Abstract

Energy is continuously transformed in the environment through the metabolic activities of organisms. Catabolic reactions generate energy (energy-yielding) which are used to fuel anabolic reactions for maintenance and growth (energy-requiring). These transformations of energy (i.e., bioenergetics) underpin most biogeochemical cycles on Earth and allow the delivery of a wide range of life-supporting ecosystem services. It has long been understood that the amount and types of energy available in an environment influence the rates of biological activity and the complexity of interactions in that system. Traditionally, energy fluxes and stocks have not been described in a quantitative manner, and it is not well-understood how physicochemical theorems such as thermodynamic principles are manifested in environmental systems. Theoretical ecological frameworks (Odum, 1969; Addiscott, 1995) have suggested that the more complex ecosystems become in terms of their food webs, the more efficient they are, i.e., relatively less energy is wasted when utilizing resources. However, this has not been rigorously tested experimentally, but in recent years, scientists in a number of fields have increasingly shown interest in quantifying how bioenergetics constrain and define ecosystem functioning. For example, organic matter in soils has distinct energetic signatures, e.g., energy densities and activation energies (Barré et al., 2016; Williams et al., 2018), and microbial bioenergetics provides empirical data for mechanistic models of carbon turnover in soils, work that is relevant to climate change (Sparling, 1983; Herrmann et al., 2014; Barros et al., 2016; Bölscher et al., 2017). Furthermore, geochemists have quantified the amount of chemolithotrophic energy available for microorganisms in a number of extreme environments to infer the dominant metabolic activities (e.g., McCollom and Shock, 1997; Shock et al., 2010; Osburn et al., 2014). These activities are challenging to monitor due to their inaccessibility and incredibly slow rates of energy processing. Although all of these efforts represent significant progress in the field of biogeochemistry, bioenergetics analysis of natural systems is still in its infancy. Nonetheless, there is increasing interest in using bioenergetics tools to better characterize biogeochemical cycling in water, soils, and sediments in terrestrial, freshwater, and marine ecosystems.