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1020  Unit 5  Environmental Exchange

     called countercurrent multiplication. Countercurrent refers to                The removal of sodium and chloride ions from the tubu-
     the fact that the exchange takes place between fluids moving in          lar fluid in the ascending limb raises the osmotic concentra-
     opposite directions: Tubular fluid in the descending limb flows          tion of the peritubular fluid around the thin descending limb
     toward the renal pelvis, while tubular fluid in the ascending limb       (Figure 26–13b). Recall that the thin descending limb is perme-
     flows toward the cortex. Multiplication refers to the fact that the ef-  able to water but not to solutes. As tubular fluid travels deeper
     fect of the exchange increases as movement of the fluid continues.       into the medulla within the thin descending limb, osmosis
                                                                              moves water into the peritubular fluid. Solutes remain behind.
          The two parallel limbs of the nephron loop have very dif-           As a result, the tubular fluid at the turn of the nephron loop has
     ferent permeability characteristics. The thin descending limb is         a higher osmotic concentration than it did at the start.
     permeable to water but relatively impermeable to solutes. The
     thick ascending limb is relatively impermeable to both water                  The pumping mechanism of the thick ascending limb is
     and solutes, but it contains active transport mechanisms that            highly effective. Almost two-thirds of the sodium and chloride
     pump sodium and chloride ions from the tubular fluid into the            ions that enter it are pumped out of the tubular fluid before
     peritubular fluid of the medulla.                                        that fluid reaches the DCT. In other tissues, differences in sol-
                                                                              ute concentration are quickly resolved by osmosis. However,
          A quick overview of countercurrent multiplication will              osmosis cannot take place across the wall of the thick ascending
     help you make sense of the details:                                      limb, because the epithelium there is impermeable to water. So,
                                                                              as Na+ and CI– are removed, the solute concentration in the
      	 Sodium and chloride ions are pumped out of the thick                  tubular fluid decreases. Tubular fluid arrives at the DCT with
                                                                              an osmotic concentration of only about 100 mOsm/L. This
        ascending limb and into the peritubular fluid. This dilutes           value is one-third the concentration of the peritubular fluid of
        the tubular fluid.                                                    the renal cortex.

      	 The pumping action increases the osmotic concentration in                  The rate of ion transport across the thick ascending limb
                                                                              is proportional to an ion’s concentration in tubular fluid. As
        the peritubular fluid around the thin descending limb.                a result, more sodium and chloride ions are pumped into the
                                                                              medulla at the start of the thick ascending limb, where NaCl
      	 This creates a small concentration difference between the             concentrations are highest, than near the cortex. This regional
                                                                              difference in the rate of ion transport is the basis of the concen-
        tubular fluid and peritubular fluid in the renal medulla.             tration gradient within the medulla.

      	 The concentration difference results in an osmotic flow of            The Concentration Gradient of the Medulla

        water out of the thin descending limb and into the peritu-            Normally, the maximum solute concentration of the per-
        bular fluid. As a result, the solute concentration increases          itubular fluid near the turn of the nephron loop is about
        in the thin descending limb.                                          1200 mOsm/L (Figure 26–13b). Sodium and chloride ions
                                                                              pumped out of the loop’s ascending limb make up about two-
      	 The arrival of the highly concentrated solution in the thick          thirds of that gradient (750 mOsm/L). The rest of the concen-
                                                                              tration gradient results from the presence of urea.
        ascending limb speeds up the transport of sodium and
        chloride ions into the peritubular fluid.                                  To understand how urea arrives in the medulla, let’s look
                                                                              ahead to events in the last segments of the collecting system
	26       Solute pumping at the ascending limb leads to higher sol-           (Figure 26–13c). The thick ascending limb of the nephron loop,
     ute concentrations in the descending limb, which then brings             the DCT, and the collecting ducts are all impermeable to urea.
     about increased solute pumping in the ascending limb. Notice             As water is reabsorbed, the concentration of urea gradually
     that this process is a simple positive feedback loop that multi-         increases in the tubular fluid. When the tubular fluid reaches
     plies the concentration difference between the hypotonic tubu-           the papillary duct, it typically contains urea at a concentration
     lar fluid in the ascending limb and the hypertonic peritubular           of about 450 mOsm/L. Because the papillary ducts are perme-
     fluid in the renal medulla.                                              able to urea, the urea concentration in the deepest parts of the
                                                                              medulla also averages 450 mOsm/L.
          We can now take a closer look at the mechanics of the pro-
     cess. Figure 26–13a diagrams ion transport across the epithe-            Benefits of Countercurrent Multiplication
     lium of the thick ascending limb. Active transport at the apical
     surface moves sodium, potassium, and chloride ions out of the            The countercurrent mechanism performs two functions:
     tubular fluid. The carrier is called a Na–K/2 Cl transporter,
     because each cycle of the pump carries a sodium ion, a potas-            	1.	 It efficiently reabsorbs solutes and water before the tubular
     sium ion, and two chloride ions into the tubular cell. Then                   fluid reaches the DCT and collecting system.
     cotransport carriers pump potassium and chloride ions into the
     peritubular fluid. However, potassium ions are removed from
     the peritubular fluid as the sodium–potassium exchange pump
     pumps sodium ions out of the tubular cell. The potassium ions
     then diffuse back into the lumen of the tubule through potas-
     sium leak channels. The net result is that Na+ and CI– enter the
     peritubular fluid of the renal medulla.
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