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INTRODUCTION.

THE share which worms have taken in the formation of the layer of vegetable mould, which covers the whole surface of the land in every moderately humid country, is the subject of the present volume. This mould is generally of a blackish colour and a few inches in thickness. In different districts it differs but little in appearance, although it may rest on various subsoils. The uniform fineness of the particles of which it is composed is one of its chief characteristic features; and this may be well observed in any gravelly country, where a recently-ploughed field immediately adjoins one which has long remained undisturbed for pasture, and where the vegetable mould is exposed on the sides of a ditch or hole. The subject may appear an insignificant one, but we shall see that it possesses some interest; and the maxim "de minimis non curat lex," does not apply to science. Even ?lie de Beaumont, who generally undervalues small agencies and their accumulated effects, remarks: "La couche tr?s-mince de la terre v?g?tale est un monument d'une haute antiquit?, et, par le fait de sa permanence, un objet digne d'occuper le g?ologue, et capable de lui fournir des remarques int?ressantes." Although the superficial layer of vegetable mould as a whole no doubt is of the highest antiquity, yet in regard to its permanence, we shall hereafter see reason to believe that its component particles are in most cases removed at not a very slow rate, and are replaced by others due to the disintegration of the underlying materials.

As I was led to keep in my study during many months worms in pots filled with earth, I became interested in them, and wished to learn how far they acted consciously, and how much mental power they displayed. I was the more desirous to learn something on this head, as few observations of this kind have been made, as far as I know, on animals so low in the scale of organization and so poorly provided with sense-organs, as are earth-worms.

In the year 1837, a short paper was read by me before the Geological Society of London, "On the Formation of Mould," in which it was shown that small fragments of burnt marl, cinders, &c., which had been thickly strewed over the surface of several meadows, were found after a few years lying at the depth of some inches beneath the turf, but still forming a layer. This apparent sinking of superficial bodies is due, as was first suggested to me by Mr. Wedgwood of Maer Hall in Staffordshire, to the large quantity of fine earth continually brought up to the surface by worms in the form of castings. These castings are sooner or later spread out and cover up any object left on the surface. I was thus led to conclude that all the vegetable mould over the whole country has passed many times through, and will again pass many times through, the intestinal canals of worms. Hence the term "animal mould" would be in some respects more appropriate than that commonly used of "vegetable mould."

Ten years after the publication of my paper, M. D'Archiac, evidently influenced by the doctrines of ?lie de Beaumont, wrote about my "singuli?re th?orie," and objected that it could apply only to "les prairies basses et humides;" and that "les terres labour?es, les bois, les prairies ?lev?es, n'apportent aucune preuve ? l'appui de cette mani?re de voir." But M. D'Archiac must have thus argued from inner consciousness and not from observation, for worms abound to an extraordinary degree in kitchen gardens where the soil is continually worked, though in such loose soil they generally deposit their castings in any open cavities or within their old burrows instead of on the surface. Hensen estimates that there are about twice as many worms in gardens as in corn-fields. With respect to "prairies ?lev?es," I do not know how it may be in France, but nowhere in England have I seen the ground so thickly covered with castings as on commons, at a height of several hundred feet above the sea. In woods again, if the loose leaves in autumn are removed, the whole surface will be found strewed with castings. Dr. King, the superintendent of the Botanic Garden in Calcutta, to whose kindness I am indebted for many observations on earth-worms, informs me that he found, near Nancy in France, the bottom of the State forests covered over many acres with a spongy layer, composed of dead leaves and innumerable worm-castings. He there heard the Professor of "Am?nagement des For?ts" lecturing to his pupils, and pointing out this case as a "beautiful example of the natural cultivation of the soil; for year after year the thrown-up castings cover the dead leaves; the result being a rich humus of great thickness."

In the year 1869, Mr. Fish rejected my conclusions with respect to the part which worms have played in the formation of vegetable mould, merely on account of their assumed incapacity to do so much work. He remarks that "considering their weakness and their size, the work they are represented to have accomplished is stupendous." Here we have an instance of that inability to sum up the effects of a continually recurrent cause, which has often retarded the progress of science, as formerly in the case of geology, and more recently in that of the principle of evolution.

Although these several objections seemed to me to have no weight, yet I resolved to make more observations of the same kind as those published, and to attack the problem on another side; namely, to weigh all the castings thrown up within a given time in a measured space, instead of ascertaining the rate at which objects left on the surface were buried by worms. But some of my observations have been rendered almost superfluous by an admirable paper by Hensen, already alluded to, which appeared in 1877. Before entering on details with respect to the castings, it will be advisable to give some account of the habits of worms from my own observations and from those of other naturalists.

Nature of the sites inhabited--Can live long under water--Nocturnal--Wander about at night--Often lie close to the mouths of their burrows, and are thus destroyed in large numbers by birds--Structure--Do not possess eyes, but can distinguish between light and darkness--Retreat rapidly when brightly illuminated, not by a reflex action--Power of attention--Sensitive to heat and cold--Completely deaf--Sensitive to vibrations and to touch--Feeble power of smell--Taste--Mental qualities--Nature of food--Omnivorous--Digestion--Leaves before being swallowed, moistened with a fluid of the nature of the pancreatic secretion--Extra-stomachal digestion--Calciferous glands, structure of--Calcareous concretions formed in the anterior pair of glands--The calcareous matter primarily an excretion, but secondarily serves to neutralise the acids generated during the digestive process.

EARTH-WORMS are distributed throughout the world under the form of a few genera, which externally are closely similar to one another. The British species of Lumbricus have never been carefully monographed; but we may judge of their probable number from those inhabiting neighbouring countries. In Scandinavia there are eight species, according to Eisen; but two of these rarely burrow in the ground, and one inhabits very wet places or even lives under the water. We are here concerned only with the kinds which bring up earth to the surface in the form of castings. Hoffmeister says that the species in Germany are not well known, but gives the same number as Eisen, together with some strongly marked varieties.

Earth-worms abound in England in many different stations. Their castings may be seen in extraordinary numbers on commons and chalk-downs, so as almost to cover the whole surface, where the soil is poor and the grass short and thin. But they are almost or quite as numerous in some of the London parks, where the grass grows well and the soil appears rich. Even on the same field worms are much more frequent in some places than in others, without any visible difference in the nature of the soil. They abound in paved court-yards close to houses; and an instance will be given in which they had burrowed through the floor of a very damp cellar. I have seen worms in black peat in a boggy field; but they are extremely rare, or quite absent in the drier, brown, fibrous peat, which is so much valued by gardeners. On dry, sandy or gravelly tracks, where heath with some gorse, ferns, coarse grass, moss and lichens alone grow, hardly any worms can be found. But in many parts of England, wherever a path crosses a heath, its surface becomes covered with a fine short sward. Whether this change of vegetation is due to the taller plants being killed by the occasional trampling of man and animals, or to the soil being occasionally manured by the droppings from animals, I do not know. On such grassy paths worm-castings may often be seen. On a heath in Surrey, which was carefully examined, there were only a few castings on these paths, where they were much inclined; but on the more level parts, where a bed of fine earth had been washed down from the steeper parts and had accumulated to a thickness of a few inches, worm-castings abounded. These spots seemed to be overstocked with worms, so that they had been compelled to spread to a distance of a few feet from the grassy paths, and here their castings had been thrown up among the heath; but beyond this limit, not a single casting could be found. A layer, though a thin one, of fine earth, which probably long retains some moisture, is in all cases, as I believe, necessary for their existence; and the mere compression of the soil appears to be in some degree favourable to them, for they often abound in old gravel walks, and in foot-paths across fields.

Beneath large trees few castings can be found during certain seasons of the year, and this is apparently due to the moisture having been sucked out of the ground by the innumerable roots of the trees; for such places may be seen covered with castings after the heavy autumnal rains. Although most coppices and woods support many worms, yet in a forest of tall and ancient beech-trees in Knole Park, where the ground beneath was bare of all vegetation, not a single casting could be found over wide spaces, even during the autumn. Nevertheless, castings were abundant on some grass-covered glades and indentations which penetrated this forest. On the mountains of North Wales and on the Alps, worms, as I have been informed, are in most places rare; and this may perhaps be due to the close proximity of the subjacent rocks, into which worms cannot burrow during the winter so as to escape being frozen. Dr. McIntosh, however, found worm-castings at a height of 1500 feet on Schiehallion in Scotland. They are numerous on some hills near Turin at from 2000 to 3000 feet above the sea, and at a great altitude on the Nilgiri Mountains in South India and on the Himalaya.

It has often been said that under ordinary circumstances healthy worms never, or very rarely, completely leave their burrows at night; but this is an error, as White of Selborne long ago knew. In the morning, after there has been heavy rain, the film of mud or of very fine sand over gravel-walks is often plainly marked with their tracks. I have noticed this from August to May, both months included, and it probably occurs during the two remaining months of the year when they are wet. On these occasions, very few dead worms could anywhere be seen. On January 31, 1881, after a long-continued and unusually severe frost with much snow, as soon as a thaw set in, the walks were marked with innumerable tracks. On one occasion, five tracks were counted crossing a space of only an inch square. They could sometimes be traced either to or from the mouths of the burrows in the gravel-walks, for distances between 2 or 3 up to 15 yards. I have never seen two tracks leading to the same burrow; nor is it likely, from what we shall presently see of their sense-organs, that a worm could find its way back to its burrow after having once left it. They apparently leave their burrows on a voyage of discovery, and thus they find new sites to inhabit.

Morren states that worms often lie for hours almost motionless close beneath the mouths of their burrows. I have occasionally noticed the same fact with worms kept in pots in the house; so that by looking down into their burrows, their heads could just be seen. If the ejected earth or rubbish over the burrows be suddenly removed, the end of the worm's body may very often be seen rapidly retreating. This habit of lying near the surface leads to their destruction to an immense extent. Every morning during certain seasons of the year, the thrushes and blackbirds on all the lawns throughout the country draw out of their holes an astonishing number of worms, and this they could not do, unless they lay close to the surface. It is not probable that worms behave in this manner for the sake of breathing fresh air, for we have seen that they can live for a long time under water. I believe that they lie near the surface for the sake of warmth, especially in the morning; and we shall hereafter find that they often coat the mouths of their burrows with leaves, apparently to prevent their bodies from coming into close contact with the cold damp earth. It is said that they completely close their burrows during the winter.

The circulatory system is well developed. Worms breathe by their skin, as they do not possess any special respiratory organs. The two sexes are united in the same individual, but two individuals pair together. The nervous system is fairly well developed; and the two almost confluent cerebral ganglia are situated very near to the anterior end of the body.

Hoffmeister, however, states that worms, with the exception of a few individuals, are extremely sensitive to light; but he admits that in most cases a certain time is requisite for its action. These statements led me to watch on many successive nights worms kept in pots, which were protected from currents of air by means of glass plates. The pots were approached very gently, in order that no vibration of the floor should be caused. When under these circumstances worms were illuminated by a bull's-eye lantern having slides of dark red and blue glass, which intercepted so much light that they could be seen only with some difficulty, they were not at all affected by this amount of light, however long they were exposed to it. The light, as far as I could judge, was brighter than that from the full moon. Its colour apparently made no difference in the result. When they were illuminated by a candle, or even by a bright paraffin lamp, they were not usually affected at first. Nor were they when the light was alternately admitted and shut off. Sometimes, however, they behaved very differently, for as soon as the light fell on them, they withdrew into their burrows with almost instantaneous rapidity. This occurred perhaps once out of a dozen times. When they did not withdraw instantly, they often raised the anterior tapering ends of their bodies from the ground, as if their attention was aroused or as if surprise was felt; or they moved their bodies from side to side as if feeling for some object. They appeared distressed by the light; but I doubt whether this was really the case, for on two occasions after withdrawing slowly, they remained for a long time with their anterior extremities protruding a little from the mouths of their burrows, in which position they were ready for instant and complete withdrawal.

When the light from a candle was concentrated by means of a large lens on the anterior extremity, they generally withdrew instantly; but this concentrated light failed to act perhaps once out of half a dozen trials. The light was on one occasion concentrated on a worm lying beneath water in a saucer, and it instantly withdrew into its burrow. In all cases the duration of the light, unless extremely feeble, made a great difference in the result; for worms left exposed before a paraffin lamp or a candle invariably retreated into their burrows within from five to fifteen minutes; and if in the evening the pots were illuminated before the worms had come out of their burrows, they failed to appear.

From the foregoing facts it is evident that light affects worms by its intensity and by its duration. It is only the anterior extremity of the body, where the cerebral ganglia lie, which is affected by light, as Hoffmeister asserts, and as I observed on many occasions. If this part is shaded, other parts of the body may be fully illuminated, and no effect will be produced. As these animals have no eyes, we must suppose that the light passes through their skins, and in some manner excites their cerebral ganglia. It appeared at first probable that the different manner in which they were affected on different occasions might be explained, either by the degree of extension of their skin and its consequent transparency, or by some particular incident of the light; but I could discover no such relation. One thing was manifest, namely, that when worms were employed in dragging leaves into their burrows or in eating them, and even during the short intervals whilst they rested from their work, they either did not perceive the light or were regardless of it; and this occurred even when the light was concentrated on them through a large lens. So, again, whilst they are paired, they will remain for an hour or two out of their burrows, fully exposed to the morning light; but it appears from what Hoffmeister says that a light will occasionally cause paired individuals to separate.

When a worm is suddenly illuminated and dashes like a rabbit into its burrow--to use the expression employed by a friend--we are at first led to look at the action as a reflex one. The irritation of the cerebral ganglia appears to cause certain muscles to contract in an inevitable manner, independently of the will or consciousness of the animal, as if it were an automaton. But the different effect which a light produced on different occasions, and especially the fact that a worm when in any way employed and in the intervals of such employment, whatever set of muscles and ganglia may then have been brought into play, is often regardless of light, are opposed to the view of the sudden withdrawal being a simple reflex action. With the higher animals, when close attention to some object leads to the disregard of the impressions which other objects must be producing on them, we attribute this to their attention being then absorbed; and attention implies the presence of a mind. Every sportsman knows that he can approach animals whilst they are grazing, fighting or courting, much more easily than at other times. The state, also, of the nervous system of the higher animals differs much at different times, for instance, a horse is much more readily startled at one time than at another. The comparison here implied between the actions of one of the higher animals and of one so low in the scale as an earth-worm, may appear far-fetched; for we thus attribute to the worm attention and some mental power, nevertheless I can see no reason to doubt the justice of the comparison.

Although worms cannot be said to possess the power of vision, their sensitiveness to light enables them to distinguish between day and night; and they thus escape extreme danger from the many diurnal animals which prey on them. Their withdrawal into their burrows during the day appears, however, to have become an habitual action; for worms kept in pots covered by glass plates, over which sheets of black paper were spread, and placed before a north-east window, remained during the day-time in their burrows and came out every night; and they continued thus to act for a week. No doubt a little light may have entered between the sheets of glass and the blackened paper; but we know from the trials with coloured glass, that worms are indifferent to a small amount of light.

Worms appear to be less sensitive to moderate radiant heat than to a bright light. I judge of this from having held at different times a poker heated to dull redness near some worms, at a distance which caused a very sensible degree of warmth in my hand. One of them took no notice; a second withdrew into its burrow, but not quickly; the third and fourth much more quickly, and the fifth as quickly as possible. The light from a candle, concentrated by a lens and passing through a sheet of glass which would intercept most of the heat-rays, generally caused a much more rapid retreat than did the heated poker. Worms are sensitive to a low temperature, as may be inferred from their not coming out of their burrows during a frost.

Worms do not possess any sense of hearing. They took not the least notice of the shrill notes from a metal whistle, which was repeatedly sounded near them; nor did they of the deepest and loudest tones of a bassoon. They were indifferent to shouts, if care was taken that the breath did not strike them. When placed on a table close to the keys of a piano, which was played as loudly as possible, they remained perfectly quiet.

Although they are indifferent to undulations in the air audible by us, they are extremely sensitive to vibrations in any solid object. When the pots containing two worms which had remained quite indifferent to the sound of the piano, were placed on this instrument, and the note C in the bass clef was struck, both instantly retreated into their burrows. After a time they emerged, and when G above the line in the treble clef was struck they again retreated. Under similar circumstances on another night one worm dashed into its burrow on a very high note being struck only once, and the other worm when C in the treble clef was struck. On these occasions the worms were not touching the sides of the pots, which stood in saucers; so that the vibrations, before reaching their bodies, had to pass from the sounding board of the piano, through the saucer, the bottom of the pot and the damp, not very compact earth on which they lay with their tails in their burrows. They often showed their sensitiveness when the pot in which they lived, or the table on which the pot stood, was accidentally and lightly struck; but they appeared less sensitive to such jars than to the vibrations of the piano; and their sensitiveness to jars varied much at different times.

It has often been said that if the ground is beaten or otherwise made to tremble, worms believe that they are pursued by a mole and leave their burrows. From one account that I have received, I have no doubt that this is often the case; but a gentleman informs me that he lately saw eight or ten worms leave their burrows and crawl about the grass on some boggy land on which two men had just trampled while setting a trap; and this occurred in a part of Ireland where there were no moles. I have been assured by a Volunteer that he has often seen many large earth-worms crawling quickly about the grass, a few minutes after his company had fired a volley with blank cartridges. The Peewit seems to know instinctively that worms will emerge if the ground is made to tremble; for Bishop Stanley states that a young peewit kept in confinement used to stand on one leg and beat the turf with the other leg until the worms crawled out of their burrows, when they were instantly devoured. Nevertheless, worms do not invariably leave their burrows when the ground is made to tremble, as I know by having beaten it with a spade, but perhaps it was beaten too violently.

The whole body of a worm is sensitive to contact. A slight puff of air from the mouth causes an instant retreat. The glass plates placed over the pots did not fit closely, and blowing through the very narrow chinks thus left, often sufficed to cause a rapid retreat. They sometimes perceived the eddies in the air caused by quickly removing the glass plates. When a worm first comes out of its burrow, it generally moves the much extended anterior extremity of its body from side to side in all directions, apparently as an organ of touch; and there is some reason to believe, as we shall see in the next chapter, that they are thus enabled to gain a general notion of the form of an object. Of all their senses that of touch, including in this term the perception of a vibration, seems much the most highly developed.

In worms the sense of smell apparently is confined to the perception of certain odours, and is feeble. They were quite indifferent to my breath, as long as I breathed on them very gently. This was tried, because it appeared possible that they might thus be warned of the approach of an enemy. They exhibited the same indifference to my breath whilst I chewed some tobacco, and while a pellet of cotton-wool with a few drops of millefleurs perfume or of acetic acid was kept in my mouth. Pellets of cotton-wool soaked in tobacco juice, in millefleurs perfume, and in paraffin, were held with pincers and were waved about within two or three inches of several worms, but they took no notice. On one or two occasions, however, when acetic acid had been placed on the pellets, the worms appeared a little uneasy, and this was probably due to the irritation of their skins. The perception of such unnatural odours would be of no service to worms; and as such timid creatures would almost certainly exhibit some signs of any new impression, we may conclude that they did not perceive these odours.

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 1/4 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.

L?on Fredericq states that the digestive fluid of worms is of the same nature as the pancreatic secretion of the higher animals; and this conclusion agrees perfectly with the kinds of food which worms consume. Pancreatic juice emulsifies fat, and we have just seen how greedily worms devour fat; it dissolves fibrin, and worms eat raw meat; it converts starch into grape-sugar with wonderful rapidity, and we shall presently show that the digestive fluid of worms acts on starch. But they live chiefly on half-decayed leaves; and these would be useless to them unless they could digest the cellulose forming the cell-walls; for it is well known that all other nutritious substances are almost completely withdrawn from leaves, shortly before they fall off. It has, however, now been ascertained that some forms of cellulose, though very little or not at all attacked by the gastric secretion of the higher animals, are acted on by that from the pancreas.

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.

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, 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 Dionaea; for here animal matter is digested and converted into peptone not within a stomach, but on the surfaces of the leaves.

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 1 1/2 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.

Clapar?de 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, lamellae 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 lamellae. After the formation and expulsion of a large concretion, new lamellae 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 lamellae.

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 lamellae 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 lamellae; 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. 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.

On the other hand, the following considerations render it highly probable that the carbonate of lime, which is excreted by the glands, aids the digestive process under ordinary circumstances. Leaves during their decay generate an abundance of various kinds of acids, which have been grouped together under the term of humus acids. We shall have to recur to this subject in our fifth chapter, and I need here only say that these acids act strongly on carbonate of lime. The half-decayed leaves which are swallowed in such large quantities by worms would, therefore, after they have been moistened and triturated in the alimentary canal, be apt to produce such acids. And in the case of several worms, the contents of the alimentary canal were found to be plainly acid, as shown by litmus paper. This acidity cannot be attributed to the nature of the digestive fluid, for pancreatic fluid is alkaline; and we have seen that the secretion which is poured out of the mouths of worms for the sake of preparing the leaves for consumption, is likewise alkaline. The acidity can hardly be due to uric acid, as the contents of the upper part of the intestine were often acid. In one case the contents of the gizzard were slightly acid, those of the upper intestines being more plainly acid. In another case the contents of the pharynx were not acid, those of the gizzard doubtfully so, while those of the intestine were distinctly acid at a distance of 5 cm. below the gizzard. Even with the higher herbivorous and omnivorous animals, the contents of the large intestine are acid. "This, however, is not caused by any acid secretion from the mucous membrane; the reaction of the intestinal walls in the larger as in the small intestine is alkaline. It must therefore arise from acid fermentations going on in the contents themselves . . . In Carnivora the contents of the coecum are said to be alkaline, and naturally the amount of fermentation will depend largely on the nature of the food."

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." 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. Clapar?de 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.

Manner in which worms seize objects--Their power of suction--The instinct of plugging up the mouths of their burrows--Stones piled over the burrows--The advantages thus gained--Intelligence shown by worms in their manner of plugging up their burrows--Various kinds of leaves and other objects thus used--Triangles of paper--Summary of reasons for believing that worms exhibit some intelligence--Means by which they excavate their burrows, by pushing away the earth and swallowing it--Earth also swallowed for the nutritious matter which it contains--Depth to which worms burrow, and the construction of their burrows--Burrows lined with castings, and in the upper part with leaves--The lowest part paved with little stones or seeds--Manner in which the castings are ejected--The collapse of old burrows--Distribution of worms--Tower-like castings in Bengal--Gigantic castings on the Nilgiri Mountains--Castings ejected in all countries.

IN the pots in which worms were kept, leaves were pinned down to the soil, and at night the manner in which they were seized could be observed. The worms always endeavoured to drag the leaves towards their burrows; and they tore or sucked off small fragments, whenever the leaves were sufficiently tender. They generally seized the thin edge of a leaf with their mouths, between the projecting upper and lower lip; the thick and strong pharynx being at the same time, as Perrier remarks, pushed forward within their bodies, so as to afford a point of resistance for the upper lip. In the case of broad flat objects they acted in a wholly different manner. The pointed anterior extremity of the body, after being brought into contact with an object of this kind, was drawn within the adjoining rings, so that it appeared truncated and became as thick as the rest of the body. This part could then be seen to swell a little; and this, I believe, is due to the pharynx being pushed a little forwards. Then by a slight withdrawal of the pharynx or by its expansion, a vacuum was produced beneath the truncated slimy end of the body whilst in contact with the object; and by this means the two adhered firmly together. That under these circumstances a vacuum was produced was plainly seen on one occasion, when a large worm lying beneath a flaccid cabbage leaf tried to drag it away; for the surface of the leaf directly over the end of the worm's body became deeply pitted. On another occasion a worm suddenly lost its hold on a flat leaf; and the anterior end of the body was momentarily seen to be cup-formed. Worms can attach themselves to an object beneath water in the same manner; and I saw one thus dragging away a submerged slice of an onion-bulb.

The edges of fresh or nearly fresh leaves affixed to the ground were often nibbled by the worms; and sometimes the epidermis and all the parenchyma on one side was gnawed completely away over a considerable space; the epidermis alone on the opposite side being left quite clean. The veins were never touched, and leaves were thus sometimes partly converted into skeletons. As worms have no teeth and as their mouths consist of very soft tissue, it may be presumed that they consume by means of suction the edges and the parenchyma of fresh leaves, after they have been softened by the digestive fluid. They cannot attack such strong leaves as those of sea-kale or large and thick leaves of ivy; though one of the latter after it had become rotten was reduced in parts to the state of a skeleton.

Worms seize leaves and other objects, not only to serve as food, but for plugging up the mouths of their burrows; and this is one of their strongest instincts. They sometimes work so energetically that Mr. D. F. Simpson, who has a small walled garden where worms abound in Bayswater, informs me that on a calm damp evening he there heard so extraordinary a rustling noise from under a tree from which many leaves had fallen, that he went out with a light and discovered that the noise was caused by many worms dragging the dry leaves and squeezing them into the burrows. Not only leaves, but petioles of many kinds, some flower-peduncles, often decayed twigs of trees, bits of paper, feathers, tufts of wool and horse-hairs are dragged into their burrows for this purpose. I have seen as many as seventeen petioles of a Clematis projecting from the mouth of one burrow, and ten from the mouth of another. Some of these objects, such as the petioles just named, feathers, &c., are never gnawed by worms. In a gravel-walk in my garden I found many hundred leaves of a pine-tree drawn by their bases into burrows. The surfaces by which these leaves are articulated to the branches are shaped in as peculiar a manner as is the joint between the leg-bones of a quadruped; and if these surfaces had been in the least gnawed, the fact would have been immediately visible, but there was no trace of gnawing. Of ordinary dicotyledonous leaves, all those which are dragged into burrows are not gnawed. I have seen as many as nine leaves of the lime-tree drawn into the same burrow, and not nearly all of them had been gnawed; but such leaves may serve as a store for future consumption. Where fallen leaves are abundant, many more are sometimes collected over the mouth of a burrow than can be used, so that a small pile of unused leaves is left like a roof over those which have been partly dragged in.

A leaf in being dragged a little way into a cylindrical burrow is necessarily much folded or crumpled. When another leaf is drawn in, this is done exteriorly to the first one, and so on with the succeeding leaves; and finally all become closely folded and pressed together. Sometimes the worm enlarges the mouth of its burrow, or makes a fresh one close by, so as to draw in a still larger number of leaves. They often or generally fill up the interstices between the drawn-in leaves with moist viscid earth ejected from their bodies; and thus the mouths of the burrows are securely plugged. Hundreds of such plugged burrows may be seen in many places, especially during the autumnal and early winter months. But, as will hereafter be shown, leaves are dragged into the burrows not only for plugging them up and for food, but for the sake of lining the upper part or mouth.

When worms cannot obtain leaves, petioles, sticks, &c., with which to plug up the mouths of their burrows, they often protect them by little heaps of stones; and such heaps of smooth rounded pebbles may frequently be seen on gravel-walks. Here there can be no question about food. A lady, who was interested in the habits of worms, removed the little heaps of stones from the mouths of several burrows and cleared the surface of the ground for some inches all round. She went out on the following night with a lantern, and saw the worms with their tails fixed in their burrows, dragging the stones inwards by the aid of their mouths, no doubt by suction. "After two nights some of the holes had 8 or 9 small stones over them; after four nights one had about 30, and another 34 stones." One stone--which had been dragged over the gravel-walk to the mouth of a burrow weighed two ounces; and this proves how strong worms are. But they show greater strength in sometimes displacing stones in a well-trodden gravel-walk; that they do so, may be inferred from the cavities left by the displaced stones being exactly filled by those lying over the mouths of adjoining burrows, as I have myself observed.

Work of this kind is usually performed during the night; but I have occasionally known objects to be drawn into the burrows during the day. What advantage the worms derive from plugging up the mouths of their burrows with leaves, &c., or from piling stones over them, is doubtful. They do not act in this manner at the times when they eject much earth from their burrows; for their castings then serve to cover the mouths. When gardeners wish to kill worms on a lawn, it is necessary first to brush or rake away the castings from the surface, in order that the lime-water may enter the burrows. It might be inferred from this fact that the mouths are plugged up with leaves, &c., to prevent the entrance of water during heavy rain; but it may be urged against this view that a few, loose, well-rounded stones are ill-adapted to keep out water. I have moreover seen many burrows in the perpendicularly cut turf-edgings to gravel-walks, into which water could hardly flow, as well plugged as burrows on a level surface. It is not probable that the plugs or piles of stones serve to conceal the burrows from scolopendras, which, according to Hoffmeister, are the bitterest enemies of worms, or from the larger species of Carabus and Staphylinus which attack them ferociously, for these animals are nocturnal, and the burrows are opened at night. May not worms when the mouth of the burrow is protected be able to remain with safety with their heads close to it, which we know that they like to do, but which costs so many of them their lives? Or may not the plugs check the free ingress of the lowest stratum of air, when chilled by radiation at night, from the surrounding ground and herbage? I am inclined to believe in this latter view: firstly, because when worms were kept in pots in a room with a fire, in which case cold air could not enter the burrows, they plugged them up in a slovenly manner; and secondarily, because they often coat the upper part of their burrows with leaves, apparently to prevent their bodies from coming into close contact with the cold damp earth. Mr. E. Parfitt has suggested to me that the mouths of the burrows are closed in order that the air within them may be kept thoroughly damp, and this seems the most probable explanation of the habit. But the plugging-up process may serve for all the above purposes.

Whatever the motive may be, it appears that worms much dislike leaving the mouths of their burrows open. Nevertheless they will reopen them at night, whether or not they can afterwards close them. Numerous open burrows may be seen on recently-dug ground, for in this case the worms eject their castings in cavities left in the ground, or in the old burrows instead of piling them over the mouths of their burrows, and they cannot collect objects on the surface by which the mouths might be protected. So again on a recently disinterred pavement of a Roman villa at Abinger the worms pertinaciously opened their burrows almost every night, when these had been closed by being trampled on, although they were rarely able to find a few minute stones wherewith to protect them.

In the first place 227 withered leaves of various kinds, mostly of English plants, were pulled out of worm-burrows in several places. Of these, 181 had been drawn into the burrows by or near their tips, so that the foot-stalk projected nearly upright from the mouth of the burrow; 20 had been drawn in by their bases, and in this case the tips projected from the burrows; and 26 had been seized near the middle, so that these had been drawn in transversely and were much crumpled. Therefore 80 per cent. had been drawn in by the tip, 9 per cent. by the base or foot-stalk, and 11 per cent. transversely or by the middle. This alone is almost sufficient to show that chance does not determine the manner in which leaves are dragged into the burrows.

Of the above 227 leaves, 70 consisted of the fallen leaves of the common lime-tree, which is almost certainly not a native of England. These leaves are much acuminated towards the tip, and are very broad at the base with a well-developed foot-stalk. They are thin and quite flexible when half-withered. Of the 70, 79 per cent. had been drawn in by or near the tip; 4 per cent. by or near the base; and 17 per cent. transversely or by the middle. These proportions agree very closely, as far as the tip is concerned, with those before given. But the percentage drawn in by the base is smaller, which may be attributed to the breadth of the basal part of the blade. We here, also, see that the presence of a foot-stalk, which it might have been expected would have tempted the worms as a convenient handle, has little or no influence in determining the manner in which lime leaves are dragged into the burrows. The considerable proportion, viz., 17 per cent., drawn in more or less transversely depends no doubt on the flexibility of these half-decayed leaves. The fact of so many having been drawn in by the middle, and of some few having been drawn in by the base, renders it improbable that the worms first tried to draw in most of the leaves by one or both of these methods, and that they afterwards drew in 79 per cent. by their tips; for it is clear that they would not have failed in drawing them in by the base or middle.

The leaves of a foreign plant were next searched for, the blades of which were not more pointed towards the apex than towards the base. This proved to be the case with those of a laburnum for on doubling the terminal over the basal half, they generally fitted exactly; and when there was any difference, the basal half was a little the narrower. It might, therefore, have been expected that an almost equal number of these leaves would have been drawn in by the tip and base, or a slight excess in favour of the latter. But of 73 leaves pulled out of worm-burrows, 63 per cent. had been drawn in by the tip; 27 per cent. by the base, and 10 per cent. transversely. We here see that a far larger proportion, viz., 27 per cent. were drawn in by the base than in the case of lime leaves, the blades of which are very broad at the base, and of which only 4 per cent. had thus been drawn in. We may perhaps account for the fact of a still larger proportion of the laburnum leaves not having been drawn in by the base, by worms having acquired the habit of generally drawing in leaves by their tips and thus avoiding the foot-stalk. For the basal margin of the blade in many kinds of leaves forms a large angle with the foot-stalk; and if such a leaf were drawn in by the foot-stalk, the basal margin would come abruptly into contact with the ground on each side of the burrow, and would render the drawing in of the leaf very difficult.

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