Nutrient Acquisition by Animals

Learning Objectives

  1. Compare and contrast complete and incomplete digestive tracts
  2. Identify and explain variation of digestive tract function in animal lineages, including teeth, gizzard, crop, cecum, rumen, and appendix
  3. Describe the steps of mechanical and chemical digestion, and nutrient absorption using the human digestive system as a model

Animal Digestive Systems

The information below was adapted from OpenStax Biology 34.1

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 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).

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. This type of digestive system is also called an incomplete digestive tract. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly fish, and sea anemones) use this type of digestion. Gastrovascular cavities 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 is a more advanced system: it consists of one tube with a mouth at one end and an anus at the other.  This type of digestive system is also called a complete digestive tract. 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.

Part A shows a hydra, which has a vase-shaped body with tentacles around the rim. The hydra’s mouth is located between the tentacles, at the top of the vase. Next to the hydra is a jellyfish medusa, which is bell shaped with tentacles hanging down from the edge of the bell. The mouth, in the lower middle part of the body, opens into the gastrovascular cavity. Part B shows a nematode, which has a long, tube-like body that is wide at one end and tapers down to a tail at the other. The mouth is in the center of the wide end. It opens into an esophagus, then a pharynx. The pharynx empties into a long intestine, which ends at the anus a short distance before the tail.

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. 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 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.

The basic components of the human and rabbit digestive system are the same: each begins at the mouth. Food is swallowed through the esophagus and into the kidney-shaped stomach. The liver is located on top of the stomach, and the pancreas is underneath. Food passes from the stomach to the long, winding small intestine. From there it enters the wide large intestine before passing out the anus. At the junction of the small and large intestine is a pouch called the cecum. The small and large intestines are much longer in rabbits than in humans, and the cecum is much longer as well.

(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. Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their body weight low; this translates to eating and passing food often. In additional contrast to humans, rather than mechanical digestion by teeth, the bird gizzard serves to store and mechanically grind. 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.

Illustration shows an avian digestive system. Food is swallowed through the esophagus into the crop, which is shaped like an upside-down heart. From the bottom of the crop food enters a tubular proventriculus, which empties into a spherical gizzard. From the gizzard, food enters the small intestine, then the large intestine. Waste exits the body through the cloaca. The liver and pancreas are located between the crop and gizzard. Rather than a single cecum, birds have two caeca at the junction of the small and large intestine.

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.

 

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. The horny beak, lack of jaws, and the smaller tongue of the birds can be traced back to their dinosaur ancestors digesting seed. Seed-eating birds have beaks that are shaped for grabbing seeds and the two-compartment stomach allows for delegation of tasks.

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. The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down 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.

Illustration shows the digestive system of a goat. Food passes from the mouth, through the esophagus and into the rumen. It circulates clockwise through the rumen, then moves forward, and down into the small, pouch-shaped reticulum. From the reticulum the food, which is now cud, is regurgitated. The animal chews the cud, and then swallows it into the coiled omasum, which sits between the reticulum and the rumen. After circulating through the omasum the food enters the small intestine, then the large intestine. Waste is excreted through the anus.

Ruminant animals, such as goats and cows, have four stomachs. The first two stomachs, the rumen and the reticulum, contain prokaryotes and protists that are able to digest cellulose fiber. The ruminant regurgitates cud from the reticulum, chews it, and swallows it into a third stomach, the omasum, which removes water. The cud then passes onto the fourth stomach, the abomasum, where it is digested by enzymes produced by the ruminant.

 

The video below compares and contrasts different vertebrate digestive systems (starting at 8:59):

https://www.youtube.com/watch?v=7s23mLohwg4&feature=youtu.be&t=529

Digestive Processes

The information below was adapted from OpenStax Biology 34.3

Obtaining nutrition and energy from food is a multi-step process. For ingestive feeders (animals that swallow food), the first step is ingestion, the act of taking in food. This is followed by digestion, absorption, and elimination. In the following sections, each of these steps will be discussed in detail.

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 break down of food into small organic fragments. It is important to break down 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 below.

Pathways for the breakdown of starch and glycogen, sucrose, and lactose are shown. Starch and glycogen, which are both polysaccharides, are broken down into the disaccharide maltose. Maltose is then broken down into the monosaccharaide glucose. Sucrose, a disaccharide, is broken down by sucrose into the monosaccharides glucose and fructose. Lactose, also a disaccharide, is broken down by lactase into glucose and galactose.

Digestion of carbohydrates is performed by several enzymes. Starch and glycogen are broken down into glucose by amylase and maltase. Sucrose (table sugar) and lactose (milk sugar) are broken down by sucrase and lactase, respectively.

 

Digestion of Carbohydrates
Enzyme Produced By Site of Action Substrate Acting On End Products
Salivary amylase Salivary glands Mouth Polysaccharides (Starch) Disaccharides (maltose), oligosaccharides
Pancreatic amylase Pancreas Small intestine Polysaccharides (starch) Disaccharides (maltose), monosaccharides
Oligosaccharidases Lining of the intestine; brush border membrane Small intestine Disaccharides Monosaccharides (e.g., glucose, fructose, galactose)

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 break down 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 below.

Protein digestion begins in the stomach, where pepsin breaks proteins down into fragments, called peptides. Further digestion occurs in the small intestine, where a variety of enzymes break peptides down into smaller peptides, and then into individual amino acids. Several of the protein-digesting enzymes found in the small intestine are secreted from the pancreas. Amino acids are absorbed from the small intestine into the blood stream. The liver regulates the distribution of amino acids to the rest of the body. A small amount of dietary protein is lost in the feces.

Protein digestion is a multistep process that begins in the stomach and continues through the intestines.

 

Digestion of Protein
Enzyme Produced By Site of Action Substrate Acting On End Products
Pepsin Stomach chief cells Stomach Proteins Peptides
  • Trypsin
  • Elastase Chymotrypsin
 Pancreas Small intestine Proteins Peptides
Carboxypeptidase Pancreas Small intestine Peptides Amino acids and peptides
  • Aminopeptidase
  • Dipeptidase
 Lining of intestine Small intestine Peptides Amino acids

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 break down 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 break down 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.

Illustration shows a row of absorptive epithelial cells that line the intestinal lumen. Hair-like microvilli project into the lumen. On the other side of the epithelial cells are capillaries and lymphatic vessels. In the intestinal lumen, lipids are emulsified by the bile. Lipases break down fats, also known as triglycerides, into fatty acids and monoglycerides. Fats are made up of three fatty acids attached to a 3-carbon glycerol backbone. In monoglycerides, two of the fatty acids are removed. The emulsified lipids form small, spherical particles called micelles that are absorbed by the epithelial cells. Inside the epithelial cells the fatty acids and monoglyerides are reassembled into triglycerides. The triglycerides aggregate with cholesterol, proteins, and phospholipids to form spherical chylomicrons. The chylomicrons are moved into a lymph capillary, which transports them to the rest of the body.

Lipids are digested and absorbed in the small intestine.

 

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.

Steps in mechanical and chemical digestion are shown. Digestion begins in the mouth, where chewing and swallowing mechanically breaks down food into smaller particles, and enzymes chemically digest carbohydrates. In the stomach, mechanical digestion includes peristaltic mixing and propulsion. Chemical digestion of proteins occurs, and lipid-soluble substances such as aspirin are absorbed. In the small intestine, mechanical digestion occurs through mixing and propulsion, primarily by segmentation. Chemical digestion of carbohydrates, lipids, proteins and nucleic acid occurs. Peptides, amino acids, glucose, fructose, lipids, water, vitamins, and minerals are absorbed into the bloodstream. In the large intestine, mechanical digestion occurs through segmental mixing and mass movement. No chemical digestion occurs except for digestion by bacteria. Water, ions, vitamins, minerals, and small organic molecules produced by bacteria are absorbed into the bloodstream.

Mechanical and chemical digestion of food takes place in many steps, beginning in the mouth and ending in the rectum.

 

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.

This video gives an overview of the digestive process in humans: