ALIMENTARY CANAL, in anatomy. The alimentary canal, strictly speaking, is the whole digestive tract from the mouth to the anus. From the one orifice to the other the tube is some 25 to 30 ft. long, and the food, in its passage, passes through the following parts one after the other:—mouth, pharynx, oesophagus, stomach, small intestines, caecum, large intestines, rectum and anus. Into this tube at various points the salivary glands, liver and pancreas pour their secretions by special ducts. As the mouth (q.v.) and pharynx (q.v.) are separately described, the detailed description will here begin with the oesophagus or gullet.
The oesophagus (Gr. οἴσω, I will carry, and φαγεῖν, to eat), a muscular tube lined with mucous membrane, stretches from the lower limit of the pharynx, at the level of the cricoid cartilage, to the cardiac orifice of the stomach. It is about 10 in. long (25 cm.) and half to one inch in diameter. At first it lies in the lower part of the neck, then in the thorax, and lastly, for about an inch, in the abdomen. As far as the level of the fourth or fifth thoracic vertebra it lies behind the trachea, but when that tube ends, it is in close contact with the pericardium, and, at the level of the tenth thoracic vertebra, passes through the oesophageal opening of the diaphragm (q.v.), accompanied by the two vagi nerves, the left being in front of it and the right behind. In the abdomen it lies just behind the left lobe of the liver. Both in the upper and lower parts of its course it lies a little to the left of the mid line. Its mucous membrane is thrown into a number of longitudinal pleats to allow stretching.
The stomach (Gr. στόμαχος) is an irregularly pear-shaped bag, situated in the upper and left part of the abdomen. It is somewhat flattened from before backward and so has an anterior and posterior surface and an upper and lower border. When moderately distended the thick end of the pear or fundus bulges upward and to the left, while the narrow end is constricted to form the pylorus, by means of which the stomach communicates with the small intestine. The cardiac orifice, where the oesophagus enters, is placed about a third of the way along the upper border from the left end of the fundus, and, between it and the pylorus, the upper border is concave and is known as the lesser curvature. From the cardiac to the pyloric orifice, round the lower border, is the greater curvature. The stomach has in front of it the liver (see fig. 1), the diaphragm and the anterior abdominal wall, while behind it are the pancreas, left kidney, left adrenal, spleen, colon and mesocolon. These structures form what is known as the stomach chamber. When the stomach is empty it contracts into a tubular organ which is frequently sharply bent, and the transverse colon ascends to occupy the vacant part of the stomach chamber.
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| From A. Birmingham; Cunningham’s Text-Book of Anatomy. |
| Fig. 1.—The Abdominal Viscera in situ, as seen when the abdomen is laid open and the great omentum removed (drawn to scale from a photograph of a male body aged 56, hardened by formalin injections).
The ribs on the right side are indicated by Roman numerals; it will be observed that the eighth costal cartilage articulated with the sternum on both sides. The subcostal, intertubercular, and right and left Poupart lines are drawn in black, and the mesial plane is indicated by a dotted line. The intercostal muscles and part of the diaphragm have been removed, to show the liver and stomach extending up beneath the ribs. The stomach is moderately distended, and the intestines are particularly regular in their arrangement. |
The last inch of the stomach before reaching the pylorus is usually tubular and is known as the pyloric canal. Before reaching this there is a bulging known as the pyloric vestibule (see D. J. Cunningham, Tr. R. Soc. of Edinib. vol. xlv. pt. 1, No. 2). The pylorus is an oval opening, averaging half an inch in its long axis but capable of considerable distension; it is formed by a special development of the circular muscle layer of the stomach, and during life is probably tightly closed. The mucous membrane of the stomach is thrown into pleats or rugae when the organ is not fully distended, while between these it has a mammillated appearance.
Superficial to the mucous coat is a sub-mucous, consisting of loose connective tissue, while superficial to this are three coats of unstriped muscle, the inner oblique, the middle circular and the outer longitudinal. The peritoneal coat is described in the article on the coelom and serous membranes.
The small intestine is a tube, from 22 to 25 ft. long, beginning at the pylorus and ending at the ileo-caecal valve; it is divided into duodenum, jejunum and ileum.
The duodenum is from 9 to 11 in. long and forms a horseshoe or C-shaped curve, encircling the head of the pancreas. It differs from the rest of the gut in being retroperitoneal. Its first part is horizontal and lies behind the fundus of the gall-bladder, passing backward and to the right from the pylorus. The second part runs vertically downward in front of the hilum of the right kidney, and into this part the pancreatic and bile ducts open. The third part runs horizontally to the left in front of the aorta and vena cava, while the fourth part ascends to the left side of the second lumbar vertebra, after which it bends sharply downward and forward to form the duodeno-jejunal flexure.
The jejunum forms the upper two-fifths of the rest of the small intestine; it, like the ileum, is thrown into numerous convolutions and is attached by the mesentery to the posterior abdominal wall. (See Coelom and Serous Membranes.)
The ileum is the remaining three-fifths of the small intestine, though there is no absolute point at which the one ends and the other begins. Speaking broadly, the jejunum occupies the upper and left part of the abdomen below the subcostal plane (see Anatomy: Superficial and Artistic), the ileum the lower and right part. About 3 ft. from its termination a small pouch, known as Meckel’s diverticulum, is very occasionally found. At its termination the ileum opens into the large intestine at the ileo-caecal valve.
The caecum is a blind sac occupying the right iliac fossa and extending down some two or three inches below the ileo-caecal junction. From its posterior and left surface the vermiform appendix protrudes, and usually is directed upward and to the left, though it not infrequently hangs down into the true pelvis. This worm-like tube is blind at its end and is usually 3 or 4 in. long, though it has been seen as long as 10 in. Its internal opening into the caecum is about 1 in. below that of the ileum. On transverse section it is seen to be composed of (1) an external muscular coat, (2) a submucous coat, (3) a mass of lymphoid tissue, which appears after birth, and (4) mucous membrane. In many cases its lumen is wholly or partly obliterated, though this is probably due to disease (see R. Berry and L. Lack, Journ. Anat. & Phys. vol. xl. p. 247). Guarding the opening of the ileum into the caecum is the ileo-caecal valve, which consists of two cusps projecting into the caecum; of these the upper forms a horizontal shelf, while the lower slopes up to it obliquely. Complete absence of the valve has been noticed, and in one such case the writer found that no abdominal inconvenience had been recorded during life. The caecum is usually completely covered by peritoneum, three special pouches of which are often found in its neighbourhood; of these the ileo-colic is just above the point of junction of the ileum and caecum, the ileocaecal just below that point, while the retro-caecal is behind the caecum. At birth the caecum is a cone, the apex of which is the appendix; it is bent upon itself to form a U, and sometimes this arrangement persists throughout life (see C. Toldt, “Die Formbildung d. menschl. Blinddarmes,” Sitz. der Wiener Akad. Bd. ciii. Abteil. 3, p. 41) .
The ascending colon runs up from the caecum at the level of the ileo-caecal valve to the hepatic flexure beneath and behind the right lobe of the liver; it is about 8 in. long and posteriorly is in contact with the abdominal wall and right kidney. It is covered by peritoneum except on its posterior surface (see fig. 1).
The transverse colon is variable in position, depending largely on the distension of the stomach, but usually corresponding to the subcostal plane (see Anatomy: Superficial and Artistic). On the left side of the abdomen it ascends to the splenic flexure, which may make an impression on the spleen (see Ductless Glands), and is bound to the diaphragm opposite the eleventh rib by a fold of peritoneum called the phrenico-colic ligament. The peritoneal relations of this part are discussed in the article on the coelom and serous membranes.
The descending colon passes down in front of the left kidney and left side of the posterior abdominal wall to the crest of the ilium; it is about 6 in. long and is usually empty and contracted while the rest of the colon is distended with gas; its peritoneal relations are the same as those of the ascending colon, but it is more likely to be completely surrounded.
The iliac colon stretches from the crest of the ilium to the inner border of the psoas muscle, lying in the left iliac fossa, just above and parallel to Poupart’s ligament. Like the descending, it is usually uncovered by peritoneum on its posterior surface. It is about 6 in. in length.
The pelvic colon lies in the true pelvis and forms a loop, the two limbs of which are superior and inferior while the convexity reaches across to the right side of the pelvis. In the foetus this loop occupies the right iliac fossa, but, as the caecum descends and enlarges and the pelvis widens, it is usually driven out of this region. The distal end of the loop turns sharply downward to reach the third piece of the sacrum, where it becomes the rectum. To this pelvic colon Sir F. Treves (Anatomy of the Intestinal Canal, London, 1885) has given the name of the omega loop. Formerly the iliac and pelvic colons were spoken of as the sigmoid flexure, but Treves and T. Jonnesco (Le Colon pelvien pendant la vie intra-utérine, Paris, 1892) have pointed out the inapplicability of the term, and to the latter author the modern description is due.
The rectum, according to modern ideas, begins in front of the third piece of the sacrum; formerly the last part of the Ω (or omega) loop was described as its first part. It ends in a dilatation or rectal ampulla, which is in contact with the back of the prostate in the male and of the vagina in the female and is in front of the tip of the coccyx. The rectum is not straight, as its name would imply, but has a concavity forward corresponding to that of the sacrum and coccyx.
When viewed from in front three bends are usually seen, the upper and lower of which are sharply concave to the left, the middle one to the right. At the end of the pelvic colon the mesocolon ceases, and the rectum is then only covered by peritoneum at its sides and in front; lower down the lateral covering is gradually reflected off and then only the front is covered. About the junction of the middle and lower thirds of the tube the anterior peritoneal covering is also reflected off on to the bladder or vagina, forming the recto-vesical pouch in the male and the pouch of Douglas in the female. This reflexion is usually about 3 in. above the anal aperture, but may be a good deal lower.
The anal canal is the termination of the alimentary tract, and runs downward and backward from the lower surface of the rectal ampulla between the levatores ani muscles. It is about an inch long and its lateral walls are in contact, so that in section it appears as an antero-posterior slit (see J. Symington, Journ. Anat. and Phys. vol. 23, 1888).
Structure of the Intestine.—The intestine has four coats: serous, muscular, submucous and mucous. The serous or peritoneal coat has already been described wherever it is present. The muscular coat consists of unstriped fibres arranged in two layers, the outer longitudinal and the inner circular (see fig. 2). In the large intestine the longitudinal fibres, instead of being arranged evenly round the tube as they are in the small, are gathered into three longitudinal bands called taeniae (see fig. 1); by the contraction of these the large intestine is thrown into a series of sacculi or slight pouches. The taeniae in the caecum all lead to the vermiform appendix, and form a useful guide to this structure. In the rectum the three taeniae once more become evenly arranged over the whole surface of the bowel, but more thickly on the anterior and posterior parts. The circular layer is always thicker than the longitudinal; in the small intestine it decreases in thickness from the duodenum to the ileum, but in the large it gradually increases again, so that it is thickest in the duodenum and rectum.
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| From A. Birmingham; Cunningham’s Text-Book of Anatomy. |
| Fig. 2.—Diagram to show the structure of the small and large intestine and the duodenum. |
The submucous coat is very strong and consists of loose areolar tissue in which the vessels break up.
The mucous coat is thick and vascular (see fig. 2); it consists of an epithelial layer most internally which forms the intestinal glands (see Epithelial, Endothelial and Glandular Tissues). External to this is the basement membrane, outside which is a layer of retiform tissue, and this is separated from the submucous coat by a very thin layer of unstriped muscle called the muscularis mucosae. In the duodenum and jejunum the mucous membrane is thrown into a series of transverse pleats called valvulae conniventes (see fig. 3); these begin about an inch from the pylorus and gradually fade away as the ileum is reached. About 4 in. from the pylorus the common bile and pancreatic ducts form a papilla, above which one of the valvulae conniventes makes a hood and below which a vertical fold, the frenulum, runs downward. The surface of the mucous membrane of the whole of the small intestine has a velvety appearance, due to the presence of closely-set, minute, thread-like elevations called villi (see fig. 2). Throughout the whole length of the intestinal tract are minute masses of lymphoid tissue called solitary glands (see fig. 2); these are especially numerous in the caecum and appendix, while in the ileum they are collected into large oval patches, known as agminated glands or Peyer’s patches, the long axes of which, from half an inch to 4 in. long, lie in the long axis of the bowel. They are always found in that part of the intestine which is farthest from the mesenteric attachment. In the interior of the rectum three shelf-like folds, 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 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 Ichthyology, 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 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 mesentery 
Fig. 5.—Intestinal Tract of Canis vulpes. S, cut end of duodenum; C, caecum; R, cut end of rectum.grew out with these folds, but the presence of adjacent organs, the disturbance due to the outgrowth of the liver, and the secondary relations brought about between different portions of the gut, as the out-growing loops invaded each other’s localities, disturbed the primitive simplicity. Three definite regions of outgrowth, however, became conspicuous and are to be recognized in the actual disposition of the gut in existing birds and mammals. The first of these is the duodenum. In the vast majority of birds, and in some of the simpler mammals, the portion of the gut immediately distal of the stomach grows out into a long and narrow loop (fig. 4, d), the proximal and distal ends of which are close together, whilst the loop itself may remain long and narrow, or may develop minor loops on its course. In mammals generally, however, the duodenum is complex and is not so sharply marked off from the distal portion of the gut as in birds. The second portion is Meckel’s tract. It consists of the part generally known as the small intestines, the jejunum and ileum of human anatomy, and
stretches from the distal end of the duodenum to the caecum or caeca. It is the chief absorbing portion of the gut, and in nearly all birds and mammals is the longest portion. It represents, however, only a very small part of the primitive straight gut, corresponding to not more than two or three somites of the embryo. This narrow portion grows out to form the greater part of what is called the pendent loop in mammalian embryology. Its anterior or proximal end lies close to the approximated
proximal and distal ends of the duodenal loop, whilst its distal end passes into the hind-gut at the colic caecum or caeca. In the embryos of all birds and mammals, the median point of Meckel’s tract, the part of the loop which has grown out farthest from the dorsal edge of the mesentery, is marked by the diverticulum caecum vitelli, the primitive connexion of the cavity of the gut with the narrowing stalk of the yolk-sac (fig. 4, y.) Naturally, in birds where the yolk-sac is of great functional importance this diverticulum is large, and in a majority of the families of birds persists throughout life, forming a convenient point of orientation. In mammals, no doubt in association with the functional reduction of the yolk-sac, this diverticulum, which is known as Meckel’s diverticulum, has less importance, and whilst it has been observed in a small percentage of adult human subjects has not been recognized in the adult condition of any lower Mammalia.
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| Fig. 6.—Intestinal Tract of Macropus bennetti. S, cut end of duodenum; R, cut end of rectum; C, caecum; C2, accessory caecum; C.L, colic loop of hind-gut. |
Fig. 7.—Intestinal Tract of Tapir. S, cut end of duodenum; R, cut end of rectum; C, caecum; CL, colon. |
In birds, Meckel’s tract falls into minor folds or loops, the disposition of which forms a series of patterns remarkably different in appearance and characteristic of different groups. In fig. 4 an extremely primitive type is represented. In mammals Meckel’s tract remains much more uniform; it may be short, or increase enormously in length, but in either case it falls into a fairly symmetrical shape, suspended at the circumference of a nearly circular expanse of mesentery. Where it is short it is thrown into very simple minor loops (figs. 5, 6 and 7); where it is long, these minor loops form a convoluted mass (figs. 8 and 9).
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| Fig. 8.—Intestinal Tract of Giraffe. S, cut end of duodenum; R, cut end of rectum; C, caecum; P.C.L, post-caecal loop; S.P, spiral loop; SF, third loop of hind-gut. |
The third portion of the gut should be termed the hind-gut and lies between the caecum or caeca and the anus, corresponding to the large intestines, colon and rectum of human anatomy. It is formed from a much larger portion of the primitive straight gut than the duodenum and Meckel’s tract together, and its proximal portion, in consequence, lies very close to the origin of the duodenum. In the vast majority of birds, the hind-gut in the adult is relatively extremely short, often being only from
one-eighth to one-thirtieth of the whole length of the gut. A certain number of primitive birds, however, have retained a relatively long condition of the hind-gut (fig. 4), the greatest relative length occurring in struthious birds, and particularly in the ostrich, where the hind-gut exceeds in length the duodenum and Meckel’s tract together. Mammals may be contrasted with birds as a group in which the hind-gut is always relatively long, sometimes extremely long, and in which, moreover, there is a strong tendency to differentiation of the hind-gut into regions the characters of which are of systematic importance. The first region is the colon, which forms a very simple expansion in mammals such as Carnivora (fig. 5), where the whole hind-gut is relatively short, or a series of simple loops in mammals in which the whole gut has a primitive disposition (e.g. Marsupialia, fig. 6). In the odd-toed Ungulata, the colon (fig. 7) forms an enormously long loop, the two limbs of which are closely approximated and the calibre of which is very large. 
Fig. 9.—Intestinal Tract of Gorilla. S, cut end of duodenum; R, cut end of rectum; C, vermiform appendix of caecum; X, X2, X3, cut ends of factors of the portal vein.In Ruminantia (fig. 8) the colon is still more highly differentiated, displaying first a simple wide loop, then a complicated watchspring-like coil, and finally a very long, irregular portion. In the higher Primates (fig. 9) it forms one enormous very wide loop, corresponding to the ascending, transverse and descending colons of human anatomy, and a shorter distal loop, the omega loop of human anatomy. Other striking patterns are displayed in other mammalian groups.
The second region of the hind-gut is usually known as the rectum, and although it is sometimes lengthened it is typically little longer than the portion of the primitive straight gut that it represents.
Adaptations of the Intestinal Tract to Function.—The chief business of the gut is to provide a vascular surface to which the prepared food is applied so that the nutritive material may be absorbed into the system. Overlying and sometimes obscuring the morphological patterns of the gut, are many modifications correlated with the nature of the food and producing homoplastic resemblances independent of genetic affinity. Thus in birds and mammals alike there is a direct association of herbivorous habit with great relative length of gut. The explanation of this, no doubt, is simply that the vegetable matter which such creatures devour is in a form which requires not only prolonged digestive action, but, from the intimate admixture of indigestible material, a very large absorbing surface. In piscivorous birds and mammals, the gut is very long, with a thick wall and a relatively small calibre, whilst there is a general tendency for the regions of the gut to be slightly or not at all defined. Fish, as it is eaten by wild animals, contains a large bulk of indigestible matter, and so requires an extended absorbing surface; the thick wall and relatively small calibre are protections against wounding by fish bones. In frugivorous birds the gut is strikingly short, wide and simple, whilst a similar change has not taken place in frugivorous mammals. Carnivorous birds and mammals have a relatively short gut. In birds, generally, the relation of the length and calibre of the gut to the size of the whole creature is striking. If two birds of similar habit and of the same group be compared, it will be found that the gut of the larger bird is relatively longer rather than relatively wider. The same general rule applies to Meckel’s tract in mammals, whereas in the case of the hind-gut increase of capacity is given by increase of calibre rather than by increased length.
The Colic Caeca.—These organs lie at the junction of the hind-gut with Meckel’s tract and are homologous in birds and mammals although it happens that their apparent position differs in the majority of cases in the two groups. In most birds, the hind-gut is relatively very short, and the caecal position, accordingly, is at a very short distance from the posterior end of the body, whereas in most mammals the hind-gut is very long and the position of the caecum or caeca is relatively very much farther from the anus. Next, in most birds, the caeca when present are paired, whereas in most mammals there is only a single caecum. On the other hand, in certain birds (herons) as a normal occurrence, and in many birds as an individual variation, only a single caecum occurs. In some mammals, e.g. many armadillos, in Hyrax and the manatee, the caeca are normally paired; in many other (e.g. some rodents and marsupials) in addition to the normal caecum there is a reduced second caecum, whilst in quite a number of forms the relation of the caecum, ileum and colon at their junction is readily intelligible on the assumption that the caeca were originally paired. The origin and many of the peculiarities of the ileo-caecal valve find their best explanation on this hypothesis.
The caeca are hollow outgrowths of the wall of the gut, the blind ends being directed forwards. The caecal wall is in most cases highly glandular and contains masses of lymphoid tissue. In birds and in mammals this tissue may be so greatly increased as to transform the caecum into a solid or nearly solid sac, the calibre of which is for the most part smaller than that of the unmodified caecum. In some birds, the whole area of the caecum may be modified in this way; in mammals, it is generally the terminal portion, which then becomes the vermiform appendix, familiar in the anthropoid apes, in man and in some rodents. It is difficult to see in this modification merely a degeneration; not improbably it is the formation of a new glandular organ.
The caeca exhibit almost every gradation of development, from relatively enormous size to complete absence, and there is no definite, invariable connexion between the nature of the food and the degree of their development. In the case of birds, it may be said that on the whole the caeca are generally large in herbivorous forms and generally small in insectivorous, frugivorous, carnivorous and piscivorous forms, but there are many exceptions. Thus, owls and falcons have a diet that is closely similar, and yet owls have a pair of very long caeca, whilst in the Falconidae these organs are much reduced and apparently functionless. The insectivorous and omnivorous rollers,motmots and bee-eaters have a pair of large caeca, whilst in passerine birds of similar habit the caeca are vestigial glandular nipples. It is impossible to doubt that family history dominates in this matter. Certain families tend to retain the caeca, others to lose them, and direct adaptation to diet appears only to accelerate or retard these inherited tendencies. So also in mammals, no more than a general relation between diet and caecal development can be shown to exist, although the large size of the single caecum of mammals is more closely associated with a herbivorous as opposed to a carnivorous, frugivorous, piscivorous or omnivorous diet than is the case in birds. There is no relationship between diet and the complete or partial presence of both members of the primi-pair of caeca in mammals, the occurrence of the pair being rather an “accident” of inheritance than in any direct relation to function.
Literature.—T. W. Bridge, in The Cambridge Natural History (vol. vii).; D. S. Jordan, A Guide to the Study of Fishes; R. Owen, Anatomy of Vertebrates; M. Weber, Die Säugethiere; W. H. Flower, The Organs of Digestion in Mammalia; R. Wiedersheim, Lehrbuch der vergleichenden Anatomie der Wirbelthiere; A. Oppel, Lehrbuch der vergleichenden mikroskopischen Anatomie der Wirbelthiere; Chalmers Mitchell, “The Intestinal Tract of Birds,” Transactions of the Linn. Soc. of London (vol. viii., 1901); and “On the Intestinal Tract of Mammals,” Transactions of the Zool. Soc. of London (vol. xvii., 1905). (In the two latter memoirs a fuller list of literature is given.) (P. C. M.)