The Part of the Poultry Digestive System That Stores and Moistens Feed

Chapter 15. Animal Nutrition and the Digestive System

15.1 Digestive Systems

Learning Objectives

By the end of this section, you will be able to:

• Explain the processes of digestion and absorption
• Compare and contrast different types of digestive systems
• Explain the specialized functions of the organs involved in processing food in the body
• Describe the ways in which organs work together to digest food and absorb nutrients

Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are a number of processes that modify food within the animal body in order to make the nutrients and organic molecules accessible for cellular function. As animals evolved in complexity of form and function, their digestive systems have also evolved to accommodate their various dietary needs.

Herbivores, Omnivores, and Carnivores

Herbivores are animals whose primary food source is plant-based. Examples of herbivores, as shown in Figure 15.2 include vertebrates like deer, koalas, and some bird species, as well as invertebrates such as crickets and caterpillars. These animals have evolved digestive systems capable of handling large amounts of plant material. Herbivores can be further classified into frugivores (fruit-eaters), granivores (seed eaters), nectivores (nectar feeders), and folivores (leaf eaters).

Figure 15.2.  Herbivores, like this (a) mule deer and (b) monarch caterpillar, eat primarily plant material. (credit a: modification of work by Bill Ebbesen; credit b: modification of work by Doug Bowman)

Carnivores are animals that eat other animals. The word carnivore is derived from Latin and literally means "meat eater." Wild cats such as lions, shown in Figure 35.3 a and tigers are examples of vertebrate carnivores, as are snakes and sharks, while invertebrate carnivores include sea stars, spiders, and ladybugs, shown in Figure 15.3 b . Obligate carnivores are those that rely entirely on animal flesh to obtain their nutrients; examples of obligate carnivores are members of the cat family, such as lions and cheetahs. Facultative carnivores are those that also eat non-animal food in addition to animal food. Note that there is no clear line that differentiates facultative carnivores from omnivores; dogs would be considered facultative carnivores.

Figure 15.3.  Carnivores like the (a) lion eat primarily meat. The (b) ladybug is also a carnivore that consumes small insects called aphids. (credit a: modification of work by Kevin Pluck; credit b: modification of work by Jon Sullivan)

Omnivores are animals that eat both plant- and animal-derived food. In Latin, omnivore means to eat everything. Humans, bears (shown in Figure 15.4 a ), and chickens are example of vertebrate omnivores; invertebrate omnivores include cockroaches and crayfish (shown in Figure 15.4 b ).

Figure 15.4.  Omnivores like the (a) bear and (b) crayfish eat both plant and animal based food. (credit a: modification of work by Dave Menke; credit b: modification of work by Jon Sullivan)

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 15.5 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 break down the food. The food particles are engulfed by the cells lining the gastrovascular cavity.

The alimentary canal, shown in Figure 15.5 b , 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.

Figure 15.5.  (a) A gastrovascular cavity has a single opening through which food is ingested and waste is excreted, as shown in this hydra and in this jellyfish medusa. (b) An alimentary canal has two openings: a mouth for ingesting food, and an anus for eliminating waste, as shown in this nematode.

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 15.6 ab . 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 break down 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 blood stream 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.

Figure 15.6.
(a) Humans and herbivores, such as the (b) rabbit, have a monogastric digestive system. However, in the rabbit the small intestine and cecum are enlarged to allow more time to digest plant material. The enlarged organ provides more surface area for absorption of nutrients. Rabbits digest their food twice: the first time food passes through the digestive system, it collects in the cecum, and then it passes as soft feces called cecotrophes. The rabbit re-ingests these cecotrophes to further digest them.

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 15.7, 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.

Figure 15.7.  The avian esophagus has a pouch, called a crop, which stores food. Food passes from the crop to the first of two stomachs, called the proventriculus, which contains digestive juices that break down food. From the proventriculus, the food enters the second stomach, called the gizzard, which grinds food. Some birds swallow stones or grit, which are stored in the gizzard, to aid the grinding process. Birds do not have separate openings to excrete urine and feces. Instead, uric acid from the kidneys is secreted into the large intestine and combined with waste from the digestive process. This waste is excreted through an opening called the cloaca.

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 15.9. 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 break down 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.

Figure 15.9.
Digestion of food begins in the (a) oral cavity. Food is masticated by teeth and moistened by saliva secreted from the (b) salivary glands. Enzymes in the saliva begin to digest starches and fats. With the help of the tongue, the resulting bolus is moved into the esophagus by swallowing. (credit: modification of work by the National Cancer Institute)

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 15.10. 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.

Figure 15.10.  The esophagus transfers food from the mouth to the stomach through peristaltic movements.

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 15.11. 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.

Figure 15.11.  The human stomach has an extremely acidic environment where most of the protein gets digested. (credit: modification of work by Mariana Ruiz Villareal)

Which of the following statements about the digestive system is false?

1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.

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 15.12, 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 blood stream 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.

Figure 15.12.  Villi are folds on the small intestine lining that increase the surface area to facilitate the absorption of nutrients.

Which of the following statements about the small intestine is false?

1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.

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 15.11. 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 break down the food particles in the chyme into glucose, triglycerides, and amino acids. Some 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 15.11. Here, hydrolysis of nutrients is continued while most of the carbohydrates and amino acids are absorbed through the intestinal lining. The bulk of chemical digestion and nutrient absorption occurs in the jejunum.

The ileum, also illustrated in Figure 15.11 is the last part of the small intestine and here the bile salts and vitamins are absorbed into blood stream. 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 15.13, 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 15.13. 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.

Summary

Different animals have evolved different types of digestive systems specialized to meet their dietary needs. Humans and many other animals have monogastric digestive systems with a single-chambered stomach. Birds have evolved a digestive system that includes a gizzard where the food is crushed into smaller pieces. This compensates for their inability to masticate. Ruminants that consume large amounts of plant material have a multi-chambered stomach that digests roughage. Pseudo-ruminants have similar digestive processes as ruminants but do not have the four-compartment stomach. Processing food involves ingestion (eating), digestion (mechanical and enzymatic breakdown of large molecules), absorption (cellular uptake of nutrients), and elimination (removal of undigested waste as feces).

Many organs work together to digest food and absorb nutrients. The mouth is the point of ingestion and the location where both mechanical and chemical breakdown of food begins. Saliva contains an enzyme called amylase that breaks down carbohydrates. The food bolus travels through the esophagus by peristaltic movements to the stomach. The stomach has an extremely acidic environment. An enzyme called pepsin digests protein in the stomach. Further digestion and absorption take place in the small intestine. The large intestine reabsorbs water from the undigested food and stores waste until elimination.

Exercises

1. Which of the following statements about the digestive system is false?
   1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
   2. Food enters the large intestine before the small intestine.
   3. In the small intestine, chyme mixes with bile, which emulsifies fats.
   4. The stomach is separated from the small intestine by the pyloric sphincter.
2. Which of the following statements about the small intestine is false?
   1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
   2. The inside of the small intestine has many folds, called villi.
   3. Microvilli are lined with blood vessels as well as lymphatic vessels.
   4. The inside of the small intestine is called the lumen.
3. Which of the following is a pseudo-ruminant?
   1. cow
   2. pig
   3. crow
   4. horse
4. Which of the following statements is untrue?
   1. Roughage takes a long time to digest.
   2. Birds eat large quantities at one time so that they can fly long distances.
   3. Cows do not have upper teeth.
   4. In pseudo-ruminants, roughage is digested in the cecum.
5. The acidic nature of chyme is neutralized by ________.
   1. potassium hydroxide
   2. sodium hydroxide
   3. bicarbonates
   4. vinegar
6. The digestive juices from the liver are delivered to the ________.
   1. stomach
   2. liver
   3. duodenum
   4. colon
7. How does the polygastric digestive system aid in digesting roughage?
8. How do birds digest their food in the absence of teeth?
9. What is the role of the accessory organs in digestion?
10. Explain how the villi and microvilli aid in absorption.

Answers

1. B
2. C
3. D
4. B
5. C
6. C
7. Animals with a polygastric digestive system have a multi-chambered stomach. The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down the cellulose and ferment the ingested food. The abomasum is the "true" stomach and is the equivalent of a monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary for ruminants to digest plant material.
8. Birds have a stomach chamber called a gizzard. Here, the food is stored, soaked, and ground into finer particles, often using pebbles. Once this process is complete, the digestive juices take over in the proventriculus and continue the digestive process.
9. Accessory organs play an important role in producing and delivering digestive juices to the intestine during digestion and absorption. Specifically, the salivary glands, liver, pancreas, and gallbladder play important roles. Malfunction of any of these organs can lead to disease states.
10. The villi and microvilli are folds on the surface of the small intestine. These folds increase the surface area of the intestine and provide more area for the absorption of nutrients.

Glossary

alimentary canal: tubular digestive system with a mouth and anus
anus: exit point for waste material
bile: digestive juice produced by the liver; important for digestion of lipids
bolus: mass of food resulting from chewing action and wetting by saliva
carnivore: animal that consumes animal flesh
chyme: mixture of partially digested food and stomach juices
digestion: mechanical and chemical break down of food into small organic fragments
duodenum: first part of the small intestine where a large part of digestion of carbohydrates and fats occurs
endocrine system: system that controls the response of the various glands in the body and the release of hormones at the appropriate times
esophagus: tubular organ that connects the mouth to the stomach
essential nutrient: nutrient that cannot be synthesized by the body; it must be obtained from food
gallbladder: organ that stores and concentrates bile
gastric inhibitory peptide: hormone secreted by the small intestine in the presence of fatty acids and sugars; it also inhibits acid production and peristalsis in order to slow down the rate at which food enters the small intestine
gastrin: hormone which stimulates hydrochloric acid secretion in the stomach
gastrovascular cavity: digestive system consisting of a single opening
gizzard: muscular organ that grinds food
herbivore: animal that consumes strictly plant diet
ileum: last part of the small intestine; connects the small intestine to the large intestine; important for absorption of B-12
ingestion: act of taking in food
jejunum: second part of the small intestine
lactase: enzyme that breaks down lactose into glucose and galactose
large intestine: digestive system organ that reabsorbs water from undigested material and processes waste matter
lipase: enzyme that chemically breaks down lipids
liver: organ that produces bile for digestion and processes vitamins and lipids
maltase: enzyme that breaks down maltose into glucose
mineral: inorganic, elemental molecule that carries out important roles in the body
monogastric: digestive system that consists of a single-chambered stomach
omnivore: animal that consumes both plants and animals
pancreas: gland that secretes digestive juices
pepsinogen: inactive form of pepsin
pepsin: enzyme found in the stomach whose main role is protein digestion
peristalsis: wave-like movements of muscle tissue
proventriculus: glandular part of a bird's stomach
rectum: area of the body where feces is stored until elimination
roughage: component of food that is low in energy and high in fiber
ruminant: animal with a stomach divided into four compartments
salivary amylase: enzyme found in saliva, which converts carbohydrates to maltose
small intestine: organ where digestion of protein, fats, and carbohydrates is completed
somatostatin: hormone released to stop acid secretion when the stomach is empty
sphincter: band of muscle that controls movement of materials throughout the digestive tract
stomach: saclike organ containing acidic digestive juices
villi: folds on the inner surface of the small intestine whose role is to increase absorption area
vitamin: organic substance necessary in small amounts to sustain life

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