5-2-6. The Small and Large IntestinesLearning Objectives
The word intestine is derived from a Latin root meaning “internal,” and indeed, the two organs together nearly fill the interior of the abdominal cavity. In addition, called the small and large bowel, or colloquially the “guts,” they constitute the greatest mass and length of the alimentary canal and, with the exception of ingestion, perform all digestive system functions. The Small IntestineChyme released from the stomach enters the StructureThe coiled tube of the small intestine is subdivided into three regions. From proximal (at the stomach) to distal, these are the duodenum, jejunum, and ileum (Figure 1). The shortest region is the 25.4-cm (10-in) Of particular interest is the
Small Intestine
The The Parasympathetic nerve fibers from the vagus nerve and sympathetic nerve fibers from the thoracic splanchnic nerve provide extrinsic innervation to the small intestine. The superior mesenteric artery is its main arterial supply. Veins run parallel to the arteries and drain into the superior mesenteric vein. Nutrient-rich blood from the small intestine is then carried to the liver via the hepatic portal vein. HistologyThe wall of the small intestine is composed of the same four layers typically present in the alimentary system. However, three features of the mucosa and submucosa are unique. These features, which increase the absorptive surface area of the small intestine more than 600-fold, include circular folds, villi, and microvilli (Figure 2). These adaptations are most abundant in the proximal two-thirds of the small intestine, where the majority of absorption occurs. Circular folds Also called a plica circulare, a Villi Within the circular folds are small (0.5–1 mm long) hairlike vascularized projections called Microvilli As their name suggests, Intestinal Glands In addition to the three specialized absorptive features just discussed, the mucosa between the villi is dotted with deep crevices that each lead into a tubular The submucosa of the duodenum is the only site of the complex mucus-secreting The roles of the cells in the small intestinal mucosa are detailed in Table 1. Table 1. Cells of the Small Intestinal Mucosa
Intestinal MALT The lamina propria of the small intestine mucosa is studded with quite a bit of MALT. In addition to solitary lymphatic nodules, aggregations of intestinal MALT, which are typically referred to as Peyer’s patches, are concentrated in the distal ileum, and serve to keep bacteria from entering the bloodstream. Peyer’s patches are most prominent in young people and become less distinct as you age, which coincides with the general activity of our immune system.
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Watch this animation that depicts the structure of the small intestine, and, in particular, the villi. Epithelial cells continue the digestion and absorption of nutrients and transport these nutrients to the lymphatic and circulatory systems. In the small intestine, the products of food digestion are absorbed by different structures in the villi. Which structure absorbs and transports fats? Mechanical Digestion in the Small IntestineThe movement of intestinal smooth muscles includes both segmentation and a form of peristalsis called migrating motility complexes. The kind of peristaltic mixing waves seen in the stomach are not observed here. If you could see into the small intestine when it was going through segmentation, it would look as if the contents were being shoved incrementally back and forth, as the rings of smooth muscle repeatedly contract and then relax. Segmentation in the small intestine does not force chyme through the tract. Instead, it combines the chyme with digestive juices and pushes food particles against the mucosa to be absorbed. The duodenum is where the most rapid segmentation occurs, at a rate of about 12 times per minute. In the ileum, segmentations are only about eight times per minute (Figure 3).
Segmentation
When most of the chyme has been absorbed, the small intestinal wall becomes less distended. At this point, the localized segmentation process is replaced by transport movements. The duodenal mucosa secretes the hormone The ileocecal valve, a sphincter, is usually in a constricted state, but when motility in the ileum increases, this sphincter relaxes, allowing food residue to enter the first portion of the large intestine, the cecum. Relaxation of the ileocecal sphincter is controlled by both nerves and hormones. First, digestive activity in the stomach provokes the Chemical Digestion in the Small IntestineThe digestion of proteins and carbohydrates, which partially occurs in the stomach, is completed in the small intestine with the aid of intestinal and pancreatic juices. Lipids arrive in the intestine largely undigested, so much of the focus here is on lipid digestion, which is facilitated by bile and the enzyme pancreatic lipase. Moreover, intestinal juice combines with pancreatic juice to provide a liquid medium that facilitates absorption. The intestine is also where most water is absorbed, via osmosis. The small intestine’s absorptive cells also synthesize digestive enzymes and then place them in the plasma membranes of the microvilli. This distinguishes the small intestine from the stomach; that is, enzymatic digestion occurs not only in the lumen, but also on the luminal surfaces of the mucosal cells. For optimal chemical digestion, chyme must be delivered from the stomach slowly and in small amounts. This is because chyme from the stomach is typically hypertonic, and if large quantities were forced all at once into the small intestine, the resulting osmotic water loss from the blood into the intestinal lumen would result in potentially life-threatening low blood volume. In addition, continued digestion requires an upward adjustment of the low pH of stomach chyme, along with rigorous mixing of the chyme with bile and pancreatic juices. Both processes take time, so the pumping action of the pylorus must be carefully controlled to prevent the duodenum from being overwhelmed with chyme. Disorders of the…Small Intestine: Lactose IntoleranceLactose intolerance is a condition characterized by indigestion caused by dairy products. It occurs when the absorptive cells of the small intestine do not produce enough lactase, the enzyme that digests the milk sugar lactose. In most mammals, lactose intolerance increases with age. In contrast, some human populations, most notably Caucasians, are able to maintain the ability to produce lactase as adults. In people with lactose intolerance, the lactose in chyme is not digested. Bacteria in the large intestine ferment the undigested lactose, a process that produces gas. In addition to gas, symptoms include abdominal cramps, bloating, and diarrhea. Symptom severity ranges from mild discomfort to severe pain; however, symptoms resolve once the lactose is eliminated in feces. The hydrogen breath test is used to help diagnose lactose intolerance. Lactose-tolerant people have very little hydrogen in their breath. Those with lactose intolerance exhale hydrogen, which is one of the gases produced by the bacterial fermentation of lactose in the colon. After the hydrogen is absorbed from the intestine, it is transported through blood vessels into the lungs. There are a number of lactose-free dairy products available in grocery stores. In addition, dietary supplements are available. Taken with food, they provide lactase to help digest lactose. The Large IntestineThe StructureThe large intestine runs from the appendix to the anus. It frames the small intestine on three sides. Despite its being about one-half as long as the small intestine, it is called large because it is more than twice the diameter of the small intestine, about 3 inches. SubdivisionsThe large intestine is subdivided into four main regions: the cecum, the colon, the rectum, and the anus. The ileocecal valve, located at the opening between the ileum and the large intestine, controls the flow of chyme from the small intestine to the large intestine. Cecum The first part of the large intestine is the Colon The cecum blends seamlessly with the
Large Intestine
Homeostatic ImbalancesColorectal CancerEach year, approximately 140,000 Americans are diagnosed with colorectal cancer, and another 49,000 die from it, making it one of the most deadly malignancies. People with a family history of colorectal cancer are at increased risk. Smoking, excessive alcohol consumption, and a diet high in animal fat and protein also increase the risk. Despite popular opinion to the contrary, studies support the conclusion that dietary fiber and calcium do not reduce the risk of colorectal cancer. Colorectal cancer may be signaled by constipation or diarrhea, cramping, abdominal pain, and rectal bleeding. Bleeding from the rectum may be either obvious or occult (hidden in feces). Since most colon cancers arise from benign mucosal growths called polyps, cancer prevention is focused on identifying these polyps. The colonoscopy is both diagnostic and therapeutic. Colonoscopy not only allows identification of precancerous polyps, the procedure also enables them to be removed before they become malignant. Screening for fecal occult blood tests and colonoscopy is recommended for those over 50 years of age. Rectum Food residue leaving the sigmoid colon enters the Anal Canal Finally, food residue reaches the last part of the large intestine, the HistologyThere are several notable differences between the walls of the large and small intestines (Figure 5). For example, few enzyme-secreting cells are found in the wall of the large intestine, and there are no circular folds or villi. Other than in the anal canal, the mucosa of the colon is simple columnar epithelium made mostly of enterocytes (absorptive cells) and goblet cells. In addition, the wall of the large intestine has far more intestinal glands, which contain a vast population of enterocytes and goblet cells. These goblet cells secrete mucus that eases the movement of feces and protects the intestine from the effects of the acids and gases produced by enteric bacteria. The enterocytes absorb water and salts as well as vitamins produced by your intestinal bacteria. AnatomyThree features are unique to the large intestine: teniae coli, haustra, and epiploic appendages (Figure 6). The
Teniae Coli, Haustra, and Epiploic Appendages
Figure 6 The stratified squamous epithelial mucosa of the anal canal connects to the skin on the outside of the anus. This mucosa varies considerably from that of the rest of the colon to accommodate the high level of abrasion as feces pass through. The anal canal’s mucous membrane is organized into longitudinal folds, each called an Depressions between the anal columns, each called an Bacterial FloraMost bacteria that enter the alimentary canal are killed by lysozyme, defensins, HCl, or protein-digesting enzymes. However, trillions of bacteria live within the large intestine and are referred to as the Digestive Functions of the Large IntestineThe residue of chyme that enters the large intestine contains few nutrients except water, which is reabsorbed as the residue lingers in the large intestine, typically for 12 to 24 hours. Thus, it may not surprise you that the large intestine can be completely removed without significantly affecting digestive functioning. For example, in severe cases of inflammatory bowel disease, the large intestine can be removed by a procedure known as a colectomy. Often, a new fecal pouch can be crafted from the small intestine and sutured to the anus, but if not, an ileostomy can be created by bringing the distal ileum through the abdominal wall, allowing the watery chyme to be collected in a bag-like adhesive appliance. Mechanical Digestion In the large intestine, mechanical digestion begins when chyme moves from the ileum into the cecum, an activity regulated by the ileocecal sphincter. Right after you eat, peristalsis in the ileum forces chyme into the cecum. When the cecum is distended with chyme, contractions of the ileocecal sphincter strengthen. Once chyme enters the cecum, colon movements begin. Mechanical digestion in the large intestine includes a combination of three types of movements. The presence of food residues in the colon stimulates a slow-moving Chemical Digestion Although the glands of the large intestine secrete mucus, they do not secrete digestive enzymes. Therefore, chemical digestion in the large intestine occurs exclusively because of bacteria in the lumen of the colon. Through the process of Absorption, Feces Formation, and DefecationThe small intestine absorbs about 90 percent of the water you ingest (either as liquid or within solid food). The large intestine absorbs most of the remaining water, a process that converts the liquid chyme residue into semisolid Feces are eliminated through contractions of the rectal muscles. You help this process by a voluntary procedure called The process of defecation begins when mass movements force feces from the colon into the rectum, stretching the rectal wall and provoking the defecation reflex, which eliminates feces from the rectum. This parasympathetic reflex is mediated by the spinal cord. It contracts the sigmoid colon and rectum, relaxes the internal anal sphincter, and initially contracts the external anal sphincter. The presence of feces in the anal canal sends a signal to the brain, which gives you the choice of voluntarily opening the external anal sphincter (defecating) or keeping it temporarily closed. If you decide to delay defecation, it takes a few seconds for the reflex contractions to stop and the rectal walls to relax. The next mass movement will trigger additional defecation reflexes until you defecate. If defecation is delayed for an extended time, additional water is absorbed, making the feces firmer and potentially leading to constipation. On the other hand, if the waste matter moves too quickly through the intestines, not enough water is absorbed, and diarrhea can result. This can be caused by the ingestion of foodborne pathogens. In general, diet, health, and stress determine the frequency of bowel movements. The number of bowel movements varies greatly between individuals, ranging from two or three per day to three or four per week.
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By watching this animation you will see that for the various food groups—proteins, fats, and carbohydrates—digestion begins in different parts of the digestion system, though all end in the same place. Of the three major food classes (carbohydrates, fats, and proteins), which is digested in the mouth, the stomach, and the small intestine? Chapter ReviewThe three main regions of the small intestine are the duodenum, the jejunum, and the ileum. The small intestine is where digestion is completed and virtually all absorption occurs. These two activities are facilitated by structural adaptations that increase the mucosal surface area by 600-fold, including circular folds, villi, and microvilli. There are around 200 million microvilli per square millimeter of small intestine, which contain brush border enzymes that complete the digestion of carbohydrates and proteins. Combined with pancreatic juice, intestinal juice provides the liquid medium needed to further digest and absorb substances from chyme. The small intestine is also the site of unique mechanical digestive movements. Segmentation moves the chyme back and forth, increasing mixing and opportunities for absorption. Migrating motility complexes propel the residual chyme toward the large intestine. The main regions of the large intestine are the cecum, the colon, and the rectum. The large intestine absorbs water and forms feces, and is responsible for defecation. Bacterial flora break down additional carbohydrate residue, and synthesize certain vitamins. The mucosa of the large intestinal wall is generously endowed with goblet cells, which secrete mucus that eases the passage of feces. The entry of feces into the rectum activates the defecation reflex. Interactive Link QuestionsExercise 1Watch this animation that depicts the structure of the small intestine, and, in particular, the villi. Epithelial cells continue the digestion and absorption of nutrients and transport these nutrients to the lymphatic and circulatory systems. In the small intestine, the products of food digestion are absorbed by different structures in the villi. Which structure absorbs and transports fats? Show/Hide Solution Answers may vary. Exercise 2By watching this animation, you will see that for the various food groups—proteins, fats, and carbohydrates—digestion begins in different parts of the digestion system, though all end in the same place. Of the three major food classes (carbohydrates, fats, and proteins), which is digested in the mouth, the stomach, and the small intestine? Show/Hide Solution Answers may vary. Review QuestionsExercise 3In which part of the alimentary canal does most digestion occur?
Show/Hide Solution B Exercise 4Which of these is most associated with villi?
Show/Hide Solution B Exercise 5What is the role of the small intestine’s MALT?
Show/Hide Solution D Exercise 6Which part of the large intestine attaches to the appendix?
Show/Hide Solution A Critical Thinking QuestionsExercise 7Explain how nutrients absorbed in the small intestine pass into the general circulation. Show/Hide Solution Nutrients from the breakdown of carbohydrates and proteins are absorbed through a capillary bed in the villi of the small intestine. Lipid breakdown products are absorbed into a lacteal in the villi, and transported via the lymphatic system to the bloodstream. Exercise 8Why is it important that chyme from the stomach is delivered to the small intestine slowly and in small amounts? Show/Hide Solution If large quantities of chyme were forced into the small intestine, it would result in osmotic water loss from the blood into the intestinal lumen that could cause potentially life-threatening low blood volume and erosion of the duodenum. Exercise 9Describe three of the differences between the walls of the large and small intestines. Show/Hide Solution The mucosa of the small intestine includes circular folds, villi, and microvilli. The wall of the large intestine has a thick mucosal layer, and deeper and more abundant mucus-secreting glands that facilitate the smooth passage of feces. There are three features that are unique to the large intestine: teniae coli, haustra, and epiploic appendages. ReferencesThe Nutrition Source. Fiber and colon cancer: following the scientific trail [Internet]. Boston (MA): Harvard School of Public Health; c2012 [cited 2013 Apr 3]. Available from: http://www.hsph.harvard.edu/nutritionsource/nutrition-news/fiber-and-colon-cancer/index.html. Centers for Disease Control and Prevention (US). Morbidity and mortality weekly report: notifiable diseases and mortality tables [Internet]. Atlanta (GA); [cited 2013 Apr 3]. Available from: http://www.cdc.gov/mmwr/preview/mmwrhtml/mm6101md.htm?s_cid=mm6101md_w. Glossaryanal canal anal column anal sinus appendix ascending colon bacterial flora brush border cecum circular fold colon descending colon duodenal gland duodenum epiploic appendage external anal sphincter feces flatus gastrocolic reflex gastroileal reflex haustrum haustral contraction hepatopancreatic ampulla hepatopancreatic sphincter ileocecal sphincter ileum internal anal sphincter intestinal gland intestinal juice jejunum lacteal large intestine left colic flexure main pancreatic duct major duodenal papilla mass movement mesoappendix microvillus migrating motility complex motilin pectinate line rectal valve rectum right colic flexure saccharolytic fermentation sigmoid colon small intestine tenia coli transverse colon Valsalva’s maneuver villus
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