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Triacylglycerols Are Highly Concentrated Energy Stores - Biochemistry - NCBI Bookshelf

NCBI Book sh elf. A ser v ice of t h e Na t ion a l Libr a r y of Medicin e, Na t ion a l In st it u t es of Hea lt h .

Berg JM, Ty moczko JL, Stry er L. Biochemistry . 5th edition. New Y ork: W H Freeman; 2002.

Section 22.1 Stores

Triacylglycerols Are Highly Concentrated Energy

Triacylglycerols are highly concentrated stores of metabolic energy because they are reduced and anhydrous . The y ield from the complete ox idation of fatty acids is about 9 kcal g-1 (38 kJ g-1 ), in contrast with about 4 kcal g-1 (1 7 kJ g-1 ) for carbohy drates and proteins. The basis of this large difference in caloric y ield is that fatty acids are much more reduced. Furthermore, triacy lgly cerols are nonpolar, and so they are stored in a nearly anhy drous form, whereas much more polar proteins and carbohy drates are more highly hy drated. In fact, 1 g of dry gly cogen binds about 2 g of water. Consequently , a gram of nearly anhydrous fat stores more than six times as much energy as a gram of hydrated glycogen, which is likely the reason that triacy lgly cerols rather than gly cogen were selected in ev olution as the major energy reserv oir. Consider a ty pical 7 0-kg man, who has fuel reserv es of 1 00,000 kcal (420,000 kJ) in triacy lgly cerols, 25,000 kcal (1 00,000 kJ) in protein (mostly in muscle), 600 kcal (2500 kJ) in gly cogen, and 40 kcal (1 7 0 kJ) in glucose. Triacy lgly cerols constitute about 1 1 kg of his total body weight. If this amount of energy were stored in gly cogen, his total body weight would be 55 kg greater. The gly cogen and glucose stores prov ide enough energy to sustain biological function for about 24 hours, whereas the triacy lgly cerol stores allow surv iv al for sev eral weeks. In mammals, the major site of accumulation of triacy lgly cerols is the cy toplasm of adipose cells (fat cells). Droplets of triacy lgly cerol coalesce to form a large globule, which may occupy most of the cell v olume (see Figure 22.1 ). Adipose cells are specialized for the sy nthesis and storage of triacy lgly cerols and for their mobilization into fuel molecules that are transported to other tissues by the blood. The utility of triacy lgly cerols as an energy source is dramatically illustrated by the abilities of migratory birds, which can fly great distances without eating. Ex amples are the American golden plov er and the ruby -throated sparrow. The golden plov er flies from Alaska to the southern tip of South America; a large segment of the flight (3800 km, or 2400 miles) is ov er open ocean, where the birds cannot feed. The ruby -throated hummingbird can fly nonstop across the Gulf of Mex ico. Fatty acids prov ide the energy source for both these prodigious feats. Figure Triacy lgly cerols fuel the long migration flights of the American Golden Plov er (Pluvialis dominica ). [Gerard Fuehrer/V isuals Unlimited.]

22.1.1 Dietary Lipids Are Digested by Pancreatic Lipases

Most lipids are ingested in the form of triacy lgly cerols but must be degraded to fatty acids for absorption across the intestinal epithelium. Recall that lipids are not easily solubilized, y et they must be in order to be degraded. Triacy lgly cerols in the intestinal lumen are incorporated into micelles formed with the aid of bile salts (Figure 22.3), amphipathic molecules sy nthesized from cholesterol in the liv er and secreted from the gall bladder. Incorporation of lipids into micelles orients the ester bonds of the lipid toward the surface of the micelle, rendering the bonds more susceptible to digestion by pancreatic lipases that are in aqueous solution. If the production of bile salts is inadequate owing to liv er disease, large amounts of fats (as much as 30 g day -1 ) are ex creted in the feces. This condition is referred to as steatorrhea, from the Greek steato , fat. Figure 22.3 Gly cocholate. Bile salts, such as gly cocholate, facilitate lipid digestion in the intestine. The lipases digest the triacy lgly cerols into free fatty acids and monoacy lgly cerol (Figure 22.4). These digestion products are carried in micelles to the intestinal epithelium where they are absorbed across the plasma membrane. Figure 22.4 Action of Pancreatic Lipases. Lipases secreted by the pancreas conv ert triacy lgly cerols into fatty acids and monoacy lgly cerol for absorption into the intestine.
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Triacylglycerols Are Highly Concentrated Energy Stores - Biochemistry - NCBI Bookshelf

22.1.2 Dietary Lipids Are Transported in Chylomicrons

In the intestinal mucosal cells, the triacy lgly cerols are resy nthesized from fatty acids and monoacy lgly cerols and then packaged into lipoprotein transport particles called chylomicrons , stable particles ranging from approx imately 1 80 to 500 nm in diameter (Figure 22.5). These particles are composed mainly of triacy lgly cerols, with apoprotein B-48 as the main protein component. Protein constituents of lipoprotein particles are called apolipoproteins . Chy lomicrons also function in the transport of fat-soluble v itamins and cholesterol. Figure 22.5 Chy lomicron Formation. Free fatty acids and monoacy lgly cerols are absorbed by intestinal epithelial cells. Triacy lgly cerols are resy nthesized and packaged with other lipids and apoprotein B48 to form chy lomicrons, which are then released into the ly mph (more...) The chy lomicrons are released into the ly mph sy stem and then into the blood. These particles bind to membrane-bound lipoprotein lipases, primarily at adipose tissue and muscle, where the triacy lgly cerols are once again degraded into free fatty acids and monoacy lgly cerol for transport into the tissue. The triacy lgly cerols are then resy nthesized inside the cell and stored. In the muscle, they can be ox idized to prov ide energy , as will be discussed shortly . By agreement with the publisher, this book is accessible by the search feature, but cannot be browsed. Copy right 2002, W. H. Freeman and Company .
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