The Formation of Vegetable Mould Through the Action of Worms. Чарльз Дарвин
the secretion with which the leaves are moistened being alkaline, and from its acting both on the starch-granules and on the protoplasmic contents of the cells, we may infer that it resembles in nature not saliva, [40] but pancreatic secretion; and we know from Fredericq that a secretion of this kind is found in the intestines of worms. As the leaves which are dragged into the burrows are often dry and shrivelled, it is indispensable for their disintegration by the unarmed mouths of worms that they should first be moistened and softened; and fresh leaves, however soft and tender they may be, are similarly treated, probably from habit. The result is that they are partially digested before they are taken into the alimentary canal. I am not aware of any other case of extra-stomachal digestion having been recorded. The boa-constrictor is said to bathe its prey with saliva, but this is doubtful; and it is done solely for the sake of lubricating its prey. Perhaps the nearest analogy may be found in such plants as Drosera and Dionæa; for here animal matter is digested and converted into peptone not within a stomach, but on the surfaces of the leaves.
Calciferous Glands.—These glands (see Fig. 1), judging from their size and from their rich supply of blood-vessels, must be of much importance to the animal. But almost as many theories have been advanced on their use as there have been observers. They consist of three pairs, which in the common earth-worm debouch into the alimentary canal in advance of the gizzard, but posteriorly to it in Urochæta and some other genera. [41a] The two posterior pairs are formed by lamellæ, which, according to Claparède, are diverticula from the œsophagus. [41b] These lamellæ are coated with a pulpy cellular layer, with the outer cells lying free in infinite numbers. If one of these glands is punctured and squeezed, a quantity of white pulpy matter exudes, consisting of these free cells. They are minute, and vary in diameter from 2 to 6 μ. They contain in their centres a little excessively fine granular matter; but they look so like oil globules that Claparède and others at first treated them with ether. This produces no effect; but they are quickly dissolved with effervescence in acetic acid, and when oxalate of ammonia is added to the solution a white precipitate is thrown down. We may therefore conclude that they contain carbonate of lime. If the cells are immersed in a very little acid, they become more transparent, look like ghosts, and are soon lost to view; but if much acid is added, they disappear instantly. After a very large number have been dissolved, a flocculent residue is left, which apparently consists of the delicate ruptured cell-walls. In the two posterior pairs of glands the carbonate of lime contained in the cells occasionally aggregates into small rhombic crystals or into concretions, which lie between the lamellæ; but I have seen only one case, and Claparède only a very few such cases.
The two anterior glands differ a little in shape from the four posterior ones, by being more oval. They differ also conspicuously in generally containing several small, or two or three larger, or a single very large concretion of carbonate of lime, as much as 1½ mm. in diameter. When a gland includes only a few very small concretions, or, as sometimes happens, none at all, it is easily overlooked. The large concretions are round or oval, and exteriorly almost smooth. One was found which filled up not only the whole gland, as is often the case, but its neck; so that it resembled an olive-oil flask in shape. These concretions when broken are seen to be more or less crystalline in structure. How they escape from the gland is a marvel; but that they do escape is certain, for they are often found in the gizzard, intestines, and in the castings of worms, both with those kept in confinement and those in a state of nature.
Claparède says very little about the structure of the two anterior glands, and he supposes that the calcareous matter of which the concretions are formed is derived from the four posterior glands. But if an anterior gland which contains only small concretions is placed in acetic acid and afterwards dissected, or if sections are made of such a gland without being treated with acid, lamellæ like those in the posterior glands and coated with cellular matter could be plainly seen, together with a multitude of free calciferous cells readily soluble in acetic acid. When a gland is completely filled with a single large concretion, there are no free cells, as these have been all consumed in forming the concretion. But if such a concretion, or one of only moderately large size, is dissolved in acid, much membranous matter is left, which appears to consist of the remains of the formerly active lamellæ. After the formation and expulsion of a large concretion, new lamellæ must be developed in some manner. In one section made by my son, the process had apparently commenced, although the gland contained two rather large concretions, for near the walls several cylindrical and oval pipes were intersected, which were lined with cellular matter and were quite filled with free calciferous cells. A great enlargement in one direction of several oval pipes would give rise to the lamellæ.
Besides the free calciferous cells in which no nucleus was visible, other and rather larger free cells were seen on three occasions; and these contained a distinct nucleus and nucleolus. They were only so far acted on by acetic acid that the nucleus was thus rendered more distinct. A very small concretion was removed from between two of the lamellæ within an anterior gland. It was imbedded in pulpy cellular matter, with many free calciferous cells, together with a multitude of the larger, free, nucleated cells, and these latter cells were not acted on by acetic acid, while the former were dissolved. From this and other such cases I am led to suspect that the calciferous cells are developed from the larger nucleated ones; but how this was effected was not ascertained.
When an anterior gland contains several minute concretions, some of these are generally angular or crystalline in outline, while the greater number are rounded with an irregular mulberry-like surface. Calciferous cells adhered to many parts of these mulberry-like masses, and their gradual disappearance could be traced while they still remained attached. It was thus evident that the concretions are formed from the lime contained within the free calciferous cells. As the smaller concretions increase in size, they come into contact and unite, thus enclosing the now functionless lamellæ; and by such steps the formation of the largest concretions could be followed. Why the process regularly takes place in the two anterior glands, and only rarely in the four posterior glands, is quite unknown. Morren says that these glands disappear during the winter; and I have seen some instances of this fact, and others in which either the anterior or posterior glands were at this season so shrunk and empty, that they could be distinguished only with much difficulty.
With respect to the function of the calciferous glands, it is probable that they primarily serve as organs of excretion, and secondarily as an aid to digestion. Worms consume many fallen leaves; and it is known that lime goes on accumulating in leaves until they drop off the parent-plant, instead of being re-absorbed into the stem or roots, like various other organic and inorganic substances. [46] The ashes of a leaf of an acacia have been known to contain as much as 72 per cent. of lime. Worms therefore would be liable to become charged with this earth, unless there were some special means for its excretion; and the calciferous glands are well adapted for this purpose. The worms which live in mould close over the chalk, often have their intestines filled with this substance, and their castings are almost white. Here it is evident that the supply of calcareous matter must be super-abundant. Nevertheless with several worms collected on such a site, the calciferous glands contained as many free calciferous cells, and fully as many and large concretions, as did the glands of worms which lived where there was little or no lime; and this indicates that the lime is an excretion, and not a secretion poured into the alimentary canal for some special purpose.
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