Increasing microbial carbon use efficiency with warming predicts soil heterotrophic respiration globally

The degree to which climate warming will stimulate soil organic carbon (SOC) losses via heterotrophic respiration remains uncertain, in part because different or even opposite microbial physiology and temperature relationships have been proposed in SOC models. We incorporated competing microbial carbon use efficiency (CUE)–mean annual temperature (MAT) and enzyme kinetic–MAT relationships into SOC models, and compared the simulated mass‐specific soil heterotrophic respiration rates with multiple published datasets of measured respiration. The measured data included 110 dryland soils globally distributed and two continental to global‐scale cross‐biome datasets. Model–data comparisons suggested that a positive CUE–MAT relationship best predicts the measured mass‐specific soil heterotrophic respiration rates in soils distributed globally. These results are robust when considering models of increasing complexity and competing mechanisms driving soil heterotrophic respiration–MAT relationships (e.g., carbon substrate availability). Our findings suggest that a warmer climate selects for microbial communities with higher CUE, as opposed to the often hypothesized reductions in CUE by warming based on soil laboratory assays. Our results help to build the impetus for, and confidence in, including microbial mechanisms in soil biogeochemical models used to forecast changes in global soil carbon stocks in response to warming.     

Thermal adaptation of soil microbial respiration to elevated temperature

In the short‐term heterotrophic soil respiration is strongly and positively related to temperature. In the long‐term, its response to temperature is uncertain. One reason for this is because in field experiments increases in respiration due to warming are relatively short‐lived. The explanations proposed for this ephemeral response include depletion of fast‐cycling, soil carbon pools and thermal adaptation of microbial respiration. Using a > 15 year soil warming experiment in a mid‐latitude forest, we show that the apparent ‘acclimation’ of soil respiration at the ecosystem scale results from combined effects of reductions in soil carbon pools and microbial biomass, and thermal adaptation of microbial respiration. Mass‐specific respiration rates were lower when seasonal temperatures were higher, suggesting that rate reductions under experimental warming likely occurred through temperature‐induced changes in the microbial community. Our results imply that stimulatory effects of global temperature rise on soil respiration rates may be lower than currently predicted.     

增温和刈割对高寒草甸土壤呼吸及其组分的影响

评估土壤呼吸及其组分对增温等全球变化的响应对于预测陆地生态系统碳循环至关重要。本研究利用红外线辐射加热器(Infrared heater)装置在青藏高原高寒草甸生态系统设置增温和刈割野外控制实验。通过测定2018年生长季(5—9月)土壤呼吸和异养呼吸,探究增温和刈割对土壤呼吸及其组分的影响。研究结果表明：(1) 单独增温使土壤呼吸显著增加31.65% (P<0.05),异养呼吸显著增加27.12% (P<0.05),土壤自养呼吸没有显著改变(P>0.05);单独刈割对土壤呼吸和自养呼吸没有显著影响(P>0.05),单独刈割刺激异养呼吸增加32.54% (P<0.05);(2) 增温和刈割之间的交互作用对土壤呼吸和异养呼吸没有显著影响(P>0.05),但是对自养呼吸的影响是显著的(P<0.05),土壤呼吸和异养呼吸的季节效应显著(P<0.05);(3)土壤呼吸及其组分与土壤温度均成显著指数关系,与土壤湿度呈显著的正相关关系(P<0.05),处理影响它们的响应敏感性。本研究表明青藏高原东缘高寒草甸土壤碳排放与气候变暖存在正反馈。     

The Impact of Invasive Earthworms on Soil Respiration and Soil Carbon Within Temperate Hardwood Forests

Improving current understanding of the factors that control soil carbon (C) dynamics in forest ecosystems remains an important topic of research as it plays an integral role in the fertility of forest soils and the global C cycle. Invasive earthworms have the potential to alter soil C dynamics, though mechanisms and effects remain poorly understood. To investigate potential effects of invasive earthworms on forest C, the forest floor, mineral soil, fine root biomass, litterfall and microbial litter decay rates, and total soil respiration (TSR) over a full year were measured at an invaded and uninvaded deciduous forest site in southern Ontario. The uninvaded site was approximately 300 m from the invaded site and a distinct invasion front between sites was present. Along the invasion front, the biomass of the forest floor was negatively correlated with earthworm abundance and biomass. There was no significant difference between litterfall, microbial litter decay, and TSR between the invaded and uninvaded sites, but fine root biomass was approximately 30% lower at the invaded site. There was no significant difference in total soil C pools (0–30 cm) between the invaded and uninvaded sites. Despite profound impacts on forest floor soil C pools, earthworm invasion does not significantly increase TSR, most likely because increased heterotrophic respiration associated with earthworms is largely offset by a decrease in autotrophic respiration caused by lower fine root biomass.     

帽儿山3种森林生态系统土壤动物与土壤呼吸及其相互关系研究

土壤呼吸是森林生态系统碳循环的关键过程,土壤动物可通过自身代谢及影响微生物活动调控土壤呼吸,因此研究土壤动物与土壤呼吸的相互关系对进一步揭示生态系统碳循环的规律和机理具有重要意义。通过野外定点,以帽儿山3种森林生态系统的土壤呼吸及土壤动物为研究对象,探讨不同森林生态系统的土壤呼吸、土壤动物个体密度和生物量的时间变化规律及二者相互关系。结果表明:(1)3种森林生态系统土壤总呼吸速率与土壤异养呼吸速率均呈现先增强后减弱的时间动态变化(P<0.05),且不同森林生态系统土壤异养呼吸速率差异显著(P<0.05),表现为硬阔叶林最高,红松人工林最低;(2)3种森林生态系统土壤动物生物量也具有显著的时间动态变化(P<0.05),均在9月份达到最大,且不同森林生态系统土壤动物个体密度显著不同(P<0.05),蒙古栎林土壤动物个体密度显著小于红松人工林与硬阔叶林;(3)通过回归分析可得,土壤动物数量及生物量的增加抑制了土壤呼吸速率,尤其在生长季初期、末期。研究表明土壤动物可通过抑制微生物生命活动和降低根系呼吸从而对土壤总呼吸及异养呼吸产生负反馈作用,三者是不可分割的整体,与土壤温度、水分等环境因子共同调控着土壤呼吸。     

武夷山不同海拔土壤呼吸对变暖和变冷的响应

由于人类活动,我国亚热带地区正面临剧烈的气候变化,这可能对土壤呼吸有潜在影响.本研究选择武夷山国家公园内针叶林(1442 m)和常绿阔叶林(645 m)为对象,通过土柱置换试验模拟变暖(针叶林置换到常绿阔叶林)和变冷(常绿阔叶林置换到针叶林),探讨模拟变暖和变冷对土壤碳过程的影响,测定两个海拔样地的原位和置换处理的微气...     

Soil microbial responses to warming and increased precipitation and their implications for ecosystem C cycling

A better understanding of soil microbial ecology is critical to gaining an understanding of terrestrial carbon (C) cycle–climate change feedbacks. However, current knowledge limits our ability to predict microbial community dynamics in the face of multiple global change drivers and their implications for respiratory loss of soil carbon. Whether microorganisms will acclimate to climate warming and ameliorate predicted respiratory C losses is still debated. It also remains unclear how precipitation, another important climate change driver, will interact with warming to affect microorganisms and their regulation of respiratory C loss. We explore the dynamics of microorganisms and their contributions to respiratory C loss using a 4-year (2006–2009) field experiment in a semi-arid grassland with increased temperature and precipitation in a full factorial design. We found no response of mass-specific (per unit microbial biomass C) heterotrophic respiration to warming, suggesting that respiratory C loss is directly from microbial growth rather than total physiological respiratory responses to warming. Increased precipitation did stimulate both microbial biomass and mass-specific respiration, both of which make large contributions to respiratory loss of soil carbon. Taken together, these results suggest that, in semi-arid grasslands, soil moisture and related substrate availability may inhibit physiological respiratory responses to warming (where soil moisture was significantly lower), while they are not inhibited under elevated precipitation. Although we found no total physiological response to warming, warming increased bacterial C utilization (measured by BIOLOG EcoPlates) and increased bacterial oxidation of carbohydrates and phenols. Non-metric multidimensional scaling analysis as well as ANOVA testing showed that warming or increased precipitation did not change microbial community structure, which could suggest that microbial communities in semi-arid grasslands are already adapted to fluctuating climatic conditions. In summary, our results support the idea that microbial responses to climate change are multifaceted and, even with no large shifts in community structure, microbial mediation of soil carbon loss could still occur under future climate scenarios.     

Effects of three years of soil warming and shading on the rate of soil respiration: substrate availability and not thermal acclimation mediates observed response

In a number of recent field studies, the positive response of soil respiration to warming has been shown to decline over time. The two main differing hypotheses proposed to explain these results are: (1) soil microbial respiration acclimates to the increased temperature, and (2) substrate availability within the soil decreases with warming so reducing the rate of soil respiration. To investigate the relative merits of these two hypotheses, soil samples (both intact cores and sieved samples) from a 3-year grassland soil-warming and shading experiment were incubated for 4 weeks at three different temperatures under constant laboratory conditions. We tested the hypothesis that sieving the soils would reduce differences in substrate availability between warmed and control plot samples and would therefore result in similar respiration rates if microbial activity had not acclimated to soil warming. In addition, to further test the effect of substrate availability, we compared the respiration rates of soils taken from shaded and unshaded plots. Both soil warming and shading significantly reduced respiration rates in the intact cores, especially under higher incubation temperatures. However, sieving the soil greatly reduced these differences suggesting that substrate availability, and not microbial acclimation to the higher temperatures, played the dominant role in determining the response of heterotrophic soil respiration to warming. The effect of shading appeared to be mediated by reduced plant productivity affecting substrate availability within the soil and hence microbial activity. Given the lack of evidence for thermal acclimation of microbial respiration, there remains the potential for prolonged carbon losses from soils in response to warming.     

Interactive effects of wildfire and permafrost on microbial communities and soil processes in an Alaskan black spruce forest

Boreal forests contain significant quantities of soil carbon that may be oxidized to CO₂ given future increases in climate warming and wildfire behavior. At the ecosystem scale, decomposition and heterotrophic respiration are strongly controlled by temperature and moisture, but we questioned whether changes in microbial biomass, activity, or community structure induced by fire might also affect these processes. We particularly wanted to understand whether postfire reductions in microbial biomass could affect rates of decomposition. Additionally, we compared the short‐term effects of wildfire to the long‐term effects of climate warming and permafrost decline. We compared soil microbial communities between control and recently burned soils that were located in areas with and without permafrost near Delta Junction, AK. In addition to soil physical variables, we quantified changes in microbial biomass, fungal biomass, fungal community composition, and C cycling processes (phenol oxidase enzyme activity, lignin decomposition, and microbial respiration). Five years following fire, organic surface horizons had lower microbial biomass, fungal biomass, and dissolved organic carbon (DOC) concentrations compared with control soils. Reductions in soil fungi were associated with reductions in phenol oxidase activity and lignin decomposition. Effects of wildfire on microbial biomass and activity in the mineral soil were minor. Microbial community composition was affected by wildfire, but the effect was greater in nonpermafrost soils. Although the presence of permafrost increased soil moisture contents, effects on microbial biomass and activity were limited to mineral soils that showed lower fungal biomass but higher activity compared with soils without permafrost. Fungal abundance and moisture were strong predictors of phenol oxidase enzyme activity in soil. Phenol oxidase enzyme activity, in turn, was linearly related to both ¹³C lignin decomposition and microbial respiration in incubation studies. Taken together, these results indicate that reductions in fungal biomass in postfire soils and lower soil moisture in nonpermafrost soils reduced the potential of soil heterotrophs to decompose soil carbon. Although in the field increased rates of microbial respiration can be observed in postfire soils due to warmer soil conditions, reductions in fungal biomass and activity may limit rates of decomposition.     

Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage—a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte‐dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year‐round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock‐dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short‐ and long‐term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon‐rich permafrost ecosystems.     

寒温带岛状林沼泽土壤呼吸速率和季节变化

2011年生长季内利用静态箱-气相色谱法,研究了寒温带典型湿地白桦(Betula platyphylla)岛状林沼泽、兴安落叶松(Larix gmelinii)岛状林沼泽土壤呼吸速率的季节动态及其主要环境因子,利用壕沟隔断法对土壤呼吸各组分间的差异进行研究。结果表明:生长季白桦和兴安落叶松岛状林沼泽土壤呼吸速率具有明显的季节性规律,土壤呼吸总速率分别为368.60和312.46 mg m-2h-1,异养呼吸速率分别为300.57和215.70 mg m-2h-1,占土壤呼吸总速率的81.5%和69.0%;自养呼吸速率为68.03和96.76 mg m-2h-1,占土壤呼吸总速率的18.5%和31.0%。不同处理条件下的土壤呼吸在季节变化上表现基本一致,高峰期都发生在夏季;土壤呼吸与温度呈极显著相关性,但与土壤湿度的相关性较差。生长季白桦和兴安落叶松岛状林沼泽土壤呼吸总量分别为12.64和10.61 t/hm2。     

火烧对森林土壤有机碳的影响研究进展

对国内外火烧影响森林土壤有机碳动态的研究成果进行了综合述评。较多研究表明低强度火烧不会造成土壤有机碳贮量的明显变化,但火烧非常强烈而彻底,土壤有机碳明显减少。有限研究表明火烧对森林土壤呼吸的影响结果有增加、降低或无影响,因火烧强度、火后观测时间、森林类型、火烧迹地上植被恢复进程和气候条件等而异。同时,火烧对土壤有机碳组分(活性有机碳和黑碳)也具有不同程度的影响。随着全球变化研究的深入,火烧作为森林主要管理措施对大气CO2浓度影响亦愈来愈受重视,今后应着重开展以下几方面研究:(1)扩大气候和经营管理的变化对森林土壤有机碳贮量时空动态影响研究;(2)深入探讨火烧影响土壤CO2释放的过程及机理;(3)加强火烧历史和频率对黑碳影响的研究;(4)从广度和深度上加强火烧等经营措施对亚热带森林土壤碳动态影响的研究。     

Substrate availability and not thermal acclimation controls microbial temperature sensitivity response to long-term warming

Microbes are responsible for cycling carbon (C) through soils, and predicted changes in soil C stocks under climate change are highly sensitive to shifts in the mechanisms assumed to control the microbial physiological response to warming. Two mechanisms have been suggested to explain the long-term warming impact on microbial physiology: microbial thermal acclimation and changes in the quantity and quality of substrates available for microbial metabolism. Yet studies disentangling these two mechanisms are lacking. To resolve the drivers of changes in microbial physiology in response to long-term warming, we sampled soils from 13- and 28-year-old soil warming experiments in different seasons. We performed short-term laboratory incubations across a range of temperatures to measure the relationships between temperature sensitivity of physiology (growth, respiration, carbon use efficiency, and extracellular enzyme activity) and the chemical composition of soil organic matter. We observed apparent thermal acclimation of microbial respiration, but only in summer, when warming had exacerbated the seasonally-induced, already small dissolved organic matter pools. Irrespective of warming, greater quantity and quality of soil carbon increased the extracellular enzymatic pool and its temperature sensitivity. We propose that fresh litter input into the system seasonally cancels apparent thermal acclimation of C-cycling processes to decadal warming. Our findings reveal that long-term warming has indirectly affected microbial physiology via reduced C availability in this system, implying that earth system models including these negative feedbacks may be best suited to describe long-term warming effects on these soils.     

Soil Carbon Storage Response to Temperature: an Hypothesis

Recently, global and some regional observations of soil carbonstocks and turnover times have implied that warming may notdeplete soil carbon as much as predicted by ecosystem models.The proposed explanation is that microbial respiration of carbonin ‘old’ mineral pools is accelerated less by warmingthan ecosystem models currently assume. Data on the sensitivityof soil respiration to temperature are currently conflicting.An alternative or additional explanation is that warming increasesthe rate of physico-chemical processes which transfer organiccarbon to ‘protected’, more stable, soil carbonpools. These processes include adsorption reactions, some ofwhich are known to have positive activation energies. Theoretically,physico-chemical reactions may be expected to respond more towarming than enzyme-mediated microbial reactions. A simple analyticalmodel and a complex multi-pool soil carbon model are presented,which separate transfers between pools due to physico-chemicalreactions from those associated with microbial respiration.In the short-term, warming depletes soil carbon. But in thelong-term, carbon losses by accelerated microbial respirationare offset by increases in carbon input to the soil (net production)and any acceleration of soil physico-chemical ‘stabilization’reactions. In the models, if net production rates are increasedin response to notional warming by a factor of 1.3, and microbialrespiration (in all pools) by 1.5, then soil carbon at equilibriumremains unchanged if physico-chemical reactions are acceleratedby a factor of about 2.2 (50% more than microbial reactions).Equilibrium soil carbon increases if physico-chemical reactionsare over 50% more sensitive to warming than soil respiration.Copyright 2001 Annals of Botany Company Soil organic matter, carbon, respiration, temperature, stabilization, decomposition, model     

Pathways regulating decreased soil respiration with warming in a biocrust‐dominated dryland

A positive soil carbon (C)‐climate feedback is embedded into the climatic models of the IPCC. However, recent global syntheses indicate that the temperature sensitivity of soil respiration (R_S) in drylands, the largest biome on Earth, is actually lower in warmed than in control plots. Consequently, soil C losses with future warming are expected to be low compared with other biomes. Nevertheless, the empirical basis for these global extrapolations is still poor in drylands, due to the low number of field experiments testing the pathways behind the long‐term responses of soil respiration (R_S) to warming. Importantly, global drylands are covered with biocrusts (communities formed by bryophytes, lichens, cyanobacteria, fungi, and bacteria), and thus, R_S responses to warming may be driven by both autotrophic and heterotrophic pathways. Here, we evaluated the effects of 8‐year experimental warming on R_S, and the different pathways involved, in a biocrust‐dominated dryland in southern Spain. We also assessed the overall impacts on soil organic C (SOC) accumulation over time. Across the years and biocrust cover levels, warming reduced R_S by 0.30 μmol CO₂ m^?2 s^?1 (95% CI = ?0.24 to 0.84), although the negative warming effects were only significant after 3 years of elevated temperatures in areas with low initial biocrust cover. We found support for different pathways regulating the warming‐induced reduction in R_S at areas with low (microbial thermal acclimation via reduced soil mass‐specific respiration and β‐glucosidase enzymatic activity) vs. high (microbial thermal acclimation jointly with a reduction in autotrophic respiration from decreased lichen cover) initial biocrust cover. Our 8‐year experimental study shows a reduction in soil respiration with warming and highlights that biocrusts should be explicitly included in modeling efforts aimed to quantify the soil C–climate feedback in drylands.     

Predominant role of water in regulating soil and microbial respiration and their responses to climate change in a semiarid grassland

Climate change can profoundly impact carbon (C) cycling of terrestrial ecosystems. A field experiment was conducted to examine responses of total soil and microbial respiration, and microbial biomass to experimental warming and increased precipitation in a semiarid temperate steppe in northern China since April 2005. We measured soil respiration twice a month over the growing seasons, soil microbial biomass C (MBC) and N (MBN), microbial respiration (MR) once a year in the middle growing season from 2005 to 2007. The results showed that interannual variations in soil respiration, MR, and microbial biomass were positively related to interannual fluctuations in precipitation. Laboratory incubation with a soil moisture gradient revealed a constraint of the temperature responses of MR by low soil moisture contents. Across the 3 years, experimental warming decreased soil moisture, and consequently caused significant reductions in total and microbial respiration, and microbial biomass, suggesting stronger negatively indirect effects through warming‐induced water stress than the positively direct effects of elevated temperature. Increased evapotranspiration under experimental warming could have reduced soil water availability below a stress threshold, thus leading to suppression of plant growth, root and microbial activities. Increased precipitation significantly stimulated total soil and microbial respiration and all other microbial parameters and the positive precipitation effects increased over time. Our results suggest that soil water availability is more important than temperature in regulating soil and microbial respiratory processes, microbial biomass and their responses to climate change in the semiarid temperate steppe. Experimental warming caused greater reductions in soil respiration than in gross ecosystem productivity (GEP). In contrast, increased precipitation stimulated GEP more than soil respiration. Our observations suggest that climate warming may cause net C losses, whereas increased precipitation may lead to net C gains in the semiarid temperate steppe. Our findings highlight that unless there is concurrent increase in precipitation, the temperate steppe in the arid and semiarid regions of northern China may act as a net C source under climate warming.     

干湿交替格局下黄土高原小麦田土壤呼吸的温湿度模型

全球气候变化的直接后果是气温升高,同时还可能引起强降雨增多和干旱频发,形成干湿交替的格局.土壤呼吸在全球变化过程中发挥着重要作用.以黄土高原沟壑区小麦田土壤为研究对象,采用3个全自动多通量箱以及相应的气象监测系统,对土壤呼吸和环境因子全天候连续测定,利用已有的单因子模型、双因子模型对测定的土壤呼吸与气温和湿度的关系进行了拟合,通过优化,根据实际情况提出E-Q(exponential-quadratic)模型.结果表明:(1)干湿交替格局下,基于气温的单因子模型(指数模型,幂函数模型和线性模型)不适合模拟土壤呼吸;(2)基于土壤湿度的单因子模型中,二次曲线模型最适合模拟干湿交替格局下土壤呼吸的响应情况;(3)基于气温和土壤湿度的双因子模型中,E-Q模型SR=aebT(c+dW+fW2)g,既能反映土壤呼吸随气温的正向指数变化,又能表现土壤湿度对土壤呼吸的双向调节作用,解释了土壤呼吸73.05%的变化情况,比其他双因子模型和单因子模型更能有效描述干湿交替情况下土壤呼吸对气温和土壤湿度协同变化的响应特征.     

Rapid turnover of DOC in temperate forests accounts for increased CO2 production at elevated temperatures

The evidence for the contribution of soil warming to changes in atmospheric CO₂ concentrations and carbon stocks of temperate forest ecosystems is equivocal. Here, we use data from a beech/oak forest on concentrations and stable isotope ratios of dissolved organic carbon (DOC), phosphate buffer-extractable organic carbon, soil organic carbon (SOC), respiration and microbial gross assimilation of N to show that respired soil carbon originated from DOC. However, the respiration was not dependent on the DOC concentration but exceeded the daily DOC pool three to four times, suggesting that DOC was turned over several times per day. A mass flow model helped to calculate that a maximum of 40% of the daily DOC production was derived from SOC and to demonstrate that degradation of SOC is limiting respiration of DOC. The carbon flow model on SOC, DOC, microbial C mobilization/immobilization and respiration is linked by temperature-dependent microbial and enzyme activity to global warming effects of CO₂ emitted to the atmosphere.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Interactions with successional stage and nutrient status determines the life‐form‐specific effects of increased soil temperature on boreal forest floor vegetation

The boreal forest is one of the largest terrestrial biomes and plays a key role for the global carbon balance and climate. The forest floor vegetation has a strong influence on the carbon and nitrogen cycles of the forests and is sensitive to changes in temperature conditions and nutrient availability. Additionally, the effects of climate warming on forest floor vegetation have been suggested to be moderated by the tree layer. Data on the effects of soil warming on forest floor vegetation from the boreal forest are, however, very scarce. We studied the effects on the forest floor vegetation in a long‐term (18 years) soil warming and fertilization experiment in a Norway spruce stand in northern Sweden. During the first 9 years, warming favored early successional species such as grasses and forbs at the expense of dwarf shrubs and bryophytes in unfertilized stands, while the effects were smaller after fertilization. Hence, warming led to significant changes in species composition and an increase in species richness in the open canopy nutrient limited forest. After another 9 years of warming and increasing tree canopy closure, most of the initial effects had ceased, indicating an interaction between forest succession and warming. The only remaining effect of warming was on the abundance of bryophytes, which contrary to the initial phase was strongly favored by warming. We propose that the suggested moderating effects of the tree layer are specific to plant life‐form and conclude that the successional phase of the forest may have a considerable impact on the effects of climate change on forest floor vegetation and its feedback effects on the carbon and nitrogen cycles, and thus on the climate.