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of great value as to the life of the population, the influence of external
conditions, etc.
The most important investigation of this kind carried out at
Rothamsted was organised by Mr. Cutler.[3] A team of six workers was
assembled, and for 365 days without a break they counted every day
the ciliates, the amœbæ, the flagellates, and the bacteria in a plot of
arable ground, distinguishing no less than seventeen different kinds of
protozoa. The conclusions arrived at were carefully tested by the
Statistical Department.
Of the protozoa the flagellates were found to be the most numerous,
the amœbæ came next, and the ciliates were by far the fewest. The
numbers of each organism varied from day to day in a way that
showed conclusively the essentially trophic nature of the protozoan
population. The numbers of amœbæ—especially Dimastigamœba and
of a species called α—were sharply related to the numbers of
bacteria: when the amœbae were numerous the bacteria were few,
and vice versa. Detailed examination showed that the amœbæ were
probably the cause of the fluctuations in the bacterial numbers, but Mr.
Cutler has not yet been able to find why the amœbæ fluctuated; it
does not appear that temperature, moisture content, air supply or food
supply were determining causes. The flagellates and ciliates also
showed large fluctuations, amounting in one case—Oicomonas—to a
definite periodicity, apparently, however, not related to bacterial
numbers, or, so far as can be seen, to external conditions of moisture,
temperature and food supply, and showing no agreement with the
fluctuations of the amœbæ. However, one cannot be certain that lack
of agreement between curves expressing protozoan numbers and
physical factors implies absence of causal relationships: the
observations (though the best that can yet be made) are admittedly
not complete. If we saw only the end of the bough of a tree, and could
see no connection with a trunk, we might have much difficulty in
finding relationships between its motion and the wind; whatever the
direction of the wind it would move backwards and forwards in much
the same way, and even when the wind was blowing along the plane
of its motion it would just as often move against the wind as with it.
Meanwhile evidence was obtained that the twenty-four hour interval
adopted by the protozoological staff was too long for bacteria, and
accordingly the Bacteriological Department, under Mr. Thornton,
refined the method still further. Bacterial counts were made every two
hours, day and night, for several periods of sixty or eighty hours
without a break. The shape of the curve suggests that two hours is
probably close enough, and for the present counts at shorter intervals
are not contemplated. But there is at least one maximum and one
minimum in the day, although the bacterial day does not apparently
correspond with ours, nor can any relationship be traced with the
diurnal temperature curve.
The nitrate content of the soil was simultaneously determined by Mr.
Page and found to vary from hour to hour, but the variations did not
sharply correspond with the bacterial numbers; this, however, would
not necessarily be expected. The production of nitrate involves various
stages, and any lag would throw the nitrate and bacterial curves out of
agreement. There is a suggestion of a lag, but more counts are
necessary before it can be regarded as established.
Examination of these and other nitrate curves obtained at
Rothamsted has brought out another remarkable phenomenon. No
crop is growing on these plots, and no rain fell during the eighty hours,
yet nitrate is disappearing for a considerable part of the time. Where is
it going to? At present the simplest explanation seems to be that it is
taken up by micro-organisms. A similar conclusion had to be drawn
from a study of the nitrogen exhaustion of the soil. The whole of the
nitrate theoretically obtainable from the organic matter of the soil is not
obtained in the course of hours or even days; in one of our
experiments at Rothamsted nitrification is still going on, and is far from
complete, even after a lapse of fifty-three years. The explanation at
present offered is that part of the nitrate is constantly being absorbed
by micro-organisms and regenerated later on.
Now what organisms could be supposed to absorb nitrates from the
soil? Certain bacteria and fungi are known to utilise nitrates, and one
naturally thinks of algæ as possible agents also. Dr. Muriel Bristol was
therefore invited to study the algæ of the soil. Her account is given in
Chapter VI. She has found them not only on the surface, but scattered
throughout the body of the soil, even in the darkness of 4 inches, 5
inches, or 6 inches depth, where no light can ever penetrate, and
where photosynthesis as we understand it could not possibly take
place. Some modification in their mode of life is clearly necessary, and
it may well happen that they are living saprophytically. Dr. Bristol has
not yet, however, been able to count the algæ in the soil with any
certainty, although she has made some estimates of the numbers.
The quantitative work on the soil population indicates other
possibilities which are being investigated. There is not only a daily
fluctuation in the numbers, but so far as measurements have gone, a
seasonal one also. There seems to be some considerable uplift in
numbers of bacteria, protozoa, and possibly algæ and fungi in the
spring-time, followed by a fall in summer, a rise in autumn, and a fall
again in winter. At present we are unable to account for the
phenomenon, nor can we be sure that it is general until many more
data are accumulated.
In the cases of the protozoa and the algæ, there was a definite
reason for seeking them in the soil.
Another section of the population, the fungi, was simply found, and
at present we have only limited views as to their function. The older
workers considered that they predominated in acid soils, while
bacteria predominated in neutral soils. Present-day workers have
shown that fungi, including actinomycetes, are normal inhabitants of
all soils. The attempts at quantitative estimations are seriously
complicated by the fact that during the manipulations a single piece of
mycelium may break into fragments, each of which would count as
one, while a single cluster of spores might be counted as thousands.
Little progress has therefore been made on the quantitative lines
which have been so fruitful with protozoa. Dr. Brierley gives, in
Chapters VII. and VIII., a critical account of the work done on fungi.
In addition to the organisms already considered there are others of
larger size. The nematodes are almost visible to the unaided eye,
most of them are free living and probably help in the disintegration of
plant residues, though a few are parasitic on living plants and do much
injury to clover, oats, and less frequently to onions, bulbs, and
potatoes. Further, there are insects, myriapods and others, the effects
of which in the soil are not fully known. Special importance attaches to
the earthworms, not only because they are the largest in size and in
aggregate weight of the soil population, but because of the great part
they play in aerating the soil, gradually turning it over and bringing
about an intimate admixture with dead plant residues, as first
demonstrated by Darwin. Earthworms are the great distributors of
energy material to the microscopic population. Systematic quantitative
work on these larger forms is only of recent date, and Dr. Imms, in
Chapter IX., discusses our present knowledge.
TABLE I.
Soil Population, Rothamsted, 1922.
(The figures for algæ and fungi are first approximations only, and have considerably
less value than those for bacteria and protozoa.)

Approximate Weight
per Acre of—
Numbers Dry MatterNitrogen
per Gram Living in in
of Soil. Organisms. Organisms. Organisms.
Bacteria— lb. lb. lb.
High level 45,000,000 50
25 }
2 0·2
Low level 22,500,000
Protozoa—
Ciliates—
High level 1,000 — — —
Low level 100 — — —
Amœbæ—
High level 280,000 320
Low level 150,000 170 } 12 1·2
Flagellates—
High level 770,000 190
Low level 350,000 85 } 7 0·7
Algæ (not blue-green) [100,000] 125 6 0·6
Blue-green Say
Not known. — 6 Say 0·6
Fungi—
High level [1,500,000] 1700
Low level [700,000] 800 } 60 6·0
93 9·3
= 4 parts nitrogen per
1,000,000 of soil.

Larger Organisms.

Numbers Approximate Weight

per Acre.[D] per Acre of—
Living Dry Matter Nitrogen
 Organisms. in in
Organisms. Organisms.
Un- Un- Un- Un-
Ma- ma- Ma- ma- Ma- ma- Ma- ma-
nured. nured. nured. nured. nured. nured. nured. nured.
Oligochaeta
(Limicolae)— lb. lb. lb. lb. lb. lb.
Nematoda,
etc. 3,609,000 794,000 9 2 3 1 — —
Myriapoda 1,781,000 879,000 203 99 85 42 4 2
Insects 7,727,000 2,475,000 34 16 14 6 1 1
Earthworms 1,010,000 458,000 472 217 108 50 10 5
Total 210 99 15 9
Total organic matter (dry weight) in this soil = 126,000 lb. per acre.
Total nitrogen = 5700 lb. per acre. (1 lb. nitrogen per acre = 0·4 parts per 1,000,000
of soil.)
[D] To a depth of 9 inches. The weight of soil is approximately 1,000,000 kilos.

Are there any other members of the soil population that are of
importance? As already shown, the method of investigating the soil
population in use at Rothamsted is to find by chemical methods the
changes going on in the soil; to find by biological methods what
organisms are capable of bringing about these changes; and then to
complete the chain of evidence by tracing the relationships between
the numbers or activities of these organisms and the amount of
change produced. The list as we know it to-day is given in Table I.
The method, however, does not indicate whether the account is
fairly complete, or whether there are other organisms to be found. We
might, of course, trust to empirical hunting for organisms, or to chance
discoveries such as led Goodey to find the mysterious Proteomyxan
Rhizopods, which cannot yet be cultured with certainty, so that they
are rarely found by soil workers. It is possible that there are many
such organisms, and it is even conceivable that these unknown forms
far outnumber the known. The defect of the present method is that it
always leaves us in doubt as to the completeness of the list, and so
we may have to devise another.
Reverting to Table I., it obviously serves no purpose to add the
numbers of all the organisms together. We can add up the weights of
living organisms, of their dry matter or nitrogen, so as to form some
idea of the proportion of living to non-living organic matter, and this
helps us to visualise the different groups and place them according to
their respective masses. But a much better basis for comparing the
activities of the different groups would be afforded by the respective
amounts of energy they transform, if these could be determined. It is
proposed to attempt such measurements at Rothamsted. The results
when added would give the sum of the energy changes effected by
the soil population as we know it: the figure could be compared with
the total energy change in the soil itself as determined in a
calorimeter. If the two figures are of the same order of magnitude, we
shall know that our list is fairly well complete; if they are widely
different, search must be made for the missing energy transformers.
There are, of course, serious experimental difficulties to be overcome,
but we believe the energy relationships will afford the best basis for
further work on the soil population.
Finally, it is necessary to refer to the physical conditions obtaining in
the soil. These make it a much better habitat for organisms than one
might expect. At first sight one thinks of the soil as a purely mineral
mass. This view is entirely incorrect. Soil contains a considerable
amount of plant residues, rich in energy, and of air and water. The
usual method of stating the composition of the soil is by weight, but
this is misleading to the biologist because the mineral matter has a
density some two and a half times that of water and three times that of
the organic matter. For biological purposes composition by volume is
much more useful, and when stated in this way the figures are very
different from those ordinarily given. Table II. gives the results for two
Broadbalk arable plots, one unmanured and the other dunged; it
includes also a pasture soil.
The first requirement of the soil population is a supply of energy,
without which it cannot live at all. All our evidence shows that the
magnitude of the population is limited by the quantity of energy
available. The percentage by weight of the organic matter is about two
to four or five, and the percentage by volume runs about four to
twelve. Not all of this, however, is of equal value as source of energy.
About one-half is fairly easily soluble in alkalis, and may or may not be
of special value, but about one-quarter is probably too stable to be of
use to soil organisms.
 A second requirement is water with which in this country the soil is
usually tolerably well provided. Even in prolonged dry weather the soil
is moist at a depth of 3 inches below the surface. It is not uncommon
to find 10 per cent. or 20 per cent. by volume of water present, spread
in a thin film over all the particles, and completely saturating the soil
atmosphere.
TABLE II.
Volume of Air, Water and Organic Matter in 100 Volumes of
Rothamsted Soil.
In Pore Space.
Values Commonly
Solid Matter. Pore Obtained.
Mineral. Organic. Space. Water. Air.
(1) 62 4 34 23 11
(2) 51 11 38 30 8
(3) 41 12 47 40 7

(1) Arable, no manure applied to soil. (2) Arable, dung applied to soil. (3) Pasture.

The air supply is usually adequate owing to the rapidity with which
diffusion takes place. Except when the soil is water-logged, the
atmosphere differs but little from that of the one we breathe. There is
more CO2, but only a little less oxygen.[8] The mean temperature is
higher than one would expect, being distinctly above that of the air,
while the fluctuations in temperature are less.[5]
The reaction in normal soils is neutral to faintly alkaline; pH values
of nearly 8 are not uncommon. Results from certain English soils are
shown on p. 18.
The soil reaction is not easily altered. A considerable amount of acid
must accumulate before any marked increase in intensity of pH value
occurs; in other words, the soil is well buffered. The same can be said
of temperature, of water, and of energy supply. Like the reaction, they
alter but slowly, so that organisms have considerable time in which to
adapt themselves to the change.
Hydrogen Ion Concentration and Soil Fertility.
pH
Alkaline 10 Sterile: Alkali soil.
 9

8 Fertile: Arable.

Neutral 7

5 Potato Scab fails.

Nitrification hindered.
4 Barley fails.

Acid 3 Sterile: Peat.

A SELECTED BIBLIOGRAPHY.
[1] Berthelot, Marcellin, “Fixation directe de l’azote atmosphérique libre par
certains terrains argileux,” Compt. Rend., 1885, ci., 775-84.
[2] Boussingault, J. B., and Léwy, “Sur la composition de l’air confiné dans
la terre végétale,” Ann. Chim. Phys., 1853, xxxvii., 5-50.
[3] Cutler, D. W., Crump, L. M., and Sandon, H., “A Quantitative
Investigation of the Bacterial and Protozoan Population of the Soil, with an
Account of the Protozoan Fauna,” Phil. Trans. Roy. Soc., Series B, 1922,
ccxi., 317-50.
[4] Hellriegel, H., and Wilfarth, H., “Untersuchungen über die
Stickstoffnahrung der Gramineen und Leguminosen,” Zeitsch. des Vereins
f. d. Rübenzucker-Industrie, 1888.
[5] Keen, B. A., and Russell, E. J., “The Factors determining Soil
Temperature,” Journ. Agric. Sci., 1921, xi., 211-37.
[6] Lawes, J. B., and Gilbert, J. H., “On Agricultural Chemistry, Especially
in Relation to the Mineral Theory of Baron Liebig,” Journ. Roy. Agric. Soc.,
1851, xii., 1-40.
[7] Liebig, Justus, “Chemistry in its Application to Agriculture and
Physiology,” 1st and 2nd editions (1840 and 1841), 3rd and 4th editions
(1843 and 1847); “Natural Laws of Husbandry,” 1863.
[8] Russell, E. J., and Appleyard, A., “The Composition of the Soil
Atmosphere,” Journ. Agric. Sci., 1915, vii., 1-48; 1917, viii., 385-417.
 [9] Russell, E. J., and Hutchinson, H. B., “The Effect of Partial Sterilisation
of Soil on the Production of Plant Food,” Journ. Agric. Sci., 1909, iii., 111-
14; Part II., Journ. Agric. Sci., 1913, v., 152-221.
[10] Schloesing, Th., and Müntz, A., “Sur la Nitrification par les ferments
organisés,” Compt. Rend., 1877, lxxxiv., 301-3; 1877, lxxxv., 1018-20; and
1878, lxxxvi., 892-5. “Leçons de chimie agricole,” 1883.
[11] Warington, R., “On Nitrification,” Part I., Journ. Chem. Soc., 1878,
xxxiii., 44-51; Part II, Journ. Chem. Soc., 1879, xxxv., 429-56; Part III.,
Journ. Chem. Soc., 1884, xlv., 637-72; Part IV., Journ. Chem. Soc., 1891,
lix., 484-529.
[12] Way, J. T., “On the Composition of the Waters of Land Drainage and of
Rain,” Journ. Roy. Agric. Soc., 1856, xvii., 123-62.
[13] Winogradsky, S., “Recherches sur les organismes de la nitrification,”
Ann. de l’Inst. Pasteur, 1890, iv., 1e Mémoire, 213-31; 2e Mémoire, 257-75;
3e Mémoire, 760-71.
“Recherches sur l’assimilation de l’azote libre de l’atmosphère par les
microbes.” Arch. des Sci. Biolog. St. Petersburg, 1895, iii, 297-352.
For further details and fuller bibliography, see E. J. Russell, “Soil Conditions and
Plant Growth,” Longmans, Green & Co.
 CHAPTER II.
SOIL BACTERIA.

A. Occurrence and Methods of Study.

To understand the development of our knowledge of soil bacteria,

it must be remembered that bacteriology is under the disadvantage
that it started as an applied science. Although bacteria were first
seen by Leeuwenhoeck about the middle of the seventeenth century,
and some of their forms described by microscopists of the eighteenth
and early nineteenth centuries, it was only with the work of Pasteur
on fermentation, and of Duvaine, Pasteur, and their contemporaries
on disease bacteria, that bacteriology may be said to have started.
From the outset, therefore, attention has been directed mainly to the
bacteria in their specialised relationship to disease or to fermentation
and similar processes. As a result, little research was done on the
pure biology of the bacteria, so that even now many of the most
fundamental and elementary problems concerning them are quite
unsolved.
In their work on fermentations and disease bacteria, the earlier
workers were assisted by the fact that under both sets of conditions
the causative bacteria exist, as a rule, either in practically pure
culture, or else in preponderating numbers. The study and
elucidation of such a mixed micro-population as exists in the soil,
became possible only when methods had been devised for isolating
the different kinds of bacteria, and thus studying them apart from
each other. It was the development of the gelatine plate method of
isolating pure cultures by Koch[36] in 1881 that made the study of the
soil bacteria practicable. The plating method opened up two lines of
research. In the first place, it provided a simple means of isolating
organisms from the mixed population of the soil, and thus enabled a
qualitative study to be made of each organism in pure culture, and,
in the second place, from it was developed a counting technique for
estimating differences in bacterial numbers between samples of soil,
from which has sprung much of our knowledge of the quantitative
side.
The earliest studies of the soil bacteria consisted of such
estimations of numbers, and showed that the soil contained a very
numerous population of bacteria. About 20,000,000 bacteria per
gram of soil is now considered a fair average number. The number
and variety of bacteria existing in the soil is so enormous that the
method of separating out all the different forms, and of discovering
their characters and functions, has proved impracticable. In practice,
therefore, the problem has been approached from the biochemical
standpoint. That is to say, the special chemical changes that the
bacteria produce in the soil have first been investigated, and this has
been followed by the isolation and study of the various groups of
bacteria that bring about the changes under investigation.
The method commonly employed in isolating the organisms that
produce a given chemical change in the soil is called the “elective”
method. The soil is inoculated into a culture medium that will
especially favour the group of bacteria to be isolated, to the
exclusion of others. For example, if it is desired to isolate the
organisms that attack cellulose, a medium is made up containing no
other organic carbon compounds except cellulose. Such a selective
medium encourages the growth of the group of organisms to be
investigated, so that after several transfers to fresh medium a culture
is obtained containing only two or three different types of organisms.
These are separated by plating and pure cultures obtained.
Another difficulty which has not yet been completely overcome is
that of adequately describing an organism when it is isolated. The
morphology of bacteria is not the constant thing that is seen in the
more stable higher organisms. In many cases the appearance of a
single strain is entirely different on different media, and may be quite
altered by such conditions as changes in acidity of the medium or
temperature of incubation. Even on a single medium remarkable
changes in morphology occur, at any rate, in some bacteria. This is
well seen in a cresol-decomposing organism under investigation at
Rothamsted. In cultures a few days old this organism develops as
bent and branching rods; these rods then break up into chains of
cocci and short rods, which separate, and in old cultures all the
organisms may be in the coccoid form (Fig. 1). It is claimed by
Löhnis[47b] that the possession of a complex life-cycle of changing
forms is a universal character in the bacteria. The instability of shape
in many bacteria makes it necessary to standardise very carefully
the cultural conditions under which they are kept when their
appearance is described.

Culture 15 hours old. Culture 3 days old.

Fig. 1.—Change in appearance, in culture, of a cresol decomposing bacterium.

The inadequacy of mere morphology as a basis for describing

bacteria led to the search for diagnostic characters, based on the
biochemical changes that they produced in their culture media, and
the appearance of their growth in the mass on various media. These
characters unfortunately have also proved to be very much
influenced by the exact composition of the medium and other
conditions of culture. Recently an attempt has been made by the
American Society of Bacteriologists to standardise the diagnostic
characters used in describing bacteria, and also the media and
cultural conditions under which they are grown for the purpose of
description. The need for such precautions, however, was not
sufficiently realised by the early workers, many of whose
descriptions cannot now be referred to any definite organism.
 The large number of organisms found in the soil, and the difficulty
and labour of adequately describing them, is such that even now we
have no comprehensive description of the common soil bacteria that
appear on gelatine platings. A careful study based on modern
methods of characterisation has been made of certain selected
groups of bacteria, and it is hoped that the laborious systematic work
of describing the common forms will gradually be completed.
Several attempts have been made to classify the bacteria that
appear commonly on gelatine platings. This work was commenced
by Hiltner and Stormer in Germany, and continued by Chester,
Harding, and Conn in America. Conn[10], [14] found that the common
organisms fell into the following main groups:—
(1) Large spore-forming bacteria, related to Bacillus subtilis, which
form about 5-10 per cent. of the numbers. He adduced
evidence[12], [13] that these organisms exist in the soil mainly as
spores, so that they may not form an important part of the active soil
population.
(2) Short non-sporing organisms, related to Pseudomonas
fluorescens, that are rapid gelatine liquefiers. These form another 10
per cent. of the numbers.
(3) Short rod forms that liquefy gelatine slowly or not at all, and
develop colonies very slowly. These form 40-75 per cent. of the
numbers, and may therefore be of considerable importance in the
soil.
(4) A few micrococci also occur.
These groups comprise the larger portion of the bacterial flora of
the soil, but, in addition to these organisms, that develop on the
media commonly used for plating, there are special and important
groups that appear only on special media, either owing to their being
unable to grow on ordinary media or because they get swamped by
other forms. Examples of such groups are the ammonia and nitrite
oxidising bacteria, the nitrogen fixing groups, the cellulose
decomposing organisms, and the sulphur bacteria.
In order that we may apply the results of the study of a definite
organism to other localities, a knowledge of the geographical
distribution of the soil bacteria is clearly needed. We have,
unfortunately, very little knowledge of the distribution of soil
organisms. The common spore-forming groups appear to be
universally distributed. Thus Barthel, in a study of the bacterial flora
of soils from Greenland and the island of Disko, obtained soil
organisms belonging to the groups of Bacillus subtilis, B.
amylobacter, B. fluorescens, B. caudatus, and B. Zopfii, which are
common groups in European soil, indicating that the general
constitution of the bacterial flora of the soil in arctic regions is not
widely different from that of Western Europe. Bredemann, who made
an extensive study of the Bacillus amylobacter group, obtained soil
samples from widely scattered localities, and found these organisms
in soil from Germany, Holstein, Norway, Italy, Morocco, Teneriffe,
Russia, Japan, China, the East Indies, Samoa, Illinois, Arizona,
German East Africa, and the Cameroons. Some soil organisms, on
the other hand, are apparently absent from certain districts. This may
be due to the conditions, such as climatic environment, being
unfavourable to them. A study has recently been made at
Rothamsted of the distribution over Great Britain of a group of
bacteria that are capable of decomposing phenol and cresol. One of
these organisms, apparently related to the acid-fast B. phlœi, has an
interesting distribution. It has been found in 50 per cent. of the soils
samples examined from the drier region, where the annual rainfall is
less than 30 inches, but in only 20 per cent. of the samples in the
wetter parts of Britain. Another example of limited distribution is
found in the case of Bacillus radicicola, the organism that produces
tubercles on the roots of leguminous plants. The distribution of the
varieties of this organism follows that of the host plants with which
they are associated, so that when a new leguminous crop is
introduced into a country, nodules may not appear on the roots
unless the soil be specially inoculated with the right variety of
organism. In cases where a group of soil organisms is widely
distributed over the globe, it may yet be absent from many soils
owing to the soil conditions not suiting it. Thus, phenol decomposing
bacteria, though abundant in the neighbourhood of Rothamsted, are
yet absent from field plots that have been unmanured for a
considerable period. The occurrence of the nitrifying organisms and
the nitrogen fixing Azotobacter is also very dependent on the soil
conditions.
 Owing to the method by which our knowledge of soil bacteria has
been acquired, by studying first the chemical changes in the soil and
then the bacteria that produce them, it is natural for us to divide them
into physiological groups according to the chemical changes that
they bring about. This grouping is the more reasonable since so little
is known as to the true relationships of the different groups of
bacteria that a classification based on morphology is well-nigh
impossible. In considering the activities of bacteria in the soil, it is
convenient to group the changes which they bring about into the two
divisions into which they naturally fall in the economy of the
organisms.
In the first place, there are the changes that result in a release of
energy, which the bacteria utilise for their vital processes.
In the second place, there are the processes by which the bacteria
build up the material of their bodies. These building up processes
involve an intake of energy for their accomplishment.
It will be convenient to deal first with the release of energy for their
own use by bacteria, and its consequences.
 B. Activities Connected with the Acquirement of Energy.

Unlike the green plants, most bacteria are unable to obtain the
energy that is required for their metabolism from sunlight. They must,
therefore, make use of such chemical changes as will involve the
release of energy.
As an example of the acquirement of energy in this way may be
taken the oxidation of methane by B. methanicus. This organism,
described by Söhngen, obtains its energy supply by the conversion
of methane into CO2 and H2O.
CH4 + 2O2 = CO2 + 2H2O 220 Cal.
A further example is the acetic organism that obtains its energy
through the oxidation of alcohol to acetic acid.
C2H6O + O2 = C2H4O2 + H2O 115 Cal.
The decomposition processes brought about by micro-organisms
in obtaining energy are usually oxidations, but this is not necessarily
so, as can be seen in case of the fermentation of sugar into
alcohol.[E]
C6H12O6 = 2C2H6O + 2CO2 50 Cal.
[E] These examples are from Orla-Jensen (Centralblatt f. Bakt., II., Bd. 22,
p. 305).

By far the greater part of the decomposition of organic matter is

brought about by bacteria in the process of acquiring energy. In the
soil, nearly the whole of the material utilised by bacteria as a source
of energy is derived ultimately from green plants. The energy
materials left in the soil by the plant fall into two groups, the non-
nitrogenous compounds, which are mainly carbohydrates and their
derivatives, and the nitrogenous compounds, principally derived from
proteins.

(1) Decomposition of Non-nitrogenous Compounds.

 The simpler carbohydrates and starches are attacked and
decomposed by a large variety of bacteria. The addition of such
substances to soil causes a rapid increase in bacterial numbers. In
nature the sugars are in all probability among the first plant
constituents to be destroyed during the decay processes.
A large proportion of plant tissues consist of cellulose and its
derivatives. These compounds are consequently of great importance
in the soil. Unfortunately our knowledge of the processes by which
cellulose is broken down in the soil is very inadequate. The early
experimental study of cellulose decomposition, such as that of
Tappeiner[60] and Hoppe-Seyler,[33] was mostly carried out under
conditions of inadequate aeration, and the products of
decomposition were found to include methane and CO2, and
sometimes fatty acids and hydrogen. The bacteriology of this
anaerobic decomposition was studied by Omelianski,[54] who
described two spore-bearing organisms, one of which attacked
cellulose with the production of hydrogen, and the other with the
production of methane. Both species also produce fatty acids and
CO2. It is probable that these organisms operate in the soil under
conditions of inadequate aeration. In swamp soils, in which rice is
grown, it has been shown that methane, hydrogen, and CO2 are
evolved in the lower layers. In these soils, however, the methane and
hydrogen are oxidised when they reach the surface layers. This
oxidation is also effected by micro-organisms. Bacteria capable of
deriving energy by the oxidation of hydrogen gas have been isolated
and studied by Kaserer,[37] and by Nabokich and Lebedeff,[52] while
Söhngen[57] has isolated an organism which he named Bacillus
methanicus, that was capable of oxidising methane.
Under normal conditions in cultivated soils, however, the
decomposition of cellulose takes place in the presence of an
adequate air supply, and so follows a different course from that
studied by Omelianski. Our knowledge of this aerobic decomposition
is very scanty. A number of bacteria, capable of decomposing
cellulose aerobically, are known. A remarkable organism was
investigated by Hutchinson and Clayton,[30] who named it
Spirochæta cytophaga. This organism, which they isolated from
Rothamsted soil, though placed among the Spirochætoidea, is of
doubtful affinities. During the active condition it exists for the most
part as thin flexible rods tapered at the extremities. This form passes
into a spherical cyst-like stage, at first thought to be a distinct
organism (Fig. 2). Spirochæta cytophaga is very aerobic, working
actively, only at the surface of the culture medium. It is very selective
in its action. It appears unable to derive energy from any
carbohydrate other than cellulose. Indeed, many of the simple
carbohydrates, especially the reducing sugars, are toxic to the
organism in pure culture. An extensive study of aerobic cellulose
decomposition by bacteria was made by McBeth and Scales,[50] who
isolated fifteen bacteria having this power. Five of these were spore-
forming organisms. Unlike Spirochæta cytophaga, they are all able
to develop on ordinary media such as beef agar or gelatine, and are
thus not nearly so selective in their food requirements.

Fig. 2.—Spirochæta cytophaga. Changes occurring in culture. (After Hutchinson

and Clayton.)

We are at present ignorant as to which organisms are most

effective in decomposing cellulose in the soil under field conditions,
or what are the conditions best suited to their activity. It is possible
that fungi also help in the decomposition of cellulose to a great
extent. This subject of the decomposition of cellulose offers one of
the most promising fields of research in soil bacteriology. The
difficulty of the subject is further increased by our present ignorance
of the chemical aspect of cellulose decomposition. It has been
supposed that the early decomposition products are simpler sugars,
but these are not found under conditions in which cellulose is being
decomposed by pure cultures of the bacteria mentioned above.
Hutchinson and Clayton found that their organism produced volatile
acids, mucilage, and a carotin-like pigment. The organisms isolated
by McBeth and Scales also produce acids, and in some cases yellow
pigments. It is known, however, that the decomposition products of
cellulose can be utilised as energy supply for other organisms, such
as nitrogen fixing bacteria.
When plant remains decompose in the soil there are ultimately
produced brown colloidal bodies collectively known as humus. The
processes by which this humus is produced are not yet properly
understood. Humus is of great importance in the soil, in rendering
the soil suitable for the growth of crops. It affects the physical
properties of the soil to a great extent. In the first place, it improves
the texture of the soil, making heavy clay soils more friable, and
loose sandy soils more coherent. Secondly, it has great water-
retaining powers, so that soils rich in organic matter suffer
comparatively little during periods of drought. And lastly, it exerts a
strong buffering effect against soil acids. Now, it is one of the
problems of present-day farming that soil is becoming depleted of its
humus. This is due to the increasing scarcity of farmyard manure in
many districts, and the consequent use of mineral fertilisers to
supply nitrogen, potash, and phosphate to the crop. A need has
therefore arisen for a substitute for farmyard manure, by means of
which the humus content of soils may be kept up in districts where
natural manure is scarce.
Fig. 3.—Cellulose decomposed by S. cytophaga in media with increasing amounts
of nitrogen. (After Hutchinson and Clayton.)

X-axis: Milligrams of nitrogen supplied as sodium-ammonium

phosphate.
Y-axis: Milligrams of cellulose decomposed in 21 days.

It is well known that if fresh unrotted manure or straw be added to

the soil, it often produces harmful effects on the succeeding crop.
The problem, therefore, was to develop a method by which fresh
straw, before application to the soil, could be made to rot down to a
mixture of humus compounds such as occur in well-rotted farmyard
manure. The solution of this problem came as a result of an
investigation by Hutchinson and Richards,[30b] at Rothamsted, into
food requirements of the cellulose decomposing bacteria. They
realised that since more than 10 per cent. of the dry weight of
bacteria consists of nitrogen, it would be necessary to supply the
cellulose decomposing bacteria with a supply of nitrogen, in order
that they should attain their greatest activity. Experiments with