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Lecture 2 DIGESTION & ABSORPTION OF FUELS

GI Tract Functions

The survival of an organism depends on its ability to transduce energy derived from food and stored as chemical energy, to electrical energy. In addition to digestion, the GI tract also acts as a transport organ and has a metabolic function where food is altered before being transported.
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GI Tract Functions

Digestion - breakdown of complex macromolecules to di- & monomeric molecules. Absorption - fuels traverse GI track to cells & tissues of the body. Fuel sources.
Carbohydrates. Lipids. Proteins.

GI tract Functions

GI track functions.
Digestion & absorption. Transport organ - fluids, electrolytes & proteins. Metabolic function - alters food before adsorption.

Clinical considerations.
Lactose intolerance. Coeliac disease.
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Digestion & Absorption of Fuels General Principles of Digestion

GI tract activities.
Lubrication and homogenization. Secretion of enzymes. Secretion of electrolytes, acids or bases. Secretion of bile acids. Transport of digested material into enterocytes (intestinal epithelial cells) and thence to blood and lymph.
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Digestion & Absorption of Fuels General Principles of Digestion

GI tract is organized by functional requirements.

Salivary glands. Stomach. Pancreas. Liver/gall bladder. Small bowel. Large bowel.
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Digestive Enzymes

Zymogens - secreted enzymes that are initially inactive.

Activated in gut lumen.
Lipases Proteases

Enzymes in gut can be activated by several mechanisms.

pH changes pepsinogen to pepsin at pH < 4. Self activation. Endopepsidases - bound to duodenal mucosa.
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Digestion & Absorption of Fuels Digestive Enzymes & Zymogens

Hydrolases = all digestive enzymes.

Hydrolyses = a breakdown process initiated by hydrolase enzyme activity. Products of hydrolases:
Proteins = amino acids, di- and tripeptides. Carbohydrates = di- and monosaccharides. Lipids different mixtures.
Fatty acids. Glycerol. Mono- and diacylglycerols.
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Carbohydrates = major source of daily energy requirements.

Types of dietary carbohydrates (plant and animal starches):
Starches polysaccharides of plant origin. Sucrose disaccharides and lactose. Glucose monosaccharides, animal derived equivalent of starches..

Recommendations = carbohydrates should be > 50% of daily average requirements.

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Carbohydrate Digestion & Absorption

Mechanisms of carbohydrate adsorption.

Both active and passive processes are involved.
For monosaccharides simple diffusion is an option but it is a very slow process.

Control of osmotic load and prevention of fluid shifts in gut.

Monomer transport is rate limiting = compensatory mechanisms exist to prevent pooling of fluid in gut. > monomers in gut = > osmolality = > water movement into the gut = < brush border disaccharidase activity = < monomers = < osmolality.
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Carbohydrate Digestion & Absorption

Carrier mechanisms for monosaccharides (glucose, fructose and galactose).

Two carrier-mediated mechanisms (specificity).
Na+- dependent co-transporter = brush border membrane bound protein binds Na+ and glucose at separate sites.
Glucose and galactose. Na+ transported down its concentration gradient. Glucose carried along against its gradient. Linked to Na+/K+- dependent ATPase = sodium removal from cell in exchange for K+ driven by ATP hydrolysis + > electrochemical gradient which drives the process..
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Carbohydrate Digestion & Absorption

Carrier mechanisms for monosaccharides (glucose, fructose and galactose).

Na+-independent facilitated diffusion.
Fructose transport. In conjunction with glucose transporter (GLUT-5).
Located on serosal side of enterocyte membrane. Moves glucose into capillaries. Fructose moves down its concentration gradient.

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Digestion & Absorption of Fuels Lipids

Introduction.
90% of dietary lipid intake = triglycerides (TAG) with remainder as:
Cholesterol. Cholesterol ester. Phospholipids. Free fatty acids (FFA).

Hydrophobic problem of fats.

Precludes water soluble enzymes. Fat globules = limited surface area.
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Digestion & Absorption of Fuels Lipids

Emulsification process lipids cannot be broken down by water-soluble enzymes.

An emulsifaction process has to take place. Digestion of fats begins in the stomach.
Heat = liquification of lipids. Peristoltic movements = lipid emulsion. Aided by salivary and gastric lipases.

Surfactants caused by > FFA = > smaller lipid particles = > surface area = lipid dispersion throughout aqueous phase of GI tract = emulsification process.
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Digestion & Absorption of Fuels Lipids

Duodenum (upper part of small intestine) = major lipid digestion area where pancreatic enzymes and bile salts act upon the lipid emulsion. Lipid emulsion from stomach present. Pancreas = > secretion of:
Pancreatic lipase. Co-lipase.

Gall bladder secretion of bile salts = > emulsification.

Bile salts = < pancreatic lipase activity which is overcome by co-lipase.
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Digestion & Absorption of Fuels Lipids

Duodenum = major lipid digestion area where pancreatic enzymes and bile salts act upon the lipid emulsion.. Co-lipase = pancreas secretion.
Binds lipase to lipid surface and activates the lipase enzyme.

Lipase.
Acts on bonds of TAGs. Product = 2-monoglycerol (2-MAG) = facilitates and > adsorption by enterocytes.

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Digestion & Absorption of Fuels Lipids

Bile salts = detergents = > gut adsorption of lipids.

Has both hydrophobic and hydrophylic structural components due to cholic acid. Helps form micelles = < in size than fat emulsion droplets = > surface area = more rapid digestion and adsorption.
Micelle size dependent on bile acid concentration.

Bile salts readsorbed in ileum and transported to liver.

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Digestion & Absorption of Fuels Lipids

Fat emulsion droplets turn into micellar structures.

Facilitate transport of lipids in an aqueous environment. Bile salt micelles can solubilize other lipids. Transfers lipid digest to brush border of enterocyte where digest is transported into intestinal epithelial cell.

Adsorption by:
Diffusion. Fatty acid binding protein.

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Digestion & Absorption of Fuels Lipids

Free fatty acid (FFA) chain length determins type of transport mechanism.
Short-chain FFAs = < 10 carbons.
Pass by diffusion to epithelial cell and then to hepatic portal blood supply.

Long-chain FFAs = > 12 carbons.

Transported by a FFA-binding protein.

Water insoluble lipids are poorly adsorbed.

Only 30-40% of cholesterol is adsorbed.
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Importance = < lipid overload in liver after meals.

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Digestion & Absorption of Fuels Proteins

Introduction.
Efficient process = little loss. Sources of protein load received by gut.
70-100 g = dietary. 35-200 g = endogenous proteins from :
Secreted enzymes. Cell turnover.

Protein hydrolysis by peptidases.

Peptidases = hydrolysis of peptide bonds. Steps.
Edopeptidases = internal peptide bonds leading from large polypeptides to small oligopeptides.. Exopeptidases = act on small oligopeptides to produce:
Amino acids. D- and tripeptides.
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Digestion & Absorption of Fuels Proteins

Protein hydrolysis.
Protein hydrolysis begins in stomach.
pH = 1-2 by secreted HCl = protein denaturation = > protein surface area = protein unfolding. Pepsinogen A and B inactive > activation by:
Autoactivation by pH change < 5. By active pepsin. Products = > large peptide fragments and amino acids.

Increased protein digests = > cholecystokinin (hormone) release in duodenum = > secretion of pancreatic digestive enzymes.
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Digestion & Absorption of Fuels Proteins

Protein hydrolysis.
Pancreatic enzymes released as zymogens into duodenum.
Trypsinogen trypsin.
Activated by duodenal enteropeptidase. Regulated in pancreas by inhibitory peptide.

Trypsin.
Activates its zymogen. Activates all other pancreatic zymogens. Chymotrypsin. Elastase. Carboxy peptides A and B.
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Digestion & Absorption of Fuels Proteins

Protein hydrolysis.
Pancreatic proteases released as zymogens into duodenum.
Substrate specificity of peptide bond cleavage.
Trypsin cleaves at lysine and argenine. Chymotrypsin cleaves aromatic amino acids. Elastase cleaves small hydrophobic amino acids = 2-8 amino acid residues.

Pancreatic sodium bicarbonate (NaHCO3).

Neutralizes acid contents from stomach. > alkaline conditions = creating optimal pancreatic protease activity.

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Digestion & Absorption of Fuels Proteins

Protein hydrolysis.
Transport of free amino acids to portal blood system.
Small intestine enzymes complete the digestion of proteins.
Endopeptidases. Dipeptidases. Aminopeptidases.

Products.
Free amino acids. Di- and tripeptides. Adsorbed into enterocyte by specific carrier-mediated transport. Further digestion to amino acids inside enterocytes. Free amino acids inside enterocyte transfered across the contraluminal membrane into the portal blood system.
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