Equine Digestion and Absorption of Carbohydrates

By: Patricia Ochonski English 202C

I. Introduction
Over the last 60 million years, horses have evolved to more efficiently consume forages. Forage is a term used to describe plant material, such as grass or hay (Figure 1). The molars (back teeth) of the horse have evolved to grind grasses, and the hoof has become stronger to allow horses to travel large distances while grazing. However, the most important anatomical difference over the last 60 million years has been the evolution of the horses hindgut. The hindgut is the second half of the gastrointestinal tract -- the cecum and the colon. The hindgut digests via fermentation, the process by which bacteria breaks structural carbohydrates down to be absorbed by the body. By understanding this process, horse owners are able to make better decisions regarding the diet of their horse.

Figure 1: The picture on the left shows pasture grass, a fresh forage, and the picture on the right shows timothy hay, a preserved forage. Both are rich in structural and non-structural carbohydrates. 1

Nutrients are the molecular building blocks of all living things. They include water, protein, carbohydrates, fats, minerals, and vitamins. Nutrients can be further divided into those that provide energy, and those that do not. Energy is any phenomenon capable of conversion into work. Carbohydrates, proteins, and fats are the three energy-providing nutrients. Carbohydrates are the most important energy-providing nutrient for the horse, and can be split into two types: structural and non-structural. Structural Carbohydrates Structural carbohydrates include cellulose and hemicellulose, and act as the structural part of a plant the stem and the roots. Structural carbohydrates are often referred to as fiber, and are incapable of being digested and absorbed by many animals, including humans. The hindgut of the horse, however, is able to successfully break down these structural carbohydrates through fermentation.

Figure 2: The basic structure of a plant. The head of the plant, which is made up of the beard and the kernel, and the leaves are all sources of non-structural carbohydrates such as sugars and starches. The stem and the roots of the plant are rich in structural carbohydrates such as cellulose and hemicellulose.

Non-Structural Carbohydrates Non-structural carbohydrates include sugars and starches and can be found in the leaves of plants. Nonstructural carbohydrates are found predominately in gains, such as oats or barley and do not require fermentation in order to be broken down. Instead, they are digested and absorbed in the small intestine.

Leafy grasses are rich in non-structural carbohydrates 2

II. Digestion and Absorption of Structural Carbohydrates

Figure 3: Overview of the equine gastrointestinal tact

Mouth The first step in digestion is the ingestion of food. Horses use their front teeth to rip grass from the ground or to pick up pieces of hay. The skull of the horse has evolved to be able to chew forages. Looking at Figure 4, the horse has a long cheekbone that allows for more muscle attachment. These muscles give the horse the strength to chew forages. The process of chewing food is referred to as mastication. Additionally, the pre-molars and molars at large and flat in shape, and are able to break down forage faster. After the forage is properly chewed and formed into a bolus (a ball of food), it is swallowed and travels down the esophagus until it reaches the stomach. Figure 4: Skull of a Horse

Long Jaw Bone

Pre-Molars and Molars

Esophagus and Stomach The esophagus is essentially a tube that acts as connection between the mouth and the stomach. The stomach is a muscular organ that serves as the site of mechanical digestion. Here, enzymes, biological molecules that break down larger molecules, and acids are secreted. These enzymes and acids break down proteins, fats, and nonstructural carbohydrates into smaller components to allow them to be absorbed further on in the gastrointestinal tract. Structural carbohydrates remain fully in tact at this point. Small Intestine After proteins, fats, and non-structural carbohydrates have been properly digested in the stomach, the bolus moves on to the small intestine. The small intestine is the site of absorption of broken down proteins, fats, and non-structural carbohydrates. It is composed of many folds of tissue as well as finger-like projections of tissue called microvilli, which serve to increase the overall surface area of the tissue. A large surface area is needed to allow efficient absorption of the majority of nutrients that pass through the small intestine. At this point in the digestive system, structural carbohydrates remain in tact and are not absorbed. The Cecum and Colon The remaining structural carbohydrates and other undigested foodstuffs enter the cecum and colon (also known as the large intestine), collectively referred to as the hindgut. The hindgut of the horse is where bacteria that naturally reside there (Figure 5) break down structural carbohydrates in the process of fermentation. The product of this process are volatile fatty acids (VFAs), which are absorbed by the lining of the colon to be used as an energy source in the horses body.

Figure 5: The microbial population in the hindgut of the horse is composed of thousands of different species of bacteria. Together, these bacteria break down structural carbohydrates into VFAs, allowing them to be absorbed into the body through the lining of the colon.

The hindgut of the horse has the feature to separate particles of different sizes as they travel through it if a particle is too big it is retained and is broken down further. This allows the horse to extract as many nutrients from these particles as possible. Figure 6 provides a detailed description of each of the three barriers. Figure 6: The Junctions of the Hindgut
1. Junction between the cecum and the colon: prevents large particles from entering the colon. 2. Pelvic flexure: located at the junction of the dorsal and ventral colon; the colon decreases in circumference, preventing the passage of fiber and other food particles, and the tissue becomes much smaller. This is a common location of impaction colic. 3. Junction of the dorsal and transverse colon: prevents larger particles that still need to be fermented from escaping and aids in water retention.

Horses are able to eat a variety of plants

III. Summary
Carbohydrates, the main energy sources in horses, can be broken down into two groups: structural and nonstructural. Structural carbohydrates include cellulose and hemicellulose, molecules that make up the stem and roots of a plant. They are digested and absorbed via fermentation by microbes in the horses hindgut. Nonstructural carbohydrates, such as sugars and starches, are mainly found in the leaves of plants, digested in the stomach, and absorbed in the small intestine. The hindgut of the horse is equipped with junctions that prevent the passage of food particles before they are properly fermented 1) the junction between the cecum and colon, 2) the pelvic flexure, and 3) the junction of the dorsal and transverse colon.

IV. Glossary
Bolus: ball of food material that travels through the gastrointestinal tract. Energy: any phenomenon capable of conversion into work. Enzymes: biological molecules that break down larger molecules. Fermentation: the process by which bacteria break down structural carbohydrates in the hindgut. Forage: plant material, such as grass or hay. Hindgut: a collective term for the cecum and colon; the site of microbial fermentation. Mastication: the process of chewing food. Microvilli: finger-like projections of tissue in the small intestine that serve to increase the overall surface area of the tissue. Non-structural carbohydrates: include sugars and starches and can be found in the leaves of plants; absorbed in the small intestine. Nutrients: the molecular building blocks of all living things. Small intestine: located after the stomach and before the colon; the site of absorption of broken down proteins, fats, and non-structural carbohydrates. Stomach: located after the esophagus and before the small intestine; a muscular organ that serves as the site of mechanical digestion. Structural carbohydrates include cellulose and hemicellulose, and act as the structural part of a plant the stem and the roots; digested and absorbed in the colon. Volatile Fatty Acids (VFAs): what structural carbohydrates are broken down into through microbial fermentation in the hindgut.