BIO2305 The Digestive System The Digestive System The alimentary canal or gastrointestinal (GI) tract digests and absorbs food Alimentary canal: Oral cavity Pharynx Esophagus Stomach Small intestine Large intestine Rectum Accessory digestive organs: Teeth Tongue Salivary glands Liver Gallbladder Pancreas Organization of the Digestive Tract Mucosa - lines digestive tract (mucous epithelium) Moistened by glandular secretions Lamina propria and epithelium form mucosa Submucosa - layer of dense irregular connective tissue Muscularis externa - smooth muscle arranged in circular and longitudinal layers Serosa - serous membrane covering most of the muscularis externa 1 Digestive System Activities The GI tract is a “disassembly” line Nutrients become more available to the body in each step Ingestion – taking food into the digestive tract Mechanical digestion – chewing, churning food, segmentation Propulsion – swallowing and peristalsis Chemical digestion – catabolic breakdown of food Absorption – movement of nutrients from the GI tract to the blood or lymph Excretion – elimination of indigestible solid wastes Basic Processes of the Digestive System Motility Movement of digestive materials Visceral smooth muscle Tonic contractions Sustained, long-lasting Smooth muscle sphincters and stomach Phasic contractions Rhythmic cycles of contraction and relaxation initiated by pacemaker cells (modified smooth muscles cells) Last a few seconds Peristalsis moves bolus forward Segmentation mixes 2 Autonomous Smooth Muscle GI Pacemaker cells (modified smooth muscles cells) are called Interstitial cells of Cajal (ICCs) Slow wave potentials digestive tract’s basic electrical rhythm cycles between depolarizations and repolarizations temporal summation slowly brings the cell membrane to threshold Action potentials are transmitted throughout smooth muscle via gap junctions Rate and frequency of action potentials are effected of various mechanical, neural, and hormonal factors Motility Peristalsis – waves that move a bolus forward Segmentation – to churn and fragment a bolus No net forward movement Ingestion and Mechanical Digestion Food is ingested Mechanical digestion begins (chewing) Propulsion is initiated by swallowing Salivary amylase begins chemical breakdown of starch The pharynx and esophagus serve as conduits to pass food from the mouth to the stomach Uvula guards opening to pharynx 3 Salivary Glands (three types) Parotid, sublingual, and submandibular glands produce saliva Stimulated by thought of food or ingested food Secreted from serous and mucous cells of salivary glands Watery solution includes electrolytes, buffers, glycoproteins, antibodies, enzymes Functions include: Lubrication, moistening, and dissolving Salivary Amylase – a salivary enzyme that initiates of digestion of complex carbohydrates Strong sympathetic stimulation inhibits salivation and results in dry mouth Swallowing (Deglutition) Process Involves the coordinated activity of the tongue, soft palate, pharynx, esophagus and 22 separate muscle groups Three phases: Buccal phase – tongue pushes bolus against soft palate and forced into the oropharynx, triggering swallowing reflex – controlled by the medulla and lower pons Pharyngeal – esophageal sphincter relaxes while epiglottis closes Esophageal – bolus moves into esophagus propelled by peristalsis and into stomach Functions of the Stomach Receives and holds ingested food Digests food both physically and chemically Delivers chyme to the small intestine Enzymatically digests proteins with pepsin Secretes intrinsic factor required for absorption of vitamin B12 4 Stomach Lining The stomach is exposed to the harshest conditions in the digestive tract To keep from digesting itself, the stomach has a mucosal barrier with: A thick coat of bicarbonate-rich mucus on the stomach wall Epithelial cells that are joined by tight junctions Gastric glands that have cells impermeable to HCl Glands of the Stomach Gastric glands have a variety of secretory cells Mucous neck cells secrete mucus Parietal cells secrete HCl and intrinsic factor Chief cells secrete pepsinogen HCl converts pepsinogen to pepsin in the stomach Pepsin activates more pepsin via positive feedback Enteroendocrine cells secrete gastrin, histamine, cholecystokinin (CCK) Regulation of Gastric Secretion Neural and hormonal mechanisms regulate the release of gastric juice Stimulatory and inhibitory events occur in three phases: Cephalic (reflex) phase: prior to food entry Gastric phase: once food enters the stomach Intestinal phase: as partially digested food enters the duodenum 5 Cephalic Phase Cephalic phase prepares stomach to receive ingested material Directed by CNS (parasympathetic nervous system) via the vagus nerve (CN X) Stimulated by sight, smell, taste, or thought of food Accelerates gastric juices Inhibited by loss of appetite, depression Gastric Phase Enhanced secretion of gastric juices due to the arrival of food in the stomach Homogenize and acidify chyme Production of pepsinogen - digestion of proteins Stimulated by Stomach distension - activation of stretch receptors Chemoreceptors detects peptides, caffeine, and rising pH Neural - plexuses Hormonal - secretion of gastrin Inhibitory events include: pH lower than 2 Emotional upset that overrides the parasympathetic division Intestinal phase Intestinal phase – release of hormones controls the rate of gastric emptying Excitatory phase – distension of duodenum, presence of partially digested foods Releases enterogastrones that inhibit gastric secretion: CCK, GIP, Secretin Inhibited by low pH, presence of fatty, acidic, or hypertonic chyme, and/or irritants in the duodenum 6 Release of Gastric Juice Regulation of HCl Secretion Hydrochloric acid (HCl) secretion is stimulated by ACh, histamine, and gastrin through second-messenger systems Release of HCl: Is low if only one ligand binds to parietal cells Is high if all three ligands bind to parietal cells Antihistamines block Histamine H2 receptors and decrease HCl release 7 Gastric Contractile Activity Peristaltic waves move toward the pylorus at the rate of 3 per minute This basic electrical rhythm (BER) is initiated by pacemaker cells Most vigorous peristalsis and mixing occurs near the pylorus Chyme is either: Delivered in small amounts to the duodenum or Forced backward into the stomach for further mixing Regulation of Gastric Emptying Gastric emptying is regulated by: The neural enterogastric reflex Hormonal (enterogastrone) mechanisms These mechanisms inhibit gastric secretion and duodenal filling Carbohydrate-rich chyme quickly moves through the duodenum Fat-laden chyme is digested more slowly causing food to remain in the stomach longer Gastrointestinal Hormones Gastrin Release is stimulated by presence of protein in stomach Secretion inhibited by accumulation of acid in stomach Acts in several ways to increase secretion of HCl and pepsinogen Enhances gastric motility, stimulates ileal motility, relaxes ileocecal sphincter, induces mass movements in colon Helps maintain well-developed, functionally viable digestive tract lining Secretin Presence of acid in duodenum stimulates release Inhibits gastric emptying in order to prevent further acid from entering duodenum until acid already present is neutralized Inhibits gastric secretion to reduce amount of acid being produced Stimulates pancreatic duct cells to produce large volume of aqueous NaHCO3 secretion Stimulates liver to secrete NaHCO3 rich bile which assists in neutralization process Along with CCK, is trophic to exocrine pancreas CCK Inhibits gastric motility and secretion Stimulates pancreatic acinar cells to increase secretion of pancreatic enzymes Causes contraction of gallbladder Along with secretin, is trophic to exocrine pancreas Implicated in long-term adaptive changes in proportion of pancreatic enzymes in response to prolonged diet changes Important regulator of food intake GIP Glucose-dependent insulinotrophic peptide Stimulates insulin release by pancreas 8 Regulation of Gastric Emptying Digestion And Absorption In The Stomach Preliminary digestion of proteins by pepsin Permits digestion of carbohydrates Very little nutrient absorption Although lipid soluble drugs (e.g.: alcohol, aspirin) can cross the gastric mucosa and enter the blood stream Small Intestine Important digestive and absorptive functions Secretions and buffers provided by pancreas, liver, gall bladder Three subdivisions: Duodenum Jejunum Ileum Ileocecal sphincter - transition between small and large intestine 9 Small Intestine Structural modifications of the small intestine wall increase surface area Plicae circulares – deep circular folds of the mucosa and submucosa Villi – fingerlike extensions of the mucosa Microvilli – tiny projections of absorptive mucosal cells’ plasma membranes Small Intestine The epithelium of the mucosa is made up of: Absorptive cells and goblet cells Enteroendocrine cells Interspersed T cells called intraepithelial lymphocytes (IELs) Cells of intestinal crypts secrete intestinal juice Secreted in response to distension or irritation of the mucosa Slightly alkaline and isotonic with blood plasma Largely water, enzyme-poor, but contains mucus Small Intestine Glands of the duodenum Moisten chyme Help buffer acids Maintain digestive material in solution Hormones Secretin - produces alkaline buffers, increase bile by liver and pancreas Cholecystokinin (CCK) – increase pancreatic enzymes, stimulates contraction of gall bladder, reduces hunger sensation GIP – stimulates release of insulin, inhibits gastric secretion and motility 10 Activities of Major Digestive Tract Hormones The Liver The largest gland in the body Performs metabolic and hematological regulation and produces bile Histological organization Lobules containing single-cell thick plates of hepatocytes Lobules unite to form common hepatic duct Duct meets cystic duct to form common bile duct 11 The Liver Hexagonal-shaped liver lobules are the structural and functional units of the liver Composed of hepatocytes (liver cells) Hepatocytes’ functions include: Production of bile Processing blood-borne nutrients Storage of fat-soluble vitamins Detoxification Production of plasma proteins Secreted bile flows between hepatocytes toward the bile ducts in the portal triads Composition of Bile A yellow-green, alkaline solution containing bile salts, bile pigments, cholesterol, neutral fats, phospholipids, and electrolytes Bile salts are cholesterol derivatives that: Emulsify fat Facilitate fat and cholesterol absorption Help solubilize cholesterol Enterohepatic circulation recycles bile salts The chief bile pigment is bilirubin (stercobilin), a waste product of heme The Gallbladder Thin-walled, green muscular sac on the ventral surface of the liver Stores and concentrates bile by absorbing its water and ions Releases bile via the cystic duct, which flows into the bile duct 12 Regulation of Bile Release Acidic, fatty chyme causes the duodenum to release: Cholecystokinin (CCK) and secretin into the bloodstream Bile salts and secretin transported in blood stimulate the liver to produce bile Vagal stimulation causes weak contractions of the gallbladder Cholecystokinin (CCK): The gallbladder to contract The hepatopancreatic sphincter to relax As a result, bile enters the duodenum Regulation of Bile Release 13 The Pancreas Pancreatic duct penetrates duodenal wall at duodenal papilla Endocrine functions: Modulate Blood Glucose Levels: Insulin (to ↓BGL) and glucagon (to ↑BGL) Endocrine hormones enter venous bloodstream Exocrine functions: Digestion: Pancreatic juice secreted into small intestine to break down all categories of foodstuff Acini – clusters of secretory acinar cells that contain zymogen granules with digestive enzymes Exocrine juices enter duodenum through duodenal papilla, controlled by hepatopancreatic sphincter Composition of Pancreatic Juice Water solution of enzymes and electrolytes (primarily HCO3–) Neutralizes acid chyme Provides optimal environment for pancreatic enzymes Enzymes are released in inactive form and activated in the duodenum Examples include: Trypsinogen is activated to trypsin Procarboxypeptidase is activated to carboxypeptidase Active enzymes secreted: Pancreatic amylase, lipases, and nucleases These enzymes require ions or bile for optimal activity Regulation of Pancreatic Secretion Secretin and CCK are released when fatty or acidic chyme enters the duodenum CCK and secretin enter the bloodstream Upon reaching the pancreas: CCK induces the secretion of enzyme-rich pancreatic juice Secretin causes secretion of bicarbonate-rich pancreatic juice Vagal stimulation also causes release of pancreatic juice 14 Regulation of Pancreatic Secretion Digestion in the Small Intestine As chyme enters the duodenum: Carbohydrates and proteins are only partially digested No fat digestion has taken place Digestion continues in the small intestine Chyme is released slowly into the duodenum Because it is hypertonic and has low pH, mixing is required for proper digestion Required substances needed are supplied by the liver Virtually all nutrient absorption takes place in the small intestine Motility in the Small Intestine The most common motion of the small intestine is segmentation It is initiated by intrinsic pacemaker cells (Cajal cells) Moves contents steadily toward the ileocecal valve After nutrients have been absorbed: Peristalsis begins with each wave starting distal to the previous Meal remnants, bacteria, mucosal cells, and debris are moved into the large intestine Control of Motility Local enteric neurons of the GI tract coordinate intestinal motility Cholinergic neurons cause: Contraction and shortening of the circular muscle layer Shortening of longitudinal muscle Distension of the intestine Other impulses relax the circular muscle The gastroileal reflex and gastrin: Relax the ileocecal sphincter Allow chyme to pass into the large intestine 15 Functions of the Large Intestine Reabsorb water and compact material into feces Absorb vitamins produced by bacteria Store fecal matter prior to defecation Functions of the Large Intestine Other than digestion of enteric bacteria, no further digestion takes place Vitamins, water, and electrolytes are reclaimed Major function: propulsion of fecal material toward the anus Though essential for comfort, the colon is not essential for life Motility of the Large Intestine Haustral contractions Slow segmenting movements that move the contents of the colon Haustra sequentially contract as they are stimulated by distension Presence of food in the stomach: Activates the gastrocolic reflex Initiates peristalsis that forces contents toward the rectum The Rectum Last portion of the digestive tract Terminates at the anal canal Internal and external anal sphincters Defecation reflex triggered by distention of rectal walls 16 Defecation Distension of rectal walls caused by feces: Stimulates contraction of the rectal walls Relaxes the internal anal sphincter Voluntary signals stimulate relaxation of the external anal sphincter and defecation occurs Regulation of Digestion Intrinsic control by local centers Autonomous smooth muscle pacesetter cells Intrinsic nerve plexuses and sensory receptors Extrinsic control ANS GI hormones Mechano- and chemoreceptors respond to: Stretch, osmolarity, and pH Presence of substrate, and end products of digestion They initiate reflexes that: Activate or inhibit digestive glands Mix lumen contents and move them along Nervous Control of the GI Tract Intrinsic controls Nerve plexuses near the GI tract initiate short reflexes Short reflexes are mediated by local enteric plexuses (gut brain) Extrinsic controls Long reflexes arising within or outside the GI tract Involve CNS centers and extrinsic autonomic nerves Parasympathetic reflexes 17 Enteric Nervous System Composed of two major intrinsic nerve plexuses: Submucosal nerve plexus – regulates glands and smooth muscle in the mucosa Myenteric nerve plexus – Major nerve supply that controls GI tract motility Segmentation and peristalsis are largely automatic involving local reflex arcs Linked to the CNS via long autonomic reflex arc Control of the Digestive System Local mechanisms coordinate response to changes in pH, physical, chemical stimuli Neural and hormonal mechanisms coordinate glands Hormonal mechanisms enhance or inhibit smooth muscle contraction Digestion And Absorption Of Nutrients Disassembling organic food into smaller fragments Hydrolyzing carbohydrates, proteins, lipids and nucleic acids for absorption 18 Chemical Digestion: Carbohydrates Begins in the mouth: Salivary and pancreatic enzymes catabolize into disaccharides and trisaccharides Brush border enzymes catabolize into monosaccharides Absorption of monosaccharides occurs across the intestinal epithelia Absorption: via cotransport with Na+, and facilitated diffusion Enter the capillary bed in the villi Transported to the liver via the hepatic portal vein Enzymes used: salivary amylase, pancreatic amylase, and brush border enzymes Chemical Digestion: Proteins Low pH destroys tertiary and quaternary structure Enzymes used include pepsin, trypsin, chymotrypsin, and elastase Liberated amino acids are absorbed Absorption: similar to carbohydrates Enzymes used: pepsin in the stomach Enzymes acting in the small intestine: Pancreatic enzymes – trypsin, chymotrypsin, and carboxypeptidase Brush border enzymes – aminopeptidases, carboxypeptidases, and dipeptidases Lipid Digestion and Absorption Lipid digestion utilizes lingual and pancreatic lipases Bile salts improve chemical digestion by emulsifying lipid drops Lipid-bile salt complexes called micelles are formed Micelles diffuse into intestinal epithelia Small micelles enter bloodstream and are sent to liver Large micelles enter lymphatic system as chylomicrons 19 Chemical Digestion: Fats Absorption – diffusion into intestinal cells where they: Combine with proteins and extrude chylomicrons Enter lacteals and are transported to systemic circulation via lymph Glycerol and short chain fatty acids are: Absorbed into the capillary blood in villi Transported via the hepatic portal vein Enzymes/chemicals used: bile salts and pancreatic lipase Fatty Acid Absorption Fatty acids and monoglycerides enter intestinal cells via diffusion They are combined with proteins within the cells Resulting chylomicrons are extruded They enter lacteals and are transported to the circulation via lymph Chemical Digestion: Nucleic Acids Enzymatic catabolism: Pancreatic ribonucleases and deoxyribonuclease in the small intestines Absorption: Active transport via carrier proteins Absorbed in villi and transported to liver via hepatic portal vein 20 Absorption of Water Water: nearly all (95%) that is ingested is reabsorbed Net osmosis occurs whenever a concentration gradient is established by active transport of solutes into the mucosal cells Water uptake is coupled with solute uptake, and as water moves into mucosal cells, substances follow along their concentration gradients Absorption of Vitamins Vitamins: Water-soluble vitamins are absorbed by diffusion Fat-soluble vitamins are absorbed as part of micelles Vitamin B12 requires intrinsic factor for absorption Electrolyte Absorption Most ions are actively absorbed along the length of small intestine Na+ is coupled with absorption of glucose and amino acids Ionic iron is transported into mucosal cells where it binds to ferritin Anions passively follow the electrical potential established by Na+ K+ diffuses across the intestinal mucosa in response to osmotic gradients Ca2+ absorption: Is related to blood levels of ionic calcium Is regulated by vitamin D and parathyroid hormone (PTH) 21