:, lung:li, 'I'"eril!' iiIllIilGI~~dI Ii!l!ed illi1i'll \~ilh 110% f!~IJ!~r!!~;bl!!rr~r,~dl f~~mli!~ii!1 ;:!Irld emh-dd~d 1,1'1p1!raHi!lr.• ~n~ S"~mr ~{tiQ!lJ!i M!F~s:w~m:~ with ~,e;mo1l!tox:y~in iJnd '!1o.snn QW iI y, p>f!slis illiIl"ibiJIdy, ·TheiliJ1lil9~ shown aw·!!' reprll:selltllw!E! ,af eli:'jl!l1rimel1ls r~peatlm twice. SC,[)11.e rs ·indl.cate' .200 JI.IiI1I. ba

REPORTS
F'!!g. 3. fJQg!$~~FI of the~!il"' A Hilm m1l1oryr'~llsl! 10 yo pem'S' 0092 or t092 6p£(J. {AJ~~h:e w@'re !!Inrilflr,oclladl or ird@[t@d ]111"'
trtU10lQl[V"mcl at Olanous times,
~Ur'!giS, ~@rI~ rl~m'Q\Ii@~ "Ild

I~H
-,'(;09::1 -IOOil~: ,Opt.

im"'

liIler~d lr-;l(ted

·U'iRNA!l.ateil'

Woorn'i'.cm,

!'I$ai~ed" <lindre~ative (yto.'kil1E1 Ira mlsCI'iipt 'qjU'aiiiilitie-~ {oml~~red\lfith, th(iise IC!f llnl!'diEl(t~d mf(@ WEre d i!'1!?!" minedl by QRT·PCR us-in£! Hlle
l~fS~lfa

and

DO. frlNA

Ulmbi'UII",
WGSB'!:·

llll.Ct !i'!1e~h(ld(22) gl!l~ I1iOIF" Iillalilzi!!dl to 9 ~y()erald'dv,d l"'" 3= ph,o:'i~ha~e ,!rlehydrogenase for ililf'eded mice (\J.llilit~,. II." iin· 'terae'lIk~r1; MIP" mauo:pl1agle ii'ift:i'IiM1'aU):ry ~f~t6ln; aildTirU.
tUlT1a UiI@C~o$~fa dIlr.

r, p~s#"s 0:192

(Illal~k) (If ~Il"

{B,

PCNA a.rllibody '~wfnan;d hemato);'~ID il:!'JullttlrSltain !lJIJ tulf!giS in~@dlsd With Y. p~!'i:l< 009'2 or '0092: ~(i afier 48 hours. PCNA-p!»ii~w~lls. (anow~ are presellit 1'n lITe wn9~ of l:Jpla •. inrecJte(l mk-e but !;;Irrgelyab~iI1~ in the \lIil.d4ype iIli1"utiion" ExIlra·· C:0924~d~(tlilcl li.m,g~ oorrelatf.! wiiWil the Pfe!le-J1C@ !Or Ib~d:@fia. 5(ale b.ilr~,50 ~m.
rnll!IcleaJr '£!Itarnll,:l'la~ ~t.alr1i~ iuntliie

and ,0

DNA symhcsiis (N}. \;VhOl)e<! s jhe (lei I:,;of\'",Hd~ ~ypC'~nfcclc[J lungs were <Ji~1110S1 lJ.mifor,~nly PtNA-m:gal,ivc (:r-ig. :;:B)I,llal'G,c numbers of P(,NA.-posi~iv.c cells WOlle presetJI ~n 8,rdainlict:t·CJd mice (FIg, Jel,. indic.uil:~~ <l!::~hccell pro~ it~'<J:lion and ~Jll;r<J!lifln (If ti;;;!'ILI;Cinmhc Iun~:\.L~n~g !l.;'p;li r O'It tl~bi;1,1~(,l o r~hc ; .~I:IIQc,i{lni.. fi.nth~r (..~\'id!,)l1t~ [h'l~ ~~I!'l~ilic.~ ,Ulllongtxp.I(~i:I'IOC:-cl0d lT1ic~.~ru Il!lil. ~, COJI:>lX!lUCrl~'~ ,):~rw.~,)', oJ in:lrlaml m,Jtion or d:rI!r~; butiastcad are lhc I"CsLlII. .. ~e of s,YSI~lilic :>pn.~ of Ihe OO,cI:Crii!. ~ P:lia alone OOllOOI's the ablillty of ~~'peMi:. to f ca.lI.IISC PIl'C"LlI1fbOI1rIC jjIJg1le. we hYII~olhe.s ized dl:Ji1 C'x~:iii.:dmen~IJ:~1 i!~t1li:Cl~OI1J pJa expressionmid-

way during the aborted pu~molli.uy disease would be $.'i.l~rkicnl to hJl"lali:lc IlonpJ1cumulak intection into a pneumonic oac, Totest this, \!,'C acllar~ecl ~lIcletrncyc Iine-rcspcn si ve 11ro:lno~cr
sysiem ( 15) to exogcl1~~yoonlr>o!J gene c'xpres.;~

or

i~l)~l'I!Sli"~dun n~ intetlio:n (MalC<j.:bl~and MO[~liIi>l;h:: figl'i~ S.~ ,~~'Id$21. We cuit\lf!.:)d !he <!Hld j'J,.ph~' solnlam of ~~ p~'_"li.'i; C';llrrying [Jirl Llilld.,r ~O:i'i[ru~ or the 'it:i['ac'ycl iuk~-['~';.>;,'P(m;si V~ p'ru:ri'iou.l::r
s ~'l

One ~lYBlO~hl;."5b the ~ncdmni)1m by whkh {Dr l]~ 1)1:!:~is(iOn 4f)[(~ P~"I.l/a. stmin YPi 38i) In Pia ~i:u;:mml~s Ihe im'l.ll,\.i\ic cml.lilrC'of l<: I'~;sli:.~ls p/~,=['C~ Ilg eondidons U .e.• absence 01' ilI1hyda:olclmcydinc. or ATe), l'hir1)1-sh; h(lurs a:I:\cr IIm11 he protease converts host plas mil10gell in~o p!J;).'lmin "vhile dcaro~il1g [I~c 1)I01$.min inhi~Tto;r intrn!liils.d inoc~lllmioll. we lndaecd f~i(f Ci'>iPlCSI."l1;·mltij}lasmin, rcJJC<~i'iDlg bacteria t~{lm the en" :;,ion by injcc!.ing ATe 1Hld dlCII :tbl.IQ\\',cd the u ,Ip[" U 6, J f). rmk~d. !'l.'"C~nl ~)~.\ogr~~'i,io~~. rec!~"n,~=t.u;:H;rj." tn~hc'p/(l· !WIl!llCll~ 0 r~hc ~u ~.\CIJ~,*:;",d M,ltc l,; .. wbl~~lll,N;l !1,[mpm~\~'i.i\'c cvid"'!I~~ Illb., :;;~~{1\:V1f! l<l~ t~bl-ij!l d!.;'P~hioUli:;; <Ill U 1"I1I.i.'O,m; ~) r iU~mHml,': conll'ol. :tor :[.II VOIa:ih.llllig iuar~t~OI:IUlla u 'l:n,tl'm.I.T~r.miilar lQ 1.h,1l or t1in;; import;lI]l {,1)1 of l)athlJg~~D~ (1./5 ]I)). ,md lhlWi I.h(; ~ub... rc f.oqIJt.!.jIl ji.x:tcd fI1ll e, ~l{l~"C,\'I,~';OI]O~ Ale \w:;. ade :;;i on 0:1"ihe {'{IflgulmioH cascade Il'ifly be fL IJolllbk: Ill'lillisl.~TtlJ ilnd :12'1.1 "x.pn.~~ioll \.... up-regtll{llC(n, C ~:s ,i['"ul~llce rnccl'h1lrlis:ru ·,oflsisLlC:l1i .i,d'l tti1is;, we w the comlitlQrI of these mice quOC;kliy cenvcrtcd show thall~lc pl<L'lmiiJog.cfl ...tCI,ivmin,g aCiivity to a disease C\.. dt~ all~dlJc l:CaU.ifCS Or[)IlCllUlllO<~]k plague: rilJ~K!'~:Iicililferntkm or bacteria W~dl die- Qf !filii :is essential IA)Y. In.:'Mis vlfl!.llcnce in the p.l~DnOrl3ry )i,~i;s,i_";ll' '(l'1itll~(;rUfilli and MC'L~I(X:":> fLiJld vclopn~clli. of vlisi.llIc ~:ni:croco.~Oflict>Fi~, 4., A ( (11~(1 B,. unresuictcd illllf:lH1!t.I'UIliOry iinfilirni.c. [Jig. SJ~. Addliti.onrnll:y. m'lJ'il:l(ogcl~) dCjJ~ilion tissue dama,ge t Pig. 4C), and &iIGriicnctilim,c W can be ,dk:t:ecl.l)d ill Ihe !lmgs (I r mike '~fIf"e<:1cd wiih diller the w~lldLtype or the t1pJa strain, b~l dea!h, 11in.ls.the absence ()i~'.PIa sla],t:; dlC'dc\lcIIhe p;1UJ.'1'11nde; x ~en~ ;fir librin( O.g,iCfI ~ imnma G~~nle~~I, ordis~OIi;~ in Ihe e.9:~lyanti"in'n!;Ul1111![1iLmy ~~osL~il1 is s ~b8la nli~l~Iy a~~!:rod ( fig. 54). We ing '~~la.W dees !lot eliruinate Ille po'tcJl~i.ll or~h~ mit cm~lro'! exclude, hO\VC·VCf. lilt' pOS8 iili h'~y [Iwt {1l:g'lI!li~11~ C•mse pnl;'ulVilonic p~'lg\~c. lfll.irwalc· W Iy, 'nh~hloot;;k~n !he rmgrc~~l\ii[lrl iuJ(.-'(:lLfl'i'! Lly (lilt of O!hill!i" '.;u:gl."!~' f f~.<1 OIeli \'i'~}'1]1,(1;:- <11"0 'C(unribu:rc ~,('Spirn1.Oryroute i~,:;olnlP],l'[dY[\(~'II~!Nibk by [he 'In I]"K~~k:\'cCklpnt'l.'I1I!. of prrir~.laryPltL'UIIilOI~~t p!;!:gl;l~. ~l;.ilil~uhdJ!.~, 1.1.1~nk (if Ill:l diJri~lgj'lnciIUti()lilic r cxp[\''5Ki,{lii~ of Pia.

,t

or nb!~

5,11"

REPORTS A,
]00 •• FI_I-, .,_""I_"1
aDll il!liC~

._"'I_........

1j:),~pliIJ' +1I:1i'O

___ !P\1It:",ptilll' ·\iD!il"C

I

'I
'I

P:l..r~' ~A1iC
~!Qt"f',~, "'oA,l'(:

'i "
i
ij

f:

!lio,'1
I

..... f~,lPtll\ ..J.,A'.ro

.0,1
I

:;!a'1
2J:1'~1

Q~!~~F9~-F~~-F~~~48 Ii! 7.2h

il~MItq

:1
':a:

AiCG':!li6!'1
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s
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.~

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,!If
,4,

'1

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I

C:OD!2 Pm·,i!tI~,·A,l(:

.' p!!il!,',IP'/~+A"O

iii)

211, :111

,t

40; hi

72h

,IOOUOOM1h ,AiIi'C@M,111

\\flm~ iirrlfl'dMintJrilD'iaSoi!~y ,l.IIith 'y: pes1is C-092. (b1ack~ or (IJ'92 8p/ll + 1~~·plllprepalred in the .ah~f1(:e' of AiI'C (~:ep. ressed st1litld. AD.E; 3,6 hcurs, mke rlilre(~ed~..mhthe iP~~·p/(jl stlraln we re <ltlmi ni5t~['edl IP!limphat,e·bl!l Ueredl ",o~liIlianl (PIE!~H(bWe) ~r A1i'C ~~) by it'U~I)~riOO.I'I~II, il'!aie<O~f!I bid. CFlJlilil~h~ !l!Ilfl~~ 11\) and ~pl.e~.rI8) W't,r'e de~MTfIinet! .at. ( v.ariOI!l~ lj!f'iile~ S)rmboll5. (Inc! lines: as 'iii! I"ig!. 1. () IWI'li!j hi~tolc~ or mioo i!llr@!l:tEdl!With ~h~ P~~~~p&:lS!trainCj Uf~teill ,,11I' wlith er ~L1I119iS 1N~,ril gJrepared as [OJ Fi:~. 2:. S;c:;i~

Fig!.~. Contro1 ,(!If priimary pn'~L1moorik: P1agILle! rGgre~iora by p ~,~ 'te'tr:a.qr(liI'lH~por!r~~~ jilrtilmoter ~yste'llilif! ¥:. pestis, (A alld IB) ~m:il!l(~en ~f Fi'!l! d!.!rilll91irl!nlil~~i'I~, ililJ@lctiert Mic,e<

'Mpts

.:tAre

0l"1'S, 200 ~um'. I~D) .~!.epF@$:'Si()nti~ pJ~ e;.;pres.si(J!l mtdr,N'a¥ !ihrOll! gh jIlnll umeni!;: illa'~l!!eirdE(ti(lrt Survival or Hll;(ii? iD'l[e~too wi~ 00'92 AprrJ piUIS, PlE"pl~illl ~e p{ooliru:!'llred stMeror Ihe dl!lrall'i:t1Jl ,or the e;(perim:'€nt (blil(k} hean~IifL'It to ,death (MT'O) ~ 3.1 dalfS,l, g!rl till? piff'-repfe5.Sed staill<l mo:r Ih~ duration tl:nlue) (MID '" 5 J, dia,y.d. or 'in ith;e- pl(l'".l ndured :\jla~ rOi" the fir~t day 'r!Cillowe<i b;Vthe ,pkl'~ePfe~oo SWt!il f·~r th~ :rcma~noor oif the ~;i;p@r]liIIIl!lIIlt:(r@dl~(MTD "" ~.6 d~ys), Se.(? lI&Iteri~:ls. ~l'Id ,Mlmds r!Or detam~'l'tiJ~' bar' ~Ii!~thr th~ gr~f!h i:!pprQ;I;i~,tE~ th,~p@ri!'lld 111wh:ih p/tJ l~d'l!Citi(l1fl ends 'lII'l:d r@p:.F'8:Ssio.n D!!liQln1s•

pas .~rc

pl~uc nllll)" ~1C'~P 'c.xpll.l~n how ); Pi~~lf:;;l'Cqlllin.xi !

Ill,;; Ilb~ Iliay IOeaLjSIi: a mpud. severe re .. ,p.~rmOr-li infeclion aud, be IrmlSll1Ii.ucd ~ronnrcr.;{m to

p~f:j!{mbythe

aerosoJ reutc, whereas :~~ p.owur/oc.

flr~n-.,t(}sis ami :~:('m(f:m.wl#j(~~did not, l:nt1;:l:~

C':\.tin.g~y,lhcali~'Cd syndrolllc we observed 'whl~ ;)I.. P'.la·.!lei!<'Ii~ iye SH<l~n i.~ ~ 111iI,_lr~{l'C"o1I!it" dr,.":i(:I~ip~ In!l!'i li\[Jm~h\C '~,j,1Idy 20!JN,~llt~UY M;,1!nt;:'llWla:i,H~
(.1]1d~~m~lic" which :M,;n)t\ol-u~-qll.iR".{j pblg;ul.,~ .~ in

ins the ~nfeclioll (:F~g.4.1)): in faCl.lhe k~nelic::l, more closely resemble the'll5\: of fl ~/,t~ ~11 leeti,on, Tbis SUgg'C5ISI~lil~CXOg~110HS iinhirbitlloll of P~a dllJ'illS 1)~ill~1I)' ItnCclnnonioc 1~lagLJe11m)' indeed prolOng [ile :i1l1lViv:l1 of'~bc:lffC\:'lCd ind h"id~lal,c.'l,p'Ul.ding ~hcwindo\%' dlu'.in,g \vh ~dl au(ih Kl.iC'.~could be .8~I:cocs.'j~id ,~dmi!li."h,;T{,\d lQ Iy ~1l.;'<TtiN di."L,';l!\ll;:'. t Rer.ef1l'fiu::esand, IN(I'I~$
l. R D, Pe~I)'.JI,[I. j;~lBer5l(m. C~'fI,M'K.fobia1. !!lev. :!i~. 3:5

1/,. sanno:lfir~ IL v. S:lnioif1lOl),. r, til. i(e:!~OV, i.G, Dm!dlllll', A. p. llini~!'OO~.J., M'ed. ~~ 45" 04.,(I u'996), 12. 1. M. !)all ef Am.~, rrop. Mw. Hyg. SlllM U99I1}, B. 'i'!!. W. t'U!i1eom. S. [I. 'C~!lT~, v. L iWl'~r,~, E. GoI..d'rnO!~. ~ Nali ~(ld. Sd.~lSA l!!lZ.~'1~6 ((000). 1.4. 1.. Eo Celi5, P. f.lI~5lI!m, A.C!elil, H. V.. Uiei5ep, 8;, G>es5er, ~ ~,gIj'; ~~@,. 1 11.5- R. M'l. H. !)I;Ji~Ji;I. lIlr,Jd~iC .~dfb· ~. ~5, 1l!Ol (l99:n.

u.

6'..

nwm

!;u:hcd in rm£iJ :)~r:>Lilix~Iol\: ;;~joc/al .ILlng d~::;c,~'Se I could (fCYC~op lhe critieel role for PI;a :i'LIgg(',~t.... l~lm j~s iiilhibitlon could om~ra.lhcmlX:lJticad'I<l"t~~. P'lrlk'u~<lr~y because the ropM pro,~·>iOIn rn~~lJmojtt~ plrtguc leaves .IliuJ:,,;: i.irli~\:for ~nccl.i\',~ i,1iCa.tnU~rlt \Vc ieslc,j thb, c1>lflCri~llcni.aliy by inIccting mice' ""illrl the PI;-u·j,l1dm:lbllc }: t)t'stl.~ stmin YIPI Jf.l prepsrcd in the presence of Ale, \:VC then provided .t\T('lol~le nnirnals ror only the liM d:l}' oflhe inlection, a llowing the KrC ~obe eleared and t,t(~ e",.pl)C'~lt~{l:I~ o be repres~oo ~ for ~h1;'lti;!In .. illdcr ,pI' lh~ C1!>pcr.iniClll. Th" tUf]I!C

16. I\\. '~kkOmen~~",t,

{.200u.

tirol, Mi,I'~,

411)', 1I)'i!]
!!i';!'. :1!i'll7

~2n

U9'i171
2. T. 'Ii.. !Il\!II~by f.r ~L,JaWlA 2:83, "all f.!@oo)<. 3- Ii t vlb~ 1. B, lII)i!O~•.,W(L ~; Micrcbit)/. ~9'. 6?
12001;;),

:l7, Cl J!L :S:m~~ndil' •. I. D. Gogum, ffl!~ct OO.rr:loo.

U989h,

113. !. It::. ~~~rl'J

~r., fJ!je,.,;

1l'Imi!t!!',

39ille

or

~. X. Z:. Huall1g. Nt, ~'. Nl~lid1,

5,

<I, ias tlOO6:t IL E.. 'Lin!!lteir",. im fil~~l(j

t, IE, lil1ldler. €Jffr. Me'd.

11if!J.

,rJf!IJ~;~all$~jj(itlik~tllli\d

..~. rut:~s, M, fJ;JI;!'dIJ" Ed~, ftifLor & i'r.mois." IM~ Yll)11;, ~~). ~fiic H50)-U5>9. 6. T. LC·cy~. It (. Aber, N, fn'{jl, J. Mea 321., 16 ~rUR!i~,

n:~a9~.

7- II, IPliiill!lliU.,e( ~t, li'~~lfre ~'~l, 5~;I {~OOD,

Lln'~il dCl,1illl
M:iilk:

I;k:lIJ}'cd wh~:n IhcPlu-im:l~uo~'iI ?Jib. Mvl.it'Ji!:d to ,a, .P~!I-p:pT",;:&<:.:;d ~13'~C Jm'·,

W~IL~

8.. 0" A. !iodeii!1de er Ill" ,~W!r1l1'2~liOO4 UW:::l}" 9. !'i. L. ~kM, A. M. fri@cl:wle~, IK•. 1. 10a,,;~ Mrtm1l.\ Ptlfii~ 2it nt. ~199~t
:tb. !j, Wl!t~ e~'1Il. V.:lrtmi!:1!1I, 220'

S, T. SmHq, t. fIqIo. Mal. 19'7, S!IJ(1.~,OOll ,20l M. ~ !'Lid::,N "", t Clin. lnl!!:'Sl. l.H, :~,~6 {mO<l}, 21. 1,..'1", ?hi• .!! rf~ Qn Pla,gue' (i!,e~91i!~ li!f N'~ntii'l,!:,. Ge~m, 1l9'2:M. ill A.r;~ Z2. ~ieti! EIiOlr.;Ie.rll~in,A.8I Plijm 7700 Sf.Q~ePH' ~~~rWn $~~ VW~~~ l tI¥;wll~ B!i~~.lerTil', fe>IEiI'('i.y, 0\199?~. H. !j\Ii?iIII~ nU. Wlr~~it l', ~!l.':mr. OlImdl], \I'ogi!l r~ help!lIlIi dl5l:.IJ.~icm; [, mlil!1n~ forlh~ Y.. pesfi~

(lOOS), 19. L. l, ro~nso.,., t;, til, Ii~~r~m, IF.M. SIW, '1\10 CMn,

~~ffl~'

wm:a

DiisGtl~
]ij]t5
Ij'j~.

~~iM!1;

"nd thll W~!]i]ill\l~ i!!J!lwerii'ly l!:Ii~ II!~ardi (!X1!' ~tllJiROO far ~i~iIliIO!ioc:aJs~ntm,. Wili5 :5IJ~rXJrtel!! b'j ~!Ii! 9r.!ol ~.~ A!tI!itWl

512

REPORTS
·to tl1~ MiIil\le:51 1l'e9X!nal finler III iE);CeU~~e rllJ e I1Jt'!lddelii~~1'10!1 [m~rgirn~ Irlfedi~~. [)i~~~!1IR~~ ~MME~ ~1i'Id' my NilH gr~i'I!. AI~n9B, fl\~ OO:RCC i~ ~'i!JPIKII:L~dl NIH granl.Iiilt::525i4. W.W.L 'w,a~ ~~pportl!(j by

m:p.pllfled b~ 111e "'Df! lm;tjMilI:[!!I1 NIIlSA 1'3;:2: G!Ml7i'J61 I~ 111,;: W~~li1gl!!):n '1iJ1i1il!!l1i'~' S{IhIg~ III ~idinf!.
I!l!\i!~

!:lOO Je:g Fig~ S~ tI(I ~4

f~bll!~ S:il ,~li'IdSl
~~feJetI(~,

b.~tile aiPi')J!tJr.m~#ti~~~ FellllwihililllWilr,iWI d' ~~~ MRCE,lml'! WJI.!i. tillld l~orndili~l)n" j),nmlhor!' NIH N!.l1l1l*l'iJi. R~eilf<cih 5efVl:l!l' "A~~ fN!':5A) FlI2 M{!I6%8INIL I'AP.

Sl!fPlil'ordn!li Oi!'l!i,lnli! Ma1erlal

nrrr.9I.[en~l!ID~gJlo~~'lf{OO!1JI!mlll!!i~1R1'!I.i'.!i;8:1~9JDCl ~a.tl!l!iia'~ a!1!dMe<!bOO~

>6 Ifjc'ttmDI!I' 2006> <I~(epl.tJd1.4 IrlrtlMlkr' :!Il06

1O,1l!261s;;;iemc~.LH i19:~

A, Vilru:s, inl a FUinglus, in a Plant:' Thlr'eI8-,Way Sym,biosis, Re1qUlilr'e,d,lor' Thermal Teleranea

A ml!~'uii!liSit:i( ~srodi!Jti(J'!l1 b~IO'!'~@~ (I fun~lal @lIId(lphyt@!:!l'ld a tropi'Cilil iJ€lr!ic gl!'~~(!Uil~:S
'l'HgflEilll:;i'll$; Ul' '[grlI)W oiilt 111)(111oH t(!li1!p!MalJUlr@s" s chalrgC:l~ri~ooa VhlkS ~r(lilli1 th~5 fYI'IQ'!,1!S tlili,~t fs, ilT'iI'(I~~d in ~he mmUglU$'t11( llill1era(liJ()n. l't1i11ga[i5<l1llal'€'~ ,c,uredl of th~ Vb~UIS1'Jlr~ ~l1lalb~~oo (()Fif'er 1iiI,~atlol~raniCe, ]bun l1e;;llt tll,ler a iii c::e 1$ Ir~l(lre,d a rl~r til~ viOJlsi~ rreimtrtJdUic~dl. 1 he 'lliirlils~il'l~ected fungI.! scolII~"il'rs, h~,,!'too [e-ran('@ !l.Qt(lnlYll)il~ I"Kllti1o''e rIlIClI)!J('lOt h(ls[ but ats(I [I) a f1udh:()t host.

VkLiS.

S.2~,lli~ .:U-I'&i.rm.!!\<llt~~'i! (RNA ].I'~:; in\!o.lvcd in re,jJ~ieaJtiou!. rlk'>both of k~ ORE's lire :snmnil~i;

w~

1:I(JI~n

identity wi!h ~hc RdR-po or a virus or d!.e .tln~g;al I"'~!hll~n IJisi.'ll/u ~k~'rlj":tft"h'(J. Th~:.;c ~w·oORf:-: o\'("I'I"l~ and eou l{l be C;NIPrcSSOO .<IiS a ,~i n~glc (pK'I.'b..'in lby Ii,Ul1Clll1 iIUn:g_ <I" comillon C'x~lw.lsi\Qn :>trn:(1)gy or v~ral ro.J~lic""!\CI1. lho lWO ORE... or ~ti\lA .2 have !m ;;;imii~~rjlyto 'W!)' p:r~will '~~Ih
Ikllm\l~l~ul!clioJt 1\); :~nli'ioMd;->RNA mill),(imi:I'U~I.~. I.l~c: 5' il".il1di> .(2~ bp) bmh RNA:> ~~re

~() vhi"1ilre~JiC&"lC.~. The ilirs.~. ORFIII. hill> 29W'1l tlImillO add ScqH~Jle.e idcn.tit)1 wilh lIPlit!llivc ~rn!·\-de~;.em~dCiu :R.N.A I"'i'ill)'mc~c ~ltdR;I~~· lrom lite rabhil. hC'li~lor.u:ht~kdisease v:~ru~"The aani~lo acid ,'1cq,UCI1.CC orthc seeend, ORHb, ha... 33% .

which

:SI,Jln:g~$t$~~at.

ihe !;i111:<iertyiog med'la'ilismin,~ol~s

pa~ln¥"a!f:i cl)f1l~r\l'ed

b~M~i'! th~~'

Qr

b\i(l

conserved,

V~ru:s particles

rllrif~(;d

rrom

~ ~()L1PSof pta fitS.

krtlQW~lmutu",lisli:c endophyll" .. EJ.McMr~ll.!;1j.tUtfN,!, IUIDP,h),!ic 111118<1ooml1l1lO:llIy BIlOW \"~thitl , I)illllYl !issues and 'C";lH h<~ !lUllllIa ~is~ic i~~ btl! 11.I."1pl'lC1liOtYlJoC' hills beCIl associatod w ilh Ih~s. ViL'If.!s(9t ~omClcases.as Ihey allow p~;ml Jdalllation FUllga11virus g:cmome.~ are cmnmonly cmllW C'XlJ:1e~neen.\' immnCtllS (I~ A wlrml.- Jiulgill pused of dcable-suandcd ])tNA (d~R.NA) (1m. ,~ytl1b[osis between a!mp"k'a~ panic £I'JSS [rorn ge<nillCnll;:itI SOl ls, Dicl!mltJl~/hml Imw:!hw:iUm., Ll:u:ge m.o~'Il;lulcs 01" dsRNA do IlOl. llon'l1l<l~:I:t O(iCtJ!JI' in Illlijl~1 cdL~ and, dlC'ilx,-:Rnrc,nheir pl'~1l1CC and jhc ~lIJ1gll:S OmnrW10 I'mWI1.!;tI'fJ{tl<Il1nW"s i~ s i~v-! or" viral i~f1;'Cl~~I:! UKUJ:lg a pn;11OCoi a ,t~n bmh o~OI!~~is;l'I1\l00,grDw .,ul~l:igh:sJn~1~l'l'lC'm~~I':1.'" I{}j' rltlld'dl: ,I(;id CXIIl1IC'~llCm wilh~!~ric'hm~[llror ill YCHOW:.lIOIIC NaliL1lm~rmx (YNP) Ln F~rd d::>R fl. un ..\lie dl'[I,.t:led n.hc 1:iI~.es!,:Ulec{lora virus .1L1'ldiaoomlo:ry CXJlC'r:inimll~s have shown I]lm when mGt zones are hCiucd IIIIp 10 ,6:5"C. 111(ln.~ ~n C iJlu,.. he-mw. The dliRNA band illg prm:C.I1r1 t symh[ol.ic pl:1J:1IS ~hhcr become Shr.D'I'dcd ::md cOllll1oislSonwo ,$GWTilcms,afalxll~~1,2: runt! 1.8 kb, ell ~-orolic or sjm,pl~idie, wh(!r,~assymbiotic plllilllS A :smaJllcr scgm~m. less I]mul kb in ilcllgilJI. was wkrmc: !IlK]' survive the healreg·illuc. Whenl \l'ari~blc ntl 11t'CSClloc lind size Utl the isoletes g rown sepaf<Ltc~y,,~lcit~ll1!l"lhe fUl1glJs nor the platl~ analyzed. and. l!ale.', was cO"Jlfkl1rooo lobe a subalone :us,tilMe to ,giI'OIill' 011 ICmJ~ml.lll"C'>'lboVoC 31{"IC,. :gclromk: clement. most .~ilk:dy ,[I delectivc RNA. but s}'il1tblmicaUy, Ihe,,. arc able 10 lOk:i.'81C etcir (t.ig. 51 and Pi@,I. to. and .I). Us~ng lagged ~mldol11J hC:II:am.cf p.rbncl:'S. 'We l,rmISCrilx:d the v.t1ed tCI1fpcmllwcs. ,~Ildl'C' aJbfiCltcc -0 f ,hem SliCSS, symMotr:c p~;)j~HS ~liltV>C' cililall.ce<l £mw~h ralC v UL'II.IS reverse Irn~l.'Kr.iJ)1n~C (Irn. io now-eo by wilh '(amp,,~d \vllh nOI1=>ymbt()~ic ~dmlt~ m1icl also mflll,~iliic;]l!ion tlIlld ,ln~~Ing. Seq ~cllec <In.lIysis, 1)c~~eoJll'edIlml each (If ti~c UVQ :RNA :;;CglllCnl::l show ,~igniHc,ml dfO'l~~1I ~oil:Cronct p~, COH[:!.JIin:..... 1.\"''0 O'fJo1;ln l."C.'ldi!1:!;l: Ihum"5 ,(O~F:-;) ]Fln~IHll vim:'I,.'l'i myc'wif!Js;~ C'II~ Hwt:hlll\II,C or ff'}kU1l'.-i~ll:JWd !:\yrnb;.iQ!\~~':\, 'Dl~ l]!..'Sl km;t\\'.!! "')!\'lnn.~ IC}k~of Ihisi~ the hypm.iu,b'i !l:m~a~liI,."l:lI,mk'!; ~hc A IFi!il. :1.. I?r@Jj~n.(~0 r albStlnc@'(jf v iru Il."libtx: (h),p!J'!I'iJ;ild~lct;), rIll!.: dl~~lnlllbj ilghl. bp C1UIhTV in diirl8i,emt flilrallili:'i of C. ~.i.uII.gWl~. OJ'fJiffJm'C:hit~ Imm:;ilim t J f- \finf.'> i-.cglJll~ • p,r'O'wberat-a,. d~l:e(ted by el.h'id~ lutPoiUl ofh'lYI}MiruJ.«''Il,CC hm, l'M:CfI00 It~nsn<litcd iil]m br,o m'i'iJ'e!staillii 111 9 (PJ, C'xpetimcl1.wrllly hl ~~e,,;m~O'lh~f pall'aogcu k; limgj Morllmem bioi ui5il19 IRiNA 1 (lJ) {5 ,~'l' HOWC"~'I.lit. dm' cn",._-c~ of nmYC{n'iru:;.._~,>mb a IiIdl RfiiJA, 2 {() tfa I1stll'ip1lS ~~. nmliiliil.l:is~.1c ,lim!;!11 egttiophytc'S j:> lij'likD~m''''l, TI 1i..'1',1.: :thll! 'IoIinls as: probe!>, and IFI1~ is olll,)< lDi'lC report of ~,nnyef)\~ilrus.h:m1 ihe w-c] ~~ I

c.

pmJjllA-'mlcj are s:imiJilr 1.<Q~.hQS-~ o~her :LUI!Dga:1 of viruses; 5tpl'ieric-aJI lind -27 11m inl diameter llf.ig. This virus, i,~ trnnsmhl:C"d vCltiC<ll.lIy in dlC coeidicspores, We IlroJ'iIO$Ii! mlmin~ this: virus ulmlJi'mitl ~~Ic<lnm~ lOic1L,mcC' virus {(,lh1V~~o

S:n,

.~tshost 0 r origin and hs l>llclm~yp;:. TIle ahili~y OrdlC rUll:gIJSIO confer hcal m':li!rnncc ~o i~s host plant is l1Cialcdw dlC IllflCSCJltt of CTI~lrV. \Vl.l'd~y,]C' iroJ~.~C'Sof C
l't'n,OC!
!)j·M~/r'J(.'I'Wu (:Qrllwi

u~Cid \I~ rus in h ~~ll the ~i~cQ1\, =IS: l;;'vic;I.t;Il1;:"d}'lhci rh i,gh co:ntt11~1,;,)ILoll of d:;;N,Ni\ b 1[--2~Igtg o:i'.lyophillizcd myCt:liu!'l:I~'.However, !Ill. isolate tlhmiub(.'iI ~mlllil scctori I1g (ChfiUgC :1 n Ul.10liph.ologyJ OU'.fL,...ild~lypc cO]fm), cmu~limxi a V~~i 101\'"'0' liiICr ofthc: virus, ~",i]'idic&lcd by a tffW oolll.!C'~urnJtio.1"nof .as.RNA (~II02 ~.gi'[lorlyophi= lizcd rnyccHmn), These t\\IO isolal.ei were ·t(fcl1llieal by s,~mj1le ,'YCqlncncc repC<li1 ,(SSEU .u'Il'Tllys:1s wi(hlwo singlCi)rttnCrpo.~Ymcmliic chfilin l'Cne-

mg.

Ikm (PC'R) l:c"aclncms rmd by s.equenlcc31lm~ysis of lite rONA ITSl-5.K1-I1'S2 ut'giOIl (rIgS 54 and SS}, Dl:'i>h;:c<uion and rlrCct:ing-1 imwrlll,gc),cics <!!.N Im:O\'lllJlto d ism p:t \,~i!ll!s Inil!Jt1c~lI;."~ ::):~ht!s, my(I I;C'Hun~ or ":hc isolmc ob:~<lin'l.'tl by ~!;.'C'~~rillgwm;;

(I

peR !dSll'1g p:rijm~r5:s~ediijc kI~

l~i1t IBlQ~ IDI:'I'l$lQII, rnwel ~Qbei1l_'S toI~e ~Qi!!ild<rtl~, ~ p~ OfillOE!b. ,2180. III'd'rnof'E:, OK 7l40i2. U5J\, ~parl'

",em., G~ 6otan~, liInjil'efslty af Wa\~m.gtcn, Seatt~, WA '9'3:'1:95. USA. ~~pi!lI'I:I'fIml ~ MOCroll.'bidcgy, Cfi!i1111i1 S1:~ M LJIJi[iii\~rsiw, ~lmll, MT 51517l17"USA..~ll!I.S., 1Jl'dlogk<ll SUI" ~r, SeM!'t'le: A 981]5" U.sA. W

~ s~ctiOI'l 'ilf th@ RNA, 2 ~O)+ 11:iM' 1lsol.ate o~lhle fllll'!gu~ obtam:ea ri)y$oc~rii'lg W~I~ mClde 'Yirus~ Wh~'I!1inls wa') re,lnlmdu(:ed iUilto the- vi~YSilr'~~ lW~iM~'thlQ~~11l Ilyphil ~n~l~lQIDQs:fiS fAro withlhe mld t~ Mft}, n~ w!t(Hy~ isolal!e ,of th~hllll9!l1s !iom~tnml'£ (Olll:ai 11~011!llibg.lllmmiic '~rragm~!It tlf the virus that hybrid~l!~10 III e R.NoA, probE! s 1
(anro.w).

rl1~ (Vir)

by fr,ee.~II1Q,,;thamllig.

"To ~!bom ~:o~~e~ppden~e!IlIG~dlbe 1

mroo~'fI(k@IlIOO1~"lIIfg

add~ess:ed. ,E-Plait

REPORTS
rfOZi~1!1 at :'lOPe, .~l!1d~!l1li;\!IM'~d to Clomalll:;li!l;:ly of ~~'C\' ires, The ~~!R"](!~ .,hs!J'!!!~~ofoC'l'hTV ~H!his. is:O~;I'~ \\,~~ ~1!JI[JImN

IYOillhi!ized,

by dsRNA. e;x.tmcliO{»l. Nornilcm

(ri,g~ ~~
\\~f!,;:

blg~I~.llg. RT-P('R
(110

(l)Crim'Cll~1I1y~hc

abil i~y of ;the wi Id-tyrloCOlnd

~ll~Vt

lI!l~~

c!wrol1~nil;;rog:ow
~ IlrOl!l!f

Qbt.~!,.Q~

wkS.~~

P'!rlidi;;~
q_",,-

\1!,0~

~'''i!.;"SS~~

11t.~l ~(II~c!:;'If;lW hy l,!~il1gUwm~~~iOu! ~~Jlwl~an~¥l f)). rl~nl"i~ll:ot;_' (1. 1!I:la!~ld Dh'l~c \!inllJ;-inf,~'(:t.,;d\'11~1d.tYJ~ !::jolni,,: of w~
~'~II"Il.!S"~1~C !$(I!i;)!A,~ lQ I;;'Gnf,~:

D. IGlIl',ugino51l1m planllsafter ~,~at·;nf\~~S e\l'.:p~fim'~,j'It

Fii~. ~2·. (1fllp) Ri~:!lres;en~:t!ti'll"E!' tin,eo

Ulermal stD~lsiml!!lati)!'$. Rhizo= $ph~lletem~~~atl!Jr~ was maru!iltaiMdi·at 6,5"'( (or 10 hQ~u:s rildl ~ :3 7°C fer 1.4 hours/day ~or 14 .clay:').under 'gffi~!Illolll~e ~mul~tiD n~, Piallli:s were nons.ymbJotic: (NoS) and symbiiotic ~Vith tm,E!' wikHyPE!' v;r!!l1!:-~nrlea.t~di!5m1a:te '(]II C, pfOtubemfo (WU, Hal! hy!)f(Jm}'ciwN'esistaililt hntate nell'll~!i i~~~ted '7f~htb~ lJiF\Jl$tl'u!()ll!Jh tlypn'-!l t!naiSbJ~(AJm), er ~e l;Iirus:-t~ee Ilygr~m~cil!"l·.r~'$ta['lt isOOte (l,IFJ;. ~BMOOm)' ThE! i1~slQ'gram prt's.ents tlile I'IU rnber of ""La r:lt:~oChlCllmti~, de<ld" and .a~iv'El .'lit tiil@ <end ,oflh@ @i(lJ'Ilrim,@nIL Ihe 5m.aU LeUe'lS ,(]II'II~(]Ip o'i"tifui'El baiFSindiCill'e' st~llj$lka~ din~r-

whh

ih1,: r"IJlglll~iOlliL:l1IIh:d lirliOC'Il]iliih:~nl ro:illi~Hlp"'ni~m"" S Q (I n~';ll~x;, d'~j'}.wtn,;n:m!ii 110111 I1ml~ymbiQ~ic- phlli\~ "l!l1d

as high :~ (iJ5 C fQr 2~\'ock~ ,(I flo h{l!u""

pl:mt5
il1{l(:ukLt~

wil.h 111~ \'hl~fn,,:c

iSi!_l~!lICof
.1lliid:

'mhe
didd

finigti:.;;

bo._·,Cm'ii:C' ~l1ri''if'ckd mtd

chlotolit

tFig.. 2). ToO~,()~l!ii'llni.hi1i.CiihTV WiS i~lvolv''iI im heal !OtCi'8~lCt in Lit'!.':'f)lall1r.rlm~1: S)'~~lbiosis..wc rein""
lroduced t~IC \ inlS ~ilW Ill;.! \'il'us:- riIX.".C It.m~1

ib

II:!

isoiJ.1[c and lCSI.cd its <fuilhyw conl:br heal. toleraace, To 'pro,iic_l'c 3 s(,'leclllble m;:ui\.c:I'.·Ihc \I ieus~ltc i~o!mcw,l.s 1"u1ii,lbn~I~ld with .. 'pC'D4 'VCCIOJ' c'OI!H1ini ~!~ a h)lgnI!nydn"'n.;iS.i:;.t:;,1ne~ ~cnI:;i (I 3l
b)' rcstri clio~~ ~Hl~ym\1-n~!,_~d iallL;'d il~'h;!!:rOlltOIl i mENU) ROlLl1s1bmmiion (1-1). Vil1l$4C{lnu.1i.ll~filg ~~'~!d4YIlt;hJgro~llyci'l.~a:>i~i\'(l (\V~I and "un~ ~ro~ hYl;lmmy~i n-nJ:su~I:oiUlt ~VF 'I i~~nhHC'-~ of tn'Dml~~·tut1 WL;'m cu.hurucl 0.1'1 5urlig~Q P~Lli d~s;h!ci>Siml ~ih)\V(ld 1m ll,nd:CI1!lO hyp~m~ .Wmoion or al1J,liS'WmOS:lIS,(F ~g.3 A). The mY0l:IIn.lIlll1 from nile area of ansstomosis \l!.'1IS !>LlbnJ.~I.urodtwice with stl1gJe liOlli:dioSIJOrcs gl'lQ\'\I!l ml hy,g.rom,ycil~. OOl~tm,hliflg rimes. T1hirly-fl\!c hygro~nydJH·l'SiSlalll, i~ollil~es o'b~liD1cd~n this W'I}i were $1!:1\.'CIl.Cd for ~hcil'll~R_NA 11jiOnic- s, but on~y one \VOIS foond~('I ~m\'\O! ;l{;qla:i.!'I.,d lhe: virus (F~. I .md JEI),lhis Inlllga~.isol~nl!. n~cw~y ill feeted b}'~I!!Ill~lliIi mmSI& moo,,,,,",\'il'i~(11]]\I'V'\.n). ' \\'O]S tested lor it" abilil)' toocrmlC-r 1!C"Jl w[cm!l'Ce hy tile same C:""1IC'~1'I'!t~!1ml ! ~p~ll'rmchindic.. ~cdI abovo, The ~1'i..';Il~IR~ ,c",pcr~ ili!ll~!ll (;OnntlM(ld Ihm Eht:iso~~ ncw~y in~t'(;lyd wit1i~ Clll!TV cm~(t)a}ij li!~ ~JI'tC Ic\'d (Dnk~lt l.o~crance as dim ocmJ~JtUiI by 'Ille w~l!tMypc ltiob.le I!Fig.2}. Pre\! iOLlSly. we immd Ihm some 00lc:i1Ic.l:a:l

c:.

'~!lo(~~rsJm!writies
l€S.t, P -c O.ol).

kihi"s(!u'a!"il:

inrIKl'ecl' ~s(ll~oo(Iff C. pro·

lJ'klll:5.;rree

IFiig. 3,. (A) AH»sto~J, of lihe !i\likl'~lPel,!jmS'"

tub'e'{Q\~"'i.WO ,~m! the ~,eosist.a IiiItisolalle

hi)!\g mm)'rill~
(lJ,~)

to

~nm::11ll1oo' a ¥il'lliSrilmf~led fulY!lromyciilil-~e1i1st.mtiso .. lkJoo (An). ~B)~ ~~(e..

5p1lrl2'Milatij(ln

p,u~ (1J~tu~ h;isolate

to

pllOOUl:e

m:; lAd

~e.:dy 'ililroctN wfthlhJ~ rttain~d the' ~~romyd~~fiOO ~nd the mo[IPrneJ!IO!!!,li'oi'lhl! VF ~iSolale. Fiig. 4. ('liepJ R~'pre~!IIltlti\lllro'"
lifiIa~o {S~ll1oom lyrcper.s:.rcon;, var, Rliu;g;~s) ptalnt$afli~rdne Ih~t-~re~

'~~lPerlmtir'it I'la!lUlWlIl!ri!! $yntliliolli.c (N5) and ~'bio~~

i1 en-

wilt! the~~~ildi~ 1l'iro$"ililfu(.t~dl ~~1300 'of C .p,rolubetflwMtt). trn!.llhygrom,yanruesi:iitalf1l~olaitE! irlE!!.'l'ly imlldi:lId will'l:

Clld~phylcs iiStj)!~J1Cdrom U10J'tOOOns coukl be r Irnmli:m::d 10 cudko~mld :sti~lflLmct[,o];l as ~ml!~mIiSb" J). Thas, we tl!'llocilhe OlbiJIily (Df Ihe ( C f»vlUb(~mJ'fl iSOL.'llliCS&Ooonl~r h~l. mtcuallc.em !ommo (.:)okm ..llJ.~ ~,l'f)pf::r.~i{'()n}"Usiing a sI~gh~ly modiijed pl'Olocol fur .he heaHI[I:'CS:i!, ~J"i.perimenl Il.l ~i,we obmi!1~d~il1illliU'1\,"SIi.~IIIS, dmsc oinlru'lleG to wilh D, Aflmginosllm '~Hg. 4)0.I ~OWc\'Cf. iI, w.... not ~ ~)(Il'h.~iblcw <lIt",in IOOi% nl~:~I~colm~~z;llio.n of r~lll'{!O !)!;Lu!:5, ~]lcl til ~ m .. y C'xpl.ah! l!:Ie

the< vii lIS 'mwu:gh hyphtd (l!l<i!slOtr'!(r~,11l@Lrd orlhe virl!l51ofree lhygmrnll"irIr~esislal'lt.i:ro~Cit!!' (VA. Iilihirospfulere

h;ghcr FlI'I:1flOl1Licm of dt:ad r~!lm:;'~,{lJcm~~..u wilh Uk; "'Ill er An flllllgUs.., C'onn~)~I'I;(i1 ..... ,ilb ~h~ (.,.pc~TIllliCI'1I1 lli:Oillg D. Itlli'lfgimJ.,:um. GivcrJ lImt C :similru-lfUl!llhlaiJSI lc 1.J.c1K:I~ts~o boiha ftl,Gn~orotarrld
iii

un.

temper atJ!,u@ w~s m~inltill~ n@dl (It.

:~ ~e~aUlr!l;'(2i6"'t~ kI~]);I hourslday fo r 14 ,~afs uf!:d~r'gr~~f'!hrDL!SJ.l 'Ci~!'Idi= .~ :I 'tnon~.. (Bottoln))' Tlhe h~$l'Og',nfl :z IPreS~f1B the l1~fif1Ib~r '(If p1a:i'i1Sd'e~~ (white) .and alive (bla(klo at tile ,end of thf! experilil'llmit lIhe ~11Il,"l~ ~ell~H5 ,(!!nWiP of ~~ ba,:s,ifldl~ate' :stal'Vs~'G1I1 dSf'lerernce5 !;iml1aritiie'S'~I~h~r~s or ,exadtll!si:, Po:: O,O!)}.
Ia)

6·SaCf'or 10 heurs and ambieiFil tern-

i!! t!i!!!
i!Iiiii

=
c

/1HHilbr::mrfl.,

when ill roctoo

willh

cn·rv., provide.'!i

14,
12', -

101 B:
6;

~OtlpS

.~
2'
~I +-.....l...~...,j;,,--r-"--"--"T".....I:.-.....L.-~....:L....

W,t

Ani

VI"

__ ...I....""I

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REPORTS
csized ttlm 'cnc._l'ol~hyte~ mij,Y pl'Ot~. Iheir hQSI pl.m~!I. by :i!C;tr\i~J1Igin~ Ih~ Ja nmg~ng reactive (),"Yl¥J~~~~~i~ (llI.OS) g~Nl',atJ;-d by !he 1)1:l·11~.
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6. T. ZJl w, 'G, .1. BotiIm!,,~,!Il!Jp'alll!~,o.JlIY 81.147 (1997:1. t. II. I!. A~IT\, '1', ~ tee. ,!!.l'a1. f!la~t Mia. 1l1ieRltl; ji,4, 496

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~~I100 ~!f l!lile Si!!1tuei :;:(II)~m NOOI!~ rolii ~:l.tiOOJ,'h"e Il'iSUfJm~mi:l!l SUoIIri9(f rm1illilit~, ~jjdl g,an'!!; rrom 1lIli~ US. 'UeolOgical)jjM!/, Ih~' Nt5F (99n'~,2 and '(M!l;1!,II6n,. and!n~ liS iI.ii'JIrlf I!~\lldl
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:~fer~Qe<i· 10!Ombu

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2006; ~m~lJ)d: 12 ()~oom'lbt~2:0!'Jt6 1O,112li15i:ie~~e.ln6.B7

The INaurlal Basis 10'1Loss .Aversion in D"8Iciision~'Mllakinlg IUnder Ri:sk

P100lple typic.<Illty ex nib~t glreal~iI' selil"litnviLylo InSiS.eS tina n ~o ,e-q,ui;valeni~ ganlns \linen ilI1IiJlkiflgl d,ocisioflS,. We investigated neu ra ~ mlT,~I~t.e'5r 1'o's'S. ¥e'rsionl while nndi;vldmllbl ciedci oo~~,etine r to (I' a ,alm~ptor re'jec;lgilrrnbl~5, that oneredl ClI50i50 dlilnce of !ililin;inlgl'~H ~Dldl'lg :money .. A broad ~t or ,alr-e:1Ii~ (i IlIdud]119 !mid bralil'l dopa I'IIIln.erglic. reQliclI1:5, d their targets) s noweclim: rMsilnS! OIctiv'it.y as, an poleilUal gail1l~irm'e'aJ~d, Pe,t,e,nt1i<lI. losses WI~re represented by de.c:reiilS]1il9 ac:Hv~ty in ~vera~ or thes.e $o'!me gahll·s.e-niS.iti'lle.'! reas. Ffn.:!lly, 11ldhrid ual d~~fei(1(.'rl!Ces]n be'nIDi'll()ralID'S:S (I\JIe'rSiOIli \'IIew'e prediltt~d by a me-aSlU'E!' or m!@ll!ri!l less ;1!v€,rs,i()!llilfl sewfal regio,m,•. ~ndud'iftl![!lJhe \I'!!il~ral s;trl~ti!!1i!ii

pcricl'lCoo'·lIti.H~y ~ tn. H. 14) ra~hcr !h.m )\pccirka'Uy in\fcsti,g<l.ti.ng which h'mlin sy~ !c.~n~l\'"'P!1.'M.'i!l~ pou.."ll~iil!.~ I~ se~ VC!'l\!Ll~:!,l~in.'lwhen. a decision is beillg mruk ("decision'· lltuHt~I),.8'1.'Ino:Jl~ ("e,l(

Imnialc cuteomes {··allticbP.[l~cdl'"tnilHr},l (11.111 or Ih:e actual experience gainling OJ: los,ilflg

or

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cxperiCll!c(.1(.L ,me] decsien unlities {l~bl di,\'C1i,'gc in 0 r.JHl>!!lic'w~)I:ii,which misc,'l the po~ibililY 111.11,
Ih~ cmli'"~,,polf~din~,brain
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dilih U5t lnjhc (·um~llt scud)" Wl' m.i~n~"l! In isollalC ael~vity ru.",",odflk·'{lWilli. Ili~,; evalaatinn ora ~fnubrc \.... dtoo.singwhclltCir or IWI. TO aCQC'pl. hcu II lb'! .• rl'cd:ii,Ql1 ilnillilY) wilhom Ihl~ cx~elil~ton. Ilml.~bc g.amb~c W01l ld be i~m,~cdi.ru.dy resolved, ~!1Id1 prehO!lliil (OIFt.e-:Il. This allo\vcd u.s to lest whcthct" l1,clIml n:::ll)m~sC$ dill :11141 ihe e\'a~m:u;llln or IlOtc:nd<l1 O'!.locmllC'5 <Ire .a~lY ~Qdskl~"'~, ~Ljc:'I~~wh(!l~!cr to h~.~ th~ 'COl~C2!..1ll ~o.'lS il'VCrnl on: Peoptcflil"C :nl~'iU: si fI1I,illarTO p!fUI,cms ~)re\'.iiou$~y l\;JJOrtOO nl1 stud iies 'V~'SI. III n.lle SIOC!.: nnmtC:l or 1.0 rn:.cepl ,<I oD'0'!l11lkip<llICdawj c;..pcrf.cnce~ £rLltoomc.."'. :1.Ci1iSitivew the ross,ibil i!)' of klisi,ll£ obj~ or new job,. :in,Yolvc thc possib:ili,IY of Ilw,ney dmn lllcy m~elodlc po....... Onc rllnd~nncnl"~ qllcslion r\ll' I~le sty(]y 01" ibU:li~y ol:gahJins g<!.i~lillg or 100'~llg Nlatll'!,"Cw the su:ltus, q~IO. Whcn th~ smile objOC:l" or allllomU$ of mom>:>' 'll. S .~). docL"ion-lfiaking Is wlu::du;:r :1005;Jj\:Cr:il,[o1l. l"C'ilr,:.:;;IS ~~ICed with $.W!ch dcd:ilirul~,. mQSl I]coplc ;]!l'C .he cngagam.'Ul of di~linct cmali{lua:1 !lU'O'CC~SCS Thl!5. ~)1."Qplle•.ypic'dHy rcql!il'c iJ. 'lwtcnti<!~ .gllln of m;1~k_c(]ly rifJ.< ,~VC~, For i~~s!mtt:!::. pr.J{lplc IyWJ'- at ~ca.",~:s H10 to m,Lkc U~l' Ipr !l:;o;;«]~~l!IIl'!,; w ;:! \,vhlC'nro.t(:!¥~~i~ ]~ssC'S "~rcro~~,,,idcl\I;,'\J,h tms lJoCcn il;:<lny v.:j'L.'1:;'t ~mllb~~'!ii~1,1I (lfr~T ,I 'O/SO ch,..l"U;;c \.'If ilah,'nu~11 I~'i of S.50 'lx.\Ctnl~ !i~c ..u'bj(.i£,'~i\'c :o;ill,lil1:J.;':o!tC'd Im! '1;IlIh'lltc~, .fi~I~!'iiti\'i!.)I lO!(l. .... i:;; ~ ~:'i g<1,i~l~ n~ 01.' ~{ki:ing.!lm1ll~}', ~Inlll':\!.'i IillC i1111i10UI1l '[im~ JriVI,"U by Ilc.g<lIiive l.,nmirYII!:'l. 8WIt'h ,,~ (JC;.ia.· r o ~IllJ}.l.cl oftw;l\I;,:;; its ruughJy !wi!;e IIhul of g<lim;. p ~uld: be gaill.li(] i:;lUI kll~t [witt 1Il1(~ mmlUl~l th~a. Simi:l~miy. P~"{)P~'~ dt-:utKmd ;l;~b~~ltInl.l n~Qrc ,m:.\:i.cly ,~[6). -n]]S lull 10111 n.;dica:-; ~lm.t CXpa'>LlI\.' I~} ..~ly ill!Cn;:tl:.;i:ll~prn~llI!ia~ ~Q~ sJm:uld Ix;: [~'il;ocJml.1d could ~ 10:;\ lC,ll}. 11 SO/SO' C:huIlCCIO .ci Ih~r g<iifl 111'l£iney to p~11 winh objcclL'i I.hal. thl:Y have ~'\:n S100 or lo~ S50J (nProi:'Il'CClnh~OJ-:--y. (~lc mosa. g iH;~llimn "!h:n[i,lcy would hm,vc been wI,n~nlg1.0 with :lUCiii:a.'>'Cd acth,ilynnl bmif1 s:truCluiCS t~{)tlghl s~,oO;:SS~LI~ ~ciha\' icYrnll model 01.· dOCiSiOf1rmaki~'I8, I)aylo <JCq1uirc th()~eoojccls in the [jrsl plil~ (6). IiO a\'C\.lim!i: ncgatrvc crnlllOil:l ill docisiollr·mak ing llmdk:r ri::!ik ,and ll1lccl1."IiimY II. 2). CXIJlain.~ tiS:k ,Loss <lversio'll abo has been li~d to c;x!~bill "', I:>ud'l ~s Ihc am.ygd!JIJ or",m~rior ills WIIll; cmnpill"C :mVCn>ionl ~i1;r ··mi'{,('-£i" 'tWliilIIOti;:.;)gamblcs Ll~il'l!!l wid..:: :Ulin!ijC or ~OIlQnli.c '11' ith p 7, .IoN ~]. A hCmf.lli\"~I:)r, :kk,;S t1I.vcffiiull could ~b'hfl\'iru::> ull:tside U:IC laoomiory ( 7, II(),., F~lrthcr; lo.ss,uv\"IT'S'iOll L~s(!>cn,in reHcci. run !l!;ymrne:triic rc::;POIlS.CiO I~'>CS VII!fS'tI!;: l:drcf1.as yom~g as <Lgc gains wilhin <L sillgl~ 'sYSilClIIl I~mt c{ld~s for the ll)eiPafil!lI!lEmt ,al' P~:!,elii~ogy. UniV!!ll~it)! of (alirorqia, 1O.~ tradi ll£ lx:havior 0:1:' L'iO~l ,dl~ slllbjcct~ve va~!1ieor the pctcntilal g<.1I'~Mc. slIch live' lV}1I!Uld c<lipuchinlnonkcys tJO). wh~ch SlIg.Ang,~~ (VCliJI), ii'r....!lI!; ii!l~,kJ~951S63, ! ~ Ange~~~ CA '9OOIjl,S-:l!.S 63:, 1I),A. ~1!i~dtu~MI Sdool of rl'0n<l~m~nI, >IS venlrolned~al lucli:OHtal C<ll"i:e'x (V.I\,lPFC)I gegls;; Iha~ illl11Qy reneel, 11 !~mdamc~~I,OIIlealllllC or UOJlo~ U.oWes.mod ~l~la. ~.&:5..A~geles:. (A9()09'H14Bl; .. how rotl!nt i<d olLtcomes,;'Ire "--~~ by Ill!:: orb~IOrl'l.'llll l c0I1ex lOf(') .mel ycn1J;Oll slri<ltWim U:s:A lBQUI!II Ilese~n;h, 1~litl,J~1!, UClA, 1.05 ,An~I1!~, (',I!i. (J'J. J'9,.2.(.I), 900'll5, 1J15.A.~!)~~liIlmlmt!1lIIP~IiJii31Jy Biooii!hilIliQra~ rl; nmlc hl<lin" ~mll To c,x~~Jno lic !he n!(;UJ;ll~ !I!'s~t;;mi!'i lim!. pr'oo!;."""s; Pre ...~ou~ neumu~n~gi.~g s:!uuijc.'5 p r rc..pi[l!"~~' ')(i!:mCll~. UCl!\ !l.O$ A~gel:~, 01 oo@9'5, U~. [0 mn]:lel)a~y ~.(1.1Sor ~O£."'OC,';S, hi1l;\I'e Ibc~~~d 01] ,It::· doci.;ion 'ulilil}', w~ coliCCled fU.l1c'litililal! IN1~nt;;n:ic: "To ~hom ~:o~~e~ppcten~e 1!IlIG~dlbe add~ess:ed, ,E·P1ait Jl)lIId r.I(k:@IJ(!<I.{1d"d ti''iI'i'tl' ~1,s._«1Ci1U~td willi, I.he :l1.llit,np~~~iCiI] or .inil'i· £Csommioc il:ll~~~llg lr~mI) dlill~u ,,;,,~!iI~ paniici.

.or

REPORTS
creased t<l!v-cmging O\'Cr ~JII levels g.1in). ITl" ft;,mQ,j; ecided wheiher t{l sccept orrqj:C>C'1 III ~~ d !'>t~';:I~~,Oil g!TIl!P of tml~!l 'N~gijQ~J.-'; !ldudi~!g ~hc ~ g<!:mib~c~~h~t On~t"O~ a ,5IVSO ch;Lv~~e or cidwf "tij..~~m. VNWfC, \~ma,l~ ACe ;ia1(,!rawdi~1 g;;'!.i~i~~~g~Jli,;' ;I1~9!)i!1! (If !llm~"'1' ot !~i~g <I!.'1r¥!hlJr :aHIDI,ml I( fi~.IA j, V/~, -r;t) ~1I.1~Ol,.lmgC" p,llli.iclll;.!DIQz.o orc, .n[l~t ofwhid~,uIS,{l~Oi;!~ fOT.g<lin is, ti~"I{)~v~d d~:cn~l~ing ~I(:Livily !Iii i:h~liilz..,:orth1.~r[},1Cll1ti3J,10!i:;, '~o R~i1~d OITIi.~h':: li:1Jjcel.i.\'c~uulrdClavlllle.. of O!td! ~ ss increased ~Fig. 2: ;u~d rig, s:n. i\ w!:!ju!lclil'ln g",:mJbk'!lilllhcr~h"'l 1.0 mlr nr~ a nx.cd deci sinn

or

Ihe bel'itif'(l;.18il~le .g... n~blc$ (S"iJl: S~4 100 $40; ~!f;S:: S5, I{I SK). ln ,<Ii wh()lc-brnr~~ ml"~ys.~.lhIlJl; were ~!Q flJ'jilim1f' l.h'I!. ~1~rlo\l!I\;"'d sUg!ru F!iI;W'I'II}'~llrJr~ ,L't;~ ,i\ri~y lor th~ WOTl'lI.~<hUIlb!k"'1L" ~ol,\pal\l~ 10 dt:JIl ~hr ihe bl,.'-,..t &1Il1M~$ t com:':C'il::t1 P > 1[)I,4iu aU,
'I!(I.",1:I

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I:CWlrdilig h;tl'",,«IiI!1.td 1L'C'livilY in dk: al'l'lygclaI1'l.mi{B ills",l.. based t.!lllpl\.'\·~ous s,tm:l~~".l\(if J::"q'h:ri~'iiiCtr] ul.i.~ity!lIid risk
l.'L\,~rs,k;lj,

~jX""I;:inc p~dic!iol1

oRier ttl fbU,r;)W for :;epa~~t: ~~i.nlIUil\''S. or It\:lil<ll re'jXii~:!:e..~ro~i"iiil;~ ful~t1 1,000000~,~ile si~:> oflhc P'OI~ilt:ia~ g..1in and loss \c\'C1:'C ITI1Ii1ipLlIatro iu~dc~ ~~.Ild.cilt!Ij', wIth £,Ili'l1~ i:<i.nli:-h~..\:!i frmn :s JJ) to Sot!} Hu inctt!lUcl1t:l1, of 52) and ~OSi1Ci1.rn~12.ill£ Ifl"ilU S5 110 £20 (in iaererneras I), \'I,lc chOS'C '~h~c mn:&,~ lil1.;'Cau~c:p~viGu~ ~:tudic.s ~ndiJ(;OI~c~m poop!e arc, ~ r!~~ awrnw.~.f;Ollghl)' lwice "l> ~n~itiw III Ii:}!;~~, a .. ~n g:,16~bS U. j~. tli!~Iz.;, wi,; 1,;;X~i:'t1,,'d ~h.;,!II:' fot mR"~ ~dci!l.llm!>.lh~~ m~lg~~ ~'l!l!nbil~ WOWlld c~ich a ('I.r wide IiTIlVgcoJ .0001il.lIdel1,.i:om M~l!ilH{:"C~p!m~re f ~o undi ITch;lncc to M!p:il~ n,:rjI..'lCli,on.To i~llrocluC{l i'fltXl'mi ",~.comJ.l'l~ibk' jJll.)'olT. '~H: :;m]Hw~ld par.., J ~ir:ip:llm,., wilh. $30 otlC \,\,.edk.lxlforc ~Jlnl1ing:.uN! IO~~I p:lrti.cipants 111m one dI~Ci..s901'1[mUll each or l}m.'C ~911l ni1t]l,rmlS wouM be hOblm,cd rorrc<ll

and VMPfC (F ig. .3 !.ind robl!!S u h~ o:rd...r hJ enlSlIIre thill, l1QiCillia~ lO~--d'CII!J1..cr1

\'"Cinl1l~

S:1:!'uillllli'i'il.

l.l J •. 1H~

W~ l'iIC'tI'QFlm~ ,U'uru\C't'~i)nl)l~e:;

ors

mlco.rre.c~d I1H'Cgho~,d r I) <: 0,101. IhCi'e were :liiO 0 I\CSpm\Sl~ wer,e !lot reing obscueed by the o:;,'cralll !>i,~.ni'l:l('allt \'o.>:cl-: h~ the mny,~daiill ruld Oil II)' 1\\'0 ll!!.ls:ili'!ic 1;.'}i.pec[I.'(1 value of ll1.e g.1mhk": •. we :>;i 11!£lc!imcmmcctcd 'l,.'O';xclh:. the iu~~l!ibi, comin B)-' coma~d acll\ II)' cvolo:cd by the \.\1\Offl! 1)O:;:~i,t~lc ~mi:>on,m 111C)1<1Il1'C: thm:hokl,~herl:l 'were luge j;;lOlmble;,\1 ~.~I!: S m '!.O, S 16: ~O!l8: S ~7 to S20l ,1I1d dU~l~r:;. of acl i\'<11 ion till!' '~hcbc:;~ 'IIC1)1U"sIht.: "~~or~"'l tFig.
'1., (A] AniUl~t'.;,b~m! ©~ th~ eV~M"re.'lgllild

Ihlll foeused Oil these ar'~li.~.E\'oetl!III.Ivei)' I

Hbe:mI

Inollcy.

\Vc assessed bcilavi.orai SCills1 11\'11)'to g9,ntlS and 1Qe1$($by fh~iDlg ~L IOgi~toc l'CgfCs$ion In each
p.ml'llicipa:nfs. .1lCCCfl~<lbi jud§lncnt'i. Ili~y ool~ccr~'<i

task d@st~ iii. Do rii ng !il"C.cih 'triall tlfJ~ IP;;!lftil:ipall1t was ~lf"esemted [OJ 3 .s, wIth Oil di:sp!la1y ')hQ\Ving~he ~i:re of the potential gaiin Un glFee'" a IilIdl lO!l;s (illl r,ed). Afri:.elr th~ .[lccelltm n~,ied re~jlII) rI~, aJ'lrari~hte iii!t<er'"
val 'was pt''elemttedto, a[lO'l'll for optimi!!l~ d!N:DIIIV!li[III00.n of fMRiI rt:flji)OIllS€S t~t?a(h

d uri ~!g ~ming:, ~lsi!'lJglilt' !l.i7J.~ of" thr:! g,.in .JaW loss as .illc.iepclld":nt vmiablcs. Based em Ilhi5 analysls, \.,,'c cOlnpLn~d a tncasuuc orbdl8J\ ~(lml IO)lS <!J\'C~iO!l i. <J~ tlilC l';JJ~io or t1!C (,oImOll.!lC) J~~ respom!:ii~~ 10 Ihc J',!!iJi'l1 1\.'1\,;P01]"(:,whic:h yic-W«l ,~
I!. = 1_93 {u;.m!lC:(t99t{) 6.75), This tlind~ i:l,'I~ .it;; ttlmiz;;l):m \.\fill'! tlsc ol:t..ttvmnmh" Ilmi. par~idp:iiillll,~ Vllcre, Oli~il\'Crng.c" ilidiffl~relit 'uo.g;JI:rnblc:s;
ml1l:xJ ian

trial (2.?).G~mbllil50 were I'!~t l~s~I~~ I(!Yr!i19 scaf!" tiling. iflhli! vallJ!lS o[ Igtii n ilnd less fOf ea;rnm8i[ ~~~C'e samp,1ed hom 'Ule

g,
5, :~-

r,V(; f~ffilillliill,yzca tl:le in~agil1cg dfIlBfL m idcmi~~,' ~.giOllS, ~~il~oscoCilvilI:t:i.oll (;QJ:IlCIOlCd with ~hc size o:f th~ l}()l>C'llll IT! g.li 11 or toss, using ram~ll~~I,~i.Cl:egl\."l\,'iO'1S (,1/). ThL~,m Rly:;;is isoll;n~ a ~e~ :~"Cgions lleo;)poJl~i\"'c too iLhc ;i.i~ oi"' IJoGlcnli;d f,_lft!.i~!~ whJ.:n r:;:\',dlI:;1!~i~!g g;1Il~bh;;,s(a\'croging ()\'tir lC"vc~ ~Q~) l(Fj~• .2 '_II1.d jig. S:21 Tlu;glliu.~ .rc¥!~n:;l \'(; nl.'~WOlk iI~ChldQ~1I{,£ ocm:. pn;-ViOllsiy ~how'II1I~) Hl'l'OciOk,d wi,:h Ih(;·,uuHi.eip.IIJol1 .m!! be: rul'l.'"liPIof rfLOIJCL:;;;I)" rewards, inc:lwl~liug [he d{lP.;al 2md vcntrnl MiclQluill1!. V[\.'U?f(', V(ntrolatcm.1 [lFC. <lI,ntcrior c~ilgtllalc concx lACe}, or-c. and
brain

garimll:k!l:5!i m<'lIDriix,. ilS ShOl.lIH ~'U1!r-e 'fo:rIMlO e.xi3il1llpl.e gi;lrnbl~~ a gCliililble in whkh I.her(J~elillifli !#I:ilnwas IWiCClhc ,fUl,mul1l eseh ceil iilil thii5 16 x 16 Oif~hc11OICil~:i:allloss,~.Firg.ll) and lha~p"llb"licipiilll'" I rma~rTl!;"'a$ll=!feSf;i'l~ d~rWCIIC slower to decide whelhcr or nm to ~CCCU)~ ~rlg·S(.aFlJlli~'H\b~uhoe~ta Ihcsc ~;:Ijwblcs (Fig,. ~'Cl 'nlC:OC: bcI1lllvkl'J'fII1 dma were col!liIpm::iinro 11 4, x also ,ao.;--o.ru weil ,,,,,,'~th 'prc."..ious [i~~dings it. . .),. 4 l~tr'.1l( 'rOfa!i!alYS'i~. f!ll

P(Cbabili!ly 0:1 aoceilla_li!~

rrom

(Q"JTbrn~~ons of ~ail!lS 1Ild

Ilm!ie'S 1!;'Em2' presented.

IS~,

or

jil1llerstirn1Ulll!I$il1it~nvaL. OJ,) (olm",ood'ed h:eal~rni3ip o~ ~robab~lmyof '!lI8,l1lIble aw~pti3iI1~~ .~~ ,ea.:h ~~l ,of ga~I'l~~os.."5 (n!dindkat.es inig:h '!fill~illlgnes5, t.tl ilc~e;pl th e ga m'b~e.and Ib~!I.Ieill'ldi~al'e~ ~O~\I'wi llil'lgnE!)s 10 "H!(ep~ the' IgaDl1bie,). ~c:) Co!lor·-coo!ed healtmap' of rel'lpo.nse tim.f!> hedllnrlkatoe~ Slmll€f Jrt'~po,n~ tj'mes~

or

!!i!ild 1b~11.i~ ~!ldkat~ra~ter lFig. 2. Whol@~br~illl

oiIliIOli)f$lHlf pgwamelrh: re-

f~sp()n:s~'limes),

SpO!1~~ t.o sire !Jf pm~!11tijill~

lo~.." (leU) I),r gllijn

dOI)3l1lil1.(:rgic nli.db.mi]l~gio~s. n,el'( wc~ 1\,) rcgi.tm... tha.t sho",",l:'iI dCCn."il:>lUl!:j ilcii vai~,orl ilS,

g.m,iu~sinCfCUscd"
.~flmsEl\"l!."j;sLotl is

Ibc~h',c lII.""Sponse(c'.g,..

dri'\!eu~ by auegalnv,;: al~ rCJI'. viSi ~1n>cc. d iro:mnin

for!), Ihell 'Om: w()I.IM CN:pet:1 incre.1.'!;ing.u::q,~\'ily

Ihc ::;i:lJ;.;

kiglhU. S~alil5tica,l maps were Ilrojecledl onilo .a n awrage wrt'G:ail ~uriare ~vi~hthe use ,(If' Im!JIttifi(llJdai maAJIngl in (MET soh\!,t'<lI~ US); mrolil i!Ii~ dllde:d t(l show lj!er1lra~

h... i.n ~iun:s a._~ia!ed \~'itlil dlC'Se CIIlroliPJ1~,<!5

sUo~

(y "" :00)

ar,~ in·

0.1' !Il.(! 'IXlIi;'.'J!ia~ ~, illcrca."C'5. COfttl:<l!ji sirla~l~ ctW-a 00i1!. JIilllf'fl'laJp,s 1.0~hi... 1'I~l;,'.dictigH.11.(l!imuil1 ~iIOgir)lliS!'ihm\lcd ~g~ fl~ !::on,lf(;ooJl;!1 f~r m!.!Ui" rI il'icmtU}, int'IL .. :;;~ing~Lt~ i\l,UI ion dl;lfong c\'l'I~lli;Jaimb pie 'C;OIilil p;ilr~0Il15 <lit th~ whme- br:tlin level by mel'lll S e~· usllllr·bas,ed d of g(lnnbk~ 'Ik~I~k:.';~"JC n.W '[ia~rotClil;jJ~ lo~ i~.-, at P < (D,ns. U"'~ Len Il~,mi~pliler-e'; IRIH..right hemiS\piliere. ;

GlI!lIs...sianr.mdem i~eld cOH'oclinn (1:9)

516

REPORTS
g~:llib~e.~in the v'erllrnl :slflOl!itlll1m ;md V~\IP:fC. A~-

~hQ~gh mill results in fMIU must be inl~flN~C<! \.,'i~b 1(:!lI.~tiOll~.t!!!~ ~I;~~dt!l.,~rq ~m~:sif'~~nl with ~lw il;om::l~l~ion Ill,., :IOSi.'iC<~,\IIW l:l!lill.8 ~rc Il;:Qt;IL;-d [.;y 1.11";:mlllc 'l~l~chanliMn r.Jn:iwr '~hlln lJy hoWl ~cll"'mlleuuoch;!Jni&'f!lf;" Moroo\,1.".i. Ih V~ 'lgl~!'C'" ,g<ltc n.;pfC'1'>cn!.~[iQnof d~-cision lI1:i~il)' 'Ipp~arn
~tl b(~,[it~rn,;:s.tiJ'i[¢d. by

r~gion s '~J·;hihi~oo <1 p;att~L1l, \"r "nclI!ml

~Q;~

aver-

lil'i~ sa nfl~

H'I:i1 it.

(:I'i£'l!l.gcod bya r;lIi,!:!-'~ ~) ~x1Jl'l~rlC'.['ic~drewa.UtI:!: (1/), These re;MlIlt~ iSul)l)on ~plte...ia us

11(;(1

m 1 c~,n.;Luilry

!i.i~n.·';~hmil'l,~he {m;~,~!i~ e] :o;!ol1-; (If the decrease ill OI~li\'ity fOr i~~~l:,'{L"il!~ ~1[)~'iS \V"ih'l Mlmi~~f ~"'!l I ~h~~ :;:Iop~of the in~~.,,~:ICin ~I~lil\'.ily ~lr irtcreasil'i!5 Wlin.o;.in :JI,rlllljority ofpmti~ip[u[!b; t:-oiraatUlIK los~ >ga~1'l for 14 Olll ~ r Hi r"lr!ic ipa,nls. P = O.OU4: VMVF'C: Im.s > g.;,lIill for 13 Dlill of If, rgrli,C'I~lI1tS., P = (lo-1:~ ~. ]l:IoJ\f(;"II' IO!llIOn; dil'(':(;'IJy
3iS:I~'ti~

Ih'l~rel!aluO:r'blhibP \lo'1.."CmJ bet

11L:'Llru~.~.(l.l;t-; !IVC(:·

Plil;\'~Q(!i.." t>!lI!dies have ShO\'\I11 lIW!t .<lnt]ci.~l!;ld or ~.x~rif;n4;QO ~Q;"'M,.~ gij \'1;': ri'i.i; to ,;!C,iij\f~~i9~in ~iQ'Il:;~h .... h;,I1.\'1!,; blX!l~1~'1oll0C!l!'~~,wv'h 1~~1~'i\·~ ~1~lotim'l~.:>~cin !1!;~il.1C'mny~tbfuJ. a!1i.1",1l,lf ;n~~II'!l or n. J 7, IN~" In (',Qll~[fi~i,ihC' p.n;a~lll :.imly dt~11;· (l!,...:trnt"~ Ihal, in 'th~ c\lpg~e1o;l f,olJcisinn-!'Il,tk;il'lg. o pq~"n~i~.I IO~jj~ ,nv [iL.lI'l\.~n~cd by dj.;~!Ti.'a ..iL1g ;.'I1;;'ruviny ip:i ~gnO:L1S Hml ;)I,O~!~l ~~ otidc for sabj~ri¥j." \I~~II[JC' rnLh~i'Ihmnby iniell\.~~ili!5 3~i.v,i:Ly in
I

d1IJ~j:Ciirelhel\\'!(:~n,l)re~iU and previcru.':i ii.."Sull$ It'in~o:rcct> the impmtanc,e di:;lj~lg~h.hilil& ru'~OI1g: c}\~~ricncoo. .,unici~med, and doci~t{)n. lIIiiHl)' in ","'COllOm ic th~\"ric:;(iF choice 0 5). It is: biliil!crnIi \'Cltud ,..~ri<mm~ H:~g. j'. bilmCllni Imer.lII 4 IXIs~ib Ic~h ..t amygdala ~nsa~'llIIC'JH foU' ex~ a ~~d ~W!IlI:Jr.iUJfPFC pI'IJ"!1"'!11P'I~!:!lc!1itm)'mCl~Oir ~llCricnct:ldJ 1()t1~runiOC!.' !lcg;~lh e I~rodlic~im~ I'!ll, II I]; 2·l)t r.,1fhCf 'Uh~ll ~~g.;'!,r~~\';dWI~,~\'htuG", Ihe ,UC;I.)\ and riglll in!l:llor pa!~c:~1IC'(l!'WX ,!1g,:,. S6 i[HI.!Ol!lll,otll'lll ... :~n'[hi; PU("li('11't \\'l~tdy \$i: (lI]J S7 ,uid 1,1b:~C2). Tile£~ ~~ull"" ik'ntl;:lf"~I1~~~(~ IHiCkof In~ln'l!.;-d S wh(~n:!h~ n~UI1lIIiL'Spallf;!,;: w g'l.i~I~M ~m;~'iS, W,;l" cmrel.!II.cd wi.1hbcimviom~ less OI:VoCl~~OH. Unc~~ [Il>il~ dirrerencc.'l illllx!ha\'ku' WC1l'C ~JU'(I~~ly !~H'.\" 'Iwhich wa.\1 dCll-igmx! 10 iSlf}~me ,ooc:i:.;:ion l!li~i,ty) 'I~K'Ctlld~", th~ CU~lp\lI;:nioll of Il~'(;dicl~on CiITO~. r.d ~~CC!~~~bi, :gn:"ali!o!f b<c'I~<l\'~oml10.."''' "'!~rI1i{m \~'a" d ~(:i!(;d ~l}' di rtc!iCun~,:\~!1 UICI!JI 1·~"'flOn.~. a~~in~~d wiiiJJ! !!!,liI;01lCtlll,\lI,lr;d~'l.,;ili\'~!}' nOI 'mly The ~lt,)iuftlllm."i""of d~ei~ioll ~nd(;r I'isk \~,';~'l UIC' ~~"'t'ru M",dy n;;pli~"lt~ !iI~~ C(lI.~n:mm iTI\"L;!oilig~u[l.!duu :I. !l:,'lC'~~1fI1 ~1.i!lI.d}' y DeM;lr1tiino b ~o IClI\S~ bUll31SID III .g~Linl\~Fo;ri!ICl.t;~Ih..ing .glli,ns, b~h:l\ ior.d P"lWtt! of rid; ~I.\'~!Nlt}]lI'm lirl~~d we observed II ~.i.!!lI]I~.fiem,m co[ruhu~o.tl ..~'Ilh g;lITllbll'S Illl:'lt !1)~r !1 50/5{} chance lOr g;lining or 1!1 ~1. P'j ~,.who fOlUfilc;l LilffLla.mygda 1:1 ~CI IVuly roml ared 'IV i~cl ehoiecs 0 rr.isky Rlf]n:nbl:es ftiU'i'il~d. Io.~illg m.fm~y and shows 111m. UJ'i is pill!l:JiI] of b-c:lIH\'ior"J.l1 loss a:vc:rs~ol" in I.he M:J'iJ.orimm:at bchilvkJ.r L~ diirectly ~icdto~hc; bJ'aJUI'S gucalcr Cl1:I:iLex and M1p!.."Ui or rrol1 ml cortex (Ilg. S4). 0.111 I'L" losses and SlJ:l)C Oc'llDOOI11lCS rrmn.. xlas ~:ij1.". :>C11lSil i\'illy lO potcntno II lasses mlmn gains, IhC'!>C' I.I{)!,vcvcu:, nll~n::Hcctionl in ii~kmlilllldl.a\i when !]~C other hllnd, a.s JlOtcn~i8 ~~QsSC'SIIncrcasoo.~m i exicnsive 1C,t ofm'l.:as shQ\~"'-"'<I more rnpi:dily a results p'rodde ev ldence in rilvof or one of UrIC IllIJIdemtc-11flOl_mbllilygambles are rmu~] a.s losses versus !,!.:!iin.>;: h~ .becI1 alilribrult~[Jill pmspecl. d~ool}' f1tm.dIlHl£lul (~li 11.<;,f prospCCil 'L~~C(!Il)jU. 1), 1 1[1 dC(:~ca. illng response to n~mmlj ng roSSC$<I!mllOl18 s namely Ih:n thcfunl::li[)n~h;n n~I's !Hone;), to ind~voc.hmls \'>'ho \VCI'C I1lOfC ~o."i.,"'O!!vmc S 5 )1, prilTl<lri~yro t~u::n:clkctocm i~! ct!U'\.'murcomlC value ii:i. Ib, ftU1cljm~ fur 1D!'i~1!\.'\'1J!1IllQS ....2) (IIU] SOOOl1d"f~ g;'~nl ( Nnt:~bly. ~lCl1e.regions '~nwr~.ll1~'%Cd nmny of !'ILI~ljl,.'JCti\~C~r<l.IL!~ n!J[!!k(''A:lJ.y !i.tot..'tlIK';: I~";I;'~. ~I~c~~:ca.,> tlun shewed all, overall decrease in ily to (jj,stolitiOl~ill probab~'lilly \I!!'CiSI~IM~ .ml.hc.t" than g..'lin.~ [see a:llil.o (") 1. MOl\.'O,,'o:::r. ItlookuiOll

studies s~~owii1~ increased n,na decreased ~C~ ~iv.~y ill dl~ ;:-.lrial.lIIlJlfor experienced m(iIl:e~my g&ins ,md losses, ~e."~'Cli\'cll' lIt IJt '~Vc nc:.<!,invcst~mcd whelher :indh'id'Lml dil: IcwncCl1, hn bmin <tc!i\"~I,y dWlllllg docision-m1!kin,g were P::~'!l:(.'Jd u;_D~ndi\iidual dulYerenOt.'l' inlJclm.vim, lI~;;i'l11l \lI.ihn~!J-br.tin ~umb'~~ ~o idctu~ty [,L.'lgaor!:>

pcrrOJ'~,'i.'tI .. wholl.!--hrrni b! fi}L'i'Lh I. n;;!(in.~km IlfilfI!).'$ h w i.Ih these n1Ca.»IlIb"CS (21). This atm~)!sb re ...~~lcrI si2n~lk<lnl. eorrclstlens between bcil!l'l,'iornI, and ncumllos~ av,clfSklll h~ SCi. er.d 1:c.!:!:iOJl~. hlciudin::: sio~!
II

~[ilId ibcha,viO<GH ~O~:>!I",~rsion,wc ~

l'Cfjimit-l

'1,,,,~ocl.m\f..'tI

wil.h irn~g!tilJ\.o.: eJ:l'tQ:!it)fl~,nlb~

or

I~

mg.

nlcLJ:I~~1 ~c:tIVity \\' ith incrc.1S VJ~.Q Ilmcnt ~tliloss, The m..sociaa~on oJ o.£Cl'Cascd bdm\,li(mJ!~ loss
;)IVCF-'iol1 \JJ,,'~~h dtclt:O'I:«:d n~W1['01In.>s.,pOI]SK'S ~n b [JJh!o!\.'i~>:ll ..al1dg,~ili!~Lltiil]~ dl;jCi~im~-lliItik d i!l~ is C'o.n:si_ ...~ll[W~lh [~u::l{mg-smndll'lg nmUIilI] Illm t ~OIi:lC I.olill'ii:" ()rrisk~uklng~nffLy hll\'e IJ~.(':il' roo:~s.

amllysis (1 i I i'lJ,gSCsts tlilJ~ indi\ idl!~1d~llcrollce$ in heh,I\'rot,)1 loss il\'Crs~O:n {<I$ il1lfctu;-C(!b}! will"

!lhall 10 pml~

loss n versien,

]1] (m~IfTh.'\'l,\\'C asked

Itlll~i:~-

ill lite [l1\,""''C'J11 SIi1.u.I),lO c'vaTl!mc b.1lilloccd

in scl1~.I:ioJ1 s~d!;:illg by illdivilhml)i \\'ho~la:vc fI dii,nhli:>ll~d 'Physio:llJgkii~ rcSpOiilS1::' to s.~inmla~ lion (,11,. Exam~m:lIion or Itgocms oJ:' UlltOl:CSI. h~lhc ;«lruMlJ!I11 ~md VMPFC ~i'(lmlhc gaTTI/loss 0011jLlfl.Gl.ioll ollalys,w, O~l,g., ,3) re:v~l!oo 111m ahc.<;C'
F'iig. ~.. (lll'WjlJl!'I~~Qm! il[l~lY~i$ ~e-

,(50150 ~ lll<linillos.<; .gtIll1hles,\'\.'hid~,anowl,,:-d y~m ingnC:oill. 10 m;lC'Cp~ ~lli1>H~d g,lmbK"Sl~m drllwn ~ rriIHI1rril)'by illd uvidu:aJ dirl;"n;nt~l;j .in !1,;;ur1!1181;01l. ijj~lllah:' ~]lL;roll;;: nnOSc'i 11~rsion. nlll~,I.IDhough sill\'lly 10, rok~mial !(t~~!l!_ Altlt()~lgh Ihe pR.":;cm amygdm.!l!, <JlCtU\"Iniol:! mill)' p.1I'i)' II rok ~Hlsome :o<tudy fOCi[L~':S 0.11 I~si'im'Clllii,Q'Jl III [he COI.'1lc;d.of j]eei"iaJ'lS ~1~td.f1in:;ik., il. dfrc:s llol.np'p.;.\.'tJ:" 001 be il mX.'fiVLt) OOJ11J)Qu!.':m:111 10.,,$ flv~rstonl. mixed gflmblcs, ICOcll!. \\'Ork~13:' iOCllllndlJiiu the

cocfBidcl1ot ono~ avcrsl.oJl (i.e., d'IC n'Llio ofscn· si~ivily 10 Ilosses versus g.."IitlS)is, Iligihliy oom:lmcd across risky i!i~[jrlsldess,~Oll~e."'IS(.2'3). ·nH::I'C~'t):t1C. \\I,e :;:llIln~isc' rll1l;~ a ::..imiJar l'Iilech<J.nis~:n l'Iila)' COJ1~~tllile 10 c}ll~e~mm~i~~'lll[io'nlS 01":1:055 av~rs!OI1,

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d~~es lhigh'lig11~dustm th~ ~P«~~ hOOt!m~5 .~ th~ right. l. I~ R, ri~lhI:.~B)IHi!atrnap5.IM'~ ,created by- av,e~a'9inlgl parame~~,r '8'Slllm:U~ vmU5 ~t.@~Fl! ~'I!ithin @aiCflt
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dmtt!rim l~!! [D~IJIlCtiD n map' fm of th~ :116;OOU;s(~r 16 g,{ilmblflS ,~a{lrd In ~he gaflilliboss m1ll~)I11x;[(11.0r ,mlDllgl r~netts stilenglB:i af nell!ral ~e~Jl'!lns..e ~i1l.ch llon.di~Q!iiI, s;u!;;b for that da rik rred rep.reselll~ the :stmnI~e~t,utiva.liol'l allitl Wlrk blUe' ~ep~e-

~at loss iI'I,!\~jon 'b~hav:l(liI'al klS5~rs'!O~ ll1 v~nlr:aL SlriatlllmJ [Mcmlreat Nemokisjc:a~ ~1i1S:tiltit.e ,00000rdhU!s (y, y; .d! 3,6, 6.3, 3.9: (El1ter ~~'glFavilY ifl millirm~tBrst Regms'ion "!lie .anti P '~l'u;~ wer~ (OR'lJlultedl ~\ii~h fl'le I!l~' oJ rnhtli5t Jegre:~ n by ireralw@liy re.'I@~[Jhred h~cm SQ~F@~ 00 p'rw,~t the in'OuerJCie,~[ Qll!Ilie[S; Iw'l\l~r, timis, regre51lion elsa li€1ll1lainecl h1:ghly ~i'gnifi{(ml ~ fi.OOO) \lihen t~
e;l;trelililill

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REPORTS
The pJ;T;sc!lt~1!lts illll!$Il<!i~~ how :fIl1o}l!mimO1:g,i~g I;;I!~11!G' used to dqf\,'JC"l~3i~~$~. r~Clicl~o!'!,-,; '''I.G1!mnil]~rfo!~"I ~J~h"'\iio~! ~11;'(nri¥'ll.: il~ .h~!l. '1-W. ~
Rf!'rli!n!llIce~ ;[Ina! INob!~ lL A. l\1er~~. O. ~atm~l1!!iIn, '.lIiI"k !..ImM; :5, 297 ~1991!). Ol. I!}. :~'hn~~m"i\, Iyt.r~k.y,. ~erli{{l 01:7,Z6J,

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22. I"o!i. Zi;tc!l:ffilll!'lm. () ... M. Kuh'Im~~\ 1. ~n;, 69. '999 (20001. 2l. Eo Jo!im~"" S. Gi.nli!l.er:"A. ~~!!J'!I"~".·i~P'J;{ir!g ~~ ,~.. IIiI~ of Ikk~: ilVtn;lan" Unrtiliwi@ flu ihI~ SlliIily 61 ~bor (1m D\1clll~5iDnP.aper filO. 20],5, :Soo~1 Sdence' !':_es~r,dJl

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5. M. AJj}de-,'I_!aO!!j,. H_B~~IO!!n" c, !!'<!r.aidiriY, M'ana!l~~m·

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m'i~~ltilk1jLl

25. 8. D~ M~rli~D" 11). Kurn1lf~ 1'1., Il. !~fJ'jmDLir" R 1. Dolin!, S{"""~ lU" 6t1<il(~&:). 26. E. (o~gd'<H'I, T. (.a m~,8fhc:w. ('~g'!l:jtNrufD>li.. Rflo(. 4".

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~b W'. W·.IH!Mlili!U!lh. K. Iifo1lJ1! •• L Vl~~lumc, ttM. lM'. "n, 175 (2011)[1. 1!{I. M. I(_ ~n, \1'. l~k!oli1ii'1tin.;ii'~.3~;!;n" L It ~nltos, j',.1'b'lt. ErM. 114. 51{ ~200£il. 11.11. IHo. C. llriN~~; I . .AA~!QfI, I). :t::~~fli~r!~ilf1!. ~'~. A.
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Ill. KIn Il!~CI\ C. III, .Adamj;" G. W. iFDI!1Q. e, fklmrneI, J. ~.egu:mi.;!!::11, IICl!5~(lQOD, M. It DI!Igiid\n.\ H. M_ LlXh. '11'. ,A,. S'bellger, l- A. F·i8l, C'C/9lfL it1f~Ci. 8tltDv" trJ~mm. :~, :2 7 a,om). lB. ~ L!!t1an,s, 1J'Ii. fOIli~, CM. Ad~!m. ]. IL lti!.!f!eiI, Il'llilifl1r1m .• I'IWlON!f!lJf-l l,1t 3683 ~200lJ. II). KiL'llmllma~1 P. ,. Wjj~Jief, R. SaM, Q,1. EcM. ::1.12, 3:75 n,9'\l1l,L (, f. Camm~,. J. IIo:fQrt«. .Rts.~,Z; U\l 1200s.~, II. KahlLll6 DJ.,N,wfM 3i3~ 98:3: 12002}. C. M. Kv:1)nw, Ill. ~fl!Il!I!l!!n, Nr;:{lIIOll ,4,7, ',\'63 ~2:005.1. Iii. Kill jjll!OI'l, G!. W. f',om~,. S.. ill. l6eillIllIMJt, C. M,.A!'i\!!JI1i'I!, 10. ioIc!mm~r" NWiQJ'mrJ!)11 :t:e,2U '(20031.

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Ilmalml~}' divide ,is)''lH!lllClric,uU y. pmr;b;ill:!;l nn~ f.;~!,."nltQR which fl;'un'Linl.. O1U,lchl;O to rh!..lhub. s :md one !;~{'HJ'itl~hl~lSt. hich inilimes dinbp.'Il~ia\,.. tion. '[his SIX'I\,:'(nWpo;I" <ISYL11JHlt;lil.1t mU(;mll!;;, is;
C:OIllIJ[)Jk;d, by Ili!l~

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by an orfe~ood :spfndt~ end local. :s~U-ue!il~~val.sig,n~l:; from d"l@ :;t~ml ()ell

micm!u~f.m ur

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dhrbio,r'!i5. cam! bes;pedl'ied

Adl!l~t stem celb ,Clfte'll dMd,e .CI:symm'e1i:ric.<IUy tID prodl!l.u~ one seLF~renelWedl:stem ceWl iI:n~ Olil'~ c.et~ ti1I!1's, 1fIlI:"rnMil1!ing bll!th pOIPl!!1aliQn~. The a~ymmetric ,Ol!I~((lme ,of ~~1IIlI (Ell

to. Ihe hu~ NU 1~l2LJt nne dnl!lglnl.'1j' (\t~ I inhclt.i.b. Ihe flI'mclnnem 1:0 [he hWLh.whe:refll.. s the otheris displaced !lway lH Tbe stereotyped oricl1Ilmio,tQ of the mitotic Sililldl,c fls set lip l)ythe positinnhlg ccntm:~Kll]K:S dt:lrilJg in~e~h<JSC ,wrg.~A) {(.iI, GSC's

or

I'CIIni!li~l

micl1lcd W""a1:d~hcnkilc til rol!glww

dlC

IIIh::he.. H!IlUl' we :~mlJi~'i'that die'llel[Qpllll~llt,[lUy pnlgr<llililmed' a~vmll1e1triir;; belhi:!nliofllli1d ~lnll~riil:arH;e ef cell cycle. ~n G. phil~e. Ihe :sil1:J;;k0(;111rnS(llllC ~tl IIIlcthew and! di.!u'\! hler ~entH:IS(ilFlUilSum:fe-rlies t fu~s:t~reotyped ~piindlle orie-nliltio 1m .1md ~svmrJN~t:riC: Ipc~~too uenr ·~tI~ ~nood"oo with the hub.. \1\f1l10'1I Ihe 'll!I!IU:iIl'irljl~of S't~rl'iI (ell. divis-ions ilfl~he [)f(}s,r;p.hlJ(J m'~~ 't;i'e'.fml i,il1l~. h~ mOl~her ~etn~'l;loome r'ElITl.lhll::l dl!J)IiC<!il~t1 centrooomcs T f>'CI);]I,UC in G:;i! phll:l.C. ,onc ~nl(hOlf~d f!~"r th~fl~,h~' 1i'Ii:~it~tih~ daughter (~Ii'!~OS(l!i!i!~ migfatt:S t.o,th!~ QPplJiS:i~i!' s-ide ~~! th-e sm)'s ~ei\;1to Ihe hub, ~'\,'lli':l'l:!nsthe o~l1,i;!rmigl'l:BJtr;:s In ~hc oppnositc :l.idc IJf the cl;:1l. Cl;,':n~rl1\').onu;:~ ,celilbd'o!l'!i:spf!'ldt~ro'rmarriQI'!· •. in. till,': (iSle/! _'il!.;'p~ur;.I~'ull!il~~lall)learly ill i'l~",r:;W.illil ~d~~ vt; !I powm~~to h.. ,~1il\'r't!~ ":\}~Lmn.l"-~~· phillM;, r.lllhcr ~:I<LrD oil []!!,,: GrProri!,l'i'li; tr,m~;Llitn~. d~"'1Nk;m .cctbi Il~,~ima~rl ImJl~ln.'duul~~ 0 r h~ghI)' d Ii"!:{;rcnl.~ mc:d b:u~ i'ihO:rl·JI~\ ,~od ~:k'Llly,.pwducingOJlc C~iL.mmlnIO ~'(~ G$C:;; W.~l!l rLJllly!01;;-p.flL11k\d i:;ll:.in] ;:~nI(1 ont: diJ!fCiL\:U· so it i~ .0(:.11:;, I hrough~)otItll:Iii .. Ii Ii:; of [he C!rgll.~ ; CCrm\1ii:}IlIiK'l> willtOLli. i:L :>pine! Ie ,t .:;.500 Ci SCs ill. tiill~l1g ocll t TI1(, 3:>ymnnctric Ol!IOOJltt' of ilislflll. To Il'IfllinU1iul lh~ crilica:l, ~alill1ltt bclw~cnl lhc o· ~o Jrdily..o:ld i:l1rllcs COLlifll.cd, -40o/r, of Ihe s:tcfI1Iccll d~,,,is,kJ;.DIS caD~ be :;poei iucd by reglula,t\::d (}SCs hilt! h,,'Q CCrDlro$Ojl~ thill, \VC~ StCpmml.cd slCln cell aile! eli n:1..'liC~~lilatil'lg ,e~I, POlJtI~.mions.. spin~d~c oricIDDmioi'i, StiCh t~>IIIthe two d'ol~hlcr !o o('lpo'silc sidc.~ II~eeel~. DlLt no Sjlilld I~'WiflS ccll~m'C I~r<lOOO .in di fl"erent mk.rocJw iromllli:nts tlltu ,,;iLher s,pc!Cify stem, eell i(k~lli~ I i>lelii~ celli iy )lei <I..'l:ScnlMcdj Ii). lDelpar:lilDE!n! of De'!le[~;pID!!m~1 Bicr>OgY, 5!,mrord Un'~iI'ni~hc) Oiifll~,~~' di!1'"Cticm:ilflliiofl ql. I)jlfC'tI;!lCCS bctWI;.'\C'll~hc'nn,tll~'l:" ;;Ind dmJ,ghtcr 5ity5d1100t of lIliE!dione-, ~'tanford G!\ 943(11)-5129, USA. centrosome:>:: l.mdcrli~ the s!lc«.'oiwcclb.::ha\'io:r. .of'o.w_Jf/Mtl ~nmlc gel"ll111~ne:f>1.cm cclls (GSCs) ~DiliPafil!!mmt ,or MdeC'li!L~1' Gene,li,cs and (ell B[degy, aremnain~;)jintCd l.h1.'OLIgh OlItlacilmcltLto smmlJic of~he ccnll'OOO:rl'I'CS ill Dl7.'I."fJ{Jlrirl'lmllk: GS('!f,. i,l1~n~rs;\y 01 (hit~gQl. (hIC~9f!, Il ~~6a7, li SA ~{k,:~~I7Ii' me~l~o~ 'G~ne.tics"Stan!ora 11lIni!j\f,r~;ijSthoo1 of I'!.!!~dicine, h!!llb ~lIs. whidl 'ro:J1Slal\!W the stem Cte'll lI1i;;;:hc'. Ute moll1.Cf cc:nlmSOJl1~nOml<lll)' l'el'll>!i ns an:S.~amlfolild. GIl 9430:5<-5,)2'9, USA. dl~'CdIJOo the hul1--GSC ~!1tocl:litoc and ~s illl'W!'ilod Ill!!b cdb sern.tc dlC' sjgIllilHJ:lg ligand l-'l~. \~'Mch ·Pr,e~eP1t .1ld'dlieS£; Life S6etf1ces!rnlivl.I!e, ROOflIl, 54031 Imi·, m::.h .. !A.~ ~11'"lianus kinil&(.~gTml~r.ul.'\du~ 1.O~ 'the' (is " wJ~!t'a" .Iie .oml,ghtcr o~nlm8m]1~ <!lid ~-er~l\y gl M1~h~m. ,!\ii1fi ~bQ:r, 2:l!iOW~~hl~il1!1',i A'I'I1ilI!t~, ;u;'!U\'<lID!' ~ .. Irm)!o1cri:p •.ion lJAK~STAn pnl!h· Jl.1O\~ ,~way I"pm lhiCllu'b a~!cl i~inh~ri'h,:'d h~' Aim Mor, MI 48:1!..83-2.216, U'lA. ~hl~ ~II '~in~11 cmtmti:i'l~ 10 d:~~rcrr~l]a~ion" ol.hi:t < M way in~;hc neighboring ~Ii,;';ffi!n OcU:> In ~Io~i~)' fTo IWmom ()o~res)il'Qnde!"ce' sholiJ~dbe add~et ,['·mail: ali.d daughter C~rrnl.rut;o.llil';S,\"'~~R:' d:ill]b.~tunallj' yw lIi~cll1ll¥@Urnlidl~,~dl!h r'l'.MJ(.~ rulLilr®@!n~crd..oow i(M..T.r..) S:[~nil.ecll i:JcllI[ily Q4. 51, .IJ1'rJ.wphila Itl\~.tc GSC~

or

J,..

or

518

REPORTS
o r green nl~m1l;:il~ pr:ricemrin/AK/W4S0 C'-9wrmi!nI!tO {GfP-PA('T) 1~L'}m f).1n"111pifil(.J p~TiQ1;,I1!ri~.· !.!W labc·ll:.d by
111"11~iCI1tC]"p;r;-.cs$~ml ~"cI1!i

second eel I,,cycle" GC'~er.dly" Ihe oonrooomc

to

pro~~j'l.

l!wIH~b

\'\,r,,~, ~§lh~~.

,d~~1 "!j,,,,lh~~1:'I~;h~ ~mfC!S(lme i

rroJ.;i!IllJ~. 10 ~I~~ Il~~b w;t."I 'GFr"ll1C.'g:!illiv~ >9$% (.Df { lilk ~]n;lt~~ill. ~nd!".'[1W-~Ui:'iIIl.1l~k(jJJ14 eomrol ,t?) GS['j; iu t~lI; '~Ylm~l d~'~)ld~) (H"iM:' L D ml(1,IE), ~ ~ iltdiernillg Ih:d LIte d11ughkr t"!L;rI!Rkimn~ l.nigr:Jllc {;'\II"Th'G PACT dO:l1Nlin. which i~ n¥'t~Gl;;lim)' mtd ~dnd"'lla ~h, ,oollnpL.1r ID>C3li(!twiC1H. b. ~nopt.. <l\,V.j,II}' rrom the hl~h.!CiSC' i.!l~C'rthoo ,('hlring ,~,y~l' . p [lm~od ,illro colltnok':S on] }'d.l.lf,iJlg C'()!1UWNQI'IlL;' metric {iSC di\'i!'iioras. (li;apl.i¢!iliioI1

~'mindlby the m'lC~. ,"cnDht'01lg~l InlJhillie c ro"nCl~ or GS(, divisions, SQITlt" 'C;S .s m;)j~11i)<lin(;)d ~ytol~I.. ~qIlik (iFr-rAC'T, .~_"p!o,~itJUy !!!1 U
Im'v:l1;" ~~I~I.~ng

y~~.Ib{'(~n di!ul.L'd cul

\1I.re
I;;CglIi!'OS.ol1t~,'l,

d!lI!~h~ GfP-PAfl' hmd ~n]~ {fig, 2Cgfu"l:'Il collllu,1I'I~

OlIJSQ O~]ili;,':f'\~l;'d &l!l1C

GS{.":-;. ",;,ill~ hvo ~ bek;cl ..

!IItd dpc,~~ol

iI,:xch~t:ig~ wilh

~hc

l;Jhding.

t~IC l1)o~h¢r 1'<~1~'kT ,.m the dl;r~lgliil";T U

ulrs~ cy-cl1; ad~

~lI,£g';'~lillg 1&i.;:Lllh~}' mtl i!1 the ,~~krahe d~pk[itm orcyl(;ipJ~I~lltic

rotUDli'ili5).

t'y[opllf15rii1it: poo~ ~1'). 13011'1 the itlotii':l" :ilK! dau~lw.'!r tx'.i"Il.MS!tltllCS ;In:: J'I.lbe~.!.XIby tiFP· r)J\Ci hllhcl~ USil cell cyclle IitlCi' hCf11, sltoc.k {f:i~, IBJ.~n tllC' second ecll C~CllC.b.hc daugJucr cClllroiilCillinC rcteins GFP-I,)~\eT. wi~e:I.'Ca3 the ~l101hC'll'c:cn~roso:me ,us11.01lii'Ltl!C'lcd.1ht!~ disIJI,g.uillllhl,g lire nlO1.hcr >lnd d<llllghicr (''"Cll~J)Olimn~ Wiy, 1B). A short ~W! ~~ 'G r '"·PACT I;lx:prei1~ .~~m ind!oc;ed~' <I ;l,S-hour hlJOlJt ~~h("l!(':k., ~1 20 J(]IH,ofthe(iSC~ h,L\H GFP-I~I("tI ,~nl:rm.om¢S..

cCIl,lrol'it.utiJL~,qmfiru~koJ liI'ilHI'Ilie i'liult GSC~

in Hie

GF.P-I)A;lT (Fig. 2C. n:d

i'I[d~~n:rCi"i:i1lill:UlYatlmjil'i motltcl'

('(1llro50Jtt~';'I

o'VC:, linllc. Cenl'Driolcs IIsl~::mbloo th.rii"iilg calfly ~mblyo~e:I'IC'Sis were lilbclcd MS,~I~ the NIT[ 40 0:114 dro,L\er Ii) dL'ivclite
C:!lf)UCSSt('l~l

of'G FP-P/,\C1

ncr ,I

or

in bla....~odcmH1.a£lC' cmb:l')'o.~. sluJttil'l~ off' <iller gCnt1b.1n<i C~~~li\[cn {91.ln~hc firsl cellcycle

The [IHmhet CellitrosOI'i1,~~I~ GSC~, Ii~~'· ~Ilrcd to ~'1\l,~iiUjjii1 ro'blii:.l. ilrl~qilm~ micro!u b~iJ!C ~rrn:'y:;:. Ullrn;iOLnIC~iilrn.l1 ana I~s~~or Ihc .ases uc'V\,"a~ecl t~lmlite, ccnlJnsml'lt' pro:!'!hml '!o lite huh W,:LO;: co:rmllo:nly :ti1$.OCWttcri wilh ~mul)'

aft~r~h!,;' d~~p~I.'l'!.io~ t~l!.')C}'linll"~lnic 1'!{l!O~ (If GFr-PACT i~ 1110'GSC5. 11O!h !he mo~I!l.'r ,mti d"LU~:IICt (:~IUml';Qimll,.-;_'; shnuld b~ h,bc!l,)dI, 111
Ml~stlq]ucnt

or

~nLCI'OlllbWlIb;IJ]ml!gholl!l t~nco~1 cyclic,.

OCnlfl:J!'iOm~

~r;lipm;

Dr

in ases were !he OlU)ic;d ti~

N~LlCll.II;,'11

:-;tllrl.'d

in ~r!<ll and

(If

[(';S,I,L,::;_ E111L,.,,\,l,l ~\;,l1aI'l'H>OD1k';,>; \\'q'c;

I1v~ \cv.ild·~j'pe ~,rlit'.:;di,,~ dOl'ic

Inl~

iHdic.aling the cll!.p~k:m;ti{!11 ccnl!l'O$OJn~~ d~ll," of ing, [he \\'i~!d¢iWof G FIP'··P:ACT C;~I~!rl;.':.'isi{m. B)i 12 hOllin. o:anCf lw<n ~hod .. >9{}% or'1I1I; hdJdLXi
('iSC';\1

ceM cyck~.$. ml~Y Ihe

nmlIhCl'

cemrneell

to the adlli:,'ltll1." juncli.r1-1J.'l 'lJChvce~ dlC

somCl:>:>hou~d ix~ labeled (:Fig.1At ~Il l1!lOM '(iSCH in [I].C Mi{:om;li

Ci'f ~,lt'Or

:ha(B

I.wo GI.:;P-R'I'llo5.il.i\'1.!C'('lnr"ll>mIi!C~,.

~!~.

clicrning that 1.11,(;:)' had JlIt),gn;::.;:;~,d, the fil to ph~S(!cf'ihe ~1n,I.cclI1cycle: after (iF'[l"·P/H..T iu.· C"Orpoml ion ~Fig. r, e ~iIld By .~;.:w 2~ imW'S .!'I.I1crl'LeiIl shoek, the number

~.Y~I", unef the depl!.>:tioJl c~f c},[OP,I;J51tiIic (!Fr,· P/\;[T, I.hel:lbdcd cenunsomc was po~i~iofll.:d
next to I.hl.! li'llb·GSC

~he (iSC's. Ni~~~ 1I~~~ plmirnmJ C(l!~trosoIV~C..\1 of ~lrp;:;a~d 10 be ill iln'l,!,;.ifJll~[;c~~I~\'"based on rlU(:]£1l[r .U1110I]iIhol!ugy ;:JmJ flliCWLl).llblllk' mnmg-l.'~~K'i,l TYpit"'Jmy. ihcsc

ilt't(~rphai>C

CCri'[rooolillll,:~S

orGsC's, "\I~th!WO GFP--posil:iI\'C O!I1!tro."«lm.1lS had dOCL'eJiSOO.whcrc.'1.'"thc~lllnlibcr Qif GSC's wi!~loae (iFP"pmili\!c anu one Gfl~-nC'g,a~iv,cO<'nlll'Ollmne ~!<1!d iJla\~a"'I,.>d. )il.1,gg~ting IUOg!1l;.~ion inw!he lFiig. 1.G'rP-Labeledi dilIJJ9ilterr r::entJ'o=

pro-.xh1b:d~oDhc hiiib were ru..ss(IICim~ with liU· interfaee, :Jnd~hc ~1D]lil· beled ccnl.msome had meved 3WClY fmm 11:IC ~ncro~s, i~,icrol!JIbllll'S tPiy. 3A and 11g. stA), :111, &mu: .sru~npl!l!5.,. l~k;ro!:LlbulcsSipperl~d to C'X[Cifld 1 hull (Fig. 2. B snd C). Tit!! I"rcqucncy o:I'GSCs, ~rl)m~hc C'C'~1~If)QSOJ1':(,l toward rhc adheren s thm :lim:! the proxnr.na), b~lnol. dL~IJ~,. centrosome labeled relVlai~wd co:nstaJlII over time for jenerions ,(F:ig. 3J\.', armwhcads). The o~hi:,'r

10 d;!Y!' (U

larvae

~o

day=3 adl!Jlt!'i.l. '~llI!gg~'l1l.~'diabl}'

!\1;,'cn ~'(l(l! VI11<1~ccrnmSOnll.'S

allpoc:lll1.>d 'to

ing !i~,H!h~ mother

oen!l'D,\"lO!]l\O'$ "00

~IIL>; ~n milO~ie prnphiikw UI,,!;l, S

Un h\\~d

be: in
011

semes m~~rat.eaway' frolml Ihe niir:he. {A} St.er,e<ol)'l]leti ]p(llSitioJl!ng 'Clf ~@I1lro~orn'e-J!,in m.aleGS( during inte.· phaS-e :sets. UI~ 'rhe Orteif1ta:~Oii'l ,(If tine !ill! it(lll'i!c sp,,]ll1dle r,g(fi<1ljlitedr.rom, {o}l. Ried., (entrosOflle; blue, ilIub; g.rBe'JI". ~l,iool~ii'i!, B)P~b~(il'la<s-e ~ stfal'i\i~' t~ lli:otl.ls ~ls IiIUlfftib~re!!l1 ln oHi@r or '~eliler<lJIJ1oru. TraJniSiellil: e~ress](m Q'f

A,

~ab~t '~h~ d"u:ght:eJ ,~!!~ItOiSO!11@. Ea(!i1 'O~!'IlrOOom:e ((irdl!') (on:ta'in$ l\l{(I (:elil" (j,Fp·PAn labels
fte~~y

a 3Sems [00

'o~ntri!lil>es (3). II'n tile 1iirslcell c;yde ,ilifit.l'f ~e GFp·PACT IPU~, bolfl!he
~o~h~r ('l+~)
!lIil(l

dallgM~r

(2...:;I}

'OMlf~~am!.~ o1i~ G~P"i:lIooitil;1~.h'lU1e ~rniildl ~ell c.yde, Ihe' lmolher (>e'lIlroS,()l!'ll!i!S

U,+4

(J~

2+4) ,,~ GFP=f1.eg~~~,

wh,w~~s~h@< dfllL!9nwr ('~l1t]f'OS(llm'~ (3+·4.): afl~ GF!'l·IP05iliVt~. ,(( aiiildl [I-)

Tesliis tips wirth ~(lbek:d'61510 (doUoo

m.rIUne} in the

.cyd.eaH-er the' GFp·P\!\Ci pulse ((0, lL2 ~'!:H.!rS ilr-tU n!-at shock] or Ute :51!lt/cm:d (,e;l~ ~yde ((0)0,. 24 Ihoo rs an9\r heat sh,ock]. Greeli1. GFP-PAU; red, 'Y.
tl!lbllillin ((e!lJtroo()rm~s ~~ :sIl.Q\\lfl 'Mt~ ar.rO'lli"heads.) alnrl 11'08",111111 lib,. litO; blue, (iii VQI~ ~g,~nm (;@Llsl" Sca:w ~~r, ]0 JAm. ~,E)fr-cq,l!enW (I,r ,lallJ.e'~ !!l uk:ooles O1l'er lime anN h!iNJ~ shod:. Blue, ~!sCsilil thefnrs:l
'CIl!l!~O~moe~):

(h

phase o~'lIiIe fir~t'Ciel~

~e:
time: p~,~;t1hC'·~t~Ii!~!a! ni!!i~j) !I;.eliltm50111U~ ~o '~he' hulJi; Ibla(k" Hie 'f>iIl{vnd !I:~11 next {y{I~el'.lIYitll the l'Il!llth~r (,~~UQ;~me dl,st~l t(), th~ hlJ!b; gf~n. th~ s~ond ~!.lU ~de. with neftl1~F (ellltro50rne nexllo tlhe hub . .on Ly th~e tiSCs that Ilad hllGl'(en lro:5orTle-:s, with on;~ Olr both Cf!:JtlrO:!1Qmes labeled l.'i'itliiGlfP\.PA.cr, ~\II~,re cli9f11l!:!d.

c!!l'l[ [yde (hwlalbe.l-ed\ mi,en~ed' ~~,GiSCsin th~ :s;~nd ,a~l~(yd~ (Ofl~ labewdl~lf!d Q~~ !lml.i!be~1!'d (entrQsvrlilid 'i'lii'~hthe ~aJoo~d (daughter) umtrosClime dif>ti!~ {roml [he< Ilnlb. R:ilIre~y oD'>er'IJed lCIutlIHiI"iU1!:5 ~\IIeirepin ~ th @ ~irs~.(l:!Il,cyde,wit~ n.:ei.lher

REPORTS

1)C:lYgilter lFilg. ,2:. M11Ioo GSCs ill the niri:he maililt~il1 ,ti~llt)(iQte:s as.s~iiJlibledlmiilll1l1' ~'l:,U Ig,erl!:1'l<iltiom:l<lrl.tElir. (A~ :5tr.ateg)!, to, la be'l the niioth:err (emitrm!;om:e: -,GFP'· l PAO expre!5$ed dlJlril'lm1 ear.ly ,~Il!rYQgene~~ 1JI$'ng~e NIG'I'40 driv,e-r lIfIinks the [l!tnlrioles (1 and 2:) retatnadin lli'ii!' molh'Br (]lmilro~ome& atflieor C th~ dlep,i@tnolil of (ytll,p!~,~mic mp·M(J.~Bne5.ti~ tip hOmla .ne~'I!ilyed,oS>e'd malte ~vi~.~ 'G,s(s oonrtllijllllng i(j1j;:P·i.a,belt><! lIIiIothercel1trQsomes Iprm:]rTI<ll M the huo·GSC ]ntE'~~ -; foll!1>8'", Rm, y·,tu blll]1TI (~elntrosomes a rre ~ho\'ln \lIith anmllme"ds) and F'il''> lilll (ill ab, 11i); ~ ,~re~n.. GIF~P'\!!!i(r; blue, Va;sa {germ ceU-s). S~a\e bar, 10 (C) I'reql!le!iil(Y i:l:f ~~be'l <llli ,(!!~U:Qm~s ©ver tim~'fr~'m NGIT40lUA:S~F:P"IPACil' flie;s, UtAS, up:Stn::~lm .i!t:tiv@t]n'g $iequence. IIHUll! Hi~llm,i'ilS;, GSC:s in lhe S!e~o:nd er later cell, cy,::1.e a'fi:er GFp··PAH ~ ,tI'i!<,p~iHiiD:n (~,f1~Y O'I'U! c@fltrOs,O!in@ \~S l.i!b€'~~dl mth GH). 1 p:rc~ima~. lalleled :;J 'cerltru5;orne liI:u:i!le,d prox.]lIIiIal ~O' the hu'b~5.(intt~rrill[l!l;1 diis;!:a~.~il!be!ed c@r1trO$orn@ 'i ~.o~ated distaJlrro:m the hub 4SSCifiterfa ~e;] IfIlii:'lClri, neillil,e:r of til,e hvo ell!IiItiro:'lClme~ ~QCM@dnear the h:lI!i:l-GSCiI'U@riaw. Rll?d,col.ll!Jirll'ls, GSC:sin 'Ih@ first i!:1l:M cyd!! (l!leUl .~ O,!f1tm51omes were llalbel.ed by GlFiJI).20ri, one' (mtmmme dOI;t:' tIl the hlllb·GS(c: 'ill! interra'C!E!'; II m~:'iimiji, neither CE'Dlltrmlorne' ,diose 'In the thuil.-GS( InterFace. 6reeliiil "Ollil mil, GS's thall. still. had qrlQP~"smi,c; GIFp·PACT.O 1m ~y SO tu 600f0o~ th,~ to't:al fIiIale J:iil: G,SCs \\!'ere la,be~ed by' NG,r4I)JGFI~·IP,#la. TilHl' r'e\malint~1190i '1.0 .~Q~tI could mnl:ailil u. 4 iII'i~tl'i,err lI:einbii!Dles;assemb~~d iln ~alrly 'embFYOi]riN,.e~is berlOIDe GI~P·P'Acr expfe·~i:Olii. Olflly tililiii5@ GSCs ·~h.i!t hd two (CrUrQS:illllfl@~, \Viltili O!'1e or beltih c@ntro,rom!s, 1(lllel@,d with GFp·IPAU, were (~~Iiill~,d. InlJiI1lbe~ of GS(s S(@flOO per t'm,~ pelnt, N.

.._.......,I' :

..- 1'#'
If-

wn.

11

~,

.,

.. k

I'

.1." -

i!

Filg.3'_ (U\lOCIsom~s, n~t~o lh~ ~ub barber robust fIiIh:rutUlbll!l[e ,8 rra:f5. W Eloonm mluogaph ,ilIncii Mi.') Slilmm~lry diagram ,~f a Pffl~ imsl (:l;!"mros:oo;~illl a GiSC. Armll-

Im!>OJ:llL~ nrlHy W]I~~S

illmlyli 'I~11ul. ffimhor~hem !n

:Itlm.~lTImil1, HflIl~rp,li!~I!li(; mkilt]~ultllllk: ihe ~1i!JbLG5(, imCftlfl£lr. .he daughter ecmrosomes mflr,Y :i:nilially

~ead5 in fA') snOl'l ,i;ll tmkrol.ub~e

lh~~fi!iln~ 'rr!!lmtoo

()I,i~lrO$OO'i~

.~

the .a.diile'll'lli:'> jllllil[tiOO. • B.edl"on, ~j;(~og rapiQ O1I!1!d(B') $l!l1'i!'il'll1a1)l dria,''~ralflll of ~ dista'l IOOl1iIJooome iii! .:! GiSC. Rled, hUlll; Illue', mkrQtulilllJ'le!J..; '~r~m a'dhi!',reniSjli!!l1(lioosl ytUow;
1IIl:!'!:let.l1i; pink. !lelltniD!le; grayint pla!;fIliI membrane. Sral'e ibar, 1 jJm.

un.

3sS()CifLtOO miCU{);4IlbH~eS am'!. be rl\'C~o C5~fl.b~jSihi~lg robust micro.ub!.l~e ,ITllli.Iy II only l<IIteri 11 '~heccH cycle. O:msblcn~ \\'ilh Ihe idea Ihm astral IliIkrollii bal:e.~m1chC!<r (he meihcr ecmreserses to the. hllllYGSC :~ntcrrfL{;.e, .Mmdle~- versus (lmiightercc.nll1C!-Sounepmi!kmi ng was rnndmnizcd ill GS(,s lil:u \'V"CfC hOlI10byg.tl\~S ntl1!3m rbi,' n.munStlmhl' (cm~j' (8).. all i!l!tc~,d ct!ntrooonmi rrol~i!l reGlLlli~l ill ,1!]chQl:,a."5~r.d micRH'lhW!I~N!() C(,~I1itm· ~fnr;;", (10 1.J'~1 An;)!.Iy:s;~, mnlhL;T ,lIld d,rughtoC'r
I;""W
1'n.1OVC.

IlfNC

or

Q,."rlTntl:l!:.mllC'J,; <lfl(~r ll".l!11SI.;,111 ~""'DlI'!.: •..;..~ion of

Grp-

PACT r('H.;"Ikd 'c,1I<tt. io~.··d~'m h~m~~JL)'goWl~' IlIiIu· If~Dll GSCs w~clC:r,_; Olil~ or~h~ [we! CCrl[nX..;{)I'U~e-5

,Vi)s, I.)\')silron,oo oc;(~ ~oDhc hub. it was C&~JUifLlly

rnndom whcth~rl ~lC Inol ~1C'1" f the dilLl,ghtC';, O cemrosome ::>~,)'C'd 1lC'.,>I, tolhc !mb (f~g. iI). In ~'liIdll.tiol1i. on >25% Or!Q,tf~ Irl~loo esc:;,. I'IdihL.'t'" of the two centresomes \\:a:> l'tc'.'d.iO th.;: hub,
~~;Iilkh is cou~s:islcH! \!i:hh our previous, ·ohsC'r'!.'.rrL·lions (,!Fig. 4) (6).

~h~ir ~:ob LISI mil;:!(>ot\!lm~(;l;;urm}'s COntlllli'lIIg i:l1lJ! adied mk'-ro~lIJIblll~ nlimillgp.:u;,;.i~~I~o or pic!'::in.!!l ~licmldc,ur~uirfrtJ;:c"
~I~ COinlmsl:,.

Ih~mppallt:'li'l~y inh::rrhu~C'

of !'he .rive ,dic'iI.mJctnJt~l'I1IC'c:o. .iu! cd I:. I.h!ul. ~~~ ~it'd+ W

Ou!' resalts indkaie Ilmllhc two ceilltrooomes in, .iJRlfi~pMlamale: GSC~ h3... d~I\I:;;,I'Cnldmf~ e Ibm !md assoeiaicd microl.ubulb (Fig. 3B and Ag. SIC). The ren!<l.~ni~g llulOl".'dist..~eemrnecters a:no lilll!~" Tile HillIlthot:F cemmsome slays M~[l1!;;'_""PllCll:rOO W 1~~ ce]1k;.in milnll(: ~lror'"""~¢. ~C1>;,l~~l~Bic jl,Ui,c'lipu. Wi!:11 !he ,!iJ;;h~ anI(.! .Us in· in ~!cri~"dby l.h~ cd~~i31 sel t:lt"ncw~ sb,;m cell b~d 0)1 micm~Ib;~k: :1ml¥~ cll!1t:J1.ining liU!ficlkd l"ilicl',o~lIbuICt; (rig, SID). 1l~!,J:"iJie"rrair»Jl'iC:t tt.ri· ~iJ.k:. Thus, GSClo: c~m 1111111 I~.:liIill old c~lill:rio.l~ !i'n~

r~w