放牧对草地生态系统影响的研究进展

全球草地占据30%左右的陆地面积, 在全球气候变化、碳氮及养分循环、保持水土、调节畜牧业生产等方面具有重要的作用。目前草地的主要利用方式之一就是放牧, 不同的牲畜种类、放牧强度、年限、历史和制度等, 会影响草地植物群落、生物多样性及土壤微生物, 进而影响草地生态系统结构、功能和过程。该文围绕放牧对草地生态系统结构、功能和过程的影响, 1)回顾了20世纪50年代到现在各个历史阶段放牧对草地生态系统影响的研究; 2)利用文献计量分析的方法, 剖析了放牧对草地影响研究的热点内容、重要区域和关键词等; 3)阐明了放牧对草地植物生长、群落特征、碳氮及养分循环、生产力及土壤质量等的各方面影响的研究进展及国内相关研究的优势及存在的主要问题和不足; 4)基于上述分析, 从草地放牧精准管理、经典假说验证、放牧和全球变化研究相结合等方面, 提出未来研究的前沿方向和优先领域。该文在系统总结放牧对草地生态系统影响的研究进展、研究优势及存在问题的基础上, 提出未来的研究应与全球变化相结合, 为我国的草地放牧生态学研究、适应性管理和可持续利用等提供科学基础。     

Advances in the effect of nitrogen deposition on grassland litter decomposition

Atmospheric nitrogen deposition has increased in the last several decades due to anthropogenic activities and global changes. Increasing nitrogen deposition has become an important factor regulating carbon cycle in grassland ecosystems. Litter decomposition, a key process of carbon and nutrient cycling in terrestrial ecosystems, is the main source of soil carbon pool and the basis of soil fertility maintenance. Elevated nitrogen deposition could affect litter decomposition by raising soil nitrogen availability, increasing the quantity and quality of litter inputs, and altering soil microorganism and soil conditions. Litter decomposition are complex biological, physical and chemical processes, which were affected by abiotic, biological factors and their interactions. The effects of nitrogen deposition on litter decomposition and the underlying mechanisms were discussed in this paper, including the aspactes of soil nitrogen availability, litter production, litter quality, microclimate, soil microorganism and enzyme activities. The main research contents, directions, methods and existing problems of litter decomposition in grasslands were discussed. We also discussed the prospect of future directions to study the interaction and feedback between nitrogen deposition and grassland ecosystem carbon cycling process.     

青藏高原高寒草地生物多样性与生态系统功能的关系

生物多样性和生态系统功能(BEF)之间的关系是目前陆地生态系统生态学研究的热点, 对于生态系统的高效利用与管理意义重大, 而且对于退化生态系统功能的恢复及生物多样性的保护有重要的指导作用。高寒草地是青藏高原生态系统的主体, 近年来, 在气候变化与人为干扰等因素的驱动下, 高寒草地生态系统功能严重衰退。为此, 本文在综述物种多样性和生态系统功能及其相互关系研究进展的基础上, 首先从地下生态学过程研究、全球变化对生态系统多功能性的影响等方面解析了目前关于草地生物多样性和生态系统功能研究中存在的问题。继而, 从不同草地类型、草地退化程度、放牧、模拟气候变化、刈割、施肥、封育和补播等干扰利用方式对高寒草地物种多样性与生态系统功能的影响进行了全面的评述。并指出了高寒草地BEF研究中存在的不足, 今后应基于物种功能多样性开展高寒草地BEF研究, 全面且综合地考虑非生物因子(养分资源、外界干扰、环境波动等)对生物多样性与生态系统功能之间关系的影响, 关注尺度效应和要素耦合在全球气候变化对高寒草地BEF研究中的作用。最后, 以高寒草地BEF研究进展和结论为支撑依据, 综合提出了高寒草地资源利用和生物多样性保护的措施与建议: 加强放牧管理, 保护生物多样性; 治理退化草地, 维持生物多样性功能; 加强创新保护理念, 增强生态系统功能。     

增温和放牧对草地土壤和生态系统呼吸的影响

草地生态系统作为世界陆地生态系统的主体类型,其土壤呼吸和生态系统呼吸是陆地生态系统碳循环的重要组成部分,土壤呼吸是未经扰动的土壤由于代谢活动而产生CO2的过程,生态系统呼吸包括地下部分的土壤呼吸和地上部分植被的自养呼吸。研究增温和放牧对草地土壤和生态系统呼吸的影响,可为预测未来气候变化条件下的全球碳收支以及草地的可持续经营与管理提供重要的科学依据。该文扼要综述了关于草地土壤和生态系统呼吸对增温和放牧的响应方面的研究。结果表明:草地土壤和生态系统呼吸对增温和放牧的响应非常复杂,受多种因素的综合影响,无论是增温还是放牧对草地土壤和生态系统呼吸的影响均具有不确定性,因草地类型、增温幅度、增温时间、放牧强度、放牧频度和放牧方式的不同而不同。在此基础上,指出了以后应加强研究的方向,草地的利用离不开放牧,对于未来气候变化条件下的草地,温度升高和放牧这两个因素必然是同时存在的,以前多数实验是单独研究增温或放牧对它们的影响,然而,这两者对草地生态系统的影响并非可加的,因此,需要加强增温与放牧的耦合试验,同时加强关于生态系统呼吸不同组分对两者的响应的研究,以便更好地理解增温和放牧的影响机制。另外,草地土壤和生态系统呼吸对增温和放牧的响应会随着时间的推移而发生变化,因而加强长期连续的试验观测很有必要。     

Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta‐analysis

Livestock grazing activities potentially alter ecosystem carbon (C) and nitrogen (N) cycles in grassland ecosystems. Despite the fact that numerous individual studies and a few meta‐analyses had been conducted, how grazing, especially its intensity, affects belowground C and N cycling in grasslands remains unclear. In this study, we performed a comprehensive meta‐analysis of 115 published studies to examine the responses of 19 variables associated with belowground C and N cycling to livestock grazing in global grasslands. Our results showed that, on average, grazing significantly decreased belowground C and N pools in grassland ecosystems, with the largest decreases in microbial biomass C and N (21.62% and 24.40%, respectively). In contrast, belowground fluxes, including soil respiration, soil net N mineralization and soil N nitrification increased by 4.25%, 34.67% and 25.87%, respectively, in grazed grasslands compared to ungrazed ones. More importantly, grazing intensity significantly affected the magnitude (even direction) of changes in the majority of the assessed belowground C and N pools and fluxes, and C : N ratio as well as soil moisture. Specifically,light grazing contributed to soil C and N sequestration whereas moderate and heavy grazing significantly increased C and N losses. In addition, soil depth, livestock type and climatic conditions influenced the responses of selected variables to livestock grazing to some degree. Our findings highlight the importance of the effects of grazing intensity on belowground C and N cycling, which may need to be incorporated into regional and global models for predicting effects of human disturbance on global grasslands and assessing the climate‐biosphere feedbacks.     

放牧对草地生态系统服务和功能权衡关系的影响

内蒙古草原是我国北方的重要生态屏障和绿色畜牧业基地, 放牧是草原生态系统的主要利用和管理方式, 在放牧管理中充分发挥生态系统某一项或几项服务和功能最大利用价值时, 往往会与其他服务(功能)发生冲突, 需要权衡多项生态系统服务和功能, 制定合理的放牧管理制度。该研究以内蒙古锡林郭勒典型草原为例, 通过设置不放牧、轻度放牧、中度放牧以及重度放牧的放牧梯度, 从多项生态系统服务和功能权衡的角度比较了最适放牧管理强度。结果显示, 在放牧管理的草地生态系统服务和功能的权衡中, 权衡、协同、不相关关系同时存在, 如土壤呼吸速率与植物群落净生长量、生物多样性与植物群落净光合速率表现为权衡关系, 植物群落净生长量分别与土壤含水量、植物群落净光合速率及草地蒸散速率存在协同关系, 土壤有机碳含量与其他服务或功能间呈不相关关系; 放牧能不同程度地削弱多项生态系统服务及功能间的权衡关系(冲突对立关系); 中度放牧条件下的多项生态系统服务及功能的协同性最佳。     

放牧牲畜粪便降解及其对草地土壤养分动态的影响研究进展

放牧牲畜粪便沉积是影响草地土壤养分动态的重要途径之一,粪便降解过程调控着其养分返还效率,从而可能对草地土壤养分平衡和植被生长的养分供应等产生重要影响。针对放牧牲畜排粪行为特性、牲畜粪便的物质组成及其降解过程、以及粪便养分归还对土壤养分动态的影响等进行了系统论述,阐明了牲畜粪便降解与其养分迁移转化的关系,以及粪便养分输入对放牧草地生态系统养分生物地球化学循环的影响效应和可能的作用机制,以期为加深对牲畜粪便降解的养分动态变化过程的认知和厘清粪便-植物-土壤体系养分迁移和转化的影响机制积累理论基础,进而为优化牲畜粪便管理模式、维持土壤养分平衡和促进草地生态系统的健康协调和可持续发展提供科学依据。     

干旱半干旱草地生态系统与土壤水分关系研究进展

研究干旱半干旱草地生态系统与土壤水分关系和相互作用机理对于揭示草地生态系统稳定性及其水土关键要素的变化过程具有重要意义。从不同界面、不同尺度综述了草地生态系统对土壤水分的影响及草地生态系统的响应与适应机制,总结了草地生态系统与土壤水分关系模型研究的相关进展,并分析了气候变化对草地生态系统和土壤水分关系的影响。草地生态系统通过影响水文过程和生态过程来影响土壤水分,土壤水分在植物生长发育、形态、生理生态过程、种间关系、群落组成和结构以及草地生态系统功能等方面对草地生态系统产生影响;充分揭示草地生态系统-土壤水分相互作用机理是模型研究的关键;气候变化对草地生态系统植物与土壤水分关系具有重要影响。今后应加强以下研究:1)开展草地不同优势种和植物功能型与土壤水分关系的研究,找出能反映植物对土壤水分响应的性状指标,阈值响应点及适应机制;2)注重对不同时间和空间尺度上的转换和比较;3)加强个体、群体和生态系统尺度草地植物生长模型的研究及其与土壤-植被-大气水分传输模型的耦合;4)加强草地生态系统与土壤水分关系对气候变化响应的研究。     

Response and adaptation of terrestrial ecosystem processes to climate warming

陆地生态系统包含一系列时空连续、尺度多元且互相联系的生态学过程。由于大部分生态学过程都受到温度调控, 因此气候变暖会对全球陆地生态系统产生深远的影响。近年来, 全球变化生态学的基本科学问题之一是陆地生态系统的关键过程如何响应与适应全球气候变暖。围绕该问题, 该文梳理了近年来的研究进展, 重点关注植物生理生态过程、物候期、群落动态、生产力及其分配、凋落物与土壤有机质分解、养分循环等过程对温度升高的响应与适应机理。通过定量分析近20年来发表于主流期刊的相关论文, 展望了该领域的前沿方向, 包括物种性状对生态系统过程的预测能力, 生物地球化学循环的耦合过程, 极端高温与低温事件的响应与适应机理, 不对称气候变暖的影响机理和基于过程的生态系统模拟预测等。基于这些研究进展, 该文建议进一步研究陆地生态系统如何适应气候变暖, 更多关注我国的特色生态系统类型, 并整合实验、观测或模型等研究手段开展跨尺度的合作研究。     

Effects of grazing intensity and grazing exclusion on litter decomposition in the temperate steppe of Nei Mongol,China

Aims Grazing intensity and grazing exclusion affect ecosystem carbon cycling by changing the plant community and soil micro-environment in grassland ecosystems. The aims of this study were: 1) to determine the effects of grazing intensity and grazing exclusion on litter decomposition in the temperate grasslands of Nei Mongol; 2) to compare the difference between above-ground and below-ground litter decomposition; 3) to identify the effects of precipitation on litter production and decomposition. Methods We measured litter production, quality, decomposition rates and soil nutrient contents during the growing season in 2011 and 2012 in four plots, i.e. light grazing, heavy grazing, light grazing exclusion and heavy grazing exclusion. Quadrate surveys and litter bags were used to measure litter production and decomposition rates. All data were analyzed with ANOVA and Pearson’s correlation procedures in SPSS. Important findings Litter production and decomposition rates differed greatly among four plots. During the two years of our study, above-ground litter production and decomposition in heavy-grazing plots were faster than those in light-grazing plots. In the dry year, below-ground litter production and decomposition in light-grazing plots were faster than those in heavy-grazing plots, which is opposite to the findings in the wet year. Short-term grazing exclusion could promote litter production, and the exclusion of light-grazing could increase litter decomposition and nutrient cycling. In contrast, heavy-grazing exclusion decreased litter decomposition. Thus, grazing exclusion is beneficial to the restoration of the light-grazing grasslands, and more human management measures are needed during the restoration of heavy-grazing grasslands. Precipitation increased litter production and decomposition, and below-ground litter was more vulnerable to the inter-annual change of precipitation than above-ground litter. Compared to the light-grazing grasslands, heavy-grazing grasslands had higher sensitivity to precipitation. The above-ground litter decomposition was strongly positively correlated with the litter N content (R² = 0.489, p < 0.01) and strongly negatively correlated with the soil total N content (R² = 0.450, p < 0.01), but it was not significantly correlated with C:N and lignin:N. Below-ground litter decomposition was negatively correlated with the litter C (R² = 0.263, p < 0.01), C:N (R² = 0.349, p < 0.01) and cellulose content (R² = 0.460, p < 0.01). Our results will provide a theoretical basis for ecosystem restoration and the research of carbon cycling.     

Responses of freshwater ecosystems to global change: research progress and outlook

全球变化已经通过提高水温、改变降水格局和水流状况、促进物种入侵、增加极端事件, 对不同的淡水生态系统造成严重的威胁。该文将全球变化背景下淡水生态学的主要研究内容归纳为: (1)全球变化各要素对个体、种群、群落及至生态系统水平的影响; (2)全球变化过程中生态系统生物地球化学循环的改变; (3)淡水生态系统对全球变化的适应对策。最近10-15年淡水生态系统与全球变化研究快速发展, 取得的重要突破有: (1)阐明淡水生态系统结构与功能对全球气候变化尤其是水温升高的响应过程与机制; (2)揭示淡水生态系统(湿地、湖泊、河流等)是全球碳循环的重要组成部分, 在全球变化因素的影响下呈现有机碳埋藏减少和矿化速率提高。今后的研究中, 需要进一步加强对淡水生态系统全要素的系统观测与整合; 开展以“河流”为介质耦合多系统的碳输运和转化过程研究; 强化基础理论研究揭示淡水生态系统对全球变化的适应机制。     

Functional substitutability of native herbivores by livestock for soil carbon stock is mediated by microbial decomposers

Grazing by large mammalian herbivores impacts climate as it can favor the size and stability of a large carbon (C) pool in the soils of grazing ecosystems. As native herbivores in the world's grasslands, steppes, and savannas are progressively being displaced by livestock, it is important to ask whether livestock can emulate the functional roles of their native counterparts. While livestock and native herbivores can have remarkable similarity in their traits, they can differ greatly in their impacts on vegetation composition which can affect soil-C. It is uncertain how these similarities and differences impact soil-C via their influence on microbial decomposers. We test competing alternative hypotheses with a replicated, long-term, landscape-level, grazing-exclusion experiment to ask whether livestock in the Trans-Himalayan ecosystem of northern India can match decadal-scale (2005–2016) soil-C stocks under native herbivores. We evaluate multiple lines of evidence from 17 variables that influence soil-C (quantity and quality of C-input from plants, microbial biomass and metabolism, microbial community composition, eDNA, veterinary antibiotics in soil), and assess their inter-relationships. Livestock and native herbivores differed in their effects on several soil microbial processes. Microbial carbon use efficiency (CUE) was 19% lower in soils under livestock. Compared to native herbivores, areas used by livestock contained 1.5 kg C m⁻² less soil-C. Structural equation models showed that alongside the effects arising from plants, livestock alter soil microbial communities which is detrimental for CUE, and ultimately also for soil-C. Supporting evidence pointed toward a link between veterinary antibiotics used on livestock, microbial communities, and soil-C. Overcoming the challenges of sequestering antibiotics to minimize their potential impacts on climate, alongside microbial rewilding under livestock, may reconcile the conflicting demands from food-security and ecosystem services. Conservation of native herbivores and alternative management of livestock is crucial for soil-C stewardship to envision and achieve natural climate solutions.     

极端降水和极端干旱事件对草原生态系统的影响

当前人类活动的加剧显著地影响着全球大气循环的格局。大气循环的多个模型均预测未来全球气候变化的显著特征是极端降水事件和极端干旱事件发生的频率会显著增加。水分是干旱、半干旱区草原植物生长发育的限制性资源, 而草原生态系统是陆地生态系统中对降水格局变化非常敏感的系统。但是, 关于极端降水事件和极端干旱事件对草原生态系统结构和功能的影响还是以分散的个案研究为主, 甚至关于极端气候事件的定义迄今也不尽相同。为此, 该文在分析极端气候事件定义及其研究方法的基础上, 总结了极端降水事件和极端干旱事件对草原生态系统土壤水分和养分状况、植物生长发育和生理特性、群落结构、生产力和碳循环过程的影响, 并提出了未来极端气候事件研究中应重点关注的5个重要方向, 以及控制试验研究的2个关键科学问题, 对开展全球变化背景下草原生态系统对极端气候事件响应机制的研究具有指导意义。     

大气氮沉降影响草地植物物种多样性机制研究综述

大气氮沉降对草地生态系统结构和功能的影响已成为全球变化生物学研究重点。大气氮沉降导致草地群落物种多样性降低已成为全球普遍现象,但其生物学机制还不清楚,因此有必要系统梳理大气氮沉降对全球不同草地生态系统的研究结果,以便在氮沉降背景下为我国草地生态系统的研究和管理制定科学决策。系统综述了氮沉降降低草地群落物种多样性的可能机制,主要包括资源竞争排斥、群落更新限制、土壤酸化及其离子毒害、养分失衡、氮素本身的毒害、次生胁迫。氮沉降导致草地物种多样性降低是多种机制综合作用的结果,每种机制在不同时空具有不同的相对贡献。同时,与欧洲酸性土壤草地和美国高草草原相比,我国草地土壤类型和植被属性具有明显差异。因此,应根据我国草地生态系统的特征、不同植物功能利用养分策略,从土壤养分变化、根系养分吸收转运、叶片生理过程等方面的整合研究思路,探讨氮沉降影响我国草地群落物种多样性的生物学机制,为我国草地生态系统的科学管理提供理论依据。     

放牧对呼伦贝尔草地植物和土壤生态化学计量学特征的影响

放牧通过畜体采食、践踏和排泄物归还影响草地群落组成、植物形态和土壤养分,植物通过改变养分利用策略适应环境变化。通过分析呼伦贝尔草原放牧和围封样地中的群落植物和土壤的碳氮磷养分及化学计量比,探讨放牧对生态系统化学计量学特征和养分循环速率的影响机制。结果如下:(1)群落尺度上,放牧和围封草地植物叶片C、N和P的含量没有显著差异;但是在种群尺度上,放牧草地植物叶片N含量显著高于围封草地;(2)放牧草地土壤全C、全N、有机C、速效P含量,低于围封草地,硝态N含量高于围封草地;土壤全P和铵态N指标没有显著差异;(3)放牧草地植物C∶N比显著低于围封草地,植物残体分解速率较快,提高了生态系统养分循环速率。     

Impacts of elevated carbon dioxide concentration on terrestrial ecosystems: problems and prospective

理解生态系统对过去、现在和未来CO₂浓度变化的响应,对于在生态进化的时间尺度上认识和预测全球变化的后果至关重要。过去三十多年来CO₂浓度升高相关的科学问题主要集中在对植物生长和生产力的影响, 碳氮周转, 生态系统渐进式氮限制(PNL)形成, 与其他胁迫因子(O₃污染、氮沉降、升温、干旱)之间的交互作用等方面。尽管生态学家在数据累积、基础理论上取得了一定进展, 但是仍然存在较大不确定性和大量未知有待解决。该文探究了近30年来CO₂浓度升高对陆地生态系统影响研究的国际研究进展、重点领域及热点, 回顾了CO₂浓度升高对植物影响的模拟实验研究发展, 重点论述了CO₂浓度升高对粮食产量及品质、碳固定、水分利用效率、生态系统氮利用和土壤微生物响应等国际前沿动态研究中存在的主要问题与不足, 在此基础上展望了未来研究中值得关注的前沿研究方向。     

多年放牧对不同类型草原植被及土壤碳同位素的影响

放牧是草原牧区常见的人类活动,多年放牧对草原植被及土壤的碳过程产生较大的影响.本研究采集不同类型草原多年放牧前后植被及土壤样品,对室内碳同位素进行分析,研究了不同草原生态系统Δ¹³C(碳同位素分馏值)差异及其影响因素.结果表明: 放牧强度对植被Δ¹³C值的影响显著,0～5 cm表层土壤Δ¹³C值在放牧前后变化显著,而对深层土壤(>5 cm)影响不显著.多年放牧后大部分植被Δ¹³C值显著升高,高海拔地区升高的幅度较大.可见,放牧行为对不同草地生态系统类型、不同土壤深度以及不同海拔生态系统碳过程产生的影响差异显著.针对不同类型的草原,放牧应采取多样化的管理方式.     

放牧对全球草地生态系统碳氮磷化学计量特征影响的Meta分析

为明确全球尺度下放牧管理措施对草地生态系统碳(C)、氮(N)、磷(P)化学计量特征的影响,提高草地生态系统管理水平,本研究选取国内外83篇中英文文献进行Meta分析,并通过亚组分析探讨了放牧家畜组合(羊单牧、牛单牧和牛羊混牧)和放牧强度(轻度、中度、重度)对草地生态系统叶片、凋落物、根系,以及土壤C、N、P化学计量特征的影响。结果表明: 放牧会显著降低叶片和凋落物C含量、C/N、C/P,增加N、P含量及N/P;显著降低根系和土壤C、N含量,C/P和N/P,增加P含量和C/N。叶片、凋落物化学计量特征变化对牛、羊单独放牧响应更为明显,而根系、土壤化学计量特征变化则对混牧响应更为明显,重度放牧会对草地生态系统化学计量特征产生更大的影响。放牧会降低土壤N含量,增加P含量,表明放牧对草地N、P含量的影响路径不同。进一步研究N、P含量变化对放牧活动不平衡响应机制,将放牧方式、强度的影响纳入草地生态系统预测、管理模型,能够有效提高草地生态系统管理水平。     

放牧对草地土壤氮素循环关键过程的影响与机制研究进展

放牧是人类管理利用草地生态系统的最主要途径之一.食草动物的采食、践踏、卧息和排泄物归还等干扰不仅会改变草地地上植物群落,也会对土壤养分循环产生显著的影响.随着人类活动的加剧,放牧强度和频率也在逐渐增加,从而对草地土壤氮素循环关键过程产生重要影响.放牧主要通过改变土壤的物理性质、土壤氮库以及微生物的组成和结构,进而影响氮素在土壤中的迁移与转化.适度放牧会促进土壤氮素的矿化过程和硝化过程,加快氮素的周转,有利于植物吸收可利用氮素,而对于反硝化的影响与草地的水热条件和土壤类型等密切相关.目前,关于放牧强度对土壤氮素循环关键过程影响的研究结果不一致,其影响机制尚不明晰,尤其对于不同类型的草原仍存在较大的差异.本研究在大量查阅国内外已有研究结果的基础上,论述了放牧对土壤氮素循环关键过程的影响效应,总结了放牧对土壤氮素循环的影响机制,指出了目前研究过程中存在的不足,并对未来研究中值得重点关注和深入研究的科学问题进行了探讨与展望,为进一步理解放牧对草地土壤氮素循环的影响提供参考.     

Spatial patterns of soil and ecosystem respiration regulated by biological and environmental variables along a precipitation gradient in semi-arid grasslands in China

Precipitation is a key environmental factor in determining ecosystem structure and function. Knowledge of how soil and ecosystem respiration responds to climate change (e.g., precipitation) and human activities (e.g., grazing or clipping) is crucial for assessing the impacts of climate change on terrestrial ecosystems and for improving model simulations and predictions of future global carbon (C) cycling in response to human activities. In this study, we examined the spatial patterns of soil and ecosystem respiration along a precipitation gradient from 167.7 to 398.1 mm in a semi-arid grassland. Our results showed that soil and ecosystem respiration increased linearly with increasing mean annual precipitation. The trends were similar to those of shoot biomass, litter and soil total C content along the precipitation gradient. Our results indicated that precipitation was the primary controlling factor in determining the spatial pattern of soil and ecosystem respiration in semi-arid grasslands in China. The linear/nonlinear relationships in this study describing the variations of the ecosystem carbon process with precipitation can be useful for model development, parameterization and validation at the regional scale to improve predictions of how carbon processes in grasslands respond to climate change, land use and grassland management.