Page:EB1911 - Volume 01.djvu/709

 may be, from the pharynx to the stomach, varying in length with the length of the neck and thoracic regions in different animals, and in calibre with the nature of the food. It is almost invariably lined with a many-layered epithelium, forming a tough coating, readily repaired and not easily damaged by hard food masses. It is occasionally separated from the stomach by a slight constriction which may be capable of contraction so as to prevent regurgitation. There are few exceptions to this structural and functional simplicity. In fishes (see, Anatomy) the swim-bladder is developed as a dorsal outgrowth of the oesophagus and may remain in open connexion with it. In certain Teleosteis (e.g. Lutodeira) it is longer than the length it has to traverse and is thrown into convolutions. In many other fish, particularly Selachiis, a set of processes of the lining wall project into the cavity near the stomach and have been supposed to aid in preventing food particles, or living creatures swallowed without injury, escaping backwards into the mouth. In some egg-eating snakes the sharp tips of the ventral spines (hypapophyses) of the posterior cervical vertebrae penetrate the wall of the oesophagus and are used for breaking the shells of the eggs taken as food. In some aquatic Chelonians, the food of which consists chiefly of seaweeds, the lining membrane is produced into pointed processes backwardly directed. In birds this region frequently presents peculiarities. In Opisthocomus it forms an enormously wide double loop, hanging down over the breast-bone, which is peculiarly flattened and devoid of a keel in the anterior portion. In many birds part of the oesophagus may be temporarily dilated, forming a “crop,” as for instance in birds of prey and humming birds. In the flamingo, many ducks, storks, and the cormorant the crop is a permanent although not a highly specialized enlargement. Finally, in the vast majority of seed-eating birds, in gallinaceous birds, pigeons, sandgrouse, parrots and many Passeres, particularly the finches, the crop is a permanent globular dilatation, in which the food is retained for a considerable time, mixed with a slight mucous secretion, and softened and partly macerated by the heat of the body. Many birds feed their young from the soft contents of the crop, and in pigeons, at the breeding season, the cells lining the crop proliferate rapidly and are discharged as a soft cheesy mass into the cavity, forming the substance known as pigeon’s milk. Amongst Mammalia, in Rodentia, Carnivora, elephants and ruminants, the wall of the oesophagus contains a layer of voluntary muscle, by the contraction of which these animals induce anti-peristaltic movements and can so regurgitate food into the mouth.

Stomach.—Where the oesophagus passes into the stomach, the lining wall of the alimentary tract changes from a many-layered epithelium to a mucous epithelium, consisting of a single layer of endodermal cells, frequently thrown into pits or projecting as processes; from being chiefly protective, it has become secretory and absorbing, and maintains this character to the distal extremity where it passes into the epiblast of the proctodaeum. In most cases the course of the alimentary canal from the distal end of the oesophagus to the cloaca or anus is longer than the corresponding region of the body, and the canal is therefore thrown into folds. The fundamental form of the stomach is a sac-like enlargement of the canal, the proximal portion of which is continuous with the line of the oesophagus, but the distal portion of which is bent in the proximal portion, the whole forming an enlarged bent tube. At the distal end of the tube the intestinal tract proper begins, and the two regions are separated by a muscular constriction. In fishes the stomach is generally in one of two forms; it may be a simple bent tube, the proximal limb of which is almost invariably much wider than the distal, anteriorly directed limb; or the oesophagus may pass directly into an expanded, globular or elongated sac, from the anterior lateral wall of which, not far from the oesophageal opening, the duodenum arises. In Batrachia and Reptilia the stomach is in most cases a simple sac, marked off from the oesophagus only by increased calibre. In the Crocodilia, however, the anterior portion of the stomach is much enlarged and very highly muscular, the muscles radiating from a central tendinous area on each of the flattened sides. The cavity is lined by a hardened secretion and contains a quantity of pebbles and gravel which are used in the mechanical trituration of the food, so that the resemblance to the gizzard of birds is well marked. This muscular chamber leads by a small aperture into a distal, smaller and more glandular chamber. In birds the stomach exhibits two regions, an anterior glandular region, the proventriculus, the walls of which are relatively soft and contain enlarged digestive glands aggregated in patches (e.g. some Steganopodes), in rows (e.g. most birds of prey) or in a more or less regular band. The distal region is larger and is lined in most cases by a more or;less permanent lining which is thick and tough in birds with a muscular gizzard, very slight in the others. In many birds, specially those feeding on fish, the two regions of the stomach are of equal width, and are indistinguishable until, on opening the cavity, the difference in the character of the lining membrane becomes visible. In other birds the proventriculus is separated by a well-marked constriction from the posterior and larger region. In graminiferous forms the latter becomes a thick-walled muscular gizzard, the muscles radiating from tendinous areas and the cavity containing pebbles or gravel.

In mammals, the primitive form of the stomach consists of a more or less globular or elongated expansion of the oesophageal region, forming the cardiac portion, and a forwardly curved, narrower pyloric portion, from which the duodenum arises. The whole wall is muscular, and the lining membrane is richly glandular. In the Insectivora, Carnivora, Perissodactyla, and in most Edentata, Chiroptera, Rodentia and Primates, this primitive disposition is retained, the difference consisting chiefly in the degrees of elongation of the stomach and the sharpness of the distal curvature. In other cases the cardiac portion may be prolonged into a caecal sac, a condition most highly differentiated in the blood-sucking bat, Desmodeus, where it is longer than the entire length of the body. There are two cardiac extensions in the hippopotamus and in the peccary. In many other mammals one, two or three protrusions of the cardiac region occur, whilst in the manatee and in some rodents the cardiac region is constricted off from the pyloric portion. In the Artiodactyla the stomach is always complex, the complexity reaching a maximum in ruminating forms. In the Suidae a cardiac diverticulum is partly constricted from the general cavity, forming an incipient condition of the rumen of true ruminants; the general cavity of the stomach shows an approach to the ruminant condition by the different characters of the lining wall in different areas. In the chevrotains, which in many other respects show conditions intermediate between non-ruminant artiodactyles and true ruminants, the oesophagus opens into a wide cardiac portion, incompletely divided into four chambers. Three of these, towards the cardiac extremity, are lined with villi and correspond to the rumen or paunch; the fourth, which lies between the opening of the oesophagus and the pyloric portion of the stomach, is the ruminant reticulum  and its wall is lined with very shallow “cells.” A groove runs along its dorsal wall from the oesophageal aperture to a very small cavity lined with low, longitudinally disposed folds, and forming a narrow passage between the cardiac and pyloric divisions; this is an early stage in the development of the omasum, psalterium or manyplies of the ruminant stomach. The fourth or true pyloric chamber is an elongated sac with smooth glandular walls and is the abomasum, or rennet sack. In the camel the rumen forms an enormous globular paunch with villous walls and internally showing a trace of division into two regions. It is well marked off from the reticulum, the “cells” of which are extremely deep, forming the well-known water-chambers. The psalterium is sharply constricted off from the reticulum and is an elongated chamber showing little trace of the longitudinal ridges characteristic of this region; it opens directly into the relatively small abomasum. In the true ruminants, the rumen forms a capacious, villous reservoir, nearly always partly sacculated, into which the food is passed rapidly as the animal grazes. The food is subjected to a rotary movement in the paunch, and is thus repeatedly subjected to