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The Formation of Vegetable Mould through the action of worms with observations, a non-fiction book by Charles Darwin |
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Chapter 1. Habits Of Worms |
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_ CHAPTER I. HABITS OF WORMS 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.
Footnotes: {7} 'Bidrag till Skandinaviens Oligochaetfauna,' 1871. {8} 'Die bis jetzt bekannten Arten aus der Familie der Regenwurmer,' 1845. 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. {9} 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.
{9} There is even some reason to believe that pressure is actually favourable to the growth of grasses, for Professor Buckman, who made many observations on their growth in the experimental gardens of the Royal Agricultural College, remarks ('Gardeners' Chronicle,' 1854, p. 619): "Another circumstance in the cultivation of grasses in the separate form or small patches, is the impossibility of rolling or treading them firmly, without which no pasture can continue good." 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. Earth-worms must be considered as terrestrial animals, though they are still in one sense semi-aquatic, like the other members of the great class of annelids to which they belong. M. Perrier found that their exposure to the dry air of a room for only a single night was fatal to them. On the other hand he kept several large worms alive for nearly four months, completely submerged in water. {10} During the summer when the ground is dry, they penetrate to a considerable depth and cease to work, as they do during the winter when the ground is frozen. Worms are nocturnal in their habits, and at night may be seen crawling about in large numbers, but usually with their tails still inserted in their burrows. By the expansion of this part of their bodies, and with the help of the short, slightly reflexed bristles, with which their bodies are armed, they hold so fast that they can seldom be dragged out of the ground without being torn into pieces. {11} During the day they remain in their burrows, except at the pairing season, when those which inhabit adjoining burrows expose the greater part of their bodies for an hour or two in the early morning. Sick individuals, which are generally affected by the parasitic larvae of a fly, must also be excepted, as they wander about during the day and die on the surface. After heavy rain succeeding dry weather, an astonishing number of dead worms may sometimes be seen lying on the ground. Mr. Galton informs me that on one such occasion (March, 1881), the dead worms averaged one for every two and a half paces in length on a walk in Hyde Park, four paces in width. He counted no less than 45 dead worms in one place in a length of sixteen paces. From the facts above given, it is not probable that these worms could have been drowned, and if they had been drowned they would have perished in their burrows. I believe that they were already sick, and that their deaths were merely hastened by the ground being flooded.
{10} I shall have occasion often to refer to M. Perrier's admirable memoir, 'Organisation des Lombriciens terrestres' in 'Archives de Zoolog. exper.' tom. iii. 1874, p. 372. C. F. Morren ('De Lumbrici terrestris Hist. Nat.' 1829, p. 14) found that worms endured immersion for fifteen to twenty days in summer, but that in winter they died when thus treated. {11} Morren, 'De Lumbrici terrestris Hist. Nat.' &c., 1829, p. 67. 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 {12} 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.
{12} 'De Lumbrici terrestris Hist. Nat.' &c., p. 14.
{13} Histolog. Untersuchungen uber die Regenwurmer. 'Zeitschrift fur wissenschaft. Zoologie,' B. xix., 1869, p. 611. 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. Senses.--Worms are destitute of eyes, and at first I thought that they were quite insensible to light; for those kept in confinement were repeatedly observed by the aid of a candle, and others out of doors by the aid of a lantern, yet they were rarely alarmed, although extremely timid animals. Other persons have found no difficulty in observing worms at night by the same means. {14}
{14} For instance, Mr. Bridgman and Mr. Newman ('The Zoologist,' vol. vii. 1849, p. 2576), and some friends who observed worms for me.
{15} 'Familie der Regenwurmer,' 1845, p. 18.
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 (Tringa vanellus, Linn.) seems to know instinctively that worms will emerge if the ground is made to tremble; for Bishop Stanley states (as I hear from Mr. Moorhouse) 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 0.25 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. It may be presumed that all animals which feed on various substances possess the sense of taste, and this is certainly the case with worms. Cabbage-leaves are much liked by worms; and it appears that they can distinguish between different varieties; but this may perhaps be owing to differences in their texture. On eleven occasions pieces of the fresh leaves of a common green variety and of the red variety used for pickling were given them, and they preferred the green, the red being either wholly neglected or much less gnawed. On two other occasions, however, they seemed to prefer the red. Half-decayed leaves of the red variety and fresh leaves of the green were attacked about equally. When leaves of the cabbage, horse-radish (a favourite food) and of the onion were given together, the latter were always, and manifestly preferred. Leaves of the cabbage, lime-tree, Ampelopsis, parsnip (Pastinaca), and celery (Apium) were likewise given together; and those of the celery were first eaten. But when leaves of cabbage, turnip, beet, celery, wild cherry and carrots were given together, the two latter kinds, especially those of the carrot, were preferred to all the others, including those of celery. It was also manifest after many trials that wild cherry leaves were greatly preferred to those of the lime-tree and hazel (Corylus). According to Mr. Bridgman the half-decayed leaves of Phlox verna are particularly liked by worms. {16}
{16} 'The Zoologist,' vol. vii. 1849, p. 2576. 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. {17} 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 Perichaeta 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.
{17} 'Familie der Regenwurmer,' p. 13. Dr. Sturtevant states in the 'New York Weekly Tribune' (May 19, 1880) that he kept three worms in a pot, which was allowed to become extremely dry; and these worms were found "all entwined together, forming a round mass and in good condition." 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 {18} 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.
{18} 'De Lumbrici terrestris Hist. Nat.' p. 19. Leon Fredericq states {19} 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. {20} 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. {21}
{19} 'Archives de Zoologie experimentale,' tom. vii. 1878, p. 394. When I wrote the above passage, I was not aware that Krukenberg ('Untersuchungen a. d. physiol. Inst. d. Univ. Heidelberg,' Bd. ii. p. 37, 1877) had previously investigated the digestive juice of Lumbricus. He states that it contains a peptic, and diastatic, as well as a tryptic ferment. {20} On the action of the pancreatic ferment, see 'A Text-Book of Physiology,' by Michael Foster, 2nd edit. pp. 198-203. 1878. {21} Schmulewitsch, 'Action des Sucs digestifs sur la Cellulose.' Bull. de l'Acad. Imp. de St. Petersbourg, tom. xxv. p. 549. 1879. 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, {22} 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.
{22} Claparede doubts whether saliva is secreted by worms: see 'Zeitschrift fur wissenschaft. Zoologie,' B. xix. 1869, p. 601.
{23} Perrier, 'Archives de Zoolog. exper.' July, 1874, pp. 416, 419. {24} 'Zeitschrift fur wissenschaft. Zoologie,' B. xix, 1869, pp. 603-606.
Claparede 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. {25} 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.
{25} De Vries, 'Landwirth. Jahrbucher,' 1881, p. 77.
{26} M. Foster, 'A Text-Book of Physiology,' 2nd edit. 1878, p. 243.
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." {27} 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. Claparede 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.
{27} M. Foster, ut sup. p. 200. _ |
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