Digestion & absorption of Lipids
المؤلف:
Peter J. Kennelly, Kathleen M. Botham, Owen P. McGuinness, Victor W. Rodwell, P. Anthony Weil
المصدر:
Harpers Illustrated Biochemistry
الجزء والصفحة:
32nd edition.p528-529
2025-12-09
73
The major lipids in the diet are triacylglycerols and, to a lesser extent, phospholipids. These are hydrophobic molecules and have to be hydrolyzed and emulsified to very small droplets (micelles, 4-6 nm in diameter) before they can be absorbed. The fat-soluble vitamins, A, D, E, and K, and a variety of other lipids (including cholesterol and carotenes) are absorbed dis solved in the lipid micelles. Absorption of carotenes and fat soluble vitamins is impaired on a very low-fat diet.
Hydrolysis of triacylglycerols is initiated by lingual and gastric lipases, which attack the sn-3 ester bond forming 1,2-diacylglycerols and free fatty acids, which act as emulsifying agents. Pancreatic lipase is secreted into the small intestine and requires a further pancreatic protein, colipase, for activity. It is specific for the primary ester links—that is, positions 1 and 3 in triacylglycerols—resulting in 2-monoacylglycerols and free fatty acids as the major end products of luminal triacylglycerol digestion. Inhibitors of pancreatic lipase are used to inhibit triacylglycerol hydrolysis in the treatment of severe obesity. Pancreatic esterase in the intestinal lumen hydrolyzes monoacylglycerols, but they are poor substrates, and only ~25% of ingested triacylglycerol is completely hydrolyzed to glycerol and fatty acids before absorption (Figure 1). Bile salts, formed in the liver and secreted in the bile, permit emulsification of the products of lipid digestion into micelles together with dietary phospholipids and cholesterol secreted in the bile (about 2 g/d) as well as dietary cholesterol (about 0.5 g/d). Micelles are less than 1 μm in diameter, and soluble, so they allow the products of digestion, including the fat-soluble vitamins, being transported through the aqueous environment of the intestinal lumen to come into close contact with the brush border of the mucosal cells, allowing uptake into the epithelium. The bile salts remain in the intestinal lumen, where most are absorbed from the ileum into the enterohepatic circulation.
Fig1. Digestion and absorption of triacylglycerols. The values given for percentage uptake may vary widely but indicate the relative importance of the three routes shown.
Within the intestinal epithelium, 1-monoacyglycerols are hydrolyzed to fatty acids and glycerol and 2-monoacylglycerols are reacylated to triacylglycerols via the monoacylglycerol pathway. Glycerol released in the intestinal lumen is absorbed into the hepatic portal vein; glycerol released within the epithelium is reutilized for triacylglycerol synthesis via the normal phosphatidic acid pathway. Long-chain fatty acids are esterified to triacylglycerol in the mucosal cells and together with the other products of lipid digestion, secreted as chylomicrons into the lymphatics, entering the bloodstream via the thoracic duct. Short- and medium chain fatty acids are mainly absorbed into the hepatic portal vein as free fatty acids.
Cholesterol is absorbed dissolved in lipid micelles and is mainly esterified in the intestinal mucosa before being incorporated into chylomicrons. Plant sterols and stanols (in which the B ring is saturated) compete with cholesterol for esterification, but are poor substrates, so that there is an increased amount of unesterified cholesterol in the mucosal cells. Unesterified cholesterol and other sterols are actively transported out of the mucosal cells into the intestinal lumen. This means that plant sterols and stanols effectively inhibit the absorption of not only dietary cholesterol, but also the larger amount that is secreted in the bile, so lowering the whole body cholesterol content, and hence the plasma cholesterol concentration.
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