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THE FORMATION OF VEGETABLE MOULD THROUGH THE ACTION OF WORMS TEXT

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middle; and 23 per cent. by the basal part. If they had been drawn 19953 characters on this page
middle; and 23 per cent. by the basal part. If they had been drawn indifferently by any point, the proportion for the apical, middle and basal parts would have been 33.3 per cent. for each; but, as we have just seen, it might have been expected that a much larger proportion would have been drawn in by the basal than by any other part. As the case stands, nearly three times as many were drawn in by the apex as by the base. If we consider the broad triangles by themselves, 59 per cent. were drawn in by the apex, 25 per cent. by the middle, and 16 per cent. by the base. Of the narrow triangles, 65 per cent. were drawn in by the apex, 14 per cent, by the middle, and 21 per cent. by the base; so that here those drawn in by the apex were more than 3 times as many as those drawn in by the base. We may therefore conclude that the manner in which the triangles are drawn into the burrows is not a matter of chance. In eight cases, two triangles had been drawn into the same burrow, and in seven of these cases, one had been drawn in by the apex and the other by the base. This again indicates that the result is not determined by chance. Worms appear sometimes to revolve in the act of drawing in the triangles, for five out of the whole lot had been wound into an irregular spire round the inside of the burrow. Worms kept in a warm room drew 63 triangles into their burrows; but, as in the case of the pine-leaves, they worked in a rather careless manner, for only 44 per cent. were drawn in by the apex, 22 per cent. by the middle, and 33 per cent. by the base. In five cases, two triangles were drawn into the same burrow. It may be suggested with much apparent probability that so large a proportion of the triangles were drawn in by the apex, not from the worms having selected this end as the most convenient for the purpose, but from having first tried in other ways and failed. This notion was countenanced by the manner in which worms in confinement were seen to drag about and drop the triangles; but then they were working carelessly. I did not at first perceive the importance of this subject, but merely noticed that the bases of those triangles which had been drawn in by the apex, were generally clean and not crumpled. The subject was afterwards attended to carefully. In the first place several triangles which had been drawn in by the basal angles, or by the base, or a little above the base, and which were thus much crumpled and dirtied, were left for some hours in water and were then well shaken while immersed; but neither the dirt nor the creases were thus removed. Only slight creases could be obliterated, even by pulling the wet triangles several times through my fingers. Owing to the slime from the worms’ bodies, the dirt was not easily washed off. We may therefore conclude that if a triangle, before being dragged in by the apex, had been dragged into a burrow by its base with even a slight degree of force, the basal part would long retain its creases and remain dirty. The condition of 89 triangles (65 narrow and 24 broad ones), which had been drawn in by the apex, was observed; and the bases of only 7 of them were at all creased, being at the same time generally dirty. Of the 82 uncreased triangles, 14 were dirty at the base; but it does not follow from this fact that these had first been dragged towards the burrows by their bases; for the worms sometimes covered large portions of the triangles with slime, and these when dragged by the apex over the ground would be dirtied; and during rainy weather, the triangles were often dirtied over one whole side or over both sides. If the worms had dragged the triangles to the mouths of their burrows by their bases, as often as by their apices, and had then perceived, without actually trying to draw them into the burrow, that the broader end was not well adapted for this purpose—even in this case a large proportion would probably have had their basal ends dirtied. We may therefore infer—improbable as is the inference—that worms are able by some means to judge which is the best end by which to draw triangles of paper into their burrows. The percentage results of the foregoing observations on the manner in which worms draw various kinds of objects into the mouths of their burrows may be abridged as follows:— Nature of Drawn into the Drawn in, by or Drawn in, by or Object. burrows, by or near the near the base. near the apex. middle. Leaves of 80 11 9 various kinds —of the Lime, 79 17 4 basal margin of blade broad, apex acuminated —of a Laburnum, 63 10 27 basal part of blade as narrow as, or sometimes little narrower than the apical part —of the 34 ... 66 Rhododendron, basal part of blade often narrower than the apical part —of Pine-trees, ... ... 100 consisting of two needles arising from a common base Petioles of a 76 ... 24 Clematis, somewhat pointed at the apex, and blunt at the base —of the Ash, 48.5 ... 51.5 the thick basal end often drawn in to serve as food —of Robinia, 44 ... 56 extremely thin, especially towards the apex, so as to be ill-fitted for plugging up the burrows Triangles of 62 15 23 paper, of the two sizes —of the broad 59 25 16 ones alone —of the narrow 65 14 21 ones alone If we consider these several cases, we can hardly escape from the conclusion that worms show some degree of intelligence in their manner of plugging up their burrows. Each particular object is seized in too uniform a manner, and from causes which we can generally understand, for the result to be attributed to mere chance. That every object has not been drawn in by its pointed end, may be accounted for by labour having been saved through some being inserted by their broader or thicker ends. No doubt worms are led by instinct to plug up their burrows; and it might have been expected that they would have been led by instinct how best to act in each particular case, independently of intelligence. We see how difficult it is to judge whether intelligence comes into play, for even plants might sometimes be thought to be thus directed; for instance when displaced leaves re-direct their upper surfaces towards the light by extremely complicated movements and by the shortest course. With animals, actions appearing due to intelligence may be performed through inherited habit without any intelligence, although aboriginally thus acquired. Or the habit may have been acquired through the preservation and inheritance of beneficial variations of some other habit; and in this case the new habit will have been acquired independently of intelligence throughout the whole course of its development. There is no _à priori_ improbability in worms having acquired special instincts through either of these two latter means. Nevertheless it is incredible that instincts should have been developed in reference to objects, such as the leaves of petioles of foreign plants, wholly unknown to the progenitors of the worms which act in the described manner. Nor are their actions so unvarying or inevitable as are most true instincts. As worms are not guided by special instincts in each particular case, though possessing a general instinct to plug up their burrows, and as chance is excluded, the next most probable conclusion seems to be that they try in many different ways to draw in objects, and at last succeed in some one way. But it is surprising that an animal so low in the scale as a worm should have the capacity for acting in this manner, as many higher animals have no such capacity. For instance, ants may be seen vainly trying to drag an object transversely to their course, which could be easily drawn longitudinally; though after a time they generally act in a wiser manner, M. Fabre states {89a} that a Sphex—an insect belonging to the same highly-endowed order with ants—stocks its nest with paralysed grass-hoppers, which are invariably dragged into the burrow by their antennæ. When these were cut off close to the head, the Sphex seized the palpi; but when these were likewise cut off, the attempt to drag its prey into the burrow was given up in despair. The Sphex had not intelligence enough to seize one of the six legs or the ovipositor of the grasshopper, which, as M. Fabre remarks, would have served equally well. So again, if the paralysed prey with an egg attached to it be taken out of the cell, the Sphex after entering and finding the cell empty, nevertheless closes it up in the usual elaborate manner. Bees will try to escape and go on buzzing for hours on a window, one half of which has been left open. Even a pike continued during three months to dash and bruise itself against the glass sides of an aquarium, in the vain attempt to seize minnows on the opposite side. {89b} A cobra-snake was seen by Mr. Layard {90} to act much more wisely than either the pike or the Sphex; it had swallowed a toad lying within a hole, and could not withdraw its head; the toad was disgorged, and began to crawl away; it was again swallowed and again disgorged; and now the snake had learnt by experience, for it seized the toad by one of its legs and drew it out of the hole. The instincts of even the higher animals are often followed in a senseless or purposeless manner: the weaver-bird will perseveringly wind threads through the bars of its cage, as if building a nest: a squirrel will pat nuts on a wooden floor, as if he had just buried them in the ground: a beaver will cut up logs of wood and drag them about, though there is no water to dam up; and so in many other cases. Mr. Romanes, who has specially studied the minds of animals, believes that we can safely infer intelligence, only when we see an individual profiting by its own experience. By this test the cobra showed some intelligence; but this would have been much plainer if on a second occasion he had drawn a toad out of a hole by its leg. The Sphex failed signally in this respect. Now if worms try to drag objects into their burrows first in one way and then in another, until they at last succeed, they profit, at least in each particular instance, by experience. But evidence has been advanced showing that worms do not habitually try to draw objects into their burrows in many different ways. Thus half-decayed lime-leaves from their flexibility could have been drawn in by their middle or basal parts, and were thus drawn into the burrows in considerable numbers; yet a large majority were drawn in by or near the apex. The petioles of the Clematis could certainly have been drawn in with equal ease by the base and apex; yet three times and in certain cases five times as many were drawn in by the apex as by the base. It might have been thought that the foot-stalks of leaves would have tempted the worms as a convenient handle; yet they are not largely used, except when the base of the blade is narrower than the apex. A large number of the petioles of the ash are drawn in by the base; but this part serves the worms as food. In the case of pine-leaves worms plainly show that they at least do not seize the leaf by chance; but their choice does not appear to be determined by the divergence of the two needles, and the consequent advantage or necessity of drawing them into their burrows by the base. With respect to the triangles of paper, those which had been drawn in by the apex rarely had their bases creased or dirty; and this shows that the worms had not often first tried to drag them in by this end. If worms are able to judge, either before drawing or after having drawn an object close to the mouths of their burrows, how best to drag it in, they must acquire some notion of its general shape. This they probably acquire by touching it in many places with the anterior extremity of their bodies, which serves as a tactile organ. It may be well to remember how perfect the sense of touch becomes in a man when born blind and deaf, as are worms. If worms have the power of acquiring some notion, however rude, of the shape of an object and of their burrows, as seems to be the case, they deserve to be called intelligent; for they then act in nearly the same manner as would a man under similar circumstances. To sum up, as chance does not determine the manner in which objects are drawn into the burrows, and as the existence of specialized instincts for each particular case cannot be admitted, the first and most natural supposition is that worms try all methods until they at last succeed; but many appearances are opposed to such a supposition. One alternative alone is left, namely, that worms, although standing low in the scale of organization, possess some degree of intelligence. This will strike every one as very improbable; but it may be doubted whether we know enough about the nervous system of the lower animals to justify our natural distrust of such a conclusion. With respect to the small size of the cerebral ganglia, we should remember what a mass of inherited knowledge, with some power of adapting means to an end, is crowded into the minute brain of a worker-ant. _Means by which worms excavate their burrows_.—This is effected in two ways; by pushing away the earth on all sides, and by swallowing it. In the former case, the worm inserts the stretched out and attenuated anterior extremity of its body into any little crevice, or hole; and then, as Perrier remarks, {93} the pharynx is pushed forwards into this part, which consequently swells and pushes away the earth on all sides. The anterior extremity thus serves as a wedge. It also serves, as we have before seen, for prehension and suction, and as a tactile organ. A worm was placed on loose mould, and it buried itself in between two and three minutes. On another occasion four worms disappeared in 15 minutes between the sides of the pot and the earth, which had been moderately pressed down. On a third occasion three large worms and a small one were placed on loose mould well mixed with fine sand and firmly pressed down, and they all disappeared, except the tail of one, in 35 minutes. On a fourth occasion six large worms were placed on argillaceous mud mixed with sand firmly pressed down, and they disappeared, except the extreme tips of the tails of two of them, in 40 minutes. In none of these cases, did the worms swallow, as far as could be seen, any earth. They generally entered the ground close to the sides of the pot. A pot was next filled with very fine ferruginous sand, which was pressed down, well watered, and thus rendered extremely compact. A large worm left on the surface did not succeed in penetrating it for some hours, and did not bury itself completely until 25 hrs. 40 min. had elapsed. This was effected by the sand being swallowed, as was evident by the large quantity ejected from the vent, long before the whole body had disappeared. Castings of a similar nature continued to be ejected from the burrow during the whole of the following day. As doubts have been expressed by some writers whether worms ever swallow earth solely for the sake of making their burrows, some additional cases may be given. A mass of fine reddish sand, 23 inches in thickness, left on the ground for nearly two years, had been penetrated in many places by worms; and their castings consisted partly of the reddish sand and partly of black earth brought up from beneath the mass. This sand had been dug up from a considerable depth, and was of so poor a nature that weeds could not grow on it. It is therefore highly improbable that it should have been swallowed by the worms as food. Again in a field near my house the castings frequently consist of almost pure chalk, which lies at only a little depth beneath the surface; and here again it is very improbable that the chalk should have been swallowed for the sake of the very little organic matter which could have percolated into it from the poor overlying pasture. Lastly, a casting thrown up through the concrete and decayed mortar between the tiles, with which the now ruined aisle of Beaulieu Abbey had formerly been paved, was washed, so that the coarser matter alone was left. This consisted of grains of quartz, micaceous slate, other rocks, and bricks or tiles, many of them from 1/20 to 1/10 inch in diameter. No one will suppose that these grains were swallowed as food, yet they formed more than half of the casting, for they weighed 19 grains, the whole casting having weighed 33 grains. Whenever a worm burrows to a depth of some feet in undisturbed compact ground, it must form its passage by swallowing the earth; for it is incredible that the ground could yield on all sides to the pressure of the pharynx when pushed forwards within the worm’s body. That worms swallow a larger quantity of earth for the sake of extracting any nutritious matter which it may contain than for making their burrows, appears to me certain. But as this old belief has been doubted by so high an authority as Claparède, evidence in its favour must be given in some detail. There is no _à priori_ improbability in such a belief, for besides other annelids, especially the _Arenicola marina_, which throws up such a profusion of castings on our tidal sands, and which it is believed thus subsists, there are animals belonging to the most distinct classes, which do not burrow, but habitually swallow large quantities of sand; namely, the molluscan Onchidium and many Echinoderms. {97} If earth were swallowed only when worms deepened their burrows or made new ones, castings would be thrown up only occasionally; but in many places fresh castings may be seen every morning, and the amount of earth ejected from the same burrow on successive days is large. Yet worms do not burrow to a great depth, except when the weather is very dry or intensely cold. On my lawn the black vegetable mould or humus is only about 5 inches in thickness, and overlies light-coloured or reddish clayey soil: now when castings are thrown up in the greatest profusion, only a small proportion are light coloured, and it is incredible that the worms should daily make fresh burrows in every direction in the thin superficial layer of dark-coloured mould, unless they obtained nutriment of some kind from it. I have observed a strictly analogous case in a field near my house where bright red clay lay close beneath the surface. Again on one part of the Downs near Winchester the vegetable mould overlying the chalk was found to be only from 3 to 4 inches in thickness; and the many castings here ejected were as black as ink and did not effervesce with acids; so that the worms must have confined themselves to this thin superficial layer of mould, of which large quantities were daily swallowed. In another place at no great distance the castings were white; and why the worms should have burrowed into the chalk in some places and not in others, I am unable to conjecture. Two great piles of leaves had been left to decay in my grounds, and months after their removal, the bare surface, several yards in diameter, was so thickly covered during several months with castings that they formed an almost continuous layer; and the large number of worms which lived here must have subsisted during these months on nutritious matter contained in the black earth. The lowest layer from another pile of decayed leaves mixed with some
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PUBLIC-DOMAIN BOOK The Formation of Vegetable Mould Through the Action of Worms Charles Darwinprimary
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