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          potassium balance are maintained. Most               Intercalated cell  Plasma
          potassium  in  excreted urine reaches  the
  VetBooks.ir  tubular fluid by tubular secretion in the   Urine  H O +  CO 2     CO 2
                                                                2
          collecting duct.
            Unregulated secretion of aldosterone    H +      H +        HCO 3 –   HCO 3 –
          by adrenal tumors can cause significant
          reductions in plasma potassium concen-                Carbonic
          trations, and these can threaten life.                anhydrase
          Such individuals may also have moder-
          ate  degrees  of sodium  retention  and   Figure  23-12.  Formation and secretion of
          increases in ECF volume, but this is usu-  hydrogen and bicarbonate ions by intercalated
          ally not as severe as the changes in    cells. Formation of hydrogen and bicarbonate from
          plasma potassium. The more moderate     water and carbon dioxide in the cell is promoted by
          changes in sodium balance are because   the action of intracellular carbonic anhydrase.
          sodium transport by other nephron seg-
          ments is regulated by other factors that   Some of the secreted hydrogen ions com-
          can counterbalance the sodium‐retain-   bine with HPO  ions in the tubular fluid to
                                                                −2
          ing effects of aldosterone.                          4
                                                            −
                                                                                −2
                                                  form H PO . In this manner, HPO  acts as
                                                                                4
                                                         2
                                                            4
                                                  an intratubular buffer to reduce the concen-
          Urine Acidification                     tration of free hydrogen ions and prevent the
                                                  urine pH from dropping too low. The phos-
          Intercalated cells in the walls of collect-  phate ions originally entered the tubular fluid
          ing ducts are capable of the active trans-  as part of the glomerular filtrate. Ammonia
          port of hydrogen ions into the tubular   may also serve as an intratubular buffer
          lumen to acidify the urine. This system   (forming ammonium ions), and it is secreted
          can generate a difference between blood   into the tubular fluid by the collecting duct.
          and urine of about 3 pH units, so if blood
          pH is 7.4, urine with a pH of 4.4 can be   Regulation of Acid‐Base Balance
          formed. The hydrogen ions to be secreted
          are generated in intercalated cells by the
          hydration of carbon dioxide under the   The pH of blood plasma and other extracel-
          influence of carbonic anhydrase. A bicar-  lular fluids is maintained relatively constant
          bonate ion is also generated during this   within a narrow range (about 7.35 to 7.45).
          process, and these are secreted into the   It is important to maintain this constancy
          extracellular fluids at the base of the cell,   because enzyme activity and metabolic
          from where the bicarbonate can diffuse   processes require close control of pH for
          into the plasma (Fig. 23‐12).           optimal function. Three major mechanisms
            The secretion of hydrogen ions by     function together to prevent the develop-
          intercalated cells is regulated in part by   ment of an abnormal pH. These are: (1)
          the concentrations of carbon dioxide and   extracellular and intracellular chemical
          bicarbonate ions in the plasma and other   buffers; (2) ventilatory control of plasma
          extracellular fluids. If carbon dioxide con-  carbon dioxide; and (3) urinary excretion of
          centration increases or if bicarbonate ion   bicarbonate or an acid urine as needed.
          concentration decreases, the rate of hydro-
          gen  ion  secretion  accelerates,  and  urine   Extracellular and Intracellular Buffers
          becomes more acidic. The importance and
          relevance of  this regulation  by  carbon   A chemical buffer system acts to maintain
          dioxide or bicarbonate to the regulation of   a constant pH by either donating or remov-
          overall acid‐base balance is discussed in a   ing free hydrogen ions in a solution (see
          later section of this chapter.          Chapter 2).
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