Fig. 189.—Transverse section through Branchiura sowerbyi. × 20. d.br, Dorsal branchia; i, intestine; n, nerve-cord; v.br, ventral branchia.
Nervous System.—The central nervous system of the Oligochaeta is very uniform in its structure in the entire group. The only family which is at all anomalous is that of the Aphaneura. In Aeolosoma there appears to be only a pair of cerebral ganglia, which retain the primitive position of these organs in being still in direct connexion with the epidermis. In all other Oligochaeta there are a pair of cerebral ganglia, connected by a circumoesophageal commissure with a ventral ganglionated cord. From the cerebral ganglia arises a system of nerve-fibres and nerve-cells, which represents the stomatogastric nerves of other Invertebrates.
Senses and Sense-Organs.—The only organs that can be regarded with anything like probability as sense-organs are the pigmented eyes of certain Naids and the tactile cells of many worms. The latter are usually elongated cells provided at their free extremity with a stiff process; they occur associated in groups, and often these bundles of cells have a segmental arrangement. The head end of many of the lower Oligochaeta, for instance the genus Aeolosoma, has delicate processes projecting here and there; these appear to be also of a tactile nature, and are of course connected with cells of the epidermis. The eyes of certain Naids are little more than lenticular bodies embedded in a mass of pigment. In the genus Eudrilus and in many Eudrilidae are peculiar integumental bodies, which were independently discovered by Dr. Horst[[411]] and myself, and compared by us to the Pacinian bodies of Mammals. Whether these structures are connected with nerves or not is doubtful. In spite of the poor development and the simplicity of their sense organs, the higher Oligochaeta at any rate can feel, and can distinguish light from darkness. Darwin[[412]] came to the conclusion that "light affects worms by its intensity and its duration." And furthermore, it is only the anterior end of the body which is thus affected. Of the sense of hearing these animals appear to be utterly devoid. Some kept by Darwin "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." But it is always necessary to discriminate between sound and vibrations passing through any solid body, which would appeal rather to a sense of touch. Here worms are most sensitive. It is quite easy, by digging with some vigour, to arouse the worms in the neighbourhood, who will crawl to the surface and away from the scene of action; a proceeding on their part which is sometimes put down to a desire to escape from their enemy the mole.
Smell appears to be another sense which is somewhat deficient. But worms are epicures, and exhibit a decided taste and preference for certain articles of diet. Like their fellow tiller of the soil, the agricultural labourer, worms have a keen relish for onions, which, however, they must recognise by the smell. They prefer green cabbage to red, celery to both, and raw meat appears to be the greatest delicacy that can be offered to them. It is only substances they are not likely to meet with, such as perfumes, tobacco, and paraffin, that produce no impression upon the worm's sense of smell.
Coelom and Vascular System.—When an earthworm is dissected the various organs are seen to lie in a fairly spacious cavity, which is interrupted and divided into a series of chambers by the mesenteries or septa which stretch across from wall to wall of the body, and correspond roughly in their position to the grooves which separate the body externally. This cavity, common to all the higher animals, is known as the coelom; it is lined by cells, which cover the intestines as well as the inside of the body-wall; and upon the intestine assume the form so characteristic of the group, namely, that of large yellow cells loaded with secreted matters, and called "chloragogen-cells" by Claparède. The coelom communicates with the exterior by means of the dorsal pores, the nephridia, and the ducts of the reproductive organs. As in all animals which possess a coelom, the reproductive tissues, ova and sperm, are developed on its walls.
Fig. 190.—Sparganophilus tamesis; general anatomy, × 3. (After Benham.) I-XVIII, segments. 1, 4, 6, Perivisceral vessels (6 is one of the hearts); 2, 3, 7, dorsal vessel; 5, spermatheca; 8, sperm sacs; 9, intestino-tegumentary vessels; 10, ovary; 11, 12, integumentary vessels.
The vascular system of the Oligochaeta forms a system of perfectly closed vessels, which ramify into fine capillary networks in the body-wall, in the coats of the alimentary canal, and upon the other organs of the body. The main trunks are a dorsal and a ventral longitudinal, which communicate directly in the anterior end of the body by large transverse contractile trunks, the so-called hearts (see Fig. 190, 6). The dorsal vessel is also contractile, but not the ventral, or, when it occurs, the subnervian. The vascular system has many degrees of complexity in different families; it is simpler in the smaller aquatic forms. The blood is usually red, and the pigment which is suspended in the plasma is haemoglobin. The blood is corpusculated.
Excretory Organs.—There appears to be a great deal more variation in the structure of the excretory system than there is in many other groups. For a long time only Lumbricus and a few of the aquatic genera were known as regards their excretory systems. In these there is a pair of excretory organs or nephridia in nearly all the segments. These are much coiled tubes, in which it is always possible to recognise three divisions. The nephridium commences with an orifice of a funnel-like character, fringed with long cilia, and opening into the body-cavity; from this springs a tube, which immediately perforates the septum lying between the segment which contains the funnel and the following one; this tube has the peculiarity first pointed out by Claparède of being excavated in the substance of cells; the glandular part of the nephridium is a row of cells which are bored through by a continuous canal, the walls of which are here and there furnished with cilia. It often happens that the main canal gives off minute lateral ramifications, which may even form a kind of network round the principal canal. The terminal section of the nephridium is a muscular sac which opens on to the exterior by a pore, and from which the products of excretion are from time to time evacuated by contractions of its walls. This is a brief statement of the main facts in the structure of those Oligochaeta in which there is a single pair of nephridia to each segment of the body; small differences of more or less importance occur. In Chaetogaster, for example, there is no trace of a funnel; in some genera the terminal sac is much reduced or unusually extended, being even sometimes provided with a caecum of moderate dimensions. In Acanthodrilus novae-zelandiae and a few other species the point of opening of the nephridia varies from segment to segment, though it always bears some relation to the chaetae. In these species the nephridia which open more dorsally are a little different in structure from those which open more ventrally. One set have a caecum, and the other have not.
The nephridia of the terrestrial forms are enveloped by a richly developed network of blood capillaries, which is absent in the smaller aquatic genera.