Macfarlane Craig, Adams Mark A, Hansen Lee D
School of Plant Biology (Botany), The University of Western Australia, 35 Stirling Highway, Crawley 6009, Australia.
Proc Biol Sci. 2002 Jul 22;269(1499):1499-507. doi: 10.1098/rspb.2002.2030.
The enthalpy balance model of growth uses measurements of the rates of heat and CO(2) production to quantify rates of decarboxylation, oxidative phosphorylation and net anabolism. Enthalpy conversion efficiency (eta(H)) and the net rate of conservation of enthalpy in reduced biosynthetic products (R(SG)DeltaH(B)) can be calculated from metabolic heat rate (q) and CO(2) rate (R(CO2)). eta(H) is closely related to carbon conversion efficiency and the efficiency of conservation of available electrons in biosynthetic products. R(SG)DeltaH(B) and eta(H) can be used, together with biomass composition, to describe the rate and efficiency of growth of plant tissues. q is directly related to the rate of O(2) consumption and the ratio q:R(CO2) is inversely related to the respiratory quotient. We grew seedlings of Eucalyptus globulus at 16 and 28 degrees C for four to six weeks, then measured q and R(CO2) using isothermal calorimetry. Respiratory rate at a given temperature was increased by a lower growth temperature but eta(H) was unaffected. Enthalpy conversion efficiency - and, therefore, carbon conversion efficiency - decreased with increasing temperature from 15 to 35 degrees C. The ratio of oxidative phosphorylation to oxygen consumption (P/O ratio) was inferred in vivo from eta(H) and by assuming a constant ratio of growth to maintenance respiration with changing temperature. The P/O ratio decreased from 2.1 at 10-15 degrees C to less than 0.3 at 35 degrees C, suggesting that decreased efficiency was not only due to activity of the alternative oxidase pathway. In agreement with predictions from non-equilibrium thermodynamics, growth rate was maximal near 25 degrees C, where the calculated P/O ratio was about half maximum. We propose that less efficient pathways, such as the alternative oxidase pathway, are necessary to satisfy the condition of conductance matching whilst maintaining a near constant phosphorylation potential. These conditions minimize entropy production and maximize the efficiency of mitochondrial energy conversions as growing conditions change, while maintaining adequate finite rates of energy processing.
生长的焓平衡模型利用热量和二氧化碳产生速率的测量值来量化脱羧、氧化磷酸化和净合成代谢的速率。焓转化效率(η(H))以及还原生物合成产物中焓的净守恒速率(R(SG)ΔH(B))可根据代谢热速率(q)和二氧化碳速率(R(CO2))计算得出。η(H)与碳转化效率以及生物合成产物中可用电子的守恒效率密切相关。R(SG)ΔH(B)和η(H)可与生物量组成一起用于描述植物组织的生长速率和效率。q与氧气消耗速率直接相关,q:R(CO2)的比值与呼吸商呈反比。我们将蓝桉幼苗在16℃和28℃下培养4至6周,然后使用等温量热法测量q和R(CO2)。较低的生长温度会提高给定温度下的呼吸速率,但η(H)不受影响。从15℃到35℃,焓转化效率以及因此的碳转化效率随温度升高而降低。通过η(H)并假设随温度变化生长呼吸与维持呼吸的比值恒定,可在体内推断氧化磷酸化与氧气消耗的比值(P/O比值)。P/O比值从10 - 15℃时的2.1降至35℃时小于0.3,这表明效率降低不仅是由于交替氧化酶途径的活性。与非平衡热力学的预测一致,生长速率在25℃左右达到最大值,此时计算出的P/O比值约为最大值的一半。我们认为效率较低的途径,如交替氧化酶途径,对于满足电导匹配条件同时维持接近恒定的磷酸化电位是必要的。随着生长条件的变化,这些条件可使熵产生最小化并使线粒体能量转换效率最大化,同时保持足够的有限能量处理速率。