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DIGESTIVE PHYSIOLOGY Anatomy: 1) MOUTH = Buccal Cavity = lips, teeth, tongue Lips obtain food, extremely tactile and

d sensitive Teeth incisors cut forage; molars grind feed Circular motion sharp points floating Tongue distributes feed within mouth, forms bolus Digestive Processes: 1. MASTICATION = chewing = reduction of food particle size; increase surface area; expose inner portions of grain and/or forage 2. SALIVATION = addition of saliva from salivary glands to food; ONLY occurs during mastication (no base salivary flow); contains ENZYMES (probably amylase) and high concentration of buffer 2) ESOPHAGUS one-way tube composed primarily of smooth muscle; approximately 4-5 feet long. Digestive Process: transport food bolus from mouth to stomach ***Under normal conditions, the horse CANNOT vomit, belch, etc.** 3. STOMACH muscular mixing vat; composes about 8% of total GIT; contains 2-3 gallons Structures/regions CARDIAC SPHINCTER - one-way valve the controls the entry of bolus from esophagus; prevent ingesta and gas from entering the esophagus. SACCUS CECUS the non-glandular portion of the stomach; located in the anterior 1/3; contains microbes GLANDULAR REGION contains Chief cells and Parietal cells which produce pepsin and HCl; very low pH PYLORIC VALVE semi-one-way valve which controls the exit of ingesta from the stomach **Under grazing conditions, the stomach is NEVER empty, and the production of gastric juices is CONTINUOUS.

After 48-60 hours of fasting, production of gastric juices will decline. Digestive Processes: 1. Slight microbial digestion begins to breakdown fibrous cell walls 2. Proteolysis (protein peptides) begins 3. HCl increases acidity; weakens cell walls 4) SMALL INTESTINE primary site of enzymatic digestion and absorption of soluble nutrients; 30% of GIT; ~ 70 feet long; contains ~ 24 gallons 3 Segments 1. Duodenum 1st segment; fixed part; addition of pancreatic enzymes and bile 2. Jejunum middle segment; mesenteric part (not fixed; attached via thin, pliable mesentery); addition of succous entericus (gastric juice); high number of well developed microvilli increased surface area; location of fibrous bands (mixing of ingesta and enzymes); probable main site of starch and fat digestions and absorption 3. Ileum final segment; mesenteric part, restricted at cecal end; high number of well developed microvilli; microbe population that is proteolytic in activity; addition of buffer to reduce acidity; probable final site of major protein digestion and absorption SECRETIONS: 1. Pancreatic enzymes produced in the pancreas and flow through a duct system to the duodenum; flow is high volume with low concentrations of amylase, protease, and lipase enzymes; flow is CONTINUOUS (decrease after 48 hours of starvation) 2. Bile produced by the liver and flows directly to the duodenum (Horse has no gall bladder); enables lipase enzymes to penetrate and degrade large fat molecules; constant flow which is NOT influenced by feed (or fat) intake 3. Succous Entericus produced by glands within the jejunum and ileum; contains amylase, protease, and lipase enzymes; glands also secrete large concentrations of buffer and mucous; flow tends to be continuous, but does respond to volume, particle size, and moisture content of ingesta 4. Microbes the small intestine does house a limited number of specialized microbes, i.e. cleavage of disaccharides to monosaccharide or cleavage of a specific amino acid

Digestive Processes: 1. Soluble CHO are ~ 90% digested and absorbed in limited quantities 2. Dietary fat is ~100% digested and absorbed in limited quantities 3. Available protein (not bound to fiber) is ~50-70% digested and absorbed 4. Almost ALL vitamins and many minerals (calcium, zinc, copper, magnesium, manganese) are absorbed **LIMITATIONS are primarily related to volume of feed and rate of passage. 5) CECUM large, muscular mixing vat; 12-15% of GIT; 4 ft. long; 7-9 gallons ILEOCECAL VALVE separates the foregut and hindgut; 3-way valve allows ingesta to enter cecum from ileum; allow ingesta to leave cecum and enter large intestine; CAN contract and allow ingesta to by-pass the cecum Environment Ph neutral; vast, varied population of microbes; very liquid Digestive Processes: 1. Initiation of fermentation Insol CHO VFA, CH4, H2O; absorption of VFA Begin protein NH3+, C-chains, R-groups; absorption of NH3+ 2. Begin synthesis of B Vitamins, possible synthesis of amino acids 6) LARGE INTESTINE digestion, absorption and excretion of nutrients; 40-50% of GIT; 40-50 gallons 3 PARTS: 1. Large ventral colon makes up about 50% of large intestine; continuation and completion of microbial digestion; absorption of VFA, B vitamins, and ?AA? 2. Large dorsal colon conservation of electrolytes (Na, Cl, K); absorption of P 3. Small Colon final portion of digestive system; primary function is to CONSERVE WATER, ~80% of water is removed from ingesta at this point **PELVIC FLEXURE narrowing of the intestine between the ventral and dorsal colon (8 diameter3-4 diameter); slows passage of ingesta; allows heavy particles (sand) to settle out; can create a jam of undigestible fiber = impaction colic

DIGESTIVE PROCESSES In general the digestive system of the horse can be separated into two parts the FOREGUT and the HINDGUT. The FOREGUT is composed of the mouth, esophagus, stomach, and small intestine and digestion involves both physical and chemical processes. Physical processing = MASTICATION Mastication physically reduces the feed particle size by crushing/ shredding/tearing the kernel/stem/leaf. This exposes the inner, high nutrient, portions of the feed to the chemical digestion to come. It also increases the amount of surface area that will be directly exposed to this chemical activity. Any impairment in mastication will result in an overall decrease in digestive efficiency. Chemical processing involves exposure of the masticated feed to chemicals enzymes and acids. ENZYMES break susceptible feed particles into nutrients that can be absorbed and utilized by the body. Enzymes are produced by the: Salivary Glands amylase Chief Cells (stomach) protease Pancreas amylase, protease, lipase Intestinal Glands amylase, protease, lipase. ACIDS weaken/destroy cell walls and the bonds creating large, complex molecules. Acids act primarily on structural proteins, exposing other substrates to enzymatic digestion. Acids are produced by the Parietal Cells (stomach) primarily HCl. BILE is neither an enzyme or and acid, but it is necessary for proper digestion of dietary fat. Bile is an emulsifier. It breaks up large fat globules into smaller units that can be attacked by lipase. Bile is produced by the LIVER and flows directly into the duodenum of the small intestine. The horse has no GALL BLADDER, and can not store bile. The production and flow of bile is continuous and, under normal circumstances, is NOT influenced by dietary intake or dietary fat level. **Contrary to popular belief, MICROBIAL DIGESTION does occur in the foregut! Microbes (bacteria, protozoa, microflora) are found in the mouth, stomach, and small intestine.

MOUTH primarily bacteria; attack structural protein; expose inner feed particles; very slight impact on digestion. STOMACH a variety of microbes are found in the Saccus Cecus (non-glandular portion); attack structural proteins; also produce VFA and lactic acid VFA provide energy for stomach cells Lactic acid see acid effects above SMALL INTESTINE specific microbes are found in the jejunum and ileum; those in the jejunum attack specific disaccharides monosaccharide; those in the ileum attack specific peptides amino acids.

The HINDGUT is composed of the cecum, large intestine, rectum and anus and digestion involves MICROBIAL ACTION, primarily fermentation. Proteolysis and microbial synthesis also occur. FERMENTATION is the microbial breakdown of insoluble CHO (digestible fiber). Fiber VFA + CH4 + CO2 + H2O Begins in the cecum and is completed in the ventral colon; Very time consuming, but highly efficient. VFA are used in and/or absorbed from both the cecum and ventral colon. CH4 and CO2 remain in suspension. PROTEOLYSIS is the microbial breakdown of structural, fiber-bound protein. This type of protein largely escaped digestion in the foregut. Protein NH3 + Carbon chains + CO2 + H2O + R-groups MICROBIAL SYNTHESIS is the microbial production of compounds, including most of the B-vitamins and some amino acids. Occurs primarily in the ventral colon. Carbon chains + R-groups vitamins NH3 + carbon chains + R-groups amino acids

NUTRIENT DIGESTION AND ABSORPTION WATER: Very little water is removed from the GIT until it reaches the small colon. In the small colon, 80+% of the water ingested, or created, is absorbed into the body. CARBOHYDRATES: Soluble CHO are those CHO which are susceptible to digestion by enzymes produced by the digestive glands of the horse. STARCH is the primary form of sol-CHO found in common feedstuffs. Starch is a long, simple chain of sugar molecules held together with simple bonds. These bonds are easily broken by amylase enzymes. Starch sugars (primarily glucose) DIGESTION: 1) Begins in the mouth with the addition of saliva; Initial breakdown is very small and may function primarily to provide a source of energy for the microbes and cells of the stomach. 2) In the small intestine, amylase enzymes are added from the pancreas (duodenum) and from the intestinal glands (jejunum); starch is broken down into compound and complex sugars: >the compound sugars are further broken down into simple sugars (glucose) and absorbed from the jejunum and ileum; >the complex sugars usually contain fructose or lactose. The bonds connecting these sugars are NOT susceptible to amylase. Breaking these bonds requires enzymes produced by specific microbes found in the crypts of the jejunum. ABSORPTION: The equine digestive system is designed for 100% absorption of sugars in the small intestine. Insoluble CHO = Digestible fiber are those structural CHO which are NOT susceptible to digestion from the animals enzymes. BACTERIA present in the cecum and large colon produce cellulose, and enzyme specific for the breakdown of cellulose. CELLULOSE and hemicellulose are the main digestible insol-CHO found in fiber. (LIGNIN is the main indigestible insol-CHO). Cellulose is a very long, complex chain of sugars. The chain is branched and may even contain cross

branches between chains. The bonds connecting the sugars are also complex, and require the stronger cellulose enzymes in order to be separated. DIGESTION: FERMENTATION = the bacterial digestion of fiber: Fiber VFA + CH4 + CO2 + H2O. Fermentation begins in the cecum and continues in the large ventral colon. The process is very slow (~24-36 hours) and consistent, if fiber is available. Gaseous products (CH4 and CO2) typically remain in suspension as small bubbles, and are expelled with the feces. ABSORPTION: VFA = volatile fatty acids; very short chain fatty acids which are easily transported across membrane walls; the most common are acetic (2-C), proprionic (3-C), and butyric (4-C); the relative percentages of each depends on the make-up of the fiber; absorbed from the ventral colon. STARCH DIGESTION IN THE CECUM AND VENTRAL COLON: The cecum and ventral colon contain a very small population of starch digesting bacteria. These bacteria are specifically designed to convert starch to ATP via anaerobic phosphorylation. The waste product produced is lactic acid. **If the fermentative bacteria get a hold of starch, the process is greatly ACCELERATED, producing large amount of GAS and LACTIC ACID. This creates accumulation of gas in pockets and lowers pH which results in the death of bacteria.

PROTEIN: Protein digestion in the horse is a fairly long, drawn out process when compared to the ruminant. Digestion is very dependent on the QUALITY and AVAILABILITY of the protein in the feed. Non-structural protein = available protein = non-fiber protein: Available protein is that protein found inside of the grain kernel; the soft, inner portion of the stem; or the inner portion of the leaf that is exposed during MASTICATION. This type of protein is very susceptible to protease enzyme activity. Structural protein = fiber protein = cell wall protein: Fiber protein is that protein contained in the fibrous cells forming the outer covering of the

kernel, stem, and leaf. This protein is often bound to cellulose. Therefore, it is only susceptible to BACTERIAL BREAKDOWN. FOREGUT DIGESTION Stomach: Saccus Cecus microbes present in this region attach the cellulose portion of the feed exposing more of the proteins Glandular Region ingesta is now exposed to HCl which weakens the bonds between cells and weakens the cell membranes; this exposes more protein and makes the bonds between the amino acids more susceptible to enzymatic activity; Addition of pepsin and pepsinogen to the long chain, complex proteins begins the enzymatic digestive process. These enzymes break the chains between specific amino acids resulting in shorter chains termed PEPTIDES. Small Intestine: Duodenum addition of protease enzymes from the pancreas. Jejunum addition of protease enzymes from succous entericus; thorough mixing of enzymes with ingesta via fibrous band action; peptides broken down into smaller chains, and some to individual AA. Ileum continued exposure to protease enzymes releases more individual AA which are absorbed. Presence of specific microbes attach specific bonds holding specific AA together (structural protein), which allows for absorption of these specific AA. HINDGUT DIGESTION Cecum: Structural proteins enter the cecum and are exposed to bacteria degradation. Complete degradation of protein occurs: AA NH3 + C-chains + R-group + CO2 + CH4 +H2O. Ventral Colon: Degradation of protein is complete. Absorption of NH3, C-chains, etc. occurs. HOWEVER, the ventral colon contains synthesizing bacteria. These bacteria are capable of creating AA, including EAA. Absorption of these created AA is fair in the young, growing horse, and probably quite poor in the mature horse. MUCH of the AA created pass out via the feces.

FAT: Fat digestion in the horse is very efficient under normal feeding conditions. Common feedstuffs are low (<3.5%) in crude fat and 100% of fat digestion and absorption occurs in the small intestine. DIGESTION: Small Intestine: Addition of bile from the liver and lipase from the pancreas occurs in the duodenum. Bile breaks down (emulsifies) the large fat globules into small portions. This allows the lipase enzymes to attack the very long chain fat molecules. More lipase is added from the intestinal glands in the jejunum. Fat (bile) (lipase) Free Fatty Acids ABSORPTION: Free Fatty Acids (FFA) are absorbed from the jejunum and ileum.

VITAMINS: The majority of dietary vitamins are digested and absorbed in the foregut. Fat soluble vitamins (A, D, E, K) are acquired via fat digestion/absorption. Vitamin C is absorbed directly from the small intestine. The B vitamins are synthesized by microbes in the cecum and large ventral colon, and absorbed in the large dorsal colon. MINERALS: Mineral digestion and absorption occurs throughout the GIT. Digestion includes the separation of minerals from the feedstuffs, and then individual minerals are absorbed. FOREGUT: Most macro minerals and the majority of the microminerals are absorbed in the foregut Ca, Cu, Zn, Mn, Mg, Se, etc. HINDGUT: The hindgut absorbs specific minerals electrolytes: Na, Cl, K; and P, and CONSERVES others as the need arises.

Anatomical Peculiarities that Predispose a Horse to Colic 1. Esophagus and Cardiac Sphincter The combination of the esophagus and cardiac sphincter create a ONE-WAY passage to the stomach. The horse cannot belch or vomit under normal conditions. Therefore, the equine stomach can fairly easily become distended from a buildup of gas, through overeating, or from a backup of ingesta. RUPTURE of the equine stomach is achievable. 2. Fibrous Bands in the small intestine Fibrous bands, located primarily in the jejunum, separate the small intestine into SEGMENTS. Passage of ingesta through these segments is slowed to allow for the mixing of enzymes and the absorption of nutrients. Overfeeding can cause distension of the segments resulting in pain. Torsion of the intestine and/or complete blockage may result in the rupture and/or death of a segment. 3. Ileo-cecal valve and the cecum Movement of ingesta into and out of the cecum is controlled by the ileocecal valve. Movement into the cecum is determined by pressure in the SMALL INTESTINE, and movement out is controlled by pressure within the CECUM. Thus, it is possible for BOTH valves to open at the same time. This can allow ingesta and/or gas to move back into the small intestine creating distension, pain, torsion, etc. In addition, the cecum is very loosely attached. The strong muscular contractions of the cecum can result in a lot of movement. If the ileo-cecal valves are closed, due to gas buildup, torsion and/or displacement of the cecum can occur. 4. Pelvic Flexure The pelvic flexure is a physical narrowing of the large intestine between the ventral and dorsal large colon designed to slow the passage of ingesta to allow for time to complete microbial digestion and synthesis, and absorption of nutrients. This restriction allows HEAVY PARTICLES to settle out of the ingesta and collect in the area. These heavy particles (sand, dirt, etc.) may place enough weight on the lining to restrict blood flow resulting in the death of the tissue. IN addition, the narrowing of the intestinal lumen may allow the buildup of indigestible fiber. If unabated, this buildup can create an impaction ceasing the movement of ingesta through this part of the intestine.

RATE OF PASSAGE The horse is designed, anatomically, to consume small frequent meals composed of relatively low quality roughage-based feed. CP ~ 8-15%, Fat ~ 3%, Sol-CHO < 10%, Fiber > 20% Efficient digestion of this diet takes TIME. IN the ruminant, the greatest volume and the slowest passage of ingesta occurs in the FOREGUT, where microbial and chemical digestion takes place. Volume of the hindgut is relatively small and passage is relatively fast. Little digestion occurs in the hindgut, and the most important absorption is the conservation of water. The horse is just the opposite. The foregut has the LEAST volume (~35-40%) and the most RAPID passage of ingesta. Chemical digestion is restricted to the foregut. The hindgut has the GREATEST volume (>60%) and the SLOWEST movement of ingesta. Significant microbial digestion and absorption of nutrients occurs in the hindgut. Therefore, when we consider the EFFICIENCY of digestion in the horse, feed intake (volume) and rate of passage (time) are RESTRICTIVE FACTORS. STOMACH: Volume: In the mature 1100# horse, the stomach accounts for ~ 8% of the GIT, or ~ 3-4 gallons. Under normal conditions, the stomach will contain ~ 1 gallon of liquid at all times. Roughage will require an equal, or greater, amount of saliva added to move it to the stomach. Little of the liquid is absorbed into the roughage, but it is needed to smooth the passage of the ingesta. o 6# hay + 6+# saliva (1 gallon) + 1 gallon already in the stomach = 3+ gallons of volume. Grain-based supplements require up to TWICE as much saliva as roughage (on a weight basis). Processed grain (mastication or mechanical) will ABSORB significant amounts of liquid from saliva. In addition, there is still a required amount of saliva needed to smooth the passage of the ingesta from the mouth to the stomach. o 3# grain + 3# absorbed + 6# saliva + 1 gallon in stomach = 3+ gallons of volume in stomach.

Therefore, feeding as little as 3# of grain, or 6# of hay may fill the stomach to capacity. Adding additional feed, until this has had a chance to move out of the stomach, may result in overloading the stomach and reducing digestive efficiency. Time: Food remains in the stomach for only 10-15 minutes. During this time, the ingesta MUST be exposed to microbial action in the saccus cecus and to HCl and pepsin in the glandular region. Overloading the stomach will result in limited digestive activity, resulting in a decrease in protein digestion. SMALL INTESTINE: Volume: The small intestine makes up about 30% of the GIT, is ~ 70 feet long, and holds ~ 24 gallons. In addition to the 4 gallons of ingesta from the stomach, the addition of bile, pancreatic enzymes and intestinal enzymes will add another 2+ gallons of liquid. As the stomach empties, every 15 minutes, an additional 4 gallons (plus the 2 gallons of liquid) is moved into the small intestine. Assuming a continuous intake of feed, the small intestine will fill to capacity in ~ 60 minutes. Time: Under normal conditions, ingesta moves through the small intestine in 60-90 minutes. Duodenum movement through the duodenum is very quick. Addition of bile and pancreatic enzymes to the ingesta is the only process occurring. Jejunum contains the digestive segments separated by the fibrous bands. The digestive segments restrict the movement of ingesta. o Allow ~ - gallon in at one time. o Peristaltic muscle contractions mix the feed stuffs with bile, pancreatic enzymes, and the intestinal juice added in each segment. o Each segment attacks specific nutrients with certain enzymes. o Absorption from each segment is also specific.

o There are 10-12 segments, and the ingesta, under normal conditions spends 2-5 minutes in each segment. Ileum contains a tremendous surface area created by extensive CRYPTS lining the intestine. o Movement through the ileum is fairly constant. o Absorption of many (most) nutrients occurs. o Time (rate of passage) is influenced by the intake of feed pushing the current contents out of the small intestine. PRACTICALITY Given a 1000# horse fed two meals per day at 1% BW grain and 1.5% BW hay; 5# grain/feeding and 7.5# hay/feeding. Most horses will consume the grain first in ~ 15-20 minutes. Hay consumption will take ~ 30-45 minutes. Prior to feeding, the stomach contains ~ 1 gallon of fluid. As the horse begins eating ~ # bolus of grain is swallowed every 1-2 minutes. Within 6-12 minutes the stomach is full. Additional feed forces the contents of the stomach to move into the small intestine in less than the 10-15 minute time needed for proper digestion. The last 1-2# of grain probably does NOT have an opportunity for complete digestion in the stomach. At 15 minutes, the horse begins to consume hay. Hay is passed as # bolus every 12 minutes. As the hay moves into the stomach, it forces the grain out. Processing roughage takes more time in the stomach than grain. Therefore, the 1st 1-2# of hay will probably NOT spend an adequate amount of time in the stomach. Ten to fifteen minutes after the horse has begun his meal, the grain ingesta begins to enter the small intestine. Ten minutes later. ALL of the grain is in the small intestine. Within 5 minutes, the 1st bolus enters the 1st digestive segment. Within 30-45 minutes ALL segments contain grain ingesta. Thirty minutes after the beginning of the meal, roughage begins to enter the small intestine. Within 5 minutes, the roughage begins to push the grain ingesta through the digestive segments.

Roughage moves SLOWER, but has MORE BULK. 1# roughage will displace > 2# grain. THEREFORE, the final 1-2# of grain may NOT have adequate time for digestion and absorption to occur.