We welcome the opportunity to respond to . We hope that this exchange will help to clarify some of the strengths and weaknesses of the Metabolic theory of ecology (MTE), and will point towards fruitful areas for future research.The MTE has been formulated based on the premise that the structure and dynamics of ecological communities are inextricably linked to individual metabolism. The individual metabolic rate is the rate at which an organism takes up energetic and material resources from the environment, transforms them into useable forms, and allocates them to the fitness-enhancing processes of survival, growth, and reproduction. Interactions between organisms and their environment (including other organisms) are therefore constrained by metabolic rate. Physiologists have long known that there are three primary factors that control metabolic rate: body size, body temperature, and resource availability. MTE builds on this earlier work by providing a quantitative framework to better understand how these three variables combine to affect metabolic rate, and how metabolic rate, in turn, influences the ecology and evolution of populations, communities, and ecosystems (Brown et al. 2004).Gillooly et al. (2001) developed a model for the scaling of metabolic rate that combines the effects of body size and temperature (West et al. 1997, Gillooly et al. 2002, Charnov and Gillooly 2003, Brown et al. 2004. The model leads to a single equation for individual metabolic rate:where b o is a normalization constant, M is body mass, and T is absolute temperature in degrees Kelvin. The size-dependence, M 3/4 , is attributed to geometric constraints on the delivery of energy and materials to cells through biological distribution networks (West et al. 1997). The Boltzmann-Arrhenius term, e (E=kT ; characterizes the exponential effect of temperature, where E is the average activation energy of the respiratory complex (Â0.65 eV), and k is Boltzmann's constant (8.62 )10(5 eV K (1 ) (Gillooly et al. 2006). This simple analytical expression yields quantitative predictions on metabolic rate that are supported by empirical data for a broad assortment of taxonomic groups (Gillooly et al. 2001). raise issues with the MTE that deserve attention and/or clarification. These issues can be divided into two major points of contention: (1) they argue that the proposed mechanisms underlying the size-and temperature-dependencies of individual metabolic rate in Eq. 1 are invalid; and (2) that attempts to link individual metabolic rate to higher levels of biological organization (populations, communities, ecosystems) using MTE are overly simplistic and should therefore be abandoned. Here we respond to these criticisms, and then conclude by discussing the philosophical issues that underlie them.
Points of contention