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 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.” [49]
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.” [50] 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. Claparède 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.
CHAPTER II.
HABITS OF WORMS—continued.
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.