Hoppeler H, Weibel E R
Department of Anatomy, University of Berne, Bühlstrasse 26, CH-3000 Berne 9, Switzerland.
Acta Physiol Scand. 2000 Apr;168(4):445-56. doi: 10.1046/j.1365-201x.2000.00696.x.
Environmental oxygen is transported by the respiratory cascade to the site of oxidation in active tissues. Under conditions of heavy exercise it is ultimately the working skeletal muscle cells that determine the aerobic demand as over 90% of energy is spent in muscle cells. Oxygen is transported in the circulation bound to haemoglobin of erythrocytes while substrates are transported in the plasma. The supply of oxygen must be continuous because there are only minimal oxygen stores in the body of most mammalian species while substrates are stored in significant quantities both within muscle cells as also in organismic substrate stores. The pathways for oxygen and substrates ultimately converge in muscle mitochondria. In mammals, a structural limitation of carbohydrate and lipid transfer from the microvascular system to muscle cells is reached at a moderate work intensity (i.e. at less than 50% of VO2max). At higher work rates intracellular substrate stores must be used for oxidation. It is therefore not surprising to find larger intramyocellular carbohydrate and lipid stores in 'athletic' species as well as in endurance-trained human athletes. The transfer limitations for carbohydrates and lipids presumably occur on the level of the sarcolemma. These findings imply that the design of the respiratory cascade from lungs to muscle mitochondria has to be analysed with regard to satisfying the demand for oxygen of the working muscle cells. Substrate stores are replenished at low flux rates during periods of rest and are stored intracellularly. They are therefore locally available to mitochondria for aerobic work at high intensities.
环境中的氧气通过呼吸级联被输送到活跃组织中的氧化部位。在剧烈运动的情况下,最终是工作的骨骼肌细胞决定有氧需求,因为超过90%的能量消耗在肌肉细胞中。氧气在循环中与红细胞中的血红蛋白结合运输,而底物则在血浆中运输。氧气供应必须持续不断,因为大多数哺乳动物体内的氧气储备极少,而底物则大量储存在肌肉细胞内以及机体的底物储备中。氧气和底物的运输途径最终在肌肉线粒体中汇合。在哺乳动物中,在中等工作强度(即低于最大摄氧量的50%)时,碳水化合物和脂质从微血管系统向肌肉细胞的转运就会达到结构上的限制。在更高的工作强度下,细胞内的底物储备必须用于氧化。因此,在“运动型”物种以及耐力训练的人类运动员中发现更大的细胞内碳水化合物和脂质储备也就不足为奇了。碳水化合物和脂质的转运限制可能发生在肌膜水平。这些发现意味着,必须分析从肺到肌肉线粒体的呼吸级联设计,以满足工作肌肉细胞对氧气的需求。底物储备在休息期间以低通量率补充,并储存在细胞内。因此,它们在局部可被线粒体用于高强度的有氧工作。