Completing the FACE of elevated CO₂ research

We appraise the present geographical extent and inherent knowledge limits, following two decades of research on elevated CO2 responses in plant communities, and ask whether such research has answered the key question in quantifying the limits of compensatory CO2 uptake in the major biomes. Our synthesis of all ecosystem-scale (between 10 m2 and 3000 m2 total experimental plot area) elevated CO2 (eCO2) experiments in natural ecosystems conducted worldwide since 1987 (n = 151) demonstrates that the locations of these eCO2 experiments have been spatially biased, targeting primarily the temperate ecosystems of northern America and Europe. We consider the consequences, suggesting fundamentally that this limits the capacity of the research to understand how the world's major plant communities will respond to eCO2. Most notably, our synthesis shows that this research lacks understanding of impacts on tropical forests and boreal regions, which are potentially the most significant biomes for C sink and storage activity, respectively. Using a meta-analysis of the available data across all biomes, we show equivocal increases in net primary productivity (NPP) from eCO2 studies, suggesting that global validation is needed, especially in the most important biomes for C processing. Further, our meta-analysis identifies that few research programs have addressed eCO2 effects on below-ground C storage, such that at the global scale, no overall responses are discernable. Given the disparity highlighted in the distribution of eCO2 experiments globally, we suggest opportunities for newly-industrialized or developing nations to become involved in further research, particularly as these countries host some of the most important regions for tropical or sub-tropical forest systems. Modeling approaches that thus far have attempted to understand the biological response to eCO2 are constrained with respect to collective predictions, suggesting that further work is needed, which will link models to in situ eCO2 experiments, in order to understand how the world's most important regions for terrestrial C uptake and storage will respond to a future eCO2 atmosphere

Root oxygen mitigates methane fluxes in tropical peatlands

Tropical peatlands are a globally important source of methane, a potent greenhouse gas. Vegetation is critical in regulating fluxes, providing a conduit for emissions and regular carbon inputs. However, plant roots also release oxygen, which might mitigate methane efflux through oxidation prior to emission from the peat surface. Here we show, using in situ mesocosms, that root exclusion can reduce methane fluxes by a maximum of 92% depending on species, likely driven by the significant decrease in root inputs of oxygen and changes in the balance of methane transport pathways. Methanotroph abundance decreased with reduced oxygen input, demonstrating a likely mechanism for the observed response. These first methane oxidation estimates for a tropical peatland demonstrate that although plants provide an important pathway for methane loss, this can be balanced by the influence of root oxygen inputs that mitigate peat surface methane emissions

Plant community composition and an insect outbreak influence phenol oxidase activity and soil-litter biochemistry in a sub-Arctic birch-heath

Rates of decomposition in Arctic soils are regulated by temperature and moisture, but substrate availability is dictated by vegetation inputs, which are also subject to biotic influences. Here, we examine how leaf and litter inputs from individual dwarf shrub species influence soil enzyme activity in a sub-Arctic heath community in Abisko, Sweden. We further consider how foliar damage via insect herbivory (and outbreak) affects the soil community and decomposition. During the peak growing season (July 2011), we assessed how shrub community composition (Empetrum hermaphroditum, Vaccinium myrtillus, V. uliginosum and V. vitis-idaea) determined litter and soil phenol oxidase activity. A periodic severe outbreak of autumn moth larvae (Epirrita autumnata) affected this community in the following year (July 2012), and we used this to investigate its impact on relationships with phenol oxidase activity, soil respiration, soluble NH4 + and soluble phenolics; the soluble factors being directly associated with inputs from insect larval waste (frass). Pre-outbreak (2011), the strongest relationship observed was higher phenol oxidase activity with E. hermaphroditum cover. In the outbreak year (2012), phenol oxidase activity had the strongest relationship with damage to the deciduous species V. myrtillus, with greater herbivory lowering activity. For the other deciduous species, V. uliginosum, soil NH4 + and phenolics were negatively correlated with foliar larval damage. Phenol oxidase activity was not affected by herbivory of the evergreen species, but there was a strong positive relationship observed between E. hermaphroditum community abundance and soil respiration. We highlight the dominant role of E. hermaphroditum in such sub-Arctic shrub communities and show that even during insect outbreaks, it can dictate soil processes

Root herbivores drive changes to plant primary chemistry, but root loss is mitigated under elevated atmospheric CO2

Above- and belowground herbivory represents a major challenge to crop productivity and sustainable agriculture worldwide. How this threat from multiple herbivore pests will change under anthropogenic climate change, via altered trophic interactions and plant response traits, is key to understanding future crop resistance to herbivory. In this study, we hypothesized that atmospheric carbon enrichment would increase the amount (biomass) and quality (C:N ratio) of crop plant resources for above- and belowground herbivore species. In a controlled environment facility, we conducted a microcosm experiment using the large raspberry aphid (Amphorophora idaei), the root feeding larvae of the vine weevil (Otiorhynchus sulcatus), and the raspberry (Rubus idaeus) host-plant. There were four herbivore treatments (control, aphid only, weevil only and a combination of both herbivores) and an ambient (aCO2) or elevated (eCO2) CO2 treatment (390 versus 650 ± 50 μmol/mol) assigned to two raspberry cultivars (cv Glen Ample or Glen Clova) varying in resistance to aphid herbivory. Contrary to our predictions, eCO2 did not increase crop biomass or the C:N ratio of the plant tissues, nor affect herbivore abundance either directly or via the host-plant. Root herbivory reduced belowground crop biomass under aCO2 but not eCO2, suggesting that crops could tolerate attack in a CO2 enriched environment. Root herbivory also increased the C:N ratio in leaf tissue at eCO2, potentially due to decreased N uptake indicated by lower N concentrations found in the roots. Root herbivory greatly increased root C concentrations under both CO2 treatments. Our findings confirm that responses of crop biomass and biochemistry to climate change need examining within the context of herbivory, as biotic interactions appear as important as direct effects of eCO2 on crop productivity

Environmental and microbial controls on microbial necromass recycling, an important precursor for soil carbon stabilization

There is an emerging consensus that microbial necromass carbon is the primary constituent of stable soil carbon, yet the controls on the stabilization process are unknown. Prior to stabilization, microbial necromass may be recycled by the microbial community. We propose that the efficiency of this recycling is a critical determinant of soil carbon stabilization rates. Here we explore the controls on necromass recycling efficiency in 27 UK grassland soils using stable isotope tracing and indicator species analysis. We found that recycling efficiency was unaffected by land management. Instead, recycling efficiency increased with microbial growth rate on necromass, and was highest in soils with low historical precipitation. We identified bacterial and fungal indicators of necromass recycling efficiency, which could be used to clarify soil carbon stabilization mechanisms. We conclude that environmental and microbial controls have a strong influence on necromass recycling, and suggest that this, in turn, influences soil carbon stabilization

Can digital image classification be used as a standardised method for surveying peatland vegetation cover?

The ability to carry out systematic, accurate and repeatable vegetation surveys is an essential part of long-term scientific studies into ecosystem biodiversity and functioning. However, current widely used traditional survey techniques such as destructive harvests, pin frame quadrats and visual cover estimates can be very time consuming and are prone to subjective variations. We investigated the use of digital image techniques as an alternative way of recording vegetation cover to plant functional type level on a peatland ecosystem. Using an established plant manipulation experimental site at Moor House NNR (an Environmental Change Network site), we compared visual cover estimates of peatland vegetation with cover estimates using digital image classification methods, from 0.5 m × 0.5 m field plots. Our results show that digital image classification of photographs taken with a standard digital camera can be used successfully to estimate dwarf-shrub and graminoid vegetation cover at a comparable level to field visual cover estimates, although the methods were less effective for lower plants such as mosses and lichens. Our study illustrates the novel application of digital image techniques to provide a new way of measuring and monitoring peatland vegetation to the plant functional group level, which is less vulnerable to surveyor bias than are visual field surveys. Furthermore, as such digital techniques are highly repeatable, we suggest that they have potential for use in long-term monitoring studies, at both plot and landscape scales

Land use not litter quality is a stronger driver of decomposition in hyperdiverse tropical forest

In hyperdiverse tropical forests, the key drivers of litter decomposition are poorly understood despite its crucial role in facilitating nutrient availability for plants and microbes. Selective logging is a pressing land use with potential for considerable impacts on plant-soil interactions, litter decomposition, and nutrient cycling. Here, in Borneo's tropical rainforests, we test the hypothesis that decomposition is driven by litter quality and that there is a significant "home-field advantage," that is positive interaction between local litter quality and land use. We determined mass loss of leaf litter, collected from selectively logged and old-growth forest, in a fully factorial experimental design, using meshes that either allowed or precluded access by mesofauna. We measured leaf litter chemical composition before and after the experiment. Key soil chemical and biological properties and microclimatic conditions were measured as land-use descriptors. We found that despite substantial differences in litter quality, the main driver of decomposition was land-use type. Whilst inclusion of mesofauna accelerated decomposition, their effect was independent of land use and litter quality. Decomposition of all litters was slower in selectively logged forest than in old-growth forest. However, there was significantly greater loss of nutrients from litter, especially phosphorus, in selectively logged forest. The analyses of several covariates detected minor microclimatic differences between land-use types but no alterations in soil chemical properties or free-living microbial composition. These results demonstrate that selective logging can significantly reduce litter decomposition in tropical rainforest with no evidence of a home-field advantage. We show that loss of key limiting nutrients from litter (P & N) is greater in selectively logged forest. Overall, the findings hint at subtle differences in microclimate overriding litter quality that result in reduced decomposition rates in selectively logged forests and potentially affect biogeochemical nutrient cycling in the long term

Microbial responses to warming enhance soil carbon loss following translocation across a tropical forest elevation gradient

Tropical soils contain huge carbon stocks, which climate warming is projected to reduce by stimulating organic matter decomposition, creating a positive feedback that will promote further warming. Models predict that the loss of carbon from warming soils will be mediated by microbial physiology, but no empirical data are available on the response of soil carbon and microbial physiology to warming in tropical forests, which dominate the terrestrial carbon cycle. Here we show that warming caused a considerable loss of soil carbon that was enhanced by associated changes in microbial physiology. By translocating soils across a 3000 m elevation gradient in tropical forest, equivalent to a temperature change of ± 15 °C, we found that soil carbon declined over 5 years by 4% in response to each 1 °C increase in temperature. The total loss of carbon was related to its original quantity and lability, and was enhanced by changes in microbial physiology including increased microbial carbon‐use‐efficiency, shifts in community composition towards microbial taxa associated with warmer temperatures, and increased activity of hydrolytic enzymes. These findings suggest that microbial feedbacks will cause considerable loss of carbon from tropical forest soils in response to predicted climatic warming this century

Do soil depth and plant community composition interact to modify the resistance and resilience of grassland ecosystem functioning to drought?

While the effect of drought on plant communities and their associated ecosystem functions is well studied, little research has considered how responses are modified by soil depth and depth heterogeneity. We conducted a mesocosm study comprising shallow and deep soils, and variable and uniform soil depths, and two levels of plant community composition, and exposed them to a simulated drought to test for interactive effects of these treatments on the resilience of carbon dioxide fluxes, plant functional traits, and soil chemical properties. We tested the hypotheses that: (a) shallow and variable depth soils lead to increased resistance and resilience of ecosystem functions to drought due to more exploitative plant trait strategies; (b) plant communities associated with intensively managed high fertility soils, will have more exploitative root traits than extensively managed, lower fertility plant communities. These traits will be associated with higher resistance and resilience to drought and may interact with soil depth and depth heterogeneity to amplify the effects on ecosystem functions. Our results showed that while there were strong soil depth/heterogeneity effects on plant-driven carbon fluxes, it did not affect resistance or resilience to drought, and there were no treatment effects on plant-available carbon or nitrogen. We did observe a significant increase in exploitative root traits in shallow and variable soils relative to deep and uniform, which may have resulted in a compensation effect which led to the similar drought responses. Plant community compositions representative of intensive management were more drought resilient than more diverse “extensive” communities irrespective of soil depth or soil depth heterogeneity. In intensively managed plant communities, root traits were more representative of exploitative strategies. Taken together, our results suggest that reorganization of root traits in response to soil depth could buffer drought effects on ecosystem functions

Sticky Dead Microbes: rapid abiotic retention of microbial necromass in soil

Microbial necromass dominates soil organic matter. Recent research on necromass and soil carbon storage has focused on necromass production and stabilization mechanisms but not on the mechanisms of necromass retention. We present evidence from soil incubations with stable-isotope labeled necromass that abiotic adsorption may be more important than biotic immobilization for short-term necromass retention. We demonstrate that necromass adsorbs not only to mineral surfaces, but may also interact with other necromass. Furthermore, necromass cell chemistry alters necromass-necromass interaction, with more bacterial tracer retained when there is yeast necromass present. These findings suggest that the adsorption and abiotic interaction of microbial necromass and its functional properties, beyond chemical stability, deserve further investigation in the context of soil carbon sequestration