Digestive Processes Six essential activities

1. 2. 3. 4. 5. 6. Ingestion Propulsion Mechanical breakdown Digestion Absorption Defecation

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Figure 23.2 Gastrointestinal tract activities.

Ingestion
Mechanical breakdown Chewing (mouth) Churning (stomach) Segmentation (small intestine) Digestion Food Pharynx Esophagus Propulsion Swallowing (oropharynx) Peristalsis (esophagus, stomach, small intestine, large intestine) Stomach Absorption Lymph vessel Small intestine Large intestine Mainly H2O Feces Defecation
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Blood vessel

Anus

GI Tract Regulatory Mechanisms 1. Mechanoreceptors and chemoreceptors

Respond to stretch, changes in osmolarity and pH, and presence of substrate and end products of digestion Initiate reflexes that
Activate or inhibit digestive glands Stimulate smooth muscle to mix and move lumen contents

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GI Tract Regulatory Mechanisms 2. Intrinsic and extrinsic controls

Short reflexes - enteric nerve plexuses (gut brain) respond to stimuli in GI tract Long reflexes respond to stimuli inside or outside GI tract; involve CNS centers and autonomic nerves Hormones from cells in stomach and small intestine stimulate target cells in same or different organs to secrete or contract

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Figure 23.4 Neural reflex pathways initiated by stimuli inside or outside the gastrointestinal tract.

External stimuli (sight, smell, taste, thought of food) Visceral afferents

Central nervous system

Long reflexes
Extrinsic visceral (autonomic) efferents

Internal (GI tract) stimuli

Chemoreceptors, osmoreceptors, or mechanoreceptors

Local (intrinsic) nerve plexus ("gut brain")

Effectors: Smooth muscle or glands

Short reflexes Gastrointestinal wall (site of short reflexes) Lumen of the alimentary canal Response: Change in contractile or secretory activity

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Gastric Gland Secretions Glands in fundus and body produce most gastric juice Parietal cell secretions
Hydrochloric acid (HCl)
pH 1.53.5 denatures protein, activates pepsin, breaks down plant cell walls, kills many bacteria

Intrinsic factor
Glycoprotein required for absorption of vitamin B12 in small intestine

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Gastric Gland Secretions Chief cell secretions

Pepsinogen - inactive enzyme
Activated to pepsin by HCl and by pepsin itself (a positive feedback mechanism)

Lipases
Digest ~15% of lipids

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Gastric Gland Secretions Enteroendocrine cells

Secrete chemical messengers into lamina propria
Act as paracrines
Serotonin and histamine

Hormones
Somatostatin (also acts as paracrine) and gastrin

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Mucosal Barrier Harsh digestive conditions in stomach Has mucosal barrier to protect
Thick layer of bicarbonate-rich mucus Tight junctions between epithelial cells
Prevent juice seeping underneath tissue

Damaged epithelial cells quickly replaced by division of stem cells

Surface cells replaced every 36 days

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Homeostatic Imbalance Gastritis

Inflammation caused by anything that breaches mucosal barrier

Peptic or gastric ulcers

Erosions of stomach wall
Can perforate peritonitis; hemorrhage

Most caused by Helicobacter pylori bacteria Some by NSAIDs

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Figure 23.16 Photographs of a gastric ulcer and the H. pylori bacteria that most commonly cause it.

Bacteria

Mucosa layer of stomach

A gastric ulcer lesion

H. pylori bacteria

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Regulation of Gastric Secretion Neural and hormonal mechanisms Gastric mucosa up to 3 L gastric juice/day Vagus nerve stimulation secretion Sympathetic stimulation secretion Hormonal control largely gastrin
Enzyme and HCl secretion Most small intestine secretions - gastrin antagonists
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Regulation of Gastric Secretion Three phases of gastric secretion

Cephalic (reflex) phase conditioned reflex triggered by aroma, taste, sight, thought Gastric phase lasts 34 hours; gastric juice released
Stimulated by distension, peptides, low acidity, gastrin (major stimulus) Enteroendocrine G cells stimulated by caffeine, peptides, rising pH gastrin

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Stimuli of Gastric Phase

Gastrin enzyme and HCl release
Low pH inhibits gastrin secretion (as between meals)

Buffering action of ingested proteins rising pH gastrin secretion Three chemicals - ACh, histamine, and gastrin stimulate parietal cells through secondmessenger systems
All three are necessary for maximum HCl secretion

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Regulation of Gastric Secretion Intestinal phase

Stimulatory component
Partially digested food enters small intestine brief intestinal gastrin release

Inhibitory effects (enterogastric reflex and enterogastrones)

Chyme with H+, fats, peptides, irritating substances inhibition

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Enterogastric Reflex Three reflexes act to

Inhibit vagal nuclei in medulla Inhibit local reflexes Activate sympathetic fibers tightening of pyloric sphincter no more food entry to small intestine

Decreased gastric activity protects small intestine from excessive acidity

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Intestinal Phase Enterogastrones released

Secretin, cholecystokinin (CCK), vasoactive intestinal peptide (VIP)
All inhibit gastric secretion

If small intestine pushed to accept more chyme dumping syndrome

Nausea and vomiting Common in gastric reduction for weight loss

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Figure 23.17 Neural and hormonal mechanisms that regulate release of gastric juice. Stimulatory events
Cephalic phase 1 Sight and thought of food Cerebral cortex Conditioned reflex Hypothalamus and medulla oblongata Vagus nerve

Inhibitory events
Lack of stimulatory impulses to parasympathetic center Cerebral cortex 1 Loss of appetite, depression

2 Stimulation of taste and smell receptors

1 Stomach distension activates stretch receptors

Vagovagal reflexes

Medulla

Vagus nerve

Gastrin secretion declines Overrides parasympathetic controls

G cells

Gastric phase

Local reflexes

2 Food chemicals G cells (especially peptides and caffeine) and rising pH activate chemoreceptors

Sympathetic nervous system activation

1 Excessive acidity (pH < 2) in stomach 2 Emotional stress

Gastrin release to blood Stomach secretory activity Enterogastric reflex Local reflexes Vagal nuclei in medulla Pyloric sphincter Release of enterogastrones (secretin, cholecystokinin, vasoactive intestinal peptide) 1 Distension of duodenum; presence of fatty, acidic, or hypertonic chyme; and/or irritants in the duodenum

Intestinal phase

1 Presence of partially digested foods in duodenum or distension of the duodenum when stomach begins to empty

Intestinal (enteric) gastrin release to blood

Brief effect

Stimulate Inhibit

2 Distension; presence of fatty, acidic, partially digested food in the duodenum

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Response of the Stomach to Filling Stretches to accommodate incoming food

Pressure constant until 1.5 L food ingested
Reflex-mediated receptive relaxation
Coordinated by swallowing center of brain stem

Gastric accommodation
Plasticity (stress-relaxation response) of smooth muscle (see Chapter 9)

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Gastric Contractile Activity Peristaltic waves move toward pylorus at rate of 3 per minute
Basic electrical rhythm (BER) set by enteric pacemaker cells (formerly interstitial cells of Cajal) Pacemaker cells linked by gap junctions entire muscularis contracts

Distension and gastrin increase force of contraction

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Figure 23.19 Deglutition (swallowing).

Slide 1

Pyloric valve closed

Pyloric valve closed

Pyloric valve slightly opened

1 Propulsion: Peristaltic

2 Grinding: The most

3 Retropulsion: The pyloric

waves move from the fundus toward the pylorus.

vigorous peristalsis and mixing action occur close to the pylorus.

end of the stomach acts as a pump that delivers small amounts of chyme into the duodenum, simultaneously forcing most of its contained material backward into the stomach.

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Regulation of Gastric Emptying As chyme enters duodenum

Receptors respond to stretch and chemical signals Enterogastric reflex and enterogastrones inhibit gastric secretion and duodenal filling

Carbohydrate-rich chyme moves quickly through duodenum Fatty chyme remains in duodenum 6 hours or more
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Figure 23.20 Neural and hormonal factors that inhibit gastric emptying.
Presence of fatty, hypertonic, acidic chyme in duodenum

Duodenal enteroendocrine cells

Chemoreceptors and stretch receptors

Secrete

Target

Enterogastrones (secretin, cholecystokinin, vasoactive intestinal peptide)

Via short reflexes

Via long reflexes

Enteric neurons Duodenal stimuli decline

CNS centers sympathetic activity; parasympathetic activity

Contractile force and rate of stomach emptying decline

Initial stimulus Physiological response

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Stimulate Inhibit

Result

Regulation of Bile Secretion Bile secretion stimulated by

Bile salts in enterohepatic circulation Secretin from intestinal cells exposed to HCl and fatty chyme

Hepatopancreatic sphincter closed unless digestion active bile stored in gallbladder

Released to small intestine ~ only with contraction

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Regulation of Bile Secretion Gallbladder contraction stimulated by

Cholecystokinin (CCK) from intestinal cells exposed to acidic, fatty chyme Vagal stimulation (minor stimulus)

CCK also causes

Secretion of pancreatic juice Hepatopancreatic sphincter to relax

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Regulation of Pancreatic Secretion CCK induces secretion of enzyme-rich pancreatic juice by acini Secretin causes secretion of bicarbonaterich pancreatic juice by duct cells Vagal stimulation also causes release of pancreatic juice (minor stimulus)

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Figure 23.28 Mechanisms promoting secretion and release of bile and pancreatic juice.

Slide 1

1 Chyme enter -ing duodenum causes duodenal enteroendocrine cells to release cholecystokinin (CCK) and secretin. 2 CCK (red dots) and secretin (yellow dots) enter the bloodstream. 3 CCK induces secretion of enzyme-rich pancreatic juice. Secretin causes secretion of HCO3 -rich pancreatic juice.
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4 Bile salts and, to a lesser extent, secretin transported via bloodstream stimulate Liver to produce bile more rapidly. 5 CCK (via blood stream) causes gallbladder to contract and Hepatopancreatic Sphincter to relax. Bile Enters duodenum. 6 During cephalic and gastric phases, vagal Nerve stimulates gallbladder to contract weakly.
CCK secretion Secretin secretion

Digestion in the Small Intestine Chyme from stomach contains

Partially digested carbohydrates and proteins Undigested fats

36 hours in small intestine

Most water absorbed ~ All nutrients absorbed

Small intestine, like stomach, no role in ingestion or defecation

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Requirements for Digestion and Absorption in the Small Intestine Slow delivery of acidic, hypertonic chyme Delivery of bile, enzymes, and bicarbonate ions from liver and pancreas Mixing

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Motility of the Small Intestine Segmentation

Most common motion of small intestine Initiated by intrinsic pacemaker cells Mixes/moves contents toward ileocecal valve Intensity altered by long & short reflexes; hormones
Parasympathetic ; sympathetic

Wanes in late intestinal (fasting) phase

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Motility of the Small Intestine Peristalsis

Initiated by rise in hormone motilin in late intestinal phase; every 90120 minutes Each wave starts distal to previous
Migrating motor complex

Meal remnants, bacteria, and debris moved to large intestine From duodenum ileum ~ 2 hours

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Figure 23.3a Peristalsis and segmentation.

From mouth

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Peristalsis: Adjacent segments of alimentary tract organs alternately contract and relax, moving food along the tract distally.

Motility of the Small Intestine Local enteric neurons coordinate intestinal motility Cholinergic sensory neurons may activate myenteric plexus
Causes contraction of circular muscle proximally and of longitudinal muscle distally Forces chyme along tract

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Motility of the Small Intestine Ileocecal sphincter relaxes, admits chyme into large intestine when
Gastroileal reflex enhances force of segmentation in ileum Gastrin increases motility of ileum

Ileocecal valve flaps close when chyme exerts backward pressure

Prevents regurgitation into ileum

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Digestion Digestion
Catabolic; macromolecules monomers small enough for absorption

Enzymes
Intrinsic and accessory gland enzymes break down food

Hydrolysis
Water is added to break bonds

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Digestion of Carbohydrates Only monosaccharides can be absorbed Monosaccharides absorbed as ingested

Glucose, fructose, galactose

Digestive enzymes
Salivary amylase, pancreatic amylase, and brush border enzymes (dextrinase, glucoamylase, lactase, maltase, and sucrase) Break down disaccharides sucrose, lactose, maltose; polysaccharides glycogen and starch

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Digestion of Carbohydrates Starch digestion

Salivary amylase (saliva) oligosaccharides at pH 6.75 7.00 Pancreatic amylase (small intestine) breaks down any that escaped salivary amylase oligosaccharides Brush border enzymes (dextrinase, glucoamylase, lactase, maltase, sucrase) oligosaccharides monosaccharides

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Figure 23.32 Flowchart of digestion and absorption of foodstuffs. (1 of 4)

Foodstuff Starch and disaccharides

Enzyme(s) and source

Site of action Path of absorption Glucose and galactose are absorbed via cotransport with sodium ions. Fructose passes via facilitated diffusion. All monosaccharides leave the epithelial cells via facilitated diffusion, enter the capillary blood in the villi, and are transported to the liver via the hepatic portal vein.

Salivary amylase Pancreatic amylase

Mouth
Small intestine

Carbohydrate digestion

Oligosaccharides and disaccharides

Lactose

Maltose

Sucrose

Galactose

Glucose

Fructose

Brush border enzymes in small intestine (dextrinase, glucoamylase, lactase, maltase, and sucrase)

Small intestine

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Digestion of Proteins
Source is dietary, digestive enzymes, mucosal cells; digested to amino acid monomers Begins with pepsin in stomach at pH 1.5 2.5
Inactive in high pH of duodenum

Pancreatic proteases
Trypsin, chymotrypsin, and carboxypeptidase

Brush border enzymes

Aminopeptidases, carboxypeptidases, and dipeptidases

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Figure 23.33 Protein digestion and absorption in the small intestine.

Slide 1

Lumen of intestine Pancreatic proteases Na+

Amino acids of protein fragments Brush border enzymes Apical membrane (microvilli) 1 Proteins and protein fragments are digested to amino acids by pancreatic proteases (trypsin, chymotrypsin, and carboxypeptidase), and by brush border enzymes (carboxypeptidase, aminopeptidase, and dipeptidase) of mucosal cells. 2 The amino acids are then absorbed by active transport into the absorptive cells, and move to their opposite side. Amino acid carrier

Na+

Absorptive epithelial cell

Capillary
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3 The amino acids leave the villus epithelial cell by facilitated diffusion and enter the capillary via intercellular clefts.

Figure 23.33 Protein digestion and absorption in the small intestine.

Slide 2

Lumen of intestine Pancreatic proteases Na+

Amino acids of protein fragments Brush border enzymes Apical membrane (microvilli) 1 Proteins and protein fragments are digested to amino acids by pancreatic proteases (trypsin, chymotrypsin, and carboxypeptidase), and by brush border enzymes (carboxypeptidase, aminopeptidase, and dipeptidase) of mucosal cells.

Na+

Absorptive epithelial cell

Capillary
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Figure 23.33 Protein digestion and absorption in the small intestine.

Slide 3

Lumen of intestine Pancreatic proteases Na+

Amino acids of protein fragments Brush border enzymes Apical membrane (microvilli) 1 Proteins and protein fragments are digested to amino acids by pancreatic proteases (trypsin, chymotrypsin, and carboxypeptidase), and by brush border enzymes (carboxypeptidase, aminopeptidase, and dipeptidase) of mucosal cells. 2 The amino acids are then absorbed by active transport into the absorptive cells, and move to their opposite side. Amino acid carrier

Na+

Absorptive epithelial cell

Capillary
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Figure 23.33 Protein digestion and absorption in the small intestine.

Slide 4

Lumen of intestine Pancreatic proteases Na+

Amino acids of protein fragments Brush border enzymes Apical membrane (microvilli) 1 Proteins and protein fragments are digested to amino acids by pancreatic proteases (trypsin, chymotrypsin, and carboxypeptidase), and by brush border enzymes (carboxypeptidase, aminopeptidase, and dipeptidase) of mucosal cells. 2 The amino acids are then absorbed by active transport into the absorptive cells, and move to their opposite side. Amino acid carrier

Na+

Absorptive epithelial cell

Capillary
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3 The amino acids leave the villus epithelial cell by facilitated diffusion and enter the capillary via intercellular clefts.

Figure 23.32 Flowchart of digestion and absorption of foodstuffs. (2 of 4)

Foodstuff Proteins

Enzyme(s) and source

Site of action Path of absorption Amino acids are absorbed via cotransport with sodium ions. Some dipeptides and tripeptides are absorbed via cotransport with H+ and hydrolyzed to amino acids within the cells. Infrequently, transcytosis of small peptides occurs. Amino acids leave the epithelial cells by facilitated diffusion, enter the capillary blood in the villi, and are transported to the liver via the hepatic portal vein.

Large polypeptides

Pepsin (stomach glands) in presence of HCl Pancreatic enzymes (trypsin, chymotrypsin, carboxypeptidase) Brush border enzymes (aminopeptidase, carboxypeptidase, and dipeptidase)

Stomach

Protein digestion

Small intestine

Small polypeptides, small peptides

Small intestine

Amino acids (some dipeptides and tripeptides)

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Digestion of Lipids Pre-treatmentemulsification by bile salts

Does not break bonds

Enzymespancreatic lipases
Fatty acids and monoglycerides

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Figure 23.34 Emulsification, digestion, and absorption of fats.

Fat globule

Slide 1

Bile salts

1 Bile salts in the duodenum emulsify large fat globules (physically break them up into smaller fat droplets). 2 Digestion of fat by the pancreatic enzyme lipase yields free fatty acids and monoglycerides. These then associate with bile salts to form micelles which ferry them to the intestinal mucosa. Micelles made up of fatty acids, monoglycerides, and bile salts 3 Fatty acids and monoglycerides leave micelles and diffuse into epithelial cells. There they are recombined and packaged with other fatty substances and proteins to form chylomicrons. 4 Chylomicrons are extruded from the epithelial cells by exocytosis. The chylomicrons enter lacteals and are carried away from the intestine in lymph.

Fat droplets coated with bile salts

Epithelial cells of small intestine

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Lacteal

Figure 23.32 Flowchart of digestion and absorption of foodstuffs. (3 of 4)

Foodstuff Unemulsified triglycerides

Enzyme(s) and source

Site of action Path of absorption Fatty acids and monoglycerides enter the intestinal cells via diffusion. Fatty acids and monoglycerides are recombined to form triglycerides and then combined with other lipids and proteins within the cells. The resulting chylomicrons are extruded by exocytosis. The chylomicrons enter the lacteals of the villi and are transported to the systemic circulation via the lymph in the thoracic duct. Some short-chain fatty acids are absorbed, move into the capillary blood in the villi by diffusion, and are transported to the liver via the hepatic portal vein.

Lingual lipase

Mouth

Gastric lipase

Stomach

Fat digestion

Emulsification by the detergent action of bile salts ducted in from the liver Pancreatic lipases Monoglycerides (or diglycerides with gastric lipase) and fatty acids

Small intestine

Small intestine

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Digestion of Nucleic Acids Enzymes

Pancreatic ribonuclease and deoxyribonuclease nucleotide monomers Brush border enzyme nucleosidases and phosphatases free bases, pentose sugars, phosphate ions

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Figure 23.32 Flowchart of digestion and absorption of foodstuffs. (4 of 4)

Foodstuff

Enzyme(s) and source

Site of action Path of absorption Units enter intestinal cells by active transport via membrane carriers. Units are absorbed into capillary blood in the villi and transported to the liver via the hepatic portal vein.

Nucleic acids
Pancreatic ribonuclease and deoxyribonuclease Brush border enzymes (nucleosidases and phosphatases) Small intestine

Nucleic acid digestion

Small intestine

Pentose sugars, N-containing bases, phosphate ions

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Absorption ~ All food; 80% electrolytes; most water absorbed in small intestine
Most prior to ileum
Ileum reclaims bile salts

Most absorbed by active transport blood

Exception - lipids

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Absorption of Carbohydrates Glucose and galactose

Secondary active transport (cotransport) with Na+ epithelial cells Move out of epithelial cells by facilitated diffusion capillary beds in villi

Fructose
Facilitated diffusion to enter and exit cells

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Absorption of Carbohydrates Glucose and galactose

Secondary active transport (cotransport) with Na+ epithelial cells Move out of epithelial cells by facilitated diffusion capillary beds in villi

Fructose
Facilitated diffusion to enter and exit cells

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Absorption of Protein Amino acids transported by several types of carriers

Most coupled to active transport of Na+

Dipeptides and tripeptides actively absorbed by H+-dependent cotransport; digested to amino acids within epithelial cells Enter capillary blood by diffusion

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Homeostatic Imbalance Whole proteins not usually absorbed Can be taken up by endocytosis/exocytosis
Most common in newborns food allergies
Usually disappear with mucosa maturation

Allows IgA antibodies in breast milk to reach infant's bloodstream passive immunity

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Absorption of Lipids
Absorption of monoglycerides and fatty acids
Cluster with bile salts and lecithin to form micelles Released by micelles to diffuse into epithelial cells Combined with lecithin, phospholipids, cholesterol, & coated with proteins to form chylomicrons Enter lacteals; transported to systemic circulation Hydrolyzed to free fatty acids and glycerol by lipoprotein lipase of capillary endothelium
Cells can use for energy or stored fat

Absorption of short chain fatty acids

Diffuse into portal blood for distribution
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Absorption of Nucleic Acids Absorption

Active transport across epithelium bloodstream

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Absorption of Vitamins In small intestine

Fat-soluble vitamins (A, D, E, and K) carried by micelles; diffuse into absorptive cells Water-soluble vitamins (vitamin C and B vitamins) absorbed by diffusion or by passive or active transporters. Vitamin B12 (large, charged molecule) binds with intrinsic factor, and is absorbed by endocytosis

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Absorption of Vitamins In large intestine

Vitamin K and B vitamins from bacterial metabolism are absorbed

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Absorption of Electrolytes
Most ions actively along length of small intestine Iron and calcium are absorbed in duodenum Na+ coupled with active absorption of glucose and amino acids Cl transported actively K+ diffuses in response to osmotic gradients; lost if poor water absorption Usually amount in intestine is amount absorbed

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Absorption of Electrolytes Iron and calcium absorption related to need

Ionic iron stored in mucosal cells with ferritin When needed, transported in blood by transferrin

Ca2+ absorption regulated by vitamin D and parathyroid hormone (PTH)

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Absorption of Water 9 L water, most from GI tract secretions, enter small intestine
95% absorbed in the small intestine by osmosis Most of rest absorbed in large intestine

Net osmosis occurs if concentration gradient established by active transport of solutes Water uptake coupled with solute uptake
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Malabsorption of Nutrients Causes

Anything that interferes with delivery of bile or pancreatic juice Damaged intestinal mucosa (e.g., bacterial infection; some antibiotics)

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Malabsorption of Nutrients Gluten-sensitive enteropathy (celiac disease)

Immune reaction to gluten Gluten causes immune cell damage to intestinal villi and brush border Treated by eliminating gluten from diet (all grains but rice and corn)

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