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322       Tissues and organs



             Kidney: functions                                transported back into the blood by resorption,
                                                              to prevent losses of valuable metabolites and
             A. Functions of the kidneys                      electrolytes. In the proximal tubule, organic
                                                              metabolites (e. g., glucose and other sugars,
             The kidneys’ main function is excretion of
             water and water-soluble substances (1). This     amino acids, lactate, and ketone bodies) are
             is closely associated with their role in regulat-  recovered by secondary active transport (see
             ing the body’s electrolyte and acid–base bal-    p. 220). There are several group-specific
             ance (homeostasis, 2; see pp. 326 and 328).      transport systems for resorbing amino acids,
             Both excretion and homeostasis are subject to    with which hereditary diseases can be
             hormonal control. The kidneys are also in-       associated (e. g., cystinuria, glycinuria, and
                                                                                        –
                                                                                              +
             volved in synthesizing several hormones (3;      Hartnup’s disease). HCO 3 ,Na ,phophate,
             see p. 315). Finally, the kidneys also play a role  and sulfate are also resorbed by ATP-depend-
             in the intermediary metabolism (4), particu-     ent (active) mechanisms in the proximal tu-
             larly in amino acid degradation and gluconeo-    bule. The later sections of the nephron mainly
             genesis (see p. 154).                            serve for additional water recovery and regu-
                                                                                            –
                                                                                    +
                The kidneys are extremely well-perfused       lated resorption of Na and Cl (see pp. 326,
             organs, with about 1500 L of blood flowing       328). These processes are controlled by hor-
             through them every day. Approximately 180 L      mones (aldosterone, vasopressin).
             of primary urine is filtered out of this. Re-       Secretion. Some excretable substances are
             moval of water leads to extreme concentra-       released into the urine by active transport in
                                                                                                          +
             tion of the primary urine (to approximately      the renal tubules. These substances include H
                                                                    +
             one-hundredth of the initial volume). As a       and K ions, urea, and creatinine, as well as
                                                              drugssuch aspenicillin.
             result, only a volume of 0.5–2.0 L of final
             urine is excreted per day.                          Clearance. Renalclearanceisusedasaquan-
                                                              titative measure of renal function. It is defined
                                                              as the plasma volume cleared of a given sub-
             B. Urine formation
                                                              stance per unit of time. Inulin, a fructose poly-
             The functional unit of the kidney is the neph-   saccharide with a mass of ca. 6 kDa (see p. 40)
             ron. It is made up of the Malpighian bodies or   that is neither actively excreted nor resorbed
             renal corpuscles (consisting of Bowman’s cap-    butisfreely filtered, hasa clearanceof
             sules and the glomerulus), the proximal tu-      120mL  min   –1  in healthy individuals.
             bule, Henle’s loop, and the distal tubule,
             which passes into a collecting duct. The hu-     Further information
             man kidney contains around one million
             nephrons. The nephrons form urine in the         Concentrating urine and transporting it
             following three phases.                          through membranes are processes that re-
                Ultrafiltration. Ultrafiltration of the blood  quire large amounts of energy. The kidneys
             plasma in the glomerulus gives rise to primary   therefore have very high energy demands. In
             urine, which is isotonicwithplasma.The pores     the proximal tubule, the ATP needed is ob-
             in the glomerular basal membrane, which are      tained from oxidative metabolism of fatty
             made up of type IV collagen (see p. 344), have   acids, ketone bodies, and several amino acids.
             an effective mean diameter of 2.9 nm. This al-   To a lesser extent, lactate, glycerol, and citric
             lows all plasma components with a molecular      acid arealso used. In thedistaltubuleand
             mass of up to about 15 kDa to pass through       Henle’s loop, glucose is the main substrate
             unhindered. At increasing masses, molecules      for the energy metabolism. The endothelial
             are progressively held back; at masses greater   cells in the proximal tubule are also capable
             than 65 kDa, they are completely unable to       of gluconeogenesis. The substrates for this are
             enter the primary urine. This applies to almost  mainly the carbohydrate skeletons of amino
             all plasma proteins—which in addition, being     acids. Their amino groups are used as ammo-
             anions, are repelled by the negative charge in   nia for buffering urine (see p. 311). Enzymes
             the basal membrane.                              for peptide degradation and the amino acid
                Resorption.   All  low-molecular    weight    metabolism occur in the kidneys at high lev-
             plasma components enter the primary urine        els of activity (e. g., amino acid oxidases,
             via glomerular filtration. Most of these are     amine oxidases, glutaminase).


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