Adaptive differences in plant physiology and ecosystem paradoxes: Insights from metabolic scaling theory

The link between variation in species-specific plant traits, larger scale patterns of productivity, and other ecosystem processes is an important focus for global change research. Understanding such linkages requires synthesis of evolutionary, biogeograpahic, and biogeochemical approaches to ecological research. Recent observations reveal several apparently paradoxical patterns across ecosystems. When compared with warmer low latitudes, ecosystems from cold northerly latitudes are described by (1) a greater temperature normalized instantaneous flux of CO₂ and energy; and (2) similar annual values of gross primary production (GPP), and possibly net primary production. Recently, several authors attributed constancy in GPP to historical and abiotic factors. Here, we show that metabolic scaling theory can be used to provide an alternative 'biotically driven' hypothesis. The model provides a baseline for understanding how potentially adaptive variation in plant size and traits associated with metabolism and biomass production in differing biomes can influence whole-ecosystem processes. The implication is that one cannot extrapolate leaf/lab/forest level functional responses to the globe without considering evolutionary and geographic variation in traits associated with metabolism. We test one key implication of this model -that directional and adaptive changes in metabolic and stoichiometric traits of autotrophs may mediate patterns of plant growth across broad temperature gradients. In support of our model, on average, mass-corrected whole-plant growth rates are not related to differences in growing season temperature or latitude. Further, we show how these changes in autotrophic physiology and nutrient content across gradients may have important implications for understanding: (i) the origin of paradoxical ecosystem behavior; (ii) the potential efficiency of whole-ecosystem carbon dynamics as measured by the quotient of system capacities for respiration, R, and assimilation, A; and (iii) the origin of several 'ecosystem constants'-attributes of ecological systems that apparently do not vary with temperature (and thus with latitude). Together, these results highlight the potential critical importance of community ecology and functional evolutionary/physiological ecology for understanding the role of the biosphere within the integrated earth system.