The Digestive System Lecture 1

Bill Sellers mailto:Bill.Sellers@ed.ac.uk November 30, 1998

Contents
1 Introduction 2 Objectives 3 General Characteristics 4 The Mouth 4.1 Teeth . . . . . 4.2 Tongue . . . . 4.3 Swallowing . . 4.4 Salivary Glands 5 Oesophagus 6 Stomach 7 The Small Intestine 7.1 Duodenum . . . . . . . . . . . . . 7.2 Jejunum and Ileum . . . . . . . . 8 Pancreas 9 Liver 10 Gall Bladder 11 Large Intestine 11.1 Caecum . . . . . . . . . . . . . . 11.2 Colon . . . . . . . . . . . . . . . 11.3 Rectum . . . . . . . . . . . . . . 12 Anus 1 1 2 3 4 4 5 5 5 6

13 Portal Circulation

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1 Introduction
The purpose of these lectures is to describe the structures involved in digestion and to relate these structures to their functions. The information I will provide will follow very closely what is given in Hole and Koos, chapter 12, but because my research interest is in the interaction of form and function over the last 100 million years or so, I will also try to provide an evolutionary context for some of these observations.

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2 Objectives

8 The objectives of these three lectures are as follows 8 (taken straight from the textbook): 8 9 10 10 11 11 11 12 12 1 1. Name and describe the location of the organs of the digestive system and their major parts. 2. Describe the structure and general function of each digestive organ and the liver. 3. Describe the structure of the wall of the alimentary canal. 4. Explain how the contents of the alimentary canal are mixed and moved. 5. Describe the mechanism of swallowing.

6. Explain how the products of digestion are absorbed.

3 General Characteristics
The major components of the digestive system that we will cover can be seen in gure 1. As you can see, the digestive system in humans is basically a long tube and associated glandular structures. Food is ingested cranially and processed incrementally as it passes caudally. This processing is both mechanical and chemical, leading to the absorption of some of the chemical components of the food items whilst other, indigestible components are passed straight through. You will probably already be familiar with the basic structure and the location of the major components, but we will cover each organ in detail. The muscular tube through which the food passes is called the alimentary canal. The exact histological appearance of this tube varies down the length of the tube, but it basically consists of four concentric layers (see gure 2). The innermost layer is called the mucosa. This itself consists of three thin layers: an epithelium lining the inner surface, some underlying connective tissue (lamina propria), and a small amount of smooth muscle, confusingly named the muscularis mucosae. The next layer is the submucosa. This is loose connective tissue containing blood and lymph vessels and nerves. The next layer is a thicker muscular layer called the muscularis. In most regions this consists of two layers of smooth muscle: the inner one with bres arranged circularly, and the outer one with bres arranged longitudinally. Rhythmic contraction of these muscles pushes the contents of the tube steadily caudally. This movement is called peristalsis. This mechanical arrangement of bres running perpendicularly is very common in biological tubes whether it is hydraulic skeletons in roundworms or or armadillo penises. The precise angulation of the bres depends on the exact mechanical properties required. 2

Figure 1: Overview of digestive tract

Figure 2: Idealized cross-section of gut tube

The outermost layer of the the gut tube is a thin covering of visceral peritoneum - very similar in structure to the pleura round the lungs. It is called the serosa and consists of an outer epithelium with connective tissue underneath. Movement in the gut tube is not continuous. At points, food is not moved, but kept in one place and mixed. Obvious places where this occurs are in the mouth and stomach. Peristaltic movements are largely intrinsic to the smooth muscle of the gut tube, however there is both sympathetic and parasympathetic nerve supply. The parasympathetic supply (rest and digest) tends to increase peristaltic frequency and increase secretion of enzymes and mucus. The sympathetic supply tends to decrease inhibit this and also contracts the arterioles, reducing the local blood supply. The sympathetic supply also controls directly specic muscular structures called sphincters. These are regions of thickened circular muscle that block the tube when contracted, preventing the movement of gut contents. The anal sphincter is slightly more complicated, with a contribution from skeletal muscle allowing a degree of conscious control of defecation. 3

Figure 3: The mouth

4 The Mouth
The mouth (see gure 3) is extremely complicated anatomically because it has to cope with a variety of different functions. Speaking, eating, breathing are the obvious three, but certainly in humans and other primates it is also an important manipulation and tactile organ. It is also important in non-verbal communication for display and sexual behaviour. Humans and other so-called haplorhine primates are unusual among mammals in that the sides of the upper lip have fused which means that we do not have the wet, dog-like noses present amongst most mammals. The most important digestive role of the mouth is mastication. This is the physical breakdown of food by chewing. It is a complex process requiring interactions between the highly muscular tongue, cheeks and lips to position the food bolus between the teeth, and strong crushing and shearing movements of the tooth surfaces controlled by the mus-

cles of mastication pulling on the jaws. The exact conguration of jaws and teeth is highly variable amongst mammals, and is often used as a means of classifying different mammalian groups. Humans, as omnivores, do not have specializations for specic food items. This means that they have very loose temporomandibular joints, and a great deal of movement exibility (depression, elevation, protrusion, retraction, circumduction, left and right deviation). Mammals with more specic diets have more specialized dentition and restricted jaw movements. Carnivora, for example, can only depress and elevate to any great extent and have scissor like teeth for shearing food. Individual teeth also vary in shape and function.

formation into a bolus may be more important than the classic explanation of increasing surface area. It is likely that our digestive system can cope with large particle sizes, but we all know how difcult eating dry cream-crackers is.

4.2 Tongue
The tongue has several important functions in eating. Most obviously, the tongue contains the taste receptors. These help identify the food value of what has been put into the mouth. Things that taste pleasant are likely to be nutrition, and things that taste unpleasant are likely to be poisonous. Things that are eaten, whether they are plants or animals, are involved in an escalating evolutionary arms race which their predators. In the case of plants, many species produce various toxic and otherwise unpalatable compounds to reduce their desirability as food items. Interestingly, at least two species, humans and common chimpanzees have realized that these compounds can have valuable medicinal effects and can be considerably more toxic to various parasitic organisms than to the host, and they actively seek out certain plants when they feel unwell. In addition, many of the chemicals that give certain spices a hot taste are actually plant toxins. The other major functions of the tongue are mechanical. It moves the bolus of food between the teeth whilst chewing, and in swallowing it moves the food to the back of the buccal cavity to initiate swallowing. The tongue contains three sets of intrinsic muscles with bres that run orthogonally in the three anatomical directions (proximodistally, transversely and supero-inferiorly). These allow the tongue to atten or elongate. There are also extrinsic muscles that attach the tongue to the lower jaw and throat which allow it to move around within the mouth (pro- and retraction, lateral deviation, elevation and depression). It can also roll within the mouth. 4

4.1 Teeth
Humans have a dental formula of 2:1:2:3 . This 2:1:2:3 means that an adult human has 2 incisors, 1 canine, 2 premolars and 3 molars on their upper jaw, and the same on their lower jaw. Commonly, the 3rd molar may be missing, or may never erupt. Incisors are single cusped, sharp edged cutting teeth used for biting chunks off a larger food item. They are also used for removing the skins of fruit. Human canines are extremely small compared to most other primates, although they are still classically pointed. In carnivores, canines are used to immobilize prey, but in most primates they are more important for communication (aggression and as a secondary sexual characteristic). In humans, their role is largely as an accessory incisor. The premolars are bicuspid, with attened surfaces and the molars generally have 4 cusps on the upper jaw and 5 on the lower jaw although there is a certain amount of variation here. These so called cheek-teeth are where most of the chewing takes place. A combination of occlusive, puncture-crush forces and side-to-side shearing forces reduces the size of the food particles so that the surface area is increased to help digestion, and to produce a soft, deformable, moist, lubricated bolus that can be easily swallowed and moved by peristalsis. This

4.4 Salivary Glands

Salivary glands in the mouth secrete saliva (spit) (see gure 5). This uid moistens food and contains enzymes that start off starch digestion. It also contains a variety of other chemicals with antibacterial and anti-inammatory functions. There are many small salivary glands lining the tongue, cheeks and palate. In addition there are three pairs of large, discrete glands: the submandibular, parotid and sublingual glands. Salivary glands contain two sorts of secretory cells: serous cells that secrete a watery uid containing digestive enzymes; and mucous cells that secrete a viscous uid called mucus whose major role is to stick food particles together to form the bolus. In section, the serous cells stain strongly with H&E, but the mucous cells remain largely unstained (gure 6). The parotid and submandibular glands form a branching tree structure around their ducts, and saliva is secreted through a single duct per gland. The parotid duct enters the buccal cavity near the upper second molar and the submandibular duct opens underneath the tongue quite close to the midline. The sublingual glands have many, separate small ducts. Problems with saliva production can lead to the formation of salivary stones which can block the ducts and need to be removed surgically.

Figure 4: Sagittal section of mouth and pharynx

4.3 Swallowing
Swallowing is a complex series of movements that moves food from the buccal to the oesophagus (see gure 4). It is made more complicated by the dual role (respiration and digestion) of a number of the structures in this region it is important that food does not fall into the trachea and the lungs and also important that it does not get pushed up into the nasopharynx. During swallowing the tongue pushes the bolus backwards into the oropharynx. Other muscles raise the soft palate and this action closes off the nasopharynx. The back of the tongue and muscles in the walls of the pharynx start to push the bolus downwards by peristalisis and this action pushes the epiglottis, a sprung, cartilagenous ap, downward so that it covers the entrance to the trachea whilst the food passes downwards to the oesophagus. The epiglottis does not produce a perfect seal and choking occurs as a reex reaction of particles of food do accidentally fall into the trachea. 5

5 Oesophagus
The oesophagus is a straight, muscular tube connecting between the pharynx and the stomach. It descends through the thorax behind the trachea and anterior to the thoracic aorta. The vagus nerves exit the thoracic cavity through the diaphragm next to the oesophagus. The wall of the oesophagus is muscular (gure 7), and mucous glands in the mucosa serve to keep the stratied, squamous epithelia moist and lubricated. In the resting state the oesophagus is deeply folded and it distends greatly to allow the passage of food. At the oesophageal hiatus (the hole in the di-

Figure 7: Cross-section of the oesophagus aphragm), the muscular wall of the oesophagus is thickened. These bres are usually contracted to Figure 5: Diagram showing the locations of the prevent reux of stomach contents. This action is helped by a local thickening of muscle in the dimajor salivary glands aphragm which acts externally on the oesophagus to help prevent both regurgitation and the physical movement of the top of the stomach into the thorax.

6 Stomach
The stomach is a large, classically J-shaped, baglike organ that hangs under the diaphragm on the left-hand side (gure 8). It has a capacity of about one litre, although, this can increase markedly. In humans the stomach is used to mix food with gastric juices, initiate the digestive process and store food before passage to the small intestine. In many mammals, notably ruminants, but also some leaf eating primates, it is also a site for the fermentation of vegetable matter. This means that the stomach contains a large number of micro-organisms that are used to help break down otherwise indigestible vegetable matter and neutralize toxins. This role can be enhanced by having a multi-chambered stomach that helps maintain optimal fermentation conditions, and in some cases allows controlled reux so that food can be chewed repeatedly. The stomach can be divided into a number of re6

Figure 6: Diagram of a salivary gland

Figure 8: The stomach gions. The cardiac region is near the oesophageal opening. The fundus is the area just above this opening that is often seen containing air on abdominal X-rays. The pylorus is the area near the exit of the duodenum. This contains the powerful pyloric sphincter which is thickening of the muscle wall that prevents reux from the small intestine back into the stomach. The rest (and largest part) of the stomach is the body. The stomach wall consists of a thick mucosa that Figure 9: Diagram showing the variations in the contains gastric pits (gures 9 and 10). These stomach wall are tubular gastric glands that in general contain three types of secretory cells. Mucous cells secrete mucus, chief cells secrete digestive chemicals and parietal cells secrete a strong acid. This combined product is the aforementioned gastric juice. The role of the acid is to kill bacteria present in the food. The digestive chemicals start breaking down large food molecules into smaller component molecules that can be absorbed, and the mucus helps prevent damage to the stomach wall by lubricating against mechanical damage, and neutralizing the effects of both the acid and digestive chemicals. The stomach is a very muscular organ. It has three layers of muscle in the muscularis rather than the normal two. In addition to the normal exterior Figure 10: Cross-section of stomach wall longitudinal muscle, and the deeper circular muscle, it has an inner layer of oblique muscle. These 7

muscles are used to churn the food to help the mixing of food with gastric juice which produces a mixture called chyme. A small amount of this is squirted at intervals through the pylorus into the small intestine.

7 The Small Intestine

The small intestine is a tubular long, tubular structure running from the exit of the stomach to the beginning of the large intestine. It is mostly concerned with continued food digestion and absorption of the products of digestion. It is divided into a number of sections.

Figure 11: The small intestine

7.1 Duodenum
The rst section of the small intestine is the duodenum. This is a short (about 25cm long) C-shaped tube that runs horizontally to the right from the exit of the stomach, wraps itself around the head of the pancreas, passing over the top of the right kidney and then passes to the left before becoming the next section of the small intestine (gure 11). It is attached to the posterior abdominal wall rather that being suspended on a fold of peritoneum like most of the rest of the abdominal alimentary canal. This condition is known as retroperitoneal. The duodenum is histologically distinct from the rest of the small intestine because it contains extra (Brunners) glands that secrete an alkaline mucus that is used to neutralize the acidic chyme that is produced in the stomach. It is also the region that receives large ducts from the major digestive organs: the liver and the pancreas.

Figure 12: Diagram showing the variation between different areas of the small intestine to the abdominal wall and the gut tube acts rather like a hose-pipe left underneath a carpet. At intervals the length if this hose is such that it loops away from the oor taking the covering carpet with it. So, the jejunum and ileum, several metres long, hang in loops in the abdominal cavity, suspended by a mesentery which consists of a double layer of peritoneum which is continuous with the visceral peritoneum surrounding the tube and with the visceral peritoneum attached to the posterior abdominal wall. The blood and nerve supply to and the lymph drainage from the loops of gut runs between these two folds. 8

7.2 Jejunum and Ileum

The next two sections of the small intestine are the jejunum and the ileum. There is little real distinction between the two parts the jejunum is simply the proximal 2/5ths of this section (gure 12). The jejunum and ileum are suspended by a fold of peritoneum. The parietal peritoneum is attached

Figure 13: Cross section of ileum The role of the small intestine is digestion and absorption (gure 13). The gut wall consists of millions of small, nger-like projections called villi. Between the villi are intestinal glands which contain mucus secreting goblet cells. The villi contain a rich blood supply for transporting away digestion products, and also blind-ended lacteals which drain into the lymph vessels which also carry away digestion products. The internal wall of the small intestine is raised in circular folds called plicae circulares. These help to increase the surface area available for absorption. The villi further increase the surface area and the cells in the columnar epithelium that line the villi have microvilli on their lumenal surface again increases the surface area available for absorption. The small intestine is an extremely active tissue. The constant movement of intestinal contents and the energetic constraints of food absorption mean that cells are quickly lost from the tips of the villi. There is a constant migration of cells from the intestinal glands (also known as crypts) towards the 9

Figure 14: Diagram showing the relations of the liver and pancreas tips of the villi, so that within the crypts, there are often signs of active cell division. The small intestine ends at the ileocaecal valve which is a sphincter that separates the contents of the small intestine from the large intestine.

8 Pancreas
The pancreas, in the context of this series of lectures, is an organ for the production and secretion of digestive enzymes (gure 14) this is its exocrine function. It also has an important endocrine role which will be dealt with elsewhere. It is a long, thin organ that sits underneath the stomach with its head region closely associated to the interior of the C of the duodenum. The pancreatic duct runs the length of the body of the pancreas and merges with the common bile duct before entering the duodenum. The contents of the duct is pancreatic juice which is a secretion produced by acinar cells (gure 15).

Figure 16: Diagram showing the structure of the liver Figure 15: Section of pancreas

9 Liver
The liver is the largest gland in the body (gure 14). This is a somewhat misleading statement since many of its functions are not glandular. However, it is an extremely large and important organ. You dont live long with no liver! The liver takes up the space on the right-hand side that is occupied by the stomach on the left. It is rmly pushed up against the right dome of the diaphragm where the inferior vena cava, which passed through the liver, enters the thoracic cavity. It is held in place by folds of peritoneum. This peritoneal folding produces the falciform ligament anteriorly which attaches to the ventral abdominal wall and the coronary ligament superiorly which attaches to the diaphragm. The liver itself is divided into four lobes, with the left and right lobe being by far the largest, with the caudate and quadrate lobes being small lobes near the vena cava and the gall bladder respectively. Microscopically, the liver is organized into hepatic lobules. These consist of a small central vein with columns of hepatic cells radiating outwards from this central point (gures 16 and 17). Blood ows radially inwards towards the central vein between the columns from both hepatic arteries and branches of the hepatic portal vein. The hepatic portal vein drains the portal venous sys-

Figure 17: Section of liver tem and contains high concentrations of digestion products that are processed by cells in the liver (hepatocytes). Also present in the lobules are hepatic macrophages that engulf most of the bacterial cells that have entered through the intestinal wall. Also within the lobules are bile canals. These are ne tubes that drain the secretory product bile out to the periphery of the lobule where the tubes unite to form hepatic ducts which eventually unite to form the common hepatic duct that carries bile out of the liver.

10 Gall Bladder
The gall bladder is a small sac-like container that attaches by a short tube (the cystic duct) to the common hepatic duct (gure 14). Once the com-

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the lower right quadrant of the abdomen. In humans it is rather insignicant, but in many other mammals it is the alternative site for fermentation of food. When this is the case, it can be larger than the stomach. In humans, a small, blind-ended tube hangs off the caecum. This is the vermiform1 appendix which has no digestive role, although it may have a role as part of the lymphatic system in ghting off infection.

11.2 Colon
The colon is a long tube that functions as the main area for water and electrolyte absorption. In many cases, this is actually re-absorption since large quantities of both are secreted into the gut higher up in the alimentary canal to aid digestion. The colon is the region where these substances are recovered. Descriptively, it is divided into four regions. 1. The ascending colon is the rst part of the colon which ascends vertically from the caecum in the lower right quadrant up towards the liver in the upper right quadrant. It is retroperitoneal. 2. The transverse colons hangs down from a complex mesentery more or less horizontally from the end of the ascending colon in the upper right quadrant to the descending colon in the upper left quadrant. 3. The descending colon runs retroperitoneally, vertically down from the upper left quadrant to the lower left quadrant. 4. The sigmoid colon is a short length of colon that is once again attached to a mesentery that connects the end of the descending colon to the rectum. It is mostly contained within the pelvic cavity.
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Figure 18: The large intestine mon hepatic duct has joined the cystic duct it is renamed the common bile duct which continues to join with the pancreatic duct just before it empties into the duodenum. The bile duct stores the bile produced by the liver, and it is able to contract to force bile into the duodenum as required by digestion.

11 Large Intestine
The large intestine is the last part of the alimentary canal. The large in the name refers to its diameter rather than its length since the small intestine is longer in humans. The large intestine consists of three functionally distinct parts, all those these are further subdivided for descriptive purposes. These major divisions are the caecum, the colon and the rectum (gure 18).

11.1 Caecum
The caecum is a sac-like structure at the beginning of the large intestine, situated retroperitoneally in 11

Vermiform means worm-like.

Histologically, the wall of the colon contains a thick mucosa containing many tubular, mucus secreting glands (gure 19). Absorption occurs primarily in the proximal half of the colon. The mucus is mainly lubricating and protective to allow the easy movement of the gut contents distally.

11.3 Rectum
The nal region of the large intestine is the rectum. This is a short tube that lies next to the sacrum within the pelvic cavity (gure 20).

12 Anus
The external opening of the rectum is through the anus. This forms the last 2 to 4 cm of the alimentary canal. The mucous membrane lining the anal canal is folded into a series of 6 to 8 longitudinal anal columns. Free movement of faecal matter through the anal canal is prevented by a series of ring-like sphincters. The internal anal sphincter is a ring of smooth muscle which is under involuntary control. The external anal sphincter is skeletal muscle under voluntary control. retroperitoneal.

13 Portal Circulation
The blood supply of most of the digestive system is entirely normal blood ows in from nearby arteries and is drained into nearby veins. However, the venous drainage for the parts of the gut involved in substantial amounts of absorption of digestion products have a different venous drainage (gure 21). Normal venous drainage where the blood is conveyed directly back to the right atrium of the heart is referred to as systemic. Venous drainage from the gut drains rst to the liver, and only after that does it pass to the heart. This drainage from one organ system directly to another is referred to as the portal system. The reason for this separate drainage is that the composition 12

Figure 19: Cross-section of colon

Figure 20: The rectum and anus of blood is greatly altered by digestion products, and these products need to be processed before the blood can be allowed to mix back with the rest of the blood supply. This processing occurs within the liver, so the liver has two blood inputs: the hepatic artery which supplies oxygenated arterial blood that provides the liver with the oxygen that it needs; and the hepatic portal vein which is deoxygenated, venous blood drained from the distal oesophagus, the stomach, the duodenum, jejunum, ileum, the caecum, appendix, colon and the proximal part of the rectum. It also drains blood from the spleen, the pancreas and the gall bladder. There are some areas where the portal and systemic system anastamose. These are generally not important, but in certain situations (notably portal hypertension) they can open up to allow an alternative route for blood draining the gut. These include the junctions between the areas of the oesophagus and the rectum that are drained by portal and systemic veins, and umbilical vein remnants left over from the fetal circulation.

Figure 21: Diagram of the hepatic portal system

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