Digestion and Absorption: Carbohydrate, Lipid, Protein Flashcards
Amylose
- polysaccharide
- plant CHO
- storage linear chain with alpha 1,4 glycosidic bonds
Amylopectin
- polysaccharide
- amylose + branches with alpha 1,6 glycosidic bonds
Glycogen
- polysaccharide
- animal CHO storage
- similar to amylopectin except more branches
Cellulose
- polysaccharide
- linear chain with beta 1, 4 glycosidic bonds
Oligosaccharides
- 4-10 (avergage 8) glucose units
- alpha limit dextrin = branced (1 or 2 alpha 1,6 bonds)
Early Digestion of Carbohydrates
- Initial hydrolysis begins with salivary (or lingual) alpha amylase
- The saliva also contains mucin, a glycoprotein that serves as a lubricant and helps disperse polysaccharides. Because this initial hydrolysis is quantitatively small, the predominant hydrolysis is catalyzed by pancreatic alpha-amylase, which is secreted in large excess, into the small intestine, relative to starch intake.
Disaccharides and Moosaccharides
- Monosaccharides are carbohydrates containing at least three carbon atoms. The number of carbons is indicated by the prefix for the sugar such that hexoses contain six carbons and pentoses contain five carbons.
- glucose
- fructose
- sucrose
- lactose
- The state of the oxygen on the carbon in the 1-position determines whether a sugar can react with oxidized compounds (e.g., copper or iron).
-If the oxygen is not attached to some other structure, that sugar is a reducing sugar since the hydroxyl group (OH) on that carbon can donate electrons to reduce copper or iron. (lactose)
•Sucrose, a disaccharide of glucose and fructose, is non-reducing since the first carbon of the two sugar residues are joined leaving no free hydroxyl group to act as a reducing agent.
Dietary Carbohydrates
- Dietary carbohydrates provide a major component of the daily caloric requirements accounting for >50% of calories in a typical US diet. Their complete oxidation to CO2 and H2O yields 4 cal/g. When there are insufficient amounts of carbohydrates in the diet, such as on a high protein diet, glucose is produced endogenously from amino acids or even galactose or fructose.
- Dietary polysaccharides are consumed mostly as starch, the storage form for carbohydrates in plants. Starch consists of either only linear chains of glucose molecules linked by alpha-1,4-glycosidic bonds (amylose) or of linear chains with occasional branch points created by the formation of alpha-1,6-glycosidic bonds (amylopectin).
- Glycogen, the principal animal polysaccharide, is similar in structure to amylopectin except that it has many more branch points. Glycogen, itself, is a minor dietary source of carbohydrates.
- Polysaccharides are hydrolyzed to mono-, di- and oligo saccharides by glucosidases, which are a special group of enzymes that hydrolyze glycosidic bonds. Cellulose, nondigestible carbohydrate (fiber), contains alpha-1,4-glycosidic bonds that cannot be hydrolyzed in mammalian intestine.

•Since carbohydrates are absorbed as monosaccharides, the disaccharides (and oligosaccharides) must be processed further by glycosidase complexes that lie on the brush border surface of intestinal epithelial cells

Polysaccharides
•Polysaccharides are polymers of monosaccharides held together by glycosidic bonds.
- Sucrose (table sugar) consists of one molecule each of glucose and fructose.
- Lactose (milk sugar) contains instead one molecule each of galactose and glucose. Maltose and trehalose each contain two molecules of glucose.
Amylases
- Amylases, principally endosaccharidases, mostly cleave in the middle of the molecule, as opposed to exosaccharidases that remove terminal glucosyl units.
- Amylases are specific for alpha- 1,4-bonds in the linear chains of the molecule and do not attack branch points.
- The primary cleavage products from the action of amylase include: glucose, when only the terminal sugar is removed; maltose, a disaccharide of glucose; maltotriose, a trisaccharide of glucose; and oligosaccharides that on average contain eight glucosyl units. Oligosaccharides that contain one or two branches are alpha-limit dextrins.



Absorption of Monosaccharides
- The major monosaccharides resulting from carbohydrate digestion are glucose, galactose and fructose.
- Absorption of these and other minor monosaccharides are carrier-mediated processes that exhibit substrate specificity, stereospecificity and saturation kinetics.
- At least two types of monosaccharide transporters move monosaccharides from the intestinal lumen into the epithelial cell.
GLUT - 5
- A Na+ -independent, facilitated diffusion type of monosaccharide transporter (GLUT-5) facilitates absorption primarily of fructose, but also some glucose.
- Xylose is also absorbed by GLUT-5. Because xylose is not metabolized in the body, its appearance in the blood serves as an indicator of successful monosaccharide absorption.
- Hexose sugars enter via the Na+ - independent transporter by virtue of the carbohydrate concentration gradient.
SGLT - 1
- A Na+ -cotransporter (SGLT- 1) has high specificity for glucose and galactose and promotes “active” sugar absorption.
- The driving force for the Na+ -dependent transport is derived from the maintenance of low intracellular concentrations of Na+ by the action of the Na+ ,K+ -ATPase. Hydrolysis of ATP provides energy to export three Na+ ions in exchange for two K+ ions.
- The high gradient of Na+ between the intestinal lumen and the cytoplasm provides the driving force for active carbohydrate transport.
GLUT - 2
- Intracellular carbohydrate concentrations are kept low by transport out of the cytoplasm to capillaries via Na+ -independent GLUT-2 transporter in the contraluminal plasma membrane.
- These monosaccharides then travel via the portal system to the liver where the bulk of these carbohydrates are metabolized.
- Some glucose also continues to other tissues for energy metabolism.
Dietary Lipids
•On average, fat (lipid) comprises 37% of calories in the American diet.
Fat provides 9 cal/gm.
Dietary lipids are primarily (90%) triacylglycerols (TAGs; also referred to as triglycerides).
•The remainder includes cholesterol esters, phospholipids, essential unsaturated fatty acids and fat-soluble vitamins (A, D, E, K).
Digestion of Dietary Lipids
- Normally, essentially all (98%) of the dietary fat is absorbed, and most is transported to adipose for storage.
- The poor water solubility of lipids presents problems for digestion because lipids are not easily accessible to the digestive enzymes in the aqueous phase, and lipolytic products tend to aggregate into larger complexes that make poor contact with the cell surface. This latter problem is overcome by “solubilization” of lipid products with amphipathic (i.e., containing both hydrophobic and hydrophilic portions) bile acids.
- Aside from the solubility aspects, the general principle of dietary lipid assimilation is that of hydrolyzing large non-absorbable molecules into smaller units.
Six Steps of Lipid Digestion and Absorption
- Minor Digestion
- Major Digestion
- Bile Acid
- Passive Absorption
- Reesterification
- Assembly and Export
Step 1: Minor Digestion
- TAGS in mouth and stomach by lingual (acid-stable) lipase
- triggers release of CCK in duodenum
- Digestion of lipids continues in the stomach catalyzed by an acid-stable gastric lipase, which is released from the gastric mucosa. Generally the rate of hydrolysis is slow because of solubility problems.
- However, some lipase can convert TAGs into fatty acids and DAGs at the water-lipid interface.
- The importance of this initial hydrolysis is that a fraction of the water-immiscible TAGs is converted to amphipathic products that cause dispersion of the lipid phase into smaller droplets (emulsification).
- This process provides more sites for association of enzyme molecules, both lingual/gastric lipase in the stomach, and eventually pancreatic lipase in the intestinal lumen.
Step 2: Major Digestion
- all lipids in lumen of duodenum/jejunum; pancreatic lipolytic enzymes
- The entry of acid chyme and free fatty acids into the duodenal lumen stimulates endocrine cells to release cholecystokinin (CCK) into the bloodstream.
- CCK binds to its G-protein linked receptor on the gall bladder (CCK-A) causing contraction and hence release of bile salts into the intestinal lumen.
- CCK binding in human pancreas to G-protein linked CCK-B receptors causes secretion into the intestinal lumen of digestive (lipolytic) enzymes (e.g., trypsinogen).
- CCK also elicits the release of enteropeptidase into the lumen, where it activates trypsin from its zymogen form, trypsinogen.
- Other endocrine cells release secretin into the circulation. This hormone causes the pancreas to secrete bicarbonate-rich fluid that neutralizes the gut lumen pH for optimal enzymatic activity.
- These events are similar to that described for the effect of amino acids during protein digestion.
- The major enzyme for TAG hydrolysis is pancreatic lipase.
Step 3: Bile Acid
- Bile acid facilitated formation of mixed micelles; present lipolytic products to mucosal surface, followed by enterohepatic bile acid recycling
- Bile Acids —> Bile Salts
- 75% end up recycled
- Bile acids can be considered as “biological detergents” that are synthesized exclusively in the liver from cholesterol, stored in the gallbladder as bile salts, and secreted into the duodenum to form micelles with the products of lipid digestion.
- Individuals who have bile duct obstruction absorb dietary lipids poorly, and instead eliminate them in the feces (steatorrhea).
- Shortly after a meal, bile salts/acids from the gallbladder and liver are released into the lumen of the upper small intestine.
- Bile acids act in the absorption of lipids by reversibly forming micelles, equilibrium structures with well-defined sizes that are much smaller than emulsion droplets. The arrangement of amphipathic bile acids in micelles is such that the hydrophobic portions (hydrocarbon rings) are removed from contact with water, while hydrophilic groups (hydroxyls and carboxylate) remain exposed to the water.
- Bile acids form micelles with other lipids, such as 2-monoacylglycerol, phospholipids, fatty acids, cholesterol and fat-soluble vitamins.
- These mixed micelles have disk-like shapes, wherein the phospholipids and fatty acids form a bilayer and the bile acids occupy the edge positions, rendering the edge of the disk hydrophilic.
Step 4: Passive Absorption
- lipolytic products from mixed micelle into intestinal epithelial cell
- Uptake of lipids by the epithelial cells occurs in two ways: by passive diffusion through the apical membrane of duodenal and jejunal enterocytes of fatty acids and monoacylglycerols following micelle breakdown and by transport protein-mediated import of lipid.
- Absorption is virtually complete for fatty acids and 2-monoacylglycerols, which are slightly water-soluble. It is less efficient for water-insoluble lipids; for example, only 30% of dietary cholesterol is absorbed.
- After absorption of lipid digestion products, the micelles remain behind to solubilize other lipid products, thus acting as a type of “shuttle.”
- Bile acids are not absorbed at this point but instead travel the length of the small intestine and are absorbed in the terminal ileum by an ATP requiring active transport process.
Step 5: Reesterification
- : 2-monoacylglycerol, lysolecithin, cholesterol with FFA inside intestinal enterocyte
- Within the intestinal cells, the fate of absorbed fatty acids depends on chain length.
- Fatty acids of medium chain length (6-12 carbon atoms) pass through the cell into the portal blood without modification.
- Long-chain fatty acids (>12 carbon atoms) become bound to a cytoplasmic intestinal fatty acid-binding protein (FABP) and are transported to the smooth endoplasmic reticulum where they are re-esterified into TAGs.
- Glycerol for this process is derived from the absorbed 2-monoacylglycerols.
- Cholesterol exists in the diet both as free cholesterol and as cholesterol ester.
- Prior to its absorption, cholesterol ester is hydrolyzed to free cholesterol by pancreatic esterase, and micellar solubilization is an obligatory step for its absorption.
- Before packaging into chylomicrons, cholesterol is re-esterified with fatty acid in the intestinal epithelial cell by acylCoA:cholesterol acyl transferase (ACAT).
•Lysophospholipids are also re-acylated with fatty acyl CoA to phospholipids, which form the surface of chylomicrons, a lipoprotein that transports in the circulation most lipids derived from the diet.

