Nikinmaa M
Department of Biology, University of Turku, Finland.
J Exp Biol. 1997 Jan;200(Pt 2):369-80. doi: 10.1242/jeb.200.2.369.
Two major strategies are apparent for the regulation of gas transport by vertebrate blood except in the myxinoids, which seem to have little scope for such regulation. In lampreys and teleost fish, haemoglobins have low buffering capacities and large Bohr/Haldane effects. Na+/H+ exchange plays an important role in the control of haemoglobin oxygen-affinity in these vertebrate groups. The large Bohr/Haldane effect also facilitates carbon dioxide transport: the blood (or erythrocyte) pH increases upon deoxygenation, thus increasing the concentration of bicarbonate formed at a given carbon dioxide tension. In lampreys, the bicarbonate permeability of the erythrocyte membrane is low. As a consequence, extracellular acid loads cannot be buffered by haemoglobin. In contrast, teleost erythrocytes possess a functional anion exchange, allowing extracellular proton loads to be buffered by haemoglobin. However, because the buffering capacity of teleost haemoglobins is low, buffering of extracellular acid loads is less effective in teleost fish than in elasmobranch fish and in air-breathing vertebrates whose haemoglobins have a high buffering capacity. However, the high buffering capacity of the haemoglobins diminishes the possibility of regulating haemoglobin oxygen-affinity via secondarily active Na+/H+ exchange, because intracellular pH changes, caused by proton efflux, remain small.
除了盲鳗类似乎几乎没有这种调节的空间外,脊椎动物血液对气体运输的调节有两种主要策略。在七鳃鳗和硬骨鱼中,血红蛋白具有低缓冲能力和大的玻尔/哈代效应。钠/氢交换在控制这些脊椎动物群体中血红蛋白的氧亲和力方面起着重要作用。大的玻尔/哈代效应也有利于二氧化碳的运输:血液(或红细胞)在脱氧时pH值升高,从而在给定的二氧化碳张力下增加碳酸氢盐的浓度。在七鳃鳗中,红细胞膜的碳酸氢盐通透性较低。因此,细胞外酸负荷不能被血红蛋白缓冲。相比之下,硬骨鱼红细胞具有功能性阴离子交换,使细胞外质子负荷能够被血红蛋白缓冲。然而,由于硬骨鱼血红蛋白的缓冲能力较低,与血红蛋白具有高缓冲能力的板鳃亚纲鱼类和空气呼吸脊椎动物相比,硬骨鱼对细胞外酸负荷的缓冲效果较差。然而,血红蛋白的高缓冲能力降低了通过继发性主动钠/氢交换调节血红蛋白氧亲和力的可能性,因为质子外流引起的细胞内pH变化仍然很小。