陆地生态系统过程对气候变暖的响应与适应

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。     

Combined effects of global climate change and regional ecosystem drivers on an exploited marine food web

Changes in climate, in combination with intensive exploitation of marine resources, have caused large‐scale reorganizations in many of the world's marine ecosystems during the past decades. The Baltic Sea in Northern Europe is one of the systems most affected. In addition to being exposed to persistent eutrophication, intensive fishing, and one of the world's fastest rates of warming in the last two decades of the 20th century, accelerated climate change including atmospheric warming and changes in precipitation is projected for this region during the 21st century. Here, we used a new multimodel approach to project how the interaction of climate, nutrient loads, and cod fishing may affect the future of the open Central Baltic Sea food web. Regionally downscaled global climate scenarios were, in combination with three nutrient load scenarios, used to drive an ensemble of three regional biogeochemical models (BGMs). An Ecopath with Ecosim food web model was then forced with the BGM results from different nutrient‐climate scenarios in combination with two different cod fishing scenarios. The results showed that regional management is likely to play a major role in determining the future of the Baltic Sea ecosystem. By the end of the 21st century, for example, the combination of intensive cod fishing and high nutrient loads projected a strongly eutrophicated and sprat‐dominated ecosystem, whereas low cod fishing in combination with low nutrient loads resulted in a cod‐dominated ecosystem with eutrophication levels close to present. Also, nonlinearities were observed in the sensitivity of different trophic groups to nutrient loads or fishing depending on the combination of the two. Finally, many climate variables and species biomasses were projected to levels unseen in the past. Hence, the risk for ecological surprises needs to be addressed, particularly when the results are discussed in the ecosystem‐based management context.     

Adding climate change to the mix: responses of aquatic ectotherms to the combined effects of eutrophication and warming

The threat of excessive nutrient enrichment, or eutrophication, is intensifying across the globe as climate change progresses, presenting a major management challenge. Alterations in precipitation patterns and increases in temperature are increasing nutrient loadings in aquatic habitats and creating conditions that promote the proliferation of cyanobacterial blooms. The exacerbating effects of climate warming on eutrophication are well established, but we lack an in-depth understanding of how aquatic ectotherms respond to eutrophication and warming in tandem. Here, I provide a brief overview and critique of studies exploring the cumulative impacts of eutrophication and warming on aquatic ectotherms, and provide forward direction using mechanistically focused, multi-threat experiments to disentangle complex interactions. Evidence to date suggests that rapid warming will exacerbate the negative effects of eutrophication on aquatic ectotherms, but gradual warming will induce physiological remodelling that provides protection against nutrients and hypoxia. Moving forward, research will benefit from a greater focus on unveiling cause and effect mechanisms behind interactions and designing treatments that better mimic threat dynamics in nature. This approach will enable robust predictions of species responses to ongoing eutrophication and climate warming and enable the integration of climate warming into eutrophication management policies.     

Global‐change drivers of ecosystem functioning modulated by natural variability and saturating responses

Humans are altering global environment at an unprecedented rate through changes in biodiversity, climate, nitrogen cycle, and land use. To address their effects on ecosystem functioning, experiments most frequently explore one driver at a time and control as many confounding factors as possible. Yet, which driver exerts the largest influence on ecosystem functioning and whether their relative importance changes among systems remain unclear. We analyzed experiments in the Patagonian steppe that evaluated the aboveground net primary production (ANPP) response to manipulated gradients of species richness, precipitation, temperature, nitrogen fertilization (N), and grazing intensity. We compared the effect on ANPP relative to ambient conditions considering intensity and direction of manipulations for each driver. The ranking of responses to drivers with comparable manipulation intensity was as follows: biodiversity>grazing>precipitation>N. For a similar intensity of manipulation, the effect of biodiversity loss was 4.0, 3.6, and 1.5, times larger than N deposition, decreased precipitation, and increased grazing intensity. We interpreted our results considering two hypotheses. First, the response of ANPP to changes in precipitation and biodiversity is saturating, so we expected larger effects when the driver was reduced, relative to ambient conditions, than when it was increased. Experimental manipulations that reduced ambient levels had larger effects than those that increased them. Second, the sensitivity of ANPP to each driver is inversely related to the natural variability of the driver. In Patagonia, the ranking of natural variability of drivers is as follows: precipitation>grazing>temperature>biodiversity>N. So, in general, the ecosystem was most sensitive to drivers that varied the least. Comparable results from Cedar Creek (MN) support both hypotheses and suggest that sensitivity to drivers varies among ecosystem types. Given the importance of understanding ecosystem sensitivity to predict global‐change impacts, it is necessary to design new experiments located in regions with contrasting natural variability and that include the full range of drivers.     

Soil–plant N processes in a High Arctic ecosystem,NW Greenland are altered by long‐term experimental warming and higher rainfall

Rapid temperature and precipitation changes in High Arctic tundra ecosystems are altering the biogeochemical cycles of carbon (C) and nitrogen (N), but in ways that are difficult to predict. The challenge grows from the uncertainty of N cycle responses and the extent to which shifts in soil N are coupled with the C cycle and productivity of tundra systems. We used a long‐term (since 2003) experiment of summer warming and supplemental summer water additions to a High Arctic ecosystem in NW Greenland, and applied a combination of discrete sampling and in situ soil core incubations to measure C and N pools and seasonal microbial processes that might control plant‐available N. We hypothesized that elevated temperature and increased precipitation would stimulate microbial activity and net inorganic N mineralization, thereby increasing plant N‐availability through the growing season. While we did find increased N mineralization rates under both global change scenarios, water addition also significantly increased net nitrification rates, loss of NO₃^?‐N via leaching, and lowered rates of labile organic N production. We also expected the chronic warming and watering would lead to long‐term changes in soil N‐cycling that would be reflected in soil δ¹⁵N values. We found that soil δ¹⁵N decreased under the different climate change scenarios. Our results suggest that temperature accelerates biological processes and existing C and N transformations, but moisture increases soil hydraulic connectivity and so alters the pathways, and changes the fate of the products of C and N transformations. In addition, our findings indicate that warmer, wetter High Arctic tundra will be cycling N and C in ways that may transform these landscapes in part leading to greater C sequestration, but simultaneously, N losses from the upper soil profile that may be transported to depth dissolved in water and or transported off site in lateral flow.     

Sensitivity and responses of diatoms to climate warming in lakes heavily influenced by humans

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Predicting ecosystem carbon balance in a warming Arctic: the importance of long‐term thermal acclimation potential and inhibitory effects of light on respiration

The carbon balance of Arctic ecosystems is particularly sensitive to global environmental change. Leaf respiration (R), a temperature‐dependent key process in determining the carbon balance, is not well‐understood in Arctic plants. The potential for plants to acclimate to warmer conditions could strongly impact future global carbon balance. Two key unanswered questions are (1) whether short‐term temperature responses can predict long‐term respiratory responses to growth in elevated temperatures and (2) to what extent the constant daylight conditions of the Arctic growing season inhibit leaf respiration. In two dominant Arctic species E riophorum vaginatum (tussock grass) and B etula nana (woody shrub), we assessed the extent of respiratory inhibition in the light (R _L/R _D), respiratory response to short‐term temperature change, and respiratory acclimation to long‐term warming treatments. We found that R of both species is strongly inhibited by light (averaging 35% across all measurement temperatures). In E . vaginatum both R _L and R _D acclimated to the long‐term warming treatment, reducing the magnitude of respiratory response relative to the short‐term response to temperature increase. In B . nana, both R _L and R _D responded to short‐term temperature increase but showed no acclimation to the long‐term warming. The ability to predict plant respiratory response to global warming with short‐term temperature responses will depend on species‐specific acclimation potential and the differential response of R _L and R _D to temperature. With projected woody shrub encroachment in Arctic tundra and continued warming, changing species dominance between these two functional groups, may impact ecosystem respiratory response and carbon balance.     

川西高山林线土壤活性碳、氮对短期增温的响应

随着温室效应的加剧,受低温限制的高山林线生态系统对全球气候变暖较为敏感,可能直接影响到植物的生长和土壤碳氮过程.本研究假设气候变暖会改变高山生态系统土壤活性碳氮含量,在四川省理县米亚罗高山生态系统定位站,采用开顶式模拟增温装置(OTC)模拟增温对土壤活性碳、氮的短期影响.分别于2017年4、7和10月,采集OTC以及对照样地(CK)内土壤有机层和矿质土壤层的原状土壤,测定土壤可溶性有机碳(DOC)、土壤微生物生物量碳(MBC)、土壤可溶性有机氮(DON)和土壤微生物生物量氮(MBN)含量.结果表明: 模拟增温使年均气温升高0.88 ℃,土壤有机层和矿质土壤层的年均温度分别提高0.48和0.23 ℃.模拟增温没有显著改变土壤有机质和含水量,但显著提高了矿质土壤层的pH值,同时显著降低了非生长季矿质土壤层的DOC、DON含量;季节变化对两个层次的DOC、DON和MBN含量有极显著影响,而MBC没有明显的季节动态;增温和季节交互作用对矿质土壤层的DOC和DON有显著影响.土壤有机层的MBC、MBN含量显著高于矿质土壤层.土壤活性碳、氮与土壤有机质和含水量呈极显著正相关,MBC、MBN与土壤pH呈极显著正相关,MBN与土壤温度呈显著负相关.     

Light availability regulates the response of algae and heterotrophic bacteria to elevated nutrient levels and warming in a northern boreal peatland

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Responses of an old-field plant community to interacting factors of elevated [CO2], warming, and soil moisture

Aims: The direct effects of atmospheric and climatic change factors—atmospheric[CO₂], air temperature and changes in precipitation—canshape plant community composition and alter ecosystem function.It is essential to understand how these factors interact tomake better predictions about how ecosystems may respond tochange. We investigated the direct and interactive effects of[CO₂], warming and altered soil moisture in open-top chambers(OTCs) enclosing a constructed old-field community to test howthese factors shape plant communities. Materials and methods: The experimental facility in Oak Ridge, TN, USA, made use of4-m diameter OTCs and rain shelters to manipulate [CO₂] (ambient,ambient + 300 ppm), air temperature (ambient, ambient + 3.5°C)and soil moisture (wet, dry). The plant communities within thechambers comprised seven common old-field species, includinggrasses, forbs and legumes. We tracked foliar cover for eachspecies and calculated community richness, evenness and diversityfrom 2003 to 2005. Important findings: This work resulted in three main findings: (1) warming had species-specificeffects on foliar cover that varied through time and were alteredby soil moisture treatments; (2) [CO₂] had little effect onindividual species or the community; (3) diversity, evennessand richness were influenced most by soil moisture, primarilyreflecting the response of one dominant species. We concludethat individualistic species responses to atmospheric and climaticchange can alter community composition and that plant populationsand communities should be considered as part of analyses ofterrestrial ecosystem response to climate change. However, predictionof plant community responses may be difficult given interactionsbetween factors and changes in response through time.     

Responses of soil C pools to combined warming and altered precipitation regimes: A meta-analysis

Aim

Global warming and altered precipitation substantially affect soil carbon (C) pools and can, in turn, feed back into climate change. However, how soil C pools respond to the combined effects of warming and altered precipitation remains unclear.

Location

Global.

Time period

1996–2021.

Major taxa studied

Soil organic C pools.

Method

A meta-analysis was performed using 657 observations obtained from 34 published articles that focused on both individual and combined effects of warming and altered precipitation on soil organic C (SOC), dissolved organic C (DOC) and microbial biomass C (MBC) to quantify the responses of soil C pools.

Results

Across all combined warming and increased precipitation experiments, SOC and MBC increased by an average of 4.0% and 15.4%, respectively. In contrast, warming combined with decreased precipitation led to a substantial decline in SOC and MBC by an average of 8.2% and 12.3%, respectively. The responses of DOC to combined warming and altered precipitation were marginal. The direction and magnitude of the responses to the combined treatment were more similar to those in the individual altered precipitation treatment than to those in the individual warming treatment. Furthermore, these combined effects were substantially influenced by altered precipitation magnitudes. Combined warming and altered precipitation had greater impacts on soil C pools than their individual treatments but were not substantially different from the sum of their respective individual effects, showing overall additive effects. The responses of soil C pools to combined warming and altered precipitation were observed to be more pronounced in grasslands than in forests.

Main conclusion

The results demonstrated that altered precipitation regimes often dominated over warming in regulating soil C pools under combined warming and altered precipitation and improved our understanding of soil C cycles under climate change scenarios.     

Reduced N cycling in response to elevated CO2, warming,and drought in a Danish heathland: Synthesizing results of the CLIMAITE project after two years of treatments

Field‐scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short‐term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co‐dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought‐induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.     

Belowground carbon responses to experimental warming regulated by soil moisture change in an alpine ecosystem of the Qinghai–Tibet Plateau

Recent studies found that the largest uncertainties in the response of the terrestrial carbon cycle to climate change might come from changes in soil moisture under the elevation of temperature. Warming‐induced change in soil moisture and its level of influence on terrestrial ecosystems are mostly determined by climate, soil, and vegetation type and their sensitivity to temperature and moisture. Here, we present the results from a warming experiment of an alpine ecosystem conducted in the permafrost region of the Qinghai–Tibet Plateau using infrared heaters. Our results show that 3 years of warming treatments significantly elevated soil temperature at 0–100 cm depth, decreased soil moisture at 10 cm depth, and increased soil moisture at 40–100 cm depth. In contrast to the findings of previous research, experimental warming did not significantly affect NH ₄ ⁺‐N, NO ₃ ⁻‐N, and heterotrophic respiration, but stimulated the growth of plants and significantly increased root biomass at 30–50 cm depth. This led to increased soil organic carbon, total nitrogen, and liable carbon at 30–50 cm depth, and increased autotrophic respiration of plants. Analysis shows that experimental warming influenced deeper root production via redistributed soil moisture, which favors the accumulation of belowground carbon, but did not significantly affected the decomposition of soil organic carbon. Our findings suggest that future climate change studies need to take greater consideration of changes in the hydrological cycle and the local ecosystem characteristics. The results of our study will aid in understanding the response of terrestrial ecosystems to climate change and provide the regional case for global ecosystem models.     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.     

模拟升温和放牧对高寒草甸土壤有机碳氮组分和微生物生物量的影响

土壤活性、惰性有机质库和微生物生物量在数量和分配上的变化是陆地生态系统土壤有机质贮存和动态变化的决定性因素。采用OTCs(Open top chambers)升温以及刈割+粪便归还的方法,对青藏高原东部高寒草甸土壤有机碳氮组分和微生物生物量对气候变暖和放牧的响应进行了研究。结果表明,模拟升温在短期内显著降低土壤活性有机碳Ⅰ、活性有机氮Ⅰ和惰性有机碳的含量,而由于粪便归还作用,放牧明显增加土壤活性有机碳、氮Ⅰ的含量。模拟升温和放牧对有机碳、氮组分的作用效应相互抵消,两者共同作用下有机碳、氮组分仅略有降低。单一的模拟升温或放牧没有显著改变微生物生物量碳,但是两者共同作用却能够大大增加微生物生物量碳。放牧和取样时间存在着明显的交互作用,放牧效应随时间递减。本研究表明,气候变暖对放牧草甸有机碳、氮组分影响不大;放牧过程中的牲畜粪便归还作用不容忽视。     

Using model‐data fusion to interpret past trends,and quantify uncertainties in future projections,of terrestrial ecosystem carbon cycling

Uncertainties in model projections of carbon cycling in terrestrial ecosystems stem from inaccurate parameterization of incorporated processes (endogenous uncertainties) and processes or drivers that are not accounted for by the model (exogenous uncertainties). Here, we assess endogenous and exogenous uncertainties using a model‐data fusion framework benchmarked with an artificial neural network (ANN). We used 18 years of eddy‐covariance carbon flux data from the Harvard forest, where ecosystem carbon uptake has doubled over the measurement period, along with 15 ancillary ecological data sets relative to the carbon cycle. We test the ability of combinations of diverse data to constrain projections of a process‐based carbon cycle model, both against the measured decadal trend and under future long‐term climate change. The use of high‐frequency eddy‐covariance data alone is shown to be insufficient to constrain model projections at the annual or longer time step. Future projections of carbon cycling under climate change in particular are shown to be highly dependent on the data used to constrain the model. Endogenous uncertainties in long‐term model projections of future carbon stocks and fluxes were greatly reduced by the use of aggregated flux budgets in conjunction with ancillary data sets. The data‐informed model, however, poorly reproduced interannual variability in net ecosystem carbon exchange and biomass increments and did not reproduce the long‐term trend. Furthermore, we use the model‐data fusion framework, and the ANN, to show that the long‐term doubling of the rate of carbon uptake at Harvard forest cannot be explained by meteorological drivers, and is driven by changes during the growing season. By integrating all available data with the model‐data fusion framework, we show that the observed trend can only be reproduced with temporal changes in model parameters. Together, the results show that exogenous uncertainty dominates uncertainty in future projections from a data‐informed process‐based model.