2. Digestive System

Chemical Digestion and Absorption: A Closer Look

As you have learned, the process of mechanical digestion is relatively simple. It involves the physical breakdown of food but does not alter its chemical makeup. Chemical digestion, on the other hand, is a complex process that reduces food into its chemical building blocks, which are then absorbed to nourish the cells of the body (Figure 23.28). In this section, you will look more closely at the processes of chemical digestion and absorption.

This diagram identifies the functions of mechanical and chemical digestion and absorption at each organ. Next to each organ, a callout identifies which steps of digestion take place in that particular organ.

Figure 23.28 Digestion and Absorption Digestion begins in the mouth and continues as food travels through the small intestine. Most absorption occurs in the small intestine.

Chemical Digestion

Large food molecules (for example, proteins, lipids, nucleic acids, and starches) must be broken down into subunits that are small enough to be absorbed by the lining of the alimentary canal. This is accomplished by enzymes through hydrolysis. The many enzymes involved in chemical digestion are summarized in Table 23.8.The Digestive Enzymes

Enzyme CategoryEnzyme NameSourceSubstrateProduct
Salivary EnzymesLingual lipaseLingual glandsTriglyceridesFree fatty acids, and mono- and diglycerides
Salivary EnzymesSalivary amylaseSalivary glandsPolysaccharidesDisaccharides and trisaccharides
Gastric enzymesGastric lipaseChief cellsTriglyceridesFatty acids and monoacylglycerides
Gastric enzymesPepsin*Chief cellsProteinsPeptides
Brush border enzymesα-DextrinaseSmall intestineα-DextrinsGlucose
Brush border enzymesEnteropeptidaseSmall intestineTrypsinogenTrypsin
Brush border enzymesLactaseSmall intestineLactoseGlucose and galactose
Brush border enzymesMaltaseSmall intestineMaltoseGlucose
Brush border enzymesNucleosidases and phosphatasesSmall intestineNucleotidesPhosphates, nitrogenous bases, and pentoses
Brush border enzymesPeptidasesSmall intestineAminopeptidase: amino acids at the amino end of peptidesDipeptidase: dipeptidesAminopeptidase: amino acids and peptidesDipeptidase: amino acids
Brush border enzymesSucraseSmall intestineSucroseGlucose and fructose
Pancreatic enzymesCarboxy-peptidase*Pancreatic acinar cellsAmino acids at the carboxyl end of peptidesAmino acids and peptides
Pancreatic enzymesChymotrypsin*Pancreatic acinar cellsProteinsPeptides
Pancreatic enzymesElastase*Pancreatic acinar cellsProteinsPeptides
Pancreatic enzymesNucleasesPancreatic acinar cellsRibonuclease: ribonucleic acidsDeoxyribonuclease: deoxyribonucleic acidsNucleotides
Pancreatic enzymesPancreatic amylasePancreatic acinar cellsPolysaccharides (starches)α-Dextrins, disaccharides (maltose), trisaccharides (maltotriose)
Pancreatic enzymesPancreatic lipasePancreatic acinar cellsTriglycerides that have been emulsified by bile saltsFatty acids and monoacylglycerides
Pancreatic enzymesTrypsin*Pancreatic acinar cellsProteinsPeptides

Table23.8 *These enzymes have been activated by other substances.

Carbohydrate Digestion

The average American diet is about 50 percent carbohydrates, which may be classified according to the number of monomers they contain of simple sugars (monosaccharides and disaccharides) and/or complex sugars (polysaccharides). Glucose, galactose, and fructose are the three monosaccharides that are commonly consumed and are readily absorbed. Your digestive system is also able to break down the disaccharide sucrose (regular table sugar: glucose + fructose), lactose (milk sugar: glucose + galactose), and maltose (grain sugar: glucose + glucose), and the polysaccharides glycogen and starch (chains of monosaccharides). Your bodies do not produce enzymes that can break down most fibrous polysaccharides, such as cellulose. While indigestible polysaccharides do not provide any nutritional value, they do provide dietary fiber, which helps propel food through the alimentary canal.

The chemical digestion of starches begins in the mouth and has been reviewed above.

In the small intestine, pancreatic amylase does the ‘heavy lifting’ for starch and carbohydrate digestion (Figure 23.29). After amylases break down starch into smaller fragments, the brush border enzyme α-dextrinase starts working on α-dextrin, breaking off one glucose unit at a time. Three brush border enzymes hydrolyze sucrose, lactose, and maltose into monosaccharides. Sucrase splits sucrose into one molecule of fructose and one molecule of glucose; maltase breaks down maltose and maltotriose into two and three glucose molecules, respectively; and lactase breaks down lactose into one molecule of glucose and one molecule of galactose. Insufficient lactase can lead to lactose intolerance.

This flow chart shows the steps in digestion of carbohydrates. The different levels shown are starch and glycogen, disaccharides and monosaccharides. Under each type of sugar, examples and the enzymes responsible for digestion are listed.

Figure 23.29Carbohydrate Digestion Flow Chart Carbohydrates are broken down into their monomers in a series of steps.

Protein Digestion

Proteins are polymers composed of amino acids linked by peptide bonds to form long chains. Digestion reduces them to their constituent amino acids. You usually consume about 15 to 20 percent of your total calorie intake as protein.

The digestion of protein starts in the stomach, where HCl and pepsin break proteins into smaller polypeptides, which then travel to the small intestine (Figure 23.30). Chemical digestion in the small intestine is continued by pancreatic enzymes, including chymotrypsin and trypsin, each of which act on specific bonds in amino acid sequences. At the same time, the cells of the brush border secrete enzymes such as aminopeptidase and dipeptidase, which further break down peptide chains. This results in molecules small enough to enter the bloodstream (Figure 23.31).

This diagrams shows the human digestive system and identifies the role of each organ in protein digestion. A text call-out next to each organ details the specific function.

Figure 23.30Digestion of Protein The digestion of protein begins in the stomach and is completed in the small intestine.

This flow chart shows the different steps in the digestion of protein. The four steps shown are protein, large polypeptides, short peptides and amino acids and amino acids.

Figure 23.31Digestion of Protein Flow Chart Proteins are successively broken down into their amino acid components.

Lipid Digestion

A healthy diet limits lipid intake to 35 percent of total calorie intake. The most common dietary lipids are triglycerides, which are made up of a glycerol molecule bound to three fatty acid chains. Small amounts of dietary cholesterol and phospholipids are also consumed.

The three lipases responsible for lipid digestion are lingual lipase, gastric lipase, and pancreatic lipase. However, because the pancreas is the only consequential source of lipase, virtually all lipid digestion occurs in the small intestine. Pancreatic lipase breaks down each triglyceride into two free fatty acids and a monoglyceride. The fatty acids include both short-chain (less than 10 to 12 carbons) and long-chain fatty acids.

Nucleic Acid Digestion

The nucleic acids DNA and RNA are found in most of the foods you eat. Two types of pancreatic nuclease are responsible for their digestion: deoxyribonuclease, which digests DNA, and ribonuclease, which digests RNA. The nucleotides produced by this digestion are further broken down by two intestinal brush border enzymes (nucleosidase and phosphatase) into pentoses, phosphates, and nitrogenous bases, which can be absorbed through the alimentary canal wall. The large food molecules that must be broken down into subunits are summarized Table 23.9Absorbable Food Substances

CarbohydratesMonosaccharides: glucose, galactose, and fructose
ProteinsSingle amino acids, dipeptides, and tripeptides
TriglyceridesMonoacylglycerides, glycerol, and free fatty acids
Nucleic acidsPentose sugars, phosphates, and nitrogenous bases



The mechanical and digestive processes have one goal: to convert food into molecules small enough to be absorbed by the epithelial cells of the intestinal villi. The absorptive capacity of the alimentary canal is almost endless. Each day, the alimentary canal processes up to 10 liters of food, liquids, and GI secretions, yet less than one liter enters the large intestine. Almost all ingested food, 80 percent of electrolytes, and 90 percent of water are absorbed in the small intestine. Although the entire small intestine is involved in the absorption of water and lipids, most absorption of carbohydrates and proteins occurs in the jejunum. Notably, bile salts and vitamin B12 are absorbed in the terminal ileum. By the time chyme passes from the ileum into the large intestine, it is essentially indigestible food residue (mainly plant fibers like cellulose), some water, and millions of bacteria (Figure 23.32).

This image shows the human digestive system. Next to each organ, a text callout identifies how water and digestive secretions such as saliva and bile are processed.

Figure 23.32Digestive Secretions and Absorption of Water Absorption is a complex process, in which nutrients from digested food are harvested.

Absorption can occur through five mechanisms: (1) active transport, (2) passive diffusion, (3) facilitated diffusion, (4) co-transport (or secondary active transport), and (5) endocytosis. As you will recall from Chapter 3, active transport refers to the movement of a substance across a cell membrane going from an area of lower concentration to an area of higher concentration (up the concentration gradient). In this type of transport, proteins within the cell membrane act as “pumps,” using cellular energy (ATP) to move the substance. Passive diffusion refers to the movement of substances from an area of higher concentration to an area of lower concentration, while facilitated diffusion refers to the movement of substances from an area of higher to an area of lower concentration using a carrier protein in the cell membrane. Co-transport uses the movement of one molecule through the membrane from higher to lower concentration to power the movement of another from lower to higher. Finally, endocytosis is a transportation process in which the cell membrane engulfs material. It requires energy, generally in the form of ATP.

Because the cell’s plasma membrane is made up of hydrophobic phospholipids, water-soluble nutrients must use transport molecules embedded in the membrane to enter cells. Moreover, substances cannot pass between the epithelial cells of the intestinal mucosa because these cells are bound together by tight junctions. Thus, substances can only enter blood capillaries by passing through the apical surfaces of epithelial cells and into the interstitial fluid. Water-soluble nutrients enter the capillary blood in the villi and travel to the liver via the hepatic portal vein.

In contrast to the water-soluble nutrients, lipid-soluble nutrients can diffuse through the plasma membrane. Once inside the cell, they are packaged for transport via the base of the cell and then enter the lacteals of the villi to be transported by lymphatic vessels to the systemic circulation via the thoracic duct. The absorption of most nutrients through the mucosa of the intestinal villi requires active transport fueled by ATP. The routes of absorption for each food category are summarized in Table 23.10.Absorption in the Alimentary Canal

FoodBreakdown productsAbsorption mechanismEntry to bloodstreamDestination
CarbohydratesGlucoseCo-transport with sodium ionsCapillary blood in villiLiver via hepatic portal vein
CarbohydratesGalactoseCo-transport with sodium ionsCapillary blood in villiLiver via hepatic portal vein
CarbohydratesFructoseFacilitated diffusionCapillary blood in villiLiver via hepatic portal vein
ProteinAmino acidsCo-transport with sodium ionsCapillary blood in villiLiver via hepatic portal vein
LipidsLong-chain fatty acidsDiffusion into intestinal cells, where they are combined with proteins to create chylomicronsLacteals of villiSystemic circulation via lymph entering thoracic duct
LipidsMonoacylglyceridesDiffusion into intestinal cells, where they are combined with proteins to create chylomicronsLacteals of villiSystemic circulation via lymph entering thoracic duct
LipidsShort-chain fatty acidsSimple diffusionCapillary blood in villiLiver via hepatic portal vein
LipidsGlycerolSimple diffusionCapillary blood in villiLiver via hepatic portal vein
Nucleic AcidsNucleic acid digestion productsActive transport via membrane carriersCapillary blood in villiLiver via hepatic portal vein


Carbohydrate Absorption

All carbohydrates are absorbed in the form of monosaccharides. The small intestine is highly efficient at this, absorbing monosaccharides at an estimated rate of 120 grams per hour. All normally digested dietary carbohydrates are absorbed; indigestible fibers are eliminated in the feces. The monosaccharides glucose and galactose are transported into the epithelial cells by common protein carriers via secondary active transport (that is, co-transport with sodium ions). The monosaccharides leave these cells via facilitated diffusion and enter the capillaries through intercellular clefts. The monosaccharide fructose (which is in fruit) is absorbed and transported by facilitated diffusion alone. The monosaccharides combine with the transport proteins immediately after the disaccharides are broken down.

Protein Absorption

Active transport mechanisms, primarily in the duodenum and jejunum, absorb most proteins as their breakdown products, amino acids. Almost all (95 to 98 percent) protein is digested and absorbed in the small intestine. The type of carrier that transports an amino acid varies. Most carriers are linked to the active transport of sodium. Short chains of two amino acids (dipeptides) or three amino acids (tripeptides) are also transported actively. However, after they enter the absorptive epithelial cells, they are broken down into their amino acids before leaving the cell and entering the capillary blood via diffusion.

Lipid Absorption

About 95 percent of lipids are absorbed in the small intestine. Bile salts not only speed up lipid digestion, they are also essential to the absorption of the end products of lipid digestion. Short-chain fatty acids are relatively water soluble and can enter the absorptive cells (enterocytes) directly. The small size of short-chain fatty acids enables them to be absorbed by enterocytes via simple diffusion, and then take the same path as monosaccharides and amino acids into the blood capillary of a villus.

The large and hydrophobic long-chain fatty acids and monoacylglycerides are not so easily suspended in the watery intestinal chyme. However, bile salts and lecithin resolve this issue by enclosing them in a micelle, which is a tiny sphere with polar (hydrophilic) ends facing the watery environment and hydrophobic tails turned to the interior, creating a receptive environment for the long-chain fatty acids. The core also includes cholesterol and fat-soluble vitamins. Without micelles, lipids would sit on the surface of chyme and never come in contact with the absorptive surfaces of the epithelial cells. Micelles can easily squeeze between microvilli and get very near the luminal cell surface. At this point, lipid substances exit the micelle and are absorbed via simple diffusion.

The free fatty acids and monoacylglycerides that enter the epithelial cells are reincorporated into triglycerides. The triglycerides are mixed with phospholipids and cholesterol, and surrounded with a protein coat. This new complex, called a chylomicron, is a water-soluble lipoprotein. After being processed by the Golgi apparatus, chylomicrons are released from the cell (Figure 23.33). Too big to pass through the basement membranes of blood capillaries, chylomicrons instead enter the large pores of lacteals. The lacteals come together to form the lymphatic vessels. The chylomicrons are transported in the lymphatic vessels and empty through the thoracic duct into the subclavian vein of the circulatory system. Once in the bloodstream, the enzyme lipoprotein lipase breaks down the triglycerides of the chylomicrons into free fatty acids and glycerol. These breakdown products then pass through capillary walls to be used for energy by cells or stored in adipose tissue as fat. Liver cells combine the remaining chylomicron remnants with proteins, forming lipoproteins that transport cholesterol in the blood.

This diagram shows how lipids are absorbed from the lumen of the intestine into the lacteals. The fatty acid micelles are shown to enter the epithelial cell and form chylomicrons inside the Golgi apparatus. Then, the chylomicrons are extruded from the epithelial cell and are taken up by the lacteals.

Figure 23.33Lipid Absorption Unlike amino acids and simple sugars, lipids are transformed as they are absorbed through epithelial cells.

Nucleic Acid Absorption

The products of nucleic acid digestion—pentose sugars, nitrogenous bases, and phosphate ions—are transported by carriers across the villus epithelium via active transport. These products then enter the bloodstream.

Mineral Absorption

The electrolytes absorbed by the small intestine are from both GI secretions and ingested foods. Since electrolytes dissociate into ions in water, most are absorbed via active transport throughout the entire small intestine. During absorption, co-transport mechanisms result in the accumulation of sodium ions inside the cells, whereas anti-port mechanisms reduce the potassium ion concentration inside the cells. To restore the sodium-potassium gradient across the cell membrane, a sodium-potassium pump requiring ATP pumps sodium out and potassium in.

In general, all minerals that enter the intestine are absorbed, whether you need them or not. Iron and calcium are exceptions; they are absorbed in the duodenum in amounts that meet the body’s current requirements, as follows:

Iron—The ionic iron needed for the production of hemoglobin is absorbed into mucosal cells via active transport. Once inside mucosal cells, ionic iron binds to the protein ferritin, creating iron-ferritin complexes that store iron until needed. When the body has enough iron, most of the stored iron is lost when worn-out epithelial cells slough off. When the body needs iron because, for example, it is lost during acute or chronic bleeding, there is increased uptake of iron from the intestine and accelerated release of iron into the bloodstream. Since women experience significant iron loss during menstruation, they have around four times as many iron transport proteins in their intestinal epithelial cells as do men.

Calcium—Blood levels of ionic calcium determine the absorption of dietary calcium. When blood levels of ionic calcium drop, parathyroid hormone (PTH) secreted by the parathyroid glands stimulates the release of calcium ions from bone matrices and increases the reabsorption of calcium by the kidneys. PTH also upregulates the activation of vitamin D in the kidney, which then facilitates intestinal calcium ion absorption.

Vitamin Absorption

The small intestine absorbs the vitamins that occur naturally in food and supplements. Fat-soluble vitamins (A, D, E, and K) are absorbed along with dietary lipids in micelles via simple diffusion. This is why you are advised to eat some fatty foods when you take fat-soluble vitamin supplements. Most water-soluble vitamins (including most B vitamins and vitamin C) also are absorbed by simple diffusion. An exception is vitamin B12, which is a very large molecule. Intrinsic factor secreted in the stomach binds to vitamin B12, preventing its digestion and creating a complex that binds to mucosal receptors in the terminal ileum, where it is taken up by endocytosis.

Water Absorption

Each day, about nine liters of fluid enter the small intestine. About 2.3 liters are ingested in foods and beverages, and the rest is from GI secretions. About 90 percent of this water is absorbed in the small intestine. Water absorption is driven by the concentration gradient of the water: The concentration of water is higher in chyme than it is in epithelial cells. Thus, water moves down its concentration gradient from the chyme into cells. As noted earlier, much of the remaining water is then absorbed in the colon.

2. Digestive System

Accessory Organs in Digestion: The Liver, Pancreas, and Gallbladder

Chemical digestion in the small intestine relies on the activities of three accessory digestive organs: the liver, pancreas, and gallbladder (Figure 23.24). The digestive role of the liver is to produce bile and export it to the duodenum. The gallbladder primarily stores, concentrates, and releases bile. The pancreas produces pancreatic juice, which contains digestive enzymes and bicarbonate ions, and delivers it to the duodenum.

This diagram shows the accessory organs of the digestive system. The liver, spleen, pancreas, gallbladder and their major parts are shown.

Figure 23.24Accessory Organs The liver, pancreas, and gallbladder are considered accessory digestive organs, but their roles in the digestive system are vital.

The Liver

The liver is the largest gland in the body, weighing about three pounds in an adult. It is also one of the most important organs. In addition to being an accessory digestive organ, it plays a number of roles in metabolism and regulation. The liver lies inferior to the diaphragm in the right upper quadrant of the abdominal cavity and receives protection from the surrounding ribs.

The liver is divided into two primary lobes: a large right lobe and a much smaller left lobe. In the right lobe, some anatomists also identify an inferior quadrate lobe and a posterior caudate lobe, which are defined by internal features. The liver is connected to the abdominal wall and diaphragm by five peritoneal folds referred to as ligaments. These are the falciform ligament, the coronary ligament, two lateral ligaments, and the ligamentum teres hepatis. The falciform ligament and ligamentum teres hepatis are actually remnants of the umbilical vein, and separate the right and left lobes anteriorly. The lesser omentum tethers the liver to the lesser curvature of the stomach.

The porta hepatis (“gate to the liver”) is where the hepatic artery and hepatic portal vein enter the liver. These two vessels, along with the common hepatic duct, run behind the lateral border of the lesser omentum on the way to their destinations. As shown in Figure 23.25, the hepatic artery delivers oxygenated blood from the heart to the liver. The hepatic portal vein delivers partially deoxygenated blood containing nutrients absorbed from the small intestine and actually supplies more oxygen to the liver than do the much smaller hepatic arteries. In addition to nutrients, drugs and toxins are also absorbed. After processing the bloodborne nutrients and toxins, the liver releases nutrients needed by other cells back into the blood, which drains into the central vein and then through the hepatic vein to the inferior vena cava. With this hepatic portal circulation, all blood from the alimentary canal passes through the liver. This largely explains why the liver is the most common site for the metastasis of cancers that originate in the alimentary canal.

This image shows the microscopic anatomy of the liver. The top panel shows the liver; the center panel shows a magnified view of the connective tissue and the lobules. The bottom panel shows a further magnified view of a lobule, identifying the veins, bile duct and the sinusoids.

Figure 23.25Microscopic Anatomy of the Liver The liver receives oxygenated blood from the hepatic artery and nutrient-rich deoxygenated blood from the hepatic portal vein.


The liver has three main components: hepatocytes, bile canaliculi, and hepatic sinusoids. A hepatocyte is the liver’s main cell type, accounting for around 80 percent of the liver’s volume. These cells play a role in a wide variety of secretory, metabolic, and endocrine functions. Plates of hepatocytes called hepatic laminae radiate outward from the portal vein in each hepatic lobule.

Between adjacent hepatocytes, grooves in the cell membranes provide room for each bile canaliculus (plural = canaliculi). These small ducts accumulate the bile produced by hepatocytes. From here, bile flows first into bile ductules and then into bile ducts. The bile ducts unite to form the larger right and left hepatic ducts, which themselves merge and exit the liver as the common hepatic duct. This duct then joins with the cystic duct from the gallbladder, forming the common bile duct through which bile flows into the small intestine.

hepatic sinusoid is an open, porous blood space formed by fenestrated capillaries from nutrient-rich hepatic portal veins and oxygen-rich hepatic arteries. Hepatocytes are tightly packed around the fenestrated endothelium of these spaces, giving them easy access to the blood. From their central position, hepatocytes process the nutrients, toxins, and waste materials carried by the blood. Materials such as bilirubin are processed and excreted into the bile canaliculi. Other materials including proteins, lipids, and carbohydrates are processed and secreted into the sinusoids or just stored in the cells until called upon. The hepatic sinusoids combine and send blood to a central vein. Blood then flows through a hepatic vein into the inferior vena cava. This means that blood and bile flow in opposite directions. The hepatic sinusoids also contain star-shaped reticuloendothelial cells (Kupffer cells), phagocytes that remove dead red and white blood cells, bacteria, and other foreign material that enter the sinusoids. The portal triad is a distinctive arrangement around the perimeter of hepatic lobules, consisting of three basic structures: a bile duct, a hepatic artery branch, and a hepatic portal vein branch.


Recall that lipids are hydrophobic, that is, they do not dissolve in water. Thus, before they can be digested in the watery environment of the small intestine, large lipid globules must be broken down into smaller lipid globules, a process called emulsification. Bile is a mixture secreted by the liver to accomplish the emulsification of lipids in the small intestine.

Hepatocytes secrete about one liter of bile each day. A yellow-brown or yellow-green alkaline solution (pH 7.6 to 8.6), bile is a mixture of water, bile salts, bile pigments, phospholipids (such as lecithin), electrolytes, cholesterol, and triglycerides. The components most critical to emulsification are bile salts and phospholipids, which have a nonpolar (hydrophobic) region as well as a polar (hydrophilic) region. The hydrophobic region interacts with the large lipid molecules, whereas the hydrophilic region interacts with the watery chyme in the intestine. This results in the large lipid globules being pulled apart into many tiny lipid fragments of about 1 µm in diameter. This change dramatically increases the surface area available for lipid-digesting enzyme activity. This is the same way dish soap works on fats mixed with water.

Bile salts act as emulsifying agents, so they are also important for the absorption of digested lipids. While most constituents of bile are eliminated in feces, bile salts are reclaimed by the enterohepatic circulation. Once bile salts reach the ileum, they are absorbed and returned to the liver in the hepatic portal blood. The hepatocytes then excrete the bile salts into newly formed bile. Thus, this precious resource is recycled.

Bilirubin, the main bile pigment, is a waste product produced when the spleen removes old or damaged red blood cells from the circulation. These breakdown products, including proteins, iron, and toxic bilirubin, are transported to the liver via the splenic vein of the hepatic portal system. In the liver, proteins and iron are recycled, whereas bilirubin is excreted in the bile. It accounts for the green color of bile. Bilirubin is eventually transformed by intestinal bacteria into stercobilin, a brown pigment that gives your stool its characteristic color! In some disease states, bile does not enter the intestine, resulting in white (‘acholic’) stool with a high fat content, since virtually no fats are broken down or absorbed.

Hepatocytes work non-stop, but bile production increases when fatty chyme enters the duodenum and stimulates the secretion of the gut hormone secretin. Between meals, bile is produced but conserved. The valve-like hepatopancreatic ampulla closes, allowing bile to divert to the gallbladder, where it is concentrated and stored until the next meal.

The Pancreas

The soft, oblong, glandular pancreas lies transversely in the retroperitoneum behind the stomach. Its head is nestled into the “c-shaped” curvature of the duodenum with the body extending to the left about 15.2 cm (6 in) and ending as a tapering tail in the hilum of the spleen. It is a curious mix of exocrine (secreting digestive enzymes) and endocrine (releasing hormones into the blood) functions (Figure 23.26).

This figure shows the pancreas and its major parts. A magnified view of a small region of the pancreas shows the pancreatic islet cells, the acinar cells and the pancreatic duct.

Figure 23.26 Exocrine and Endocrine Pancreas The pancreas has a head, a body, and a tail. It delivers pancreatic juice to the duodenum through the pancreatic duct.

The exocrine part of the pancreas arises as little grape-like cell clusters, each called an acinus (plural = acini), located at the terminal ends of pancreatic ducts. These acinar cells secrete enzyme-rich pancreatic juice into tiny merging ducts that form two dominant ducts. The larger duct fuses with the common bile duct (carrying bile from the liver and gallbladder) just before entering the duodenum via a common opening (the hepatopancreatic ampulla). The smooth muscle sphincter of the hepatopancreatic ampulla controls the release of pancreatic juice and bile into the small intestine. The second and smaller pancreatic duct, the accessory duct (duct of Santorini), runs from the pancreas directly into the duodenum, approximately 1 inch above the hepatopancreatic ampulla. When present, it is a persistent remnant of pancreatic development.

Scattered through the sea of exocrine acini are small islands of endocrine cells, the islets of Langerhans. These vital cells produce the hormones pancreatic polypeptide, insulin, glucagon, and somatostatin.

Pancreatic Juice

The pancreas produces over a liter of pancreatic juice each day. Unlike bile, it is clear and composed mostly of water along with some salts, sodium bicarbonate, and several digestive enzymes. Sodium bicarbonate is responsible for the slight alkalinity of pancreatic juice (pH 7.1 to 8.2), which serves to buffer the acidic gastric juice in chyme, inactivate pepsin from the stomach, and create an optimal environment for the activity of pH-sensitive digestive enzymes in the small intestine. Pancreatic enzymes are active in the digestion of sugars, proteins, and fats.

The pancreas produces protein-digesting enzymes in their inactive forms. These enzymes are activated in the duodenum. If produced in an active form, they would digest the pancreas (which is exactly what occurs in the disease, pancreatitis). The intestinal brush border enzyme enteropeptidase stimulates the activation of trypsin from trypsinogen of the pancreas, which in turn changes the pancreatic enzymes procarboxypeptidase and chymotrypsinogen into their active forms, carboxypeptidase and chymotrypsin.

The enzymes that digest starch (amylase), fat (lipase), and nucleic acids (nuclease) are secreted in their active forms, since they do not attack the pancreas as do the protein-digesting enzymes.

Pancreatic Secretion

Regulation of pancreatic secretion is the job of hormones and the parasympathetic nervous system. The entry of acidic chyme into the duodenum stimulates the release of secretin, which in turn causes the duct cells to release bicarbonate-rich pancreatic juice. The presence of proteins and fats in the duodenum stimulates the secretion of CCK, which then stimulates the acini to secrete enzyme-rich pancreatic juice and enhances the activity of secretin. Parasympathetic regulation occurs mainly during the cephalic and gastric phases of gastric secretion, when vagal stimulation prompts the secretion of pancreatic juice.

Usually, the pancreas secretes just enough bicarbonate to counterbalance the amount of HCl produced in the stomach. Hydrogen ions enter the blood when bicarbonate is secreted by the pancreas. Thus, the acidic blood draining from the pancreas neutralizes the alkaline blood draining from the stomach, maintaining the pH of the venous blood that flows to the liver.

The Gallbladder

The gallbladder is 8–10 cm (~3–4 in) long and is nested in a shallow area on the posterior aspect of the right lobe of the liver. This muscular sac stores, concentrates, and, when stimulated, propels the bile into the duodenum via the common bile duct. It is divided into three regions. The fundus is the widest portion and tapers medially into the body, which in turn narrows to become the neck. The neck angles slightly superiorly as it approaches the hepatic duct. The cystic duct is 1–2 cm (less than 1 in) long and turns inferiorly as it bridges the neck and hepatic duct.

The simple columnar epithelium of the gallbladder mucosa is organized in rugae, similar to those of the stomach. There is no submucosa in the gallbladder wall. The wall’s middle, muscular coat is made of smooth muscle fibers. When these fibers contract, the gallbladder’s contents are ejected through the cystic duct and into the bile duct (Figure 23.27). Visceral peritoneum reflected from the liver capsule holds the gallbladder against the liver and forms the outer coat of the gallbladder. The gallbladder’s mucosa absorbs water and ions from bile, concentrating it by up to 10-fold.

This figure shows the gallbladder and its major parts are labeled.

Figure 23.27Gallbladder The gallbladder stores and concentrates bile, and releases it into the two-way cystic duct when it is needed by the small intestine.

2. Digestive System

Small and Large Intestines

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 Intestine

Chyme released from the stomach enters the small intestine, which is the primary digestive organ in the body. Not only is this where most digestion occurs, it is also where practically all absorption occurs. The longest part of the alimentary canal, the small intestine is about 3.05 meters (10 feet) long in a living person (but about twice as long in a cadaver due to the loss of muscle tone). Since this makes it about five times longer than the large intestine, you might wonder why it is called “small.” In fact, its name derives from its relatively smaller diameter of only about 2.54 cm (1 in), compared with 7.62 cm (3 in) for the large intestine. As we’ll see shortly, in addition to its length, the folds and projections of the lining of the small intestine work to give it an enormous surface area, which is approximately 200 m2, more than 100 times the surface area of your skin. This large surface area is necessary for complex processes of digestion and absorption that occur within it.


The 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 23.18).

The shortest region is the 25.4-cm (10-in) duodenum, which begins at the pyloric sphincter. Just past the pyloric sphincter, it bends posteriorly behind the peritoneum, becoming retroperitoneal, and then makes a C-shaped curve around the head of the pancreas before ascending anteriorly again to return to the peritoneal cavity and join the jejunum. The duodenum can therefore be subdivided into four segments: the superior, descending, horizontal, and ascending duodenum.

Of particular interest is the hepatopancreatic ampulla (ampulla of Vater). Located in the duodenal wall, the ampulla marks the transition from the anterior portion of the alimentary canal to the mid-region, and is where the bile duct (through which bile passes from the liver) and the main pancreatic duct (through which pancreatic juice passes from the pancreas) join. This ampulla opens into the duodenum at a tiny volcano-shaped structure called the major duodenal papilla. The hepatopancreatic sphincter (sphincter of Oddi) regulates the flow of both bile and pancreatic juice from the ampulla into the duodenum.

This diagram shows the small intestine. The different parts of the small intestine are labeled.

Figure 23.18 Small Intestine The three regions of the small intestine are the duodenum, jejunum, and ileum.

The jejunum is about 0.9 meters (3 feet) long (in life) and runs from the duodenum to the ileum. Jejunum means “empty” in Latin and supposedly was so named by the ancient Greeks who noticed it was always empty at death. No clear demarcation exists between the jejunum and the final segment of the small intestine, the ileum.

The ileum is the longest part of the small intestine, measuring about 1.8 meters (6 feet) in length. It is thicker, more vascular, and has more developed mucosal folds than the jejunum. The ileum joins the cecum, the first portion of the large intestine, at the ileocecal sphincter (or valve). The jejunum and ileum are tethered to the posterior abdominal wall by the mesentery. The large intestine frames these three parts of the small intestine.

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.


The 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 23.19). These adaptations are most abundant in the proximal two-thirds of the small intestine, where the majority of absorption occurs.

Illustration (a) shows the histological cross-section of the small intestine. The left panel shows a small region of the small intestine, along with the blood vessels and the muscle layers. The middle panel shows a magnified view of a small region of the small intestine, highlighting the absorptive cells, the lacteal and the goblet cells. The right panel shows a further magnified view of the epithelial cells including the microvilli. Illustrations (b) shows a micrograph of the circular folds, and illustration (c) shows a micrograph of the villi. Illustration (d) shows an electron micrograph of the microvilli.

Figure 23.19 Histology of the Small Intestine (a) The absorptive surface of the small intestine is vastly enlarged by the presence of circular folds, villi, and microvilli. (b) Micrograph of the circular folds. (c) Micrograph of the villi. (d) Electron micrograph of the microvilli. From left to right,

Circular folds

Also called a plica circulare, a circular fold is a deep ridge in the mucosa and submucosa. Beginning near the proximal part of the duodenum and ending near the middle of the ileum, these folds facilitate absorption. Their shape causes the chyme to spiral, rather than move in a straight line, through the small intestine. Spiraling slows the movement of chyme and provides the time needed for nutrients to be fully absorbed.


Within the circular folds are small (0.5–1 mm long) hairlike vascularized projections called villi (singular = villus) that give the mucosa a furry texture. There are about 20 to 40 villi per square millimeter, increasing the surface area of the epithelium tremendously. The mucosal epithelium, primarily composed of absorptive cells, covers the villi. In addition to muscle and connective tissue to support its structure, each villus contains a capillary bed composed of one arteriole and one venule, as well as a lymphatic capillary called a lacteal. The breakdown products of carbohydrates and proteins (sugars and amino acids) can enter the bloodstream directly, but lipid breakdown products are absorbed by the lacteals and transported to the bloodstream via the lymphatic system.


As their name suggests, microvilli (singular = microvillus) are much smaller (1 µm) than villi. They are cylindrical apical surface extensions of the plasma membrane of the mucosa’s epithelial cells, and are supported by microfilaments within those cells. Although their small size makes it difficult to see each microvillus, their combined microscopic appearance suggests a mass of bristles, which is termed the brush border. Fixed to the surface of the microvilli membranes are enzymes that finish digesting carbohydrates and proteins. There are an estimated 200 million microvilli per square millimeter of small intestine, greatly expanding the surface area of the plasma membrane and thus greatly enhancing absorption.

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 intestinal gland (crypt of Lieberkühn), which is formed by cells that line the crevices (see Figure 23.19). These produce intestinal juice, a slightly alkaline (pH 7.4 to 7.8) mixture of water and mucus. Each day, about 0.95 to 1.9 liters (1 to 2 quarts) are secreted in response to the distention of the small intestine or the irritating effects of chyme on the intestinal mucosa.

The submucosa of the duodenum is the only site of the complex mucus-secreting duodenal glands (Brunner’s glands), which produce a bicarbonate-rich alkaline mucus that buffers the acidic chyme as it enters from the stomach.

The roles of the cells in the small intestinal mucosa are detailed in Table 23.7.Cells of the Small Intestinal Mucosa

Cell typeLocation in the mucosaFunction
AbsorptiveEpithelium/intestinal glandsDigestion and absorption of nutrients in chyme
GobletEpithelium/intestinal glandsSecretion of mucus
PanethIntestinal glandsSecretion of the bactericidal enzyme lysozyme; phagocytosis
G cellsIntestinal glands of duodenumSecretion of the hormone intestinal gastrin
I cellsIntestinal glands of duodenumSecretion of the hormone cholecystokinin, which stimulates release of pancreatic juices and bile
K cellsIntestinal glandsSecretion of the hormone glucose-dependent insulinotropic peptide, which stimulates the release of insulin
M cellsIntestinal glands of duodenum and jejunumSecretion of the hormone motilin, which accelerates gastric emptying, stimulates intestinal peristalsis, and stimulates the production of pepsin
S cellsIntestinal glandsSecretion of the hormone secretin


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.

Mechanical Digestion in the Small Intestine

The 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 23.20).

This diagram shows the process of segmentation in the intestines. The left panel shows the separation of chime, the middle panel shows the remixing of the chime by pushing it back together and the right panel indicates that the chime is being digested and absorbed.

Figure 23.20 Segmentation Segmentation separates chyme and then pushes it back together, mixing it and providing time for digestion and absorption.

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 motilin, which initiates peristalsis in the form of a migrating motility complex. These complexes, which begin in the duodenum, force chyme through a short section of the small intestine and then stop. The next contraction begins a little bit farther down than the first, forces chyme a bit farther through the small intestine, then stops. These complexes move slowly down the small intestine, forcing chyme on the way, taking around 90 to 120 minutes to finally reach the end of the ileum. At this point, the process is repeated, starting in the duodenum.

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 gastroileal reflex, which increases the force of ileal segmentation. Second, the stomach releases the hormone gastrin, which enhances ileal motility, thus relaxing the ileocecal sphincter. After chyme passes through, backward pressure helps close the sphincter, preventing backflow into the ileum. Because of this reflex, your lunch is completely emptied from your stomach and small intestine by the time you eat your dinner. It takes about 3 to 5 hours for all chyme to leave the small intestine.

Chemical Digestion in the Small Intestine

The 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.

The Large Intestine

The large intestine is the terminal part of the alimentary canal. The primary function of this organ is to finish absorption of nutrients and water, synthesize certain vitamins, form feces, and eliminate feces from the body.


The 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.


The 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.


The first part of the large intestine is the cecum, a sac-like structure that is suspended inferior to the ileocecal valve. It is about 6 cm (2.4 in) long, receives the contents of the ileum, and continues the absorption of water and salts. The appendix (or vermiform appendix) is a winding tube that attaches to the cecum. Although the 7.6-cm (3-in) long appendix contains lymphoid tissue, suggesting an immunologic function, this organ is generally considered vestigial. However, at least one recent report postulates a survival advantage conferred by the appendix: In diarrheal illness, the appendix may serve as a bacterial reservoir to repopulate the enteric bacteria for those surviving the initial phases of the illness. Moreover, its twisted anatomy provides a haven for the accumulation and multiplication of enteric bacteria. The mesoappendix, the mesentery of the appendix, tethers it to the mesentery of the ileum.


The cecum blends seamlessly with the colon. Upon entering the colon, the food residue first travels up the ascending colon on the right side of the abdomen. At the inferior surface of the liver, the colon bends to form the right colic flexure (hepatic flexure) and becomes the transverse colon. The region defined as hindgut begins with the last third of the transverse colon and continues on. Food residue passing through the transverse colon travels across to the left side of the abdomen, where the colon angles sharply immediately inferior to the spleen, at the left colic flexure (splenic flexure). From there, food residue passes through the descending colon, which runs down the left side of the posterior abdominal wall. After entering the pelvis inferiorly, it becomes the s-shaped sigmoid colon, which extends medially to the midline (Figure 23.21). The ascending and descending colon, and the rectum (discussed next) are located in the retroperitoneum. The transverse and sigmoid colon are tethered to the posterior abdominal wall by the mesocolon.

This image shows the large intestine; the major parts of the large intestine are labeled.

Figure 23.21Large Intestine The large intestine includes the cecum, colon, and rectum.


Food residue leaving the sigmoid colon enters the rectum in the pelvis, near the third sacral vertebra. The final 20.3 cm (8 in) of the alimentary canal, the rectum extends anterior to the sacrum and coccyx. Even though rectum is Latin for “straight,” this structure follows the curved contour of the sacrum and has three lateral bends that create a trio of internal transverse folds called the rectal valves. These valves help separate the feces from gas to prevent the simultaneous passage of feces and gas.

Anal Canal

Finally, food residue reaches the last part of the large intestine, the anal canal, which is located in the perineum, completely outside of the abdominopelvic cavity. This 3.8–5 cm (1.5–2 in) long structure opens to the exterior of the body at the anus. The anal canal includes two sphincters. The internal anal sphincter is made of smooth muscle, and its contractions are involuntary. The external anal sphincter is made of skeletal muscle, which is under voluntary control. Except when defecating, both usually remain closed.


There are several notable differences between the walls of the large and small intestines (Figure 23.22). 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.

This image shows the histological cross section of the large intestine. The left panel shows a small region of the large intestine. The center panel shows a magnified view of this region, highlighting the openings of the intestinal glands. The right panel shows a further magnified view, with the microvilli and goblet cells.

Figure 23.22Histology of the large Intestine (a) The histologies of the large intestine and small intestine (not shown) are adapted for the digestive functions of each organ. (b) This micrograph shows the colon’s simple columnar epithelium and goblet cells.


Three features are unique to the large intestine: teniae coli, haustra, and epiploic appendages (Figure 23.23). The teniae coli are three bands of smooth muscle that make up the longitudinal muscle layer of the muscularis of the large intestine, except at its terminal end. Tonic contractions of the teniae coli bunch up the colon into a succession of pouches called haustra (singular = haustrum), which are responsible for the wrinkled appearance of the colon. Attached to the teniae coli are small, fat-filled sacs of visceral peritoneum called epiploic appendages. The purpose of these is unknown. Although the rectum and anal canal have neither teniae coli nor haustra, they do have well-developed layers of muscularis that create the strong contractions needed for defecation.

This image shows the Taenia Coli, haustra and epiploic appendages, which are parts of the large intestine.

Figure 23.23 Teniae Coli, Haustra, and Epiploic Appendages

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 anal column, which house a grid of arteries and veins. Two superficial venous plexuses are found in the anal canal: one within the anal columns and one at the anus.

Depressions between the anal columns, each called an anal sinus, secrete mucus that facilitates defecation. The pectinate line (or dentate line) is a horizontal, jagged band that runs circumferentially just below the level of the anal sinuses, and represents the junction between the hindgut and external skin. The mucosa above this line is fairly insensitive, whereas the area below is very sensitive. The resulting difference in pain threshold is due to the fact that the upper region is innervated by visceral sensory fibers, and the lower region is innervated by somatic sensory fibers.

Bacterial Flora

Most 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 bacterial flora. Most of the more than 700 species of these bacteria are nonpathogenic commensal organisms that cause no harm as long as they stay in the gut lumen. In fact, many facilitate chemical digestion and absorption, and some synthesize certain vitamins, mainly biotin, pantothenic acid, and vitamin K. Some are linked to increased immune response. A refined system prevents these bacteria from crossing the mucosal barrier. First, peptidoglycan, a component of bacterial cell walls, activates the release of chemicals by the mucosa’s epithelial cells, which draft immune cells, especially dendritic cells, into the mucosa. Dendritic cells open the tight junctions between epithelial cells and extend probes into the lumen to evaluate the microbial antigens. The dendritic cells with antigens then travel to neighboring lymphoid follicles in the mucosa where T cells inspect for antigens. This process triggers an IgA-mediated response, if warranted, in the lumen that blocks the commensal organisms from infiltrating the mucosa and setting off a far greater, widespread systematic reaction.

Digestive Functions of the Large Intestine

The 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 haustral contraction. This type of movement involves sluggish segmentation, primarily in the transverse and descending colons. When a haustrum is distended with chyme, its muscle contracts, pushing the residue into the next haustrum. These contractions occur about every 30 minutes, and each last about 1 minute. These movements also mix the food residue, which helps the large intestine absorb water. The second type of movement is peristalsis, which, in the large intestine, is slower than in the more proximal portions of the alimentary canal. The third type is a mass movement. These strong waves start midway through the transverse colon and quickly force the contents toward the rectum. Mass movements usually occur three or four times per day, either while you eat or immediately afterward. Distension in the stomach and the breakdown products of digestion in the small intestine provoke the gastrocolic reflex, which increases motility, including mass movements, in the colon. Fiber in the diet both softens the stool and increases the power of colonic contractions, optimizing the activities of the colon.

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 saccharolytic fermentation, bacteria break down some of the remaining carbohydrates. This results in the discharge of hydrogen, carbon dioxide, and methane gases that create flatus (gas) in the colon; flatulence is excessive flatus. Each day, up to 1500 mL of flatus is produced in the colon. More is produced when you eat foods such as beans, which are rich in otherwise indigestible sugars and complex carbohydrates like soluble dietary fiber.

Absorption, Feces Formation, and Defecation

The 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 (“stool”). Feces is composed of undigested food residues, unabsorbed digested substances, millions of bacteria, old epithelial cells from the GI mucosa, inorganic salts, and enough water to let it pass smoothly out of the body. Of every 500 mL (17 ounces) of food residue that enters the cecum each day, about 150 mL (5 ounces) become feces.

Feces are eliminated through contractions of the rectal muscles. You help this process by a voluntary procedure called Valsalva’s maneuver, in which you increase intra-abdominal pressure by contracting your diaphragm and abdominal wall muscles, and closing your glottis.

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.

2. Digestive System


Although a minimal amount of carbohydrate digestion occurs in the mouth, chemical digestion really gets underway in the stomach. An expansion of the alimentary canal that lies immediately inferior to the esophagus, the stomach links the esophagus to the first part of the small intestine (the duodenum) and is relatively fixed in place at its esophageal and duodenal ends. In between, however, it can be a highly active structure, contracting and continually changing position and size. These contractions provide mechanical assistance to digestion.

The empty stomach is only about the size of your fist, but can stretch to hold as much as 4 liters of food and fluid, or more than 75 times its empty volume, and then return to its resting size when empty. Although you might think that the size of a person’s stomach is related to how much food that individual consumes, body weight does not correlate with stomach size. Rather, when you eat greater quantities of food—such as at holiday dinner—you stretch the stomach more than when you eat less.

Popular culture tends to refer to the stomach as the location where all digestion takes place. Of course, this is not true. An important function of the stomach is to serve as a temporary holding chamber. You can ingest a meal far more quickly than it can be digested and absorbed by the small intestine. Thus, the stomach holds food and parses only small amounts into the small intestine at a time. Foods are not processed in the order they are eaten; rather, they are mixed together with digestive juices in the stomach until they are converted into chyme, which is released into the small intestine.

As you will see in the sections that follow, the stomach plays several important roles in chemical digestion, including the continued digestion of carbohydrates and the initial digestion of proteins and triglycerides. Little if any nutrient absorption occurs in the stomach, with the exception of the negligible amount of nutrients in alcohol.


There are four main regions in the stomach: the cardia, fundus, body, and pylorus (Figure 23.15). The cardia (or cardiac region) is the point where the esophagus connects to the stomach and through which food passes into the stomach. Located inferior to the diaphragm, above and to the left of the cardia, is the dome-shaped fundus. Below the fundus is the body, the main part of the stomach. The funnel-shaped pylorus connects the stomach to the duodenum. The wider end of the funnel, the pyloric antrum, connects to the body of the stomach. The narrower end is called the pyloric canal, which connects to the duodenum. The smooth muscle pyloric sphincter is located at this latter point of connection and controls stomach emptying. In the absence of food, the stomach deflates inward, and its mucosa and submucosa fall into a large fold called a ruga.

This image shows a cross-section of the stomach, and the major parts: the cardia, fundus, body and pylorus are labeled.

Figure 23.15Stomach The stomach has four major regions: the cardia, fundus, body, and pylorus. The addition of an inner oblique smooth muscle layer gives the muscularis the ability to vigorously churn and mix food.

The convex lateral surface of the stomach is called the greater curvature; the concave medial border is the lesser curvature. The stomach is held in place by the lesser omentum, which extends from the liver to the lesser curvature, and the greater omentum, which runs from the greater curvature to the posterior abdominal wall.


The wall of the stomach is made of the same four layers as most of the rest of the alimentary canal, but with adaptations to the mucosa and muscularis for the unique functions of this organ. In addition to the typical circular and longitudinal smooth muscle layers, the muscularis has an inner oblique smooth muscle layer (Figure 23.16). As a result, in addition to moving food through the canal, the stomach can vigorously churn food, mechanically breaking it down into smaller particles.

This diagram shows the histological cross-section of the stomach. The left panel shows the stomach and the center panel shows a magnified view of a small region including the epithelium and the gastric glands. The right panel shows a further magnification of the mucosa and the different cell types are labeled.

Figure 23.16Histology of the Stomach The stomach wall is adapted for the functions of the stomach. In the epithelium, gastric pits lead to gastric glands that secrete gastric juice. The gastric glands (one gland is shown enlarged on the right) contain different types of cells that secrete a variety of enzymes, including hydrochloride acid, which activates the protein-digesting enzyme pepsin.

The stomach mucosa’s epithelial lining consists only of surface mucus cells, which secrete a protective coat of alkaline mucus. A vast number of gastric pits dot the surface of the epithelium, giving it the appearance of a well-used pincushion, and mark the entry to each gastric gland, which secretes a complex digestive fluid referred to as gastric juice.

Although the walls of the gastric pits are made up primarily of mucus cells, the gastric glands are made up of different types of cells. The glands of the cardia and pylorus are composed primarily of mucus-secreting cells. Cells that make up the pyloric antrum secrete mucus and a number of hormones, including the majority of the stimulatory hormone, gastrin. The much larger glands of the fundus and body of the stomach, the site of most chemical digestion, produce most of the gastric secretions. These glands are made up of a variety of secretory cells. These include parietal cells, chief cells, mucous neck cells, and enteroendocrine cells.

Parietal cells—Located primarily in the middle region of the gastric glands are parietal cells, which are among the most highly differentiated of the body’s epithelial cells. These relatively large cells produce both hydrochloric acid (HCl) and intrinsic factor. HCl is responsible for the high acidity (pH 1.5 to 3.5) of the stomach contents and is needed to activate the protein-digesting enzyme, pepsin. The acidity also kills much of the bacteria you ingest with food and helps to denature proteins, making them more available for enzymatic digestion. Intrinsic factor is a glycoprotein necessary for the absorption of vitamin B12 in the small intestine.

Chief cells—Located primarily in the basal regions of gastric glands are chief cells, which secrete pepsinogen, the inactive proenzyme form of pepsin. HCl is necessary for the conversion of pepsinogen to pepsin.

Mucous neck cells—Gastric glands in the upper part of the stomach contain mucous neck cells that secrete thin, acidic mucus that is much different from the mucus secreted by the goblet cells of the surface epithelium. The role of this mucus is not currently known.

Enteroendocrine cells—Finally, enteroendocrine cells found in the gastric glands secrete various hormones into the interstitial fluid of the lamina propria. These include gastrin, which is released mainly by enteroendocrine G cells.

Table 23.6 describes the digestive functions of important hormones secreted by the stomach.

Hormones Secreted by the Stomach

HormoneProduction siteProduction stimulusTarget organAction
GastrinStomach mucosa, mainly G cells of the pyloric antrumPresence of peptides and amino acids in stomachStomachIncreases secretion by gastric glands; promotes gastric emptying
GastrinStomach mucosa, mainly G cells of the pyloric antrumPresence of peptides and amino acids in stomachSmall intestinePromotes intestinal muscle contraction
GastrinStomach mucosa, mainly G cells of the pyloric antrumPresence of peptides and amino acids in stomachIleocecal valveRelaxes valve
GastrinStomach mucosa, mainly G cells of the pyloric antrumPresence of peptides and amino acids in stomachLarge intestineTriggers mass movements
GhrelinStomach mucosa, mainly fundusFasting state (levels increase just prior to meals)HypothalamusRegulates food intake, primarily by stimulating hunger and satiety
HistamineStomach mucosaPresence of food in the stomachStomachStimulates parietal cells to release HCl
SerotoninStomach mucosaPresence of food in the stomachStomachContracts stomach muscle
SomatostatinMucosa of stomach, especially pyloric antrum; also duodenumPresence of food in the stomach; sympathetic axon stimulationStomachRestricts all gastric secretions, gastric motility, and emptying
SomatostatinMucosa of stomach, especially pyloric antrum; also duodenumPresence of food in the stomach; sympathetic axon stimulationPancreasRestricts pancreatic secretions
SomatostatinMucosa of stomach, especially pyloric antrum; also duodenumPresence of food in the stomach; sympathetic axon stimulationSmall intestineReduces intestinal absorption by reducing blood flow


Gastric Secretion

The secretion of gastric juice is controlled by both nerves and hormones. Stimuli in the brain, stomach, and small intestine activate or inhibit gastric juice production. This is why the three phases of gastric secretion are called the cephalic, gastric, and intestinal phases (Figure 23.17). However, once gastric secretion begins, all three phases can occur simultaneously.

This flowchart shows the three different phases of gastric secretion. The top panel shows the cephalic phase, the middle panel shows the gastric phase and the bottom panel shows the intestinal phase.

Figure 23.17The Three Phases of Gastric Secretion Gastric secretion occurs in three phases: cephalic, gastric, and intestinal. During each phase, the secretion of gastric juice can be stimulated or inhibited.

The cephalic phase (reflex phase) of gastric secretion, which is relatively brief, takes place before food enters the stomach. The smell, taste, sight, or thought of food triggers this phase. For example, when you bring a piece of sushi to your lips, impulses from receptors in your taste buds or the nose are relayed to your brain, which returns signals that increase gastric secretion to prepare your stomach for digestion. This enhanced secretion is a conditioned reflex, meaning it occurs only if you like or want a particular food. Depression and loss of appetite can suppress the cephalic reflex.

The gastric phase of secretion lasts 3 to 4 hours, and is set in motion by local neural and hormonal mechanisms triggered by the entry of food into the stomach. For example, when your sushi reaches the stomach, it creates distention that activates the stretch receptors. This stimulates parasympathetic neurons to release acetylcholine, which then provokes increased secretion of gastric juice.

Partially digested proteins, caffeine, and rising pH stimulate the release of gastrin from enteroendocrine G cells, which in turn induces parietal cells to increase their production of HCl, which is needed to create an acidic environment for the conversion of pepsinogen to pepsin, and protein digestion. Additionally, the release of gastrin activates vigorous smooth muscle contractions. However, it should be noted that the stomach does have a natural means of avoiding excessive acid secretion and potential heartburn. Whenever pH levels drop too low, cells in the stomach react by suspending HCl secretion and increasing mucous secretions.

The intestinal phase of gastric secretion has both excitatory and inhibitory elements. The duodenum has a major role in regulating the stomach and its emptying. When partially digested food fills the duodenum, intestinal mucosal cells release a hormone called intestinal (enteric) gastrin, which further excites gastric juice secretion. This stimulatory activity is brief, however, because when the intestine distends with chyme, the enterogastric reflex inhibits secretion. One of the effects of this reflex is to close the pyloric sphincter, which blocks additional chyme from entering the duodenum.

The Mucosal Barrier

The mucosa of the stomach is exposed to the highly corrosive acidity of gastric juice. Gastric enzymes that can digest protein can also digest the stomach itself. The stomach is protected from self-digestion by the mucosal barrier. This barrier has several components. First, the stomach wall is covered by a thick coating of bicarbonate-rich mucus. This mucus forms a physical barrier, and its bicarbonate ions neutralize acid. Second, the epithelial cells of the stomach’s mucosa meet at tight junctions, which block gastric juice from penetrating the underlying tissue layers. Finally, stem cells located where gastric glands join the gastric pits quickly replace damaged epithelial mucosal cells, when the epithelial cells are shed. In fact, the surface epithelium of the stomach is completely replaced every 3 to 6 days.

Digestive Functions of the Stomach

The stomach participates in virtually all the digestive activities with the exception of ingestion and defecation. Although almost all absorption takes place in the small intestine, the stomach does absorb some drugs, such as alcohol and aspirin.

Mechanical Digestion

Within a few moments after food enters your stomach, mixing waves begin to occur at intervals of approximately 20 seconds. A mixing wave is a unique type of peristalsis that mixes and softens the food with gastric juices to create chyme. The initial mixing waves are relatively gentle, but these are followed by more intense waves, starting at the body of the stomach and increasing in force as they reach the pylorus. It is fair to say that long before your sushi exits through the pyloric sphincter, it bears little resemblance to the sushi you ate.

The pylorus, which holds around 30 mL (1 fluid ounce) of chyme, acts as a filter, permitting only liquids and small food particles to pass through the mostly, but not fully, closed pyloric sphincter. In a process called gastric emptying, rhythmic mixing waves force about 3 mL of chyme at a time through the pyloric sphincter and into the duodenum. Release of a greater amount of chyme at one time would overwhelm the capacity of the small intestine to handle it. The rest of the chyme is pushed back into the body of the stomach, where it continues mixing. This process is repeated when the next mixing waves force more chyme into the duodenum.

Gastric emptying is regulated by both the stomach and the duodenum. The presence of chyme in the duodenum activates receptors that inhibit gastric secretion. This prevents additional chyme from being released by the stomach before the duodenum is ready to process it.

Chemical Digestion

The fundus plays an important role, because it stores both undigested food and gases that are released during the process of chemical digestion. Food may sit in the fundus of the stomach for a while before being mixed with the chyme. While the food is in the fundus, the digestive activities of salivary amylase continue until the food begins mixing with the acidic chyme. Ultimately, mixing waves incorporate this food with the chyme, the acidity of which inactivates salivary amylase and activates lingual lipase. Lingual lipase then begins breaking down triglycerides into free fatty acids, and mono- and diglycerides.

The breakdown of protein begins in the stomach through the actions of HCl and the enzyme pepsin.

Its numerous digestive functions notwithstanding, there is only one stomach function necessary to life: the production of intrinsic factor. The intestinal absorption of vitamin B12, which is necessary for both the production of mature red blood cells and normal neurological functioning, cannot occur without intrinsic factor. People who undergo total gastrectomy (stomach removal)—for life-threatening stomach cancer, for example—can survive with minimal digestive dysfunction if they receive vitamin B12 injections.

The contents of the stomach are completely emptied into the duodenum within 2 to 4 hours after you eat a meal. Different types of food take different amounts of time to process. Foods heavy in carbohydrates empty fastest, followed by high-protein foods. Meals with a high triglyceride content remain in the stomach the longest. Since enzymes in the small intestine digest fats slowly, food can stay in the stomach for 6 hours or longer when the duodenum is processing fatty chyme. However, note that this is still a fraction of the 24 to 72 hours that full digestion typically takes from start to finish.

2. Digestive System

Mouth, Pharynx, and Esophagus

In this section, you will examine the anatomy and functions of the three main organs of the upper alimentary canal—the mouth, pharynx, and esophagus—as well as three associated accessory organs—the tongue, salivary glands, and teeth.

The Mouth

The cheeks, tongue, and palate frame the mouth, which is also called the oral cavity (or buccal cavity). The structures of the mouth are illustrated in Figure 23.7.

At the entrance to the mouth are the lips, or labia (singular = labium). Their outer covering is skin, which transitions to a mucous membrane in the mouth proper. Lips are very vascular with a thin layer of keratin; hence, the reason they are “red.” They have a huge representation on the cerebral cortex, which probably explains the human fascination with kissing! The lips cover the orbicularis oris muscle, which regulates what comes in and goes out of the mouth. The labial frenulum is a midline fold of mucous membrane that attaches the inner surface of each lip to the gum. The cheeks make up the oral cavity’s sidewalls. While their outer covering is skin, their inner covering is mucous membrane. This membrane is made up of non-keratinized, stratified squamous epithelium. Between the skin and mucous membranes are connective tissue and buccinator muscles. The next time you eat some food, notice how the buccinator muscles in your cheeks and the orbicularis oris muscle in your lips contract, helping you keep the food from falling out of your mouth. Additionally, notice how these muscles work when you are speaking.

The pocket-like part of the mouth that is framed on the inside by the gums and teeth, and on the outside by the cheeks and lips is called the oral vestibule. Moving farther into the mouth, the opening between the oral cavity and throat (oropharynx) is called the fauces (like the kitchen “faucet”). The main open area of the mouth, or oral cavity proper, runs from the gums and teeth to the fauces.

When you are chewing, you do not find it difficult to breathe simultaneously. The next time you have food in your mouth, notice how the arched shape of the roof of your mouth allows you to handle both digestion and respiration at the same time. This arch is called the palate. The anterior region of the palate serves as a wall (or septum) between the oral and nasal cavities as well as a rigid shelf against which the tongue can push food. It is created by the maxillary and palatine bones of the skull and, given its bony structure, is known as the hard palate. If you run your tongue along the roof of your mouth, you’ll notice that the hard palate ends in the posterior oral cavity, and the tissue becomes fleshier. This part of the palate, known as the soft palate, is composed mainly of skeletal muscle. You can therefore manipulate, subconsciously, the soft palate—for instance, to yawn, swallow, or sing (see Figure 23.7).

This diagram shows the structure of the mouth. The teeth, lips, tongue, gums and many other parts are labeled.

Figure 23.7Mouth The mouth includes the lips, tongue, palate, gums, and teeth.

A fleshy bead of tissue called the uvula drops down from the center of the posterior edge of the soft palate. Although some have suggested that the uvula is a vestigial organ, it serves an important purpose. When you swallow, the soft palate and uvula move upward, helping to keep foods and liquid from entering the nasal cavity. Unfortunately, it can also contribute to the sound produced by snoring. Two muscular folds extend downward from the soft palate, on either side of the uvula. Toward the front, the palatoglossal arch lies next to the base of the tongue; behind it, the palatopharyngeal arch forms the superior and lateral margins of the fauces. Between these two arches are the palatine tonsils, clusters of lymphoid tissue that protect the pharynx. The lingual tonsils are located at the base of the tongue.

The Tongue

Perhaps you have heard it said that the tongue is the strongest muscle in the body. Those who stake this claim cite its strength proportionate to its size. Although it is difficult to quantify the relative strength of different muscles, it remains indisputable that the tongue is a workhorse, facilitating ingestion, mechanical digestion, chemical digestion (lingual lipase), sensation (of taste, texture, and temperature of food), swallowing, and vocalization.

The tongue is attached to the mandible, the styloid processes of the temporal bones, and the hyoid bone. The hyoid is unique in that it only distantly/indirectly articulates with other bones. The tongue is positioned over the floor of the oral cavity. A medial septum extends the entire length of the tongue, dividing it into symmetrical halves.

Beneath its mucous membrane covering, each half of the tongue is composed of the same number and type of intrinsic and extrinsic skeletal muscles. The intrinsic muscles (those within the tongue) are the longitudinalis inferior, longitudinalis superior, transversus linguae, and verticalis linguae muscles. These allow you to change the size and shape of your tongue, as well as to stick it out, if you wish. Having such a flexible tongue facilitates both swallowing and speech.

As you learned in your study of the muscular system, the extrinsic muscles of the tongue are the palatoglossus, hyoglossus, styloglossus, and genioglossus muscles. These muscles originate outside the tongue and insert into connective tissues within the tongue. The mylohyoid is responsible for raising the tongue, the hyoglossus pulls it down and back, the styloglossus pulls it up and back, and the genioglossus pulls it forward. Working in concert, these muscles perform three important digestive functions in the mouth: (1) position food for optimal chewing, (2) gather food into a bolus (rounded mass), and (3) position food so it can be swallowed.

The top and sides of the tongue are studded with papillae, extensions of lamina propria of the mucosa, which are covered in stratified squamous epithelium (Figure 23.8). Fungiform papillae, which are mushroom shaped, cover a large area of the tongue; they tend to be larger toward the rear of the tongue and smaller on the tip and sides. In contrast, filiform papillae are long and thin. Fungiform papillae contain taste buds, and filiform papillae have touch receptors that help the tongue move food around in the mouth. The filiform papillae create an abrasive surface that performs mechanically, much like a cat’s rough tongue that is used for grooming. Lingual glands in the lamina propria of the tongue secrete mucus and a watery serous fluid that contains the enzyme lingual lipase, which plays a minor role in breaking down triglycerides but does not begin working until it is activated in the stomach. A fold of mucous membrane on the underside of the tongue, the lingual frenulum, tethers the tongue to the floor of the mouth. People with the congenital anomaly ankyloglossia, also known by the non-medical term “tongue tie,” have a lingual frenulum that is too short or otherwise malformed. Severe ankyloglossia can impair speech and must be corrected with surgery.

This diagram shows the structure of the tongue and different parts of the tongue are labeled.

Figure 23.8Tongue This superior view of the tongue shows the locations and types of lingual papillae.

The Salivary Glands

Many small salivary glands are housed within the mucous membranes of the mouth and tongue. These minor exocrine glands are constantly secreting saliva, either directly into the oral cavity or indirectly through ducts, even while you sleep. In fact, an average of 1 to 1.5 liters of saliva is secreted each day. Usually just enough saliva is present to moisten the mouth and teeth. Secretion increases when you eat, because saliva is essential to moisten food and initiate the chemical breakdown of carbohydrates. Small amounts of saliva are also secreted by the labial glands in the lips. In addition, the buccal glands in the cheeks, palatal glands in the palate, and lingual glands in the tongue help ensure that all areas of the mouth are supplied with adequate saliva.

The Major Salivary Glands

Outside the oral mucosa are three pairs of major salivary glands, which secrete the majority of saliva into ducts that open into the mouth:

  • The submandibular glands, which are in the floor of the mouth, secrete saliva into the mouth through the submandibular ducts.
  • The sublingual glands, which lie below the tongue, use the lesser sublingual ducts to secrete saliva into the oral cavity.
  • The parotid glands lie between the skin and the masseter muscle, near the ears. They secrete saliva into the mouth through the parotid duct, which is located near the second upper molar tooth (Figure 23.9).


Saliva is essentially (98 to 99.5 percent) water. The remaining 4.5 percent is a complex mixture of ions, glycoproteins, enzymes, growth factors, and waste products. Perhaps the most important ingredient in saliva from the perspective of digestion is the enzyme salivary amylase, which initiates the breakdown of carbohydrates. Food does not spend enough time in the mouth to allow all the carbohydrates to break down, but salivary amylase continues acting until it is inactivated by stomach acids. Bicarbonate and phosphate ions function as chemical buffers, maintaining saliva at a pH between 6.35 and 6.85. Salivary mucus helps lubricate food, facilitating movement in the mouth, bolus formation, and swallowing. Saliva contains immunoglobulin A, which prevents microbes from penetrating the epithelium, and lysozyme, which makes saliva antimicrobial. Saliva also contains epidermal growth factor, which might have given rise to the adage “a mother’s kiss can heal a wound.”

Each of the major salivary glands secretes a unique formulation of saliva according to its cellular makeup. For example, the parotid glands secrete a watery solution that contains salivary amylase. The submandibular glands have cells similar to those of the parotid glands, as well as mucus-secreting cells. Therefore, saliva secreted by the submandibular glands also contains amylase but in a liquid thickened with mucus. The sublingual glands contain mostly mucous cells, and they secrete the thickest saliva with the least amount of salivary amylase.

This image shows the location of the salivary glands with reference to the teeth. The different salivary glands are labeled.

Figure 23.9Salivary glands The major salivary glands are located outside the oral mucosa and deliver saliva into the mouth through ducts.

Regulation of Salivation

The autonomic nervous system regulates salivation (the secretion of saliva). In the absence of food, parasympathetic stimulation keeps saliva flowing at just the right level for comfort as you speak, swallow, sleep, and generally go about life. Over-salivation can occur, for example, if you are stimulated by the smell of food, but that food is not available for you to eat. Drooling is an extreme instance of the overproduction of saliva. During times of stress, such as before speaking in public, sympathetic stimulation takes over, reducing salivation and producing the symptom of dry mouth often associated with anxiety. When you are dehydrated, salivation is reduced, causing the mouth to feel dry and prompting you to take action to quench your thirst.

Salivation can be stimulated by the sight, smell, and taste of food. It can even be stimulated by thinking about food. You might notice whether reading about food and salivation right now has had any effect on your production of saliva.

How does the salivation process work while you are eating? Food contains chemicals that stimulate taste receptors on the tongue, which send impulses to the superior and inferior salivatory nuclei in the brain stem. These two nuclei then send back parasympathetic impulses through fibers in the glossopharyngeal and facial nerves, which stimulate salivation. Even after you swallow food, salivation is increased to cleanse the mouth and to water down and neutralize any irritating chemical remnants, such as that hot sauce in your burrito. Most saliva is swallowed along with food and is reabsorbed, so that fluid is not lost.

The Teeth

The teeth, or dentes (singular = dens), are organs similar to bones that you use to tear, grind, and otherwise mechanically break down food.

Types of Teeth

During the course of your lifetime, you have two sets of teeth (one set of teeth is a dentition). Your 20 deciduous teeth, or baby teeth, first begin to appear at about 6 months of age. Between approximately age 6 and 12, these teeth are replaced by 32 permanent teeth. Moving from the center of the mouth toward the side, these are as follows (Figure 23.10):

  • The eight incisors, four top and four bottom, are the sharp front teeth you use for biting into food.
  • The four cuspids (or canines) flank the incisors and have a pointed edge (cusp) to tear up food. These fang-like teeth are superb for piercing tough or fleshy foods.
  • Posterior to the cuspids are the eight premolars (or bicuspids), which have an overall flatter shape with two rounded cusps useful for mashing foods.
  • The most posterior and largest are the 12 molars, which have several pointed cusps used to crush food so it is ready for swallowing. The third members of each set of three molars, top and bottom, are commonly referred to as the wisdom teeth, because their eruption is commonly delayed until early adulthood. It is not uncommon for wisdom teeth to fail to erupt; that is, they remain impacted. In these cases, the teeth are typically removed by orthodontic surgery.
This diagram shows the arrangement of permanent and deciduous teeth in human. The permanent teeth are labeled along with the average age at which they emerge. An inset shows the arrangement of the deciduous teeth, with the age at which they emerge listed.

Figure 23.10Permanent and Deciduous Teeth This figure of two human dentitions shows the arrangement of teeth in the maxilla and mandible, and the relationship between the deciduous and permanent teeth.

Anatomy of a Tooth

The teeth are secured in the alveolar processes (sockets) of the maxilla and the mandible. Gingivae (commonly called the gums) are soft tissues that line the alveolar processes and surround the necks of the teeth. Teeth are also held in their sockets by a connective tissue called the periodontal ligament.

The two main parts of a tooth are the crown, which is the portion projecting above the gum line, and the root, which is embedded within the maxilla and mandible. Both parts contain an inner pulp cavity, containing loose connective tissue through which run nerves and blood vessels. The region of the pulp cavity that runs through the root of the tooth is called the root canal. Surrounding the pulp cavity is dentin, a bone-like tissue. In the root of each tooth, the dentin is covered by an even harder bone-like layer called cementum. In the crown of each tooth, the dentin is covered by an outer layer of enamel, the hardest substance in the body (Figure 23.11).

Although enamel protects the underlying dentin and pulp cavity, it is still nonetheless susceptible to mechanical and chemical erosion, or what is known as tooth decay. The most common form, dental caries (cavities) develops when colonies of bacteria feeding on sugars in the mouth release acids that cause soft tissue inflammation and degradation of the calcium crystals of the enamel. The digestive functions of the mouth are summarized in Table 23.4.

This diagram shows a cross-section of a human tooth elucidating its structure. The major parts of the tooth along with the blood vessels are shown.

Figure 23.11The Structure of the Tooth This longitudinal section through a molar in its alveolar socket shows the relationships between enamel, dentin, and pulp.Digestive Functions of the Mouth

Lips and cheeksConfine food between teethFood is chewed evenly during mastication
Salivary glandsSecrete salivaMoisten and lubricate the lining of the mouth and pharynxMoisten, soften, and dissolve foodClean the mouth and teethSalivary amylase breaks down starch
Tongue’s extrinsic musclesMove tongue sideways, and in and outManipulate food for chewingShape food into a bolusManipulate food for swallowing
Tongue’s intrinsic musclesChange tongue shapeManipulate food for swallowing
Taste budsSense food in mouth and sense tasteNerve impulses from taste buds are conducted to salivary nuclei in the brain stem and then to salivary glands, stimulating saliva secretion
Lingual glandsSecrete lingual lipaseActivated in the stomachBreak down triglycerides into fatty acids and diglycerides
TeethShred and crush foodBreak down solid food into smaller particles for deglutition


The Pharynx

The pharynx (throat) is involved in both digestion and respiration. It receives food and air from the mouth, and air from the nasal cavities. When food enters the pharynx, involuntary muscle contractions close off the air passageways.

A short tube of skeletal muscle lined with a mucous membrane, the pharynx runs from the posterior oral and nasal cavities to the opening of the esophagus and larynx. It has three subdivisions. The most superior, the nasopharynx, is involved only in breathing and speech. The other two subdivisions, the oropharynx and the laryngopharynx, are used for both breathing and digestion. The oropharynx begins inferior to the nasopharynx and is continuous below with the laryngopharynx (Figure 23.12). The inferior border of the laryngopharynx connects to the esophagus, whereas the anterior portion connects to the larynx, allowing air to flow into the bronchial tree.

This diagram shows the cross-section of a human face and highlights the location of the pharynx, which runs from the nostrils to the esophagus and the larynx.

Figure 23.12Pharynx The pharynx runs from the nostrils to the esophagus and the larynx.

Histologically, the wall of the oropharynx is similar to that of the oral cavity. The mucosa includes a stratified squamous epithelium that is endowed with mucus-producing glands. During swallowing, the elevator skeletal muscles of the pharynx contract, raising and expanding the pharynx to receive the bolus of food. Once received, these muscles relax and the constrictor muscles of the pharynx contract, forcing the bolus into the esophagus and initiating peristalsis.

Usually during swallowing, the soft palate and uvula rise reflexively to close off the entrance to the nasopharynx. At the same time, the larynx is pulled superiorly and the cartilaginous epiglottis, its most superior structure, folds inferiorly, covering the glottis (the opening to the larynx); this process effectively blocks access to the trachea and bronchi. When the food “goes down the wrong way,” it goes into the trachea. When food enters the trachea, the reaction is to cough, which usually forces the food up and out of the trachea, and back into the pharynx.

The Esophagus

The esophagus is a muscular tube that connects the pharynx to the stomach. It is approximately 25.4 cm (10 in) in length, located posterior to the trachea, and remains in a collapsed form when not engaged in swallowing. As you can see in Figure 23.13, the esophagus runs a mainly straight route through the mediastinum of the thorax. To enter the abdomen, the esophagus penetrates the diaphragm through an opening called the esophageal hiatus.

Passage of Food through the Esophagus

The upper esophageal sphincter, which is continuous with the inferior pharyngeal constrictor, controls the movement of food from the pharynx into the esophagus. The upper two-thirds of the esophagus consists of both smooth and skeletal muscle fibers, with the latter fading out in the bottom third of the esophagus. Rhythmic waves of peristalsis, which begin in the upper esophagus, propel the bolus of food toward the stomach. Meanwhile, secretions from the esophageal mucosa lubricate the esophagus and food. Food passes from the esophagus into the stomach at the lower esophageal sphincter (also called the gastroesophageal or cardiac sphincter). Recall that sphincters are muscles that surround tubes and serve as valves, closing the tube when the sphincters contract and opening it when they relax. The lower esophageal sphincter relaxes to let food pass into the stomach, and then contracts to prevent stomach acids from backing up into the esophagus. Surrounding this sphincter is the muscular diaphragm, which helps close off the sphincter when no food is being swallowed. When the lower esophageal sphincter does not completely close, the stomach’s contents can reflux (that is, back up into the esophagus), causing heartburn or gastroesophageal reflux disease (GERD).

This diagram shows the esophagus, going from the mouth to the stomach. The upper and the lower esophageal sphincter are labeled.

Figure 23.13Esophagus The upper esophageal sphincter controls the movement of food from the pharynx to the esophagus. The lower esophageal sphincter controls the movement of food from the esophagus to the stomach.

Histology of the Esophagus

The mucosa of the esophagus is made up of an epithelial lining that contains non-keratinized, stratified squamous epithelium, with a layer of basal and parabasal cells. This epithelium protects against erosion from food particles. The mucosa’s lamina propria contains mucus-secreting glands. The muscularis layer changes according to location: In the upper third of the esophagus, the muscularis is skeletal muscle. In the middle third, it is both skeletal and smooth muscle. In the lower third, it is smooth muscle. As mentioned previously, the most superficial layer of the esophagus is called the adventitia, not the serosa. In contrast to the stomach and intestines, the loose connective tissue of the adventitia is not covered by a fold of visceral peritoneum. The digestive functions of the esophagus are identified in Table 23.5.Digestive Functions of the Esophagus

Upper esophageal sphincter relaxationAllows the bolus to move from the laryngopharynx to the esophagus
PeristalsisPropels the bolus through the esophagus
Lower esophageal sphincter relaxationAllows the bolus to move from the esophagus into the stomach and prevents chyme from entering the esophagus
Mucus secretionLubricates the esophagus, allowing easy passage of the bolus



Deglutition is another word for swallowing—the movement of food from the mouth to the stomach. The entire process takes about 4 to 8 seconds for solid or semisolid food, and about 1 second for very soft food and liquids. Although this sounds quick and effortless, deglutition is, in fact, a complex process that involves both the skeletal muscle of the tongue and the muscles of the pharynx and esophagus. It is aided by the presence of mucus and saliva. There are three stages in deglutition: the voluntary phase, the pharyngeal phase, and the esophageal phase (Figure 23.14). The autonomic nervous system controls the latter two phases.

This figure shows the three different phases of deglutition. The left panel shows the voluntary phase, the middle panel shows the pharyngeal phase and the right panel’s shows the esophageal phase.

Figure 23.14Deglutition Deglutition includes the voluntary phase and two involuntary phases: the pharyngeal phase and the esophageal phase.

The Voluntary Phase

The voluntary phase of deglutition (also known as the oral or buccal phase) is so called because you can control when you swallow food. In this phase, chewing has been completed and swallowing is set in motion. The tongue moves upward and backward against the palate, pushing the bolus to the back of the oral cavity and into the oropharynx. Other muscles keep the mouth closed and prevent food from falling out. At this point, the two involuntary phases of swallowing begin.

The Pharyngeal Phase

In the pharyngeal phase, stimulation of receptors in the oropharynx sends impulses to the deglutition center (a collection of neurons that controls swallowing) in the medulla oblongata. Impulses are then sent back to the uvula and soft palate, causing them to move upward and close off the nasopharynx. The laryngeal muscles also constrict to prevent aspiration of food into the trachea. At this point, deglutition apnea takes place, which means that breathing ceases for a very brief time. Contractions of the pharyngeal constrictor muscles move the bolus through the oropharynx and laryngopharynx. Relaxation of the upper esophageal sphincter then allows food to enter the esophagus.

The Esophageal Phase

The entry of food into the esophagus marks the beginning of the esophageal phase of deglutition and the initiation of peristalsis. As in the previous phase, the complex neuromuscular actions are controlled by the medulla oblongata. Peristalsis propels the bolus through the esophagus and toward the stomach. The circular muscle layer of the muscularis contracts, pinching the esophageal wall and forcing the bolus forward. At the same time, the longitudinal muscle layer of the muscularis also contracts, shortening this area and pushing out its walls to receive the bolus. In this way, a series of contractions keeps moving food toward the stomach. When the bolus nears the stomach, distention of the esophagus initiates a short reflex relaxation of the lower esophageal sphincter that allows the bolus to pass into the stomach. During the esophageal phase, esophageal glands secrete mucus that lubricates the bolus and minimizes friction.

2. Digestive System

Digestive System Processes and Regulation

The digestive system uses mechanical and chemical activities to break food down into absorbable substances during its journey through the digestive system. Table 23.3 provides an overview of the basic functions of the digestive organs.

Functions of the Digestive Organs

OrganMajor functionsOther functions
MouthIngests foodChews and mixes foodBegins chemical breakdown of carbohydratesMoves food into the pharynxBegins breakdown of lipids via lingual lipaseMoistens and dissolves food, allowing you to taste itCleans and lubricates the teeth and oral cavityHas some antimicrobial activity
PharynxPropels food from the oral cavity to the esophagusLubricates food and passageways
EsophagusPropels food to the stomachLubricates food and passageways
StomachMixes and churns food with gastric juices to form chymeBegins chemical breakdown of proteinsReleases food into the duodenum as chymeAbsorbs some fat-soluble substances (for example, alcohol, aspirin)Possesses antimicrobial functionsStimulates protein-digesting enzymesSecretes intrinsic factor required for vitamin B12 absorption in small intestine
Small intestineMixes chyme with digestive juicesPropels food at a rate slow enough for digestion and absorptionAbsorbs breakdown products of carbohydrates, proteins, lipids, and nucleic acids, along with vitamins, minerals, and waterPerforms physical digestion via segmentationProvides optimal medium for enzymatic activity
Accessory organsLiver: produces bile salts, which emulsify lipids, aiding their digestion and absorptionGallbladder: stores, concentrates, and releases bilePancreas: produces digestive enzymes and bicarbonateBicarbonate-rich pancreatic juices help neutralize acidic chyme and provide optimal environment for enzymatic activity
Large intestineFurther breaks down food residuesAbsorbs most residual water, electrolytes, and vitamins produced by enteric bacteriaPropels feces toward rectumEliminates fecesFood residue is concentrated and temporarily stored prior to defecationMucus eases passage of feces through colon


Digestive Processes:

The processes of digestion include six activities: ingestion, propulsion, mechanical or physical digestion, chemical digestion, absorption, and defecation.

The first of these processes, ingestion, refers to the entry of food into the alimentary canal through the mouth. There, the food is chewed and mixed with saliva, which contains enzymes that begin breaking down the carbohydrates in the food plus some lipid digestion via lingual lipase. Chewing increases the surface area of the food and allows an appropriately sized bolus to be produced.

Food leaves the mouth when the tongue and pharyngeal muscles propel it into the esophagus. This act of swallowing, the last voluntary act until defecation, is an example of propulsion, which refers to the movement of food through the digestive tract. It includes both the voluntary process of swallowing and the involuntary process of peristalsis. Peristalsis consists of sequential, alternating waves of contraction and relaxation of alimentary wall smooth muscles, which act to propel food along (Figure 23.5). These waves also play a role in mixing food with digestive juices. Peristalsis is so powerful that foods and liquids you swallow enter your stomach even if you are standing on your head.

This image shows the peristaltic movement of food. In the left image, the food bolus is towards the top of the esophagus and arrows pointing downward show the direction of movement of the peristaltic wave. In the center image, the food bolus and the wave movement are closer to the center of the esophagus and in the right image, the bolus and the wave are close to the bottom end of the esophagus.

Figure 23.5 Peristalsis Peristalsis moves food through the digestive tract with alternating waves of muscle contraction and relaxation.

Digestion includes both mechanical and chemical processes. Mechanical digestion is a purely physical process that does not change the chemical nature of the food. Instead, it makes the food smaller to increase both surface area and mobility. It includes mastication, or chewing, as well as tongue movements that help break food into smaller bits and mix food with saliva. Although there may be a tendency to think that mechanical digestion is limited to the first steps of the digestive process, it occurs after the food leaves the mouth, as well. The mechanical churning of food in the stomach serves to further break it apart and expose more of its surface area to digestive juices, creating an acidic “soup” called chymeSegmentation, which occurs mainly in the small intestine, consists of localized contractions of circular muscle of the muscularis layer of the alimentary canal. These contractions isolate small sections of the intestine, moving their contents back and forth while continuously subdividing, breaking up, and mixing the contents. By moving food back and forth in the intestinal lumen, segmentation mixes food with digestive juices and facilitates absorption.

In chemical digestion, starting in the mouth, digestive secretions break down complex food molecules into their chemical building blocks (for example, proteins into separate amino acids). These secretions vary in composition, but typically contain water, various enzymes, acids, and salts. The process is completed in the small intestine.

Food that has been broken down is of no value to the body unless it enters the bloodstream and its nutrients are put to work. This occurs through the process of absorption, which takes place primarily within the small intestine. There, most nutrients are absorbed from the lumen of the alimentary canal into the bloodstream through the epithelial cells that make up the mucosa. Lipids are absorbed into lacteals and are transported via the lymphatic vessels to the bloodstream (the subclavian veins near the heart). The details of these processes will be discussed later.

In defecation, the final step in digestion, undigested materials are removed from the body as feces.

In some cases, a single organ is in charge of a digestive process. For example, ingestion occurs only in the mouth and defecation only in the anus. However, most digestive processes involve the interaction of several organs and occur gradually as food moves through the alimentary canal (Figure 23.6).

This image shows the different processes involved in digestion. The image shows how food travels from the mouth through the major organs. Associated textboxes list the different processes such as propulsion, chemical and mechanical digestion and absorption near the organs where they take place.

Figure 23.6 Digestive Processes The digestive processes are ingestion, propulsion, mechanical digestion, chemical digestion, absorption, and defecation.

Some chemical digestion occurs in the mouth. Some absorption can occur in the mouth and stomach, for example, alcohol and aspirin.

Regulatory Mechanisms:

Neural and endocrine regulatory mechanisms work to maintain the optimal conditions in the lumen needed for digestion and absorption. These regulatory mechanisms, which stimulate digestive activity through mechanical and chemical activity, are controlled both extrinsically and intrinsically.

Neural Controls:

The walls of the alimentary canal contain a variety of sensors that help regulate digestive functions. These include mechanoreceptors, chemoreceptors, and osmoreceptors, which are capable of detecting mechanical, chemical, and osmotic stimuli, respectively. For example, these receptors can sense when the presence of food has caused the stomach to expand, whether food particles have been sufficiently broken down, how much liquid is present, and the type of nutrients in the food (lipids, carbohydrates, and/or proteins). Stimulation of these receptors provokes an appropriate reflex that furthers the process of digestion. This may entail sending a message that activates the glands that secrete digestive juices into the lumen, or it may mean the stimulation of muscles within the alimentary canal, thereby activating peristalsis and segmentation that move food along the intestinal tract.

The walls of the entire alimentary canal are embedded with nerve plexuses that interact with the central nervous system and other nerve plexuses—either within the same digestive organ or in different ones. These interactions prompt several types of reflexes. Extrinsic nerve plexuses orchestrate long reflexes, which involve the central and autonomic nervous systems and work in response to stimuli from outside the digestive system. Short reflexes, on the other hand, are orchestrated by intrinsic nerve plexuses within the alimentary canal wall. These two plexuses and their connections were introduced earlier as the enteric nervous system. Short reflexes regulate activities in one area of the digestive tract and may coordinate local peristaltic movements and stimulate digestive secretions. For example, the sight, smell, and taste of food initiate long reflexes that begin with a sensory neuron delivering a signal to the medulla oblongata. The response to the signal is to stimulate cells in the stomach to begin secreting digestive juices in preparation for incoming food. In contrast, food that distends the stomach initiates short reflexes that cause cells in the stomach wall to increase their secretion of digestive juices.

Hormonal Controls:

A variety of hormones are involved in the digestive process. The main digestive hormone of the stomach is gastrin, which is secreted in response to the presence of food. Gastrin stimulates the secretion of gastric acid by the parietal cells of the stomach mucosa. Other GI hormones are produced and act upon the gut and its accessory organs. Hormones produced by the duodenum include secretin, which stimulates a watery secretion of bicarbonate by the pancreas; cholecystokinin (CCK), which stimulates the secretion of pancreatic enzymes and bile from the liver and release of bile from the gallbladder; and gastric inhibitory peptide, which inhibits gastric secretion and slows gastric emptying and motility. These GI hormones are secreted by specialized epithelial cells, called endocrinocytes, located in the mucosal epithelium of the stomach and small intestine. These hormones then enter the bloodstream, through which they can reach their target organs.

2. Digestive System

Overview of the Digestive System

The function of the digestive system is to break down the foods you eat, release their nutrients, and absorb those nutrients into the body. Although the small intestine is the workhorse of the system, where the majority of digestion occurs, and where most of the released nutrients are absorbed into the blood or lymph, each of the digestive system organs makes a vital contribution to this process (Figure 23.2).

This diagram shows the digestive system of a human being, with the major organs labeled.

Figure 23.2 Components of the Digestive System All digestive organs play integral roles in the life-sustaining process of digestion.

As is the case with all body systems, the digestive system does not work in isolation; it functions cooperatively with the other systems of the body. Consider for example, the interrelationship between the digestive and cardiovascular systems. Arteries supply the digestive organs with oxygen and processed nutrients, and veins drain the digestive tract. These intestinal veins, constituting the hepatic portal system, are unique; they do not return blood directly to the heart. Rather, this blood is diverted to the liver where its nutrients are off-loaded for processing before blood completes its circuit back to the heart.

At the same time, the digestive system provides nutrients to the heart muscle and vascular tissue to support their functioning. The interrelationship of the digestive and endocrine systems is also critical. Hormones secreted by several endocrine glands, as well as endocrine cells of the pancreas, the stomach, and the small intestine, contribute to the control of digestion and nutrient metabolism. In turn, the digestive system provides the nutrients to fuel endocrine function. Table 23.1 gives a quick glimpse at how these other systems contribute to the functioning of the digestive system.Contribution of Other Body Systems to the Digestive System

Body systemBenefits received by the digestive system
CardiovascularBlood supplies digestive organs with oxygen and processed nutrients
EndocrineEndocrine hormones help regulate secretion in digestive glands and accessory organs
IntegumentarySkin helps protect digestive organs and synthesizes vitamin D for calcium absorption
LymphaticMucosa-associated lymphoid tissue and other lymphatic tissue defend against entry of pathogens; lacteals absorb lipids; and lymphatic vessels transport lipids to bloodstream
MuscularSkeletal muscles support and protect abdominal organs
NervousSensory and motor neurons help regulate secretions and muscle contractions in the digestive tract
RespiratoryRespiratory organs provide oxygen and remove carbon dioxide
SkeletalBones help protect and support digestive organs
UrinaryKidneys convert vitamin D into its active form, allowing calcium absorption in the small intestine


Digestive System Organs

The easiest way to understand the digestive system is to divide its organs into two main categories. The first group is the organs that make up the alimentary canal. Accessory digestive organs comprise the second group and are critical for orchestrating the breakdown of food and the assimilation of its nutrients into the body. Accessory digestive organs, despite their name, are critical to the function of the digestive system.

Alimentary Canal Organs

Also called the gastrointestinal (GI) tract or gut, the alimentary canal (aliment- = “to nourish”) is a one-way tube about 7.62 meters (25 feet) in length during life and closer to 10.67 meters (35 feet) in length when measured after death, once smooth muscle tone is lost. The main function of the organs of the alimentary canal is to nourish the body. This tube begins at the mouth and terminates at the anus. Between those two points, the canal is modified as the pharynx, esophagus, stomach, and small and large intestines to fit the functional needs of the body. Both the mouth and anus are open to the external environment; thus, food and wastes within the alimentary canal are technically considered to be outside the body. Only through the process of absorption do the nutrients in food enter into and nourish the body’s “inner space.”

Accessory Structures

Each accessory digestive organ aids in the breakdown of food (Figure 23.3). Within the mouth, the teeth and tongue begin mechanical digestion, whereas the salivary glands begin chemical digestion. Once food products enter the small intestine, the gallbladder, liver, and pancreas release secretions—such as bile and enzymes—essential for digestion to continue. Together, these are called accessory organs because they sprout from the lining cells of the developing gut (mucosa) and augment its function; indeed, you could not live without their vital contributions, and many significant diseases result from their malfunction. Even after development is complete, they maintain a connection to the gut by way of ducts.

Histology of the Alimentary Canal

Throughout its length, the alimentary tract is composed of the same four tissue layers; the details of their structural arrangements vary to fit their specific functions. Starting from the lumen and moving outwards, these layers are the mucosa, submucosa, muscularis, and serosa, which is continuous with the mesentery (see Figure 23.3).

This image shows the cross section of the alimentary canal. The different layers of the alimentary canal are shown as concentric cylinders with major muscles and veins labeled.

Figure 23.3Layers of the Alimentary Canal The wall of the alimentary canal has four basic tissue layers: the mucosa, submucosa, muscularis, and serosa.

The mucosa is referred to as a mucous membrane, because mucus production is a characteristic feature of gut epithelium. The membrane consists of epithelium, which is in direct contact with ingested food, and the lamina propria, a layer of connective tissue analogous to the dermis. In addition, the mucosa has a thin, smooth muscle layer, called the muscularis mucosae (not to be confused with the muscularis layer, described below).

Epithelium—In the mouth, pharynx, esophagus, and anal canal, the epithelium is primarily a non-keratinized, stratified squamous epithelium. In the stomach and intestines, it is a simple columnar epithelium. Notice that the epithelium is in direct contact with the lumen, the space inside the alimentary canal. Interspersed among its epithelial cells are goblet cells, which secrete mucus and fluid into the lumen, and enteroendocrine cells, which secrete hormones into the interstitial spaces between cells. Epithelial cells have a very brief lifespan, averaging from only a couple of days (in the mouth) to about a week (in the gut). This process of rapid renewal helps preserve the health of the alimentary canal, despite the wear and tear resulting from continued contact with foodstuffs.

Lamina propria—In addition to loose connective tissue, the lamina propria contains numerous blood and lymphatic vessels that transport nutrients absorbed through the alimentary canal to other parts of the body. The lamina propria also serves an immune function by housing clusters of lymphocytes, making up the mucosa-associated lymphoid tissue (MALT). These lymphocyte clusters are particularly substantial in the distal ileum where they are known as Peyer’s patches. When you consider that the alimentary canal is exposed to foodborne bacteria and other foreign matter, it is not hard to appreciate why the immune system has evolved a means of defending against the pathogens encountered within it.

Muscularis mucosae—This thin layer of smooth muscle is in a constant state of tension, pulling the mucosa of the stomach and small intestine into undulating folds. These folds dramatically increase the surface area available for digestion and absorption.

As its name implies, the submucosa lies immediately beneath the mucosa. A broad layer of dense connective tissue, it connects the overlying mucosa to the underlying muscularis. It includes blood and lymphatic vessels (which transport absorbed nutrients), and a scattering of submucosal glands that release digestive secretions. Additionally, it serves as a conduit for a dense branching network of nerves, the submucosal plexus, which functions as described below.

The third layer of the alimentary canal is the muscularis (also called the muscularis externa). The muscularis in the small intestine is made up of a double layer of smooth muscle: an inner circular layer and an outer longitudinal layer. The contractions of these layers promote mechanical digestion, expose more of the food to digestive chemicals, and move the food along the canal. In the most proximal and distal regions of the alimentary canal, including the mouth, pharynx, anterior part of the esophagus, and external anal sphincter, the muscularis is made up of skeletal muscle, which gives you voluntary control over swallowing and defecation. The basic two-layer structure found in the small intestine is modified in the organs proximal and distal to it.

The stomach is equipped for its churning function by the addition of a third layer, the oblique muscle. While the colon has two layers like the small intestine, its longitudinal layer is segregated into three narrow parallel bands, the tenia coli, which make it look like a series of pouches rather than a simple tube.

The serosa is the portion of the alimentary canal superficial to the muscularis. Present only in the region of the alimentary canal within the abdominal cavity, it consists of a layer of visceral peritoneum overlying a layer of loose connective tissue. Instead of serosa, the mouth, pharynx, and esophagus have a dense sheath of collagen fibers called the adventitia. These tissues serve to hold the alimentary canal in place near the ventral surface of the vertebral column.

Nerve Supply

As soon as food enters the mouth, it is detected by receptors that send impulses along the sensory neurons of cranial nerves. Without these nerves, not only would your food be without taste, but you would also be unable to feel either the food or the structures of your mouth, and you would be unable to avoid biting yourself as you chew, an action enabled by the motor branches of cranial nerves.

Intrinsic innervation of much of the alimentary canal is provided by the enteric nervous system, which runs from the esophagus to the anus, and contains approximately 100 million motor, sensory, and interneurons (unique to this system compared to all other parts of the peripheral nervous system). These enteric neurons are grouped into two plexuses. The myenteric plexus (plexus of Auerbach) lies in the muscularis layer of the alimentary canal and is responsible for motility, especially the rhythm and force of the contractions of the muscularis. The submucosal plexus (plexus of Meissner) lies in the submucosal layer and is responsible for regulating digestive secretions and reacting to the presence of food (see Figure 23.3).

Extrinsic innervations of the alimentary canal are provided by the autonomic nervous system, which includes both sympathetic and parasympathetic nerves. In general, sympathetic activation (the fight-or-flight response) restricts the activity of enteric neurons, thereby decreasing GI secretion and motility. In contrast, parasympathetic activation (the rest-and-digest response) increases GI secretion and motility by stimulating neurons of the enteric nervous system.

Blood Supply

The blood vessels serving the digestive system have two functions. They transport the protein and carbohydrate nutrients absorbed by mucosal cells after food is digested in the lumen. Lipids are absorbed via lacteals, tiny structures of the lymphatic system. The blood vessels’ second function is to supply the organs of the alimentary canal with the nutrients and oxygen needed to drive their cellular processes.

Specifically, the more anterior parts of the alimentary canal are supplied with blood by arteries branching off the aortic arch and thoracic aorta. Below this point, the alimentary canal is supplied with blood by arteries branching from the abdominal aorta. The celiac trunk services the liver, stomach, and duodenum, whereas the superior and inferior mesenteric arteries supply blood to the remaining small and large intestines.

The veins that collect nutrient-rich blood from the small intestine (where most absorption occurs) empty into the hepatic portal system. This venous network takes the blood into the liver where the nutrients are either processed or stored for later use. Only then does the blood drained from the alimentary canal viscera circulate back to the heart. To appreciate just how demanding the digestive process is on the cardiovascular system, consider that while you are “resting and digesting,” about one-fourth of the blood pumped with each heartbeat enters arteries serving the intestines.

The Peritoneum

The digestive organs within the abdominal cavity are held in place by the peritoneum, a broad serous membranous sac made up of squamous epithelial tissue surrounded by connective tissue. It is composed of two different regions: the parietal peritoneum, which lines the abdominal wall, and the visceral peritoneum, which envelopes the abdominal organs (Figure 23.4). The peritoneal cavity is the space bounded by the visceral and parietal peritoneal surfaces. A few milliliters of watery fluid act as a lubricant to minimize friction between the serosal surfaces of the peritoneum.

This diagram shows the cross section of the abdomen. The peritoneum is made distinguishable from the abdominal organs through darker lines.

Figure 23.4The Peritoneum A cross-section of the abdomen shows the relationship between abdominal organs and the peritoneum (darker lines).

The visceral peritoneum includes multiple large folds that envelope various abdominal organs, holding them to the dorsal surface of the body wall. Within these folds are blood vessels, lymphatic vessels, and nerves that innervate the organs with which they are in contact, supplying their adjacent organs. The five major peritoneal folds are described in Table 23.2. Note that during fetal development, certain digestive structures, including the first portion of the small intestine (called the duodenum), the pancreas, and portions of the large intestine (the ascending and descending colon, and the rectum) remain completely or partially posterior to the peritoneum. Thus, the location of these organs is described as retroperitoneal.The Five Major Peritoneal Folds

Greater omentumApron-like structure that lies superficial to the small intestine and transverse colon; a site of fat deposition in people who are overweight
Falciform ligamentAnchors the liver to the anterior abdominal wall and inferior border of the diaphragm
Lesser omentumSuspends the stomach from the inferior border of the liver; provides a pathway for structures connecting to the liver
MesenteryVertical band of tissue anterior to the lumbar vertebrae and anchoring all of the small intestine except the initial portion (the duodenum)
MesocolonAttaches two portions of the large intestine (the transverse and sigmoid colon) to the posterior abdominal wall


2. Digestive System

Introduction of digestive system

This photograph shows two women eating apples.

Figure 23.1Eating Apples Eating may be one of the simple pleasures in life, but digesting even one apple requires the coordinated work of many organs.

The digestive system is continually at work, yet people seldom appreciate the complex tasks it performs in a choreographed biologic symphony. Consider what happens when you eat an apple. Of course, you enjoy the apple’s taste as you chew it, but in the hours that follow, unless something goes amiss and you get a stomachache, you don’t notice that your digestive system is working. You may be taking a walk or studying or sleeping, having forgotten all about the apple, but your stomach and intestines are busy digesting it and absorbing its vitamins and other nutrients. By the time any waste material is excreted, the body has appropriated all it can use from the apple. In short, whether you pay attention or not, the organs of the digestive system perform their specific functions, allowing you to use the food you eat to keep you going. This chapter examines the structure and functions of these organs, and explores the mechanics and chemistry of the digestive processes.