Physiology II Gastroinstestinal System Lecture 2 Dr Than Kyaw 30 April 2012 Ruminant and monogastric Digestion • • • • • Ruminant digestion Monogastric digestion Enzymes and hormones Digestion products Absorption and utilization Compound stomach (multichambered , polygastric, ruminant stomach) • Consume fibrous foods - forages (grasses) - roughages - other fibrous food sources Cellulose and semicellose • Plant materials other simple stoach animals are unable to digest. • Fermented in the rumen with the help of microbes (microflora) To valuable products 4 compartments Rumen (paunch) - Bacterial and chemical breakdown of fiber (anaerobic) - Occupy most of the left-side of the abdomen - Has thick, muscular wall - Papillae on walls: up to 1 cm. in length, bacteria more concentrated - Rumen capacity - 50 to 65 gallons (dairy cattle) - 5 to 10 gallons (sheep) Dorsal coronary groove Dorsal sac Posterior Dorsal blind sac Posterior ventral blind sac Ventral coronary groove Ventral sac Right longitudinal grove Papillae in Rumen Papillae in Rumen Interior surface of rumen - numerous papillae - vary in shape and size - short and pointed to long and foliate. 4 compartments Reticulum (honey combs) - interior surface – looks like honeycomb - helps to remove foreign matter from the food material. Reticulum - full Epithelium of reticulum - thrown into folds that form polygonal cells - honey-comb appearance - Numerous small papillae Reticulum - cleaned 4 compartments Omasum (book stomach) - round and muscular - “Grinds” the food material and prepares the food material for chemical breakdown. Omasum - full Inner surface of omasum - Broad longitudinal folds or leaves (lay term - 'book'). - Omasal folds - packed with finely ground ingesta - Estimated to be 1/3 of the total surface area of the forestomachs. 4 compartments Abomasum – inside view Abomasum (true stomach) • very similar to the stomach of non-ruminants • the majority of chemical breakdown of food material occurs. • mixes in digestive enzymes (pepsin) Size of young and adult ruminants Important to know the development of ruminant stomach for feeding young and adult Compartment size of young and adult ruminants % of Total Stomach Newborn Calf Adult Cow Rumen 25% 80% Reticulum 25% 6% Omasum 10% 3% Abomasum 40% 11% Capacities of digestive tract of mature sheep Compartment Capacity Reticulum Rumen Omasum Abomasum 1.2 to 2.0 quarts 5.0 to 10.0 gallons 0.5 to 1.0 quarts 2.0 to 3.0 gallons Small intestines 2.0 to 2.5 gallons (80 ft) Large intestines 1.5 to 2.0 quarts Rumen microbes (micro flora) - Bacteria: Over 60 spp - digest sugars, starch, fiber, and protein for the cow. - Protozoa: About 35 spp (size: 20 – 200 μ) - some spp swallow and digest bacteria, starch granules, and some fiber. - Bacteria and protozoa – differ greatly in size, shape, and structure Rumen microbes (micro flora) - Fungi - very small fraction - important in splitting plant fibers open to provide easy digestion by bacteria Protozoa of rumen A Protozoan A fungal spore A Protozoan dividing into two Bacteria attaching under side of the protozoan A Protist found in rumen Some protozoan and bacteria have symbotic relationship Protozoan covered with chains of bacteria Bacteria attacking a strand of Fiber taken from a cow’s rumen. Rumen microbes (micro flora) - One trilion microbes/oz of ruminal fluid - multiply in double in 11 minutes - Almost all rumen microbes – anaerobic - 2/3 of feed digestion - 90% of fiber digestion In rumen Rumen microbes (micro flora) - Rumen bacteria – good source of protein for the host - Microbes - 55% of protein and may provide up to half of total dietary need - Urea – can be utilized by microbes (synthesis of microbial protein) - Vitmins K and B-complex synthesis - adapted to a pH between roughly 5.5 and 6.5 - Abomasum pH: 2 to 4 - 3 primary zones based on their specific gravity - Gas rises to fill the upper regions - Grain and fluid-saturated roughage ("yesterday's hay") sink to the bottom - Newly arrived roughage floats in a middle layer. Ingesta flow • Digestion Compound stomach Ruminal motility - mix the ingesta - aid in eructation of gas - propel fluid and fermented foodstuffs into the omasum - Supression of motility - ruminal impaction - Cycles of contraction - 1 to 3 times/min - During feeding - highest frequency - During resting - lowest Ruminal motility Two types of contractions Primary contractions - Originate in the reticulum - Pass caudally around the rumen - Involves a wave of contraction followed by a wave of relaxation, so as parts of the rumen are contracting, other sacs are dilating Secondary contractions - occur in only parts of the rumen - usually associated with eructation. Rumination Rumination (cud chewing) - regurgitation of ingesta from the reticulum - remastication - resalivation and - redeglutition (reswallowing) Provides - effective mechanical breakdown of roughage - increases substrate surface area to fermentative microbes. Rumination Sequences - Contraction of reticulum - Relaxation esophageal sphincter - Inspiratory movement with closed glottis - Negative pressure in the thorax - Dilation of thoracic esophagus and cardia - Reverse peristalsis Rumination time = about 8 h/d 1 circle of rumination = 1 min Result Flow of ingesta Roughage + Fluid to mouth Eructation – Enormous quantity of gas by fermentation – About 30-50 liters/h in adult cattle 5 l/h in a sheep or goat - Eructation or belching - continually get rid of fermentation gases - eructation is associated with almost every secondary ruminal contraction - Eructated gas travels up the esophagus at 160 to 225 cm/s Eructation - Any interferences with eructation – life threatening - Expanding rumen rapidly interferes with breathing - Animals suffering ruminal tympany (bloat) die from asphyxiation. Fermentation in the rumen Carbohydrate fermentation • Fibrous feed (cellulose, hemicellulose, xylans) • Readily fermentable CHO (starch, sugars) • Principle end products - VFAs Volatile fatty acids - CO2 - CH4 - Heat Volatile Fatty Acids Carbohydrates Microbial Fermentation VFA’s × Glucose O CH3 C O – Acetic acid (2c) - Rumen, cecum, colon O O CH3 CH2 CH3 C O – Propionic acid (3c) CH2 CH2 C O– Butyric acid (4c) VFA Formation 2 acetate + CO2 + CH4 + heat 1 Glucose 2 propionate + water 1 butyrate + CO2 + CH4 - VFAs absorbed passively from rumen to portal blood - Provide 70-80% of ruminant’s energy needs Uses of VFA • Acetate – Energy – Fatty acid synthesis • Propionate – Energy – Gluconeogenic – glucose synthesis • Butyrate – Energy – Rumen epithelial cells convert to ketone Proportions produced depends on diet - Fibrous feed – less propionic/a - carbohydrate feed – more propionic/a Absorption of VFAs • • No evidence for active transport VFA metabolism in the rumen wall – Cells use most of the butyrate for their own energy needs – Acetate and propionate are ‘exported’ to blood Fermentation of plant protein Plant proteins Proteolytic organisms Amino/a Ammonia Organic acids Resynthesized to different microbial proteins Rumen microbes – also able to utilze non-protein nitrogens - urea, biurets, amines Fermentation of lipids • Plant contains limited amount of lipids • Lipids found in most plant = galactosyl diglycerides • Microbes do not alter lipids very much • Can synthesize some lipids • Cannot tolerate if dietary fat contains 5 – 7% of total diet Digestion in simple stomach Discussed in Lecture (1) Stomach produces: - HCl (stomach pH about 2, pareitel cell) - pepsinogen (pepsin, Chief cell) - gastric lipase (fat digestion) - mucus (protect the stomach epithelium) - gastrin (hormone – G-cell; stimulate gastric secrretions - signals S/I to prepare for arrival of food) Digestion in simple stomach - Digest mainly protein to polypeptides - Kill pathogens Protein in bolus Pepsinogen (Pro-enzyme/ Precursor of pepsin) HCl pepsin Peptide - Liquids - removed quickly (about 30 m) - Solid portion (chyme) - takes hours Small intestine 3 parts - Duodenum - Jejunum - Ileum. Smooth m/s - circular (segmental contraction) - longitudinal (contract a wave-like action – peristalsis) - further digestion and absorption of nutrients - micro villi on the epithelium Small intestine Digesta pH Functions Duodenum 2.7 - 4 Enzymes pH change Flow rate regulation very little absorption Jejunum 4–7 Enzymes Absorption Ileum 7-8 Absorption Limited fermentation • Rate of pH increase through small intestine is faster in monogastrics. Enhanced Surface Area for Increased Nutrient Absorption Intestinal villi Note: provision of Large surface areas by the micro villi for absorption Gastrointestinal Hormones • Gastrin – Origin: stomach, abomasum – Stimulus: food in stomach – Function: stimulates HCl & pepsinogen secretion, increases stomach motility • Secretin – Origin: duodenum – Stimulus: acid – Function: stimulates pancreatic secretions; slows stomach motility and acid production Gastrointestinal Hormones • Cholecystokinin (CCK) – Origin: duodenum – Stimulus: fat & protein in duodenum – Function: stimulates bile and pancreatic secretions • Also regulates appetite and feed intake • Gastric Inhibitory Protein (GIP) – Origin: duodenum – Stimulus: fats and bile – Function: inhibit stomach motility and secretion of acid and enzymes Pancreas and its secretions - long, thin delicate organ - pinkish gray, glandular Secretions of pancreas - Exocrine - enzymes and carbonates - Endocrine - hormones Pancreas and its secretions Enzymes 1. Amylase: -- carbohydrate digestion (starch, dextrin) mainly to disaccharides (maltose) -- amylase low in ruminants 2. Lipase: fat digestion - triglycerides to monoglycerides and free fatty acids Pancreas and its secretions 3. Proteases*: protein digestion Trypsinogen - converted to trypsin (by enterokinase) Chymotrypsinogen - converted to chymotrypsin (by trypsin) Procarboxypeptidase - converted to carboxypeptidase (by trypsin) 4. Nucleases Digest nucleic acids and nucleotides 5. Lecithinase Lecithin to lysolecithin Inorganic compound NaH2CO3 – neutralize acids Secretion - pH is 7.2-7.8 * Enzymes are ususally released in inactive forms in the source organs. Why? - They are capable of doing autodigestion. Pancreas and its secretions Endocrine - islets of Langerhans insulin ( beta cells) glucagon (alpha cells) somatostatin (delta cells) 1. Insulin Main functions -- Fat Break down inhibited -- promote fat deposition and glycogenesis -- enhance glucose transport across cell membrane and facilitate diffusion Blood Glucose Amino acids uptake of glucose Insulin uptake of amino/a Insulin Muscle, Liver (Stored as glycogen) Used for protein synthesis by all cells - All essential amino/a (balanced ration) are needed for protein synthesis 2. Glucagon Opposite effect of insulin Glycogen Glycogenolysis Fat (Lipolysis) glucagon Blood glucose glucagon Blood glucose 3. Somatostatin -- Act as a moderator to the metabolic effect s of insulin, glucagon and growth hormone. -- Inhibit the secretion of insulin and glucagon -- As a moderator – it also inhibit the secretion of – gastrin, secretin, cholicystokinin, pancreatic exocrine secretion and gastric acid 4. Pancreatic polypepitides -- secretion stimulated by -- ingestion of protein -- exercise -- fasting Control of insulin and glucagon secretion Insulin sensitivity Brain Kidney Intestine Erythrocytes Little response to insulin Liver Muscle Adipose tissue Leucocytes readily response to insulin Normal blood glucose value of animals (mg/dl) Horse 60-110 cow 40-80 sheep 40-80 Blood glucose level lower than other animals Pig 80-120 dog 70-120 chicken 130-270 Large Intestine (Colon) • Absorption o f VFAs • Absorption of ammonia-N – About 30 to 40% of the net transport of N into body fluid – Absorbed N may be used for: • Synthesis of nonessential amino acids • Recycling of N to the rumen – Important on low protein diets – Regulated by: • Increased by increasing N concentration of diet • Decreased by increasing the amount of carbohydrate fermented in the large intestine • Mineral absorption • Water absorption – 90% of water entering the LI is absorbed Note: No enzymatic digestion fermenting micro-organisms are not digested; voided with feces. Nutrient absorption 2 types Passive transport (by concentration gradient) - Diffusion: freely movable through lipid bilayers of cell membrane - especially smaller molecules e.g. O2, CO2 - Facilitated diffusion – molecule diffusion facilitated by the help of transport protein, e.g glucose, amino acids Active transport – use of transport protein, – need energy – e.g. N+-K + pump, glucose and amino acid absorption in GIT Nutrient absorption Transporter protein Avians Digestive System Digestive Tract - Poultry Gall bladder Gizzard Liver Cecum Esophagus Crop Proventriculus Cloaca Pancreas Large intestine Duodenum Small intestine (jejunum, ileum) Avians (Poultry) Mouth • • No teeth, rigid tongue Poorly developed salivary glands – Saliva contains amylase • Beak - adapted for prehension and mastication Avians (Poultry) Esophagus – Has an enlarged area called crop • Ingesta holding and moistening • Location for breakdown of carbohydrate by amylase • Fermentation Proventriculus (stomach) • Release of HCl and pepsin (gastric juices) • Ingesta passes through very quickly (14 seconds) Avians (Poultry) Gizzard (ventriculus) – Muscular area with a hardened lining reduces particle size • Muscular contractions every 20-30 seconds • Includes action of grit • HCl and pepsin secreted in proventriculus Small intestine – Similar to other monogastrics Avians (Poultry) Ceca and large intestine – Contain two ceca instead of one as in other monogastrics – Large intestine is very short (2-4 in) and empties into cloaca where fecal material will be voided via the vent • Water resorption • Fiber fermentation by bacteria • H2O soluble vitamin synthesis by bacteria compound Simple Avian mouth mouth mouth esop esop esop reticulum Rumen omasum abomasum Salivary glands Rumen microbes fermentation stomach crop Rumen microbes proventriculus Gastric secretion gizard Grinding, mixing S/I S/I S/I Digestion, absorption cecum cecum cecum fermentation colon colon colon Absorption of water summary (RUMEN) Telephone Cord Traumatic reticulitis in cow Wire Sponge taken from digestive system of an animal End of Lecture Major Gastrointestinal Hormones Honnone Production Action Release stimulus Gastrin Distal stomach Primary: Stimulates acid secretion from stomach glands Secondary: Stimulates gastric motility, growth of stomach epithelium Protein in stomach; high gastric pH; vagal stimulation Secretin duodenum Primary: Stimulates bicarbonate secretion from pancreas Secondary: Stimulates biliary bicarbonate secretion Acid in duodenum Cholecystokinin (CCK) Duodenum to Primary: Stimulates enzyme secretion ileum, with highest from pancreas concentration in Secondary: Inhibits gastric emptying duodenum Proteins and fats in small intestine Major Gastrointestinal Hormones Gastric inhibitory polypeptide (GIP) Duodenum and upper jejunum Primary: Inhibits gastric motility and secretory activity Secondary: Stimulates insulin secretion provided sufficient glucose is present; may be most important action in many species Carbohydrate and fat in small instine Motilin Duodenum and jejunum Primary: Probably regulates motility pattern of the gut in period between meals Secondary: May regulate tone of lower esophageal sphincter Acetylcholine