CHAPTER IX.. Soil and Subsoil (_continued_).

SOIL AND SUBSOIL (Continued).

ORGANISMS INFLUENCING SOIL CONDITIONS; BACTERIA, ETC.

MICRO-ORGANISMS OF THE SOIL.

Intimately correlated with the humus-substances of the soil, as well as with its temporary contents of the carbohydrates (cellulose, gums and sugars) from which humus is formed, is the multitudinous flora of micro-organisms always present and exercising important functions in connection with the growth of the higher plants. Extended researches by Adametz, Schloesing and Müntz, Miquel, Koch, Fraenkel, Winogradsky, Frank and many others, have thrown light upon the immense numbers and great variety of minute organisms, especially of the bacterial group, present in soils, and upon their distribution and activities in the same. It has been shown that their numbers are greatest near (although usually not at) the surface, decreasing rapidly downward and generally disappearing wholly at depths between seven and eight feet; the latter depth varying of course according to the nature and porosity of the soil, and both depth and numbers being greatest in summer.

Numbers of Bacteria in Soils.—Adametz found in one gram of soil, 38,000 bacteria at the surface, 460,000 at ten inches depth; in a loam soil at the surface 500,000, at ten inches 464,000 in each gram of earth. Of mould and similar fungous germs there were only 40 to 50 in the same, 6 species being true molds, while four were ferments, including the yeasts of wine and beer. Fraenkel found in virgin land from near Potsdam, a sudden, marked decrease at depths of from three to five feet; while in earth from inhabited places within the city of Berlin, considerable numbers were still present at eight and even ten feet, in some cases.

In the researches lately made by Hohl at the bacteriological station at Liebefeld, near Bern, it was found that in cultivated soils the number of bacteria greatly exceeds the figures given by Fraenkel. He found a gram of moist soil to contain from three to fifteen millions of bacteria. In the cultivated soil of Liebefeld he found 5,750,000, in meadow land 9,400,000, in a manure pile 44,500,000 per cubic centimeter. These figures seem high for so small a quantity of material, but taking the average size of a bacterium, a cubic centimeter might readily contain six hundred millions. (Grandeau, Ann. Sci. Agronomique, vol. 1, p. 461, 1905).

Mayo and Kinsley (Rep. Kansas Exp’t Station for 1902-3) have made elaborate investigations of the numbers and kinds of bacteria found in various soils in Kansas, in connection with different crops. It is noteworthy that in most cases their figures exceed considerably those given by European observers, as they often reach high into the millions, in one case to over fifty millions, per cubic centimeter.

The mode of statement in the paper is not always quite clear as to the manner in which the averages given were calculated. It must be remembered that these data refer to cubic centimeters of soil, or about twice the amount (1 gram) used by European observers.

Five fields with different soils were investigated; the land being described as follows: “Field No. 1 is a black loam containing considerable humus; field No. 2 is similar to field 1 but contains more humus; field No. 3 is a thin soil with clay gumbo subsoil; fields Nos. 4 and 5 are black loams, but not as rich in humus as either No. 1 or No. 2.”

The average bacterial contents of the several fields are given as follows:

Field No. 1 33,931,747 per cubic centimeter. “ No. 2 53,596,060 “ “ “ “ No. 3 78,534 “ “ “ “ No. 4 8,643,006 “ “ “ “ No. 5 3,192,131 “ “ “

“The crop records of these fields for the past ten years indicate that the crop yield has been (more or less?) directly proportional to the bacterial content of the soil of each field; field 2 has produced the largest yield, field 3 the least.”

Unfortunately no chemical analyses of any of these soils are communicated; but at the request of the writer samples of the soils of the first three fields were sent from the Kansas station for humus determinations (courteously made by Dr. H. C. Myers), which gave the following results:

Field No. 1 2.19% of Humus. “ No. 2 3.07% “ “ “ No. 3 1.85% “ “

While these humus-percentages are not directly proportional to the bacterial content, a favoring effect of high humus-content is clearly shown. The bacterial and the humus-content of these soils are sensibly, even if not directly, correlated; which might reasonably be expected, since the organic matter and the humus are the bacterial food.

The investigation also showed wide differences in the bacterial content of the same soil when different crops were growing on it. Thus in samples taken on Aug. 15, there were found in the first twelve inches of a black loam soil bearing timothy and clover, 1,380,000, in the same with alfalfa and clover, 21,091,000, with maize from one to over two millions. In soils from the western part of Kansas, the bacterial content of the same crops was much less (as doubtless is the humus-content), and it is noteworthy that the prairie buffalo grass shows throughout a relatively high bacterial content in the first foot of the soil, ranging next to alfalfa. The root bacteria living on the legumes will naturally increase the bacterial content of the soils on which they grow, more than plants which, like maize, do not directly utilize bacterial action.

Multiplication of the Bacteria.—Marshall Ward and Duclaux have made some special observations in regard to the rapidity with which certain bacteria multiply. Duclaux summarizes the final conclusion thus: taking as a basis the time of 35 minutes for the subdivision into two, which has been frequently observed by Ward, there would be four millions of bacteria produced in twelve hours. The first filaments had plenty of room in a drop culture of one cubic millimeter; but at the end their total volume amounted to the tenth part of the total volume of the drop. At the above rate, making 48 generations in 24 hours, 281,500 billions of organisms would be produced. (Grandeau, Ann. Sci. Agron. Vol. 1, 1905, p. 456).

Aerobic and Anaerobic Bacteria.—As may readily be inferred, the cultural and other surface conditions exert a potent influence both upon the kinds and abundance of the bacteria and molds; since the life-functions of some are dependent upon the presence of free oxygen (“aerobic”), while others flourish best, or only, in the absence of air (“anaerobic”), or are able to avail themselves of the presence of combined oxygen, by reduction of oxids present. Their number is found, in general, to be greatest in cultivated lands, and bacteria are there by far predominant over the moulds. On the other hand, the moulds gain precedence in woodlands and meadows, at least so far as air can gain access; while in the deeper layers of the same, as well as in peaty lands, bacterial life is always scanty. This holds particularly in respect to the nitrifying organisms, and others whose life-functions are dependent upon abundant access of oxygen (aerobic).

Food Material Required.—All bacteria, like the fungi, are dependent for their development upon the presence of adequate amounts of some organic food-material, best apparently in water-soluble form. In the soil it seems to be chiefly compounds of the carbohydrate group, especially various gums derived from the decaying plant substance, or from stable manure; in artificial cultures, glucose is mostly found to be a highly available food. When the decaying substance reaches the state of humus, the latter seems to be available as food only to comparatively few bacteria. The very abundant development of bacterial life seems to be among the most important effects produced by stable manure upon the surface soil, in establishing good tilth (“Bodengare” in German).

Functions of the Bacteria.—While there is still much uncertainty as to the exact functions performed by most of these bacteria in respect to soil-formation and plant growth, there are several kinds whose activity has been proved to be of the utmost importance in one or both directions; it having been shown that when the soil is sterilized either by heat or antiseptic agents, certain essential processes are completely suppressed until the soil is re-infected and the conditions of bacterial life restored.

Probably the chief in importance are those connected with the processes of nitrification and denitrification, bearing as they do upon the supply to plants of the most costly of the three substances furnished by fertilizers. These organisms have been first extensively studied by Winogradsky, while the conditions of their activity have been largely developed by R. Warington.

Nitrifying Bacteria.—The conversion of ammonia into nitrates is accomplished under proper conditions by two organisms, or groups of organisms; the first stage being the formation of nitrites by the round, often flagellate cells of nitrosomonas (or nitrosococcus). The second, the oxidation of the nitrites into nitrates by very minute rod-shaped bacilli, named nitrobacteria. The conditions under which these bacteria can act are quite definite in that, aside from a supply of the nitrifiable substance, a fairly high temperature (24° C. or 75° F.) and a moderate degree of moisture, there must be a free access of oxygen (air); and there must be present a base (or its carbonate) with which the acids formed by oxidation can immediately unite. In an acid medium (“sour” soils) nitrification promptly ceases; as it also does whenever the amount of base present has been fully neutralized. The bases most favorable to nitrification are lime and magnesia in the form of carbonates, an excess of which does no harm; while in the case of the carbonates of potash and soda, the amount must be strictly limited.

Conditions of Activity.—Dumont and Crochetelle found that up to .25 per cent, potassic carbonate acted favorably on the process; which was, however, completely stopped by as much as .8 per ct. Warington has shown that ammonic carbonate similarly prevents nitrification when exceeding about .37 per ct. Ammonia salts in general appear to be antagonistic to the transformation of nitrites into nitrates.

Aside from the carbonates, some neutral salts favor nitrification very markedly; while others tend to depress it. Deherain found that .5 per cent of common salt suffices to prevent nitrification altogether, while smaller amounts retard it proportionally. According to Dumont and Crochetelle, potassium chlorid acts favorably up to .3 per cent, but at .8 per cent suppresses nitrification. Earthy and alkaline sulfates, on the contrary, seem to act favorably throughout, at least up to .5 per cent. This is especially true of gypsum, which, according to Pichard, accelerates the process more than any other substance known. Taking the effect of gypsum as the maximum, he found that, other things being equal, the amounts of nitrates formed were as shown in the table below, the effect of gypsum being taken as 100:

Gypsum 100 Sodic Sulfate 47.9 Potassic Sulfate 35.8 Calcic Carbonate 13.3 Magnesic Carbonate 12.5

The above estimates are markedly confirmed by the observations of the writer in the alkali soils of California. In these, nitrates exist most abundantly when the salts contained in the soil are mainly sulfates; while wherever common salt or sodic carbonate are present in considerable amounts, the amounts of nitrate found are notably less. In saline seashore lands nitrates are usually present in traces only. Wollny has moreover shown that the nitrates themselves exert a repressive influence on nitrification.

Effects of Aeration and Reduction.—While the fostering effect of sulfates upon nitrification is very energetic in well aerated soils, they become injurious whenever by a reductive process in ill-drained lands, the sulfates are reduced to sulfids. Under such conditions the process will in any case be much impaired. On the other hand, the favoring effect of abundant aeration was strikingly shown in the experiment made by Deherain, in which a cubic meter of soil was left unmoved for several months, while a similar mass was thoroughly agitated once a week during the same time. The proportion of nitrates formed in the latter case was as 70 to 1 formed in the quiescent soil mass. It follows that the intensity of nitrification is essentially dependent upon the porosity of the soil; and that it is thus greatly favored in the pervious soil-strata of the arid regions. It also follows that thorough and frequent tillage and fallowing greatly favor nitrification; thus explaining one of the beneficial results of these operations. At the same time, it is true that we may thus in a short time seriously diminish the reserve stock of nitrogen contained in the soil in the form of humus-amids; and since nitrates are exceedingly liable to be lost from the soil in several ways, such excessive nitrification is to be avoided.

Unhumified Organic Matter does not Nitrify.—There can be little doubt that the formation of ammonia from the amido-compounds in humus is also the work of bacteria; but this, really the initial phase of the nitrogen-nutrition of plants, has not yet been fully elucidated. That, however, it is essentially only the ready-formed humus and not the unhumified debris of the soil which participate in nitrification was shown by the experiments of the writer, see chapter 19.

Denitrifying Bacteria.—Among the sources of loss of nitrates in the soil is the action of denitrifying bacteria; some of which cause merely the reduction of nitrates to nitrites and progressively to ammonia, while others cause gaseous nitrogen to be given off from nitrites and nitrates, resulting in their complete loss to the soil. While there are probably several kinds of the latter class, the most rapidly effective is an organism contained abundantly in fresh horse dung, and also on the surface of old straw. This can readily be shown by subjecting a very dilute solution (1-3 per cent.) of Chile saltpeter to the action of fresh horse dung in a close flask, when nitrogen and carbonic dioxid gases are evolved, and in a few days the nitrate has totally disappeared. In the course of time this power of horse-manure disappears; so that “rotted manure” is practically free from it and under proper conditions serves nitrification so effectively, that in the past it has served extensively for the production of saltpeter in the “niter-plantations” for the industrial purposes; the material of which was loose earth, marl and manure, kept moist and frequently forked over for better aeration. Saltpeter is similarly produced in stables, corroding the mortar of brick foundations. Nevertheless, it is necessary to avoid the use, either together or at short intervals apart, of Chile saltpeter and fresh manure; the manure if used first should be allowed to remain at least two months in the soil before saltpeter is applied.

The reduction of nitrates to nitrites and ammonia is brought about by quite a number of bacteria, mostly anaerobic, and such as consume combined oxygen in their development. Thus the butyric ferment, which in the absence of readily reducible compounds evolves free hydrogen, will in presence of nitrates reduce the latter to nitrites, or form ammonia by addition of hydrogen to nitrogen just set free by reduction. Such reductive processes of course occur chiefly in soils rich in organic matter, or ill-aerated. The ammonia so formed, while at first simply combining with any humus acids present, may in the course of time be itself reduced to the amidic condition, being thereby rendered relatively inert, until again brought into action by ammonia-forming bacteria.

Ammonia-forming Bacteria.—A large number of different bacteria appear to be concerned in the formation of ammonia from compounds of the albuminoid group, (and probably from humus). Among these is one of the most common in soils (Bacillus mycoides, root bacillus), which while forming ammonia carbonate in solutions of albumen, is also capable of reducing nitrates to nitrites and ammonia in presence of a nutritive solution of sugar.

The “hay bacillus” (B. subtilis), so abundantly developed in hay infusions, and one of the most abundant in cultivated soils, has together with B. ellenbachensis, B. megatherium, B. mycoides, and others, by some been credited with important action in favoring vegetation; so that a fairly pure culture of B. ellenbachensis has been brought out commercially in Germany under the name of “Alinit.” Rigorous culture experiments made by Stutzer and others have, however, failed to show any general benefit from the use of alinit in infecting either land or seeds. But there is no doubt of the Effects of Bacterial Life on Physical Soil Conditions.—It is apparent that all conditions favoring the life of aerobic (air-needing) bacteria tend also to produce the loose, porous state (tilth) of the surface soil so conducive to the welfare of culture plants, designated by German agriculturists as “Bodengare.” Whether or not this condition is directly due to bacterial processes, as is thought by Stutzer (Landw. Presse, 1904, No. 11) it is assuredly a highly important point to be gained, and is essentially connected with the presence of humus in adequate amounts, which is also a favoring condition of abundant bacterial life. It seems that the preference given to the shallow putting-in, or even surface application of stable manure, existing in Europe, is largely based upon the marked effect upon the looseness of the surface soil, generally credited to the physical effect of the manure substance itself, but apparently largely due to the intensity of bacterial action thus brought about.

ROOT-BACTERIA OR RHIZOBIA OF LEGUMES.—Among the most important bacteria, agriculturally, is that which enables plants of the leguminous order—(peas, beans, vetches, clovers, lupins, etc.),—to obtain their supply of nitrogen from the air independently of those contained in the soil. The source of nitrogen to plants was long a disputed question; it was at first supposed (by de Saussure) that it was obtained directly from the soil by the absorption of humus; but this was disproved, and Liebig then contended that it was derived directly from the atmosphere through the ammonia in rain water. This was then shown to be wholly inadequate; and Boussingault proved conclusively that plants do not take up nitrogen gas from the air. This was subsequently denied by Ville; but investigation at the Rothamstead agricultural station by Lawes and Gilbert definitely confirmed Boussingault’s results. At the same time they also proved very definitely that while grass and root crops deplete the soil of nitrogen, clover and other leguminous crops leave in the soil more nitrogen than was previously present, even when the entire, itself highly nitrogenous, leguminous crop is removed from the land. The improvement of lands for wheat production by rotation with clover had long ago become a practical maxim; but the cause was not understood until, in 1888, Hellriegel and Wilfarth announced that the variously-shaped excrescences or tubercles which had long been observed as frequently deforming the roots of legumes, are caused by the attacks of bacilli capable of absorbing the free nitrogen of the air and thus enabling the host-plant to acquire its needed supply by absorbing the richly nitrogenous matter thus accumulated in the excrescences. The minute rod-shaped organism was named Bacillus radicicola by Beyerinck; Rhizobium leguminosarum, by A. Frank, who has published an extensive treatise on the subject.

Uber die Pilzsymbcose der Leguminosen, Berlin, 1890.

Original figure from drawing by O. Butler, Asst. in Agr. Dep’t Univ. of California.

Microscopic examination of the nodules shows their tissues to contain partly motile, free bacteria, partly others (bacteroids), which have assumed a quiescent condition, and are of much greater dimensions than those of the motile form. These relatively thick, and sometimes forked, forms, differing somewhat in each of the group adaptations mentioned below, constitute the bulk of the cell-contents of the nodules, and ultimately serve for the nutrition of the host-plant with nitrogen. When the growth of the excrescence is completed, the swollen, quiescent bacteroids gradually collapse and become depleted of their nitrogenous substance; and finally the apparently empty husk remains or drops off, carrying with it the minute cocci which in the soil become active bacteria again. The nodules are thus found mainly on the actively-growing roots, and at the time when vegetation and assimilation are most active in the plant. In autumn, or when the plants are in fruit, the roots may be wholly destitute of nodules.

The adhesion of the nodules to the roots is mostly very loose, and their falling-off when the seedlings are carelessly transplanted, doubtless accounts for much of the difficulty generally found in transplanting legumes when once established.

The figures annexed show the various forms assumed by the nodules in different plants, and with them also the corresponding forms of the bacteroids of each. The latter, here shown magnified about 1000 times, are taken from the inaugural dissertation of D. Brock on this subject, published at Leipzig in 1891. It appears that the forms of the bacteroids are quite as much varied as are those of the nodules they form.

FIG. 23.—Bur clover.—Medicago denticulata.

FIG. 24.—Garden pea.—Pisum sativum.]

Varieties of Forms.—While these bacilli seem to be normally present in most soils, it seems to be necessary that they should adapt themselves for this symbiosis with each of several groups of the legumes in order to exert their most beneficial effects. In many soils there appears to exist a “neutral form”, which requires about a season’s time or more to adapt itself specially to the several leguminous groups so that a great advantage is gained by infecting either the seeds or the soil with the forms already adapted, when no similar plant has lately occupied the same ground. Thus the bacillus of the clover root is of little or no benefit to beans, peas or alfalfa, and the root-bacilli of each of the latter are relatively ineffectual when used to infect either of the other groups. The same is true of the bacilli of lupins and of acacias, as applied to leguminous plants of any other groups.

“Living together” beneficially; in contradistinction to parasitism, which is injurious to the host plant.

It is asserted by some observers that the root-bacilli producing differently-shaped excrescences upon different legumes are distinct species; but this view is not sustained by the experiments of Nobbe and Hiltner, and seems intrinsically improbable.

Mode of Infection.—The infection is especially effectual when applied to the seeds before sowing; and for that purpose there may be used either the turbid water made by stirring up in it some earth of a properly infected field, or else water charged with a pure culture of the appropriate kind, commercially known under the name of nitragin, now manufactured for the purpose. Or else, the field to be sown may be infected by spreading on it broadcast, and promptly harrowing in, a wagon-load of earth per acre from a properly infected field. Such earth must not be allowed to dry, or to be long exposed to light.

Specially effective (“virulent”) and hardy forms of such bacteria have been produced under artificial culture by Dr. Geo. T. Moore of the U.S. Department of Agriculture. These cultures can be sent by mail on cotton imbued with them, for the infection of seeds.

It is very important that the bacillus should be present in the earliest stages of the growth of the seedlings; otherwise the latter will undergo a longer or shorter period of starvation, unless the soil contains, or is furnished with, a sufficiency of available nitrogen to supply their immediate wants. When such a supply is very abundant, the legume crop will sometimes develop no nodules at all; but the best crops appear to be the result of a thorough infection, and abundant formation of the excrescences.

Cultural Results.—The marked results obtained in certain soils by inoculation with the legume-root bacillus are exemplified in the following table, showing results of experiments by J. F. Duggar, at the Alabama Experiment station.

TABLE SHOWING INCREASE OF PRODUCTION BY SOIL INOCULATION. ==============================+=======+========+================ PER ACRE. | TOPS. | ROOTS. | NITROGEN. | lbs. | lbs. | lbs. | Value. ------------------------------+-------+--------+-------+-------- Hairy vetch, not inoculated | 194 | 387 | 7 | $ 1.05 “ “ inoculated | 3045 | 1452 | 106 | 15.90 Crimson clover not inoculated | 106 | 266 | 4.3 | .65 “ “ inoculated | 4840 | 1452 | 143.7 | 21.25 ------------------------------+-------+--------+-------+--------

Bull. Ala. Exp’t Station, No. 96, 1898.

Such marked increases from soil inoculation cannot of course be expected in cases where the soil has previously borne leguminous crops of similar nature and therefore already contains the root bacteria. Hence Duggar found no increase of production when inoculating for cowpea, land that had borne that crop two years before and already contained the root bacteria. In the arid region, where the almost universally calcareous soils usually bear a natural growth largely composed of various leguminous plants, inoculation is likely to be less commonly effective than in the humid region east of the Mississippi, where leguminous plants are much less generally present in the native flora.

The distinctive agricultural function of supplying nitrogen to the soils on which they grow, renders inexcusable the persistence of some writers and teachers in designating all forage plants as “grasses.” Whatever excuse there may have been for this practice so long as the nitrogen-gathering function of the legumes was unknown, disappears with this discovery, and the misleading misnomer should be banished from agricultural publications and lectures, at the very least.

Other Nitrogen-Absorbing Bacteria.—An increase in the nitrogen-content of some soils, aside from the action of leguminous root-bacteria, has long been observed. As already stated, this increase was at first ascribed to certain green algæ often seen to develop on the soil surface; but it has now been shown that the nitrogen-gathering function belongs to at least two bacteria, one of which (Clostridium pastorianum) was discovered by Winogradski, the other (Azotobacter chroococcum) by Beyerinck, and has since been farther investigated by Koch, Kröber, Gerlach and Vogel, and last by Lipman and Hugo Fischer. According to the latter it seems likely that Azotobacter chroococcum lives in symbiosis with the green algæ, all of which, like the Azotobacter itself, develop with special luxuriance on calcareous soils.

Lipman (Rep. Agr. Exp’t Station, New Jersey, 1903 and 1904) describes as Azotobacter vinelandii a form somewhat different from the A. chroococcum, the nitrogen-assimilating power of which he tested quite elaborately. He exposed to air pure cultures of A. vinelandii in nutritive solution containing the proper mineral ingredients, and glucose 20 grams per liter. 100 cub. centimeters of this solution was exposed in flasks of respectively 250, 500 and 1000 cc. content, therefore having greater surface in the larger flasks. After ten days, the amounts of nitrogen fixed were found to be respectively 1.67, 3.19 and 7.90 milligrams. When mannite solution was employed instead of glucose, a similar fixation was observed; and it was also shown that the presence of combined nitrogen in the forms of nitrates or ammonium salts discouraged the fixation by the bacillus.

It was thus clearly proved that A. vinelandii at least does not need symbiosis with algæ to fix atmospheric nitrogen; but experiments with mixed cultures of the above bacillus and another (designated as No. 30 by Lipman) proved that when these two co-operate the absorption of atmospheric nitrogen is nearly doubled. As it is probable that this is the case also with other soil bacteria, the importance of this source of nitrogen to plants is obvious; provided of course that the proper nutritive ingredients are present in available form. Lipman shows that among the organic nutrients, besides the sugars, glycerine and the salts of propionic and lactic acids, and probably also others of the same groups, can serve as nourishment to the nitrogen-fixing bacteria.

DISTRIBUTION OF THE HUMUS WITHIN THE SURFACE SOIL.

The uniform distribution of the humus-contents of the surface soil, as shown in sections of the same, is by no means easily accounted for. The roots from which its substance is so largely derived are not so universally distributed as to account for it; but least of all can the rapid disappearance of the leaf-fall and other vegetable offal from the surface be accounted for without some outside agencies. Of these, the action of fungous vegetation, and of insects and earthworms, are doubtless the chief ones.

Fungi.—When we examine a decaying root, we find radiating from it a zone of deeper tint, as though from a colored solution penetrating outward. But since under normal conditions humus is insoluble, this explanation cannot stand. Microscopic examination, however, reveals that the outside limit of this zone is also the limit to which the fungous fibrils concerned in the process extend; and as these fibrils are much more finely distributed and much more numerous than the roots of any plant, it is natural that the humus resulting from their decomposition should be more evenly distributed than the roots themselves.

Kosticheff, Formation and Properties of Humus; in abstract Jour. Chem. Soc., 1891, p. 611.

Such fungous growth is not, however, confined to dead and decaying roots only. A large number of trees and shrubs, among them pines and firs, beeches, aspen and many others, also the heaths, and woody plants associated with them, appear to depend largely for their healthy development, notably in northern latitudes, upon the co-operation (“symbiosis”) of fungous fibrils that “infest” their roots, enabling them to assimilate, indirectly, the decaying organic (and inorganic) matter which would otherwise be unavailable, and at the same time converting that matter into their own substance. Fungous growths thus mediate both the decomposition and rehabilitation of the vegetable debris.

The vegetative fibrils (mycelia) of several kinds of molds are constantly present in the soil, and while consuming the dead tissue of the higher plants, spread their own substance throughout the soil mass. The same is true of the subterranean or “root” mycelia of the larger fungi, toadstools, mushrooms, which are commonly found about dead stumps and other deposits of decaying vegetable and animal offal. All these being dependent upon the presence of air for their life functions, remain within such distance from the surface as will afford adequate aeration; the depth reached depending upon the perviousness of the soil and subsoil. In the humid region this will usually be within a foot of the surface, but in the arid may reach to several feet. Ultimately these organisms contribute their substance to the store of humus in the land.

On the surface of moist soils we frequently find a copious growth of green fibrils, which may be either those of algæ, such as Oscillaria, or the early stages (prothallia) of moss vegetation. This vegetation has been credited with absorption of nitrogen from the air, thus enriching the soil; but later researches have shown this effect to be due to symbiotic bacteria (see above p. 156).

Animal Agencies.—Darwin first suggested that wherever the common earthworm (Lumbricus) finds the conditions of existence, it exerts a most important influence in the formation of the humous surface-soil layer; and the limitation imposed upon these conditions by the subsoil has doubtless a great deal to do with the sharp demarcation we often find between it and the surface soil. Briefly stated, the earthworm nourishes itself by swallowing, successively, portions of the surrounding earth, digesting a part of its organic matter and then ejecting the undigested earth in the form of “casts,” such as may be seen by thousands on the surface of the ground during or after a rain. Darwin (The Formation of Vegetable Mold, 1881), has calculated from actual observation that in humid climates and in a ground fairly stocked with these worms, the soil thus brought up may amount to from one-tenth to two-tenths of an inch annually over the entire surface; so that in half a century the entire surface foot might have been thus worked over. Aside from the mechanical effect thus achieved in loosening the soil, and the access of air and water permitted by their burrows, the chemical effects resulting from their digestive process, and the final return of their own substance to the soil mass; also their habit of drawing after themselves into their burrows leafstalks, blades of grass and other vegetable remains, renders their work of no mean importance both from the physical and chemical point of view. The uniformity, lack of structure and loose texture of the surface soil, especially of forests, as compared with subsoil layers of corresponding thickness, is doubtless largely due to the earthworms’ work. It has frequently been observed that when an unusual overflow has drowned out the earthworm population of a considerable area, the surface soil layer remains compacted, and vegetation languishes, until new immigration has restocked the soil with them. Again, the humus formed under their influence is always neutral, never acid.

Wollny (Forsch, Agr., 1890, p. 382), has shown by direct experimental cultures in boxes, with and without earthworms, surprising differences between the cultural results obtained, and this has been fully confirmed by the subsequent researches of Djemil (Ber. Physiol. Lab. Vers. Halle, 1898). In Wollny’s experiments, the ratio of higher production in the presence of the worms, varied all the way from 2.6 per cent in the case of oats, 93.9 in that of rye, 135.9 in that of potatoes, 300 in that of the field pea, and 140 in that of the vetch, to 733 per cent in the case of rape. Wollny attributes these favorable effects in the main to the increased looseness, and perviousness of the soil to air, and diminished water-holding power. Djemil’s results all point in the same direction; and he shows, moreover, that the allegation that the roots penetrate more deeply in the presence of the worms by following their burrows, is unfounded, the descending roots often passing close to and outside of these.

The work of earthworms is especially effective in loamy soils and in the humid regions. In the arid region, and in sandy soils generally, the life-conditions are unfavorable to the worm, and the perviousness elsewhere brought about by its labors already exists naturally in most cases. It is stated by E. T. Seton (Century Mag. for June, 1904) that the earthworm is practically non-existent in the arid region between the Rocky Mountains and the immediate Pacific coast, from Manitoba to Texas. In the Pacific coast region, however, they are abundant, and do their work effectually.

Insects of various kinds are also instrumental in producing, not only the uniform distribution of humus in the surface soil, but also the looseness of texture which we see in forest soils especially. Ants, wasps, many kinds of beetles, crickets, and particularly the larvæ of these, and of other burrowing creatures, often form considerable accumulations, due directly both to their mechanical activity, and to their excrements.

The work of ants is in some regions on so large a scale as to attract the attention of the most casual observer. Especially is this the case in portions of the arid region, from Texas to Montana, where at times large areas are so thickly studded with hills from three to twelve feet in diameter, and one to two feet high, that it is difficult to pass without being attacked by the insects. The “mounds” studding a large portion of the prairie country of Louisiana seem also to be due to the work of ants, although not inhabited at present.

Larger burrowing animals also assist in the task of mixing uniformly the surface soils, and aiding root-penetration, as well as, in many cases, the conservation of moisture. Seton (loc. cit.) even claims that the pocket gophers (Thomomys) in a great degree replace the activity of the earthworms in the arid region, where they, together with the voles (commonly known there as field mice), exist in great numbers. Of course the work of these animals, as well as that of the prairie dogs, ground squirrels, badgers, etc., is incompatible with cultivation. But the effects of their burrows on the native vegetation, and the indications they give of the nature of the subsoil, are eminently useful to the land-seeker.

Thus in the rolling sediment-lands of the Great Bend of the Columbia, the observer is surprised to see the “giant rye grass,” usually at home in the moist lowlands, growing preferably on the crests of the ridges bordering the horizon. Examination shows that this is due to the burrowing of badgers, whereby the roots of the grass are enabled to reach moisture at all times, even in that extremely arid region.