喷灌对藏北高寒草地生产力和物种多样性的影响

通过3a(2008—2010年)的藏北高寒草地喷灌试验,研究了不同喷灌量对草地群落生产力和物种多样性的影响。结果表明,丰水年灌溉对藏北高寒草地的影响较小;而在相对干旱年份灌溉对高寒草地生产力和物种多样性影响显著。喷灌条件下高寒草地生物量显著提高,最高增幅出现在高水(GS)样地中,达到116%。喷灌明显促进物种重要值提高,其中灌木和阔叶杂草比例增加趋势更为明显。不同喷灌条件下优势物种相对重要值均有不同程度的降低,高水处理降低幅度最大。物种多样性方面,喷灌措施能够明显促进高寒草地Simpson指数和Shannon-weiner指数增加(P0.05),E.Pielou均匀度指数无显著变化(P0.05)。Shannon-weiner指数与生物量之间存在显著正相关关系(P0.05)。未来降水增多的气候条件可以减少干旱对高寒草地带来的负面影响,有利于提高草地生产力和维持草地物种多样性,促进高寒草地畜牧业健康发展。     

西藏草地多项供给及调节服务相互作用的时空演变规律

基于统计数据及MODIS产品,计算西藏草地2000与2010年载畜支持服务、肉类供给服务、水源涵养服务和碳吸收服务的功能量。构建Pearson相关系数、总生态系统服务(TES)、供给-调节服务互竞指数(TO),分析了4项生态系统服务相互作用关系随纬度、海拔及时间变化的规律。结果表明:西藏草地两项供给服务和两项调节服务之间分别为相互协同作用,且随时空变化不大。供给服务和调节服务之间的相互作用随海拔和纬度升高时,基本由相互竞争转变为相互协同,在高纬度时则无相互作用。4项服务在低纬度及低海拔呈相互竞争关系时,其生态系统服务总体水平也较低,中纬度及中高海拔呈协同关系时,其总体水平较高。从2000年到2010年,西藏草地4项生态系统服务皆发生较大变化。在变化中,12%的草地供给服务与调节服务发生相互竞争作用,33%的草地则为相互协同关系。发生相互竞争作用的草地主要集中在中低纬度的一江两河地区,而相互协同的区域主要为藏北羌塘高原中南部。     

气候变化对藏北地区草地生产力的影响模拟

利用政府间气候变化委员会(IPCC)排放情景特别报告(SRES)的A2和B2方案,通过区域气候模式系统PRECIS与草地生态模型SPUR相联接,模拟评估未来2071—2100年藏北地区草地生产力的变化。结果表明:2种温室气体排放情景下,温度升高、太阳总辐射降低和降水量增加的区域,各类型草地地上生物量基本呈增加的趋势;降水量减少的区域,高嵩草型草地地上生物量呈减少的趋势;藏北地区的草地生产力不大可能从CO2富集上得到多大好处。     

典型森林和草地生态系统呼吸各组分间的相互关系

生态系统呼吸是陆地生态系统碳收支的重要组成部分,分析其组分间的相互关系对理解生态系统呼吸过程和精确评价生态系统碳收支具有重要意义,也是当前碳循环研究工作的一大难点。本研究利用ChinaFLUX的长白山温带针阔混交林（CBS）,鼎湖山亚热带常绿阔叶林（DHS）和海北灌丛草甸（HBGC）三个典型生态系统的通量观测数据,采用经验统计方法,分析了其在中国典型生态系统中的适用性及敏感性,揭示了生态系统呼吸各组分的动态变化特征及相互关系。结果表明：采用本研究中的呼吸组分拆分方法所获结果与理论推测及实测数据大致相同,拆分结果对净初级生产力与总初级生产力的比值（NPP/GPP）较为敏感,NPP/GPP变化0.1时,自养呼吸在生态系统呼吸中的比例（Ra/RE）改变0.05。各生态系统中,生态系统呼吸及其组分在年内均表现出明显的单峰型变化特征,在夏季生长旺盛的时节达到最大值。异养呼吸与生态系统呼吸的比值（Rh/RE）也具有明显的季节变化,但在生态系统间表现出明显差异,CBS和HBGC分别表现出先增大后减小和先减小后增大的变化趋势,DHS则相对稳定,在0.5附近波动, Ra/RE的季节动态与Rh/RE相反。在年总量上,HBGC主要通过异养呼吸向大气排放CO2,异养呼吸占生态系统呼吸的60%,而CBS和DHS的自养呼吸和异养呼吸所占比重大致相似,异养呼吸占生态系统呼吸的49%。这说明,该统计学模型可以用来进行生态系统呼吸组分的拆分,进而可以为生态系统碳循环过程的精细研究提供参考数据,但今后应加强NPP/GPP的测定,以提高生态系统呼吸拆分的精度。     

滴灌下新疆北部棉田杂草土壤种子库的时空变化

以天山北坡绿洲至沙漠边缘垂直分布的3个试验点,对比分析了漫灌和1a至4、8a的不同滴灌时间的棉田,以及不同试验点和土层深度等不同空间的杂草土壤种子库变化.结果表明,滴灌对棉田杂草种子库影响大,物种数和单位面积的种子库密度的年际间波动明显.连续滴灌改变了杂草土壤种子库的结构和组成,物种数由漫灌27种下降到1a滴灌的20种,滴灌8a后下降到15种,Shannon-Wiener多样性指数达到最低,单位面积的种子密度明显降低.随着滴灌年限的增长,狗尾草、藜、龙葵、反枝苋和凹头苋等喜旱性杂草占总种子库的比例逐渐增加,为滴灌棉田的优势杂草.狗尾草、藜、灰绿藜、龙葵、马齿苋、凹头苋、刺儿菜、黄花蒿、苦苣菜、小蓬草、荠菜、小藜、扁蓄、苘麻、田旋花、野薄荷等物种时间生态位宽度和空间生态位宽度均较大,适应较好,而虎尾草、百脉根、播娘蒿、酸模叶蓼、滨藜、野胡麻等物种生存受到明显影响.受耕作方式影响,杂草种子库主要分布于耕作层,耕作层以下46～50cm种子数最少.水平分布格局分析发现,3个地点杂草种子库的物种相似性较高,滴灌与漫灌之间种子库的物种差异要大于地理位置间的差异.     

Grasshoppers affect grassland ecosystem functioning: Spatial and temporal variation

Using experiments and monitoring, we find that grasshoppers in a grassland ecosystem impact ecosystem functioning (nutrient cycling and primary production) in different ways among sites in the ecosystem. Experiments conducted over many years at two sites (21 and 15 years, respectively) with the same grasshopper and plant species demonstrated that grasshoppers increased nitrogen availability (N) and consequently annual plant production (ANPP) at one site, and decreased N and consequently ANPP at the other site. Comparing the two sites, N increased on average by 8% and up to 21.6%, and resulting ANPP increased on average by 18.6% and up to 33.3%. Grasshoppers increase N and ANPP by preferentially feeding on slower decomposing plants, and the opposite occurs by preferentially feeding on faster decomposing plants. Monitoring 20 random sites in the ecosystem, grasshoppers consistently increased N and ANPP over 3 years at 40% of sites, consistently decreased N and ANPP at 35% of sites, and sometimes increased and decreased N and ANPP at 25% of sites. Therefore, grassland grasshoppers, and insects in many ecosystems, may strongly affect ecosystem functioning.     

藏北高寒草地NPP变化趋势及其对人类活动的响应

基于1981～2004年多年遥感监测数据和气象数据以及其它相关数据,采用CASA(Carnegie-Ames-Stanford Approach)模型估算藏北地区草地植被净第一性生产力(NPP),分析草地植被NPP变化趋势的空间格局及其对人类活动强度的响应。结果表明:近24a以来,藏北绝大部分区域(约占草地总面积的88.61%)草地植被NPP变化趋势不明显;而草地植被NPP变化趋势显著的区域仅占草地总面积的11.39%,其中显著降低约占11.30%,显著增高仅占0.09%。在藏北地区,高海拔区域有较大比例(大于26%)的草地NPP显著降低;坡度在15～30°之间区域的草地NPP变化幅度较大;而坡向对草地NPP变化趋势的影响不大。藏北地区居民点对草地NPP变化趋势的负面影响小于道路影响;从综合影响来看,离道路和居民点越近、人类活动强度及其对草地NPP变化趋势的影响越大,尤其是草地NPP显著增高区域只分布在人类活动强度最大的第一个缓冲区内。     

西藏草地生态系统植被碳贮量及其空间分布格局

在广泛收集资料的基础上,利用平均碳密度方法,估算了西藏高原草地生态系统17类草地植被的碳贮量,并分析了其空间分布格局.结果表明:(1)17类草地植被总面积为8205.194×104hm2,总碳贮量为189.367 Tg (1TgC＝1012g),平均碳密度为2307.895 kgC/hm2,不同植被类型差异较大,在395.977～20471.161kgC/hm2之间波动;(2)从草地类型分布看,高寒草原和高寒草甸是西藏分布面积最大的2类草地,分布面积占西藏草地总面积的70.210%,又是西藏草地碳贮量的主要贮库,碳贮量占西藏草地总碳贮量的79.393%;(3)在空间分布格局上,随着自藏东南向西北的延伸,草地植被总碳密度逐次降低,这一水平分布格局与西藏独特的水热分布相一致;碳密度的垂直分布规律因地区而异,但各地区均以高寒草甸或高寒荒漠的低碳密度为终点,表现出"殊途同归"的特征.     

Disentangling drought-induced variation in ecosystem and soil respiration using stable carbon isotopes

Combining C flux measurements with information on their isotopic composition can yield a process-based understanding of ecosystem C dynamics. We studied the variations in both respiratory fluxes and their stable C isotopic compositions (δ¹³C) for all major components (trees, understory, roots and soil microorganisms) in a Mediterranean oak savannah during a period with increasing drought. We found large drought-induced and diurnal dynamics in isotopic compositions of soil, root and foliage respiration (δ¹³C_res). Soil respiration was the largest contributor to ecosystem respiration (R _eco), exhibiting a depleted isotopic signature and no marked variations with increasing drought, similar to ecosystem respired δ¹³CO₂, providing evidence for a stable C-source and minor influence of recent photosynthate from plants. Short-term and diurnal variations in δ¹³C_res of foliage and roots (up to 8 and 4‰, respectively) were in agreement with: (1) recent hypotheses on post-photosynthetic fractionation processes, (2) substrate changes with decreasing assimilation rates in combination with increased respiratory demand, and (3) decreased phosphoenolpyruvate carboxylase activity in drying roots, while altered photosynthetic discrimination was not responsible for the observed changes in δ¹³C_res. We applied a flux-based and an isotopic flux-based mass balance, yielding good agreement at the soil scale, while the isotopic mass balance at the ecosystem scale was not conserved. This was mainly caused by uncertainties in Keeling plot intercepts at the ecosystem scale due to small CO₂ gradients and large differences in δ¹³C_res of the different component fluxes. Overall, stable isotopes provided valuable new insights into the drought-related variations of ecosystem C dynamics, encouraging future studies but also highlighting the need of improved methodology to disentangle short-term dynamics of isotopic composition of R _eco.     

Significant Achievements in Protection and Restoration of Alpine Grassland Ecosystem in Northern Tibet, China

Alpine grassland is a fragile ecosystem, and a large area of this grassland type has been severely degraded in Northern Tibet, to the extent that it has become the primary ecological problem in the region. Various levels of government, including the national central government, the Tibetan Autonomous Region government, and the Nagqu Prefecture government have worked together to achieve alpine grassland ecosystem protection and prevent grassland degradation. These efforts have resulted in significant ecological, social, and economic benefits in Northern Tibet.     

Significance of cold-season respiration and photosynthesis in a subarctic heath ecosystem in Northern Sweden

While substantial cold-season respiration has been documented in most arctic and alpine ecosystems in recent years, the significance of cold-season photosynthesis in these biomes is still believed to be small. In a mesic, subartic heath during both the cold and warm season, we measured in situ ecosystem respiration and photosynthesis with a chamber technique at ambient conditions and at artificially increased frequency of freeze–thaw (FT) cycles during fall and spring. We fitted the measured ecosystem exchange rates to respiration and photosynthesis models with R²-values ranging from 0.81 to 0.85. As expected, estimated cold-season (October, November, April and May) respiration was significant and accounted for at least 22% of the annual respiratory CO₂ flux. More surprisingly, estimated photosynthesis during this period accounted for up to 19% of the annual gross CO₂ uptake, suggesting that cold-season photosynthesis partly balanced the cold-season respiratory carbon losses and can be significant for the annual cycle of carbon. Still, during the full year the ecosystem was a significant net source of 120 ± 12 g C m⁻² to the atmosphere. Neither respiration nor photosynthetic rates were much affected by the extra FT cycles, although the mean rate of net ecosystem loss decreased slightly, but significantly, in May. The results suggest only a small response of net carbon fluxes to increased frequency of FT cycles in this ecosystem.     

Measurement of net ecosystem production and ecosystem respiration in a Zoysia japonica grassland,central Japan,by the chamber method

Measuring light, temperature, soil moisture, and growth provides a better understanding of net ecosystem production (NEP), ecosystem respiration (R _eco), and their response functions. Here, we studied the variations in NEP and R _eco in a grassland dominated by a perennial warm-season C₄ grass, Zoysia japonica. We used the chamber method to measure NEP and R _eco from August to September 2007. Biomass and leaf area index (LAI) were also measured to observe their effects on NEP and R _eco. Diurnal variations in NEP and R _eco were predicted well by light intensity (PPFD) and by soil temperature, respectively. Maximum NEP (NEP_max) values on days of year 221, 233, 247, and 262, were 2.44, 2.55, 3.90, and 4.17 μmol m⁻² s⁻¹, respectively. Throughout the growing period, the apparent quantum yield (α) increased with increasing NEP_max that ranged from 0.0154 to 0.0515, and NEP responded to the soil temperature changes by 44% and R _eco changes by 48%, and R _eco responded from 88 to 94% with the soil temperature diurnally. NEP’s light response and R _eco’s temperature response were affected by soil water content; more than 27% of the variation in NEP and 67% of the variation in R _eco could be explained by this parameter. NEP was strongly correlated with biomass and LAI, but R _eco was not, because environmental variables affected R _eco more strongly than growth parameters. Using the light response of NEP, the temperature response of R _eco, and meteorological data, daily NEP and R _eco were estimated at 0.67, 0.81, 1.17, and 1.56 g C m⁻², and at 2.88, 2.50, 3.51, and 3.04 g C m⁻², respectively, on days of year 221, 233, 247, and 262. The corresponding daily gross primary production (NEP + R _eco) was 3.5, 3.3, 4.6, and 4.6 g C m⁻².     

Changes of soil organic and inorganic carbon in relation to grassland degradation in Northern Tibet

The effect of livestock grazing on grassland degradation and the resulting impact on soil carbon concentration is an important factor in carbon estimation. We addressed this issue using field observations and laboratory analysis of samples from Tibetan grassland. Based on the field measurements, we investigated the soil organic carbon (SOC) and soil inorganic carbon (SIC) under two contrasting degradation states: lightly or non-degraded grasslands (LDG) and heavily degraded grasslands (HDG). We assessed their relationships with environmental factors using data collected from 99 sites across Northern Tibet during 2011–2012. Data were analyzed using a linear mixed-effects model and one-way ANOVA. The results showed that: (1) SOC concentration decreased and SIC concentration increased following grassland degradation, especially at soil depths in the range of 0–10 cm (P < 0.05); (2) the major environmental factors affecting SOC and SIC were soil pH and plant biomass; (3) spatially, the SOC density increased with the mean annual temperature and mean annual precipitation, whereas SIC exhibited the opposite trend; (4) the SOC density increased at first and then decreased with increasing grazing intensity, with an opposite trend in SIC; and (5) soil carbon storage in this region was 0.14 Pg smaller in the HDG than in the LDG. This study suggests that grassland degradation can significantly affect the vertical distribution and storage of SOC and SIC. The carbon sequestration capacity of the top 100 cm of soil in Northern Tibet was estimated as 0.14 Pg.     

模拟增温对羊草生态系统土壤呼吸速率的影响

草地是陆地生态系统的重要组成部分,研究草地系统土壤呼吸速率对全球气候变暖的响应,对预测全球碳循环具有重要作用。采用高度分别为0.5 m（T1）和1.85 m（T2）的开顶式增温箱（OTCs）对羊草生态系统进行模拟增温,仔细观察羊草的生育期,在每个生育期的同一天的晨间t₁时段（9：00-11：00）、午间t₂时段（13：30-15：30）和晚间t₃时段（17：00-19：00）监测土壤呼吸速率。分析不同增温幅度下土壤呼吸速率的变化规律,明确影响土壤碳排放的主要因素,探讨土壤呼吸速率与影响因素间存在的关系。结果显示：1）相对于空白对照C,模拟增温T1和T2导致0-10 cm土壤温度分别显著提高1.18和2.37℃;导致0-10 cm土壤湿度降低2.27%和4.57%;2）在羊草生长阶段,土壤呼吸速率呈现明显的季节性变化特征,同一天的t₁时段、t₂时段和t₃时段土壤呼吸速率峰值分别出现在结实期、抽穗期和开花期。非生长阶段土壤呼吸速率无显著差异;3）不同处理下土壤呼吸速率与近地表气温、0-10 cm土壤温度和地下生物量呈指数正相关关系,与0-10 cm的土壤湿度呈显著二次项负相关关系,与地上生物量表现为二次项正相关关系。研究结果明确了羊草生态系统中土壤碳排放对增温的响应,可为草原生态系统应对气候变化及可持续发展提供理论依据。     

Seasonal and interannual variations of ecosystem photosynthetic characteristics in a semi-arid grassland of Northern China

北方半干旱草原生态系统光合参数的季节和年际变异 生态系统表观量子效率(α)、最大光合速率(P_max)和暗呼吸速率(R_d)不仅反映了生态系统水平 光合生理特征,同时也是碳循环模型中光合过程模拟的关键参数。气候和植被因子都会影 响光合参数的季节和年际变异,但二者在光合参数调控过程中的相对贡献和作用途径尚不清晰。本研究基于连续12年(2006–2017)的涡度相关观测数据,分析了内蒙古半干旱典型草原光合参数的季节和年际变化规律;利用回归分析和结构方程模型(SEM)方法明晰了环境和生理调控的作用途径及相对贡献。结果发现,光合参数(α、P_max和R_d)均表现出单峰的季节变化趋势,并呈现明显的年际波动。温度(T_a)和土壤含水量(SWC)的变化共同影响光合参数的季节变化,而SWC主导了其年际变异。α和R_d的变化主要由T_a决定,而Pmax的变化主要受SWC的影响。SEM模型分析表明,除了直接作用外,环境因子主要通过影响冠层水平气孔导度(gc)对光合参数和碳同化生理过程进行调控。此外,叶面积指数对光合参数特别是Pmax的季节和年际变异起主要调控作用。以上结果明确了环境和植被共同决定了生态系统水平光合参数的季节和年际变异,并强调了在水分受限的草原生态系统中,植被生理调控在光合碳同化能力和碳汇功能评估中的重要作用。     

Biomass Partitioning and Its Relationship with the Environmental Factors at the Alpine Steppe in Northern Tibet

Alpine steppe is considered to be the largest grassland type on the Tibetan Plateau. This grassland contributes to the global carbon cycle and is sensitive to climate changes. The allocation of biomass in an ecosystem affects plant growth and the overall functioning of the ecosystem. However, the mechanism by which plant biomass is allocated on the alpine steppe remains unclear. In this study, biomass allocation and its relationship to environmental factors on the alpine grassland were studied by a meta-analysis of 32 field sites across the alpine steppe of the northern Tibetan Plateau. We found that there is less above-ground biomass (M_A) and below-ground biomass (M_B) in the alpine steppe than there is in alpine meadows and temperate grasslands. By contrast, the root-to-shoot ratio (R:S) in the alpine steppe is higher than it is in alpine meadows and temperate grasslands. Although temperature maintained the biomass in the alpine steppe, precipitation was found to considerably influence M_A, M_B, and R:S, as shown by ordination space partitioning. After standardized major axis (SMA) analysis, we found that allocation of biomass on the alpine steppe is supported by the allometric biomass partitioning hypothesis rather than the isometric allocation hypothesis. Based on these results, we believe that M_A and M_B will decrease as a result of the increased aridity expected to occur in the future, which will reduce the landscape’s capacity for carbon storage.     

藏北高寒草地植被和土壤对不同放牧强度的响应

放牧压力在时间和空间的不均衡分布导致局部土地过度利用和生态退化,然而当前对放牧强度的量化多采用替代性指标或对照试验,缺乏直接监测数据,也缺乏不同草地类型对放牧强度变化的响应差异性研究。以西藏自治区那曲市为研究区,利用佩戴式GPS牛羊定位器构建高精度放牧轨迹数据集模拟放牧强度,构建栅格尺度放牧强度空间分布和划分方法,结合草地群落样方调查,通过Duncan法（Duncan''s multiple comparative analysis）进行多重比较分析,探究自由放牧模式下高寒草甸和高寒草原两类区域植被和土壤对不同放牧强度的响应方式及差异性。本研究可为放牧行为环境效应监测提供新思路,并根据不同草地状况因地制宜提出放牧优化管理策略,助力高寒传统牧区的可持续发展。主要结论有：1）随着放牧强度的增大,高寒草原地上生物量先升高后降低,高强度放牧对高寒草原植被的影响大于高寒草甸。2）高放牧强度下,高寒草原土壤水分显著高于中低强度,高寒草甸土壤容重显著低于低强度。可能原因是放牧压力多集聚于水源附近。高寒草原区土壤的砂粒含量随放牧强度的增大而增加。放牧强度的增大导致草甸上层土壤有机质增加,草甸下层、草原上层、草原下层土壤有机质先增加后减小。草甸上下层土壤全磷含量在低放牧强度下显著低于中高强度。3）植被土壤变化受到自然因子和放牧活动的共同影响。高寒草甸更加耐牧,高强度放牧对高寒草原的负面影响更大,而中度放牧有利于草地尤其是高寒草原的放牧利用。4）放牧生态系统是一个环境-植物-家畜自适应系统。在放牧管理中不能仅通过控制载畜量缓解草地超载,还需要综合考虑生态系统的弹性及各营养级的适应性,合理配置放牧强度,控制季节性超载和局部超载。     

氮添加对多伦草原土壤微生物呼吸及其温度敏感性的影响

人类活动导致氮和磷输入到草原生态系统,对土壤有机碳循环产生影响,但是土壤微生物呼吸(Soil microbial respiration,Rs)及其温度敏感性(Q₁₀)对于氮沉降和磷有效性增加的响应还存在争议。因此,依托多伦草原氮添加样地(0、50 kg N hm^-2 a^-1和100 kg N hm^-2 a^-1),并添加磷进行室内恒温培养(10℃和15℃),研究氮添加和磷有效性增加对Rs及其Q₁₀的影响。结果发现:氮添加显著降低胞壁酸含量和显著增加真菌丰富度(Fungal richness, F-richness)。与N0处理相比,N50和N100处理使累积呼吸量显著降低了61.2%和67.1%,但Q₁₀显著升高了32.7%和50.8%;磷有效性增加没有对累积呼吸量及其Q₁₀产生显著影响。逐步回归结果表明,F-richness和pH值分别是累积呼吸量及其Q₁₀最重要的影响因子。研究表明氮添加...     

Temperature and substrate controls on intra-annual variation in ecosystem respiration in two subarctic vegetation types

Arctic ecosystems are important in the context of climate change because they are expected to undergo the most rapid temperature increases, and could provide a globally significant release of CO₂ to the atmosphere from their extensive bulk soil organic carbon reserves. Understanding the relative contributions of bulk soil organic matter and plant‐associated carbon pools to ecosystem respiration is critical to predicting the response of arctic ecosystem net carbon balance to climate change. In this study, we determined the variation in ecosystem respiration rates from birch forest understory and heath tundra vegetation types in northern Sweden through a full annual cycle. We used a plant biomass removal treatment to differentiate bulk soil organic matter respiration from total ecosystem respiration in each vegetation type. Plant‐associated and bulk soil organic matter carbon pools each contributed significantly to ecosystem respiration during most phases of winter and summer in the two vegetation types. Ecosystem respiration rates through the year did not differ significantly between vegetation types despite substantial differences in biomass pools, soil depth and temperature regime. Most (76–92%) of the intra‐annual variation in ecosystem respiration rates from these two common mesic subarctic ecosystems was explained using a first‐order exponential equation relating respiration to substrate chemical quality and soil temperature. Removal of plants and their current year's litter significantly reduced the sensitivity of ecosystem respiration to intra‐annual variations in soil temperature for both vegetation types, indicating that respiration derived from recent plant carbon fixation was more temperature sensitive than respiration from bulk soil organic matter carbon stores. Accurate assessment of the potential for positive feedbacks from high‐latitude ecosystems to CO₂‐induced climate change will require the development of ecosystem‐level physiological models of net carbon exchange that differentiate the responses of major C pools, that account for effects of vegetation type, and that integrate over summer and winter seasons.