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270       Tissues and organs



             Digestive processes                              stimulate pancreatic secretion and bile re-
                                                              lease.
             Gastric juice is the product of several cell
             types. The parietal cells produce hydrochloric
             acid, chief cells release pepsinogen, and acces-  B. Zymogen activation
             sory cells form a mucin-containing mucus.        To prevent self–digestion, the pancreas re-
                                                              leases most proteolytic enzymes into the du-
                                                              odenum in an inactive form as proenzymes
             A. Formation of hydrochloric acid
                                                              (zymogens). Additional protection from the
                                                 +
                                                         –
             The secretion of hydrochloric acid (H and Cl )   effects of premature activation of pancreatic
             by the parietal cells is an active process that  proteinasesisprovidedby proteinase inhibi-
             uses up ATP and takes place against a concen-    tors in the pancreatic tissue, which inactivate
             tration gradient (in the gastric lumen, with a   active enzymes by complex formation (right).
                             +
             pH of 1, the H concentration is some 10     6       Trypsinogen plays a key role among the
             times higher than in the parietal cells, which   proenzymes released by the pancreas. In the
             have a pH of 7).                                 bowel, it is proteolytically converted into ac-
                                                 +
                The precursors of the exported H ions are     tive trypsin (see p. 176) by enteropeptidase,a
             carbon dioxide (CO 2 )and water (H 2 O). CO 2    membrane enzyme on the surface of the en-
             diffuses from the blood into the parietal cells,  terocytes. Trypsin then autocatalytically acti-
             and in a reaction catalyzed by carbonate de-     vates additional trypsinogen molecules and
             hydratase (carbonic anhydrase [2]), it reacts    the other proenzymes (left).
                                 +
             with H 2 OtoformH and hydrogen carbonate
                    –
                            +
             (HCO 3 ). The H ions are transported into the
                                              +
             gastric lumen in exchange for K by a mem-        C. Fat digestion
                                +
                             +
             brane-bound H /K -exchanging ATPase [1] (a       Dueto the “hydrophobic effect” (seep. 28),
             transport ATPase of the P type; see p. 220).     water-insoluble neutral fats in the aqueous
             The remaining hydrogen carbonate is re-          environment of the bowel lumen would ag-
             leased into the interstitium in electroneutral   gregate into drops of fat in which most of the
                                                        –
             antiport in exchange for chloride ions (Cl ),    molecules would not be accessible to pancre-
                                                    –
             andfrom thereinto the blood. TheCl ions          atic lipase. The amphipathic substances in bile
             follow the secreted protons through a chan-      (bile acids, bile salts, phospholipids) create an
             nel into the gastric lumen.                      emulsion in which they occupy the surface of
                The hydrochloric acid in gastric juice is     the droplets and thereby prevent them from
             important for digestion. It activates pepsin-    coalescing into large drops. In addition, the
             ogen to form pepsin (see below) and creates      bile salts, together with the auxiliary protein
             an optimal pH level for it to take effect. It also  colipase, mediate binding of triacylglycerol
             denatures food proteins so that they are more    lipase [1] to the emulsified fat droplets. Acti-
             easily attacked by proteinases, and it kills     vation of the lipase is triggered by a confor-
             micro-organisms.                                 mation change in the C-terminal domain of
                Regulation. HCl secretion is stimulated by    the enzyme, which uncovers the active center.
             the peptide hormone gastrin, the mediator           During passage through the intestines, the
             histamine (see p. 380), and—via the neuro-       active lipase breaks down the triacylglycerols
             transmitter acetylcholine—by the autonomous      in the interior of the droplets into free fatty
             nervous system. The peptide somatostatin         acids and amphipathic monoacylglycerols.
             and certain prostaglandins (see p. 390) have     Over time, smaller micelles develop (see
             inhibitory effects. Together with cholecysto-    p. 28), in the envelope of which monoacylgly-
             kinin, secretin, and other peptides, gastrin     cerols are present in addition to bile salts and
             belongs to the group of gastrointestinal hor-    phospholipids. Finally, the components of the
             mones (see p. 370). All of these are formed in   micelles are resorbed by the enterocytes in
             the gastrointestinal tract and mainly act in the  ways that have not yet been explained.
             vicinity of thesitewhere they areformed—            Monoacylglycerols and fatty acids are re-
             i. e.,they are paracrinehormones (seep. 372).    assembled into fats again (see p. 272), while
             While    gastrin  primarily   enhances    HCl    the bile acids return to the liver (enterohe-
             secretion, cholecystokinin and secretin mainly   patic circulation; see p. 314).


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