The Formation of Vegetable Mould Through the Action of Worms. Чарльз Дарвин

The body of a large worm consists of from 100 to 200 almost cylindrical rings or segments, each furnished with minute bristles. The muscular system is well developed. Worms can crawl backwards as well as forwards, and by the aid of their affixed tails can retreat with extraordinary rapidity into their burrows. The mouth is situated at the anterior end of the body, and is provided with a little projection (lobe or lip, as it has been variously called) which is used for prehension. Internally, behind the mouth, there is a strong pharynx, shown in the accompanying diagram (Fig. 1) which is pushed forwards when the animal eats, and this part corresponds, according to Perrier, with the protrudable trunk or proboscis of other annelids. The pharynx leads into the œsophagus, on each side of which in the lower part there are three pairs of large glands, which secrete a surprising amount of carbonate of lime. These calciferous glands are highly remarkable, for nothing like them is known in any other animal. Their use will be discussed when we treat of the digestive process. In most of the species, the œsophagus is enlarged into a crop in front of the gizzard. This latter organ is lined with a smooth thick chitinous membrane, and is surrounded by weak longitudinal, but powerful transverse muscles. Perrier saw these muscles in energetic action; and, as he remarks, the trituration of the food must be chiefly effected by this organ, for worms possess no jaws or teeth of any kind. Grains of sand and small stones, from the ½0 to a little more than the 1/10 inch in diameter, may generally be found in their gizzards and intestines. As it is certain that worms swallow many little stones, independently of those swallowed while excavating their burrows, it is probable that they serve, like mill-stones, to triturate their food. The gizzard opens into the intestine, which runs in a straight course to the vent at the posterior end of the body. The intestine presents a remarkable structure, the typhlosolis, or, as the old anatomists called it, an intestine within an intestine; and Claparède [17] has shown that this consists of a deep longitudinal involution of the walls of the intestine, by which means an extensive absorbent surface is gained.

      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. [18a]

      Hoffmeister, however, states [18b] 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