Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.     

Morphological and moisture availability controls of the leaf area‐to‐sapwood area ratio: analysis of measurements on Australian trees

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Evaluation of methods for estimating soil carbon dioxide efflux across a gradient of forest disturbance

Better understanding of variation in soil carbon dioxide (CO₂) efflux caused by measurement techniques is needed, especially over gradients of site disturbance, to accurately estimate the global carbon cycle. We present soil CO₂ efflux data from a gradient of disturbance to ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests in northern Arizona, USA that were obtained using four different techniques: vented static chambers, a Licor 6400‐09, and soil CO₂ diffusion profiles using two different models (Moldrup, Millington–Quirk) to estimate soil gas diffusivity. We also compared soil CO₂ efflux measured by the Moldrup and Millington–Quirk diffusion profile methods to nighttime total ecosystem respiration (TER) data from an eddy covariance tower. We addressed four questions: (1) Does the use of a given method to measure soil CO₂ efflux bias results across a disturbance gradient? (2) Does the magnitude of difference between observed and modeled estimates of soil CO₂ differ between methods and across sites? (3) What is the spatial variability of each method at each site? (4) Which method is closest to the estimate of TER measured by the eddy covariance tower? Although soil CO₂ efflux varied significantly among methods the differences were consistent among sites. Measured and modeled total growing season fluxes were generally higher for the Licor 6400‐09 and Millington–Quirk diffusion gradient methods compared with static chamber and the Moldrup diffusion gradient methods. A power analysis showed that the larger static chamber was the most efficient method at sampling spatial variation in soil CO₂ efflux. Nighttime measurements of soil CO₂ efflux from the Moldrup diffusion gradient method were most strongly related to nighttime TER assessed with eddy covariance. The use of a single, well‐implemented method to measure soil CO₂ efflux is unlikely to create bias in comparisons across a gradient of forest disturbance.     

Evidence of increased net ecosystem productivity associated with a longer vegetated season in a deciduous forest in south‐central Indiana,USA

Observations of net ecosystem exchange (NEE) of carbon and its biophysical drivers have been collected at the AmeriFlux site in the Morgan‐Monroe State Forest (MMSF) in Indiana, USA since 1998. Thus, this is one of the few deciduous forest sites in the world, where a decadal analysis on net ecosystem productivity (NEP) trends is possible. Despite the large interannual variability in NEP, the observations show a significant increase in forest productivity over the past 10 years (by an annual increment of about 10 g C m^?2 yr^?1). There is evidence that this trend can be explained by longer vegetative seasons, caused by extension of the vegetative activity in the fall. Both phenological and flux observations indicate that the vegetative season extended later in the fall with an increase in length of about 3 days yr^?1 for the past 10 years. However, these changes are responsible for only 50% of the total annual gain in forest productivity in the past decade. A negative trend in air and soil temperature during the winter months may explain an equivalent increase in NEP through a decrease in ecosystem respiration.     

全球旱地饱和水汽压差和根区土壤水分变化对植被生产力的影响及其成因

旱地约占全球陆地面积的40%,而水分是旱地植被生长的一大限制要素。尽管土壤水分与饱和水汽压差对植被生长的重要性已经得到了广泛证实,然而目前二者对植被生产力影响的空间异质性及其形成因素仍未得到深入研究,这对研究旱地生态系统对气候变化的响应带来了挑战。为了填补这一认知空白,研究收集了多源气象、根区土壤含水率和总初级生产力产品,基于随机森林算法量化了植被总初级生产力对根区土壤含水率和饱和水汽压差的敏感性,结合土地覆盖数据和分档平均方法分析了敏感性空间异质性的形成机制。结果表明:全球旱地饱和水汽压差与植被生产力总体呈显著上升趋势;根区土壤水分对植被生长的影响以正效应主导,饱和水汽压差对植被生长的影响以负效应主导;相较于森林和灌木,饱和水汽压差对植被生长的负效应及根区土壤含水率对植被生长的正效应在农田、草地和苔原及半干旱区更为强烈;植被生产力对饱和水汽压差和根区土壤水分的敏感性在数量上总体呈显著的线性负相关性。综上,植被种类和气候条件是导致全球旱地植被生产力对土壤水分和饱和水汽压差敏感性空间异质性的重要因素。     

Interacting controls on productivity in a northern Great Plains grassland and implications for response to ENSO events

In this study we investigated the causes of annual variability in peak aboveground biomass production, net ecosystem productivity (NEP) and gross ecosystem productivity (GEP) during an 8‐year period (1999–2006) in a northern Great Plains grassland near Lethbridge, Alberta, Canada. In particular, we tested for a significant relationship between growing season precipitation and productivity and determined whether soil moisture carry‐over from the previous fall–winter could alter this relationship. We also investigated the interaction between soil moisture availability and temperature in controlling grassland productivity. There was a very strong correlation between total precipitation input and average soil moisture content during the May–October growing season. However, the growing season average soil moisture contents in 2003 and 2006 were very similar to those recorded in 1999, despite lower than normal precipitation occurring in these 2 years. This resulted from a positive difference between precipitation and evapo‐transpiration that allowed significant soil moisture to be carried‐over from the previous fall–winter during both 2003 and 2006. Strong logistic relationships were observed between soil moisture and annual productivity based on data from all years except 2003 and 2006, years which had higher productivity than was predicted from the logistic regression. Interaction between temperature and soil moisture explained this difference. Productivity values in 2003 and 2006 were high compared with 1999, a year with approximately the same soil moisture content, and this resulted from the higher average growing season temperatures that were apparent in 2003 and 2006. Analysis of weather records indicated that precipitation in the month of June was significantly higher during El Niño years than during La Niña years in Lethbridge. During the study period, aboveground biomass, NEP and GEP were generally higher in El Niño years and lower in La Niña years because of associated variation in summer precipitation.     

涡度相关法研究土壤水分状况对沙地杨树人工林生态系统能量分配和蒸散日变化的影响

为了解位于北京大兴区林场杨树人工林在不同的土壤水分环境条件下的水汽交换过程和能量的分配差异及其与环境因子关系,运用涡度相关（Eddy covariance,EC）法开路系统、常规微气象观测系统及土壤热通量板等设施对生态系统生长季内典型水分胁迫和无水分胁迫条件下蒸散日变化、能量分配以及与各环境因子的关系进行了测定分析和比较。结果表明,在水分严重胁迫日（以7月7日为例）,蒸散日变化过程为单峰曲线,全天（24h）蒸散量为2.4mm;而在无水分胁迫典型日（以7月25日为例）,蒸散日变化过程呈多峰曲线,全天蒸散量为4．5mm。能量平衡分析显示,无水分胁迫条件下潜热通量（LE）占净辐射通量（Rn）的比例远高于水分胁迫条件下潜热通量占净辐射通量的比例,说明水分充足时,能量的大部分用于蒸散。水分胁迫条件下蒸散速率与各环境因子的相关性均低于无水分胁迫条件下蒸散速率与环境因子的相关性。水分胁迫条件下,蒸散速率主要与净辐射和下垫面因子关系显著,而与其它因子的相关性较小;无水分胁迫条件下,蒸散速率与下垫面土体含水量和各气象因子均表现出较强的相关性。大气温度对于两个典型日蒸散速率的影响均很小;土壤含水量与水分胁迫日的蒸散速率几乎没有相关性,反应出土壤水分含量低至对蒸散几乎没有贡献了。     

Semiempirical modeling of abiotic and biotic factors controlling ecosystem respiration across eddy covariance sites

In this study we examined ecosystem respiration (R_ECO) data from 104 sites belonging to FLUXNET, the global network of eddy covariance flux measurements. The goal was to identify the main factors involved in the variability of R_ECO: temporally and between sites as affected by climate, vegetation structure and plant functional type (PFT) (evergreen needleleaf, grasslands, etc.). We demonstrated that a model using only climate drivers as predictors of R_ECO failed to describe part of the temporal variability in the data and that the dependency on gross primary production (GPP) needed to be included as an additional driver of R_ECO. The maximum seasonal leaf area index (LAI_MAX) had an additional effect that explained the spatial variability of reference respiration (the respiration at reference temperature T_ref=15 °C, without stimulation introduced by photosynthetic activity and without water limitations), with a statistically significant linear relationship (r²=0.52, P<0.001, n=104) even within each PFT. Besides LAI_MAX, we found that reference respiration may be explained partially by total soil carbon content (SoilC). For undisturbed temperate and boreal forests a negative control of total nitrogen deposition (N_depo) on reference respiration was also identified. We developed a new semiempirical model incorporating abiotic factors (climate), recent productivity (daily GPP), general site productivity and canopy structure (LAI_MAX) which performed well in predicting the spatio‐temporal variability of R_ECO, explaining >70% of the variance for most vegetation types. Exceptions include tropical and Mediterranean broadleaf forests and deciduous broadleaf forests. Part of the variability in respiration that could not be described by our model may be attributed to a series of factors, including phenology in deciduous broadleaf forests and management practices in grasslands and croplands.     

Canopy and leaf gas exchange of <Emphasis Type="Italic">Haloxylon ammodendron</Emphasis> under different soil moisture regimes

In order to reveal the drought resistance and adaptation of the C₄ desert plant Haloxylon ammodendron under artificially controlled soil moisture regimes, representative plants were selected to measure canopy photosynthesis using canopy photosynthetic measurement system. The results showed that appropriate soil moisture significantly enhances the canopy and leaf photosynthetic capacity, and extremely high soil moisture is not conducive to the photosynthesis of H. ammodendron.     

Differential responses of production and respiration to temperature and moisture drive the carbon balance across a climatic gradient in New Mexico

Southwestern North America faces an imminent transition to a warmer, more arid climate, and it is critical to understand how these changes will affect the carbon balance of southwest ecosystems. In order to test our hypothesis that differential responses of production and respiration to temperature and moisture shape the carbon balance across a range of spatio‐temporal scales, we quantified net ecosystem exchange (NEE) of CO₂ and carbon storage across the New Mexico Elevational Gradient, which consists of six eddy‐covariance sites representing biomes ranging from desert to subalpine conifer forest. Within sites, hotter and drier conditions were associated with an increasing advantage of respiration relative to production such that daily carbon uptake peaked at intermediate temperatures – with carbon release often occurring on the hottest days – and increased with soil moisture. Across sites, biotic adaptations modified but did not override the dominant effects of climate. Carbon uptake increased with decreasing temperature and increasing precipitation across the elevational gradient; NEE ranged from a source of ～30 g C m^?2 yr^?1 in the desert grassland to a sink of ～350 g C m^?2 yr^?1 in the subalpine conifer forest. Total aboveground carbon storage increased dramatically with elevation, ranging from 186 g C m^?2 in the desert grassland to 26 600 g C m^?2 in the subalpine conifer forest. These results make sense in the context of global patterns in NEE and biomass storage, and support that increasing temperature and decreasing moisture shift the carbon balance of ecosystems in favor of respiration, such that the potential for ecosystems to sequester and store carbon is reduced under hot and/or dry conditions. This implies that projected climate change will trigger a substantial net release of carbon in these New Mexico ecosystems (～3 Gt CO₂ statewide by the end of the century), thereby acting as a positive feedback to climate change.     

基于土壤水分模拟的干旱动态监测指标及其适用性

根据土壤水分平衡原理,采用Penman-Monteith公式计算可能蒸散,逐日滚动模拟土壤有效水分,提出基于土壤水分(SM)的逐日干旱动态监测指标。通过与《气象干旱等级》国家标准中标准化降水指数(SPI)、相对湿润度指数(MI)和综合气象干旱指数(CI)相比较,分析了SM干旱动态监测指标在广东的适用性特征。结果表明:(1)基于SM统计的干旱频发时段(11月—翌年4月)比少雨时段(10月—翌年3月)滞后1个月,SM指标能够刻画干旱相对于降水年变化的滞后效应和逐渐发展的特征,客观反映广东干旱频率季节变化特征;(2)SM干旱指标自动隐含了降水-蒸发过程的时间累积效应,物理意义清晰,能够刻画干旱发生、发展和结束过程的丰富细节,在对干旱过程发展渐进性、降水敏感性和过程完整性方面,具有较好的应用价值。     

土壤水分对冬小麦生育后期叶片气体交换及叶绿素荧光参数的影响

通过防雨棚小区栽培,控制土壤供水系数(Kw)分别为0.8、0.6、0.4、0.2,以自然状况下的小区为对照(CK),研究土壤水分条件对冬小麦生育后期叶片气体交换及叶绿素荧光参数的影响。结果表明:Kw为0.6处理的冬小麦叶片叶绿素含量与0.8处理接近,且显著高于其他处理(P<0.05);Kw为0.6处理对冬小麦叶片的气孔导度和蒸腾速率有轻度抑制,但其光合速率却高于0.8处理,而Kw为0.2处理的光合速率、气孔导度及蒸腾速率均为最低;气孔限制值在Kw为0.4处理下最高,其次为0.2处理,0.8处理下最低;冬小麦叶片的表观量子效率在Kw为0.4处理下最高,光补偿点总体上随着土壤水分含量的降低呈下降趋势,而光饱和点及最大光合速率则以Kw为0.6处理最高,其次为0.8处理,0.2处理最低;冬小麦叶片的天线转化效率Fv’/Fm’、电子传递速率ETR、实际量子效率ФPSII及光化学猝灭qP均以Kw为0.6处理最高,其次为0.8处理,0.2处理下最低;在Kw为0.2处理下,冬小麦光合作用主要受非气孔因素限制,而在0.4处理下,则主要受气孔因素限制。     

Observed and modelled historical trends in the water‐use efficiency of plants and ecosystems

Plant water‐use efficiency (WUE, the carbon gained through photosynthesis per unit of water lost through transpiration) is a tracer of the plant physiological controls on the exchange of water and carbon dioxide between terrestrial ecosystems and the atmosphere. At the leaf level, rising CO₂ concentrations tend to increase carbon uptake (in the absence of other limitations) and to reduce stomatal conductance, both effects leading to an increase in leaf WUE. At the ecosystem level, indirect effects (e.g. increased leaf area index, soil water savings) may amplify or dampen the direct effect of CO₂. Thus, the extent to which changes in leaf WUE translate to changes at the ecosystem scale remains unclear. The differences in the magnitude of increase in leaf versus ecosystem WUE as reported by several studies are much larger than would be expected with current understanding of tree physiology and scaling, indicating unresolved issues. Moreover, current vegetation models produce inconsistent and often unrealistic magnitudes and patterns of variability in leaf and ecosystem WUE, calling for a better assessment of the underlying approaches. Here, we review the causes of variations in observed and modelled historical trends in WUE over the continuum of scales from leaf to ecosystem, including methodological issues, with the aim of elucidating the reasons for discrepancies observed within and across spatial scales. We emphasize that even though physiological responses to changing environmental drivers should be interpreted differently depending on the observational scale, there are large uncertainties in each data set which are often underestimated. Assumptions made by the vegetation models about the main processes influencing WUE strongly impact the modelled historical trends. We provide recommendations for improving long‐term observation‐based estimates of WUE that will better inform the representation of WUE in vegetation models.     

Satellite chlorophyll fluorescence measurements reveal large‐scale decoupling of photosynthesis and greenness dynamics in boreal evergreen forests

Mid‐to‐high latitude forests play an important role in the terrestrial carbon cycle, but the representation of photosynthesis in boreal forests by current modelling and observational methods is still challenging. In particular, the applicability of existing satellite‐based proxies of greenness to indicate photosynthetic activity is hindered by small annual changes in green biomass of the often evergreen tree population and by the confounding effects of background materials such as snow. As an alternative, satellite measurements of sun‐induced chlorophyll fluorescence (SIF) can be used as a direct proxy of photosynthetic activity. In this study, the start and end of the photosynthetically active season of the main boreal forests are analysed using spaceborne SIF measurements retrieved from the GOME‐2 instrument and compared to that of green biomass, proxied by vegetation indices including the Enhanced Vegetation Index (EVI) derived from MODIS data. We find that photosynthesis and greenness show a similar seasonality in deciduous forests. In high‐latitude evergreen needleleaf forests, however, the length of the photosynthetically active period indicated by SIF is up to 6 weeks longer than the green biomass changing period proxied by EVI, with SIF showing a start‐of‐season of approximately 1 month earlier than EVI. On average, the photosynthetic spring recovery as signalled by SIF occurs as soon as air temperatures exceed the freezing point (2–3 °C) and when the snow on the ground has not yet completely melted. These findings are supported by model data of gross primary production and a number of other studies which evaluated in situ observations of CO₂ fluxes, meteorology and the physiological state of the needles. Our results demonstrate the sensitivity of space‐based SIF measurements to light‐use efficiency of boreal forests and their potential for an unbiased detection of photosynthetic activity even under the challenging conditions interposed by evergreen boreal ecosystems.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.