Diagnosing destabilization risk in global land carbon sinks.

Global net land carbon uptake or net biome production (NBP) has increased during recent decades1. Whether its temporal variability and autocorrelation have changed during this period, however, remains elusive, even though an increase in both could indicate an increased potential for a destabilized carbon sink2,3. Here, we investigate the trends and controls of net terrestrial carbon uptake and its temporal variability and autocorrelation from 1981 to 2018 using two atmospheric-inversion models, the amplitude of the seasonal cycle of atmospheric CO2 concentration derived from nine monitoring stations distributed across the Pacific Ocean and dynamic global vegetation models. We find that annual NBP and its interdecadal variability increased globally whereas temporal autocorrelation decreased. We observe a separation of regions characterized by increasingly variable NBP, associated with warm regions and increasingly variable temperatures, lower and weaker positive trends in NBP and regions where NBP became stronger and less variable. Plant species richness presented a concave-down parabolic spatial relationship with NBP and its variability at the global scale whereas nitrogen deposition generally increased NBP. Increasing temperature and its increasing variability appear as the most important drivers of declining and increasingly variable NBP. Our results show increasing variability of NBP regionally that can be mostly attributed to climate change and that may point to destabilization of the coupled carbon-climate system.

A trade-off between plant and soil carbon storage under elevated CO2.

Terrestrial ecosystems remove about 30 per cent of the carbon dioxide (CO2) emitted by human activities each year1, yet the persistence of this carbon sink depends partly on how plant biomass and soil organic carbon (SOC) stocks respond to future increases in atmospheric CO2 (refs. 2,3). Although plant biomass often increases in elevated CO2 (eCO2) experiments4-6, SOC has been observed to increase, remain unchanged or even decline7. The mechanisms that drive this variation across experiments remain poorly understood, creating uncertainty in climate projections8,9. Here we synthesized data from 108 eCO2 experiments and found that the effect of eCO2 on SOC stocks is best explained by a negative relationship with plant biomass: when plant biomass is strongly stimulated by eCO2, SOC storage declines; conversely, when biomass is weakly stimulated, SOC storage increases. This trade-off appears to be related to plant nutrient acquisition, in which plants increase their biomass by mining the soil for nutrients, which decreases SOC storage. We found that, overall, SOC stocks increase with eCO2 in grasslands (8 ± 2 per cent) but not in forests (0 ± 2 per cent), even though plant biomass in grasslands increase less (9 ± 3 per cent) than in forests (23 ± 2 per cent). Ecosystem models do not reproduce this trade-off, which implies that projections of SOC may need to be revised.

Exploring the synergy of enhanced weathering and Bacillus subtilis: A promising strategy for sustainable agriculture.

Climate change is one of the most urgent environmental challenges that humanity faces. In addition to the reduction of greenhouse gas emissions, safe and robust carbon dioxide removal (CDR) technologies that capture atmospheric CO2 and ensure long-term sequestration are required. Among CDR technologies, enhanced silicate weathering (ESW) has been suggested as a promising option. While ESW has been demonstrated to depend strongly on pH, water, and temperature, recent studies suggest that biota may accelerate mineral weathering rates. Bacillus subtilis is a plant growth-promoting rhizobacterium that can facilitate weathering to obtain mineral nutrients. It is a promising agricultural biofertilizer, as it helps plants acquire nutrients and protects them from environmental stresses. Given that croplands are optimal implementation fields for ESW, any synergy between ESW and B. subtilis can hold great potential for further practice. B. subtilis was reported to enhance weathering under laboratory conditions, but there is a lack of data for soil applications. In a soil-mesocosm experiment, we examined the effect of B. subtilis on basalt weathering. B. subtilis-basalt interaction stimulated basalt weathering and increased soil extractable Fe. The combined application displayed higher CDR potential compared to basalt-only application (3.7 vs. 2.3 tons CO2 ha-1) taking solid and liquid cation pools into account. However, the cumulative CO2 efflux decreased by approximately 2 tons CO2 ha-1 with basalt-only treatment, while the combined application did not affect the CO2 efflux. We found limited mobilization of cations to the liquid phase as most were retained in the soil. Additionally, we found substantial mobilization of basalt-originated Mg, Fe, and Al to oxide- and organic-bound soil fractions. We, therefore, conclude that basalt addition showed relatively low inorganic CDR potential but a high capacity for SOM stabilization. The outcomes indicated the importance of weathering rate-GHG emission integration and the high potential of SOM stabilization in ESW studies.

Enhanced silicate weathering accelerates forest carbon sequestration by stimulating the soil mineral carbon pump.

Enhanced silicate rock weathering (ERW) is an emerging strategy for carbon dioxide removal (CDR) from the atmosphere to mitigate anthropogenic climate change. ERW aims at promoting soil inorganic carbon sequestration by accelerating geochemical weathering processes. Theoretically, ERW may also impact soil organic carbon (SOC), the largest carbon pool in terrestrial ecosystems, but experimental evidence for this is largely lacking. Here, we conducted a 2-year field experiment in tropical rubber plantations in the southeast of China to evaluate the effects of wollastonite powder additions (0, 0.25, and 0.5 kg m-2) on both soil organic and inorganic carbon at 0-10 cm depth. We found that ERW significantly increased the concentration of SOC and HCO3 -, but the increases in SOC were four and eight times higher than that of HCO3 - with low- and high-level wollastonite applications. ERW had positive effects on the accrual of organic carbon in mineral-associated organic matter (MAOM) and macroaggregate fractions, but not on particulate organic matter. Path analysis suggested that ERW increased MAOM mainly by increasing the release of Ca, Si, and Fe, and to a lesser extent by stimulating root growth and microbial-derived carbon inputs. Our study indicates that ERW with wollastonite can promote SOC sequestration in stable MOAM in surface soils through both the soil mineral carbon pump and microbial carbon pump. These effects may have been larger than the inorganic CDR during our experiment. We argue it is essential to account for the responses of SOC in the assessments of CDR by ERW.

Decadal soil warming decreased vascular plant above and belowground production in a subarctic grassland by inducing nitrogen limitation.

Below and aboveground vegetation dynamics are crucial in understanding how climate warming may affect terrestrial ecosystem carbon cycling. In contrast to aboveground biomass, the response of belowground biomass to long-term warming has been poorly studied. Here, we characterized the impacts of decadal geothermal warming at two levels (on average +3.3°C and +7.9°C) on below and aboveground plant biomass stocks and production in a subarctic grassland. Soil warming did not change standing root biomass and even decreased fine root production and reduced aboveground biomass and production. Decadal soil warming also did not significantly alter the root-shoot ratio. The linear stepwise regression model suggested that following 10 yr of soil warming, temperature was no longer the direct driver of these responses, but losses of soil N were. Soil N losses, due to warming-induced decreases in organic matter and water retention capacity, were identified as key driver of the decreased above and belowground production. The reduction in fine root production was accompanied by thinner roots with increased specific root area. These results indicate that after a decade of soil warming, plant productivity in the studied subarctic grassland was affected by soil warming mainly by the reduction in soil N.

Consistent predictors of microbial community composition across spatial scales in grasslands reveal low context-dependency.

Environmental circumstances shaping soil microbial communities have been studied extensively. However, due to disparate study designs, it has been difficult to resolve whether a globally consistent set of predictors exists, or context-dependency prevails. Here, we used a network of 18 grassland sites (11 of those containing regional plant productivity gradients) to examine (i) if similar abiotic or biotic factors predict both large-scale (across sites) and regional-scale (within sites) patterns in bacterial and fungal community composition, and (ii) if microbial community composition differs consistently at two levels of regional plant productivity (low vs. high). Our results revealed that bacteria were associated with particular soil properties (such as base saturation) and both bacteria and fungi were associated with plant community composition across sites and within the majority of sites. Moreover, a discernible microbial community signal emerged, clearly distinguishing high and low-productivity soils across different grasslands independent of their location in the world. Hence, regional productivity differences may be typified by characteristic soil microbial communities across the grassland biome. These results could encourage future research aiming to predict the general effects of global changes on soil microbial community composition in grasslands and to discriminate fertile from infertile systems using generally applicable microbial indicators.

Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland.

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO2 fluxes and the fate of recently photosynthesized C through CO2 flux measurements and a 13 CO2 pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO2 emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO2 uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.

When things get MESI: The Manipulation Experiments Synthesis Initiative-A coordinated effort to synthesize terrestrial global change experiments.

Responses of the terrestrial biosphere to rapidly changing environmental conditions are a major source of uncertainty in climate projections. In an effort to reduce this uncertainty, a wide range of global change experiments have been conducted that mimic future conditions in terrestrial ecosystems, manipulating CO2 , temperature, and nutrient and water availability. Syntheses of results across experiments provide a more general sense of ecosystem responses to global change, and help to discern the influence of background conditions such as climate and vegetation type in determining global change responses. Several independent syntheses of published data have yielded distinct databases for specific objectives. Such parallel, uncoordinated initiatives carry the risk of producing redundant data collection efforts and have led to contrasting outcomes without clarifying the underlying reason for divergence. These problems could be avoided by creating a publicly available, updatable, curated database. Here, we report on a global effort to collect and curate 57,089 treatment responses across 3644 manipulation experiments at 1145 sites, simulating elevated CO2 , warming, nutrient addition, and precipitation changes. In the resulting Manipulation Experiments Synthesis Initiative (MESI) database, effects of experimental global change drivers on carbon and nutrient cycles are included, as well as ancillary data such as background climate, vegetation type, treatment magnitude, duration, and, unique to our database, measured soil properties. Our analysis of the database indicates that most experiments are short term (one or few growing seasons), conducted in the USA, Europe, or China, and that the most abundantly reported variable is aboveground biomass. We provide the most comprehensive multifactor global change database to date, enabling the research community to tackle open research questions, vital to global policymaking. The MESI database, freely accessible at doi.org/10.5281/zenodo.7153253, opens new avenues for model evaluation and synthesis-based understanding of how global change affects terrestrial biomes. We welcome contributions to the database on GitHub.

Is the climate change mitigation effect of enhanced silicate weathering governed by biological processes?

A number of negative emission technologies (NETs) have been proposed to actively remove CO2 from the atmosphere, with enhanced silicate weathering (ESW) as a relatively new NET with considerable climate change mitigation potential. Models calibrated to ESW rates in lab experiments estimate the global potential for inorganic carbon sequestration by ESW at about 0.5-5 Gt CO2  year-1 , suggesting ESW could be an important component of the future NETs mix. In real soils, however, weathering rates may differ strongly from lab conditions. Research on natural weathering has shown that biota such as plants, microbes, and macro-invertebrates can strongly affect weathering rates, but biotic effects were excluded from most ESW lab assessments. Moreover, ESW may alter soil organic carbon sequestration and greenhouse gas emissions by influencing physicochemical and biological processes, which holds the potential to perpetuate even larger negative emissions. Here, we argue that it is likely that the climate change mitigation effect of ESW will be governed by biological processes, emphasizing the need to put these processes on the agenda of this emerging research field.

Soil properties as key predictors of global grassland production: Have we overlooked micronutrients?

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

A hierarchical, multivariate meta-analysis approach to synthesising global change experiments.

Meta-analyses enable synthesis of results from globally distributed experiments to draw general conclusions about the impacts of global change factors on ecosystem function. Traditional meta-analyses, however, are challenged by the complexity and diversity of experimental results. We illustrate how several key issues can be addressed by a multivariate, hierarchical Bayesian meta-analysis (MHBM) approach applied to information extracted from published studies. We applied an MHBM to log-response ratios for aboveground biomass (AB, n = 300), belowground biomass (BB, n = 205) and soil CO2 exchange (SCE, n = 544), representing 100 studies. The MHBM accounted for study duration, climate effects and covariation among the AB, BB and SCE responses to elevated CO2 (eCO2 ) and/or warming. The MHBM revealed significant among-study covariation in the AB and BB responses to experimental treatments. The MHBM imputed missing duration (4.2%) and climate (6%) data, and revealed that climate context governs how eCO2 and warming impact ecosystem function. Predictions identified biomes that may be particularly sensitive to eCO2 or warming, but that are under-represented in global change experiments. The MHBM approach offers a flexible and powerful tool for synthesising disparate experimental results reported across multiple studies, sites and response variables.

Fertilized graminoids intensify negative drought effects on grassland productivity.

Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full-factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter-annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought-sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.

Initial soil community drives heathland fungal community trajectory over multiple years through altered plant-soil interactions.

Dispersal limitation, biotic interactions, and environmental filters interact to drive plant and fungal community assembly, but their combined effects are rarely investigated. This study examines how different heathland plant and fungal colonization scenarios realized via three biotic treatments - addition of mature heathland-derived sod, addition of hay, and no additions - affect soil fungal community development over 6 yr along a manipulated pH gradient in a large-scale experiment starting from an agricultural, topsoil removed state. Our results show that both biotic and abiotic (pH) treatments had a persistent influence on the development of fungal communities, but that sod additions diminished the effect of abiotic treatments through time. Analysis of correlation networks between soil fungi and plants suggests that the reduced effect of pH in the sod treatment, where both soil and plant propagules were added, might be due to plant-fungal interactions since the sod additions caused stronger, more specific, and more consistent connections compared with the no addition treatment. Based on these results, we suggest that the initial availability of heathland fungal and plant taxa, which reinforce each other, can significantly steer further fungal community development to an alternative configuration, overriding the otherwise prominent effect of abiotic (pH) conditions.

Phosphorus addition increased carbon partitioning to autotrophic respiration but not to biomass production in an experiment with Zea mays.

Plant carbon (C) partitioning-the relative use of photosynthates for biomass production, respiration, and other plant functions-is a key but poorly understood ecosystem process. In an experiment with Zea mays, with or without arbuscular mycorrhizal fungi (AMF), we investigated the effect of phosphorus (P) fertilization and AMF on plant C partitioning. Based on earlier studies, we expected C partitioning to biomass production (i.e., biomass production efficiency; BPE) to increase with increasing P addition due to reduced C partitioning to AMF. However, although plant growth was clearly stimulated by P addition, BPE did not increase. Instead, C partitioning to autotrophic respiration increased. These results contrasted with our expectations and with a previous experiment in the same set-up where P addition increased BPE while no effect on autotropic respiration was found. The comparison of both experiments suggests a key role for AMF in explaining these contrasts. Whereas in the previous experiment substantial C partitioning to AMF reduced BPE under low P, in the current experiment, C partitioning to AMF was too low to directly influence BPE. Our results illustrate the complex influence of nutrient availability and mycorrhizal symbiosis on plant C partitioning.

Towards comparable assessment of the soil nutrient status across scales-Review and development of nutrient metrics.

Nutrient availability influences virtually every aspect of an ecosystem, and is a critical modifier of ecosystem responses to global change. Although this crucial role of nutrient availability in regulating ecosystem structure and functioning has been widely acknowledged, nutrients are still often neglected in observational and experimental synthesis studies due to difficulties in comparing the nutrient status across sites. In the current study, we explain different nutrient-related concepts and discuss the potential of soil-, plant- and remote sensing-based metrics to compare the nutrient status across space. Based on our review and additional analyses on a dataset of European, managed temperate and boreal forests (ICP [International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests] Forests dataset), we conclude that the use of plant- and remote sensing-based metrics that rely on tissue stoichiometry is limited due to their strong dependence on species identity. The potential use of other plant-based metrics such as Ellenberg indicator values and plant-functional traits is also discussed. We conclude from our analyses and review that soil-based metrics have the highest potential for successful intersite comparison of the nutrient status. As an example, we used and adjusted a soil-based metric, previously developed for conifer forests across Sweden, against the same ICP Forests data. We suggest that this adjusted and further adaptable metric, which included the organic carbon concentration in the upper 20 cm of the soil (including the organic fermentation-humus [FH] layer), the C:N ratio and pHCaCl2 of the FH layer, can be used as a complementary tool along with other indicators of nutrient availability, to compare the background nutrient status across temperate and boreal forests dominated by spruce, pine or beech. Future collection and provision of harmonized soil data from observational and experimental sites is crucial for further testing and adjusting the metric.

Global vegetation biomass production efficiency constrained by models and observations.

Plants use only a fraction of their photosynthetically derived carbon for biomass production (BP). The biomass production efficiency (BPE), defined as the ratio of BP to photosynthesis, and its variation across and within vegetation types is poorly understood, which hinders our capacity to accurately estimate carbon turnover times and carbon sinks. Here, we present a new global estimation of BPE obtained by combining field measurements from 113 sites with 14 carbon cycle models. Our best estimate of global BPE is 0.41 ± 0.05, excluding cropland. The largest BPE is found in boreal forests (0.48 ± 0.06) and the lowest in tropical forests (0.40 ± 0.04). Carbon cycle models overestimate BPE, although models with carbon-nitrogen interactions tend to be more realistic. Using observation-based estimates of global photosynthesis, we quantify the global BP of non-cropland ecosystems of 41 ± 6 Pg C/year. This flux is less than net primary production as it does not contain carbon allocated to symbionts, used for exudates or volatile carbon compound emissions to the atmosphere. Our study reveals a positive bias of 24 ± 11% in the model-estimated BP (10 of 14 models). When correcting models for this bias while leaving modeled carbon turnover times unchanged, we found that the global ecosystem carbon storage change during the last century is decreased by 67% (or 58 Pg C).

Rainfall manipulation experiments as simulated by terrestrial biosphere models: Where do we stand?

Changes in rainfall amounts and patterns have been observed and are expected to continue in the near future with potentially significant ecological and societal consequences. Modelling vegetation responses to changes in rainfall is thus crucial to project water and carbon cycles in the future. In this study, we present the results of a new model-data intercomparison project, where we tested the ability of 10 terrestrial biosphere models to reproduce the observed sensitivity of ecosystem productivity to rainfall changes at 10 sites across the globe, in nine of which, rainfall exclusion and/or irrigation experiments had been performed. The key results are as follows: (a) Inter-model variation is generally large and model agreement varies with timescales. In severely water-limited sites, models only agree on the interannual variability of evapotranspiration and to a smaller extent on gross primary productivity. In more mesic sites, model agreement for both water and carbon fluxes is typically higher on fine (daily-monthly) timescales and reduces on longer (seasonal-annual) scales. (b) Models on average overestimate the relationship between ecosystem productivity and mean rainfall amounts across sites (in space) and have a low capacity in reproducing the temporal (interannual) sensitivity of vegetation productivity to annual rainfall at a given site, even though observation uncertainty is comparable to inter-model variability. (c) Most models reproduced the sign of the observed patterns in productivity changes in rainfall manipulation experiments but had a low capacity in reproducing the observed magnitude of productivity changes. Models better reproduced the observed productivity responses due to rainfall exclusion than addition. (d) All models attribute ecosystem productivity changes to the intensity of vegetation stress and peak leaf area, whereas the impact of the change in growing season length is negligible. The relative contribution of the peak leaf area and vegetation stress intensity was highly variable among models.

A novel method for assessing climate change impacts in ecotron experiments.

Ecotron facilities allow accurate control of many environmental variables coupled with extensive monitoring of ecosystem processes. They therefore require multivariate perturbation of climate variables, close to what is observed in the field and projections for the future. Here, we present a new method for creating realistic climate forcing for manipulation experiments and apply it to the UHasselt Ecotron experiment. The new methodology uses data derived from the best available regional climate model projection and consists of generating climate forcing along a gradient representative of increasingly high global mean air temperature anomalies. We first identified the best-performing regional climate model simulation for the ecotron site from the Coordinated Regional Downscaling Experiment in the European domain (EURO-CORDEX) ensemble based on two criteria: (i) highest skill compared to observations from a nearby weather station and (ii) representativeness of the multi-model mean in future projections. The time window is subsequently selected from the model projection for each ecotron unit based on the global mean air temperature of the driving global climate model. The ecotron units are forced with 3-hourly output from the projections of the 5-year period in which the global mean air temperature crosses the predefined values. With the new approach, Ecotron facilities become able to assess ecosystem responses on changing climatic conditions, while accounting for the co-variation between climatic variables and their projection in variability, well representing possible compound events. The presented methodology can also be applied to other manipulation experiments, aiming at investigating ecosystem responses to realistic future climate change.

Correction to: A novel method for assessing climate change impacts in ecotron experiments.

The article was published bearing a typographical error to the second author name listed. The author group regret the error and the name should be referenced and credited as Jakob Zscheischler and not the former.

Asymmetric effects of daytime and night-time warming on Northern Hemisphere vegetation.

Temperature data over the past five decades show faster warming of the global land surface during the night than during the day. This asymmetric warming is expected to affect carbon assimilation and consumption in plants, because photosynthesis in most plants occurs during daytime and is more sensitive to the maximum daily temperature, Tmax, whereas plant respiration occurs throughout the day and is therefore influenced by both Tmax and the minimum daily temperature, Tmin. Most studies of the response of terrestrial ecosystems to climate warming, however, ignore this asymmetric forcing effect on vegetation growth and carbon dioxide (CO2) fluxes. Here we analyse the interannual covariations of the satellite-derived normalized difference vegetation index (NDVI, an indicator of vegetation greenness) with Tmax and Tmin over the Northern Hemisphere. After removing the correlation between Tmax and Tmin, we find that the partial correlation between Tmax and NDVI is positive in most wet and cool ecosystems over boreal regions, but negative in dry temperate regions. In contrast, the partial correlation between Tmin and NDVI is negative in boreal regions, and exhibits a more complex behaviour in dry temperate regions. We detect similar patterns in terrestrial net CO2 exchange maps obtained from a global atmospheric inversion model. Additional analysis of the long-term atmospheric CO2 concentration record of the station Point Barrow in Alaska suggests that the peak-to-peak amplitude of CO2 increased by 23 ± 11% for a +1 °C anomaly in Tmax from May to September over lands north of 51° N, but decreased by 28 ± 14% for a +1 °C anomaly in Tmin. These lines of evidence suggest that asymmetric diurnal warming, a process that is currently not taken into account in many global carbon cycle models, leads to a divergent response of Northern Hemisphere vegetation growth and carbon sequestration to rising temperatures.