Drought may exacerbate dryland soil inorganic carbon loss under warming climate conditions.

Low moisture conditions result in substantially more soil inorganic carbon (SIC) than soil organic carbon (SOC) in drylands. However, whether and how changes in moisture affect the temperature response of SIC in drylands are poorly understood. Here, we report that the temperature sensitivity of SIC dissolution increases but that of SOC decomposition decreases with increasing natural aridity from 30 dryland sites along a 4,500 km aridity gradient in northern China. To directly test the effects of moisture changes alone, a soil moisture control experiment also revealed opposite moisture effects on the temperature sensitivities of SIC and SOC. Moreover, we found that the temperature sensitivity of SIC was primarily regulated by pH and base cations, whereas that of SOC was mainly regulated by physicochemical protection along the aridity gradient. Given the overall increases in aridity in a warming world, our findings highlight that drought may exacerbate dryland soil carbon loss from SIC under warming.

Thermal adaptation of microbial respiration persists throughout long-term soil carbon decomposition.

Soil microbial respiration is expected to show adaptations to changing temperatures, greatly weakening the magnitude of feedback over time, as shown in labile carbon substrates. However, whether such thermal adaptation persists during long-term soil carbon decomposition as carbon substrates decrease in decomposability remains unknown. Here, we conducted a 6-year incubation experiment in natural and arable soils with distinct properties under three temperatures (10, 20 and 30°C). Mass-specific microbial respiration was consistently lower under higher long-term incubation temperatures, suggesting the occurrence and persistence of microbial thermal adaptation in long-term soil carbon decomposition. Furthermore, changes in microbial community composition and function largely explained the persistence of microbial respiratory thermal adaptation. If such thermal adaptation generally occurs in large low-decomposability carbon pools, warming-induced soil carbon losses may be lower than previously predicted and thus may not contribute as much as expected to greenhouse warming.

Convergence of carbon sink magnitude and water table depth in global wetlands.

Wetlands are strategic areas for carbon uptake, but accurate assessments of their sequestration ability are limited by the uncertainty and variability in their carbon balances. Based on 2385 observations of annual net ecosystem production from global wetlands, we show that the mean net carbon sinks of inland wetlands, peatlands and coastal wetlands are 0.57, 0.29 and 1.88 tons of carbon per hectare per year, respectively, with a mean value of 0.57 tons of carbon per hectare per year weighted by the distribution area of different wetland types. Carbon sinks are mainly in Asia and North America. Within and across wetland types, we find that water table depth (WTD) exerts greater control than climate- and ecosystem-related variables, and an increase in WTD results in a stronger carbon sink. Our results highlight an urgent need to sustain wetland hydrology under global change; otherwise, wetlands are at high risk of becoming carbon sources to the atmosphere.

Temperature fluctuation promotes the thermal adaptation of soil microbial respiration.

The magnitude of the feedback between soil microbial respiration and increased mean temperature may decrease (a process called thermal adaptation) or increase over time, and accurately representing this feedback in models improves predictions of soil carbon loss rates. However, climate change entails changes not only in mean temperature but also in temperature fluctuation, and how this fluctuation regulates the thermal response of microbial respiration has never been systematically evaluated. By analysing subtropical forest soils from a 2,000 km transect across China, we showed that although a positive relationship between soil microbial biomass-specific respiration and temperature was observed under increased constant incubation temperature, an increasing temperature fluctuation had a stronger negative effect. Our results further indicated that changes in bacterial community composition and reduced activities of carbon degradation enzymes promoted the effect of temperature fluctuation. This adaptive response of soil microbial respiration suggests that climate warming may have a lesser exacerbating effect on atmospheric CO2 concentrations than predicted.

Opposing seasonal temperature dependencies of CO2 and CH4 emissions from wetlands.

Wetlands are critically important to global climate change because of their role in modulating the release of atmospheric greenhouse gases (GHGs) carbon dioxide (CO2 ) and methane (CH4 ). Temperature plays a crucial role in wetland GHG emissions, while the general pattern for seasonal temperature dependencies of wetland CO2 and CH4 emissions is poorly understood. Here we show opposite seasonal temperature dependencies of CO2 and CH4 emissions by using 36,663 daily observations of simultaneous measurements of ecosystem-scale CO2 and CH4 emissions in 42 widely distributed wetlands from the FLUXNET-CH4 database. Specifically, the temperature dependence of CO2 emissions decreased with increasing monthly mean temperature, but the opposite was true for that of CH4 emissions. Neglecting seasonal temperature dependencies may overestimate wetland CO2 and CH4 emissions compared to the use of a year-based static and consistent temperature dependence parameter when only considering temperature effects. Our findings highlight the importance of incorporating the remarkable seasonality in temperature dependence into process-based biogeochemical models to predict feedbacks of wetland GHG emissions to climate warming.

Low soil moisture suppresses the thermal compensatory response of microbial respiration.

The thermal compensatory response of microbial respiration contributes to a decrease in warming-induced enhancement of soil respiration over time, which could weaken the positive feedback between the carbon cycle and climate warming. Climate warming is also predicted to cause a worldwide decrease in soil moisture, which has an effect on the microbial metabolism of soil carbon. However, whether and how changes in moisture affect the thermal compensatory response of microbial respiration are unexplored. Here, using soils from an 8-year warming experiment in an alpine grassland, we assayed the thermal response of microbial respiration rates at different soil moisture levels. The results showed that relatively low soil moisture suppressed the thermal compensatory response of microbial respiration, leading to an enhanced response to warming. A subsequent moisture incubation experiment involving off-plot soils also showed that the response of microbial respiration to 100 d warming shifted from a slight compensatory response to an enhanced response with decreasing incubation moisture. Further analysis revealed that such respiration regulation by moisture was associated with shifts in enzymatic activities and carbon use efficiency. Our findings suggest that future drought induced by climate warming might weaken the thermal compensatory capacity of microbial respiration, with important consequences for carbon-climate feedback.

Contrasting effects of maize litter and litter-derived biochar on the temperature sensitivity of paddy soil organic matter decomposition.

Organic matter input regulates the rate and temperature sensitivity (expressed as Q 10) of soil organic matter (SOM) decomposition by changing microbial composition and activities. It remains unclear how the incorporation of litter-made biochar instead of litter affects the Q 10 of SOM decomposition. Using a unique combination of two-and three-source partitioning methods (isotopic discrimination between C3/C4 pathways and 14C labeling), we investigated: (1) how maize litter versus litter-made biochar (of C4 origin) addition influenced the Q 10 of SOM (C3 origin) under 10°C warming, and (2) how the litter or biochar amendments affected the Q 10 of 14C-labeled fresh organic matter (FOM) after long-term incubation. Compared with biochar addition, litter increased the rates and Q 10 of mass-specific respiration, SOM and FOM decomposition, as well as the contents of SOM-derived dissolved organic C (DOC) and total phospholipid fatty acids (PLFA). Litter-amended soils have much higher activities (V max) of ß-glucosidase, N-acetyl-ß-glucosaminidase, and leucine aminopeptidase, suggesting larger enzyme pools than in soils with biochar. The Q 10 of enzyme V max (1.6-2.0) and K m (1.2-1.4) were similar between litter-and biochar-amended soils, and remained stable with warming. However, warming reduced microbial biomass (PLFA) and enzyme activity (V max), suggesting decreased enzyme production associated with smaller microbial biomass or faster enzyme turnover at higher temperatures. Reductions in PLFA content and enzyme V max due to warming were larger in litter-amended soils (by 31%) than in the control and biochar-amended soils (by 4-11%), implying the active litter-feeding microorganisms have a smaller degree of heat tolerance than the inactive microorganisms under biochar amendments. The reduction in enzyme activity (V max) by warming was lower in soils with biochar than in the control soil. Our modeling suggested that the higher Q 10 in litter-amended soils was mainly caused by faster C loss under warming, linked to reductions in microbial biomass and growth efficiency, rather than the slightly increased SOM-originated substrate availability (DOC). Overall, using straw-made biochar instead of straw per se as a soil amendment lowers the Q 10 of SOM and FOM by making microbial communities and enzyme pools more temperature-tolerant, and consequently reduces SOM losses under warming.

Microbial respiratory thermal adaptation is regulated by r-/K-strategy dominance.

Microbial thermal adaptation is considered to be one of the core mechanisms affecting soil carbon cycling. However, the role of microbial community composition in controlling thermal adaptation is poorly understood. Using microbial communities from the rhizosphere and bulk soils in an 8-year warming experiment as a model, we experimentally demonstrate that respiratory thermal adaptation was much stronger in microbial K-strategist-dominated bulk soils than in microbial r-strategist-dominated rhizosphere soils. Soil carbon availability exerted strong selection on the dominant ecological strategy of the microbial community, indirectly influencing respiratory thermal adaptation. Our findings shed light on the linchpin of the dominant ecological strategy exhibited by the microbial community in determining its respiratory thermal adaptation, with implications for understanding soil carbon losses under warming.

Spatiotemporal variability of fire effects on soil carbon and nitrogen: A global meta-analysis.

A consensus about the fire-related soil carbon (C) and nitrogen (N) impacts that determine soil health and ecosystem services at the global scale remains elusive. Here, we conducted a global meta-analysis of 3173 observations with 1444, 1334, 228, and 167 observations for soil C, N, pyrogenic C (PyC), and the percent of PyC to total organic C (PyC/TOC) from 296 field studies. Results showed that fire significantly decreased soil C (-15.2%) and N (-14.6%) but increased soil PyC (40.6%) and PyC/TOC (30.3%). Stronger negative fire impacts on soil C and N were found in tropical and temperate climates than in Mediterranean and subtropical climates; stronger effects were found in forest ecosystems than in non-forest ecosystems. Wildfire and high-severity fire led to greater soil C and N losses than prescribed and low-severity fires, respectively, while they promoted greater increases in soil PyC and PyC/TOC than prescribed and low-severity fires, respectively. However, soil C and N recovered to control levels approximately 10 years after fire, which is a shorter period than previously determined. These results suggest that fire-induced PyC production should be accounted for in the C budget under global change. These results will improve our knowledge of the spatiotemporal variability of fire effects on soil C and N storage and have implications for fire management and ecosystem recovery.

Key microorganisms mediate soil carbon-climate feedbacks in forest ecosystems.

Soil microorganisms are known to significantly contribute to climate change through soil carbon (C) cycle feedbacks. However, it is challenging to incorporate these feedbacks into predictions of future patterns of terrestrial C cycling, largely because of the vast diversity of soil microorganisms and their responses to environmental conditions. Here, we show that the composition of the bacterial community can provide information about the microbial community-level thermal response (MCTR), which drives ecosystem-scale soil C-climate feedbacks. The dominant taxa from 169 sites representing a gradient from tropical to boreal forest mainly belonged to the phyla Actinobacteria and Acidobacteria. Moreover, we show that the MCTR in warm biomes and acidic soils was linked primarily to bacteria, whereas the MCTR in cold biomes and alkaline soils was primarily associated with fungi. Our results provide strong empirical evidence of linkages between microbial composition and the MCTR across a wide range of forests, and suggest the importance of specific microorganisms in regulating soil C-climate feedbacks.

Temperature adaptation of soil microbial respiration in alpine, boreal and tropical soils: An application of the square root (Ratkowsky) model.

Warming is expected to stimulate soil microbial respiration triggering a positive soil carbon-climate feedback loop while a consensus remains elusive regarding the magnitude of this feedback. This is partly due to our limited understanding of the temperature-adaptive response of soil microbial respiration, especially over broad climatic scales. We used the square root (Ratkowsky) model to calculate the minimum temperature for soil microbial respiration (Tmin , which describes the temperature adaptation of soil microbial respiration) of 298 soil samples from alpine grasslands on the Tibetan Plateau and forest ecosystems across China with a mean annual temperature (MAT) range from -6°C to +25°C. The instantaneous soil microbial respiration was determined between 4°C and 28°C. The square root model could well fit the temperature effect on soil microbial respiration for each individual soil, with R2 higher than 0.98 for all soils. Tmin ranged from -8.1°C to -0.1°C and increased linearly with increasing MAT (R2  = 0.68). MAT dominantly regulated Tmin variation when accounting simultaneously for multiple other drivers (mean annual precipitation, soil pH and carbon quality); an independent experiment showed that carbon availability had no significant effect on Tmin . Using the relationship between Tmin and MAT, soil microbial respiration after an increased MAT could be estimated, resulting in a relative increase in respiration with decreasing MAT. Thus, soil microbial respiration responses are adapted to long-term temperature differences in MAT. We suggest that Tmin  = -5 + 0.2 × MAT, that is, every 1°C rise in MAT is estimated to increase Tmin of respiration by approximately 0.2°C, could be used as a first approximation to incorporate temperature adaptation of soil microbial respiration in model predictions. Our results can be used to predict future changes in the response of soil microbial respiration to temperature over different levels of warming and across broad geographic scales with different MAT.

Flooding variations affect soil bacterial communities at the spatial and inter-annual scales.

Hydrological variations have substantial effects on the diversity and composition of soil bacterial communities in wetlands. At the spatial scale, the responses of soil bacterial diversity and composition to hydrological variations in wetlands have been extensively investigated. However, at the temporal scale, especially at the inter-annual scale, the corresponding bacterial responses are rarely reported. Therefore, we explored the effects of flooding variations on the diversity and composition of soil bacterial communities at a lakeshore wetland in two hydrological contrasting years. Three flooding variables, i.e. flooding duration (FD), total duration of the growing season (TGD), and exposure duration of the growing season (EGD), were used to characterize flooding regime. Soil bacterial communities were determined using 16S rRNA gene sequencing method. We found a very high soil bacterial diversity at the lakeshore wetland. The Shannon's indexes of soil bacterial communities varied from 5.61 to 7.11 in two years. Soil bacterial α-diversity followed a unimodal curve along the elevation gradient, and was significantly lower in the flooding year than in the drought year. Principal coordinate analysis demonstrated that the compositions of soil bacterial communities were separated in order of elevation and year along the first and second axes, respectively. The apparent habitat preferences of soil bacterial families were closely connected with their respiratory traits, and this trend was stronger at the inter-annual scale than at the spatial scale. Soil bacterial compositions were predominantly determined by the direct (by changing respiratory traits) and indirect (by changing soil pH) effects of TGD at the spatial scale, while they were simultaneously regulated by the direct effects of three flooding variables at the inter-annual scale. Our results enhance the understanding of soil microbial communities in wetlands and have large implications for developing general theories to predicting soil microbial functions.

The thermal response of soil microbial methanogenesis decreases in magnitude with changing temperature.

Microbial methanogenesis in anaerobic soils contributes greatly to global methane (CH4) release, and understanding its response to temperature is fundamental to predicting the feedback between this potent greenhouse gas and climate change. A compensatory thermal response in microbial activity over time can reduce the response of respiratory carbon (C) release to temperature change, as shown for carbon dioxide (CO2) in aerobic soils. However, whether microbial methanogenesis also shows a compensatory response to temperature change remains unknown. Here, we used anaerobic wetland soils from the Greater Khingan Range and the Tibetan Plateau to investigate how 160 days of experimental warming (+4°C) and cooling (-4°C) affect the thermal response of microbial CH4 respiration and whether these responses correspond to changes in microbial community dynamics. The mass-specific CH4 respiration rates of methanogens decreased with warming and increased with cooling, suggesting that microbial methanogenesis exhibited compensatory responses to temperature changes. Furthermore, changes in the species composition of methanogenic community under warming and cooling largely explained the compensatory response in the soils. The stimulatory effect of climate warming on soil microbe-driven CH4 emissions may thus be smaller than that currently predicted, with important consequences for atmospheric CH4 concentrations.

Consumer regulation of the carbon cycle in coastal wetland ecosystems.

Despite escalating anthropogenic alteration of food webs, how the carbon cycle in ecosystems is regulated by food web processes remains poorly understood. We quantitatively synthesize the effects of consumers (herbivores, omnivores and carnivores) on the carbon cycle of coastal wetland ecosystems, 'blue carbon' ecosystems that store the greatest amount of carbon per unit area among all ecosystems. Our results reveal that consumers strongly affect many processes of the carbon cycle. Herbivores, for example, generally reduce carbon absorption and carbon stocks (e.g. aboveground plant carbon by 53% and aboveground net primary production by 23%) but may promote some carbon emission processes (e.g. litter decomposition by 32%). The average strengths of these effects are comparable with, or even times higher than, changes driven by temperature, precipitation, nitrogen input, CO2 concentration, and plant invasions. Furthermore, consumer effects appear to be stronger on aboveground than belowground carbon processes and vary markedly with trophic level, body size, thermal regulation strategy and feeding type. Despite important knowledge gaps, our results highlight the powerful impacts of consumers on the carbon cycle and call for the incorporation of consumer control into Earth system models that predict anthropogenic climate change and into management strategies of Earth's carbon stocks. This article is part of the theme issue 'Integrative research perspectives on marine conservation'.

Rising Temperature May Trigger Deep Soil Carbon Loss Across Forest Ecosystems.

Significantly more carbon (C) is stored in deep soil than in shallow horizons, yet how the decomposition of deep soil organic C (SOC) will respond to rising temperature remains unexplored on large scales, leading to considerable uncertainties to predictions of the magnitude and direction of C-cycle feedbacks to climate change. Herein, short-term temperature sensitivity of SOC decomposition (expressed as Q 10) from six depths within the top 1 m soil from 90 upland forest sites (540 soil samples) across China is reported. Results show that Q 10 significantly increases with soil depth, suggesting that deep SOC is more vulnerable to loss with rising temperature in comparison to shallow SOC. Climate is the primary regulator of shallow soil Q 10 but its relative influence declines with depth; in contrast, soil C quality has a minor influence on Q 10 in shallow soil but increases its influence with depth. When considering the depth-dependent Q 10 variations, results further show that using the thermal response of shallow soil layer for the whole soil profile, as is usually done in model predictions, would significantly underestimate soil C-climate feedbacks. The results highlight that Earth system models need to consider multilayer soil C dynamics and their controls to improve prediction accuracy.

Biogeographic variation in temperature sensitivity of decomposition in forest soils.

Determining soil carbon (C) responses to rising temperature is critical for projections of the feedbacks between terrestrial ecosystems, C cycle, and climate change. However, the direction and magnitude of this feedback remain highly uncertain due largely to our limited understanding of the spatial heterogeneity of soil C decomposition and its temperature sensitivity. Here we quantified C decomposition and its response to temperature change with an incubation study of soils from 203 sites across tropical to boreal forests in China spanning a wide range of latitudes (18°16' to 51°37'N) and longitudes (81°01' to 129°28'E). Mean annual temperature (MAT) and mean annual precipitation primarily explained the biogeographic variation in the decomposition rate and temperature sensitivity of soils: soil C decomposition rate decreased from warm and wet forests to cold and dry forests, while Q10-MAT (standardized to the MAT of each site) values displayed the opposite pattern. In contrast, biological factors (i.e. plant productivity and soil bacterial diversity) and soil factors (e.g. clay, pH, and C availability of microbial biomass C and dissolved organic C) played relatively small roles in the biogeographic patterns. Moreover, no significant relationship was found between Q10-MAT and soil C quality, challenging the current C quality-temperature hypothesis. Using a single, fixed Q10-MAT value (the mean across all forests), as is usually done in model predictions, would bias the estimated soil CO2 emissions at a temperature increase of 3.0°C. This would lead to overestimation of emissions in warm biomes, underestimation in cold biomes, and likely significant overestimation of overall C release from soil to the atmosphere. Our results highlight that climate-related biogeographic variation in soil C responses to temperature needs to be included in next-generation C cycle models to improve predictions of C-climate feedbacks.

Human eutrophication drives biogeographic salt marsh productivity patterns in China.

Salt marshes are important natural carbon sinks with a large capacity to absorb exogenous nutrient inputs. The effects of nutrients on biogeographic productivity patterns, however, have been poorly explored in salt marshes. We conducted field surveys to examine how complex environments affect productivity of two common salt marsh plants, invasive Spartina alterniflora and native Phragmites australis, along an 18,000-km latitudinal gradient on the Chinese coastline. We harvested peak aboveground biomass as a proxy for productivity, and measured leaf functional traits (e.g., leaf area, specific leaf area [SLA], leaf nitrogen [N] and phosphorus [P]), soil nutrients (dissolved inorganic N [DIN] and available P [AP]), and salinity. We compiled data on mean annual temperature (MAT) and exogenous nutrients (both N and P). Then, we examined how these abiotic factors affect salt marsh productivity using both linear mixed effect models and structural equation modeling. Using a trait-based approach, we also examined how salt marsh productivity responds to changing environments across latitude. Exogenous nutrients (both N and P), compared with temperature and other variables (e.g., DIN, AP, salinity), were the dominant factors in explaining the biogeographic productivity patterns of both S. alterniflora and P. australis. Leaf size-related traits (e.g., leaf area), rather than leaf economic traits (e.g., SLA, leaf N and P), can be used to indicate the positive effects of exogenous nutrients on the productivity of these two species. Our results demonstrated that human eutrophication surpassed temperature as the major driver of biogeographic salt marsh productivity pattern, challenging current models in which biogeographic productivity pattern is primarily controlled by temperature. Our findings have potential broad implications for the management of S. alterniflora, which is a global invader, as it has benefited from coastal eutrophication. Furthermore, exogenous nutrient availability and leaf size need to be integrated into earth system models that are used to predict global plant productivity in salt marshes.

A tumor-targeted immune checkpoint blocker.

To direct checkpoint inhibition to the tumor microenvironment, while avoiding systemic immune activation, we have synthesized a bispecific antibody [norleucine4, d-Phe7]-melanocyte stimulating hormone (NDP-MSH)-antiprogrammed cell death-ligand 1 antibody (αPD-L1) by conjugating a melanocyte stimulating hormone (α-MSH) analog to the antiprogrammed cell death-ligand 1 to (αPD-L1) antibody avelumab. This bispecific antibody can bind to both the melanocortin-1 receptor (MC1R) and to PD-L1 expressed on melanoma cells and shows enhanced specific antitumor efficacy in a syngeneic B16-SIY melanoma mouse model compared with the parental antibody at a 5 mg/kg dose. Moreover, the bispecific antibody showed increased infiltrated T cells in the tumor microenvironment. These results suggest that a tumor-targeted PD-L1-blocking bispecific antibody could have a therapeutic advantage in vivo, especially when used in combination with other checkpoint inhibitors.

The latitudinal herbivory hypothesis revisited: To be part is to be whole.

As the big data accumulation in ecology picks up pace, we now have the opportunity to test several macroecological hypotheses, such as the latitudinal herbivory hypothesis (LHH) dated from the 1990s. The LHH proposes that plant-herbivore interactions decrease as latitude increases, that is, from lower latitudinal areas (i.e., the equator) to higher latitudinal areas (i.e., the poles). This hypothesis has been challenged in recent years. In this study, we used the greatest volume dataset of leaf herbivory from the study of Zhang et al. (Journal of Ecology, 104, 2016, 1089) to test the LHH at a global scale, based on a quantile regression model. We found that the mean annual temperature, mean annual precipitation, and potential net primary production were heterogeneously correlated with herbivory at different quantiles or variable intervals. Although the Northern Hemisphere (NH) and the global-scale trends are in accordance with the expected latitudinal variation, the Southern Hemisphere (SH) was found to exhibit inverse trends. The latitude has a negative effect on plant-herbivore interactions in the NH and on a global scale; leaf herbivory decreased more at a given latitude in higher latitudinal areas, which is attributed to harsher survival conditions in these areas. The uniformity of leaf herbivory variability along the climate and latitude gradient in the NH and on a global scale motivates that the loosening of this herbivory variability in the SH is not significant enough to dismiss the prevalence of the LHH, a testable macroecology hypothesis.