Environmental Constraints that Limit Methanogenesis

Abstract

Methanogens are active in many different ecosystems, including habitats with biologically-derived organic matter as substrates such as aquatic sediments, wetlands, agricultural or natural soils subject to inundation, sewage digesters, and the anoxic portions of animal digestive tracts. Methanogens are also present in habitats with geochemically-supplied substrates such as hot springs, hydrothermal vents, volcanically-influenced habitats, and, potentially, the deep crustal subsurface. Methanogens as a group tolerate a broad range of physicochemical conditions, including temperatures from −2 {\textdegree}C to 122 {\textdegree}C, pH values of 3.0--10.2, salinities up to halite saturation, and pressures of at least 75 MPa. Globally, variations in methane emissions can be explained to a large degree by variations in temperature and water availability. The distribution and activity of methanogens are constrained by ecological interactions that can be stimulatory or competitive, and by physicochemical factors that act at the biochemical or bioenergetic levels. In addition to the constraints placed on methanogens by physicochemical extremes, methanogen distribution and activity are constrained by the availability of energy and nutrients, the presence of inhibitory molecules (most notably oxygen), and the seawater anion, sulfate, due to competitive ecological interactions. Although methanogen tolerances to individual extremes are documented in culture, and the corresponding biochemical adaptations are understood to varying degrees, the natural environment frequently presents combinations of extreme conditions and energy limitations that may limit methanogen distribution to less than the optimally tolerated range of a single parameter. Little is understood about the compound effects of such extremes, nor the commonalities among them that will ultimately form the basis for predictive models of environmental methanogen population distribution. Future work that targets these questions, through a combination of culture work, ``omic'' analyses, in situ studies, and conceptual and quantitative models, will be needed to better understand the physiological ecology of methanogens.