Chapter6

ATouroftheCell
LectureOutline
Overview:TheImportanceofCells

Allorganismsaremadeofcells.
Manyorganismsaresinglecelled.
Eveninmulticellularorganisms,thecellisthebasicunitofstructureandfunction.

Thecellisthesimplestcollectionofmatterthatcanlive.

Allcellsarerelatedbytheirdescentfromearliercells.

A.HowWeStudyCells
1.Microscopesprovidewindowstotheworldofthecell.
Thediscoveryandearlystudyofcellsprogressedwiththeinventionofmicroscopesin
1590andtheirimprovementinthe17thcentury.
Inalightmicroscope(LM),visiblelightpassesthroughthespecimenandthenthrough
glasslenses.
Thelensesrefractlightsuchthattheimageismagnifiedintotheeyeorontoavideo
screen.

Microscopesvaryinmagnificationandresolvingpower.
Magnificationistheratioofanobjectsimagetoitsrealsize.
Resolvingpowerisameasureofimageclarity.
Itistheminimumdistancetwopointscanbeseparatedandstillbedistinguishedas
twoseparatepoints.
Resolutionislimitedbytheshortestwavelengthoftheradiationusedforimaging.

Theminimumresolutionofalightmicroscopeisabout200nanometers(nm),thesizeof
asmallbacterium.
Lightmicroscopescanmagnifyeffectivelytoabout1,000timesthesizeoftheactual
specimen.
Athighermagnifications,theimageblurs.
Techniquesdevelopedinthe20thcenturyhaveenhancedcontrastandenabledparticular
cellcomponentstobestainedorlabeledsotheystandout.
Whilealightmicroscopecanresolveindividualcells,itcannotresolvemuchofthe
internalanatomy,especiallytheorganelles.
Toresolvesmallerstructures,weuseanelectronmicroscope(EM),whichfocusesa
beamofelectronsthroughthespecimenorontoitssurface.
IG Lecture Outline 6-1

Becauseresolutionisinverselyrelatedtowavelengthused,electronmicroscopes(whose
electronbeamshaveshorterwavelengthsthanvisiblelight)havefinerresolution.
Theoretically,theresolutionofamodernEMcouldreach0.002nanometer(nm),butthe
practicallimitisclosertoabout2nm.

Transmissionelectronmicroscopes(TEMs)areusedmainlytostudytheinternal
ultrastructureofcells.
ATEMaimsanelectronbeamthroughathinsectionofthespecimen.
Theimageisfocusedandmagnifiedbyelectromagnets.
Toenhancecontrast,thethinsectionsarestainedwithatomsofheavymetals.

Scanningelectronmicroscopes(SEMs)areusefulforstudyingsurfacestructures.
Thesamplesurfaceiscoveredwithathinfilmofgold.
Thebeamexciteselectronsonthesurfaceofthesample.
Thesesecondaryelectronsarecollectedandfocusedonascreen.
Theresultisanimageofthetopographyofthespecimen.
TheSEMhasgreatdepthoffield,resultinginanimagethatseemsthreedimensional.

Electronmicroscopesrevealorganellesthatareimpossibletoresolvewiththelight
microscope.
However,electronmicroscopescanonlybeusedondeadcells.
Lightmicroscopesdonothaveashigharesolution,buttheycanbeusedtostudylive
cells.

Microscopesaremajortoolsincytology,thestudyofcellstructures.

Cytologycombinedwithbiochemistry,thestudyofmoleculesandchemicalprocessesin
metabolism,toproducemoderncellbiology.
2.Cellbiologistscanisolateorganellestostudytheirfunctions.
Thegoalofcellfractionationistoseparatethemajororganellesofthecellssotheir
individualfunctionscanbestudied.
Thisprocessisdrivenbyanultracentrifuge,amachinethatcanspinatupto130,000
revolutionsperminuteandapplyforcesofmorethan1milliontimesgravity(1,000,000g).

Fractionationbeginswithhomogenization,gentlydisruptingthecell.

Thehomogenateisspuninacentrifugetoseparateheavierpiecesintothepelletwhile
lighterparticlesremaininthesupernatant.
Astheprocessisrepeatedathigherspeedsandforlongerdurations,smallerandsmaller
organellescanbecollectedinsubsequentpellets.

Cellfractionationpreparesisolatesofspecificcellcomponents.

Thisenablesthefunctionsoftheseorganellestobedetermined,especiallybythe
reactionsorprocessescatalyzedbytheirproteins.
Forexample,onecellularfractionwasenrichedinenzymesthatfunctionincellular
respiration.
Electronmicroscopyrevealedthatthisfractionisrichinmitochondria.
Thisevidencehelpedcellbiologistsdeterminethatmitochondriaarethesiteofcellular
respiration.
Cytologyandbiochemistrycomplementeachotherincorrelatingcellularstructureand
function.
IG Lecture Outline 6-2

B.APanoramicViewoftheCell
1.Prokaryoticandeukaryoticcellsdifferinsizeandcomplexity.

Allcellsaresurroundedbyaplasmamembrane.

Thesemifluidsubstancewithinthemembraneisthecytosol,containingtheorganelles.

AllcellscontainchromosomesthathavegenesintheformofDNA.

Allcellsalsohaveribosomes,tinyorganellesthatmakeproteinsusingtheinstructions
containedingenes.
Amajordifferencebetweenprokaryoticandeukaryoticcellsisthelocationof
chromosomes.
Inaeukaryoticcell,chromosomesarecontainedinamembraneenclosedorganelle,the
nucleus.
Inaprokaryoticcell,theDNAisconcentratedinthenucleoidwithoutamembrane
separatingitfromtherestofthecell.
Ineukaryotecells,thechromosomesarecontainedwithinamembranousnuclear
envelope.

Theregionbetweenthenucleusandtheplasmamembraneisthecytoplasm.
Allthematerialwithintheplasmamembraneofaprokaryoticcelliscytoplasm.

Withinthecytoplasmofaeukaryoticcellareavarietyofmembraneboundorganellesof
specializedformandfunction.
Thesemembraneboundorganellesareabsentinprokaryotes.

Eukaryoticcellsaregenerallymuchbiggerthanprokaryoticcells.

Thelogisticsofcarryingoutmetabolismsetlimitsoncellsize.
Atthelowerlimit,thesmallestbacteria,mycoplasmas,arebetween0.1to1.0micron.
Mostbacteriaare110micronsindiameter.
Eukaryoticcellsaretypically10100micronsindiameter.

Metabolicrequirementsalsosetanupperlimittothesizeofasinglecell.

Asacellincreasesinsize,itsvolumeincreasesfasterthanitssurfacearea.
Smallerobjectshaveagreaterratioofsurfaceareatovolume.

Theplasmamembranefunctionsasaselectivebarrierthatallowsthepassageof
oxygen,nutrients,andwastesforthewholevolumeofthecell.

Thevolumeofcytoplasmdeterminestheneedforthisexchange.

Ratesofchemicalexchangeacrosstheplasmamembranemaybeinadequatetomaintain
acellwithaverylargecytoplasm.
Theneedforasurfacesufficientlylargetoaccommodatethevolumeexplainsthe
microscopicsizeofmostcells.
Largerorganismsdonotgenerallyhavelargercellsthansmallerorganismssimply
morecells.
Cellsthatexchangealotofmaterialwiththeirsurroundings,suchasintestinalcells,may
havelong,thinprojectionsfromthecellsurfacecalledmicrovilli.Microvilliincreasesurface
areawithoutsignificantlyincreasingcellvolume.
2.Internalmembranescompartmentalizethefunctionsofaeukaryoticcell.
IG Lecture Outline 6-3

 Aeukaryoticcellhasextensiveandelaborateinternalmembranes,whichpartitionthe
cellintocompartments.
Thesemembranesalsoparticipatedirectlyinmetabolism,asmanyenzymesarebuiltinto
membranes.
Thecompartmentscreatedbymembranesprovidedifferentlocalenvironmentsthat
facilitatespecificmetabolicfunctions,allowingseveralincompatibleprocessestogoon
simultaneouslyinacell.

Thegeneralstructureofabiologicalmembraneisadoublelayerofphospholipids.

Otherlipidsanddiverseproteinsareembeddedinthelipidbilayerorattachedtoits
surface.
Eachtypeofmembranehasauniquecombinationoflipidsandproteinsforitsspecific
functions.
Forexample,enzymesembeddedinthemembranesofmitochondriafunctionincellular
respiration.
C.TheNucleusandRibosomes
1.Thenucleuscontainsaeukaryoticcellsgeneticlibrary.

Thenucleuscontainsmostofthegenesinaeukaryoticcell.
Additionalgenesarelocatedinmitochondriaandchloroplasts.

Thenucleusaveragesabout5micronsindiameter.

Thenucleusisseparatedfromthecytoplasmbyadoublemembranecalledthenuclear
envelope.
Thetwomembranesofthenuclearenvelopeareseparatedby2040nm.
Theenvelopeisperforatedbyporesthatareabout100nmindiameter.
Atthelipofeachpore,theinnerandoutermembranesofthenuclearenvelopearefused
toformacontinuousmembrane.
Aproteinstructurecalledaporecomplexlineseachpore,regulatingthepassageof
certainlargemacromoleculesandparticles.
Thenuclearsideoftheenvelopeislinedbythenuclearlamina,anetworkofprotein
filamentsthatmaintainstheshapeofthenucleus.
Thereisevidencethataframeworkoffiberscalledthenuclearmatrixextendsthrough
thenuclearinterior.
Withinthenucleus,theDNAandassociatedproteinsareorganizedintodiscreteunits
calledchromosomes,structuresthatcarrythegeneticinformation.
Eachchromosomeismadeupoffibrousmaterialcalledchromatin,acomplexof
proteinsandDNA.
Stainedchromatinappearsthroughlightmicroscopesandelectronmicroscopesasa
diffusemass.
Asthecellpreparestodivide,thechromatinfiberscoilupandcondense,becomingthick
enoughtoberecognizedasthefamiliarchromosomes.

Eacheukaryoticspecieshasacharacteristicnumberofchromosomes.
Atypicalhumancellhas46chromosomes.
Ahumansexcell(eggorsperm)hasonly23chromosomes.
IG Lecture Outline 6-4

 Inthenucleusisaregionofdenselystainedfibersandgranulesadjoiningchromatin,the
nucleolus.
Inthenucleolus,ribosomalRNA(rRNA)issynthesizedandassembledwithproteins
fromthecytoplasmtoformribosomalsubunits.
Thesubunitspassthroughthenuclearporestothecytoplasm,wheretheycombineto
formribosomes.

ThenucleusdirectsproteinsynthesisbysynthesizingmessengerRNA(mRNA).
ThemRNAtravelstothecytoplasmthroughthenuclearporesandcombineswith
ribosomestotranslateitsgeneticmessageintotheprimarystructureofaspecific
polypeptide.

2.Ribosomesbuildacellsproteins.
Ribosomes,containingrRNAandprotein,aretheorganellesthatcarryoutprotein
synthesis.
Celltypesthatsynthesizelargequantitiesofproteins(e.g.,pancreascells)havelarge
numbersofribosomesandprominentnucleoli.
Someribosomes,freeribosomes,aresuspendedinthecytosolandsynthesizeproteins
thatfunctionwithinthecytosol.
Otherribosomes,boundribosomes,areattachedtotheoutsideoftheendoplasmic
reticulumornuclearenvelope.
Thesesynthesizeproteinsthatareeitherincludedinmembranesorexportedfromthe
cell.

Ribosomescanshiftbetweenrolesdependingonthepolypeptidestheyaresynthesizing.

D.TheEndomembraneSystem
Manyoftheinternalmembranesinaeukaryoticcellarepartoftheendomembrane
system.
Thesemembranesareeitherdirectlycontinuousorconnectedviatransferofvesicles,
sacsofmembrane.
Inspiteoftheseconnections,thesemembranesarediverseinfunctionandstructure.
Thethickness,molecularcompositionandtypesofchemicalreactionscarriedoutby
proteinsinagivenmembranemaybemodifiedseveraltimesduringamembraneslife.
Theendomembranesystemincludesthenuclearenvelope,endoplasmicreticulum,Golgi
apparatus,lysosomes,vacuoles,andtheplasmamembrane.
1.Theendoplasmicreticulummanufacturesmembranesandperformsmanyother
biosyntheticfunctions.

Theendoplasmicreticulum(ER)accountsforhalfthemembranesinaeukaryoticcell.

TheERincludesmembranoustubulesandinternal,fluidfilledspacescalledcisternae.

TheERmembraneiscontinuouswiththenuclearenvelope,andthecisternalspaceofthe
ERiscontinuouswiththespacebetweenthetwomembranesofthenuclearenvelope.

TherearetwoconnectedregionsofERthatdifferinstructureandfunction.
SmoothERlookssmoothbecauseitlacksribosomes.
RoughERlooksroughbecauseribosomes(boundribosomes)areattachedtotheoutside,
includingtheoutsideofthenuclearenvelope.
IG Lecture Outline 6-5

ThesmoothERisrichinenzymesandplaysaroleinavarietyofmetabolicprocesses.
EnzymesofsmoothERsynthesizelipids,includingoils,phospholipids,andsteroids.
Theseincludethesexhormonesofvertebratesandadrenalsteroids.
InthesmoothERoftheliver,enzymeshelpdetoxifypoisonsanddrugssuchasalcohol
andbarbiturates.
FrequentuseofthesedrugsleadstotheproliferationofsmoothERinlivercells,
increasingtherateofdetoxification.
Thisincreasestolerancetothetargetandotherdrugs,sohigherdosesarerequiredto
achievethesameeffect.
SmoothERstorescalciumions.
MusclecellshaveaspecializedsmoothERthatpumpscalciumionsfromthecytosol
andstorestheminitscisternalspace.
Whenanerveimpulsestimulatesamusclecell,calciumionsrushfromtheERinto
thecytosol,triggeringcontraction.
Enzymesthenpumpthecalciumback,readyingthecellforthenextstimulation.
RoughERisespeciallyabundantincellsthatsecreteproteins.
AsapolypeptideissynthesizedonaribosomeattachedtoroughER,itisthreadedinto
thecisternalspacethroughaporeformedbyaproteincomplexintheERmembrane.
Asitentersthecisternalspace,thenewproteinfoldsintoitsnativeconformation.
Mostsecretorypolypeptidesareglycoproteins,proteinstowhichacarbohydrateis
attached.
Secretoryproteinsarepackagedintransportvesiclesthatcarrythemtotheirnextstage.
RoughERisalsoamembranefactory.
Membraneboundproteinsaresynthesizeddirectlyintothemembrane.
EnzymesintheroughERalsosynthesizephospholipidsfromprecursorsinthecytosol.
AstheERmembraneexpands,membranecanbetransferredastransportvesiclestoother
componentsoftheendomembranesystem.

2.TheGolgiapparatusistheshippingandreceivingcenterforcellproducts.

ManytransportvesiclesfromtheERtraveltotheGolgiapparatusformodification
oftheircontents.

TheGolgiisacenterofmanufacturing,warehousing,sorting,andshipping.

TheGolgiapparatusisespeciallyextensiveincellsspecializedforsecretion.

TheGolgiapparatusconsistsofflattenedmembranoussacscisternaelookinglikea
stackofpitabread.
Themembraneofeachcisternaseparatesitsinternalspacefromthecytosol.
OnesideoftheGolgi,thecisside,islocatedneartheER.Thecisfacereceivesmaterial
byfusingwithtransportvesiclesfromtheER.
Theotherside,thetransside,budsoffvesiclesthattraveltoothersites.
Duringtheirtransitfromthecistothetransside,productsfromtheERareusually
modified.
TheGolgicanalsomanufactureitsownmacromolecules,includingpectinandother
noncellulosepolysaccharides.

TheGolgiapparatusisaverydynamicstructure.
IG Lecture Outline 6-6

Accordingtothecisternalmaturationmodel,thecisternaeoftheGolgiprogressfromthe
cistothetransface,carryingandmodifyingtheirproteincargoastheymove.

Finally,theGolgisortsandpackagesmaterialsintotransportvesicles.
Molecularidentificationtagsareaddedtoproductstoaidinsorting.
Productsaretaggedwithidentifierssuchasphosphategroups.TheseactlikeZIPcodes
onmailinglabelstoidentifytheproductsfinaldestination.

3.Lysosomesaredigestivecompartments.
Alysosomeisamembraneboundsacofhydrolyticenzymesthatananimalcellusesto
digestmacromolecules.

Lysosomalenzymescanhydrolyzeproteins,fats,polysaccharides,andnucleicacids.

TheseenzymesworkbestatpH5.
Proteinsinthelysosomalmembranepumphydrogenionsfromthecytosolintothelumen
ofthelysosomes.
Ruptureofoneorafewlysosomeshaslittleimpactonacellbecausethelysosomal
enzymesarenotveryactiveattheneutralpHofthecytosol.
However,massiveruptureofmanylysosomescandestroyacellbyautodigestion.

LysosomalenzymesandmembranearesynthesizedbyroughERandthentransferredto
theGolgiapparatusforfurthermodification.
Proteinsontheinnersurfaceofthelysosomalmembranearesparedbydigestionbytheir
threedimensionalconformations,whichprotectvulnerablebondsfromhydrolysis.

Lysosomescarryoutintracellulardigestioninavarietyofcircumstances.

Amoebaseatbyengulfingsmallerorganismsbyphagocytosis.
Thefoodvacuoleformedbyphagocytosisfuseswithalysosome,whoseenzymesdigest
thefood.
Asthepolymersaredigested,monomerspasstothecytosoltobecomenutrientsforthe
cell.

Lysosomescanplayaroleinrecyclingofthecellsorganellesandmacromolecules.
Thisrecycling,orautophagy,renewsthecell.
Duringautophagy,adamagedorganelleorregionofcytosolbecomessurroundedby
membrane.
Alysosomefuseswiththeresultingvesicle,digestingthemacromoleculesandreturning
theorganicmonomerstothecytosolforreuse.

Thelysosomesplayacriticalroleintheprogrammeddestructionofcellsinmulticellular
organisms.
Thisprocessplaysanimportantroleindevelopment.
Thehandsofhumanembryosarewebbeduntillysosomesdigestthecellsinthetissue
betweenthefingers.
Thisimportantprocessiscalledprogrammedcelldeath,orapoptosis.
4.Vacuoleshavediversefunctionsincellmaintenance.

Vesiclesandvacuoles(largerversions)aremembraneboundsacswithvariedfunctions.
Foodvacuolesareformedbyphagocytosisandfusewithlysosomes.
Contractilevacuoles,foundinfreshwaterprotists,pumpexcesswateroutofthecellto
maintaintheappropriateconcentrationofsalts.
IG Lecture Outline 6-7

Alargecentralvacuoleisfoundinmanymatureplantcells.
Themembranesurroundingthecentralvacuole,thetonoplast,isselectiveinits
transportofsolutesintothecentralvacuole.
Thefunctionsofthecentralvacuoleincludestockpilingproteinsorinorganicions,
disposingofmetabolicbyproducts,holdingpigments,andstoringdefensive
compoundsthatdefendtheplantagainstherbivores.
Becauseofthelargevacuole,thecytosoloccupiesonlyathinlayerbetweenthe
plasmamembraneandthetonoplast.Thepresenceofalargevacuoleincreases
surfaceareatovolumeratioforthecell.

E.OtherMembranousOrganelles
1.Mitochondriaandchloroplastsarethemainenergytransformersofcells.
Mitochondriaandchloroplastsaretheorganellesthatconvertenergytoformsthatcells
canuseforwork.
Mitochondriaarethesitesofcellularrespiration,generatingATPfromthecatabolismof
sugars,fats,andotherfuelsinthepresenceofoxygen.

Chloroplasts,foundinplantsandalgae,arethesitesofphotosynthesis.
Theyconvertsolarenergytochemicalenergyandsynthesizeneworganiccompounds
suchassugarsfromCO2andH2O.

Mitochondriaandchloroplastsarenotpartoftheendomembranesystem.
Incontrasttoorganellesoftheendomembranesystem,eachmitochondrionorchloroplast
hastwomembranesseparatingtheinnermostspacefromthecytosol.
TheirmembraneproteinsarenotmadebytheER,butratherbyfreeribosomesinthe
cytosolandbyribosomeswithintheorganellesthemselves.

BothorganelleshavesmallquantitiesofDNAthatdirectthesynthesisofthe
polypeptidesproducedbytheseinternalribosomes.

Mitochondriaandchloroplastsgrowandreproduceassemiautonomousorganelles.

Almostalleukaryoticcellshavemitochondria.
Theremaybeoneverylargemitochondrionorhundredstothousandsofindividual
mitochondria.
Thenumberofmitochondriaiscorrelatedwithaerobicmetabolicactivity.
Atypicalmitochondrionis110micronslong.
Mitochondriaarequitedynamic:moving,changingshape,anddividing.

Mitochondriahaveasmoothoutermembraneandaconvolutedinnermembranewith
infoldingscalledcristae.
Theinnermembranedividesthemitochondrionintotwointernalcompartments.
Thefirstistheintermembranespace,anarrowregionbetweentheinnerandouter
membranes.
Theinnermembraneenclosesthemitochondrialmatrix,afluidfilledspacewithDNA,
ribosomes,andenzymes.
Someofthemetabolicstepsofcellularrespirationarecatalyzedbyenzymesinthe
matrix.
ThecristaepresentalargesurfaceareafortheenzymesthatsynthesizeATP.
Thechloroplastisoneofseveralmembersofageneralizedclassofplantstructurescalled
plastids.
IG Lecture Outline 6-8

Amyloplastsarecolorlessplastidsthatstorestarchinrootsandtubers.
Chromoplastsstorepigmentsforfruitsandflowers.
Chloroplastscontainthegreenpigmentchlorophyllaswellasenzymesandother
moleculesthatfunctioninthephotosyntheticproductionofsugar.

Chloroplastsmeasureabout2microns5micronsandarefoundinleavesandother
greenorgansofplantsandalgae.
Thecontentsofthechloroplastareseparatedfromthecytosolbyanenvelopeconsisting
oftwomembranesseparatedbyanarrowintermembranespace.
Insidetheinnermostmembraneisafluidfilledspace,thestroma,inwhichfloat
membranoussacs,thethylakoids.
ThestromacontainsDNA,ribosomes,andenzymes.
Thethylakoidsareflattenedsacsthatplayacriticalroleinconvertinglighttochemical
energy.Insomeregions,thylakoidsarestackedlikepokerchipsintograna.
Themembranesofthechloroplastdividethechloroplastintothreecompartments:the
intermembranespace,thestroma,andthethylakoidspace.

Likemitochondria,chloroplastsaredynamicstructures.
Theirshapeisplastic,andtheycanreproducethemselvesbypinchingintwo.

Mitochondriaandchloroplastsaremobileandmovearoundthecellalongtracksofthe
cytoskeleton.
2.PeroxisomesgenerateanddegradeH2O2inperformingvariousmetabolicfunctions.

Peroxisomescontainenzymesthattransferhydrogenfromvarioussubstratestooxygen.
Anintermediateproductofthisprocessishydrogenperoxide(H 2O2),apoison.
TheperoxisomecontainsanenzymethatconvertsH 2O2towater.
Someperoxisomesbreakfattyacidsdowntosmallermoleculesthataretransportedto
mitochondriaasfuelforcellularrespiration.
Peroxisomesintheliverdetoxifyalcoholandotherharmfulcompounds.
Specializedperoxisomes,glyoxysomes,convertthefattyacidsinseedstosugars,which
theseedlingcanuseasasourceofenergyandcarbonuntilitiscapableof
photosynthesis.
Peroxisomesareboundbyasinglemembrane.

Theyformnotfromtheendomembranesystem,butbyincorporationofproteinsand
lipidsfromthecytosol.

Theysplitintwowhentheyreachacertainsize.

F.TheCytoskeleton

Thecytoskeletonisanetworkoffibersextendingthroughoutthecytoplasm.

Thecytoskeletonorganizesthestructuresandactivitiesofthecell.

1.Thecytoskeletonprovidessupport,motility,andregulation.

Thecytoskeletonprovidesmechanicalsupportandmaintainscellshape.

Thecytoskeletonprovidesanchorageformanyorganellesandcytosolicenzymes.

Thecytoskeletonisdynamicandcanbedismantledinonepartandreassembledin
anothertochangetheshapeofthecell.
IG Lecture Outline 6-9

 Thecytoskeletonalsoplaysamajorroleincellmotility,includingchangesincell
locationandlimitedmovementsofpartsofthecell.

Thecytoskeletoninteractswithmotorproteinstoproducemotility.
Cytoskeletonelementsandmotorproteinsworktogetherwithplasmamembrane
moleculestomovethewholecellalongfibersoutsidethecell.
Motorproteinsbringaboutmovementsofciliaandflagellabygrippingcytoskeletal
componentssuchasmicrotubulesandmovingthempasteachother.
Thesamemechanismcausesmusclecellstocontract.

Insidethecell,vesiclescantravelalongmonorailsprovidedbythecytoskeleton.

Thecytoskeletonmanipulatestheplasmamembranetoformfoodvacuolesduring
phagocytosis.

Cytoplasmicstreaminginplantcellsiscausedbythecytoskeleton.

Recently,evidencesuggeststhatthecytoskeletonmayplayaroleintheregulationof
biochemicalactivitiesinthecell.
Therearethreemaintypesoffibersmakingupthecytoskeleton:microtubules,
microfilaments,andintermediatefilaments.
Microtubules,thethickestfibers,arehollowrodsabout25micronsindiameterand200
nmto25micronsinlength.
Microtubulefibersareconstructedoftheglobularproteintubulin.
Eachtubulinmoleculeisadimerconsistingoftwosubunits.
Amicrotubulechangesinlengthbyaddingorremovingtubulindimers.
Microtubulesshapeandsupportthecellandserveastrackstoguidemotorproteins
carryingorganellestotheirdestination.

Microtubulesarealsoresponsiblefortheseparationofchromosomesduringcelldivision.

Inmanycells,microtubulesgrowoutfromacentrosomenearthenucleus.
Thesemicrotubulesresistcompressiontothecell.

Inanimalcells,thecentrosomehasapairofcentrioles,eachwithninetripletsof
microtubulesarrangedinaring.
Beforeacelldivides,thecentriolesreplicate.
Aspecializedarrangementofmicrotubulesisresponsibleforthebeatingofciliaand
flagella.
Manyunicellulareukaryoticorganismsarepropelledthroughwaterbyciliaandflagella.
Ciliaorflagellacanextendfromcellswithinatissuelayer,beatingtomovefluidoverthe
surfaceofthetissue.
Forexample,cilialiningthewindpipesweepmucuscarryingtrappeddebrisoutof
thelungs.

Ciliausuallyoccurinlargenumbersonthecellsurface.
Theyareabout0.25micronsindiameterand220micronslong.

Thereareusuallyjustoneorafewflagellapercell.
Flagellaarethesamewidthascilia,but10200micronslong.

Ciliaandflagelladifferintheirbeatingpatterns.
Aflagellumhasanundulatorymovementthatgeneratesforceinthesamedirectionasthe
flagellumsaxis.
IG Lecture Outline 6-10

Ciliamovemorelikeoarswithalternatingpowerandrecoverystrokesthatgenerateforce
perpendiculartotheciliumsaxis.

Inspiteoftheirdifferences,bothciliaandflagellahavethesameultrastructure.
Bothhaveacoreofmicrotubulessheathedbytheplasmamembrane.
Ninedoubletsofmicrotubulesarearrangedinaringaroundapairatthecenter.This9+
2patternisfoundinnearlyalleukaryoticciliaandflagella.
Flexiblewheelsofproteinsconnectouterdoubletstoeachotherandtothetwocentral
microtubules.
Theouterdoubletsarealsoconnectedbymotorproteins.
Theciliumorflagellumisanchoredinthecellbyabasalbody,whosestructureis
identicaltoacentriole.

Thebendingofciliaandflagellaisdrivenbythearmsofamotorprotein,dynein.
Additionandremovalofaphosphategroupcausesconformationchangesindynein.
Dyneinarmsalternatelygrab,move,andreleasetheoutermicrotubules.
Proteincrosslinkslimitsliding.Asaresult,theforcesexertedbythedyneinarmscause
thedoubletstocurve,bendingtheciliumorflagellum.

Microfilamentsaresolidrodsabout7nmindiameter.
Eachmicrofilamentisbuiltasatwisteddoublechainofactinsubunits.
Microfilamentscanformstructuralnetworksduetotheirabilitytobranch.

Thestructuralroleofmicrofilamentsinthecytoskeletonistobeartension,resisting
pullingforceswithinthecell.
Theyformathreedimensionalnetworkjustinsidetheplasmamembranetohelpsupport
thecellsshape,givingthecellcortexthesemisolidconsistencyofagel.
Microfilamentsareimportantincellmotility,especiallyaspartofthecontractile
apparatusofmusclecells.
Inmusclecells,thousandsofactinfilamentsarearrangedparalleltooneanother.
Thickerfilamentscomposedofmyosininterdigitatewiththethinneractinfibers.
Myosinmoleculesactasmotorproteins,walkingalongtheactinfilamentstoshortenthe
cell.
Inothercells,actinmyosinaggregatesarelessorganizedbutstillcauselocalized
contraction.
Acontractingbeltofmicrofilamentsdividesthecytoplasmofanimalcellsduringcell
division.
Localizedcontractionbroughtaboutbyactinandmyosinalsodrivesamoeboid
movement.
Pseudopodia,cellularextensions,extendandcontractthroughthereversible
assemblyandcontractionofactinsubunitsintomicrofilaments.

Microfilamentsassembleintonetworksthatconvertsoltogel.

Accordingtoawidelyacceptedmodel,filamentsnearthecellstrailing
edgeinteractwithmyosin,causingcontraction.

Thecontractionforcestheinteriorfluidintothepseudopodium,where
theactinnetworkhasbeenweakened.

Thepseudopodiumextendsuntiltheactinreassemblesintoanetwork.
Inplantcells,actinmyosininteractionsandsolgeltransformationsdrivecytoplasmic
streaming.
IG Lecture Outline 6-11

Thiscreatesacircularflowofcytoplasminthecell,speedingthedistributionofmaterials
withinthecell.

Intermediatefilamentsrangeindiameterfrom812nanometers,largerthan
microfilamentsbutsmallerthanmicrotubules.
Intermediatefilamentsareadiverseclassofcytoskeletalunits,builtfromafamilyof
proteinscalledkeratins.
Intermediatefilamentsarespecializedforbearingtension.
Intermediatefilamentsaremorepermanentfixturesofthecytoskeletonthanaretheother
twoclasses.

Theyreinforcecellshapeandfixorganellelocation.

G.CellSurfacesandJunctions
1.Plantcellsareencasedbycellwalls.

Thecellwall,foundinprokaryotes,fungi,andsomeprotists,hasmultiplefunctions.

Inplants,thecellwallprotectsthecell,maintainsitsshape,andpreventsexcessive
uptakeofwater.

Italsosupportstheplantagainsttheforceofgravity.

Thethicknessandchemicalcompositionofcellwallsdiffersfromspeciestospeciesand
amongcelltypeswithinaplant.
Thebasicdesignconsistsofmicrofibrilsofcelluloseembeddedinamatrixofproteins
andotherpolysaccharides.Thisisthebasicdesignofsteelreinforcedconcreteorfiberglass.
Amaturecellwallconsistsofaprimarycellwall,amiddlelamellawithsticky
polysaccharidesthatholdscellstogether,andlayersofsecondarycellwall.

Plantcellwallsareperforatedbychannelsbetweenadjacentcellscalledplasmodesmata.

2.Theextracellularmatrix(ECM)ofanimalcellsfunctionsinsupport,adhesion,movement,
andregulation.
Thoughlackingcellwalls,animalcellsdohaveanelaborateextracellularmatrix
(ECM).
Theprimaryconstituentsoftheextracellularmatrixareglycoproteins,especially
collagenfibers,embeddedinanetworkofglycoproteinproteoglycans.
Inmanycells,fibronectinsintheECMconnecttointegrins,intrinsicmembrane
proteinsthatspanthemembraneandbindontheircytoplasmicsidetoproteinsattachedto
microfilamentsofthecytoskeleton.
TheinterconnectionsfromtheECMtothecytoskeletonviathefibronectinintegrinlink
permittheintegrationofchangesinsideandoutsidethecell.

TheECMcanregulatecellbehavior.
Embryoniccellsmigratealongspecificpathwaysbymatchingtheorientationoftheir
microfilamentstothegrainoffibersintheextracellularmatrix.
Theextracellularmatrixcaninfluencetheactivityofgenesinthenucleusviaa
combinationofchemicalandmechanicalsignalingpathways.
Thismaycoordinatethebehaviorofallthecellswithinatissue.

3.Intercellularjunctionshelpintegratecellsintohigherlevelsofstructureandfunction.
IG Lecture Outline 6-12

 <BL1>Neighboringcellsintissues,organs,ororgansystemsoftenadhere,interact,and
communicatethroughdirectphysicalcontact.
Plantcellsareperforatedwithplasmodesmata,channelsallowingcytosoltopass
betweencells.
Waterandsmallsolutescanpassfreelyfromcelltocell.
Incertaincircumstances,proteinsandRNAcanbeexchanged.
Animalshave3maintypesofintercellularlinks:tightjunctions,desmosomes,andgap
junctions.
Intightjunctions,membranesofadjacentcellsarefused,formingcontinuousbelts
aroundcells.
Thispreventsleakageofextracellularfluid.
Desmosomes(oranchoringjunctions)fastencellstogetherintostrongsheets,muchlike
rivets.
Intermediatefilamentsofkeratinreinforcedesmosomes.
Gapjunctions(orcommunicatingjunctions)providecytoplasmicchannelsbetween
adjacentcells.
Specialmembraneproteinssurroundthesepores.
Ions,sugars,aminoacids,andothersmallmoleculescanpass.
Inembryos,gapjunctionsfacilitatechemicalcommunicationduringdevelopment.
4.Acellisalivingunitgreaterthanthesumofitsparts.
Whilethecellhasmanystructureswithspecificfunctions,allthesestructuresmustwork
together.
Forexample,macrophagesuseactinfilamentstomoveandextendpseudopodiato
capturetheirbacterialprey.
Foodvacuolesaredigestedbylysosomes,aproductoftheendomembranesystemofER
andGolgi.
Theenzymesofthelysosomesandproteinsofthecytoskeletonaresynthesizedonthe
ribosomes.
TheinformationfortheproteinscomesfromgeneticmessagessentbyDNAinthe
nucleus.
AlloftheseprocessesrequireenergyintheformofATP,mostofwhichissuppliedby
themitochondria.

Acellisalivingunitgreaterthanthesumofitsparts.

IG Lecture Outline 6-13