Gabriel Yvon-Durocher

Methane fluxes show consistent temperature dependence across microbial to ecosystem scales

Methane (CH4) is an important greenhouse gas because it has 25 times the global warming potential of carbon dioxide (CO2) by mass over a century. Recent calculations suggest that atmospheric CH4 emissions have been responsible for approximately 20% of Earth’s warming since pre-industrial times. Understanding how CH4 emissions from ecosystems will respond to expected increases in global temperature is therefore fundamental to predicting whether the carbon cycle will mitigate or accelerate climate change. Methanogenesis is the terminal step in the remineralization of organic matter and is carried out by strictly anaerobic Archaea. Like most other forms of metabolism, methanogenesis is temperature-dependent. However, it is not yet known how this physiological response combines with other biotic processes (for example, methanotrophy, substrate supply, microbial community composition) and abiotic processes (for example, water-table depth) to determine the temperature dependence of ecosystem-level CH4 emissions. It is also not known whether CH4 emissions at the ecosystem level have a fundamentally different temperature dependence than other key fluxes in the carbon cycle, such as photosynthesis and respiration. Here we use meta-analyses to show that seasonal variations in CH4 emissions from a wide range of ecosystems exhibit an average temperature dependence similar to that of CH4 production derived from pure cultures of methanogens and anaerobic microbial communities. This average temperature dependence (0.96 electron volts (eV)), which corresponds to a 57-fold increase between 0 and 30°C, is considerably higher than previously observed for respiration (approximately 0.65 eV) and photosynthesis (approximately 0.3 eV). As a result, we show that both the emission of CH4 and the ratio of CH4 to CO2 emissions increase markedly with seasonal increases in temperature. Our findings suggest that global warming may have a large impact on the relative contributions of CO2 and CH4 to total greenhouse gas emissions from aquatic ecosystems, terrestrial wetlands and rice paddies.

Warming alters the metabolic balance of ecosystems

The carbon cycle modulates climate change, via the regulation of atmospheric CO2, and it represents one of the most important services provided by ecosystems. However, considerable uncertainties remain concerning potential feedback between the biota and the climate. In particular, it is unclear how global warming will affect the metabolic balance between the photosynthetic fixation and respiratory release of CO2 at the ecosystem scale. Here, we present a combination of experimental field data from freshwater mesocosms, and theoretical predictions derived from the metabolic theory of ecology to investigate whether warming will alter the capacity of ecosystems to absorb CO2. Our manipulative experiment simulated the temperature increases predicted for the end of the century and revealed that ecosystem respiration increased at a faster rate than primary production, reducing carbon sequestration by 13 per cent. These results confirmed our theoretical predictions based on the differential activation energies of these two processes. Using only the activation energies for whole ecosystem photosynthesis and respiration we provide a theoretical prediction that accurately quantified the precise magnitude of the reduction in carbon sequestration observed experimentally. We suggest the combination of whole-ecosystem manipulative experiments and ecological theory is one of the most promising and fruitful research areas to predict the impacts of climate change on key ecosystem services.

Reconciling the temperature dependence of respiration across timescales and ecosystem types

Ecosystem respiration is the biotic conversion of organic carbon to carbon dioxide by all of the organisms in an ecosystem, including both consumers and primary producers. Respiration exhibits an exponential temperature dependence at the subcellular and individual levels, but at the ecosystem level respiration can be modified by many variables including community abundance and biomass, which vary substantially among ecosystems. Despite its importance for predicting the responses of the biosphere to climate change, it is as yet unknown whether the temperature dependence of ecosystem respiration varies systematically between aquatic and terrestrial environments. Here we use the largest database of respiratory measurements yet compiled to show that the sensitivity of ecosystem respiration to seasonal changes in temperature is remarkably similar for diverse environments encompassing lakes, rivers, estuaries, the open ocean and forested and non-forested terrestrial ecosystems, with an average activation energy similar to that of the respiratory complex (approximately 0.65 electronvolts (eV)). By contrast, annual ecosystem respiration shows a substantially greater temperature dependence across aquatic (approximately 0.65 eV) versus terrestrial ecosystems (approximately 0.32 eV) that span broad geographic gradients in temperature. Using a model derived from metabolic theory, these findings can be reconciled by similarities in the biochemical kinetics of metabolism at the subcellular level, and fundamental differences in the importance of other variables besides temperature—such as primary productivity and allochthonous carbon inputs—on the structure of aquatic and terrestrial biota at the community level.

Ecological Networks in a Changing Climate

Summary Attempts to gauge the biological impacts of climate change have typically focussed on the lower levels of organization (individuals to populations), rather than considering more complex multi-species systems, such as entire ecological networks (food webs, mutualistic and host–parasitoid networks). We evaluate the possibility that a few principal drivers underpin network-level responses to climate change, and that these drivers can be studied to develop a more coherent theoretical framework than is currently provided by phenomenological approaches. For instance, warming will elevate individual ectotherm metabolic rates, and direct and indirect effects of changes in atmospheric conditions are expected to alter the stoichiometry of interactions between primary consumers and basal resources; these effects are general and pervasive, and will permeate through the entire networks that they affect. In addition, changes in the density and viscosity of aqueous media could alter interactions among very small organisms and disrupt the pycnoclines that currently compartmentalize many aquatic networks in time and space. We identify a range of approaches and potential model systems that are particularly well suited to network-level studies within the context of climate change. We also highlight potentially fruitful areas of research with a view to improving our predictive power regarding climate change impacts on networks. We focus throughout on mechanistic approaches rooted in first principles that demonstrate potential for application across a wide range of taxa and systems.

Warming alters community size structure and ecosystem functioning

Global warming can affect all levels of biological complexity, though we currently understand least about its potential impact on communities and ecosystems. At the ecosystem level, warming has the capacity to alter the structure of communities and the rates of key ecosystem processes they mediate. Here we assessed the effects of a 4°C rise in temperature on the size structure and taxonomic composition of benthic communities in aquatic mesocosms, and the rates of detrital decomposition they mediated. Warming had no effect on biodiversity, but altered community size structure in two ways. In spring, warmer systems exhibited steeper size spectra driven by declines in total community biomass and the proportion of large organisms. By contrast, in autumn, warmer systems had shallower size spectra driven by elevated total community biomass and a greater proportion of large organisms. Community-level shifts were mirrored by changes in decomposition rates. Temperature-corrected microbial and macrofaunal decomposition rates reflected the shifts in community structure and were strongly correlated with biomass across mesocosms. Our study demonstrates that the 4°C rise in temperature expected by the end of the century has the potential to alter the structure and functioning of aquatic ecosystems profoundly, as well as the intimate linkages between these levels of ecological organization.

Global change and food webs in running waters

Riverine habitats are vulnerable to a host of environmental stressors, many of which are increasing in frequency and intensity across the globe. Climate change is arguably the greatest threat on the horizon, with serious implications for freshwater food webs via alterations in thermal regimes, resource quality and availability, and hydrology. This will induce radical restructuring of many food webs, by altering the identity of nodes, the strength and patterning of interactions and consequently the dynamics and architecture of the trophic network as a whole. Although such effects are likely to be apparent globally, they are predicted to be especially rapid and dramatic in high altitude and latitude ecosystems, which represent ‘sentinel systems’. The complex and subtle connections between members of a food web and potential synergistic interactions with other environmental stressors can lead to seemingly counterintuitive responses to perturbations that cannot be predicted from the traditional focus of studying individual species in isolation. In this review, we highlight the need for developing new network-based approaches to understand and predict the consequences of global change in running waters.

Rapid evolution of metabolic traits explains thermal adaptation in phytoplankton

Abstract Understanding the mechanisms that determine how phytoplankton adapt to warming will substantially improve the realism of models describing ecological and biogeochemical effects of climate change. Here, we quantify the evolution of elevated thermal tolerance in the phytoplankton, Chlorella vulgaris. Initially, population growth was limited at higher temperatures because respiration was more sensitive to temperature than photosynthesis meaning less carbon was available for growth. Tolerance to high temperature evolved after ≈ 100 generations via greater down‐regulation of respiration relative to photosynthesis. By down‐regulating respiration, phytoplankton overcame the metabolic constraint imposed by the greater temperature sensitivity of respiration and more efficiently allocated fixed carbon to growth. Rapid evolution of carbon‐use efficiency provides a potentially general mechanism for thermal adaptation in phytoplankton and implies that evolutionary responses in phytoplankton will modify biogeochemical cycles and hence food web structure and function under warming. Models of climate futures that ignore adaptation would usefully be revisited.

Linking community size structure and ecosystem functioning using metabolic theory

Understanding how biogeochemical cycles relate to the structure of ecological communities is a central research question in ecology. Here we approach this problem by focusing on body size, which is an easily measured species trait that has a pervasive influence on multiple aspects of community structure and ecosystem functioning. We test the predictions of a model derived from metabolic theory using data on ecosystem metabolism and community size structure. These data were collected as part of an aquatic mesocosm experiment that was designed to simulate future environmental warming. Our analyses demonstrate significant linkages between community size structure and ecosystem functioning, and the effects of warming on these links. Specifically, we show that carbon fluxes were significantly influenced by seasonal variation in temperature, and yielded activation energies remarkably similar to those predicted based on the temperature dependencies of individual-level photosynthesis and respiration. We also show that community size structure significantly influenced fluxes of ecosystem respiration and gross primary production, particularly at the annual time-scale. Assessing size structure and the factors that control it, both empirically and theoretically, therefore promises to aid in understanding links between individual organisms and biogeochemical cycles, and in predicting the responses of key ecosystem functions to future environmental change.

Five Years of Experimental Warming Increases the Biodiversity and Productivity of Phytoplankton.

Phytoplankton are key components of aquatic ecosystems, fixing CO2 from the atmosphere through photosynthesis and supporting secondary production, yet relatively little is known about how future global warming might alter their biodiversity and associated ecosystem functioning. Here, we explore how the structure, function, and biodiversity of a planktonic metacommunity was altered after five years of experimental warming. Our outdoor mesocosm experiment was open to natural dispersal from the regional species pool, allowing us to explore the effects of experimental warming in the context of metacommunity dynamics. Warming of 4°C led to a 67% increase in the species richness of the phytoplankton, more evenly-distributed abundance, and higher rates of gross primary productivity. Warming elevated productivity indirectly, by increasing the biodiversity and biomass of the local phytoplankton communities. Warming also systematically shifted the taxonomic and functional trait composition of the phytoplankton, favoring large, colonial, inedible phytoplankton taxa, suggesting stronger top-down control, mediated by zooplankton grazing played an important role. Overall, our findings suggest that temperature can modulate species coexistence, and through such mechanisms, global warming could, in some cases, increase the species richness and productivity of phytoplankton communities.

A metabolic perspective on competition and body size reductions with warming

Temperature is a key driver of ecological processes and patterns. The ramifications of temperature for ecological communities include not only its direct effects on the physiology of individuals, but also how these effects play out in the context of other processes such as competition. Apparently idiosyncratic or difficult to predict effects of temperature on competitive outcomes are well represented in the literature. General theoretical understanding of how physiological influences of temperature filter through community dynamics to determine outcomes is limited. We present a theoretical framework for predicting the effects of temperature on competition among species, based on understanding the effects of temperature on the physiological and population parameters of the species. The approach helps unify formal resource competition theory with metabolic and physiological ecology. Phytoplankton and many other ectotherms are smaller at higher temperatures. This has been observed experimentally, across geographical gradients, and as change accompanying climate warming, but it has not been explained in terms of competition. As a case study, we apply our theoretical framework to competition for nutrients among differently sized phytoplankton. Based on this analysis, we hypothesize that the prevalence of smaller phytoplankton at higher temperatures is at least partly due to an accentuated competitive advantage of smaller cells at higher temperatures with respect to nutrient uptake and growth. We examine the scope for extending the approach to understand resource competition, generally, among ectotherms of different sizes.