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272       Tissues and organs



             Resorption                                       acids. Individual amino acid groups have
                                                              group–specific amino acid transporters, some
             Enzymatic hydrolysis in the digestive tract      of which transport the amino acids into the
                                                                                                +
             breaks down foodstuffs into their resorbable     enterocytes in cotransport with Na ions (sec-
             components. Resorption of the cleavage prod-     ondary active transport), while others trans-
                                                                                   +
             ucts takes place primarily in the small intes-   port them in an Na –independent manner
             tine. Only ethanol and short–chain fatty acids   through facilitated diffusion. Small peptides
             are already resorbed to some extent in the       can also be taken up.
             stomach.
                The resorption process is facilitated by the
             large inner surface of the intestine, with its   B. Lipids
             brush–border cells. Lipophilic molecules pen-    Fats and other lipids are poorly soluble in
             etrate the plasma membrane of the mucosal        water. The larger the accessible surface
             cells by simple diffusion, whereas polar mol-    is—i. e., the better the fat is emulsified—the
             ecules require transporters (facilitated diffu-  easier it is for enzymes to hydrolyze it (see
             sion; see p. 218). In many cases, carrier-medi-  p. 270). Due to the special properties of milk,
                                        +
             ated cotransport with Na ions can be ob-         milk fats already reach the gastrointestinal
             served. In this case, the difference in the con-  tract in emulsified form. Digestion of them
             centration of the sodium ions (high in the       therefore already starts in the oral cavity
             intestinal lumen and low in the mucosal cells)   and stomach, where lipases in the saliva and
             drives the import of nutrients against a con-    gastric juice are available. Lipids that are less
             centration gradient (secondary active trans-     accessible—e. g., from roast pork—are emulsi-
             port; see p. 220). Failure of carrier systems in  fied in the small intestine by bile salts and bile
             the gastrointestinal tract can result in dis-    phospholipids. Only then are they capable of
             eases.                                           being attacked by pancreatic lipase [4] (see
                                                              p. 270).
                                                                 Fats (triacylglycerols) are mainly attacked
             A. Monosaccharides
                                                              by pancreatic lipase at positions 1 and 3 of the
             The cleavage of polymeric carbohydrates by       glycerol moiety. Cleavage of two fatty acid
             a–amylase [1] leads to oligosaccharides,         residues gives rise to fatty acids and 2-mono-
             which are broken down further by exoglyco-       acylglycerols, which are quantitatively the
             sidases (oligosaccharidases and disacchari-      most important products. However, a certain
             dases [2]) on the membrane surface of the        amount of glycerol is also formed by complete
             brush border. The monosaccharides released       hydrolysis. These cleavage products are re-
             in this way then pass with the help of various   sorbed by a non-ATP-dependent process
             sugar–specific transporters into the cells of    that has not yet been explained in detail.
             intestinal   epithelium.   Secondary    active      In the mucosal cells, long-chain fatty acids
             transport serves for the uptake of glucose       are resynthesized by an ATP-dependent ligase
             and galactose, which are transported against     [5] to form acyl-CoA and then triacylglycerols
             a concentration gradient in cotransport with     (fats; see p. 170). The fats are released into the
                          +
                +
             Na .The Na gradient is maintained on the         lymph in the form of chylomicrons (see
                                           +
                                              +
             basal side of the cells by Na /K -ATPase [3].    p. 278) and, bypassing the liver, are deposited
             Another passive transporter then releases        in thethoracic duct—i. e., theblood system.
             glucose   and   galactose   into  the  blood.    Cholesterol also follows this route.
             Fructose is taken up by a special type of trans-    By contrast, short-chain fatty acids (with
             porter using facilitated diffusion.              chain lengthsof lessthan 12Catoms) pass
                                                              directly into the blood and reach the liver via
                                                              the portal vein. Resorbed glycerol can also
             Amino acids (not illustrated)
                                                              take this path.
             Protein degradation is initiated by proteina-
             ses—bypepsins in thestomach andbytrypsin,
             chymotrypsin, and elastase in the small intes-
             tine. The resulting peptides are then further
             hydrolyzed by various peptidases into amino


           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
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