moistening with the fluids secreted by the reticulum, as it is passed over the aperture of that cavity, and is formed into a rounded bolus. Most ruminants swallow masses of hairs, and these, by the rotary action of the paunch, are aggregated into peculiar dense, rounded balls which are occasionally discharged from the mouth and are known as “hair-balls” or “bezoars.” The food bolus, when the animal is lying down after grazing, is passed into the oesophagus and reaches the mouth by anti-peristaltic contractions of the oesophagus. After prolonged mastication and mixing with saliva, it is again swallowed, but is now passed into the psalterium, which, in true ruminants, is a small chamber with conspicuous longitudinal folds. Finally it reaches the large abomasum where the last stages of gastric digestion occur.
In the Cetacea the stomach is different from that found in any other group of mammals. The oesophagus opens directly into a very large cardiac sac the distal extremity of which forms a long caecal pouch. At nearly the first third of its length this communicates by a narrow aperture into the elongated, relatively narrow pyloric portion. The latter is convoluted and constricted into a series of chambers that differ in different groups of Cetacea. In the Sirenia the stomach is divided by a constriction into a cardiac and a pyloric portion, and the latter has a pair of caeca. In most of the Marsupialia the stomach is relatively simple, forming a globular sac with the oesophageal and pyloric apertures closely approximated; in the kangaroos, on the other hand, the stomach is divided into a relatively small, caecal cardiac portion and an enormously long sacculated and convoluted pyloric region, the general arrangement of which closely recalls the large caecum of many mammals.
Intestinal Tract.—It is not yet possible to discuss the general morphology of this region in vertebrates as a group, as, whilst the modifications displayed in birds and mammals have been compared and studied in detail, those in the lower groups have not yet been systematically co-ordinated.
Fishes.—In the Cyclostomata, Holocephali and a few Teleostei the course of the gut is practically straight from the pyloric end of the stomach to the exterior, and there is no marked differentiation into regions. In the Dipnoi, a contracted sigmoid curve between the stomach and the dilated intestine is a simple beginning of the complexity found in other groups. In very many of the more specialized teleosteans, the gut is much convoluted, exhibiting a series of watchspring-like coils. In a number of different groups, increased surface for absorption is given, not by increase in length of the whole gut, but by the development of an internal fold known as the spiral valve. This was probably originally a longitudinal fold similar to the typhlosole of chaetopods. It forms a simple fold in the larval Ammocoete, and in its anterior region remains straight in some adult fish, e.g. Polypterus, but in the majority of cases it forms a complex spiral, wound round the inner wall of the expanded large intestine, the internal edge of the fold sometimes meeting to form a central column. It occurs in Cyclostomata, Selachii, Holocephali, Chondrostei, Crossopterygii, Amiidae, Lepidosteidae and Dipnoi. A set of organs peculiar to fish and known as the pyloric caeca are absent in Cyclostomata and Dipnoi, in most Selachii and in Amia, but present, in numbers ranging from one to nearly two hundred, in the vast majority of fish. These are outgrowths of the intestinal tract near the pyloric extremity of the stomach, and their function is partly glandular, partly absorbing. In a few Teleostei there is a single caecal diverticulum at the beginning of the “rectum,” and in the same region a solid rectal gland occurs in most elasmobranchs, whilst, again, in the Dipnoi a similar structure opens into the cloaca. These caeca have been compared with the colic caeca of higher vertebrates, but there is yet no exact evidence for the homology.
In the Batrachia the course of the intestinal tract is nearly straight from the pyloric end of the stomach to the cloaca, in the case of the perennibranchiates there being no more than a few simple loops between the expanded “rectum” and the straight portion that leaves the stomach. In the Caducibranchiata the anterior end of the enlarged rectum lies very close to the distal extremity of the stomach, and the gut, between these two regions, is greatly lengthened, forming a loop with many minor loops borne at the periphery of an expanse of mesentery, recalling the Meckelian tract of birds and mammals. In the tadpole this region is spirally coiled and is still longer relatively to the length of the whole tract. In Hyla and Pipa there is a small caecum comparable with the colic caecum of birds and mammals.
In Reptilia the configuration of the intestinal tract does not differ much from that in Batrachia, the length and complexity of the minor coils apparently varying with the general configuration of the body, that is to say, in reptiles with a long, narrow, and snake-like body the minor loops of the gut are relatively short and unimportant, whilst in those with a more spacious cavity, such as chelonians, many lizards and crocodiles, the gut may be relatively long and disposed in many minor coils. There is comparatively little differentiation between the mid-gut and the gut in cases where the whole gut is long; in the others the hind-gut is generally marked by an increase of calibre. A short caecal diverticulum, comparable with the colic caecum of birds and mammals, is present in many snakes and lizards and in some chelonians.
In fishes, batrachians and reptiles the intestinal tract is swung from the dorsal wall of the abdominal cavity by a mesentery which is incomplete on account of secondary absorption in places, and which grows out with the minor loops of the gut. There are also traces, more abundant in the lower forms, of the still more primitive ventral mesentery.
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| Fig. 4.—Intestinal Tract of Chauna chavaria. | |||
| c.c. | Colic caeca. | p.v. | Cut root of portal vein. |
| d. | Duodenum. | r.v. | Rectal vein. |
| g. | Glandular patch. | s. | Proventriculus. |
| l.l. | Meckel’s tract. | y. | Meckel’s diverticulum, or |
| l.i. | Hind-gut. | Yolk-sac vestige. | |
Intestinal Tract in Birds and Mammals.—There is no doubt but that the similarity of the modes of disposition of the alimentary tract in birds and mammals points to the probability of the chief morphological features of this region in these animals having been laid down in some common ancestor, although we have not, yet sufficient exact knowledge of the gut in Pisces, Batrachia, and Reptilia to find amongst these with any certainty the most probable survival from the ancestral condition. The primitive gut must be supposed to have run backwards from the stomach to the cloaca suspended from the dorsal wall of the body-cavity by a dorsal mesentery. This tract, in the course of phylogeny of the common ancestors of birds and mammals, became longer than the straight length between its extreme points and, consequently, was thrown into a series of folds. The
