青藏高原高寒草甸非生长季温室气体排放特征及其年度贡献

高寒草甸是青藏高原地区的主要植被类型,目前对其温室气体研究多集中于生长季.本文利用静态箱-气相色谱法,对非生长季高寒草甸温室气体排放特征及其与主要环境因子的关系进行了研究.结果表明:非生长季高寒草甸表现为CO2和N2O的源、CH4的汇.其中非生长季CO2通量平均值为89.33 mg·m-2·h-1,累积排放通量为280.01g· m-2;CH4通量平均值为-11.35 μg·m-2·h-1,累积吸收通量为124.74 mg·m-2;N2O通量平均值为8.02 μg·m-2·h-1,累积排放通量为39.51 mg·m-2.非生长季CO2、CH4和N2O累积排放通量分别占全年的13.33％、53.47％和62.67％.冻融期(2012年4月)CH4累积吸收通量较小,只占非生长季的4.5％;而CO2和N2O累积排放通量较大,分别占非生长季的25.8％和20.8％.非生长季CO2通量与温度(气温、5和10 cm土壤温度)和5 cm土壤湿度均存在显著正相关关系,而CH4和N2O通量仅与5 cm土壤湿度存在显著正相关.研究表明,虽然冻融期CH4累积吸收通量在非生长季累积量中比重较小,但非生长季CH4和N2O累积排放量却占全年累积排放量的1/2以上,在温室气体累积通量评估中不容忽视.     

非生长季降水对青藏高原高寒草甸优势种生物量稳定性的影响

受全球气候变化的影响,青藏高原在过去的几十年间整体上呈现暖湿化的趋势,相比于年际之间温度和降水的变化外,生长季和非生长季气候变化模式的差异可能会对生态系统产生更重要的影响,但相关的研究尚不充分。以青藏高原东部的高寒草甸为研究对象,基于2001年至2017年17年的野外观测数据,包括优势植物紫花针茅的高度、多度以及生物量、次优势物种洽草的生物量,结合生长季和非生长季平均温度和降水量的变化,通过线性回归以及结构方程模型,探究生长季/非生长季不对称气候变化对于青藏高原高寒草甸优势物种生物量稳定性的影响。研究结果表明：1)青藏高原东部年均温和年降水在过去的17年间显著增加,呈现暖湿化的趋势,但是非生长的降水却变化不明显;2)紫花针茅的高度、多度以及生物量在过去17年没有显著的趋势,但是洽草的生物量稳定性显著减少;3)非生长降水结合紫花针茅的高度、多度以及洽草的生物量稳定性促进了紫花针茅的生物量稳定性。研究结果可以为青藏高原高寒草甸在未来气候变化的背景下合理保护与利用提供科学依据。     

日尺度下水热因子变化对青藏高原高寒草原生产力的影响特征

温度和降水变化显著影响高寒生态系统植被生长和系统功能。草地生产力作为草地系统功能强弱的重要体现,对气候变化,特别是温度和降水变化十分敏感。探究高寒草原生产力如何响应气候变化,对预测未来气候变化情景下高寒草地系统功能变化意义重大。前期研究大都从年或季节尺度探究气候变化对草地生产力的影响特征,缺乏更精细时间尺度的关联分析。本研究基于1997—2020年青藏高原高寒草原长期植被观测数据及相应气候资料,应用简单线性回归及偏最小二乘回归法(Partial Least Squares regression, PLS)探究了研究区草地地上净初级生产力对日尺度温度和降水变化响应特征。结果表明：(1)近24 a来研究区年平均气温和降水量分别以0.03℃/a和4.36 mm/a的速率显著升高;(2)近24 a来研究区草地生产力显著升高(增幅为5.24 g m^-2 a^-1),且与年平均温度和降水量呈显著正相关关系;(3)日尺度分析表明,不同阶段温度和降水变化对草地生产力的影响不同,其中5—8月和9—10月的温度及5—7月和9—11月的降水是影响研究区草地生产力的气...     

青藏高原高寒草甸夏季植被特征及对模拟增温的短期响应

以青藏高原高寒草甸为研究对象,研究了草甸植被夏季生长动态特征;同时采用红外辐射器模拟增温的方法,探讨了草甸植被对增温的短期(1a)响应.结果表明:(1)高寒草甸夏季植被高度与地下生物量、总生物量相关性不显著,盖度与二者相关性极显著;高度对地上生物量影响较大(R=0.892,P＜0.01),盖度对地下生物量(R=0.883,P＜0.01)和总生物量(R=0.888,P＜0.01)影响较大.(2)高寒草甸夏季植被地上部分和地下部分表现出不同的生长模式,地上部分近似等速生长(幂指数为1.011),地下部分则表现为异速生长(幂指数为0.459),但整体呈现异速生长(幂指数为0.473).(3)高寒草甸夏季植被地上生物量(P＜0.05)在6月份较地下生物量(P＞0.05)对环境更为敏感,且一年之后地上-地下生物量均呈减小趋势,这与空气温度、土壤温度和土壤水分的显著减小密切相关.(4)红外辐射器在高寒草甸的增温度效果较好,空气、地表、土壤温度都随增温幅度增强而增加;短期增温对高寒植被有正效应(T0-T1),而温度持续升高则对植被产生负效应(T1-T2);各植被指标的方差分析都未达到显著水平,表明短期增温对该植被影响不显著.     

青藏高原海北高寒湿地土壤呼吸对水位降低和氮添加的响应

近20年来, 青藏高原高寒湿地经历了明显的气候变化, 从而导致多数湿地水位下降和氮沉降的增加。对于湿地生态系统来说, 水位下降意味着土壤通气性能的改善, 可能会导致土壤呼吸的增加; 而氮沉降的增加可能会降低土壤微生物生物量和pH值, 从而可能抑制土壤呼吸。为此, 在青海海北高寒草地生态系统国家野外科学观测研究站利用中宇宙(Mesocosm)实验方法, 探讨了青藏高原高寒泥炭型湿地土壤呼吸对水位降低和氮添加的响应。结果表明: (1)水位降低显著增强了土壤呼吸, 而氮添加对土壤呼吸的影响依赖于水位的变化: 对照水位下, 氮添加显著抑制土壤呼吸; 而水位降低时, 氮添加对土壤呼吸速率无显著影响。(2)土壤呼吸速率与地上生物量、枯落物累积量之间呈显著正相关关系, 而与根系生物量无显著相关关系。(3)水位降低显著提高了土壤呼吸的温度敏感性, 而氮添加对其无显著的影响。因此预测: 随着氮沉降的升高, 高寒泥炭湿地土壤CO₂的排放量将会减少; 然而随着暖干化背景下水位的降低, 青藏高原高寒湿地会排放更多的CO₂。     

基于年降水、生长季降水和生长季蒸散的高寒草地水分利用效率

水分利用效率是深入理解生态系统碳、水循环间耦合关系的重要指标。以前研究青藏高原的水分利用效率多基于年降水量(AP)来分析, 但植物对水分的利用主要在生长季。该研究采用以AP、生长季降水量(GSP)和生长季蒸散量(ET_gs)分别计算的年降水利用效率(PUE_a)、生长季降水利用效率(PUE_gs)和生长季水分利用效率(WUE_gs), 分析了2000-2010年间青藏高原两种主要植被类型高寒草甸和高寒草原PUE_a、PUE_gs和WUE_gs的差异及其与降水量、蒸散量和气温的关系。结果表明: (1)高寒草甸的PUE_a、PUE_gs均大于高寒草原, 但两种草地类型的WUE_gs无显著差别, 这说明两种草地类型可能存在相似的内在的水分利用效率。(2)从年际动态来看, PUE_a和PUE_gs的波动范围相似, 而WUE_gs的波动范围更大, 说明以蒸散为依据的WUE_gs可能比PUE_a和PUE_gs更敏感, 因而可能更好地反映生态系统的水分利用能力。(3)高寒草甸和高寒草原的PUE_a、PUE_gs和WUE_gs分别与AP、GSP和ET_gs呈单调递减趋势, 表明3种水分利用效率均随降水量或蒸散量的增加而降低。高寒草原的3种水分利用效率中仅WUE_gs随着气温的增加而增加, 而高寒草甸的3种水分利用效率均与气温无显著关系, 这说明相比高寒草甸, 高寒草原的水分利用效率对气温更加敏感。     

2000-2021年青藏高原生长季植被敏感性的时空变异

为揭示青藏高原陆地生态系统对气候变化敏感性的时空变异性,基于植被敏感性指数(Vegetation Sensitivity Index, VSI),使用2000—2021年青藏高原6—8月生长季MODIS EVI和ERA5再分析资料的温度、降水和太阳辐射数据,首先探究了22年里青藏高原陆地生态系统敏感性的空间变异性及其主要气候驱动因素,其次探究了青藏高原VSI在P₁(2000—2006年)、P₂(2007—2013年)和P₃(2014—2021年)时期内VSI的时间变异性,研究表明：(1)2000—2021年青藏高原生长季VSI的空间异质性较强,其中东南部灌木和森林的VSI较高,而西北部高山荒漠、高山草原和高山草甸的VSI较低;(2)22年里温度、降水和太阳辐射分别主导着青藏高原55.89%、19.24%和24.87%地区的VSI变化,其中温度主导着东南部灌木和森林的VSI,降水主导着东北大部分地区高山草甸的VSI,而太阳辐射主导着西南大部分地区高山草原的VSI。时间变异性结果表明：(3)P₁—P     

青藏高原高寒灌丛2003-2016年生长季的不同月份CO2通量对气温日较差的响应

全球气候变化引起的气温日较差（ADT）减小,将会对高寒生态系统的碳平衡造成深刻影响。基于涡度相关系统,利用2003-2016年的涡度相关系统观测资料,做了青藏高原高寒灌丛在生长季（6-9月）不同月份的ADT对CO₂通量影响的研究。结果表明：2003-2016年的生长季中,最高气温（MaxTa）和最低气温（MinTa）呈先升高后降低的单峰变化趋势,ADT没有呈现明显的变化趋势。逐日总初级生产力（GPP）和生态系统呼吸（Re）呈先增加后降低的单峰趋势,逐日净生态系统CO₂交换（NEE）呈先下降后上升的"V"型变化趋势。高寒灌丛在生长季为碳汇,整个生长季总NEE、GPP和Re平均值分别为（-161.2±30.1）、（501.9±60.2）、（340.7±54.4） gCm^-2。在高寒灌丛生长季（6-9月）的每个月份,MaxTa、MinTa和ADT分别是GPP（P<0.001）、Re（P<0.001）和NEE（P<0.01）变化的主要控制因子。高寒灌丛的ADT的增大有利于生态系统的碳固持,暗示在未来气候变化背景下ADT的减小将会削弱高寒灌丛生态系统的碳汇能力。     

青藏高原高寒灌丛非生长季节CO2通量特征

利用2003年和2004年涡度相关系统通量观测资料,对青藏高原高寒灌丛非生长季节CO2通量特征及其主要影响因子进行了分析。(1)从净生态系统CO2交换(NEE)日变化特征看,除13:00~19:00时有较小的CO2净释放以外,其余时段NEE均很小;(2)高寒灌丛非生长季月份间NEE差异明显,4月和10月是CO2净释放量较大,1月和12月CO2净释放量较小;(3)相对温带草原(高杆草大草原)草地类型,低温抑制下的青藏高原高寒灌丛生态系统非生长季节日平均CO2释放率较低;(4)高寒灌丛非生长季NEE日变化模式与5 cm土壤温度变化呈显著正相关,土壤温度是影响非生长季节青藏高原高寒灌丛NEE变化的主导气候因子,同时NEE变化还受降水的影响。     

青藏高原东北部祁连圆柏径向生长对不同类型干旱的响应

为研究祁连圆柏径向生长对不同时期(生长季前2—4月和生长季5—7月)气候因子的响应及面对不同类型(高温、缺水、高温+缺水)干旱事件的弹性(抵抗力和恢复力)变化,利用青藏高原东北部17个采样点的祁连圆柏树轮宽度资料,分析径向生长与不同时期气候因子的相关性,探究高低海拔祁连圆柏面对各类干旱事件的弹性差异。结果表明:祁连圆柏径向生长与干旱指数呈显著正相关,与生长季温度呈负相关(P<0.1)。祁连圆柏面对不同时期干旱事件的弹性存在显著差异,生长季前发生的干旱事件中,低海拔圆柏的抵抗力比高海拔增高2.3%,恢复力降低25.1%;生长季干旱事件中,低海拔圆柏的抵抗力比高海拔降低23.7%,恢复力增高107.1%。祁连圆柏面对缺水型干旱时恢复力更强,均值达到1.68,而面对高温型干旱时祁连圆柏的抵抗力更强,均值达到1.43。未来我国西部高山祁连圆柏,尤其是处于低海拔区的,其径向生长受到全球变暖造成的极端干旱事件的影响会更加显著。     

Net primary productivity and spatial distribution of vegetation in an alpine wetland,Qinghai-Tibetan Plateau

To initially describe vegetation structure and spatial variation in plant biomass in a typical alpine wetland of the Qinghai-Tibetan Plateau, net primary productivity and vegetation in relationship to environmental factors were investigated. In 2002, the wetland remained flooded to an average water depth of 25 cm during the growing season, from July to mid-September. We mapped the floodline and vegetation distribution using GPS (global positioning system). Coverage of vegetation in the wetland was 100%, and the vegetation was zonally distributed along a water depth gradient, with three emergent plant zones (Hippuris vulgaris-dominated zone, Scirpus distigmaticus-dominated zone, and Carex allivescers-dominated zone) and one submerged plant zone (Potamogeton pectinatus-dominated zone). Both aboveground and belowground biomass varied temporally within and among the vegetation zones. Further, net primary productivity (NPP) as estimated by peak biomass also differed among the vegetation zones; aboveground NPP was highest in the Carex-dominated zone with shallowest water and lowest in the Potamogeton zone with deepest water. The area occupied by each zone was 73.5% for P. pectinatus, 2.6% for H. vulgaris, 20.5% for S. distigmaticus, and 3.4% for C. allivescers. Morphological features in relationship to gas-transport efficiency of the aerial part differed among the emergent plants. Of the three emergent plants, H. vulgaris, which dominated in the deeper water, showed greater morphological adaptability to deep water than the other two emergent plants.     

青藏高原高寒湿地生态系统CO2通量

依据涡度相关系统连续观测的2005年CO2通量数据,对青藏高原东北隅的高寒湿地生态系统源/汇功能及其部分环境影响因素进行了分析.结果表明,高寒湿地生态系统为明显的碳源,在植物生长季(5～9月份)吸收230.16 gCO2·m-2,非生长季(1～4月份及10～12月份)释放546.18 gCO2·m-2,其中净排放最高在5月份,为181.49 gCO2·m-2,净吸收最高在8月份,为189.69 gCO2·m-2,年释放量为316.02 gCO2·m-2.在平均日变化中,最大吸收值出现在7月份12:00,为(0.45±0.0012) mgCO2·m-2·s-1,最大排放速率出现在8月份0:00,为(0.22±0.0090) mgCO2·m-2·s-1.生长季中6～9月份表现为明显的单峰型日变化,非生长季的变化幅度较小.净生态系统交换量(NEE)和生态系统总初级生产力(GPP)与气温、空气水气饱和亏和地表反射率等环境因素呈现相似的相关性,与地上生物量和群落叶面积指数则为线性负相关,生态系统呼吸(Res)则与上述因子的相关性呈现相反的趋势.     

青藏高原高寒湿地生态系统CO2通量

依据涡度相关系统连续观测的2005年CO₂通量数据,对青藏高原东北隅的高寒湿地生态系统源/汇功能及其部分环境影响因素进行了分析。结果表明,高寒湿地生态系统为明显的碳源,在植物生长季（5~9月份）吸收230.16 gCO₂•m^-2,非生长季（1~4月份及10~12月份）释放546.18 gCO₂•m^-2,其中净排放最高在5月份,为181.49 gCO₂•m^-2,净吸收最高在8月份,为189.69 gCO₂•m^-2,年释放量为316.02 gCO₂•m^-2。在平均日变化中,最大吸收值出现在7月份12:00,为（0.45±0.0012） mgCO2•m^-2•s^-1,最大排放速率出现在8月份0:00,为（0.22±0.0090） mgCO₂•m^-2•s^-1。生长季中6~9月份表现为明显的单峰型日变化,非生长季的变化幅度较小。净生态系统交换量（NEE）和生态系统总初级生产力（GPP）与气温、空气水气饱和亏和地表反射率等环境因素呈现相似的相关性,与地上生物量和群落叶面积指数则为线性负相关,生态系统呼吸（Res）则与上述因子的相关性呈现相反的趋势。     

CO2 flux in alpine wetland ecosystem on the Qinghai-Tibetan Plateau, China

Using the CO₂ flux data measured by the eddy covariance method in the northeast of Qinghai-Tibetan Plateau in 2005, we analyzed the carbon flux dynamics in relation to meteorological and biotic factors. The results showed that the alpine wetland ecosystem was the carbon source, and it emitted 316.02 gCO₂ · m⁻² to atmosphere in 2005 with 230.16 gCO₂ · m⁻² absorbed in the growing season from May to September and 546.18 gCO₂ · m⁻² released in the non-growing season from January to April and from October to December. The maximum of the averaged daily CO₂ uptake rates and release rates was (0.45 ± 0.0012) mgCO₂ · m⁻² · s⁻¹ (Mean ± SE) in July and (0.22 ± 0.0090) mgCO₂ · m⁻² · s⁻¹ in August, respectively. The averaged diurnal variation showed a single-peaked pattern in the growing season, but exhibited very small fluctuation in the non-growing season. Net ecosystem exchange (NEE) and gross primary production (GPP) were all correlated with some meteorological factors, and they showed a negatively linear correlation with aboveground biomass, while a positive correlation existed between the ecosystem respiration (R_es) and those factors.     

Immediate and carry-over effects of late-spring frost and growing season drought on forest gross primary productivity capacity in the Northern Hemisphere

Forests are increasingly exposed to extreme global warming-induced climatic events. However, the immediate and carry-over effects of extreme events on forests are still poorly understood. Gross primary productivity (GPP) capacity is regarded as a good proxy of the ecosystem's functional stability, reflecting its physiological response to its surroundings. Using eddy covariance data from 34 forest sites in the Northern Hemisphere, we analyzed the immediate and carry-over effects of late-spring frost (LSF) and growing season drought on needle-leaf and broadleaf forests. Path analysis was applied to reveal the plausible reasons behind the varied responses of forests to extreme events. The results show that LSF had clear immediate effects on the GPP capacity of both needle-leaf and broadleaf forests. However, GPP capacity in needle-leaf forests was more sensitive to drought than in broadleaf forests. There was no interaction between LSF and drought in either needle-leaf or broadleaf forests. Drought effects were still visible when LSF and drought coexisted in needle-leaf forests. Path analysis further showed that the response of GPP capacity to drought differed between needle-leaf and broadleaf forests, mainly due to the difference in the sensitivity of canopy conductance. Moreover, LSF had a more severe and long-lasting carry-over effect on forests than drought. These results enrich our understanding of the mechanisms of forest response to extreme events across forest types.     

青藏高原高寒草甸植被特征与温度、水分因子关系

以广布于青藏高原的高寒草甸为研究对象,进行模拟增温实验,探讨高寒草甸植被特征与温度、水分因子关系,并试图论证高寒草甸植被是否符合生物多样性代谢理论.结果表明:①高寒草甸植被物种多样性的对数与绝对温度的倒数呈显著线性递减关系,空气-地表-浅层土壤(0-20 cm)温度(R2 ＞0.6,P＜0.01)较深层土壤(40-100 cm)温度(R2＜0.5,P＜0.05)对物种多样性影响大;其植被新陈代谢平均活化能为0.998-1.85 eV,高于生物多样性代谢理论预测值0.6-0.7 eV,这是高寒草甸植被对长期低温环境适应进化的结果.②除趋势对应分析和冗余分析显示,温度对植被地上部分影响较大,而土壤水分对全株影响均较大,适当的增温与降水均可极显著促进高寒草甸植被生长.③逐步回归和通径分析显示,40 cm、60 cm深度土壤水分对植被地上部分产生直接影响,20 cm高度空气相对湿度和40 cm深度土壤温度对其产生间接影响;40 cm深度土壤温度和60 cm深度土壤水分对植被地下部分产生直接影响,红外地表温度对其产生间接影响.深层土壤温度和水分对高寒草甸植被具有影响作用,这可能与增温后冻土的融化有关,但其机理尚待进一步研究.     

A growing season climatic index to simulate gross primary productivity and carbon budget in a Tibetan alpine meadow

Accurate estimation of gross primary production (GPP) of ecosystem is needed to evaluate terrestrial carbon cycle at various spatial and temporal scales. Eddy covariance (EC) technique provides continuous measurements of net ecosystem CO₂ exchange (NEE) and can be used to separate GPP from NEE in real time series. However, seasonal and inter-annual variation and consequently ecosystem carbon budget is still very difficult to simulate from climatic and environment. To address this limitation, we develop a growing season indicator (GSI) based on low temperature and soil water stress to model and predict intra and inter-annual dynamic of gross primary productivity (GPP). Validation of this new index was conducted using continuous six-year consective EC measurement from 2004 to 2009 at a Tibetan alpine meadow. Simulated GPP agreed well with the observed GPP in terms of seasonal and inter-annual variation. The six-year correlation coefficients on seasonal scale between GSI and scalar GPP derived from EC reached more than 0.85 no matter in dry years or wet years. In addition, the temporal GPP estimation derived from GSI model was quite similar to those from observed values by EC measurement. Moreover, accumulated GSI values can predict annual variability of net ecosystem production (NEP). Higher yearly accumulated GSI corresponded to more annual NEP. When cumulative GSI arrived up to 92, the target ecosystem was a carbon sink. This is probably a threshold which Tibetan alpine meadow changes from carbon source to carbon sink. It is indicated that the GSI model is a simple, alternative approach to estimating GPP and has the potential to simulate spatial GPP in a larger scale. However, the performance of GSI model in other vegetation types or regions still needs a further verification.     

青藏高原高寒草地生态系统服务功能的互作机制

青藏高原高寒草地是我国重要的畜牧业生产基地和生态安全屏障.高寒草地退化不仅影响了当地畜牧业生产和牧民生活,而且严重地威胁着我国和东亚地区的生态安全.从高寒草地生态系统的生态、生产和生活功能角度出发,分析了青藏高原高寒草地生态系统在人口、放牧压力与资源环境承载力的相互作用关系,及其生态、生产和生活功能比例结构变化对高寒草地生态系统服务功能的影响,阐明了高寒草地生态系统服务功能的多元耦合、多维连锁和多重反馈的相互作用机制.并以藏北那曲地区为例,把2008年农牧民脱贫线和小康线作为生活功能标准,通过生态服务功能当量,确定了维持高寒草地生态系统可持续发展的生态、生产和生活功能比例结构.据此,估测了牧民的生活功能达到脱贫线和小康线标准时允许的人口承载量.结果表明,高寒草地退化使草地生态系统的人口承载量下降了60％.以那曲地区2008年的实际的牧业人口量与允许的人口承载量相比,高寒草地退化后,实际牧业人口占脱贫标准的36％,但小康标准超载了118.9％.因此,调控人口承载量是实现青藏高原高寒草地生态系统的生态、生产和生活功能协调发展的关键.     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.