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2.4.8 - Energetics of respiration

(a)  Efficiency

Respiration represents a substantial loss of carbon from a plant, and under adverse conditions can be as high as two-thirds of the carbon fixed daily in photosynthesis. Both the rate and the efficiency of respiration will therefore affect plant growth significantly. The overall process of respiration results in the release of a substantial amount of energy which may be harnessed for metabolic work. In theory, the energy released from the complete oxidation of one molecule of glucose to CO2 and H2O in respiratory reactions leads to the synthesis of 36 molecules of ATP. However, in plants, because there are alternative routes for respiration, this yield can be greatly reduced. Mechanisms for regulating respiration rates in whole plants remain unclear. Convention has it that the rate of respiration is matched to the energy demands of the cell through feed-back regulation of glycolysis and electron transport by cytosolic ATP/ADP. However, since plants have non-phosphorylating bypasses in their respiratory chain that are insensitive to ATP levels, and since PEP carboxylase and PFP might be involved in sucrose degradation, the situation in vivo is not so simple. For example, the rotenone-insensitive alternative NADH dehydrogenases requires high concentrations of NADH in the matrix before it can operate and seems to be active only when substrate is plentiful and electron flow through complex I is restricted by lack of ADP. Alternative oxidase activity also depends on carbon and ADP availability and its flux is very dependent on the degree of environmental stress of the plant. In other words, non-phosphorylating pathways act as carbon or reductant ‘overflows’ of the main respiratory pathway and will only be active in vivo when sugar levels are high and the glycolytic flux rapid, when the cytochrome chain is inhibited, or when the bypasses have been induced significantly during stress. In glycolysis, the interaction between environmental signals and key regulatory enzymes, as well as the role of PFP and its activator fructose-2,6-P2, will be important.

(b)  Allocation of respiratory energy to process physiology

One way of viewing respiratory cost for plant growth and survival is by subdividing measured respiration into two components associated with (1) growth and (2) maintenance. This distinction is somewhat arbitrary, and these categories of process physiology must not be regarded as discrete sets of biochemical events. Such energy-dependent processes are all interconnected because ATP represents a universal energy currency for both, while a common pool of substrates is drawn upon in sustaining production of that ATP. Nevertheless, cells do vary in their respiratory efficiency, while genotype × environment interactions are also evident in both generation and utilisation of products from oxidative metabolism. The benefit of a high respiration rate is that more ATP is produced, which provides vital energy for growth of new tissue and defence processes, such as antioxidant activation, metabolite transport or production of resistant protein isoforms. However, the cost of high respiration rates is that carbon is expended on respiration instead of being allocated to synthesis of new tissue, therefore limiting growth capacity. Variation in respiration rate has implications for growth and resource use efficiency in plants during drought, temperature and salinity responses of plants.