Influence of multiple global change drivers on terrestrial carbon storage: additive effects are common

The interactive effects of multiple global change drivers on terrestrial carbon (C) storage remain poorly understood. Here, we synthesise data from 633 published studies to show how the interactive effects of multiple drivers are generally additive (i.e. not differing from the sum of their individual effects) rather than synergistic or antagonistic. We further show that (1) elevated CO₂, warming, N addition, P addition and increased rainfall, all exerted positive individual effects on plant C pools at both single‐plant and plant‐community levels; (2) plant C pool responses to individual or combined effects of multiple drivers are seldom scale‐dependent (i.e. not differing from single‐plant to plant‐community levels) and (3) soil and microbial biomass C pools are significantly less sensitive than plant C pools to individual or combined effects. We provide a quantitative basis for integrating additive effects of multiple global change drivers into future assessments of the C storage ability of terrestrial ecosystems.     

Interactive effects of global change factors on soil respiration and its components: a meta‐analysis

As the second largest carbon (C) flux between the atmosphere and terrestrial ecosystems, soil respiration (Rs) plays vital roles in regulating atmospheric CO₂ concentration ([CO₂]) and climatic dynamics in the earth system. Although numerous manipulative studies and a few meta‐analyses have been conducted to determine the responses of Rs and its two components [i.e., autotrophic (Ra) and heterotrophic (Rh) respiration] to single global change factors, the interactive effects of the multiple factors are still unclear. In this study, we performed a meta‐analysis of 150 multiple‐factor (≥2) studies to examine the main and interactive effects of global change factors on Rs and its two components. Our results showed that elevated [CO₂] (E), nitrogen addition (N), irrigation (I), and warming (W) induced significant increases in Rs by 28.6%, 8.8%, 9.7%, and 7.1%, respectively. The combined effects of the multiple factors, EN, EW, DE, IE, IN, IW, IEW, and DEW, were also significantly positive on Rs to a greater extent than those of the single‐factor ones. For all the individual studies, the additive interactions were predominant on Rs (90.6%) and its components (≈70.0%) relative to synergistic and antagonistic ones. However, the different combinations of global change factors (e.g., EN, NW, EW, IW) indicated that the three types of interactions were all important, with two combinations for synergistic effects, two for antagonistic, and five for additive when at least eight independent experiments were considered. In addition, the interactions of elevated [CO₂] and warming had opposite effects on Ra and Rh, suggesting that different processes may influence their responses to the multifactor interactions. Our study highlights the crucial importance of the interactive effects among the multiple factors on Rs and its components, which could inform regional and global models to assess the climate–biosphere feedbacks and improve predictions of the future states of the ecological and climate systems.     

不同降水条件下内蒙古荒漠草原主要植物物候对长期增温和氮添加的响应

植物物候是生态系统对气候变化响应的重要指示器,是植物生产力与植被动态模拟的重要参数。但是植物物候对全球变化的响应是否存在年际间变异、年内变异、物种间变异或生境间变异,以及如何改变,目前仍然不明确。该研究基于内蒙古荒漠草原长期增温和氮添加实验平台,选择优势植物短花针茅(Stipa breviflora)、冷蒿(Artemisia frigida)和木地肤(Kochia prostrata)为研究对象,使用物候打分观测方法和Richards生长曲线拟合方法,研究了实验处理第11、12和13年(2016–2018年)期间增温和氮添加对植物物候的影响。研究结果表明:(1)短花针茅开花时间集中在第129–145天,冷蒿开花时间集中在第230–248天,木地肤开花时间集中在第194–222天。增温、氮添加和增温+氮添加均使短花针茅和木地肤开花时间趋于提前,冷蒿开花时间趋于推迟。(2)短花针茅结果时间集中在第134–148天,冷蒿结果时间集中在第241–260天,木地肤结果时间集中在第207–231天。增温、氮添加和增温+氮添加处理均使短花针茅和木地肤结果时间趋于提前,冷蒿结果时间趋于推迟。(3)短...     

Nitrification,denitrification, and related functional genes under elevated CO2: A meta-analysis in terrestrial ecosystems

Global change may have profound effects on soil nitrogen (N) cycling that can induce positive feedback to climate change through increased nitrous oxide (N₂O) emissions mediated by nitrification and denitrification. We conducted a meta-analysis of the effects of elevated CO₂ on nitrification and denitrification based on 879 observations from 58 publications and 46 independent elevated CO₂ experiments in terrestrial ecosystems. We investigated the effects of elevated CO₂ alone or combined with elevated temperature, increased precipitation, drought, and N addition. We assessed the response to elevated CO₂ of gross and potential nitrification, potential denitrification, and abundances of related functional genes (archaeal amoA, bacterial amoA, nirK, nirS, and nosZ). Elevated CO₂ increased potential nitrification (+28%) and the abundance of bacterial amoA functional gene (+62%) in cropland ecosystems. Elevated CO₂ increased potential denitrification when combined with N addition and higher precipitation (+116%). Elevated CO₂ also increased the abundance of nirK (+25%) and nirS (+27%) functional genes in terrestrial ecosystems and of nosZ (+32%) functional gene in cropland ecosystems. The increase in the abundance of nosZ under elevated CO₂ was larger at elevated temperature and high N (+62%). Four out of 14 two-way interactions tested between elevated CO₂ and elevated temperature, elevated CO₂ and increased precipitation, and elevated CO₂ and N addition were marginally significant and mostly synergistic. The effects of elevated CO₂ on potential nitrification and abundances of bacterial amoA and nirS functional genes increased with mean annual temperature and mean annual precipitation. Our meta-analysis thus suggests that warming and increased precipitation in large areas of the world could reinforce positive responses of nitrification and denitrification to elevated CO₂ and urges the need for more investigations in the tropical zone and on interactive effects among multiple global change factors, as we may largely underestimate the effects of global change on soil N₂O emissions.     

Effects of livestock grazing on grassland carbon storage and release override impacts associated with global climate change

Predicting future carbon (C) dynamics in grassland ecosystems requires knowledge of how grazing and global climate change (e.g., warming, elevated CO₂, increased precipitation, drought, and N fertilization) interact to influence C storage and release. Here, we synthesized data from 223 grassland studies to quantify the individual and interactive effects of herbivores and climate change on ecosystem C pools and soil respiration (Rs). Our results showed that grazing overrode global climate change factors in regulating grassland C storage and release (i.e., Rs). Specifically, grazing significantly decreased aboveground plant C pool (APCP), belowground plant C pool (BPCP), soil C pool (SCP), and Rs by 19.1%, 6.4%, 3.1%, and 4.6%, respectively, while overall effects of all global climate change factors increased APCP, BPCP, and Rs by 6.5%, 15.3%, and 3.4% but had no significant effect on SCP. However, the combined effects of grazing with global climate change factors also significantly decreased APCP, SCP, and Rs by 4.0%, 4.7%, and 2.7%, respectively but had no effect on BPCP. Most of the interactions between grazing and global climate change factors on APCP, BPCP, SCP, and Rs were additive instead of synergistic or antagonistic. Our findings highlight the dominant effects of grazing on C storage and Rs when compared with the suite of global climate change factors. Therefore, incorporating the dominant effect of herbivore grazing into Earth System Models is necessary to accurately predict climate–grassland feedbacks in the Anthropocene.     

Effects of three global change drivers on terrestrial C:N:P stoichiometry: a global synthesis

Over the last few decades, there has been an increasing number of controlled‐manipulative experiments to investigate how plants and soils might respond to global change. These experiments typically examined the effects of each of three global change drivers [i.e., nitrogen (N) deposition, warming, and elevated CO₂] on primary productivity and on the biogeochemistry of carbon (C), N, and phosphorus (P) across different terrestrial ecosystems. Here, we capitalize on this large amount of information by performing a comprehensive meta‐analysis (>2000 case studies worldwide) to address how C:N:P stoichiometry of plants, soils, and soil microbial biomass might respond to individual vs. combined effects of the three global change drivers. Our results show that (i) individual effects of N addition and elevated CO₂ on C:N:P stoichiometry are stronger than warming, (ii) combined effects of pairs of global change drivers (e.g., N addition + elevated CO₂, warming + elevated CO₂) on C:N:P stoichiometry were generally weaker than the individual effects of each of these drivers, (iii) additive interactions (i.e., when combined effects are equal to or not significantly different from the sum of individual effects) were more common than synergistic or antagonistic interactions, (iv) C:N:P stoichiometry of soil and soil microbial biomass shows high homeostasis under global change manipulations, and (v) C:N:P responses to global change are strongly affected by ecosystem type, local climate, and experimental conditions. Our study is one of the first to compare individual vs. combined effects of the three global change drivers on terrestrial C:N:P ratios using a large set of data. To further improve our understanding of how ecosystems might respond to future global change, long‐term ecosystem‐scale studies testing multifactor effects on plants and soils are urgently required across different world regions.     

民勤荒漠区植物物候对气候变暖的响应

近几十年来,全球气候普遍变暖.那么,荒漠地区的气候是不是响应了全球气候的这种变化?在全球气候变化过程中,荒漠区植物物候又是如何响应这种气候变化的呢?显然,研究荒漠地区植物物候对气候变化的响应对于深入研究荒漠植物物候与气候因子的关系以及荒漠地区的植物保护都具有重要意义.运用位于中国西北典型荒漠地区的民勤沙生植物园1974～2007年42种中生、旱生植物的物候观测资料进行分析.结果表明:研究区1974年以来气温抬升幅度大于其他文献的研究报道,春季物候期提前幅度明显大于其他国家文献报道;在气温变暖的过程中,不同月份的气温变化与年平均气温的变化趋势并不完全对应,物候期发生当月的平均气温对该物候期的影响＞物候期发生上月平均气温＞年平均气温;研究区位于中国典型荒漠化地区,属于干旱荒漠气候,春季气温升高较其他地区更加明显,这就是当地春季物候期提前幅度相对较大的原因所在,也是当地以及中国西北沙区近几十年来沙尘暴天气增多和沙尘暴发生日期提前的原因.植物物候变化既是植物对气候变化的综合反应过程,又是植物适应气候变化的过程,尤其是荒漠植物.因此,物候研究将会成为今后气候学和植物生态学研究的一个重要内容.     

Flowering phenology in a species-rich temperate grassland is sensitive to warming but not elevated CO2

* Flowering is a critical stage in plant life cycles, and changes might alter processes at the species, community and ecosystem levels. Therefore, likely flowering-time responses to global change drivers are needed for predictions of global change impacts on natural and managed ecosystems. * Here, the impact of elevated atmospheric CO2 concentration ([CO2]) (550 micromol mol(-1)) and warming (+2 masculineC) is reported on flowering times in a native, species-rich, temperate grassland in Tasmania, Australia in both 2004 and 2005. * Elevated [CO2] did not affect average time of first flowering in either year, only affecting three out of 23 species. Warming reduced time to first flowering by an average of 19.1 d in 2004, acting on most species, but did not significantly alter flowering time in 2005, which might be related to the timing of rainfall. Elevated [CO2] and warming treatments did not interact on flowering time. * These results show elevated [CO2] did not alter average flowering time or duration in this grassland; neither did it alter the response to warming. Therefore, flowering phenology appears insensitive to increasing [CO2] in this ecosystem, although the response to warming varies between years but can be strong.     

Climate change effects on plant biomass alter dominance patterns and community evenness in an experimental old‐field ecosystem

Atmospheric and climatic change can alter plant biomass production and plant community composition. However, we know little about how climate change‐induced alterations in biomass production affect plant species composition. To better understand how climate change will alter both individual plant species and community biomass, we manipulated atmospheric [CO₂], air temperature, and precipitation in a constructed old‐field ecosystem. Specifically, we compared the responses of dominant and subdominant species to our climatic treatments, and explored how changes in plant dominance patterns alter community evenness over 2 years. Our study resulted in four major findings: (1) all treatments, elevated [CO₂], warming, and increased precipitation increased plant community biomass and the effects were additive rather than interactive, (2) plant species differed in their response to the treatments, resulting in shifts in the proportional biomass of individual species, which altered the plant community composition; however, the plant community response was largely driven by the positive precipitation response of Lespedeza, the most dominant species in the community, (3) precipitation explained most of the variation in plant community composition among treatments, and (4) changes in precipitation caused a shift in the dominant species proportional biomass that resulted in lower community evenness in the wet relative to dry treatments. Interestingly, compositional and evenness responses of the subdominant community to the treatments did not always follow the responses of the whole plant community. Our data suggest that changes in plant dominance patterns and community evenness are an important part of community responses to climatic change, and generally, that such compositional shifts can alter ecosystem biomass production and nutrient inputs.     

Simple additive simulation overestimates real influence: altered nitrogen and rainfall modulate the effect of warming on soil carbon fluxes

Experiments and models have led to a consensus that there is positive feedback between carbon (C) fluxes and climate warming. However, the effect of warming may be altered by regional and global changes in nitrogen (N) and rainfall levels, but the current understanding is limited. Through synthesizing global data on soil C pool, input and loss from experiments simulating N deposition, drought and increased precipitation, we quantified the responses of soil C fluxes and equilibrium to the three single factors and their interactions with warming. We found that warming slightly increased the soil C input and loss by 5% and 9%, respectively, but had no significant effect on the soil C pool. Nitrogen deposition alone increased the soil C input (+20%), but the interaction of warming and N deposition greatly increased the soil C input by 49%. Drought alone decreased the soil C input by 17%, while the interaction of warming and drought decreased the soil C input to a greater extent (?22%). Increased precipitation stimulated the soil C input by 15%, but the interaction of warming and increased precipitation had no significant effect on the soil C input. However, the soil C loss was not significantly affected by any of the interactions, although it was constrained by drought (?18%). These results implied that the positive C fluxes–climate warming feedback was modulated by the changing N and rainfall regimes. Further, we found that the additive effects of [warming × N deposition] and [warming × drought] on the soil C input and of [warming × increased precipitation] on the soil C loss were greater than their interactions, suggesting that simple additive simulation using single‐factor manipulations may overestimate the effects on soil C fluxes in the real world. Therefore, we propose that more multifactorial experiments should be considered in studying Earth systems.     

Gene expression profiling: opening the black box of plant ecosystem responses to global change

The use of genomic techniques to address ecological questions is emerging as the field of genomic ecology. Experimentation under environmentally realistic conditions to investigate the molecular response of plants to meaningful changes in growth conditions and ecological interactions is the defining feature of genomic ecology. Because the impact of global change factors on plant performance are mediated by direct effects at the molecular, biochemical, and physiological scales, gene expression analysis promises important advances in understanding factors that have previously been consigned to the 'black box' of unknown mechanism. Various tools and approaches are available for assessing gene expression in model and nonmodel species as part of global change biology studies. Each approach has its own unique advantages and constraints. A first generation of genomic ecology studies in managed ecosystems and mesocosms have provided a testbed for the approach and have begun to reveal how the experimental design and data analysis of gene expression studies can be tailored for use in an ecological context.     

Putative biotic drivers of plant phenology: With special reference to pathogens and deciduousness

Plant phenology is not only manifested in the seasonal timing of vegetative and reproductive processes but also has ontogenetic aspects. The adaptive basis of seasonal phenology has been considered mainly in terms of climatic drivers. However, some biotic factors as likely evolutionary influences on plants’ phenology appear to have been under‐researched. Several specific cases of putative biotic factors driving plant phenology are outlined, involving both herbivores and pathogens. These illustrate the diversity of likely interactions rather than any systematic coverage or review. Emphasis is on woody perennials, in which phenology is often most multifaceted and complicated by the ontogenetic aspect. The complete seasonal leaf fall that characterizes deciduous plants may be a very important defense against some pathogens. Whether biotic influences drive acquisition or long‐term persistence of deciduousness is considered. In one case, of leaf rusts in poplars, countervailing influences of the rusts and climate suggest persistence. Often, however, biotic and environmental influences likely reinforce each other. The timing and duration of shoot flushing may in at least some cases contribute to defenses against herbivores, largely through brief periods of “predator satiation” when plant tissues have highest food value. Wide re‐examination of plant phenology, accommodating the roles of biotic factors and their interplays with environments as additional adaptive drivers, is advocated toward developing and applying hypotheses that are observationally or experimentally testable.     

全球环境变化对森林凋落物分解的影响

全球环境变化将对森林生态系统凋落物的分解和养分循环产生直接和间接的多重影响.就全球环境变化如全球变暖、大气CO2浓度升高、UV-B辐射增强、氮沉降等对凋落物分解影响的研究进展进行了综合述评.影响凋落物分解的内部因素为凋落物基质质量,外部因素包括生物因素(微生物和动物)和非生物因素(温度、水分和土壤性质等).全球变暖对凋落物分解的非生物作用有正效应,也有负效应.全球变暖对凋落物化学组成虽然只有轻微的影响,但可以通过影响植被的物种组成来间接改变凋落物的产量、化学性质和分解.全球变暖对凋落物分解生物作用的主要影响是增强土壤微生物活性,从而加速凋落物的分解.CO2浓度上升将增加凋落物产量,并通过影响凋落物质量(提高C/N比、木质素/N比等)和生物环境(微生物的数量和活性)而影响分解过程.UV-B辐射和大气N沉降的增加亦对凋落物分解产生直接和间接的影响,但影响效果尚不很清楚,有待进一步的研究.总起来看,全球环境变化将通过影响凋落物的分解速率而对全球碳循环产生重要影响,但由于气候变化和凋落物分解响应的复杂性以及各因子之间的相互作用,气候变化对凋落物分解的总效应尚需更深入的研究来定量化.     

土壤微生物群落对全球气候变化响应的研究进展

全球气候变化对陆地生态系统过程和功能产生重要影响,土壤微生物群落在陆地生态系统几乎所有的生物地球化学循环过程起到关键作用。本文针对气候变化对土壤微生物的影响研究结果,主要从土壤微生物活性(土壤呼吸与酶活性)和微生物群落结构对大气CO₂升高、增温、降水变化、氮沉降等全球变化单因子和多因子的直接或间接响应进行综述,并进一步阐述参与土壤碳氮循环过程的功能微生物对气候变化的响应机制与适应规律。全球变化因子改变了土壤微生物的群落组成,呈现降低、增加和无影响3种效应,且不同功能微生物也呈现不同的敏感性。多个全球变化因子对土壤微生物群落结构的交互效应可能存在加性、协同、拮抗作用,产生加和的、相互促进或抵消的整体效果。然而,目前对多种全球变化因子如三因子或四因子的组合作用,以及多因子的高阶交互作用研究较少;已有的研究地理分布不均匀,且时间和空间大尺度的研究不足;缺乏综合生态系统模型对全球变化的影响进行模拟和预测。最后指出今后的研究发展方向：进行多种全球变化因子、长时间、多生态系统点位、大空间尺度的土壤微生物群落动态研究;探究多种全球变化因子的高阶交互作用;建立综合响应的生态系统模...     

Long-term responses of boreal vegetation to global change: an experimental and modelling investigation

The response of boreal ecosystems to future global change is an uncertain but potentially critical component of the feedback between the terrestrial biosphere and the atmosphere. To reduce some of the uncertainties in predicting the responses of this key ecosystem, the climate change experiment (CLIMEX) exposed an entire undisturbed catchment of boreal vegetation to CO₂ enrichment (560 ppmv) and climate change (+ 5 °C in winter, + 3 °C in summer) for three years (1994–96). This paper describes the leaf metabolic responses of the vegetation to the experimental treatment and model simulations of possible future changes in the hydrological and carbon balance of the site. Randomized intervention analysis of the leaf gas exchange measurements for the dominant species indicated Pinus sylvestris had significantly (P < 0.01) higher photosynthetic rates and Betula pubescens and Vaccinium myrtillus had significantly (P < 0.01) lower stomatal conductances after three years treatment compared to the controls. These responses led to sustained increases in leaf water-use efficiency of all species of trees and ground shrubs, as determined from carbon isotope analyses. Photosynthesis (A) vs. intercellular CO₂ (c_i) response curves (A/c_i responses), RuBisCo analysis and leaf nitrogen data together suggested none of the species investigated exhibited down-regulation in photosynthetic capacity. At the whole ecosystem level, the improved water economy of the plants did not translate into increased catchment runoff. Modelling simulations for the site indicate this was most likely brought about by a compensatory increase in evapotranspiration. In terms of the carbon budget of the site, the ecosystem model indicates that increased CO₂ and temperature would lead to boreal ecosystems of the type used in CLIMEX, and typical of much of southern Norway, acting as moderate net sinks for CO₂.     

Climate drivers alter nitrogen availability in surface peat and decouple N2 fixation from CH4 oxidation in the Sphagnum moss microbiome

Peat mosses (Sphagnum spp.) are keystone species in boreal peatlands, where they dominate net primary productivity and facilitate the accumulation of carbon in thick peat deposits. Sphagnum mosses harbor a diverse assemblage of microbial partners, including N₂-fixing (diazotrophic) and CH₄-oxidizing (methanotrophic) taxa that support ecosystem function by regulating transformations of carbon and nitrogen. Here, we investigate the response of the Sphagnum phytobiome (plant + constituent microbiome + environment) to a gradient of experimental warming (+0°C to +9°C) and elevated CO₂ (+500 ppm) in an ombrotrophic peatland in northern Minnesota (USA). By tracking changes in carbon (CH₄, CO₂) and nitrogen (NH₄-N) cycling from the belowground environment up to Sphagnum and its associated microbiome, we identified a series of cascading impacts to the Sphagnum phytobiome triggered by warming and elevated CO₂. Under ambient CO₂, warming increased plant-available NH₄-N in surface peat, excess N accumulated in Sphagnum tissue, and N₂ fixation activity decreased. Elevated CO₂ offset the effects of warming, disrupting the accumulation of N in peat and Sphagnum tissue. Methane concentrations in porewater increased with warming irrespective of CO₂ treatment, resulting in a ~10× rise in methanotrophic activity within Sphagnum from the +9°C enclosures. Warming's divergent impacts on diazotrophy and methanotrophy caused these processes to become decoupled at warmer temperatures, as evidenced by declining rates of methane-induced N₂ fixation and significant losses of keystone microbial taxa. In addition to changes in the Sphagnum microbiome, we observed ~94% mortality of Sphagnum between the +0°C and +9°C treatments, possibly due to the interactive effects of warming on N-availability and competition from vascular plant species. Collectively, these results highlight the vulnerability of the Sphagnum phytobiome to rising temperatures and atmospheric CO₂ concentrations, with significant implications for carbon and nitrogen cycling in boreal peatlands.     

A natural heating experiment: Phenotypic and genotypic responses of plant phenology to geothermal soil warming

Under global warming, the survival of many populations of sedentary organisms in seasonal environments will largely depend on their ability to cope with warming in situ by means of phenotypic plasticity or adaptive evolution. This is particularly true in high‐latitude environments, where current growing seasons are short, and expected temperature increases large. In such short‐growing season environments, the timing of growth and reproduction is critical to survival. Here, we use the unique setting provided by a natural geothermal soil warming gradient (Hengill geothermal area, Iceland) to study the response of Cerastium fontanum flowering phenology to temperature. We hypothesized that trait expression and phenotypic selection on flowering phenology are related to soil temperature, and tested the hypothesis that temperature‐driven differences in selection on phenology have resulted in genetic differentiation using a common garden experiment. In the field, phenology was related to soil temperature, with plants in warmer microsites flowering earlier than plants at colder microsites. In the common garden, plants responded to spring warming in a counter‐gradient fashion; plants originating from warmer microsites flowered relatively later than those originating from colder microsites. A likely explanation for this pattern is that plants from colder microsites have been selected to compensate for the shorter growing season by starting development at lower temperatures. However, in our study we did not find evidence of variation in phenotypic selection on phenology in relation to temperature, but selection consistently favoured early flowering. Our results show that soil temperature influences trait expression and suggest the existence of genetically based variation in flowering phenology leading to counter‐gradient local adaptation along a gradient of soil temperatures. An important implication of our results is that observed phenotypic responses of phenology to global warming might often be a combination of short‐term plastic responses and long‐term evolutionary responses, acting in different directions.