Boutilier R G, Donohoe P H, Tattersall G J, West T G
Department of Zoology, University of Cambridge, UK.
J Exp Biol. 1997 Jan;200(Pt 2):387-400. doi: 10.1242/jeb.200.2.387.
Many amphibians encounter conditions each winter when their body temperature is so low that normal activities are suspended and the animals enter into a state of torpor. In ice-covered ponds or lakes, oxygen levels may also become limiting, thereby forcing animals to endure prolonged periods of severe hypoxia or anoxia. Certain frogs (e.g. Rana temporaria) can dramatically suppress their metabolism in anoxia but are not as tolerant as other facultative vertebrate anaerobes (e.g. turtle, goldfish) of prolonged periods of complete O2 lack. Many overwintering amphibians do, however, tolerate prolonged bouts of severe hypoxia, relying exclusively on cutaneous gas exchange. Rana temporaria overwintering for 2 months in hypoxic water (PO2 approximately 25 mmHg) at 3 degrees C progressively reduce their blood PCO2 to levels characteristic of water-breathing fish. The result is that blood pH rises and presumably facilitates transcutaneous O2 transfer by increasing Hb O2-affinity. Even after months of severe hypoxia, there is no substantial build-up of lactate as the animals continue to rely on cutaneous gas exchange to satisfy the requirements of a suppressed aerobic metabolism. Our recent experiments have shown that the skeletal muscle of frogs oxyconforms in vitro to the amount of O2 available. The cellular basis for the oxyconformation of skeletal muscle is unknown, but the hypothesis driving our continuing experiments theories that metabolic suppression at a cellular level is synonymous with suppressed ion leak across cellular membranes.
每年冬天,许多两栖动物都会遇到这样的情况:它们的体温极低,正常活动暂停,动物进入蛰伏状态。在冰封的池塘或湖泊中,氧气水平也可能成为限制因素,从而迫使动物忍受长时间的严重缺氧或无氧状态。某些青蛙(如欧洲林蛙)在无氧状态下能显著抑制其新陈代谢,但与其他兼性脊椎动物厌氧生物(如乌龟、金鱼)相比,它们对长时间完全缺氧的耐受性较差。然而,许多越冬的两栖动物确实能忍受长时间的严重缺氧,完全依靠皮肤进行气体交换。欧洲林蛙在3摄氏度的缺氧水中(氧分压约为25毫米汞柱)越冬2个月,会逐渐将其血液中的二氧化碳分压降低到水呼吸鱼类特有的水平。结果是血液pH值升高,可能通过增加血红蛋白与氧气的亲和力促进经皮氧气转移。即使经过数月的严重缺氧,动物体内也不会大量积累乳酸,因为它们继续依靠皮肤气体交换来满足被抑制的有氧代谢的需求。我们最近的实验表明,青蛙的骨骼肌在体外会根据可用氧气量进行氧顺应调节。骨骼肌氧顺应调节的细胞基础尚不清楚,但推动我们持续实验的假设认为,细胞水平的代谢抑制与细胞膜离子泄漏的抑制是同义词。