The Formation of Vegetable Mould Through the Action of Worms · Charles Darwin
CHAPTER I. HABITS OF WORMS.
The result was different when cabbage-leaves and pieces of onion were employed, both of which are devoured with much relish by worms. Small square pieces of fresh and half-decayed cabbage-leaves and of onion bulbs were on nine occasions buried in my pots, beneath about ¼ of an inch of common garden soil; and they were always discovered by the worms. One bit of cabbage was discovered and removed in the course of two hours; three were removed by the next morning, that is, after a single night; two others after two nights; and the seventh bit after three nights. Two pieces of onion were discovered and removed after three nights. Bits of fresh raw meat, of which worms are very fond, were buried, and were not discovered within forty-eight hours, during which time they had not become putrid. The earth above the various buried objects was generally pressed down only slightly, so as not to prevent the emission of any odour. On two occasions, however, the surface was well watered, and was thus rendered somewhat compact.
After the bits of cabbage and onion had been removed, I looked beneath them to see whether the worms had accidentally come up from below, but there was no sign of a burrow; and twice the buried objects were laid on pieces of tin-foil which were not in the least displaced. It is of course possible that the worms whilst moving about on the surface of the ground, with their tails affixed within their burrows, may have poked their heads into the places where the above objects were buried; but I have never seen worms acting in this manner. Some pieces of cabbage-leaf and of onion were twice buried beneath very fine ferruginous sand, which was slightly pressed down and well watered, so as to be rendered very compact, and these pieces were never discovered. On a third occasion the same kind of sand was neither pressed down nor watered, and the pieces of cabbage were discovered and removed after the second night. These several facts indicate that worms possess some power of smell; and that they discover by this means odoriferous and much-coveted kinds of food.
Pieces of the leaves of cabbage, turnip, horse-radish and onion were left on the pots during 22 days, and were all attacked and had to be renewed; but during the whole of this time leaves of an Artemisia and of the culinary sage, thyme and mint, mingled with the above leaves, were quite neglected excepting those of the mint, which were occasionally and very slightly nibbled. These latter four kinds of leaves do not differ in texture in a manner which could make them disagreeable to worms; they all have a strong taste, but so have the four first mentioned kinds of leaves; and the wide difference in the result must be attributed to a preference by the worms for one taste over another.
_Mental Qualities_.—There is little to be said on this head. We have seen that worms are timid. It may be doubted whether they suffer as much pain when injured, as they seem to express by their contortions. Judging by their eagerness for certain kinds of food, they must enjoy the pleasure of eating. Their sexual passion is strong enough to overcome for a time their dread of light. They perhaps have a trace of social feeling, for they are not disturbed by crawling over each other’s bodies, and they sometimes lie in contact. According to Hoffmeister they pass the winter either singly or rolled up with others into a ball at the bottom of their burrows. {32} Although worms are so remarkably deficient in the several sense-organs, this does not necessarily preclude intelligence, as we know from such cases as those of Laura Bridgman; and we have seen that when their attention is engaged, they neglect impressions to which they would otherwise have attended; and attention indicates the presence of a mind of some kind. They are also much more easily excited at certain times than at others. They perform a few actions instinctively, that is, all the individuals, including the young, perform such actions in nearly the same fashion. This is shown by the manner in which the species of Perichæta eject their castings, so as to construct towers; also by the manner in which the burrows of the common earth-worm are smoothly lined with fine earth and often with little stones, and the mouths of their burrows with leaves. One of their strongest instincts is the plugging up the mouths of their burrows with various objects; and very young worms act in this manner. But some degree of intelligence appears, as we shall see in the next chapter, to be exhibited in this work,—a result which has surprised me more than anything else in regard to worms.
_Food and Digestion_.—Worms are omnivorous. They swallow an enormous quantity of earth, out of which they extract any digestible matter which it may contain; but to this subject I must recur. They also consume a large number of half-decayed leaves of all kinds, excepting a few which have an unpleasant taste or are too tough for them; likewise petioles, peduncles, and decayed flowers. But they will also consume fresh leaves, as I have found by repeated trials. According to Morren {33} they will eat particles of sugar and liquorice; and the worms which I kept drew many bits of dry starch into their burrows, and a large bit had its angles well rounded by the fluid poured out of their mouths. But as they often drag particles of soft stone, such as of chalk, into their burrows, I feel some doubt whether the starch was used as food. Pieces of raw and roasted meat were fixed several times by long pins to the surface of the soil in my pots, and night after night the worms could be seen tugging at them, with the edges of the pieces engulfed in their mouths, so that much was consumed. Raw fat seems to be preferred even to raw meat or to any other substance which was given them, and much was consumed. They are cannibals, for the two halves of a dead worm placed in two of the pots were dragged into the burrows and gnawed; but as far as I could judge, they prefer fresh to putrid meat, and in so far I differ from Hoffmeister.
The half-decayed or fresh leaves which worms intend to devour, are dragged into the mouths of their burrows to a depth of from one to three inches, and are then moistened with a secreted fluid. It has been assumed that this fluid serves to hasten their decay; but a large number of leaves were twice pulled out of the burrows of worms and kept for many weeks in a very moist atmosphere under a bell-glass in my study; and the parts which had been moistened by the worms did not decay more quickly in any plain manner than the other parts. When fresh leaves were given in the evening to worms kept in confinement and examined early on the next morning, therefore not many hours after they had been dragged into the burrows, the fluid with which they were moistened, when tested with neutral litmus paper, showed an alkaline reaction. This was repeatedly found to be the case with celery, cabbage and turnip leaves. Parts of the same leaves which had not been moistened by the worms, were pounded with a few drops of distilled water, and the juice thus extracted was not alkaline. Some leaves, however, which had been drawn into burrows out of doors, at an unknown antecedent period, were tried, and though still moist, they rarely exhibited even a trace of alkaline reaction.
The fluid, with which the leaves are bathed, acts on them whilst they are fresh or nearly fresh, in a remarkable manner; for it quickly kills and discolours them. Thus the ends of a fresh carrot-leaf, which had been dragged into a burrow, were found after twelve hours of a dark brown tint. Leaves of celery, turnip, maple, elm, lime, thin leaves of ivy, and, occasionally those of the cabbage were similarly acted on. The end of a leaf of _Triticum repens_, still attached to a growing plant, had been drawn into a burrow, and this part was dark brown and dead, whilst the rest of the leaf was fresh and green. Several leaves of lime and elm removed from burrows out of doors were found affected in different degrees. The first change appears to be that the veins become of a dull reddish-orange. The cells with chlorophyll next lose more or less completely their green colour, and their contents finally become brown. The parts thus affected often appeared almost black by reflected light; but when viewed as a transparent object under the microscope, minute specks of light were transmitted, and this was not the case with the unaffected parts of the same leaves. These effects, however, merely show that the secreted fluid is highly injurious or poisonous to leaves; for nearly the same effects were produced in from one to two days on various kinds of young leaves, not only by artificial pancreatic fluid, prepared with or without thymol, but quickly by a solution of thymol by itself. On one occasion leaves of Corylus were much discoloured by being kept for eighteen hours in pancreatic fluid, without any thymol. With young and tender leaves immersion in human saliva during rather warm weather, acted in the same manner as the pancreatic fluid, but not so quickly. The leaves in all these cases often became infiltrated with the fluid.
Large leaves from an ivy plant growing on a wall were so tough that they could not be gnawed by worms, but after four days they were affected in a peculiar manner by the secretion poured out of their mouths. The upper surfaces of the leaves, over which the worms had crawled, as was shown by the dirt left on them, were marked in sinuous lines, by either a continuous or broken chain of whitish and often star-shaped dots, about 2 mm. in diameter. The appearance thus presented was curiously like that of a leaf, into which the larva of some minute insect had burrowed. But my son Francis, after making and examining sections, could nowhere find that the cell-walls had been broken down or that the epidermis had been penetrated. When the section passed through the whitish dots, the grains of chlorophyll were seen to be more or less discoloured, and some of the palisade and mesophyll cells contained nothing but broken down granular matter. These effects must be attributed to the transudation of the secretion through the epidermis into the cells.
The secretion with which worms moisten leaves likewise acts on the starch-granules within the cells. My son examined some leaves of the ash and many of the lime, which had fallen off the trees and had been partly dragged into worm-burrows. It is known that with fallen leaves the starch-grains are preserved in the guard-cells of the stomata. Now in several cases the starch had partially or wholly disappeared from these cells, in the parts which had been moistened by the secretion; while it was still well preserved in the other parts of the same leaves. Sometimes the starch was dissolved out of only one of the two guard-cells. The nucleus in one case had disappeared, together with the starch-granules. The mere burying of lime-leaves in damp earth for nine days did not cause the destruction of the starch-granules. On the other hand, the immersion of fresh lime and cherry leaves for eighteen hours in artificial pancreatic fluid, led to the dissolution of the starch-granules in the guard-cells as well as in the other cells.
From the secretion with which the leaves are moistened being alkaline, and from its acting both on the starch-granules and on the protoplasmic contents of the cells, we may infer that it resembles in nature not saliva, {40} but pancreatic secretion; and we know from Fredericq that a secretion of this kind is found in the intestines of worms. As the leaves which are dragged into the burrows are often dry and shrivelled, it is indispensable for their disintegration by the unarmed mouths of worms that they should first be moistened and softened; and fresh leaves, however soft and tender they may be, are similarly treated, probably from habit. The result is that they are partially digested before they are taken into the alimentary canal. I am not aware of any other case of extra-stomachal digestion having been recorded. The boa-constrictor is said to bathe its prey with saliva, but this is doubtful; and it is done solely for the sake of lubricating its prey. Perhaps the nearest analogy may be found in such plants as Drosera and Dionæa; for here animal matter is digested and converted into peptone not within a stomach, but on the surfaces of the leaves.
_Calciferous Glands_.—These glands (see Fig. 1), judging from their size and from their rich supply of blood-vessels, must be of much importance to the animal. But almost as many theories have been advanced on their use as there have been observers. They consist of three pairs, which in the common earth-worm debouch into the alimentary canal in advance of the gizzard, but posteriorly to it in Urochæta and some other genera. {41a} The two posterior pairs are formed by lamellæ, which, according to Claparède, are diverticula from the œsophagus. {41b} These lamellæ are coated with a pulpy cellular layer, with the outer cells lying free in infinite numbers. If one of these glands is punctured and squeezed, a quantity of white pulpy matter exudes, consisting of these free cells. They are minute, and vary in diameter from 2 to 6 _μ_. They contain in their centres a little excessively fine granular matter; but they look so like oil globules that Claparède and others at first treated them with ether. This produces no effect; but they are quickly dissolved with effervescence in acetic acid, and when oxalate of ammonia is added to the solution a white precipitate is thrown down. We may therefore conclude that they contain carbonate of lime. If the cells are immersed in a very little acid, they become more transparent, look like ghosts, and are soon lost to view; but if much acid is added, they disappear instantly. After a very large number have been dissolved, a flocculent residue is left, which apparently consists of the delicate ruptured cell-walls. In the two posterior pairs of glands the carbonate of lime contained in the cells occasionally aggregates into small rhombic crystals or into concretions, which lie between the lamellæ; but I have seen only one case, and Claparède only a very few such cases.
The two anterior glands differ a little in shape from the four posterior ones, by being more oval. They differ also conspicuously in generally containing several small, or two or three larger, or a single very large concretion of carbonate of lime, as much as 1½ mm. in diameter. When a gland includes only a few very small concretions, or, as sometimes happens, none at all, it is easily overlooked. The large concretions are round or oval, and exteriorly almost smooth. One was found which filled up not only the whole gland, as is often the case, but its neck; so that it resembled an olive-oil flask in shape. These concretions when broken are seen to be more or less crystalline in structure. How they escape from the gland is a marvel; but that they do escape is certain, for they are often found in the gizzard, intestines, and in the castings of worms, both with those kept in confinement and those in a state of nature.
Claparède says very little about the structure of the two anterior glands, and he supposes that the calcareous matter of which the concretions are formed is derived from the four posterior glands. But if an anterior gland which contains only small concretions is placed in acetic acid and afterwards dissected, or if sections are made of such a gland without being treated with acid, lamellæ like those in the posterior glands and coated with cellular matter could be plainly seen, together with a multitude of free calciferous cells readily soluble in acetic acid. When a gland is completely filled with a single large concretion, there are no free cells, as these have been all consumed in forming the concretion. But if such a concretion, or one of only moderately large size, is dissolved in acid, much membranous matter is left, which appears to consist of the remains of the formerly active lamellæ. After the formation and expulsion of a large concretion, new lamellæ must be developed in some manner. In one section made by my son, the process had apparently commenced, although the gland contained two rather large concretions, for near the walls several cylindrical and oval pipes were intersected, which were lined with cellular matter and were quite filled with free calciferous cells. A great enlargement in one direction of several oval pipes would give rise to the lamellæ.
Besides the free calciferous cells in which no nucleus was visible, other and rather larger free cells were seen on three occasions; and these contained a distinct nucleus and nucleolus. They were only so far acted on by acetic acid that the nucleus was thus rendered more distinct. A very small concretion was removed from between two of the lamellæ within an anterior gland. It was imbedded in pulpy cellular matter, with many free calciferous cells, together with a multitude of the larger, free, nucleated cells, and these latter cells were not acted on by acetic acid, while the former were dissolved. From this and other such cases I am led to suspect that the calciferous cells are developed from the larger nucleated ones; but how this was effected was not ascertained. When an anterior gland contains several minute concretions, some of these are generally angular or crystalline in outline, while the greater number are rounded with an irregular mulberry-like surface. Calciferous cells adhered to many parts of these mulberry-like masses, and their gradual disappearance could be traced while they still remained attached. It was thus evident that the concretions are formed from the lime contained within the free calciferous cells. As the smaller concretions increase in size, they come into contact and unite, thus enclosing the now functionless lamellæ; and by such steps the formation of the largest concretions could be followed. Why the process regularly takes place in the two anterior glands, and only rarely in the four posterior glands, is quite unknown. Morren says that these glands disappear during the winter; and I have seen some instances of this fact, and others in which either the anterior or posterior glands were at this season so shrunk and empty, that they could be distinguished only with much difficulty.
With respect to the function of the calciferous glands, it is probable that they primarily serve as organs of excretion, and secondarily as an aid to digestion. Worms consume many fallen leaves; and it is known that lime goes on accumulating in leaves until they drop off the parent-plant, instead of being re-absorbed into the stem or roots, like various other organic and inorganic substances. {46} The ashes of a leaf of an acacia have been known to contain as much as 72 per cent. of lime. Worms therefore would be liable to become charged with this earth, unless there were some special means for its excretion; and the calciferous glands are well adapted for this purpose. The worms which live in mould close over the chalk, often have their intestines filled with this substance, and their castings are almost white. Here it is evident that the supply of calcareous matter must be super-abundant. Nevertheless with several worms collected on such a site, the calciferous glands contained as many free calciferous cells, and fully as many and large concretions, as did the glands of worms which lived where there was little or no lime; and this indicates that the lime is an excretion, and not a secretion poured into the alimentary canal for some special purpose.
With worms not only the contents of the intestines, but their ejected matter or the castings, are generally acid. Thirty castings from different places were tested, and with three or four exceptions were found to be acid; and the exceptions may have been due to such castings not having been recently ejected; for some which were at first acid, were on the following morning, after being dried and again moistened, no longer acid; and this probably resulted from the humus acids being, as is known to be the case, easily decomposed. Five fresh castings from worms which lived in mould close over the chalk, were of a whitish colour and abounded with calcareous matter; and these were not in the least acid. This shows how effectually carbonate of lime neutralises the intestinal acids. When worms were kept in pots filled with fine ferruginous sand, it was manifest that the oxide of iron, with which the grains of silex were coated, had been dissolved and removed from them in the castings.
The digestive fluid of worms resembles in its action, as already stated, the pancreatic secretion of the higher animals; and in these latter, “pancreatic digestion is essentially alkaline; the action will not take place unless some alkali be present; and the activity of an alkaline juice is arrested by acidification, and hindered by neutralization.” {50} Therefore it seems highly probable that the innumerable calciferous cells, which are poured from the four posterior glands into the alimentary canal of worms, serve to neutralise more or less completely the acids there generated by the half-decayed leaves. We have seen that these cells are instantly dissolved by a small quantity of acetic acid, and as they do not always suffice to neutralise the contents of even the upper part of the alimentary canal, the lime is perhaps aggregated into concretions in the anterior pair of glands, in order that some may be carried down to the posterior parts of the intestine, where these concretions would be rolled about amongst the acid contents. The concretions found in the intestines and in the castings often have a worn appearance, but whether this is due to some amount of attrition or of chemical corrosion could not be told. Claparède believes that they are formed for the sake of acting as mill-stones, and of thus aiding in the trituration of the food. They may give some aid in this way; but I fully agree with Perrier that this must be of quite subordinate importance, seeing that the object is already attained by stones being generally present in the gizzards and intestines of worms.
Text from Project Gutenberg, public domain in the United States.