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فیزیولوژی تکمیلی Advanced Physiology (part 2, Digestive system) By: A. Riasi (PhD in Animal Nutrition & Physiology) Introduction The primary function of digestive system Intracellular and extracellular process in digestive system Three specific regions in most animals digestive system: Foregut Midgut Hindgut Introduction Digestive systems perform four basic digestive process: Motility Propulsive movements Mixing movements Secretion Digestion Absorption Digestive system properties The digestive system of animals consists of: The digestive tract (gastrointestinal tract) Accessory digestive organs Additional function of the digestive tract: Osmoregulation Endocrine secretion Immune function Elemination of toxins Digestive system properties Regulation of digestive function is complex and synergetic The smooth muscle cells of digestive system are single unit type The nervous system control digestive system function Intrinsic nerve plexuses Extrinsic nerves Structure of digestive tract The four layers are the same from esophagus to anal canal Structure of digestive tract Structure of digestive tract Adapted from: Anatomy and Physiology of Domestic Animal, Akers and Denbow, 2013 Intrinsic nervous system Adapted from: Sherwood et al., Animal Physiology Intrinsic nervous system Intrinsic nervous system The enteric nervous system of a rat’s stomach Adapted from: Sherwood et al., Animal Physiology Intrinsic nervous system Receptor activation alters digestive activity through neural reflexes and hormonal pathways Three different types of sensory receptors: Chemoreceptors Mechanoreceptors (pressure receptors) Osmoreceptors Salivary glands Acini is a secretary unit of salivary gland Each acini secrete fluid into collecting ducts Water Electrolytes Mucus Enzymes Salivary glands Three major parts of salivary glands Parotid glands Submaxillary glands Sublingual glands Salivary glands In the histological sections of salivary gland shown above, the cells stained pink are serous cells, while the white, foamy cells are mucus-secreting cells. Control of salivary secretion Adapted from: Sherwood et al., Animal Physiology Salivary secretion in sheep Salivary glands Total salivary Characteristics volumes (L d) Site of reflexogenic stimuli Parotids 3-8 Inferior molars 0.7-2 Mouth, esophagus, ruminoreticulum Mouth, esophagus, ruminoreticulum Palatine, buccal Pharyngeal, Submaxillary 2-6 Sublingual, labial Total volume 0.1 0.4-0.8 6-16 Serous, isotonic, strongly buffered Serous, isotonic, strongly buffered Isotonic, strongly buffered Mouth, esophagus, ruminoreticulum Mucous, hypotonic, Mouth during feeding, weakly buffered not cudding Very mucous, hypotonic, weakly buffered Mouth Swallowing Eating and swallowing are complex neuromuscular activities consisting essentially of two stages: Oropharyngeal stage Esophageal stage Adapted from: Sherwood et al., Animal Physiology Esophagus Anatomically and functionally, the esophagus is the least complex section of the digestive tube It contains the crop in poultry Esophagus There are two physiologic sphincters: Upper and lower esophageal sphincters. In ruminants, a nasopharyngeal sphincter is present. Esophagus in birds Stomach Adapted from: Sherwood et al., Animal Physiology Gastric emptying and mixing Adapted from: Sherwood et al., Animal Physiology Factors that influence the rate of gastric emptying A- Stomach factors Amount of chyme in the stomach (more effect) Vagus nerve Stomach hormone gastrine The degree of fluidity of the chyme B- Duodenum factors Fat (more effect) Acid Hypertonicity Distention The duodenal factors trigger either neural or hormonal responses Neural response is mediated through two reflexes: Intrinsic nerve plexus (short reflex) Autonomic nerves (long reflex) (These reflexes are called the enterogastric reflex) The hormonal response involves the release duodenal’s several hormones: Secretin CCK Gastric inhibitory peptide or glucose-dependent insulinotrophic peptide Avian pancreatic polypeptide (APP) (These hormones are known as enterogastrones) Stomach Two distinct areas for secretion of gastric digestive juice: Oxyntic mucosa (contain three type of cells) Mucous cells Chief cells Parietal cells or oxyntic cells Pyloric gland area (PGA): Mucous cells Chief cells Gastric secretion cells: Gastric secretion cells Exocrine cells Paracrine cells Endocrine cells Adapted from: Sherwood et al., Animal Physiology Mechanism of HCl secretion Adapted from: Sherwood et al., Animal Physiology Adapted from Animal Physiology, by Sherwood et al., 2005 Control of gastric secretion Cephalic phase (Mediated by vagus nerve and acetylcholin) Gastric phase (gastrin has the main effect) Intestinal phase (intestinal gastrin has the main effect) Bird stomach Proventriculus-gizzard processes of digestion in birds There are two type of glands into the proventriculus: Simple mucosal glands that secrete mucus Submucosal glands that secrete HCl and pepsinogen Interestingly, unlike in mammals, both HCl and pepsinogen are synthesized with in the same cell (chief or oxynticopeptic cell) Bird stomach The mucosal lining of the gizzard is covered by koilin In birds myoglobin content of the gizzard is approximately 100-fold greater than the breast muscle, and mitochondrial numbers are also elevated. Ruminant stomach Rumen development Rumen microbiology and fermentation A large community of microorganisms are presented in the forestomach. Bacteria Protozoa Fungi Ruminant stomach Rumen muscles 1 3 4 2 5 Solid lines: internal oblique fiber (ruminal pillars, lips of reticular groove, omasal pillar); broken lines: longituidal fibers; wave lines: circular fibers. At any given place, there are only two muscle layers in the stomach wall. 1= cardia; 2= reticulum; 3= rumen 4= omasum; 5= abomasum. Ruminant stomach Rumen development Undeveloped Rumen Developed Rumen Ruminant stomach Absorptive surface area is enhanced by increasing: Papillae length Papillae width Papillae density Ruminant stomach The interior surface of the rumen forms numerous papillae Ruminant stomach The 4 layers of the rumen epithelium: stratum corneum (SC), stratum granulosum (SG), stratum spinosum (SS) and stratum basal (SB). Ruminant stomach Absorption VFA’s from the ruminal wall Ruminant stomach Relationship between ruminal pH and the associated changes in the ruminal environment and ruminal epithelial function (reproduced from Penner and Beauchemin, 2010). Ruminant stomach Rumination is centrally mediated by the "gastric centers". Tactile stimulation of the reticular and ruminal epithelia is a powerful stimulus for rumination. Ruminant stomach The ruminal movements serve to: Mix the ingesta Aid in eructation of gas Propel fluid and fermented foodstuffs into the omasum. A cycle of contractions occurs 1 to 3 times per minute. Ruminant stomach Two types of contractions are identified: Primary contractions Secondary contractions Small intestine Three parts of small intestine Duodenum Jejunum Ileum Small intestine Adapted from http://www.ufrgs.br/imunovet/molecular_immunology/gastrointestinal.html) Small intestine Adapted from http://www.ufrgs.br/imunovet/molecular_immunology/gastrointestinal.html) Small intestine The Meckel’s diverticulum is found in birds small intestine Pancreas Adapted from: Sherwood et al., Animal Physiology Pancreas Exocrine functions Enzyme Substrate Action Trypsin, Chymotrypsin, Elastase Peptides Endopeptidases; cleave bonds between amino acids Carboxypeptidase and Aminopeptidase Peptides Exopeptidases; cleave bonds at the terminus of a peptide α - amylase Polysaccharides: starch and glycogen Endoglycosidase; cleaves bonds between carbohydrate monomers to produce maltose and short carbohydrate chains. Pancreatic lipase Triacylglycerols and 1,2 diacylglycerols Fatty acids, glycerol and 2 monoacylglycerol Adapted from wikivet (http://en.wikivet.net) Pancreas Control of exocrine secretions Neural controls Endocrine controls Cholecystokinin Secretin Gastrin Pancreas Liver and gallbladder Liver and gallbladder Liver blood flow 75% venous blood from the portal vein 25% from the hepatic artery Adapted from: Sherwood et al., Animal Physiology Liver and gallbladder Adapted from: Sherwood et al., Animal Physiology Liver and gallbladder Adapted from http://www.ufrgs.br/imunovet/molecular_immunology/gastrointestinal.html) Liver and gallbladder Liver and gallbladder Lymph formation in liver Liver and gallbladder Hepatic phagocytic system Liver and gallbladder Two stage for bile secretion Hepatocytes secrete bile into canaliculi Bile modification in bile ducts Liver and gallbladder Pattern and control of bile secretion Effect of cholecystokinin Effect of secretin