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Digestion and Absorption Energy and nutrient acquisition for life activities. No photosynthetic ability -in higher forms Over evolution development of the digestive system can be observed Simple devices for filteration of particles in water Locomotion to acquire food. Development of a complex system for digestion Development of a system for absorption Development of metabolic processes to manage digestion of food absorption of nutrients. Associated Organs: Sensory organs - sight, smell, taste and feel. Salivary glands - moisten food Pancreas - digestive function, metabolic regulation Liver - digestive and metabolic function, Gall bladder - digestive function Extrinsic and Intrinsic Nervous system - Control of digestion Organs of digestion: Mouth - acquisition of food - particle size reduction Esophagus - transfer of bolus Rumen complex - fermentative digestion - three chambers Stomach/ Abomasum - acid digestion reduction in particle size Small intestine - enzymatic digestion & absorption Cecum - fermentative digestion Large intestine - Water re-absorption and stool formation Structure. Organized for regulated passage: Muscular organ Vascular - to facilitate nutrient absorption Enervated - to maintain control the Gastrointestinal tract is organised in four layers. 1. Mucosa : epithelium - columnar, glandular, squamous cells, muscularis mucosa smooth muscles. 2. Submucosa - connective tissue with blood vessels, lacteals, nerves, neurons. 3. Muscularis externa - two layers of muscles circular and longitudinal. -smooth muscle 4. Adventitious coat - loose epithelium 1 Structure - smooth muscle Spindle shaped cells Bundles - 75-200 Central nucleus Mitochondria - energy transactions for contraction Vesicles - calcium required to participate in contraction cells are linked - gap junctions Protein of contraction - actin and myosin. Organized in two layers - Longitudinal and circular Structure - vasculature Many arteries provide blood to the GIT Rate of blood flow is approximately 1.5 l/min increased by action of hormones and local reflexes Blood from GIT is collected by venules to veins to the Portal vein taken to Liver Collected from liver to Inferior vena cava - to heart. Lymph - serous fluid propelled by organ rhythm Lymph ducts to the Thoracic duct to Inferior venacava Pass through lymph nodes for filteration. Structure - Enervation Controlled by two nervous systems - extrinsic & intrinsic Extrinsic - Somatic and Autonomic Somatic - control of voluntary/ striated muscles ,eg. In mouth and anus Autonomic - Sympathetic and Parasympathetic -Coordination of smooth muscle activity by reflexes - not under voluntary control Organs - brain, spinal chord neurons with long fibres Sympathetic nervous system - transmitter is Adrenalin - relaxing effect Para sympathetic system - Acetyl choline - change cell membrane permeability to allow water uptake and become tense Principal nerve is the Vagus, but there are 12 pairs of cranial nerves 2 Structure - Enervation Electrical conductivity - cell to cell Longitudinal to circular muscles Electrical conductivity influenced by chemical constituents. Rhythmic contraction activity in the GIT- unidirectional. Functions of the enteric nervous system 1. Contraction of smooth muscles - create peristaltic wave 2. Release secretions from cells in the GIT 3. Release hormones from endocrine cells 4. Control patterns of blood flow 5. Control activities of the immune cells. Intrinsic nervous system - Meisner and Aurobach Systems. Local control Complete initiated motion Influenced by substances in extra-cellular fluid. Which set off receptors in GIT. Regulatory peptides Control of digestive function - motility, intestinal absorption, cell growth, secretion of digestive enzymes, electrolytes, water, other hormones, and mucins Gastrin - acid secretion - G cells Cholecystokinin - CCK - pancreatic enzyme secretion Secretin - bicarbonate secretion Gastric inhibitory peptide Somatostatin - paracrine action Vasoactive intestinal peptide - nerurocrine 3 GIT hormones GIT hormones Gastrin - Synthesized and secreted by G cells of antrum Stimulates - acid secretion in fundic cells, pepsinogen secretion, gastric motility and trophic action on GIT cells stimulated by vagal stimulation, ingestion of food, distension, fat GIP - inhibits gastric secretion and stimulates intestinal secretions and insulin secretion CCK - secreted by I- cells of intestine into blood stream Stimulates - enzyme secretion from pancreas, contraction of the gall bladder augments secretin action Stimulated by protein and fat in duodenum Motilin - Contraction of the smooth muscle Secretin - secreted by S cells if the intestine. Target cell pancreatic acinar cells for release of bicabonate and water. Substance P - neuropeptide increases blood flow to the GIT, Inhibits acid secretions, intestinal motility and pancreatic enzyme secretion. Somatostatin - by D cells- distributed at many locations. Inhibitory in function paracrine in effect VIP - released from nerves - stimulates GIT bicarbonate secretions and relaxes sphincters Mouth and esophagus Mouth - oral cavity entry way to the digestive tract Pharynx- orderly passage of the food Esophagus - passage way to distal GIT - Lower esophageal sphincter Chewing - tearing & grinding action by teeth, facilitated by the tongue and by mixing in of saliva Saliva - many triggers Functions - moisten food, dissolve & disperse sapid substances Oral cavity lubrication, enzymatic digestion - amylase, lipase Composition 99% water and ions (Na, K, HCO3, NH3) <.5% protein, mucins 4 Rumen Mouth and esophagus Three sets of salivary glands -sublingual, parotid, submaxillary control - involuntary and voluntary control. Swallowing - reflex - under neuronal control Can be voluntary Vomiting - not reverse peristalsis, brought on by increasing pressure by action of the diaphragm Ruminants - suborder Ruminantia - animal families - Giraffidae, Cervidae, Antilocapridae, Bovidae and Camelidae eg. giraffes, deer, antelope, cattle and camels. Chew their cud, Complex of four chambers Reticulum - recepticle - connected to rumen and omasum by reticulo orifice - participates in regurgitation and passage of liquid to omasum Rumen - fermentation chamber, muscle pillars Omasum - water absorption - may have nutrient absorption ability Abomasum - comparable to the stomach in non-ruminants papilae Reticulu m Rumen stratified squamus epithelium organized as different structures in the three forestomachs of ruminants Omasum leaves 5 Table 1. Relative number and mass of microbial organisms in the rumen Number Net mass Percent of total Group per mL (mg/100 ml) microbial mass Small Bacteria 1 x1010 1600 60-90 Selenomonads 1 x 108 300 Oscillospira flagellates 1 x 106 25 Entodinia 3 x 105 300 Dasytricha+Diplodinia 3 x 104 300 Isotricha+epidinia 1 x 104 1100 Fungi 1 x104 - Ciliated protozoa 10-40 5-10 Nitrogenous feed fermentation in the rumen Table 2. Effect of dilution rate on fermentation products in a continuous fermentor with mixed rumen population and using glucose as an energy source. Dilution (turnover) rate (h) Product 0.02 0.06 0.12 Acetatea 1.18 1.11 1.13 Propionatea 0.16 0.22 0.26 Butyratea 0.23 0.18 0.15 a 1.04 Methane 1.67 1.34 a 5.59 5.19 5.03 YATPb 7.50 11.6 16.70 ATP Yglucosec 42.40 60.20 83.90 Nitrogen in cells (%) 9.90 - 12.00 Microbial crude protein/mole glucose (g) 26.1 - 61.20 Adapted from Van Soest (1994) Microbes in the Gut. In 2nd Edition Nutritional Ecology of the Ruminant. Comstock Publishing of Cornell University Press Ithaca and London. p – 253-280 a Moles produced per mole glucose fermented b VFA production during fermentation of carbohydrates c g cells/mole ATP g cells/mole glucose 6 Abomasum/Stomach The stomach 11% of GIT in mature animals Two parts: Fundus and antrum Invaginated inner lining - gastric pits Three muscle layers - circular, longitudinal and oblique distal stomach - circular muscles thicken into pyloric sphincter Gastric pits Gastric pits Numerous types of cells produce variety of products 1. Mucoid cells - mucous 2. Oxyintic/parietal cells - acid, intrinsic factor 3. Chief/peptic cells - enzymes -pepsinogen, renin 4. Enterochromafin cells - histamine Cells in antral region G - gastrin and Dsomatstatin Hersey and Sachs 1998 Gactric acid secretion, Physiol Rev 75:155-189. 7 Stomach emptying Food entry into abomasum - continuous meal eating animals - exponential emptying related to volume movement includes - propulsion - distal Grinding - demunition of food retropulsion - retention of digesta Factors affecting emptying: pH, osmolarity, fat, fatty acid, ionization, fatty acid unsaturation, Tryptophan. Liver and its function Liver : excretory, digestive, metabolic and immunity. Urea synthesis from ammonia for excretion, Detoxification of toxins and RBC pigments for excretion Bile salts synthesis - aid in fat digestion Synthesis of Proteins, glycogen, glucose and cholesterol Haemopoietic for embryo Storage of minerals and vitamins. Cells of the liver: Endothelial cells - line sinusoids - filter Kupffer cells - macrophage like function Adipocytes - fat synthesis and vitamin storage pit cells - endocrine function regulatory role hepatocytes - principal cell - protein synthesis and bile salt synthesis by conjugation of bile acid with amino acid. Stored in gall bladder Bile - surfactant for fat digestion. Enterohepatic circulation Gall bladder - intestine deconjugation in L intestine - absorbed and returned to liver. 8 Pancreatic Function Exocrine function: digestive enzymes Enzymes proteins synthesized by cells in the ascinus. Delivered to the small intestine as inactive zymogen granules. Endocrine function: Protein hormones secreted into blood. Synthesis by islet cells Stages in Protein synthesis and secretion: Synthesis, Segregation, Intracellular transport, Concentration, Intracellular storage, Discharge. Triggers: CCK, neural input and insulin Pancreatic enzymes Proteases Trypsinogens - endopeptidase, activated by entrokinase of instestine, autocatalysed and catalyses other proteases - large peptides Chymotrypsimnogens - large peptides Proelastease - connective tissue digestion - large peptides Pro-carboxypeptidase - small peptides Pancreatic amylase - starch - disaccharides TG - lipase - hydrolyses fatty acids from the top and bottom of TG Pro-co-lipase - activator for lipase Cholesterol esterase - hydrolyses cholesterol esters Phospholipase A2 - fatty acid from phospholipids Intestines Cattle - Sheep/goats SI Cecum LI capacity SI Cecum LI capacity 46 m 1m 10m 26m 0.6m 6m 252 L 356 L 29 L 44L DNAase and RNAase - digestion of DNA and RNA 9 Structure of the small intestine Sections of the small intestine: duodenal - short jejunal - extended and highly folded ileal - distal portion - under control of ileo cecal sphincter Crypt - regeneration centre of the cells on the villi Villi - finger like projections. Villus -centre have muscle , blood capillaries, lacteals, Villi and Crypt Figure 44. Microscopic appearance of normal duodenal villi in the pig as viewed from the lumen.(From and with permission of Hornich et al., 1973 and The American College of Veterinary Pathologists, Inc.). Villi: monolayer of cells origin from bottom of crypt and migrate to top and are extruded from the tip. Differentiate into many cell types goblet cells - mucous endocrine cells - hormones tuft cells - chemoreceptors paneth cells - secretory at base of crypt - Ig - G&A and secretory component Columnar cells - enterocytes 10 Endocrine cells EC SIKMo DLN- Seretonin, Substance P Secretin Cholecystokinin - Intact protein and fat Gastric inhibitory peptide Motilin Somatostatin Glucagon -like - peptide Neurotensin Enterocyte • Principal cell type - organized in a single layer • Attached to a basement membrane • Participate in membrane bound enzymatic hydrolysis • Tight junctions • Microvilli - glycocalyx - trap susbtrate • IMP - scharidases - sucrase, maltase, lactase, transport proteins eg SGLT1, CaBP, B12+IF receptor • Basal membrane - Na/K ATPase, GLUT2 11 Small intestinal digestion Luminal digestion: Pancreatic enzymes proteases, - protein to peptides amino acids amylase - starch to limit dextrins and maltose lipase and co-lipase - lipids to fatty acids, cholesterol and glycerol, nucleases - recovery of phosphorus Membrane bound digestion: disacharidases hexoses Cellular digestion: peptides to amino acids 12 Absorption Peptides and Amino acids Glucose: Carrier mediated absorption SGLT1 Saturable kinetics, Graded substrate specificity P-D gradient in expression Inducible by substrate Large protein with 7 trans membrane spans Released by GLUT2 at BLM Fascilated transport - GLUT5 - fructose at mucosal and released by GLUT2 at BLM not energy requiring AA - poorly absorbed , energy requiring Peptides - transport across mucosal membrane into enterocyte depends upon: chain length dipeptide > tripeptide>>tetrapeptide 3 types of transporters Na dependant - primary active Na independent - secondary active H+ [] gradient - tertiary active Peptides hydrolysed intracellularly and released into blood by electro-chemical gradient. Factors controlling peptide transport and hydrolysis Peptides Di P Tri P Tetra P LL Di P DD Di P DL, LD P LD P DL P transport +++ ++ + +++ poor +++ hydrolysis +++ ++ + +++ ++ + ++ Configuration of AA at COOH end of peptide is important for transport Transporters expressed in P-D gradient. All nutrients in blood transported to liver by portal vein. Blood flows through the liver to hepatic veinules and collected into hepatic vein to inferior vena cava to heart for general circulation 13 Un-absorbed components of digesta enters Large intestine Water re-absorption Fermentative digestion of digestible substrate if ileal digesta - high residual starch fermented by rumen type bacteria consumption - excretion = gross measure of nutrient extraction Formation of feces and evacuation 14