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888  Unit 5  Environmental Exchange

     the molecules release their oxygen reserves more readily,         Hemoglobin and Temperature
     so the slope of the hemoglobin saturation curve changes
     (Figure 23–20a). In other words, as pH drops, the saturation      At a given PCO2, hemoglobin releases additional oxygen if the
     declines. At a tissue PO2 of 40 mm Hg, for example, you can see   temperature increases. Changes in temperature affect the slope
     in Figure 23–20a that a pH decrease from 7.4 to 7.2 changes       of the hemoglobin saturation curve (Figure 23–20b). As the
     hemoglobin saturation from 75 percent to 60 percent. This         temperature increases, hemoglobin releases more oxygen. As the
     means that hemoglobin molecules release 20 percent more           temperature decreases, hemoglobin holds oxygen more tightly.
     oxygen in peripheral tissues at a pH of 7.2 than they do at       Temperature effects are significant only in active tissues that are
     a pH of 7.4. This effect of pH on the hemoglobin saturation       generating large amounts of heat. For example, active skeletal
     curve is called the Bohr effect.                                  muscles generate heat, and the heat warms blood that flows
                                                                       through these organs. As the blood warms, the Hb molecules re-
          Carbon dioxide is the primary compound responsible for       lease more oxygen than can be used by the active muscle fibers.
     the Bohr effect. When CO2 diffuses into the blood, it rapidly
     diffuses into red blood cells. There, an enzyme called carbonic   Hemoglobin and BPG
     anhydrase catalyzes the reaction of CO2 with water molecules:
                                                                       Red blood cells lack mitochondria. These cells produce ATP
     CO2  +          carbonic anhydrase  ÷  H+  +  HCO3−               only by glycolysis. As a result, lactic acid is formed, as we saw
                                                                       in Chapter 10. p. 344 The metabolic pathways involved in
             H2O ÷ H2CO3                                               glycolysis in an RBC also generate the compound 2,3-bisphos-
                                                                       phoglycerate (biz-fos-fo. -GLIS-er-a.t), or BPG. This compound
          The product of this enzymatic reaction, H2CO3, is called     has a direct effect on oxygen binding and release. For any partial
     carbonic acid, because it dissociates into a hydrogen ion (H1)    pressure of oxygen, the higher the concentration of BPG, the
     and a bicarbonate ion (HCO3−). The rate of carbonic acid for-     greater the release of oxygen by Hb molecules. Normal RBCs
     mation depends on the amount of carbon dioxide in solution,       always contain BPG.

     which, as noted earlier, depends on the PCO2. When the PCO2            Both BPG synthesis and the Bohr effect improve oxygen deliv-
     rises, the rate of carbonic acid formation accelerates and the    ery when the pH changes: BPG levels rise when the pH increases,
                                                                       and the Bohr effect appears when the pH decreases. The concentra-
     reaction proceeds from left to right. The hydrogen ions that are  tion of BPG can be increased by high blood pH, thyroid hormones,
                                                                       growth hormone, epinephrine, and androgens. When plasma PO2
     generated diffuse out of the RBCs, and the pH of the plasma

     drops. When the PCO2 declines, hydrogen ions diffuse out of
     the plasma and into the RBCs. As a result, the pH of the plasma

     increases as the reaction proceeds from right to left.

     Figure 23–20  The Effects of pH and Temperature on Hemoglobin Saturation.                                                               10°C  20°C

                  100 100

     80                                                                                                                                  80              38°C
     60                                                                                                                                  60              43°C

             7.6 7.4
     Oxyhemoglobin (% saturation)7.2
                                                                                                           Oxyhemoglobin (% saturation)
	23

     40 40

                      Normal blood pH range                                                                                                        Normal blood temperature

     20                                   7.35–7.45                                                                                      20        38°C

            0 20 40 60 80 100                                                 0 20 40 60 80 100

                                           PO2 (mm Hg)                                                       PO2 (mm Hg)

     a Effect of pH. When the pH decreases below normal levels,        b Effect of temperature. When the temperature increases,
        more oxygen is released; the oxygen–hemoglobin                    more oxygen is released; the oxygen–hemoglobin
        saturation curve shifts to the right. When the pH increases,      saturation curve shifts to the right.
        less oxygen is released; the curve shifts to the left.
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