Fig. 145.—Gizzard of Scaphander lignarius L.: A, showing position with regard to oesophagus (oe) and intestine (i), the latter being full of comminuted fragments of food; p, left plate; p´, right plate; p.ac, accessory plate; B, the plates as seen from the front, with the enveloping membranes removed, lettering as in A. Natural size.
Fig. 146.—Section of the stomach of Melongena, showing the gastric plates (g.p, g.p,) for the trituration of food; b.d, biliary duct; g.g, genital gland; i, intestine; l, liver; oe, oesophagus; st, stomach. (After Vanstone.)
3. The Oesophagus.—That part of the alimentary canal which lies between the pharynx and the stomach (in Pelecypoda between the mouth and stomach) is known as the oesophagus. Its exact limits are not easy to define, since in many cases the tube widens so gradually, while the muscular structure of its walls changes so slowly that it is difficult to say where oesophagus ends and stomach begins. As a rule, the oesophagus is fairly simple in structure, and consists of a straight and narrow tube. In the Pulmonata and Opisthobranchiata it often widens out into a ‘crop,’ which appears to serve the purpose of retaining a quantity of masticated food before it passes on to the stomach. In Octopus and Patella the crop takes the form of a lobular coecum. In the carnivorous Mollusca the oesophagus becomes complicated by the existence of a varying number of glands, by the action of which digestion appears to begin in some cases before the food reaches the stomach proper.
4. The Stomach.—At the posterior end of the oesophagus lies the muscular pouch known as the stomach, in which the digestion of the food is principally performed. This organ may be, as in Limax, no more than a dilatation of the alimentary canal, or it may, as is usually the case, take the form of a well-marked bag or pocket. The two orifices of the stomach are not always situated at opposite ends; when the stomach itself is a simple enlargement of the wall of one side of the alimentary canal, the cardiac or entering orifice often becomes approximated to the orifice of exit (pyloric orifice).
The walls of the stomach itself are usually thickened and strengthened by constrictor muscles. In some Nudibranchs (Scyllaea, Bornella) they are lined on the inside with chitinous teeth. In Cyclostoma, and some Bithynia, Strombus, and Trochus there is a free chitinous stylet within the stomach.[328] In Melongena (Fig. [146]) the posterior end of the oesophagus is provided with a number of hard plate-like ridges, while the stomach is lined with a double row of cuticular knobs, which are movable on their bases of attachment, and serve the purpose of triturating food.[329] Aplysia has several hard plates, set with knobs and spines, and similar organs occur in the Pteropoda. But the most formidable organ for the crushing of food is possessed by the Bullidae, and particularly by Scaphander (Fig. [145]). Here there is a strong gizzard, consisting of several plates connected by powerful cartilages, which crush the shells, which are swallowed whole.
Into the stomach, or into the adjacent portions of the digestive tract, open the ducts which connect with the so-called liver. The functions of this important organ have not yet been thoroughly worked out. The liver is a lobe-shaped gland of a brown-gray or light red colour, which in the spirally-shelled families usually occupies the greater part of the spire. In the Cephalopoda, the two ducts of the liver are covered by appendages which are usually known as the pancreatic coeca; the biliary duct, instead of leading directly into the stomach, passes into a very large coecum (see Fig. [144]) or expansion of the same, which serves as a reservoir for the biliary secretions. At the point of connexion between the coecum and stomach is found a valve, which opens for the issue of the biliary products into the stomach, but closes against the entry of food into the coecum. In most Gasteropoda the liver consists of two distinct lobes, between which are embedded the stomach and part of the intestine. In many Nudibranchiata the liver becomes ‘diffused’ or broken up into a number of small diverticula or glands connecting with the stomach and intestine. The so-called cerata or dorsal lobes in the Aeolididae are in effect an external liver, the removal of which to the outside of the body gives the creature additional stomach-room.
The Hyaline Stylet.—In the great majority of bivalves the intestine is provided with a blind sac, or coecum of varying length. Within this is usually lodged a long cylindrical body known as the hyaline or crystalline stylet. In a well-developed Mytilus edulis it is over an inch in length, and in Mya arenaria between two and three inches. The bladder-like skin of the stylet, as well as its gelatinoid substance, are perfectly transparent. In the Unionidae there is no blind sac, and the stylet, when present, is in the intestine itself. It is said to be present or absent indifferently in certain species.
The actual function performed by the hyaline stylet is at present a matter of conjecture. Haseloff’s experiments on Mytilus edulis tend to confirm the suggestion of Möbius, that the structure represents a reserve of food material, not specially secreted, but a chemical modification of surplus food. He found that under natural conditions it was constantly present, but that specimens which were starved lost it in a few days, the more complete the starvation the more thorough being the loss; it reappeared when they were fed again. Schulze, on the other hand, believes that it serves, in combination with mucus secreted by the stomach, to protect the intestine against laceration by sharp particles introduced with the food. W. Clark found that in Pholas the stylet is connected with a light yellow corneous plate, and imagined therefore that it acts as a sort of spring to work the plate in order to comminute the food, the two together performing somewhat the function of a gizzard.[330]
5. and 6. The Intestine, Rectum, and Anus.—The intestine, the wider anal end of which is called the rectum, almost invariably makes a bend forward on leaving the stomach. This is the case in the Cephalopoda, Scaphopoda, and the great majority of Gasteropoda. The exceptions are the bilaterally symmetrical Amphineura, in which the anus is terminal, and many Opisthobranchiata, in which it is sometimes lateral (Fig. [68], p. 159), sometimes dorsal (Fig. [67]). The intestine is usually short in carnivorous genera, but long and more or less convoluted in those which are phytophagous. In all cases where a branchial or pulmonary cavity exists, the anus is situated within it, and thus varies its position according to the position of the breathing organ. Thus in Helix it is far forward on the right side, in Testacella, Vaginula, and Onchidium almost terminal, in Patella at the back of the neck, slightly to the right side (Fig. [64], p. 157).