Hydrology,water availability and tundra ecosystem function in a changing climate: the need for a closer integration of ideas?

Hydrologists and ecologists studying tundra ecosystems have worked largely independently, with little cross-fertilization between disciplines. Their disciplines are, however, inextricably linked by a need to understand the dynamics and significance of the common substance water, in its liquid, solid and gaseous state within tundra environments. The impacts of predicted long-term changes in climate have particularly important consequences for the functioning of tundra systems and there is a pressing need to initiate studies that integrate hydrological and ecological methodologies and concepts. Our paper attempts to summarize existing information on the role of water within tundra ecosystems, to emphasize the fundamental links between the biotic and the physico/chemical environments and to suggest how a closer integration of ideas might be achieved. Given the breadth of the subject matter the paper is intended to be illustrative rather than comprehensive. The paper examines the physical impacts of water in its various states on the tundra environment, emphasizing in particular the causes of spatial variation in water availability to living organisms. The significance of water is discussed for a range of organism groups, including plants, invertebrates and microorganisms and its pivotal role in ecosystem function and disturbance stressed. The need to develop integrated hydological/ecological models for tundra systems on different spatial scales is emphasized.     

西南高山地区土壤异养呼吸时空动态

土壤异养呼吸是陆地和大气之间的重要通量,是决定陆地生态系统碳源汇的关键因素之一,与气候变化紧密相关。西南高山地区是响应气候变化的重点区域,研究西南高山地区土壤异养呼吸动态及其对气候变化的响应,对于评估区域碳循环对全球气候变化的贡献具有重要意义。应用生态系统模型(CEVSA)模型估算了1954-2010年西南高山地区土壤异养呼吸(HR)的时空变化,分析了其对气候变化的响应。结果表明:(1)西南高山地区1954-2010年平均异养呼吸量为422 g C·m^-2·a^-1,在空间分布上,HR自东南向西北递减,与年平均温度(r=0.721,P<0.01)、年降水量(r=0.564,P<0.01)均显著正相关;(2)在时间尺度上,西南高山地区1954-2010年 HR总量增加趋势显著(P<0.05),变化范围为197-251 Tg C/a,平均每年增加0.710 Tg C,其中主要植被类型草地、常绿针叶林和常绿阔叶林均增加趋势显著(P<0.01),增加速度分别为1.621、1.496和1.055 g C·m^-2·a^-2。(3)土壤HR的年际变化主要受温度影响,且西北部高海拔地区较东南部低海拔对温度变化更为敏感,主要植被类型温度敏感系数Q₁₀从大到小依次为草地(2.35)、常绿针叶林(2.34)、常绿阔叶林(1.93)。     

The evidence for shrub expansion in Northern Alaska and the Pan-Arctic

One expected response to climate warming in the Arctic is an increase in the abundance and extent of shrubs in tundra areas. Repeat photography shows that there has been an increase in shrub cover over the past 50 years in northern Alaska. Using 202 pairs of old and new oblique aerial photographs, we have found that across this region spanning 620 km east to west and 350 km north to south, alder, willow, and dwarf birch have been increasing, with the change most easily detected on hill slopes and valley bottoms. Plot and remote sensing studies from the same region using the normalized difference vegetation index are consistent with the photographic results and indicate that the smaller shrubs between valleys are also increasing. In Canada, Scandinavia, and parts of Russia, there is both plot and remote sensing evidence for shrub expansion. Combined with the Alaskan results, the evidence suggests that a pan-Arctic vegetation transition is underway. If continued, this transition will alter the fundamental architecture and function of this ecosystem with important ramifications for the climate, the biota, and humans.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2+/-1.2 and 7.8+/-1.1microg CH4 m-2h-1 was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8+/-1.3microg CH4 m-2h-1 on a regional basis; plants from alpine meadows contribute at least 0.13Tg CH4 yr-1 in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.     

气候变化对不同水分条件下柿幼树光合作用的影响

以两年生柿树为试材,研究了高浓度CO₂处理（700 μmol·mol^-1）、高温处理（日平均温度高于正常日平均温度约5 ℃）、高温和高浓度CO₂复合处理及对照（温度为外界环境温度,CO₂浓度380 μmol·mol^-1）对不同水分条件下柿树净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)、水分利用效率(WUE)、叶绿素含量和叶绿素荧光参数F_v/F_m、F_v/F_o的影响.结果表明:高温和高浓度CO₂复合处理使处于各水分条件下的柿树T_r、G_s减小,WUE增大;在高、中水分条件(分别为田间持水量的75%～85%和55%～65%)下高温和高浓度CO₂复合处理使柿树P_n增大,在低水分条件(为田间持水量的35%～45%)下使柿树P_n减小.高浓度CO₂处理使处于各水分条件下的柿树P_n、WUE增大,G_s、T减小.高温及高温和高浓度CO₂复合处理下WUE均受土壤水分状况的影响,并随土壤含水量的升高而升高.与对照相比,高浓度CO₂处理还提高了各水分条件下植株叶片水平的水分利用效率、叶绿素a、叶绿素b、叶绿素（a+b）、类胡萝卜素含量及F_v/F_m和F_v/F_o,缓解了水分胁迫,提高了柿树的抗逆能力.     

Advancing the long view of ecological change in tundra systems

Despite uncertainties related to sustained funding, ideological rivalries and the turnover of research personnel, long-term studies and studies espousing a long-term perspective in ecology have a history of contributing landmark insights into fundamental topics, such as population- and community dynamics, species interactions and ecosystem function. They also have the potential to reveal surprises related to unforeseen events and non-stationary dynamics that unfold over the course of ongoing observation and experimentation. The unprecedented rate and magnitude of current and expected abiotic changes in tundra environments calls for a synthetic overview of the scope of ecological responses these changes have elicited. In this special issue, we present a series of contributions that advance the long view of ecological change in tundra systems, either through sustained long-term research, or through retrospective or prospective modelling. Beyond highlighting the value of long-term research in tundra systems, the insights derived herein should also find application to the study of ecological responses to environmental change in other biomes as well.     

Carbon exchange in biological soil crust communities under differential temperatures and soil water contents: implications for global change

Biological soil crusts (biocrusts) are an integral part of the soil system in arid regions worldwide, stabilizing soil surfaces, aiding vascular plant establishment, and are significant sources of ecosystem nitrogen and carbon. Hydration and temperature primarily control ecosystem CO₂ flux in these systems. Using constructed mesocosms for incubations under controlled laboratory conditions, we examined the effect of temperature (5–35 °C) and water content (WC, 20–100%) on CO₂ exchange in light (cyanobacterially dominated) and dark (cyanobacteria/lichen and moss dominated) biocrusts of the cool Colorado Plateau Desert in Utah and the hot Chihuahuan Desert in New Mexico. In light crusts from both Utah and New Mexico, net photosynthesis was highest at temperatures >30 °C. Net photosynthesis in light crusts from Utah was relatively insensitive to changes in soil moisture. In contrast, light crusts from New Mexico tended to exhibit higher rates of net photosynthesis at higher soil moisture. Dark crusts originating from both sites exhibited the greatest net photosynthesis at intermediate soil water content (40–60%). Declines in net photosynthesis were observed in dark crusts with crusts from Utah showing declines at temperatures >25 °C and those originating from New Mexico showing declines at temperatures >35 °C. Maximum net photosynthesis in all crust types from all locations were strongly influenced by offsets in the optimal temperature and water content for gross photosynthesis compared with dark respiration. Gross photosynthesis tended to be maximized at some intermediate value of temperature and water content and dark respiration tended to increase linearly. The results of this study suggest biocrusts are capable of CO₂ exchange under a wide range of conditions. However, significant changes in the magnitude of this exchange should be expected for the temperature and precipitation changes suggested by current climate models.     

长白山高山冻原种子植物的调查

通过多次对长白山高山冻原的实地考察,统计得到冻原种子植物共计34科、94属、146种,有6种属的分布区类型(包括3个变型),北温带分布类型最多,占总属数的68.09%。在整个冻原上,草本有125种,占总数的85.62%,其中多年生草本124种。高山冻原植物中具有多种生态,生理适应方式,低矮平卧状植株、极矮小草本、密集丛生是植物的重要适应方式,细长密集须根是冻原上主要的适应方式,占植物总数的54.94%。果实中干果共计137种,其中蒴果56种。由于高山冻原上生态环境十分残酷,极易受人为因素的影响。     

西南高山地区水分利用效率时空动态及其对气候变化的响应

水分利用效率是深入理解生态系统水碳循环耦合关系的重要指标。西南高山地区是响应气候变化的重点区域,研究西南高山地区水分利用效率动态及其对气候变化的响应,对于评估区域碳水耦合关系及对全球气候变化的响应具有重要意义。应用生态系统模型CEVSA(Carbon Exchange between Vegetation,Soil,and the Atmosphere)估算了1954—2010年西南高山地区水分利用效率(Water use efficiency,WUE)的时空变化,分析了其对气候变化的响应。结果表明:(1)西南高山地区1954—2010年水分利用效率均值为1.13 g C mm-1m-2。3种主要植被类型草地、常绿针叶林和常绿阔叶林的WUE分别为1.35、1.14、0.99 g C mm-1m-2。在空间分布上,WUE与海拔显著正相关(r=0.156,P0.05),而与温度则显著负相关(r=-0.386,P0.01)。(2)在时间尺度上,1954—2010年西南高山地区整体WUE降低趋势显著(P0.01),变动区间为0.83-1.46g C mm-1m-2,平均每年下降0.006g C mm-1m-2。整体WUE年际变化与温度呈显著负相关(r=-0.727,P0.01),与降水量相关性不显著;整体WUE下降主要原因是温度上升引起的ET增加速率大于NPP增加速率。(3)1954—2010年西南高山地区3种主要植被类型草地、常绿针叶林及常绿阔叶林WUE均显著下降(P0.01),下降速度分别为-1.03×10-2、-6.17×10-3、-1.37×10-3g C mm-1m-2a-1。西南高山地区76.3%格点WUE年际变化与温度显著负相关(P0.05),34.1%格点WUE年际变化与降水量显著正相关(P0.05)。草地和常绿针叶林WUE年际变化与温度显著负相关(r=-0.889,P0.01;r=-0.863,P0.01),与降水量相关性不显著。由于西南高山地区降水较为丰富,且过去57年降水变化不显著,因此该地区WUE的时空格局主要受温度变化的影响。1954—2010年期间温度升高造成的ET增加显著高于NPP的增加是该地区WUE下降的主要原因。未来需要获取更高空间分辨率的气候、土壤、植被数据,从而更加准确和精确地模拟西南高山地区水碳循环及其耦合关系对气候变化的响应。     

Dynamic and complex microclimate responses to warming and grazing manipulations

Synthesis efforts that identify patterns of ecosystem response to a suite of warming manipulations can make important contributions to climate change science. However, cross‐study comparisons are impeded by the paucity of detailed analyses of how passive warming and other manipulations affect microclimate. Here we document the independent and combined effects of a common passive warming manipulation, open‐top chambers (OTCs), and a simulated widespread land use, clipping, on microclimate on the Tibetan Plateau. OTCs consistently elevated growing season averaged mean daily air temperature by 1.0–2.0°C, maximum daily air temperature by 2.1–7.3°C and the diurnal air temperature range by 1.9–6.5°C, with mixed effects on minimum daily air temperature, and mean daily soil temperature and moisture. These OTC effects on microclimate differ from reported effects of a common active warming method, infrared heating, which has more consistent effects on soil than on air temperature. There were significant interannual and intragrowing season differences in OTC effects on microclimate. For example, while OTCs had mixed effects on growing season averaged soil temperatures, OTCs consistently elevated soil temperature by approximately 1.0°C early in the growing season. Nonadditive interactions between OTCs and clipping were also present: OTCs in clipped plots generally elevated air and soil temperatures more than OTCs in nonclipped plots. Moreover, site factors dynamically interacted with microclimate and with the efficacy of the OTC manipulations. These findings highlight the need to understand differential microclimate effects between warming methods, within warming method across ecosystem sites, within warming method crossed with other treatments, and within sites over various timescales. Methods, sites and scales are potential explanatory variables and covariables in climate warming experiments. Consideration of this variability among and between experimental warming studies will lead to greater understanding and better prediction of ecosystem response to anthropogenic climate warming.     

Last‐century changes of alpine grassland water‐use efficiency: a reconstruction through carbon isotope analysis of a time‐series of Capra ibex horns

The ecophysiological response of an alpine grassland to recent climate change and increasing atmospheric CO₂ concentration was investigated with a new strategy to go back in time: using a time‐series of Capra ibex horns as archives of the alpine grasslands' carbon isotope discrimination (¹³Δ). From the collection of the Natural History Museum of Bern, horns of 24 males from the population of the Augstmatthorn–Brienzer Rothorn mountains, Switzerland, were sampled covering the period from 1938 to 2006. Samples were taken from the beginning of each year‐ring of the horns, representing the beginning of the horn growth period, the spring. The horns' carbon ¹³C content (Δ¹³C) declined together with that of atmospheric CO₂ over the 69‐year period, but ¹³Δ increased slightly (+0.4‰), though significantly (P<0.05), over the observation period. Estimated intercellular CO₂ concentration increased (+56 μmol mol^?1) less than the atmospheric CO₂ concentration (+81 μmol mol^?1), so that intrinsic water‐use efficiency increased by 17.8% during the 69‐year period. However, the atmospheric evaporative demand at the site increased by approximately 0.1 kPa between 1955 and 2006, thus counteracting the improvement of intrinsic water‐use efficiency. As a result, instantaneous water‐use efficiency did not change. The observed changes in intrinsic water‐use efficiency were in the same range as those of trees (as reported by others), indicating that leaf‐level control of water‐use efficiency of grassland and forests followed the same principles. This is the first reconstruction of the water‐use efficiency response of a natural grassland ecosystem to last century CO₂ and climatic changes. The results indicate that the alpine grassland community has responded to climate change by improving the physiological control of carbon gain to water loss, following the increases in atmospheric CO₂ and evaporative demand. But, effective leaf‐level water‐use efficiency has remained unchanged.