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THE CANINE GASTROINTESTINAL TRACT: Embryology and Anatomy with certain aspects of physiology and clinical application At an early stage of development, the part of the gut (enteron) that becomes the stomach and intestine passes through the body as a simple tube. The diaphragm is just beginning to develop. You can easily identify the part of the gut that is going to become the stomach because it is increased in diameter relative to the intestine and is just caudal to the developing diaphragm. In the figure below, a first indication of the developing diaphragm is labeled pleuroperitoneal fold. The dilatation of the stomach is unmarked but can be seen as an expanded part of the gut just caudal to the pleuroperitoneal fold. Fig. 1 The relationship of stomach and intestine to the lining membrane of the body cavity, the membrane that will become the peritoneum, is shown in Figure 2: 1 Fig. 2 The peritoneum that extends from the body wall to the stomach and intestine is a double layer. Between the two layers of peritoneum is loose connective tissue in association with vessels and nerves that supply the gut wall and the ducts that pass from its accessory glands, the liver and pancreas. The part of this peritoneum that extends from the abdominal surface of the diaphragm and dorsal body wall to the stomach is designated dorsal mesogastrium; it carries vessels and nerves to the stomach. The part that extends from the dorsal body wall to the intestine (enteron) is the mesentery. The mesentery conveys vessels, nerves, and the bile and pancreatic ducts to the intestine. The stomach is not perfectly cylindrical but is spindle-shaped and curved on its long axis so that it has a greater curvature dorsally and a lesser curvature ventrally. The dorsal mesogastrium passes from the diaphragm and dorsal body wall to the greater curvature of the stomach. The ventral mesogastrium passes from the lesser curvature of the stomach and neighboring part of the intestine to the ventral diaphragm and ventral body wall. Its attachment to the ventral body wall extends caudally to the level of the umbilicus. This part of the peritoneum is designated the ventral mesogastrium. The hepato-pancreatic bud, which will form the liver and part of the pancreas, proceeds as a tubular evagination from the neighboring intestine into the ventral mesogastrium (Fig. 3). 2 Fig. 3 The hepato-pancreatic bud, enters the ventral mesogastrium and begins to branch extensively, forming the liver. An extension from the first part of the bud branches to form the ventral pancreas. Ultimately the liver becomes so large as to completely separate the ventral mesogastrium into a part passing from the lesser curvature of the stomach to the liver---this becomes the lesser omentum--; and a part passing from the liver to the diaphragm and ventral body wall. This part of the ventral mesogastrium becomes the falciform ligament. The falciform ligament conveys the umbilical vein from the umbilicus to the liver. In the dog, both before and after birth, parts of the falciform ligament are lost. A small, remnant, fold can often be detected ventral to the caudal vena cava as it leaves the liver to penetrate the diaphragm. A larger part of the falciform ligament remains at the level of the umbilicus. It becomes fat-filled and is encountered in every mid-ventral abdominal incision (e.g., spay) that takes place close to the umbilibus. In the dog and cat, the umbilical vein is entirely involuted (lost) after birth. In the horse, the umbilical vein remains and is the round ligament of the liver. It is patent (carries blood) in about 80% of horses throughout life. In the remaining 20%, it remains only as a fibrous cord. With growth of the stomach, its greater curvature, which is at first dorsal, turns to the left, and its lesser curvature then faces right. Coincident with this turning of the stomach, the intestine continues to grow in length. Growth of the intestine caudal to the stomach fills the abdomen and is probably responsible for pushing the stomach forward against the developing liver and diaphragm. The stomach 3 then lies in an oblique plane (caudodorsal to cranioventral) against the abdominal face of the diaphragm and the developing liver. Fig. 4 With growth and turning of the stomach, the dorsal mesogastrium enlarges greatly. It forms a large saclike outpouching to the left, the greater omentum. On the right side, the opening into the omental sac is the omental (L., omentum) or epiploic (Gr., epiploon) foramen. With continued growth, the liver pushes up on the right side and the omental foramen becomes fairly small. On the left side, caudal to the greater curvature of the stomach, the spleen develops in the mesenchymatous connective tissue between the two peritoneal layers of the greater omentum. The part of the greater omentum between the spleen and the greater curvature of the stomach is designated the gastrosplenic ligament. See Figure 5, below, which diagrammatically represents the development of the greater omentum and the rotation of the gut around the axis of the cranial mesenteric artery. Excepting only the inset, which shows how it develops within the greater omentum, the spleen is not shown in these diagrams; 4 Fig. 5. The upper 3 figures show the development of the gut, its rotation, and the development of the greater omentum from the left side; the lower 3 figures show this from the right side. In addition there is an inset showing how the spleen develops within the greater omentum. This occurs in the greater omentum a little caudal to the omentum’s attachment to the greater curvature of the stomach. Within the abdomen, the greater omentum exists as a flattened sac. The superficial wall of the sac lies upon the ventral body wall. The deep wall of the sac is collapsed, resting upon the superficial wall ventrally. Dorsally, it is in relation to the coils of the small intestine, chiefly, the jejunum. 5 Fig. 6 The pancreas develops from two sources: (1) as the ventral pancreatic bud from the proximal part of the hepato-pancreatic bud; and (2), as a separate tubular outpouching, the dorsal pancreatic bud, branching from the duodenum caudal to the origin of the hepato-pancreatic bud. The two parts of the pancreas join together and their branching duct systems unite and communicate. The entire pancreas is within the mesoduodenum and the contiguous part of the greater omentum. The part within the mesoduodenum is the right lobe. The left lobe extends leftward in the dorsal part of the greater omentum, caudal to the stomach. It reaches the dorsal end of the spleen. The body of the pancreas is the part at the angle of junction of right and left lobes. The portal vein (see below) always passes in relation to the body of pancreas. 6 Fig. 7. Cranial view of the stomach, showing the continuity of mesoduodenum and greater omentum, which occurs at the omental foramen. The omental foramen (upper large arrow) is relatively large here; with growth of the liver, it is much smaller. See the R and L lobes and body of the pancreas. See the portal vein. Note that the hepatic branches pass to the porta of the liver with the portal vein. They are given off from the hepatic artery as it passes ventral to the portal vein. The intestine first appears as a simple tube extending from the pylorus of the stomach to the caudal part of the embryo where the anus will form. With this simple tube, the large intestine is distinguished from the small intestine after the cecum begins to develop. The cecum forms as a blind diverticulum of the first part of the ascending colon of the large intestine. When the cecum develops, the part pyloricward is the small intestine; the part analward is large intestine, which ends at the anus. Development of the cecum is indicated in the figures on page 5. This string is around the cecum. This string is around the ascending colon. 7 Fig. 8. The left figure shows the origin of the cecum from the ascending colon. The right figure shows the protrusion of the ileum into the ascending colon, forming the ileal papilla. The mesentery suspending the intestinal tube is short cranially, where it suspends the duodenum, and short caudally where it extends to the colon. The mesentery between the two short-mesentery parts is much longer. Here the intestine forms the large loop that becomes jejunum and ileum. When intestine is suspended by a short mesentery, it is too closely attached to the body wall to do any significant amount of coiling or twisting. Thus duodenum and colon are seldom involved in a twisting of the mesentery with a shutting off of blood supply and “strangulation.” The long mesentery of jejunum-ileum does permit extensive coiling, and this part of the gut can more easily twist and “strangulate.” Also, the long mesentery permits jejunum and ileum to reach the ventral body wall. The jejunum especially is the part of the gut most frequently involved in an umbilical or scrotal hernia. Fig. 9. The long loop of jejunum and ileum is at the level of the developing cranial mesenteric artery. With growth in length and its confinement within a limited space, the body cavity, the intestine begins to rotate around the axis of the cranial mesenteric artery. (see figures on page 5) 8 The direction of rotation is normally clockwise when viewed dorsally. In rare cases, the rotation can be counter-clockwise. The rotation is gradual and the developing cranial mesenteric artery is not twisted in the rotation-process. Ultimately, the rotation is about 360 degrees (some texts say 270 degrees). At the origin of the cranial mesenteric artery, fusion of the twisting mesentery Fig. 10. Left view. Normal topography of the gut near the root of the mesentery. Note that, in Fig. 10, from the caudal flexure of the duodenum, the ascending duodenum passes forward and dorsally. Its mesoduodenum fuses with the mesocolon of the desending colon, forming the duodenocolic fold. From the cranial end of duodenocolic fusion, the duodenojejunal flexure is formed. The ascending duodenum, passes to the left of the cranial mesenteric artery (Fig. 5 shows this). Note also that the transverse colon will lie immediately caudal to the deep leaf of the greater omentum (which bears the left lobe of the pancreas). There is here a fusion of a thin slip (velum omentale) of the deep leaf with the transverse mesocolon. After rotation of the intestine, the short-mesentery duodenum and colon are each in the shape of a J. The long arm of the J of the duodenum starts on the right side, at the pylorus of the stomach. The long arm of the J of the colon is on the left side and extends cranially from the anus. The long loop of jejunum and ileum extends between the two short-mesentery parts of the intestine. 9 Fig. 11 The J-shaped duodenum (Figs. 11, 12) exhibits a short cranial part, a descending duodenum, a caudal flexure (and some add to this a “transverse part”), and an ascending duodenum. Parts of the intestine that convey the ingesta caudally are described as “descending”; parts that convey the ingesta cranially are described as “ascending.” “Transverse” parts extend from one side of the body to the other. Fig. 12 The descending duodenum is the right-most part of the gut. It extends caudally from the cranial part to the caudal flexure. The ascending duodenum passes from the caudal flexure craniodorsally to the left of the cranial mesenteric artery. 10 Its suspending mesentery, which is designated mesoduodenum, fuses with the mesocolon of the descending colon, the area of fusion being designated the duodenocolic fold. At the cranial end of the duodenocolic fold, the duodenum passes over into the jejunum at the duodenjejunal flexure. The flexure is easily recognized because it is just cranial to the fused area of the duodenocolic fold; the beginning longer mesentery of the jejunum (mesojejunum) appears at the bend. From the duodenojejunal flexure, the jejunum and ileum (the two parts of the intestine are often abbreviated as jejunoileum) pass to the ileum’s joining the ascending colon. The jejunum is much the longer part. It is thrown up into numerous coils that for the most part are ventral, separated from the ventral body wall only by the thin, collapsed sac of the greater omentum. The ileum is marked by the ileocecal fold, which extends from ileum to cecum. The ileum is also somewhat more heavily muscled than the duodenum and jejunum an adaptation to its propelling the ingesta through the ileal sphincter into the ascending colon. The J-shaped, short-mesentery colon is distinguished as ascending colon, transverse colon, and descending colon. The descending colon is continuous caudally with the rectum that joins the anus. The ileum joins the short arm of the J, which is the ascending colon. The ascending colon passes cranially from the ileocolic junction to the transverse colon. Its short mesocolon is fused in part with the mesoduodenum of descending duodenum. The mesocolon of the ascending colon is shorter than the mesoduodenum of the descending duodenum that covers it. Therefore, to see the ileocolic junction, the ascending colon, and the cecum, you have to lift up the mesoduodenum of the descending duodenum. The transverse colon is also short, extending from right to left cranial to the cranial mesenteric artery. On the left, it is continued as the long arm of the J, which is the descending colon and rectum. Within the abdomen, the descending colon is the leftmost part of the intestine. The mesoduodenum that suspends the ascending duodenum is shorter than the mesocolon of the descending colon, and fused with it to form the duodenocolic fold. Therefore to see the ascending duodenum and its attachment to the mesocolon of descending colon, you have to lift up the descending colon. Fig. 13 11 Fig. 13. The path of the gut (diagrammatic) Is all of this an esoteric academic discussion of little utility in the real world of veterinary medicine? Read these paragraphs and then decide. Get yourself in the habit of knowing this relationship that results from (1) rotation of the intestine around the cranial mesenteric artery and (2) fusion of the mesenteries to form the root of the mesentery and the duodenocolic fold. The relationship to understand: If, following your ventral median abdominal incision, you want to find an obstruction in the descending colon, you run your hand down the left abdominal wall to the place where it comes to a stop: that place is where the mesentery is reflected from the dorsal body wall. You bring your hand up with the first part of the intestine that you encounter: descending colon! Follow it cranially if you want to look at transverse colon. If you suspect (following your appreciation of the signs of illness and the radiography of the animal) that the problem is in the ascending duodenum, you simply look medial to the mesocolon of descending colon. The ascending duodenum will always be there, fused to the mesocolon of the descending colon. If you want to find the descending duodenum……or the ascending colon and cecum….. or the ileum, try this: On the left side: descending colon is most superficial, next to the body wall, and, deep to it, ascending duodenum. On the right side, descending duodenum is most superficial, next to the body wall, and deep to the mesoduodenum are ileum, ascending colon, and cecum. 12 Is there a difference between obstruction of the small intestine vs. obstruction of the large? Obstruction of the descending colon and rectum may not be detected for weeks; unless the owner checks the animal’s passage of stool or, being observant, notes the enlarging abdomen. Signs of obstruction of the small intestine are fairly immediate (unwillingness to eat and vomiting within 24 hours, usually a much shorter time) and much more critical. Essential abdominal topography: 1. Stomach, a J-shaped sac lying chiefly to the left side of the body immediately caudal to the diaphragm and liver. Cardia: where the esophagus joins the stomach; cardiac ostium, the opening between esophagus and stomach; Fundus: the blind part of the stomach dorsal to the cardia; Body: from fundus to the pyloric part; Pyloric part: is made up of the pyloric antrum, pyloric canal, and pylorus. The pyloric part is the narrow, tubular, right part of the stomach. Its wider part nearest the body is the pyloric antrum; its more narrow part that joins the pylorus is the pyloric canal. Pylorus: The constricted part of the stomach that joins the cranial part of the duodenum; pyloric sphincter, the strong smooth muscle pyloric sphincter at the pylorus; pyloric ostium, the opening in the pylorus by which the ingesta enters the duodenum. Greater curvature, lesser curvature. These curvatures extend from the cardia to the pylorus on opposing margins of the stomach. The greater omentum attaches to the greater curvature; the lesser omentum, to the lesser curvature and it extends also onto the cranial part of the duodenum. 2. The small intestine consists of duodenum, jejunum, and ileum. Its diameter is smaller than the diameter of the large intestine. Duodenum: it is comprised of cranial part, descending duodenum, caudal flexure, and ascending duodenum. The ascending duodenum is continuous with the jejunum at the duodeojejunal flexure. Cranial part: the short segment between the pylorus of the stomach and the cranial flexure. Cranial flexure: the bend between the cranial part and the descending duodenum; Descending duodenum; Major duodenal papilla. A mucosal papilla near the cranial end of the descending duodenum. The common bile duct (ductus choledochus) and pancreatic duct open here; Minor duodenal papilla. It is 3 – 6 cm caudal to the major duodenal papilla in the descending duodenum; the accessory pancreatic duct opens here; 13 Caudal flexure; Ascending duodenum; Jejunum has the longest mesentery and can be located in most parts of the abdomen. It is the part of the gut most likely to be found in a scrotal or umbilical hernia. Ileum joins the ascending colon. The ileum is a part of the large loop of jejunoileum. It can be found in a scrotal or umbilical hernia but, being shorter than the jejunum, is less likely to be involved in a herniation. The ileum is joined to the cecum by the ileocecal fold. If you incise the wall of the ascending colon opposite to where the ileum joins it, the opening of the ileum, the ileal ostium, will be seen to be on a raised area of the mucosa designated the ileal papilla. 3. The large intestine is of greater diameter than the small intestine. It consists of cecum, colon, rectum, and anus. The cecum empties into the ascending colon at the cecocolic ostium. In the dog there is a fairly distinct sphincter around the ostium. The colon is divided into ascending colon, transverse colon, and descending colon. The rectum is the straight part of the large intestine that is within the pelvis; it joins the anus. The anus is the part of the large intestine that communicates with the exterior. Note: The curvature of the colon permits a right colic flexure (between ascending colon and transverse colon) and a left colic flexure (between transverse colon and descending colon) to be distinguished. But the intestine is a flexible, plastic tube and the position of these flexures (and therefore of the boundaries of the three parts of the colon) can vary slightly. Essential gastrointestinal topography: 1. Liver is caudal to the diaphragm; stomach is caudal to the liver. 2. On the right side, the mesoduodenum with descending duodenum is most lateral, next to the right abdominal wall; medial to the mesoduodenum of the descending duodenum are the cecum, terminal ileum, and ascending colon. 3. On the left side, mesocolon with descending colon is most lateral, next to the left abdominal wall; medial to the mesocolon and fused with its medial side is the ascending duodenum. 4. Coils of jejunum (and a little of the ileum) rest upon the collapsed omental sac upon the ventral body wall (see the figure at the top of page 6). 14 5. The sac formed by the greater omentum is attached cranially to the greater curvature of the stomach and the dorsal body wall. Therefore, when meeting the omentum on a ventral abdominal incision, the omentum is drawn forward, toward its gastric attachment. If you pull it caudally, you will tear it from its gastric attachment. 6. In the usual ventral median abdominal incision, you will encounter the fat-filled remnant of the falciform ligament. You simply incise to one side of its attachment to the ventral body wall in order to enter the peritoneal cavity. Structure of the Peritoneum: The peritoneum is a membrane with an epithelial part, the mesothelium, and a connective tissue part, the lamina propria. The lamina propria attaches the peritoneum to the structures that it clothes. The peritoneum that lines the walls of the abdomen and cranial pelvis is parietal peritoneum. The diaphragm is the cranial wall of the abdomen and its caudal, abdominal, surface is covered with parietal peritoneum. Nomenclature of the Peritoneum: The peritoneum covering the stomach, intestine, liver, spleen, ovary, uterus, etc., is visceral peritoneum. The peritoneum extending between parietal peritoneum of the body wall and the visceral peritoneum covering the abdominal and pelvic organs is always given a particular name: greater omentum, mesentery (mesoduodenum, mesojejunum, etc.), mesovarium, mesosalpynx, mesometrium, right triangular ligament of the liver, etc. The peritoneum extending between parietal and visceral peritoneum is sometimes referred to in general, lay terms as “connecting peritoneum.” Function of the Peritoneum: The peritoneum, moistened with the serous fluid that its mesothelium secretes, provides a smooth, relatively frictionless surface for movement of the abdominal and pelvic viscera. The serous fluid serves as a lubricant. Every time the diaphragm contracts, there is a 15 slight movement of the viscera. Peristaltic movements of stomach and intestine are falcilitated by the lubricating effect of the serous fluid. Ordinarily, the serous fluid is present as a thin film. With inflammation of the peritoneum it is produced in much greater quantity and contains inflammatory cells and a protein-containing exudate. Liver. The liver is the largest gland in the body. Lobes of the liver: quadrate, caudate, left and right. The caudate lobe consists of a caudate process and a papillary process. Left and right lobes are each subdivided by fissures into left medial and left lateral lobes; right medial and right lateral lobes. Quadrate lobe: The quadrate lobe is between the fossa for the gall bladder and the fissure for the round ligament of the liver. The fossa for the gall bladder is the depression that seats the gall bladder. The fissure for the round ligament is the first deep fissure to the left of the gall bladder. Right lobe: to the right of the quadrate lobe and subdivided into right medial and right lateral lobes. Left lobe: to the left of the quadrate lobe and subdivided into left medial and left lateral lobes. Caudate lobe: dorsal on the caudal surface of the liver. It has prominent caudate and papillary processes. Fig. 14 Left figure is from Budras, Anatomy of the Dog, 2007; Schlütersche Verlagsgesellschaft, Hanover; right figure is from Sisson. 16 The porta and the portal circulation: The porta (also designated hilus) is the “door” to the liver. It is central on the visceral face of the live and is the place where the portal vein, hepatic branches of the hepatic artery, nerves and lymphatics enter the liver and hepatic ducts conveying bile emerge from the liver. The portal vein conveys venous blood from the stomach, intestine, spleen and pancreas to the liver. The hepatic branches of the hepatic artery convey oxygenated, arterial, blood to the liver. Branches of the portal vein and hepatic artery empty into the same vessels, the fenestrated capillaries of the liver, which are also called the hepatic sinusoids. Traversing the sinusoids, the blood is acted upon by the liver cells and intravascular macrophages. Glucose, albumin, and other constituents are added or removed to restore their concentration in the blood to normal levels. Toxins are detoxified by the hepatocytes. Bacteria are removed by the macrophages. What about venous blood from the spleen? A function of the spleen is the removal of “worn-out” red blood cells. The metabolic end-product of hemoglobin metabolism is hemosiderin, which is transported from the spleen to the liver where it is utilized in the production of bile acids that are excreted as the main constituent of bile and serve to emulsify ingested fat. What about venous blood from the pancreas? The pancreatic islets secrete glucagon and insulin, hormones that determine blood sugar levels in large part (but not entirely) by their action on the hepatocytes. From the sinusoidal capillaries of the liver, the blood is collected into venous channels that unite to form hepatic veins that empty into the caudal vena cava as it passes medial to, and partly embedded within, the right lobe of the liver. A portal venous circulation is one in which the blood passes through two sets of capillaries instead of the usual single capillary bed between artery and vein. Thus blood that has traversed a first capillary bed in stomach, intestine, spleen, and pancreas then passes in the portal vein to a second capillary bed, the hepatic sinusoids. From the sinusoids this blood is returned to the heart in the caudal vena cava. Functionally, the portal venous system restores the blood draining the digestive system and spleen to normal, homeostatic, levels before its being distributed to the rest of the body in the general circulation. The blood delivered to the liver by hepatic17 branches of the hepatic artery has, of course, traversed a single set of capillaries, the Fig. 15 Caudal vena cava. The caudal vena cava is dorsal in the abdomen, to the right of the abdominal aorta. At the level of the first lumbar vertebra, it passes cranioventrally on the medial side of the right lobe of the liver in which it is partly embedded. At the level of the foramen venae cavae of the diaphragm, it passes from the liver through the diaphragm and is continued within the thorax in the dorsal margin of the plica venae cavae. It empties into the right atrium. As it passes in relation to the liver, the hepatic veins empty into the caudal vena cava. The hepatic veins are of variable size, the largest emptying into the caudal vena cava as it departs the liver to pass in the foramen venae cavae. The bile conducting system. Bile is produced by the hepatocytes and conveyed within bile canaliculi to bile ducts that are within the liver. The bile ducts of each liver lobe are collected to form a lobar hepatic duct. In the dog, the lobar hepatic ducts discharge individually into the cystic duct, the large duct leading from the gall bladder. After all lobar hepatic ducts have joined the cystic duct, the cystic duct is continued by the common bile duct (ductus choledochus) to the duodenum. The common bile duct opens on the major duodenal papilla. The small pancreatic duct discharges with the common bile duct at the major duodenal papilla. Peritoneum and ligaments of the liver. The liver is interposed between the abdominal face of the diaphragm and the stomach. Its diaphragmatic face is apposed to the diaphragm and is in part adherent to the diaphragm. Parietal peritoneum covering the abdominal surface of the diaphragm passes onto the surface of the liver at the margin of this area of adhesion. Upon the liver, the peritoneum is visceral peritoneum. The thin line of junction of the parietal peritoneum with the visceral peritoneum, where it is at the margin of the area of adhesion, is the coronary ligament of the liver (coronary = like a crown, signifying that the line of reflection encircles the area of adhesion ”like a crown”). From the coronary ligament, the right and left triangular ligaments are duplicatures of peritoneum that extend to either side along the dorsal border of the corresponding lobe of the liver. A part of the falciform ligament is evident usually only as a short fold extending ventrally from the liver to the diaphragm close to the place where the caudal vena cava departs the liver to pass through the diaphragm. Pancreas. The pancreas consists of right lobe, body, and left lobe. The right lobe is in the mesoduodenum of the descending duodenum. From the cranial end of the right lobe, the left lobe extends leftward in the dorsal part of the deep wall of the greater omentum (the deep wall is the wall of the omental bursa that is apposed to the viscera; the superficial wall is the wall of the omental bursa in contact with the floor of the abdominal cavity). The left lobe lies within the deep 18 wall just cranial to the transverse colon; its left tip is in relation to the dorsal extremity of the spleen. The body of the pancreas is at the angle of junction of right and left lobes and is in relation to the portal vein. It is just proximal to the portal vein’s passing to the porta of the liver. The pancreatic duct opens with the common bile duct at the major duodenal papilla. The accessory pancreatic duct opens alone at the minor duodenal papilla. The pancreas is shown in the figure below. Fig. 17. The celiac artery and its branches…. Arteries supplying the gastrointestinal tract and spleen. There are only three ventral arteries of the abdominal aorta and all are involved in the blood supply to the gastrointestinal tract. The arteries are: celiac, cranial mesenteric, and caudal mesenteric. 19 Celiac artery. It arises ventral to the first lumbar vertebra. It is about 2 cm long and ends in three main branches: splenic, left gastric, and hepatic. Splenic and left gastric often come off by a short common trunk. The splenic artery is large. It passes left in the dorsal part of the greater omentum, dividing into two branches that are united by an arciform vessel that passes along the hilus of the spleen. The more ventral of the two branches, or the arcade, gives off the left gastroepiploic artery that runs in the greater omentum along the left ventral part of the greater curvature of the stomach. This vessel anastomoses with the right gastroepiploic artery. Short gastric branches of the splenic artery pass to the greater curvature of the stomach and pancreatic branches supply the left lobe of the pancreas. The left gastric artery passes cranioventrally, then to the right along the lesser curvature of the stomach. Its branches supply the esophagus and lesser curvature of the stomach. It anastomoses along the lesser curvature with the right gastric artery. The hepatic artery inclines right-cranioventrally in the dorsal part of the deep wall of the greater omentum. It crosses the cranial surface of the left lobe of the pancreas, reaches the portal vein as the portal vein passes through the body of the pancreas, and continues to the right, passing ventral to the portal vein. As it passes ventral to the portal vein, it gives off hepatic branches to the liver---these pass to the porta of the liver with the portal vein. With the portal vein, the hepatic branches lie within the lesser omentum. The hepatic artery ends by dividing into the small right gastric artery and the large gastroduodenal artery. The right gastric artery passes right along the lesser curvature of the stomach and unites with the left gastric. The gastroduodenal artery gives off the cranial pancreaticododenal artery onto the mesenteric border of the descending duodenum and the right gastroepiploic artery, which passes caudal to the duodenum at the cranial flexure. The right gastroepiploic artery passes left within the greater omentum along the omental attachment to the greater curvature of the stomach. It anastomoses with the left gastroepiploic artery of the splenic. Cranial mesenteric artery. It arises from the aorta about 1 cm caudal to the origin of the celiac. It passes ventrally with the mesentery, its proximal branches passing to opposite ends of the gut: caudal pancreaticoduodenal artery to the duodenum; ileocolic artery to ileum, cecum and colon. The continuing cranial mesenteric artery passes within the mesojejunum, giving off jejunal arteries to either side and one or two to the ileum. The caudal pancreaticoduodenal artery passes in the mesoduodenum to the ascending duodenum and caudal flexure, and anastomoses with the cranial pancreaticoduodenal. The ileocolic artery passes in the mesocolon to the transverse and ascending colon, to the cecum and to the ileum. The first branch of the ileocolic is almost always the middle colic artery, which supplies the transverse colon and forms anastomotic arcades with the right colic artery that next proceeds from the ileocolic. A left branch of the middle colic passes along the mesenteric border of the descending colon. It 20 anastomoses with the left colic artery, which is a branch of the caudal mesenteric. The middle colic sometimes arises directly from the cranial mesenteric. Note: In the writer’s opinion, it is clearest, and correct, to define the middle colic artery always as the branch of the ileocolic (or sometimes of the cranial mesenteric artery directly) that anastomoses with the left colic artery branch of the caudal mesenteric. Fig. 18. Right view at root of the mesentery. Proximal branches of the cranial mesenteric artery; portal vein entering the porta of the liver; the omental (epiploic) foramen. The second and third (sometimes fourth also) branches of the ileocolic are the single right colic artery and one or two colic branches. These vessels pass to the mesenteric border of the ascending colon. The ileocolic ends by dividing into a small mesenteric ileal artery, which passes along the mesenteric border of the terminal ileum, and a large cecal artery, which crosses the medial (deep) side of the ileocolic junction. The cecal artery enters the ileocecal fold and gives off branches to both the cecum and the terminal ileum. After the last cecal branch is given off, the continuation of the cecal artery as the antimesenteric ileal artery passes within the “tail” of the ileocecal fold upon the antimesenteric border of the ileum. Note: There is always a small unnamed branch of the cecal artery that passes on the lateral (superficial) side of the ileocolic junction. This branch anastomoses with the 21 cecal artery within the ileocecal fold. The result is a vascular ring that surrounds the ileocolic junction. Caudal mesenteric artery. This artery arises from the aorta about at the level of the fifth lumbar vertebra. It inclines caudally within the mesocolon and ends at the mesenteric border by dividing into a cranially directed left colic artery and a caudally directed cranial rectal artery. Colic lymph nodes are at the place of division. The left colic artery passes cranially along the mesenteric border of the descending colon and anastomoses with the middle colic artery of the ileocolic (or of the cranial mesenteric). The cranial rectal artery passes caudally on the rectum, anastomosing with branches of the middle and caudal rectal arteries. Fig. 19. Union of the cranial and caudal mesenteric veins to form the portal vein. Tributaries of the portal vein; veins satellite to the branches of the celiac, cranial mesenteric, and caudal mesenteric arteries. With a few exceptions, veins are satellite to branches of the celiac, cranial mesenteric, and caudal mesenteric arteries; but, as explained below, no vein accompanies the main trunk of the celiac, cranial mesenteric, and caudal 22 mesenteric arteries to the caudal vena cava. Instead the large venous trunks are collected to drain into the portal vein. The caudal mesenteric vein arises from cranial rectal veins and, as a continuous trunk, passes cranially alongside the left colic artery, receiving left colic veins that proceed from the descending colon. Near its union with the cranial mesenteric vein, it receives the ileocolic vein. The cranial mesenteric vein accompanies the distal (jejunal) trunk of the cranial mesenteric artery. It collects jejunal and ileal veins, receives the caudal pancreaticoduodenal vein and joins the caudal mesenteric vein near the root of the mesentery to form the portal vein. From the root of the mesentery, the portal vein arches cranially over the dorsal aspect of the transverse colon and the body of the pancreas. On the left, it receives the large splenic vein. Near the porta of the liver it receives the gastroduodenal vein ventrally and, joined by hepatic branches of the hepatic artery passes in the lesser omentum to the porta of the liver. The splenic vein accompanies the splenic artery. It receives the left gastric vein and passes to the portal vein within the dorsal part of the deep wall of the greater omentum. No vein is satellite to the main trunk of the hepatic artery; it is its hepatic branches that accompany the portal vein to the liver. The gastroduodenal vein receives cranial pancreaticoduodenal and right gastroepiploic veins. It joins the portal vein about 1 - 2 cm before the portal vein enters the porta of the liver. Main thing to bear in mind: The portal vein is formed at the union of the cranial mesenteric and caudal mesenteric veins. It then passes dorsal to the transverse colon and turns cranioventrally to reach the porta of the liver. In passing to the porta, the portal vein with the hepatic artery and its hepatic branches lies in the lesser omentum, which forms the ventral margin of the omental foramen. Thus, when your finger is in the omental foramen, it is ventrally in contact with the lesser omentum with the portal vein, the hepatic artery and its branches to the liver. Note: The omental foramen is small after the liver is fully developed. It lies always ventral to the caudate lobe of the liver. The ventral boundary of the foramen is formed by the lesser omentum as explained above. The dorsal boundary of the foramen is the caudate lobe and the caudal vena cava, which passes in relation to the medial side of the caudate lobe. Nerves supplying the abdominal viscera. A brief introduction to nervous system function: Receptors of the body are sensitive to various specific stimuli (heat, cold, pH, electromagnetic radiation of 23 the visible spectrum, etc.). Stimulation of these receptors results in the production of nerve impulses that pass by way of sensory, afferent, neurons to the central nervous system (CNS), the brain and spinal cord. Within the brain and spinal cord, the conduction of these impulses is directed into pathways provided by interneurons (neurons that begin and end within the CNS). The interneuronal pathways modulate the excitation arriving by way of the afferent neurons in that, within the CNS, interneurons act to inhibit or facilitate the ongoing passage of excitation. From the CNS there is an outflow of impulses resulting in the contraction of muscle and the secretion of glands. “Activities” that we recognize as thinking, consciousness (awareness of the stimulus), dreaming, attention, etc. take place within the CNS itself. They are part of the circuitry within the CNS and influence the outflow of impulses that result in the contraction of muscle and the secretion of glands. The outflow from the CNS is by way of efferent nerves that stimulate muscle or cause glands to secrete. The body thus responds to stimuli by contraction of muscle and secretion of glands. The outflow from the CNS to striated muscle fibers is always by way of a single neuron that has its dendrites and cell body within the CNS. The outflow to cardiac muscle, smooth muscle, and glands is called the autonomic nervous system. For the most part, it occurs by way of a chain of two neurons. The cell body and dendrites of the first neuron of the chain are in the CNS; the cell body and dendrites of the second neuron of the chain are in a peripheral ganglion: cervicothoracic ganglion, middle cervical ganglion, etc. Some gangia are microscopic and lie within the walls of the organ innervated. Fig. 20. Diagrammatic representation of the nervous system. CNS, brain and spinal cord INPUT: afferent neurons from receptors carrying impulses to the CNS OUTPUT: Efferent neurons pass to striated muscle or to 24 effectors autonomic (heart muscle, smooth muscle, Interneurons within the CNS, some excitatory, some inhibitory. Interneuronal pathways ultimately lead to efferent neurons. 2 neuron chain to autonomic effectors single neuron to striated muscle fibers The abdominal viscera, containing no striated muscle but having lots of smooth muscle and glands are innervated by the autonomic nervous system. They also receive sensory fibers which, though they travel with autonomic fibers, are not regarded as part of the autonomic nervous system. The autonomic nervous system is in two divisions, sympathetic and parasympathetic which often have opposing effects on the cells innervated. For example, sympathetic fibers relax smooth muscle of the intestinal wall; except for its sphincters, which they stimulate. Parasympathetic fibers generally have the reverse effect. At the origin of the celiac and cranial mesenteric arteries are the celiacomesenteric ganglia and plexus. Autonomic nerves passing to the ganglia and plexus are (1) the greater and lesser splanchnic nerves, which contain sympathetic and sensory fibers; and (2) a branch from the dorsal vagal trunk, which contains parasympathetic and sensory fibers. From the gangia and plexus, perivascular nerve fibers extend in plexuses that pass with the branches of these two major arteries. It is in that way that innervation reaches the smooth muscle and glands of the abdominal viscera with sympathetic and parasympathetic fibers. Sensory fibers are also in the perivascular plexus. They supply the viscera with sensory fibers. On the cranial face of the caudal mesenteric artery is the caudal mesenteric ganglion. It receives lumbar splanchnic nerves, which contain sympathetic and sensory fibers. The cells of the ganglion give rise to R and L hypogastric nerves that pass to the pelvis in the mesorectum. Parasympathetic and sensory fibers pass in ventral branches of the second and third sacral spinal nerves and unite to form the pelvic nerve. In the dog, the pelvic nerve passes ventrally just a little caudal to the prostatic/vaginal vessels. Within the pelvis, fibers in the hypogastric nerve mix with fibers in the pelvic nerve, forming the pelvic plexus. This plexus provides sympathetic, parasympathetic and sensory fibers to the pelvic viscera. It 25 is generally described as extending cranially in the mesocolon to the descending colon and transverse colon. The anus. This important part of the alimentary canal is subject to direct visual inspection and palpation. External anal sphincter: this is the striated muscle that gives voluntary control over the passage of stool. It is well developed. It is innervated by the caudal rectal nerve, a branch of the pudendal nerve. Internal anal sphincter: a smooth muscle sphincter that is not well defined on gross inspection. It is deep to the external sphincter. Paranal sinuses (“anal sacs”). These are R and L saclike pouches, about one centimeter in diameter situated between the internal and external anal sphincter.. Each sinus stores the secretion of its glandular walls and discharges by a single duct that is ventrolateral to the mucocutaneous junction of the anal opening. The paranal sinuses function as marking devices by which the animal designates its “territory.” The sacs are subject to infection and impaction and are sometimes surgically removed. In the cutaneous area immediately surrounding the anal opening are circumanal glands that may become tumorous. Fig. 21. This figure is from Ellenberger – Baum, 1943; Springer Verlag. 26