The metabolic theory of ecology (MTE) is the ecological component of the more general West, Brown, and Enquist (WBE) Metabolic Scaling Theory and Kleiber's law. It posits that the metabolic rate of organisms is the fundamental biological rate that governs most observed patterns in ecology. MTE is part of a larger set of theory known as metabolic scaling theory that attempts to provide a unified theory for the importance of metabolism in driving pattern and process in biology from the level of cells all the way to the biosphere. MTE is based on an interpretation of the relationships between body size, body temperature, and metabolic rate across all organisms. Small-bodied organisms tend to have higher mass-specific metabolic rates than larger-bodied organisms. Furthermore, organisms that operate at warm temperatures through endothermy or by living in warm environments tend towards higher metabolic rates than organisms that operate at colder temperatures. This pattern is consistent from the unicellular level up to the level of the largest animals and plants on the planet. In MTE, this relationship is considered to be the primary constraint that influences biological processes (via their rates and times) at all levels of organization (from individual up to ecosystem level). MTE is a macroecological theory that aims to be universal in scope and application.
Fundamental concepts in MTE
Metabolism Metabolic pathways consist of complex networks, which are responsible for the processing of both energy and material. The metabolic rate of a heterotroph is defined as the rate of respiration in which energy is obtained by oxidation of a carbon compound. The rate of photosynthesis on the other hand, indicates the metabolic rate of an autotroph. According to MTE, both body size and temperature affect the metabolic rate of an organism. Metabolic rate scales as 3/4 power of body size, and its relationship with temperature is described by the Van't Hoff-Arrhenius equation over the range of 0 to 40 °C.
Stoichiometry From the ecological perspective, stoichiometry is concerned with the proportion of elements in both living organisms and their environment. In order to survive and maintain metabolism, an organism must be able to obtain crucial elements and excrete waste products. As a result, the elemental composition of an organism would be different from the exterior environment. Through metabolism, body size can affect stoichiometry. For example, small organism tend to store most of their phosphorus in rRNA due to their high metabolic rate, whereas large organisms mostly invest this element inside the skeletal structure. Thus, concentration of elements to some extent can limit the rate of biological processes. Inside an ecosystem, the rate of flux and turn over of elements by inhabitants, combined with the influence of abiotic factors, determine the concentration of elements.
Theoretical background Metabolic rate scales with the mass of an organism of a given species according to Kleiber's law where B is whole organism metabolic rate (in watts or other unit of power), M is organism mass (in kg), and Bo is a mass-independent normalization constant (given in a unit of power divided by a unit of mass. In this case, watts per kilogram):
B = B o M 3 / 4 {\displaystyle B=B_{o}M^{3/4}\,}
At increased temperatures, chemical reactions proceed faster. This relationship is described by the Boltzmann factor, where E is activation energy in electronvolts or joules, T is absolute temperature in kelvins, and k is the Boltzmann constant in eV/K or J/K:
e − E k T {\displaystyle e^{-{\frac {E}{k\,T}}}}
While Bo in the previous equation is mass-independent, it is not explicitly independent of temperature. To explain the relationship between body mass and temperature, building on earlier work showing that the effects of both body mass and temperature could be combined multiplicatively in a single equation, the two equations above can be combined to produce the primary equation of the MTE, where bo is a normalization constant that is independent of body size or temperature:
B = b o M 3 / 4 e − E k T {\displaystyle B=b_{o}M^{3/4}e^{-{\frac {E}{k\,T}}}}
According to this relationship, metabolic rate is a function of an organism's body mass and body temperature. By this equation, large organisms have higher metabolic rates (in watts) than small organisms, and organisms at high body temperatures have higher metabolic rates than those that exist at low body temperatures. However, specific metabolic rate (SMR, in watts/kg) is given by
S M R = ( B / M ) = b o M − 1 / 4 e − E k T {\displaystyle SMR=(B/M)=b_{o}M^{-1/4}e^{-{\frac {E}{k\,T}}}}
Hence SMR for large organisms are lower than small organisms.
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