在三部曲的最后一篇文章中,在前两篇论文中开发的ATP合成的统一生物热动力学理论应用于比较生理学中的一个主要问题,生物化学,和生态学——代谢尺度是物种间体重的函数。能量代谢中的种内和种间关系有明显的区别,自从Kleiber在近一个世纪前首次提出他的老鼠对大象统治以来,从一开始就存在的困惑。结果表明,异速关系[公式:见正文]中基础/标准代谢率的总体质量指数由两部分组成,从斜率(B)的相对种内恒定性中得出的一个,另一个(b')是由质量系数的种间变化引起的,A(M)与身体大小。定量分析显示出隐藏的潜在关系,然后是种间质量系数,a(M)=P0M0.10,普值P0=3.23瓦,W来自从小鼠到牛的哺乳动物的经验数据。只有在进化的生物学背景下才能理解上述关系,并为应对规则提供了生理解释。该分析还有助于从根本上理解生物学和生态学中的异速关系中如何出现比例指数的变异性和多样性。接下来,通过考虑ATP合成η的热力学效率,可以获得对哺乳动物物种代谢缩放的分子水平理解。线粒体质子泄漏是生物系统基础代谢率的主要决定因素。通过求解生物热动力学ATP理论的数学方程获得迭代解,和关键的热力学参数,例如,耦合度q,操作P/O比,并定量评估了从大鼠到牛的哺乳动物的ATP合成η的代谢效率。从老鼠到牛的2000倍体型范围内的η增加(15%),主要是由线粒体H+泄漏率降低3倍引起的,由统一的ATP理论量化。解释基础代谢的生化和机械后果,并详细讨论了产生的各种分子含义。结果扩展到最大代谢率,并在数学上解释为一般ATP理论的极限情况。指出了剖析的局限性。总之,基于ATP合成的统一生物热动力学理论的全面定量分析被证明可以解决生物学中的一个中心问题,生理学,和生态学关于能量代谢随身体大小的缩放。
In this last article of the trilogy, the unified biothermokinetic theory of ATP synthesis developed in the previous two papers is applied to a major problem in comparative physiology, biochemistry, and ecology-that of metabolic scaling as a function of body mass across species. A clear distinction is made between intraspecific and interspecific relationships in energy metabolism, clearing up confusion that had existed from the very beginning since Kleiber first proposed his mouse-to-elephant rule almost a century ago. It is shown that the overall mass exponent of basal/standard metabolic rate in the allometric relationship [Formula: see text] is composed of two parts, one emerging from the relative intraspecific constancy of the slope (b), and the other (b\') arising from the interspecific variation of the mass coefficient, a(M) with body size. Quantitative analysis is shown to reveal the hidden underlying relationship followed by the interspecific mass coefficient, a(M)=P0M0.10, and a universal value of P0=3.23 watts, W is derived from empirical data on mammals from mouse to cattle. The above relationship is shown to be understood only within an evolutionary biological context, and provides a physiological explanation for Cope\'s rule. The analysis also helps in fundamentally understanding how variability and a diversity of scaling exponents arises in allometric relations in biology and ecology. Next, a molecular-level understanding of the scaling of metabolism across mammalian species is shown to be obtained by consideration of the thermodynamic efficiency of ATP synthesis η, taking mitochondrial proton leak as a major determinant of basal metabolic rate in biosystems. An iterative solution is obtained by solving the mathematical equations of the biothermokinetic ATP theory, and the key thermodynamic parameters, e.g. the degree of coupling q, the operative P/O ratio, and the metabolic efficiency of ATP synthesis η are quantitatively evaluated for mammals from rat to cattle. Increases in η (by ∼15%) over a 2000-fold body size range from rat to cattle, primarily arising from an ∼3-fold decrease in the mitochondrial H+ leak rate are quantified by the unified ATP theory. Biochemical and mechanistic consequences for the interpretation of basal metabolism, and the various molecular implications arising are discussed in detail. The results are extended to maximum metabolic rate, and interpreted mathematically as a limiting case of the general ATP theory. The limitations of the analysis are pointed out. In sum, a comprehensive quantitative analysis based on the unified biothermokinetic theory of ATP synthesis is shown to solve a central problem in biology, physiology, and ecology on the scaling of energy metabolism with body size.