Exploring links between physiology and ecology at macro-scales: the role of respiratory metabolism in insects

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Abstract
The relationships between macro-ecological patterns and physiological investigations in insects, especially those dealing with respiratory metabolism, are assessed in an attempt to encourage the development of the interaction between macroecology and physiological ecology. First, we demonstrate that although physiological ecology has been explicitly concerned with a number of issues relating to species boundaries, many questions remain unanswered. We argue that there are essentially two ways in which the relationship between physiological tolerances and species range boundaries have been investigated. The correlational approach involves physiological inference, physiological prediction, isocline analyses and climatic matching, and has often been criticized for a lack of rigour, while the experimental approach seeks to examine experimentally the relationships between physiological variables and range edges. Second, we use the recent debate on processes underlying latitudinal patterns in body size to caution against the conflation of patterns and processes operating at intraspecific and interspecific levels, the dangers inherent in invoking single explanatory variables, and an undue focus on adaptationist (e.g. optimization) rather than nonadaptationist explanations or some combination of the two. We show that both positive and negative relationships between body size and latitude have been found at the intraspecific level and suggest that interactions between temperature-induced heterochrony, and the relationship between habitat durational stability, growing season length, and generation time can be used to explain these differences. Similar variation in documented patterns is demonstrated at the interspecific level, and the mechanisms usually proffered to explain such dines (especially the starvation/desiccation-resistance hypothesis) are discussed. Interactions between various environmental factors, such as host-plant quality, and their effects on size dines are also discussed. Third, we argue that respiratory metabolism, as a measure of ATP cost, and its spatiotemporal variation are critical to many explanations of macroecological patterns. Adaptive changes in metabolism reputedly involve both depression (stress resistance) and elevation of metabolic rate, although recent studies are increasingly calling these ideas into question. In particular, flow-through respirometry is revolutionizing results by allowing careful separation of resting (or standard) and active metabolic rates. These techniques have rarely been applied to studies of metabolic cold adaptation in insects, one of the most polemical adaptations ascribed to high-latitude and high-altitude species. We conclude by arguing that physiological investigations of species tolerances are important in the context of macroecology, especially species distributional patterns and the possible impact of climate change thereon. However, we caution that relationships between abiotic variables, species tolerances, and distributional ranges may be non-linear and subject to considerable modification by the presence of other species, and that many of the pressing questions posed by macroecology have not been addressed by insect physiologists. Nonetheless, we suggest that because an understanding of the dynamics of species distributions is of considerable importance, especially in the context of current conservation problems, insect physiological ecology has much future scope.