陆地生态系统野外增温控制实验的技术与方法

由于人类活动导致的碳排放急剧增加, 工业革命以来全球地表温度显著增加约1 ℃, 未来全球气候还将持续变暖, 到21世纪末最高可升温4 ℃。这种前所未有的气候变化不仅影响陆地植被的适应策略, 也深刻影响生态系统的结构和功能。其中陆地生态系统碳收支对全球变暖的反馈, 是决定未来气候变化强度的关键因素, 因此全球已经开展了大量的生态系统尺度的野外增温控制实验, 研究生态系统碳收支对气温升高的响应, 从而提高地球系统模型的预测精度。然而由于增温技术和方法的不同, 不同研究的结果之间难以进行比较。该文系统总结了常见的野外增温技术和方法, 包括主动增温和被动增温, 阐述了其优缺点、适用对象以及相关研究成果。同时简要介绍了野外增温控制实验的前沿研究方向——新一代野外增温技术(包括全土壤剖面增温和全生态系统增温)和基于新一代增温技术开展的野外增温联网实验。     

Pollen and climatic change

Summary There has been increasing interest shown in the literature over the possible implications of global warming on the climate, ecology and economy of the world community. Aerobiologists in Europe could play an important strategic role in monitoring and predicting ecological change and in providing useful information for climatologists. The problems of identification and co-ordination are briefly discussed.     

Asymmetric effects of daytime and nighttime warming on alpine treeline recruitment

Trees at their upper range limits are highly sensitive to climate change, and thus alpine treelines worldwide have changed their recruitment patterns in response to climate warming. However, previous studies focused only on daily mean temperature, neglecting the asymmetric influences of daytime and nighttime warming on recruitments in alpine treelines. Here, based on the compiled dataset of tree recruitment series from 172 alpine treelines across the Northern Hemisphere, we quantified and compared the different effects of daytime and nighttime warming on treeline recruitment using four indices of temperature sensitivity, and assessed the responses of treeline recruitment to warming-induced drought stress. Our analyses demonstrated that even in different environmental regions, both daytime and nighttime warming could significantly promote treeline recruitment, and however, treeline recruitment was much more sensitive to nighttime warming than to daytime warming, which could be attributable to the presence of drought stress. The increasing drought stress primarily driven by daytime warming rather than by nighttime warming would likely constrain the responses of treeline recruitment to daytime warming. Our findings provided compelling evidence that nighttime warming rather than daytime warming could play a primary role in promoting the recruitment in alpine treelines, which was related to the daytime warming-induced drought stress. Thus, daytime and nighttime warming should be considered separately to improve future projections of global change impacts across alpine ecosystems.     

寒地春玉米生长发育及产量对花前夜间增温的响应

采用田间开放式夜间增温试验方法,研究雨养农区寒地春玉米生长发育及产量对花前夜间增温的响应.结果表明:夜间增温条件下,0 ～ 10 cm耕层土壤夜间温度升高1.7℃,土壤水分略有下降;夜间增温使春玉米物候期明显提前,花前生育期缩短ld,花后生育期延长1d;夜间增温明显促进春玉米幼苗生长,提高根系长度,单株绿叶面积和棒三叶面积分别比对照提高13.5％和14.6％;与对照相比,春玉米地上生物量和籽粒产量分别显著增高8.2％和9.3％,百粒重显著增高7.1％.东北地区气候变暖尤其是日最低温度升高对春玉米的直接影响效应可能以增产为主.     

Seasonally different response of photosynthetic activity to daytime and night‐time warming in the Northern Hemisphere

Over the last century the Northern Hemisphere has experienced rapid climate warming, but this warming has not been evenly distributed seasonally, as well as diurnally. The implications of such seasonal and diurnal heterogeneous warming on regional and global vegetation photosynthetic activity, however, are still poorly understood. Here, we investigated for different seasons how photosynthetic activity of vegetation correlates with changes in seasonal daytime and night‐time temperature across the Northern Hemisphere (>30°N), using Normalized Difference Vegetation Index (NDVI) data from 1982 to 2011 obtained from the Advanced Very High Resolution Radiometer (AVHRR). Our analysis revealed some striking seasonal differences in the response of NDVI to changes in day‐ vs. night‐time temperatures. For instance, while higher daytime temperature (T_max) is generally associated with higher NDVI values across the boreal zone, the area exhibiting a statistically significant positive correlation between T_max and NDVI is much larger in spring (41% of area in boreal zone – total area 12.6 × 10⁶ km²) than in summer and autumn (14% and 9%, respectively). In contrast to the predominantly positive response of boreal ecosystems to changes in T_max, increases in T_max tended to negatively influence vegetation growth in temperate dry regions, particularly during summer. Changes in night‐time temperature (T_min) correlated negatively with autumnal NDVI in most of the Northern Hemisphere, but had a positive effect on spring and summer NDVI in most temperate regions (e.g., Central North America and Central Asia). Such divergent covariance between the photosynthetic activity of Northern Hemispheric vegetation and day‐ and night‐time temperature changes among different seasons and climate zones suggests a changing dominance of ecophysiological processes across time and space. Understanding the seasonally different responses of vegetation photosynthetic activity to diurnal temperature changes, which have not been captured by current land surface models, is important for improving the performance of next generation regional and global coupled vegetation‐climate models.     

放牧和模拟增温对藏北高寒草地植物群落特征及生产力的影响

气候变化和放牧活动对草地植物物种多样性和生产力具有重要影响。为探索藏北高寒草地植物物种多样性和生产力对增温、放牧及其交互作用的响应, 于2011年在藏北高原开始建立增温实验平台, 2016年起增设放牧、增温+放牧实验, 连续2年(2016-2017年)观测了植物群落特征、群落组成、生产力和物种多样性。结果表明, 增温和放牧对高寒草地植物高度和净初级生产力具有显著交互作用。在放牧条件下, 增温对植物高度无显著影响; 但在不放牧条件下, 增温却显著增加了植物高度。在放牧条件下, 增温对净初级生产力的影响存在年际差异, 2016年增温对生产力无显著影响, 2017年增温显著降低了植物净初级生产力; 但在不放牧条件下, 增温对植物净初级生产力无显著影响。增温和放牧对高寒草地植物物种丰富度、盖度、重要值及多样性均无显著交互作用。植物盖度在增温和放牧条件下显著降低, 杂类草物种比例显著增加, 但物种多样性均无显著变化。研究表明, 增温和放牧显著改变高寒草地群落结构。未来气候变化条件下, 放牧活动加剧有可能导致高寒草地生产力降低。     

大气和土壤增温对杉木幼苗地上物候及生长的影响

为了探讨全球变暖对杉木幼苗地上物候和生长的影响,在福建三明森林生态系统与全球变化研究站陈大观测点开展大气温度控制(ambient, open-top chamber)和土壤温度控制(ambient, ambient+4℃)双因子试验,设置对照、单独大气增温、单独土壤增温、大气和土壤同时增温4种处理,大气增温采用开顶箱被动式增温,土壤增温采用电缆增温。建立48个单株水平的根箱,每个根箱内种植1棵1年生2代半短侧枝杉木幼苗,于2016年开始对杉木幼苗地上物候和生长动态进行为期1年的研究。结果表明:(1)大气温度控制对杉木幼苗物候和生长具有显著影响;与无大气增温相比,大气增温使杉木幼苗顶芽膨胀、顶芽展开和顶芽新稍生长时间显著提前,树高生长季长度显著延长,杉木幼苗树高生长得到显著促进。(2)土壤温度控制对杉木幼苗的物候和生长均没有显著影响。(3)大气温度控制和土壤温度控制的交互作用对杉木幼苗生长具有显著影响,大气和土壤同时增温处理的杉木幼苗树高和侧枝生长显著大于单独大气增温处理。表明全球气候变暖可能对杉木生产力具有一定的促进作用。     

全球变暖与陆地生态系统研究中的野外增温装置

由于化石燃料燃烧和森林砍伐等人类活动引起的地球大气层中温室气体(主要是二氧化碳)的富集已导致全球平均温度在20世纪升高了0.6 ℃,并将在本世纪继续上升1.4~5.8 ℃。这种地质历史上前所未有的全球变暖将对陆地植物和生态系统产生深远影响,并通过全球碳循环的改变反馈于全球气候变化。作为全球变化生态学的主要研究方法之一,生态系统增温实验能够为生态模型提供参数估计和模型验证。然而由于在世界各地使用的增温装置不同,使得各个生态系统之间的结果比较和整合难以实施,增加了模型预测的不确定性。该文通过比较几种常见的野外增温装置在模拟全球变暖情形时的优缺点,指出利用不同增温装置进行全球变暖研究中应注意的一些问题;同时探讨了全球变暖控制实验研究中的一些关键性的科学问题。     

Design and performance of combined infrared canopy and belowground warming in the B4WarmED (Boreal Forest Warming at an Ecotone in Danger) experiment

Conducting manipulative climate change experiments in complex vegetation is challenging, given considerable temporal and spatial heterogeneity. One specific challenge involves warming of both plants and soils to depth. We describe the design and performance of an open‐air warming experiment called Boreal Forest Warming at an Ecotone in Danger (B4WarmED) that addresses the potential for projected climate warming to alter tree function, species composition, and ecosystem processes at the boreal‐temperate ecotone. The experiment includes two forested sites in northern Minnesota, USA, with plots in both open (recently clear‐cut) and closed canopy habitats, where seedlings of 11 tree species were planted into native ground vegetation. Treatments include three target levels of plant canopy and soil warming (ambient, +1.7 °C, +3.4 °C). Warming was achieved by independent feedback control of voltage input to aboveground infrared heaters and belowground buried resistance heating cables in each of 72‐7.0 m² plots. The treatments emulated patterns of observed diurnal, seasonal, and annual temperatures but with superimposed warming. For the 2009 to 2011 field seasons, we achieved temperature elevations near our targets with growing season overall mean differences (?T_below) of +1.84 °C and +3.66 °C at 10 cm soil depth and (?T^above) of +1.82 °C and +3.45 °C for the plant canopies. We also achieved measured soil warming to at least 1 m depth. Aboveground treatment stability and control were better during nighttime than daytime and in closed vs. open canopy sites in part due to calmer conditions. Heating efficacy in open canopy areas was reduced with increasing canopy complexity and size. Results of this study suggest the warming approach is scalable: it should work well in small‐statured vegetation such as grasslands, desert, agricultural crops, and tree saplings (<5 m tall).     

开放式昼夜不同增温对单季稻影响的试验研究

全球变暖趋势日益明朗,且存在明显的季节性差异和昼夜不对称性。水稻是我国最重要的粮食作物,研究水稻生产力对昼夜不同增温的响应与适应对我国未来粮食安全战略决策至关重要。为此,作者在江苏南京设计我国首个稻田开放式增温（FATI：Free Air Temperature Increased) 系统,在2007—2008年对水稻进行昼夜不同增温（全天增温、白天增温和夜间增温）的试验研究。结果表明,该增温系统可以形成4 m2均匀且稳定的增温范围,全天、白天和夜间增温处理平均分别可以使水稻全生育期冠层日均温升高2.0℃、0.6℃和0.9℃。白天增温使水稻全生育期冠层白天温度平均升高1.1℃,夜间增温使夜间温度平均升高1.8℃,与未来的升温幅度相似。全天、白天和夜间增温处理下,水稻冠层温度日较差变化分别为0.1℃、0.6℃和-0.9℃。同时,在该系统的3种增温情景下,水稻分蘖期、孕穗期和灌浆期的田间冠层温度日变化趋势基本与常规对照区一致,全生育期的日平均温度变化趋势也基本一致。两年的增温试验表明,不同增温情景均对水稻的生育进程、成熟期地上生物量和产量产生了明显的影响。其中全天、白天和夜间增温分别使水稻从移栽到始穗平均的日期缩短3.5 d、2 d和2.5 d,但对始穗至成熟期的影响不明显;全天、白天和夜间增温分别使水稻地上生物量降低7.7%、6.6%和2.8%,但差异均不显著;白天和夜间增温分别使水稻产量下降8.9%和4.5%,而全天增温下水稻产量略有上升,但均未达到显著水平。从产量构成来看,增温下有效穗数和结实率呈现递增趋势,每穗粒数和千粒重呈现下降趋势。增温对水稻株高的影响不明显。上述结果表明,该稻田开放式增温系统能满足水稻系统生产力对未来气候变暖响应与适应的试验研究要求,气候变暖对水稻生产力影响的相关模型分析结果也尚需进一步的田间实际增温试验验证,模型所需的相关参数也需要进一步完善。     

Development of a single mode electromagnetic resonant cavity for rewarming of cryopreserved biomaterials

An electromagnetic (EM) heating system is developed to achieve the rapid and uniform warming of cryopreserved biomaterials. Using the heating system, a rectangular resonant cavity is excited in TE101 mode at frequencies near 434 MHz. In experiments, a spherical phantom of biomaterial with a diameter of 36 mm is placed at the center of the cavity. The phantom is first cooled down to about -80 degrees C within the cavity and then thawed by EM absorption. Results show that EM warming can produce much higher warming rate than conventional water-bath warming method. The spatial temperature distribution in the phantom during EM warming is also more uniform than that during the water-bath warming.     

基于红外增温的草地生态系统响应全球变暖的研究进展

为了更好地了解全球变暖对草地生态系统的影响机制, 该文介绍了红外加热技术的原理、增温效应及其优缺点, 重点从植物物候、光合生理、生长发育、群落结构和功能、土壤特性, 特别是植物群落地下过程方面, 系统综述了基于红外加热技术模拟气候变暖对草地生态系统影响的最新研究进展, 在此基础上探讨了未来草地生态系统响应全球变暖研究拟重视的研究领域。     

Infrared heater system for warming tropical forest understory plants and soils

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.     

Adjustment of Forest Ecosystem Root Respiration as Temperature Warms

Adjustment of ecosystem root respiration to warmer climatic conditions can alter the autotrophic portion of soil respiration and influence the amount of carbon available for biomass production. We examined 44 published values of annual forest root respiration and found an increase in ecosystem root respiration with increasing mean annual temperature (MAT),but the rate of this cross-ecosystem increase (Q10 = 1.6) is less than published values for short-term responses of root respiration to temperature within ecosystems (Q10 = 2-3). When specific root respiration rates and root biomass values were examined, there was a clear trend for decreasing root metabolic capacity (respiration rate at a standard temperature) with increasing MAT. There also were tradeoffs between root metabolic capacity and root system biomass, such that there were no instances of high growing season respiration rates and high root biomass occurring together. We also examined specific root respiration rates at three soil warming experiments at Harvard Forest, USA, and found decreases in metabolic capacity for roots from the heated plots. This decline could be due to either physiological acclimation or to the effects of co-occurring drier soils on the measurement date. Regardless of the cause, these findings clearly suggest that modeling efforts that allow root respiration to increase exponentially with temperature, with Qt0 values of 2 or more, may over-predict root contributions to ecosystem CO2 efflux for future climates and underestimate the amount of C available for other uses,including net primary productivity.     

Climate sensitivity of conifer growth doesn’t reveal distinct low–high dipole along the elevation gradient in the Wolong National Natural Reserve,SW China

Recent climate warming is usually hypothesized to cause tree growth decline in the semi-arid regions where forests are particularly vulnerable to warming induced increases of water deficit. But there is still a large knowledge gap of climate warming effects on tree growth of cold temperate forest in the sub-humid region. Here we assess how climate warming has affected tree growth in the Wolong National Natural Reserve, Southwestern China, where recent warming might not cause tree growth decline because of the cold-humid climatic conditions. Tree-ring data from four co-dominant coniferous species (Larix potaninii var. macrocarpa, Tsuga chinensis, Abies faxoniana and Juniperus saltuaria) along an elevation gradient (from 2700 m to 3700 m) all imprinted temperature signals, and were both positively and significantly correlated with instrumental record of temperature data during the analyzed period of 1954–2010. Furthermore, the rising temperature since 1980 induced pervasive tree growth increases and stronger temperature signals for the coniferous species along the elevation gradient. The tree-ring chronology recorded a strong coherence with instrumental temperature since 1980 and was successful to keep up with the pace of climate warming rate. If climate warming continues, further increases in forest growth could be expected, and the terrestrial carbon sink will be strengthened for the local forest ecosystem in the future.     

大气和土壤增温对杉木幼苗叶片功能性状的影响

为了探讨杉木幼苗叶片功能性状对全球气候变暖的响应与适应机制,在福建三明森林生态系统与全球变化研究站陈大观测点,设置大气增温(开顶箱被动增温,不增温)×土壤增温(电缆增温4℃,不增温)双因子试验,对杉木幼苗叶片生理代谢、形态属性、化学计量学特征等功能性状进行为期1年的研究。结果表明,虽然土壤增温和大气增温对4月和7月的叶片净光合作用速率和水分利用效率没有显著影响,但土壤增温对11月的叶片净光合作用速率有显著促进作用,而大气增温和土壤增温对11月的水分利用效率有显著促进作用。大气增温显著增加比叶面积和叶含水率,并降低叶厚度;而土壤增温则对其均没有显著影响。土壤增温、大气增温均显著增加杉木叶氮浓度,但只有同时大气增温和土壤增温处理才显著提高叶磷浓度。研究结论表明,从叶功能性状角度看,同时大气增温和土壤增温可能更有利于促进杉木幼苗生长。     

模拟增温对土壤呼吸影响机制的研究进展与展望

土壤呼吸是土壤碳库向大气碳库输入的主要途径,而温度升高会影响土壤呼吸从而改变全球碳平衡.据预测在21世纪末,全球平均地表温度将升高0.3～4.8 ℃,因此野外自然条件下的模拟增温试验对土壤呼吸的影响是全球变化研究的热点之一.本文综述了不同时空格局下土壤呼吸对模拟增温的响应特征,指出短期增温能提高土壤呼吸,而长期增温下无统一规律,并且不同生态系统之间也存在差异;重点讨论了模拟增温对土壤呼吸的影响机制,指出增温能直接影响土壤呼吸,同时增温也能通过影响土壤水分、盐分、土壤理化性质等环境因子以及光合作用、凋落物等生物因子对土壤呼吸产生间接影响;另外,分析了土壤呼吸对增温产生适应性的形成机制,主要包括微生物、根、酶的温度适应性、水分限制、氮素过量以及呼吸底物限制.在此基础上对今后的研究方向加以展望:加强根际微生态系统的研究;重点研究不对称增温下土壤呼吸的特征及机制;关注典型物候期和不同季节典型天气土壤呼吸的测定;构建土壤呼吸响应模拟增温试验的研究网络,进行联网试验.     

模拟土壤温度升高对湿草甸小叶章生长及生理特性的影响

采用改进的电缆加热升温方法,研究了土壤温度升高1 ℃～3 ℃对湿草甸小叶章生长及生理特性的影响.结果表明：土壤温度升高明显促进了小叶章植株的生长淹水处理条件下各指标的增长程度依次为:根系总吸收面积(60%)>叶面积(37.32%)>分蘖数(25.81%)>根系体积(24.67%)>株高(18.2%);非淹水处理条件下为:根系总吸收面积(37.47%)>根系体积(36.26%)>叶面积(32.9%)>分蘖数(32.77%)>株高(12.09%).淹水处理条件下小叶章各生理指标对温度升高的响应较为显著,升温淹水处理下小叶章的总叶绿素含量、根部可溶性蛋白含量均呈增加趋势(9.8%和23.88%),而叶片可溶性蛋白和可溶性糖及根部可溶性糖含量显著下降(24.67%、22.92%和35.23%).说明土温升高1 ℃～3 ℃能够显著促进湿草甸小叶章的生长,且其生理特性对土温升高的响应程度因水分条件而异.     

Potential effects of warming and drying on peatland plant community composition

Boreal peatlands may be particularly vulnerable to climate change, because temperature regimes that currently constrain biological activity in these regions are predicted to increase substantially within the next century. Changes in peatland plant community composition in response to climate change may alter nutrient availability, energy budgets, trace gas fluxes, and carbon storage. We investigated plant community response to warming and drying in a field mesocosm experiment in northern Minnesota, USA. Large intact soil monoliths removed from a bog and a fen received three infrared warming treatments crossed with three water‐table treatments (n = 3) for five years. Foliar cover of each species was estimated annually. In the bog, increases in soil temperature and decreases in water‐table elevation increased cover of shrubs by 50% and decreased cover of graminoids by 50%. The response of shrubs to warming was distinctly species‐specific, and ranged from increases (for Andromeda glaucophylla) to decreases (for Kalmia polifolia). In the fens, changes in plant cover were driven primarily by changes in water‐table elevation, and responses were species‐ and lifeform‐specific: increases in water‐table elevation increased cover of graminoids – in particular Carex lasiocarpa and Carex livida– as well as mosses. In contrast, decreases in water‐table elevation increased cover of shrubs, in particular A. glaucophylla and Chamaedaphne calyculata. The differential and sometimes opposite response of species and lifeforms to the treatments suggest that the structure and function of both bog and fen plant communities will change – in different directions or at different magnitudes – in response to warming and/or changes in water‐table elevation that may accompany regional or global climate change.     

Allee effects under climate change

Understanding how climate change affects population dynamics is crucial for assessing future of biodiversity. Here I ask how can Allee effects, occurring when mean individual fitness is reduced in rare populations, respond to increasing temperature. Despite the role Allee effects play in ecology of invasive, threatened and harvested populations, impacts of climate change on Allee effects are practically unknown. Analysis of two population models reveals that whereas the Allee effect driven by predation generally weakens as temperature increases, the Allee effect due to need of finding mates is predicted to become stronger when warming occurs. For the former model, the metabolic theory suggests that with increasing temperature prey growth rate should increase faster than predator attack rate. Increasing temperature thus weakens the Allee effect. In the latter, gypsy moth population model, mating rate increases with warming due to enhanced female?male encounter rate and temperature‐induced modifications in female and male adult emergence distributions. However, male and female mortality rates increase, too and the net effect is strengthening of the Allee effect. These results have repercussions also for pest control, indicating that augmentation of biocontrol agents may perhaps be not as effective as using pesticides or disrupting mating.