Invertebrate Digestive Systems

Animals have evolved different types of digestive systems to aid in the digestion of the different foods they consume. The simplest example is that of a gastrovascular cavity and is found in organisms with only one opening for digestion. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly fish, and sea anemones) use this type of digestion. Gastrovascular cavities, as shown in Figure 24.1 a, are typically a blind tube or cavity with only one opening, the “mouth”, which also serves as an “anus”. Ingested material enters the mouth and passes through a hollow, tubular cavity. Cells within the cavity secrete digestive enzymes that breakdown the food. The food particles are engulfed by the cells lining the gastrovascular cavity.

The alimentary canal, shown in Figure 24.1b, is a more advanced system: it consists of one tube with a mouth at one end and an anus at the other. Earthworms are an example of an animal with an alimentary canal. Once the food is ingested through the mouth, it passes through the esophagus and is stored in an organ called the crop; then it passes into the gizzard where it is churned and digested. From the gizzard, the food passes through the intestine, the nutrients are absorbed, and the waste is eliminated as feces, called castings, through the anus.

Vertebrate Digestive Systems

Vertebrates have evolved more complex digestive systems to adapt to their dietary needs. Some animals have a single stomach, while others have multi-chambered stomachs. Birds have developed a digestive system adapted to eating unmasticated food.

Monogastric: Single-chambered Stomach

As the word monogastric suggests, this type of digestive system consists of one (“mono”) stomach chamber (“gastric”). Humans and many animals have a monogastric digestive system as illustrated in Figure 24.2ab. The process of digestion begins with the mouth and the intake of food. The teeth play an important role in masticating (chewing) or physically breaking down food into smaller particles. The enzymes present in saliva also begin to chemically breakdown food. The esophagus is a long tube that connects the mouth to the stomach. Using peristalsis, or wave-like smooth muscle contractions, the muscles of the esophagus push the food towards the stomach. In order to speed up the actions of enzymes in the stomach, the stomach is an extremely acidic environment, with a pH between 1.5 and 2.5. The gastric juices, which include enzymes in the stomach, act on the food particles and continue the process of digestion. Further breakdown of food takes place in the small intestine where enzymes produced by the liver, the small intestine, and the pancreas continue the process of digestion. The nutrients are absorbed into the bloodstream across the epithelial cells lining the walls of the small intestines. The waste material travels on to the large intestine where water is absorbed and the drier waste material is compacted into feces; it is stored until it is excreted through the rectum.

Avian

Birds face special challenges when it comes to obtaining nutrition from food. They do not have teeth and so their digestive system, shown in Figure 24.3 must be able to process un-masticated food. Birds have evolved a variety of beak types that reflect the vast variety in their diet, ranging from seeds and insects to fruits and nuts. Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their body weight low. The stomach of birds has two chambers: the proventriculus, where gastric juices are produced to digest the food before it enters the stomach, and the gizzard, where the food is stored, soaked, and mechanically ground. The undigested material forms food pellets that are sometimes regurgitated. Most of the chemical digestion and absorption happens in the intestine and the waste is excreted through the cloaca.

Evolution Connection

Birds have a highly efficient, simplified digestive system. Recent fossil evidence has shown that the evolutionary divergence of birds from other land animals was characterized by streamlining and simplifying the digestive system. Unlike many other animals, birds do not have teeth to chew their food. In place of lips, they have sharp pointy beaks. The beak, lack of jaws, and the smaller tongue of the birds can be traced back to their dinosaur ancestors. The emergence of these changes seems to coincide with the inclusion of seeds in the bird diet. Seed-eating birds have beaks that are shaped for grabbing seeds and the two-compartment stomach allows for delegation of tasks. Since birds need to remain light in order to fly, their metabolic rates are very high, which means they digest their food very quickly and need to eat often. Contrast this with the ruminants, where the digestion of plant matter takes a very long time

Ruminants

Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating large amounts of roughage or fiber. They have evolved digestive systems that help them digest vast amounts of cellulose. An interesting feature of the ruminants’ mouth is that they do not have upper incisor teeth. They use their lower teeth, tongue and lips to tear and chew their food. From the mouth, the food travels to the esophagus and on to the stomach.

To help digest the large amount of plant material, the stomach of the ruminants is a multi-chambered organ, as illustrated in Figure 24.4. The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that breakdown cellulose and ferment ingested food. The abomasum is the “true” stomach and is the equivalent of the monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary to digest plant material in ruminants. The fermentation process produces large amounts of gas in the stomach chamber, which must be eliminated. As in other animals, the small intestine plays an important role in nutrient absorption, and the large intestine helps in the elimination of waste.

Pseudo-ruminants

Some animals, such as camels and alpacas, are pseudo-ruminants. They eat a lot of plant material and roughage. Digesting plant material is not easy because plant cell walls contain the polymeric sugar molecule cellulose. The digestive enzymes of these animals cannot breakdown cellulose, but microorganisms present in the digestive system can. Therefore, the digestive system must be able to handle large amounts of roughage and breakdown the cellulose. Pseudo-ruminants have a three-chamber stomach in the digestive system. However, their cecum—a pouched organ at the beginning of the large intestine containing many microorganisms that are necessary for the digestion of plant materials—is large and is the site where the roughage is fermented and digested. These animals do not have a rumen but have an omasum, abomasum, and reticulum.

Parts of the Digestive System

The vertebrate digestive system is designed to facilitate the transformation of food matter into the nutrient components that sustain organisms.

Oral Cavity

The oral cavity, or mouth, is the point of entry of food into the digestive system, illustrated in Figure 24.5. The food consumed is broken into smaller particles by mastication, the chewing action of the teeth. All mammals have teeth and can chew their food.

The extensive chemical process of digestion begins in the mouth. As food is being chewed, saliva, produced by the salivary glands, mixes with the food. Saliva is a watery substance produced in the mouths of many animals. There are three major glands that secrete saliva—the parotid, the submandibular, and the sublingual. Saliva contains mucus that moistens food and buffers the pH of the food. Saliva also contains immunoglobulins and lysozymes, which have antibacterial action to reduce tooth decay by inhibiting growth of some bacteria. Saliva also contains an enzyme called salivary amylase that begins the process of converting starches in the food into a disaccharide called maltose. Another enzyme called lipase is produced by the cells in the tongue. Lipases are a class of enzymes that can breakdown triglycerides. The lingual lipase begins the breakdown of fat components in the food. The chewing and wetting action provided by the teeth and saliva prepare the food into a mass called the bolus for swallowing. The tongue helps in swallowing—moving the bolus from the mouth into the pharynx. The pharynx opens to two passageways: the trachea, which leads to the lungs, and the esophagus, which leads to the stomach. The trachea has an opening called the glottis, which is covered by a cartilaginous flap called the epiglottis. When swallowing, the epiglottis closes the glottis and food passes into the esophagus and not the trachea. This arrangement allows food to be kept out of the trachea.

Esophagus

The esophagus is a tubular organ that connects the mouth to the stomach. The chewed and softened food passes through the esophagus after being swallowed. The smooth muscles of the esophagus undergo a series of wave like movements called peristalsis that push the food toward the stomach, as illustrated in Figure 24.6. The peristalsis wave is unidirectional—it moves food from the mouth to the stomach, and reverse movement is not possible. The peristaltic movement of the esophagus is an involuntary reflex; it takes place in response to the act of swallowing.

A ring-like muscle called a sphincter forms valves in the digestive system. The gastro-esophageal sphincter is located at the stomach end of the esophagus. In response to swallowing and the pressure exerted by the bolus of food, this sphincter opens, and the bolus enters the stomach. When there is no swallowing action, this sphincter is shut and prevents the contents of the stomach from traveling up the esophagus. Many animals have a true sphincter; however, in humans, there is no true sphincter, but the esophagus remains closed when there is no swallowing action. Acid reflux or “heartburn” occurs when the acidic digestive juices escape into the esophagus.

Stomach

A large part of digestion occurs in the stomach, shown in Figure 24.7. The stomach is a saclike organ that secretes gastric digestive juices. The pH in the stomach is between 1.5 and 2.5. This highly acidic environment is required for the chemical breakdown of food and the extraction of nutrients. When empty, the stomach is a rather small organ; however, it can expand to up to 20 times its resting size when filled with food. This characteristic is particularly useful for animals that need to eat when food is available.

The stomach is also the major site for protein digestion in animals other than ruminants. Protein digestion is mediated by an enzyme called pepsin in the stomach chamber. Pepsin is secreted by the chief cells in the stomach in an inactive form called pepsinogen. Pepsin breaks peptide bonds and cleaves proteins into smaller polypeptides; it also helps activate more pepsinogen, starting a positive feedback mechanism that generates more pepsin. Another cell type—parietal cells—secrete hydrogen and chloride ions, which combine in the lumen to form hydrochloric acid, the primary acidic component of the stomach juices. Hydrochloric acid helps to convert the inactive pepsinogen to pepsin. The highly acidic environment also kills many microorganisms in the food and, combined with the action of the enzyme pepsin, results in the hydrolysis of protein in the food. Chemical digestion is facilitated by the churning action of the stomach. Contraction and relaxation of smooth muscles mixes the stomach contents about every 20 minutes. The partially digested food and gastric juice mixture is called chyme. Chyme passes from the stomach to the small intestine. Further protein digestion takes place in the small intestine. Gastric emptying occurs within two to six hours after a meal. Only a small amount of chyme is released into the small intestine at a time. The movement of chyme from the stomach into the small intestine is regulated by the pyloric sphincter.

When digesting protein and some fats, the stomach lining must be protected from getting digested by pepsin. There are two points to consider when describing how the stomach lining is protected. First, as previously mentioned, the enzyme pepsin is synthesized in the inactive form. This protects the chief cells, because pepsinogen does not have the same enzyme functionality of pepsin. Second, the stomach has a thick mucus lining that protects the underlying tissue from the action of the digestive juices. When this mucus lining is ruptured, ulcers can form in the stomach. Ulcers are open wounds in or on an organ caused by bacteria (Helicobacter pylori) when the mucus lining is ruptured and fails to reform.

Small Intestine

Chyme moves from the stomach to the small intestine. The small intestine is the organ where the digestion of protein, fats, and carbohydrates is completed. The small intestine is a long tube-like organ with a highly folded surface containing finger-like projections called the villi. The apical surface of each villus has many microscopic projections called microvilli. These structures, illustrated in Figure 24.8, are lined with epithelial cells on the luminal side and allow for the nutrients to be absorbed from the digested food and absorbed into the bloodstream on the other side. The villi and microvilli, with their many folds, increase the surface area of the intestine and increase absorption efficiency of the nutrients. Absorbed nutrients in the blood are carried into the hepatic portal vein, which leads to the liver. There, the liver regulates the distribution of nutrients to the rest of the body and removes toxic substances, including drugs, alcohol, and some pathogens.

The human small intestine is over 6m long and is divided into three parts: the duodenum, the jejunum, and the ileum. The “C-shaped,” fixed part of the small intestine is called the duodenum and is shown in Figure 24.8. The duodenum is separated from the stomach by the pyloric sphincter which opens to allow chyme to move from the stomach to the duodenum. In the duodenum, chyme is mixed with pancreatic juices in an alkaline solution rich in bicarbonate that neutralizes the acidity of chyme and acts as a buffer. Pancreatic juices also contain several digestive enzymes. Digestive juices from the pancreas, liver, and gallbladder, as well as from gland cells of the intestinal wall itself, enter the duodenum. Bile is produced in the liver and stored and concentrated in the gallbladder. Bile contains bile salts which emulsify lipids while the pancreas produces enzymes that catabolize starches, disaccharides, proteins, and fats. These digestive juices breakdown the food particles in the chyme into glucose, triglycerides, and amino acids. The bulk of chemical digestion of food takes place in the duodenum. Absorption of fatty acids also takes place in the duodenum.

The second part of the small intestine is called the jejunum, shown in Figure 24.8. Here, hydrolysis of nutrients is continued while most of the carbohydrates and amino acids are absorbed through the intestinal lining. Some chemical digestion and the bulk of nutrient absorption occurs in the jejunum.

The ileum, also illustrated in Figure 24.8 is the last part of the small intestine and here the bile salts and vitamins are absorbed into the bloodstream. The undigested food is sent to the colon from the ileum via peristaltic movements of the muscle. The ileum ends and the large intestine begins at the ileocecal valve. The vermiform, “worm-like,” appendix is located at the ileocecal valve. The appendix of humans secretes no enzymes and has an insignificant role in immunity.

Large Intestine

The large intestine, illustrated in Figure 24.9, reabsorbs the water from the undigested food material and processes the waste material. The human large intestine is much smaller in length compared to the small intestine but larger in diameter. It has three parts: the cecum, the colon, and the rectum. The cecum joins the ileum to the colon and is the receiving pouch for the waste matter. The colon is home to many bacteria or “intestinal flora” that aid in the digestive processes. The colon can be divided into four regions, the ascending colon, the transverse colon, the descending colon, and the sigmoid colon. The main functions of the colon are to extract the water and mineral salts from undigested food, and to store waste material. Carnivorous mammals have a shorter large intestine compared to herbivorous mammals due to their diet.

Rectum and Anus

The rectum is the terminal end of the large intestine, as shown in Figure 24.9. The primary role of the rectum is to store the feces until defecation. The feces are propelled using peristaltic movements during elimination. The anus is an opening at the far-end of the digestive tract and is the exit point for the waste material. Two sphincters between the rectum and anus control elimination: the inner sphincter is involuntary and the outer sphincter is voluntary.

Accessory Organs

The organs discussed above are the organs of the digestive tract through which food passes. Accessory organs are organs that add secretions (enzymes) that catabolize food into nutrients. Accessory organs include salivary glands, the liver, the pancreas, and the gallbladder. The liver, pancreas, and gallbladder are regulated by hormones in response to the food consumed.

The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes many plasma proteins.

The pancreas is another important gland that secretes digestive juices. The chyme produced from the stomach is highly acidic in nature; the pancreatic juices contain high levels of bicarbonate, an alkali that neutralizes the acidic chyme. Additionally, the pancreatic juices contain a large variety of enzymes that are required for the digestion of protein and carbohydrates.

The gallbladder is a small organ that aids the liver by storing bile and concentrating bile salts. When chyme containing fatty acids enters the duodenum, the bile is secreted from the gallbladder into the duodenum.

Ingestion

The large molecules found in intact food cannot pass through the cell membranes. Food needs to be broken into smaller particles so that animals can harness the nutrients and organic molecules. The first step in this process is ingestion. Ingestion is the process of taking in food through the mouth. In vertebrates, the teeth, saliva, and tongue play important roles in mastication (preparing the food into bolus). While the food is being mechanically broken down, the enzymes in saliva begin to chemically process the food as well. The combined action of these processes modifies the food from large particles to a soft mass that can be swallowed and can travel the length of the esophagus.

Digestion and Absorption

Digestion is the mechanical and chemical breakdown of food into small organic fragments. It is important to breakdown macromolecules into smaller fragments that are of suitable size for absorption across the digestive epithelium. Large, complex molecules of proteins, polysaccharides, and lipids must be reduced to simpler particles such as simple sugar before they can be absorbed by the digestive epithelial cells. Different organs play specific roles in the digestive process. The animal diet needs carbohydrates, protein, and fat, as well as vitamins and inorganic components for nutritional balance. How each of these components is digested is discussed in the following sections.

Carbohydrates

The digestion of carbohydrates begins in the mouth. The salivary enzyme amylase begins the breakdown of food starches into maltose, a disaccharide. As the bolus of food travels through the esophagus to the stomach, no significant digestion of carbohydrates takes place. The esophagus produces no digestive enzymes but does produce mucous for lubrication. The acidic environment in the stomach stops the action of the amylase enzyme.

The next step of carbohydrate digestion takes place in the duodenum. Recall that the chyme from the stomach enters the duodenum and mixes with the digestive secretion from the pancreas, liver, and gallbladder. Pancreatic juices also contain amylase, which continues the breakdown of starch and glycogen into maltose, a disaccharide. The disaccharides are broken down into monosaccharides by enzymes called maltases, sucrases, and lactases, which are also present in the brush border of the small intestinal wall. Maltase breaks down maltose into glucose. Other disaccharides, such as sucrose and lactose are broken down by sucrase and lactase, respectively. Sucrase breaks down sucrose (or “table sugar”) into glucose and fructose, and lactase breaks down lactose (or “milk sugar”) into glucose and galactose. The monosaccharides (glucose) thus produced are absorbed and then can be used in metabolic pathways to harness energy. The monosaccharides are transported across the intestinal epithelium into the bloodstream to be transported to the different cells in the body. The steps in carbohydrate digestion are summarized in Figure 24.10.

Protein

A large part of protein digestion takes place in the stomach. The enzyme pepsin plays an important role in the digestion of proteins by breaking down the intact protein to peptides, which are short chains of four to nine amino acids. In the duodenum, other enzymes—trypsin, elastase, and chymotrypsin—act on the peptides reducing them to smaller peptides. Trypsin elastase, carboxypeptidase, and chymotrypsin are produced by the pancreas and released into the duodenum where they act on the chyme. Further breakdown of peptides to single amino acids is aided by enzymes called peptidases (those that breakdown peptides). Specifically, carboxypeptidase, dipeptidase, and aminopeptidase play important roles in reducing the peptides to free amino acids. The amino acids are absorbed into the bloodstream through the small intestines. The steps in protein digestion are summarized in Figure 24.11.

Lipids

Lipid digestion begins in the stomach with the aid of lingual lipase and gastric lipase. However, the bulk of lipid digestion occurs in the small intestine due to pancreatic lipase. When chyme enters the duodenum, the hormonal responses trigger the release of bile, which is produced in the liver and stored in the gallbladder. Bile aids in the digestion of lipids, primarily triglycerides by emulsification. Emulsification is a process in which large lipid globules are broken down into several small lipid globules. These small globules are more widely distributed in the chyme rather than forming large aggregates. Lipids are hydrophobic substances: in the presence of water, they will aggregate to form globules to minimize exposure to water. Bile contains bile salts, which are amphipathic, meaning they contain hydrophobic and hydrophilic parts. Thus, the bile salts hydrophilic side can interface with water on one side and the hydrophobic side interfaces with lipids on the other. By doing so, bile salts emulsify large lipid globules into small lipid globules.

Why is emulsification important for digestion of lipids? Pancreatic juices contain enzymes called lipases (enzymes that breakdown lipids). If the lipid in the chyme aggregates into large globules, very little surface area of the lipids is available for the lipases to act on, leaving lipid digestion incomplete. By forming an emulsion, bile salts increase the available surface area of the lipids many fold. The pancreatic lipases can then act on the lipids more efficiently and digest them. Lipases breakdown the lipids into fatty acids and glycerides. These molecules can pass through the plasma membrane of the cell and enter the epithelial cells of the intestinal lining. The bile salts surround long-chain fatty acids and monoglycerides forming tiny spheres called micelles. The micelles move into the brush border of the small intestine absorptive cells where the long-chain fatty acids and monoglycerides diffuse out of the micelles into the absorptive cells leaving the micelles behind in the chyme. The long-chain fatty acids and monoglycerides recombine in the absorptive cells to form triglycerides, which aggregate into globules and become coated with proteins. These large spheres are called chylomicrons. Chylomicrons contain triglycerides, cholesterol, and other lipids and have proteins on their surface. The surface is also composed of the hydrophilic phosphate “heads” of phospholipids. Together, they enable the chylomicron to move in an aqueous environment without exposing the lipids to water. Chylomicrons leave the absorptive cells via exocytosis. Chylomicrons enter the lymphatic vessels, and then enter the blood in the subclavian vein.

Vitamins

Vitamins can be either water-soluble or lipid-soluble. Fat soluble vitamins are absorbed in the same manner as lipids. It is important to consume some amount of dietary lipid to aid the absorption of lipid-soluble vitamins. Water-soluble vitamins can be directly absorbed into the bloodstream from the intestine.

Link to Learning

This website has an overview of the digestion of protein, fat, and carbohydrates.

Elimination

The final step in digestion is the elimination of undigested food content and waste products. The undigested food material enters the colon, where most of the water is reabsorbed. Recall that the colon is also home to the microflora called “intestinal flora” that aid in the digestion process. The semi-solid waste is moved through the colon by peristaltic movements of the muscle and is stored in the rectum. As the rectum expands in response to storage of fecal matter, it triggers the neural signals required to set up the urge to eliminate. The solid waste is eliminated through the anus using peristaltic movements of the rectum.

Common Problems with Elimination

Diarrhea and constipation are some of the most common health concerns that affect digestion. Constipation is a condition where the feces are hardened because of excess water removal in the colon. In contrast, if enough water is not removed from the feces, it results in diarrhea. Many bacteria, including the ones that cause cholera, affect the proteins involved in water reabsorption in the colon and result in excessive diarrhea.

Emesis

Emesis, or vomiting, is elimination of food by forceful expulsion through the mouth. It is often in response to an irritant that affects the digestive tract, including but not limited to viruses, bacteria, emotions, sights, and food poisoning. This forceful expulsion of the food is due to the strong contractions produced by the stomach muscles. The process of emesis is regulated by the medulla.