The reasons why metabolic rate (B) scales allometrically with body mass (M) remain hotly debated. The field is dominated by correlational analyses of the relationship between B and M; these struggle to disentangle competing explanations because both B and M are confounded with ontogeny, life history, and ecology. Here, we overcome these problems by using an experimental approach to test among competing metabolic theories. We examined the scaling of B in size-manipulated and intact colonies of a bryozoan and show that B scales with M(0.5). To explain this, we apply a general model based on the dynamic energy budget theory for metabolic organization that predicts B on the basis of energy allocation to assimilation, maintenance, growth, and maturation. Uniquely, this model predicts the absolute value of B, emphasizes that there is no single scaling exponent of B, and demonstrates that a single model can explain the variation in B seen in nature.

Metabolic rate and respiratory gas exchange patterns vary significantly both between and within species, even after a number of biotic and abiotic factors are taken into account. This suggests that such variation is of evolutionary importance, but the life history implications of this variation remain relatively poorly characterized. In the present study, we examine the effect of metabolic variation on starvation and desiccation resistance in the speckled cockroach Nauphoeta cinerea. We also compare the starvation and desiccation resistance of individuals that exchange respiratory gases continuously with those that breathe discontinuously.We show that metabolic rate has no effect on survival during food and water restriction, but cockroaches exhibiting discontinuous gas exchange cycles (DGCs) live longer than those that do not and those provisioned with water lived longer than those that were not. This finding represents the first demonstration that DGCs confer a fitness benefit, and supports the oldest hypothesis for the evolution of DGCs (which suggests that DGCs arose or are maintained to reduce respiratory water loss) as we also reveal reduced water loss (both respiratory and total) in cockroaches exhibiting discontinuous gas exchange. K E Y W O R D S :DGC, discontinuous, fitness, gas, metabolism, Nauphoeta cinerea.The rate at which organisms use energy for anabolic and catabolic reactions (metabolic rate) is known to vary widely both between and within species. Metabolic rate scales positively with body mass, and variation in mass accounts for the majority (>90%) of interspecific variation (Mueller and Diamond 2001;White et al. 2006;Chown et al. 2007). However, similarly sized species of ectotherm can show metabolic rates that differ by over an order of magnitude (White et al. 2006) and individuals of the same species can differ by up to a several fold (Speakman et al. 2004). It is now well known that metabolism is influenced by many biotic and abiotic factors such as age, sex, activity, thermoregulatory strategy, reproductive, and absorptive states as well as temperature and aridity (Mueller and Diamond 2001; Addo-Bediako et al.

SUMMARYThe reasons why many insects breathe discontinuously at rest are poorly understood and hotly debated. Three adaptive hypotheses attempt to explain the significance of these discontinuous gas exchange cycles (DGCs), whether it be to save water, to facilitate gas exchange in underground environments or to limit oxidative damage. Comparative studies favour the water saving hypothesis and mechanistic studies are equivocal but no study has examined the acclimation responses of adult insects chronically exposed to a range of respiratory environments. The present research is the first manipulative study of such chronic exposure to take a strong-inference approach to evaluating the competing hypotheses according to the explicit predictions stemming from them. Adult cockroaches (Nauphoeta cinerea) were chronically exposed to various treatments of different respiratory gas compositions (O 2 , CO 2 and humidity) and the DGC responses were interpreted in light of the a priori predictions stemming from the competing hypotheses. Rates of mass loss during respirometry were also measured for animals acclimated to a range of humidity conditions. The results refute the hypotheses of oxidative damage and underground gas exchange, and provide evidence supporting the hypothesis that DGCs serve to reduce respiratory water loss: cockroaches exposed to low humidity conditions exchange respiratory gases for shorter durations during each DGC and showed lower rates of body mass loss during respirometry than cockroaches exposed to high humidity conditions.

Some insects display an intermittent pattern of gas exchange while at rest, often going hours between breaths. These discontinuous gas exchange cycles (DGCs) are known to have evolved independently within five insect orders, but their possible adaptive benefit and evolutionary origin remain an enigma. Current research is primarily concerned with testing three adaptive hypotheses: that DGCs originally evolved or are currently maintained to (1) limit respiratory water loss, (2) enhance gas exchange in subterranean environments, or (3) limit oxidative damage. These adaptive explanations fail to unite a range of apparently contradictory observations regarding the insects that display DGCs and the conditions under which they occur. Here we argue that DGCs are explained by circadian, developmental, or artificially induced reductions in brain activity. We conclude that this pattern results from the thoracic and abdominal ganglia regulating ventilation in the absence of control from higher neural centers, and it is indicative of a sleeplike state.

Floral heating is not only associated with attraction, but continues throughout the night when beetles are active inside the flower and increases again when they leave. Floral chamber temperatures similar to activity temperatures of actively endothermic beetles imply that thermogenesis is an energy reward.