one above the other, project into the cavity and correspond to the lateral concavities or kinks of the tube. They are not in the same line and the largest is usually on the right side. They are known as the plicae recti or valves of Houston. In the anal canal are four or five longitudinal folds called the columns of Morgagni. (For further details, see Quain’s Anatomy, London, 1896; Gray’s Anatomy, London, 1905; Cunningham’s Anatomy, Edinburgh, 1906.)
Embryology.—The greater part of the alimentary canal is formed by the closing-in of the entoderm to make a longitudinal tube, ventral and parallel to the notochord. This tube is blind in front and behind (cephalad and caudad), but the middle part of its ventral wall is for some distance continuous with the wall of the yolk-sac, and this part of the canal, which at first opens into the yolk-sac by a very wide aperture, is called the mid gut. The part in front of it, which lies dorsal to the heart, is the fore gut, while the part behind the aperture of the yolk-sac is the hind gut.
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| From A. Birmingham; Cunningham’s Text-Book of Anatomy. |
| Fig. 3.—Valvulae Conniventes. |
| A, As seen in a bit of jejunum which has been filled with alcohol and hardened. B, A portion of fresh intestine spread out under water. |
The pharynx, oesophagus, stomach and part of the duodenum are developed from the fore gut, a good deal of the colon and the rectum from the hind gut, while the mid gut is responsible for the rest. The cephalic part of the fore gut forms the pharynx (q.v.), and about the fourth week the stomach appears as a fusiform dilatation in the straight tube. Between the two the oesophagus gradually forms as the embryo elongates. The opening into the yolk-sac, which at first is very wide, gradually narrows, as the ventral abdominal walls close in, until in the adult the only indication of the connexion between the gut and the yolk-sac is the very rare presence (about 2%) of Meckel’s diverticulum already referred to. The stomach soon shows signs of the greater and lesser curvatures, the latter being ventral, but maintains its straight position. About the sixth week the caecum appears as a lateral diverticulum, and, until the third month, is of uniform calibre; after this period the terminal part ceases to grow at the same rate as the proximal, and so the vermiform appendix is formed. The mid gut forms a loop with its convexity toward the diminishing vitelline duct, or remains of the yolk-sac, and until the third month it protrudes into the umbilical cord. The greater curvature of the stomach grows more rapidly than the lesser, and the whole stomach turns over and becomes bent at right angles, so that what was its left surface becomes ventral. This turning over of the stomach throws the succeeding part of the intestine into a duodenal loop, which at first has a dorsal and ventral mesentery (see Coelom and Serous Membranes). The intestine now grows very rapidly and is thrown into a series of coils; the caecum ascends and passes to the right ventral to the duodenum, and presses it against the dorsal wall of the abdomen; then it descends toward its permanent position in the right iliac fossa.
From the ventral surface on the hinder (caudal) closed end of the intestinal tube the allantois grows to form the placenta and bladder (see Urinary System, Reproductive System and Placenta), and this region is the cloaca into which the alimentary, urinary and generative canals or ducts all open, but later two lateral folds appear which, by their union, divide the cloaca into a ventral and a dorsal part, the former being genito-urinary and the latter alimentary or intestinal. In this way the rectum or dorsal compartment is shut off from the genito-urinary. Later an ectodermal invagination at the hind end of the embryo develops and forms the anal canal; this is the proctodaeum, and for some time it is separated from the hind (caudal) end of the rectal part of the mesodaeum (or part of the intestinal canal formed from the mesoderm) by a membrane called the anal membrane. This is eventually absorbed and the digestive tract now communicates with the surface by the anus.
F. Wood Jones (British Medical Journal, 17th of December 1904) has given a somewhat different description of the development of the cloaca and anus, which better explains the various abnormalities met with in this region but requires further confirmation before it is generally accepted. For the development of the mouth, pharynx, lungs, liver and pancreas from the primitive alimentary canal, the reader is referred to the special articles on those structures. (For further details, see W. His, Anatomie menschlicher Embryonen (Leipzig, 1880–1885); C. S. Minot’s Embryology (New York, 1897); and J. P. M‘Murrich, Development of the Human Body (London, 1906). (F. G. P.)
Comparative Anatomy.—The primitive condition of the vertebrate alimentary canal may be described as a straight, simple tube, consisting of an anterior portion, the stomodaeum, formed by an ectodermal invagination, the mesenteron, a long median portion lined by endoderm, and a short posterior portion, the proctodaeum, formed by ectodermal invagination. In the lower vertebrates the primitive tube subserved also the purpose of respiration, and traces of the double function remain in the adult structure of all vertebrates (see Mouth, Pharynx). In fish, the pharynx, or branchial region, suddenly becomes narrower, posterior to the gill-slits, to form the oesophagus; in higher animals the oesophagus, in the adult, is separated from the primitive pharyngeal region and lies dorsal to it. Probably,. in the primitive vertebrata, the entire alimentary canal was lined with ciliated cells. Traces of this ciliation persist in many living forms. In the Ammocoete, the larval form of Petromyzon (see Cyclostomata), the whole canal is ciliated except the pharynx and the rectum; in the Dipnoi the epithelium of the stomach and the intestines is ciliated; in Selachii that of the posterior part oaf the gullet, and the spiral valve, is ciliated; extensive ciliation may occur in almost any region of the gut of the lower teleostomes, but in the higher forms (Teleostei) it is generally absent. In the latter, however, and in higher groups of vertebrates, a peculiar striated border on the columnar cells lining the intestinal tract has been held to be a final trace of ancestral ciliation.
The alimentary canal may be conveniently described in three divisions, the oesophagus or gullet, the passage by which food reaches the stomach, the stomach, typically an expanded region in which the food remains for a considerable time and is mechanically pulped, mixed with mucus and certain digestive juices (see Nutrition) and partly macerated, the intestinal tract or gut, extending from the distal end of the stomach to the cloaca or anus, in which the food is subjected to further digestive action,. but which is above all the region in which absorption of the products of digestion takes place, the refuse material together with quantities of waste matter entering the gut from the blood and liver being gradually passed towards the anus for discharge from the body.
The oesophagus is essentially merely a passage, as straight as
