Page 335 - Color Atlas of Biochemistry
P. 335

326       Tissues and organs



             Functions in the acid–base balance               B. Ammonia excretion
                                                              Approximately 60 mmol of protons are ex-
             Along with the lungs, the kidneys are partic-
             ularly involved in keeping the pH value of the   creted with the urine every day. Buffering
                                                              systems in the urine catch a large proportion
             extracellular fluid constant (see p. 288). The           +
             contribution made by the kidneys particularly    of the H ions, so that the urine only becomes
                                                              weakly acidic (down to about pH 4.8).
                                      –
             involves resorbing HCO 3 and actively excret-
             ing protons.                                        An important buffer in the urine is the
                                                              hydrogen   phosphate/dihydrogen    phosphate
                                                                           2–
                                                                                    –
                                                              system (HPO 4 /H 2 PO 4 ). In addition, ammo-
             A. Proton excretion                              nia also makes a vital contribution to buffer-
                                                              ing the secreted protons.
             The renal tubule cells are capable of secreting     Since plasma concentrations of free am-
                         +
             protons (H ) from the blood into the urine       monia are low, the kidneys release NH 3 from
             against a concentration gradient, despite the    glutamine   and   other   amino    acids.  At
                            +
             fact that the H concentrationinthe urine is
             up to a thousand times higher than in the        0.5–0.7 mM, glutamine is the most important
                                                              amino acid in the plasma and is the preferred
             blood. To achieve this, carbon dioxide (CO 2 )
             is taken up from the blood and—together with     form for ammonia transport in the blood. The
             water (H 2 O) and with the help of carbonate     kidneys take up glutamine, and with the help
                                                              of glutaminase [4], initially release NH 3 from
             dehydratase (carbonic anhydrase, [1])—con-
             verted into hydrogen carbonate (“bicarbo-        the amide bond hydrolytically. From the glu-
                                                              tamate formed, a second molecule of NH 3 can
                                     +
                         –
             nate,” HCO 3 )and one H . Formally, this yields
             carbonic acid H 2 CO 3 as an intermediate, but it  be obtained by oxidative deamination with
                                                              the help of glutamate dehydrogenase [5] (see
             is not released during the reaction.             p. 178). The resulting 2-oxoglutarate is fur-
                The hydrogen carbonate formed in car-
             bonic anhydrase returns to the plasma, where     ther metabolized in the tricarboxylic acid
                                                              cycle. Several other amino acids—alanine in
             it contributes to the blood’s base reserve. The
             proton is exported into the urine by secondary   particular, as well as serine, glycine, and
                                                              aspartate—can also serve as suppliers of am-
                                                +
             active transport in antiport for Na (bottom      monia.
             right). The driving force for proton excretion,
             as in other secondary active processes, is the      Ammonia can diffuse freely into the urine
                +
             Na gradient established by the ATPase in-        through the tubule membrane, while the am-
                               +
                                                     +
                                                  +
                                  +
             volved in the Na /K exchange (“Na /K AT-         monium ions that are formed in the urine are
                                                              charged and can no longer return to the cell.
             Pase”, see p. 220). This integral membrane       Acidic urine therefore promotes ammonia ex-
             proteinonthe basalside (towards the blood)
                                         +
             of tubule cells keeps the Na concentration in    cretion, which is normally 30–50 mmol per
                                                              day. In metabolic acidosis (e. g., during fasting
             the tubule cell low, thereby maintaining Na  +
             inflow. In addition to this secondary active H +  or in diabetes mellitus), after a certain time
                                                              increased induction of glutaminase occurs in
                                                         +
             transport mechanism, there is a V-type H -       the kidneys, resulting in increased NH 3 excre-
             transporting ATPase in the distal tubule and                                    +
             collecting duct (see p. 220).                    tion. This in turn promotes H release and
                                                              thus counteracts the acidosis. By contrast,
                An important function of the secreted H   +
                                         -
             ions is to promote HCO 3 resorption (top         when the plasma pH value shifts towards
                                                              alkaline values (alkalosis), renal excretion of
             right). Hydrogen carbonate, the most impor-      ammonia is reduced.
             tant buffering base in the blood, passes into
             the primary urine quantitatively, like all ions.
                                          –
             In the primary urine, HCO 3 reacts with H    +
             ions to form water and CO 2 ,which returns
             by free diffusion to the tubule cells and from
             there into the blood. In this way, the kidneys
                                           –
             also influence the CO 2 /HCO 3 buffering bal-
             ance in the plasma.





           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
           All rights reserved. Usage subject to terms and conditions of license.
   330   331   332   333   334   335   336   337   338   339   340