基于通量和光谱观测的中亚热带人工针叶林光能利用效率的反演

利用光谱反射率测量的光化学植被指数（PRI）估算植被光合作用的光能利用效率（LUE）,能够更好地为生态系统总初级生产力的估算及尺度扩展提供重要的技术支撑.本研究以中国通量网（ChinaFLUX）千烟洲通量观测站为研究区域,2013年9月和12月在通量塔上测量了中亚热带人工针叶林的植被反射光谱,并获取了通量塔上同步观测的气象数据和涡度相关通量数据,对两者进行回归分析.结果表明： PRI-LUE相关关系（R²=0.20,P<0.001）优于NDVI LUE.在整个观测期内,土壤水分含量（SWC）与PRI组合的二元回归模型能够提高LUE的估算精度（日间观测R²=0.29,P<0.001;正午观测R²=0.30,P<0.01）,而在秋季,饱和水汽压差（VPD）与PRI组合的二元回归模型能较好地估算正午LUE（R²=0.448, P<0.001）,表明环境因子SWC和VPD是影响PRI-LUE关系的重要因素,不同季节的二元回归模型所选择的最佳环境变量有所不同.     

C3和C4植物光合途径的适应性变化和进化

 高等植物大多为C₃植物, C₄植物和景天酸代谢(Crassulacean acid metabolism, CAM)植物是由C₃植物进化而来的。C₄途径的多源进化表明, 光合途径由C₃途径向C₄途径的转变相对简单。该文分析研究了植物光合途径的进化前景, 指出C₄植物是从C₃植物进化而来的高光效种类, 且地质时期以来降低的大气CO₂浓度和升高的大气温度以及干旱和盐渍化是C₄途径进化的外部动力。C₃植物的C₄途径的发现说明植物的光合途径并非是一成不变的, C₃和C₄植物的光合特征具有极大的可塑性, 某些环境的变化会引起植物光合途径在C₃和C₄途径之间转变。C₃植物具有的C₄途径是环境调控的产物, 是对逆境的适应性进化结果, 因而光合途径的转变也适用于干旱地区植被的适应性生存机理研究。该文还利用国外最新的C₄光合进化模型介绍了植物在进化C₄途径中所经历的7个重要时期(从分子基础到形态基础、结构基础, 再到物质代谢水平、光合酶活水平, 直到C3和C4途径协调运转时期, 最后达到形态与功能最优化阶段), 并结合全球气候变化的特点对国内外相关领域的研究进行了分析, 总结了植物光合途径的适应性转变和进化的研究成果, 为今后的相关工作提出建议。     

植被光能利用率:模型及其不确定性

光能利用率（Light use efficiency： LUE）指植物截获的光能转化为化学能的效率,表示为生产力和吸收光能之比。基于LUE概念的模型对模拟预测全球变化下碳循环、植被生产力及其潜力具有重要意义。全球变化和人类活动影响给植被生产力和碳循环的评估带来了巨大挑战。系统梳理了LUE模型的不确定性并分析其原因,以期提高生产力模拟预测的准确度。分析发现LUE模型准确度仅为62%-70%且模型间差异较大（32%）,误差随着植被类型、时间尺度和空间区域的不同存在显著差别。目前计算LUE的误差是模型不确定性的关键,原因主要在于LUE与影响因素尤其是水分的关系并不清楚。一方面不能准确区分水分胁迫指标对LUE的影响机制,另一方面无法准确模拟水分等影响因素与LUE关系的时空演变特征。未来该领域研究的重要方向是发展集成样地和区域尺度的叶绿素荧光、光化学指数等研究方法,厘定LUE与影响因素特别是的水分关系,并分析其时空演变特征。     

CO2浓度升高对茴香植株生长、精油含量和组分的影响

 采用每日定时向密封人工气候室补充CO₂的方法,研究了3种CO₂浓度（平均浓度分 别为287.11、532.88和780.46 μmol·mol^-1）对茴香 (Foeniculum vulgare)生长、精油含量和组分的影响。结果表明：随着CO₂浓度的升高,茴香的株高、花序数、花序鲜重、花序干重、全株干重 和植株的干物率均有所上升;植株可溶性糖和全碳含量不断升高,而全氮和蛋白氮含量不断减少;叶色素含量呈下降趋势,叶绿素a/b比的差异 不显著;植株精油含量（分别为1.26、1.45和1.57 ml·(100 g) ^-1 DW）和单株精油产量（分别为0.019、 0.023和0.033 ml）均随之升高。从茴 香植株的精油中鉴定出22种成分,用不同浓度的CO₂处理,精油的成分种类没有差异,成分相对含量却有差别,差异达到极显著水平的有：醎蒎 烯、鈅蒎烯、月桂烯、对聚伞花素、反式葑醇乙酸酯和顺式茴香脑;含量差异达到显著水平的有：香桧烯、水芹烯、罗勒烯、鉥萜品烯、3,4- 二甲基-2,4,6_三烯、爱草脑、葑醇乙酸酯、古巴烯、 金合欢烯和吉玛烯。茴香精油的主要成分反式茴香脑的含量（分别为55.94%、57.20%和 59.5 5%）随着CO₂浓度的升高而升高,而柠檬烯含量（29.60%、30.24%和26.12%）表现出相反的趋势,二者在不同的CO₂浓度处理之间差异均不 显著。     

基于3S的自然植被光能利用率的时空分布特征的模拟

 光能利用率（LUE）直接影响植被各层中的能量分布和光合速率,在确定环境对光合和地上部生长分配的综合限制上十分有价值,是衡量系统功能的一个重要指标。本研究以遥感图像（TM）作为数据源,获取了影响植被LUE的重要变量——叶面积指数（LAI）;用程序语言编写了描述系统碳循环和水循环的景观尺度生态系统生产力过程模型（EPPML）,对长白山自然保护区的太阳总辐射、净初级生产力（NPP）和LUE等的季节动态和空间分布进行了模拟;并用地理信息系统（GIS）手段对空间数据进行处理、分析和显示,从而实现了将植物生理生态研究的结果从小尺度向中尺度进行拓展和转换。EPPML可以比较准确地模拟长白山自然保护区景观尺度上主要植被类型的NPP和太阳总辐射,对LUE的模拟结果也大多在我国森林的LUE范围之内,但对不同植被类型LUE的验证因实测数据不足,仅做了初步比较。模拟结果表明,长白山植被的LUE与NPP的季节进程十分近似,7月可达2.9%。春、夏、秋、冬四个季节植被LUE的模拟平均值分别为0.551%、2.680%、0.551%和0.047%。植被年LUE的模拟值平均为1.075%,在-3.272%～3.556%之间变化,阔叶红松(Pinus koraiensis)林最大（1.653%）,高山流砾滩草类最小（0.146%）。阔叶红松林的LUE虽然较高,但仍有很大的增长潜力。     

CO2浓度升高对森林土壤微生物呼吸与根（际）呼吸的影响

 根呼吸与微生物呼吸的作用底物不同,二者对高浓度CO₂的响应机理及敏感程度亦不同。在大气CO₂浓度升高的背景下,精确区分根呼吸与微生物呼吸是构建森林生态系统碳循环模型和预测森林生态系统碳源/汇关系所必需的。根（际）呼吸与微生物呼吸对高浓度CO₂的响应呈增加、降低或无明显变化等不同趋势,根（际）呼吸变化主要与根生物量明显相关,细根的作用大于粗根;土壤微生物呼吸变化存在较大的不确定性,微生物量和微生物活性与土壤微生物呼吸相关或不相关。根系统对高浓度CO₂的响应会潜在地影响微生物的代谢底物,进而影响微生物呼吸强度。凡影响土壤总呼吸的生物与非生物因子都会直接或间接地影响根呼吸与土壤微生物呼吸。     

热带季节雨林林冠上方和林内近地层CO2浓度的时空动态及其成因分析

 为探讨西双版纳独特地方气候背景下,热带季节雨林CO₂浓度的时空变化特征和不同时间尺度上环境因素对森林CO₂浓度时间分布的作用,以及 为研究热带季节雨林的碳通量、净生态系统交换量(Net ecosystem exchange, NEE)等提供支持,我们利用热带季节雨林林冠上方和林内近地层 CO₂浓度连续监测资料,结合同步气象资料进行了统计分析。研究结果表明：在植被生理活动、土壤呼吸以及林内湍流的共同作用下,西双版纳 热带季节雨林CO₂浓度表现出明显的日变化、季节变化和林冠上下差异。在日尺度上,林冠上方的CO₂浓度时间变化曲线为“单峰型”,林内近 地层CO₂浓度时间变化曲线为“双峰型”,造成林内近地层傍晚第二个峰值的主要因子是地形因子作用下形成的局地环流。在季节尺度上,林冠 上方CO₂浓度主要受林冠代谢作用的影响,呈现雨季低、干季高的特点,而林内近地层的CO₂浓度则主要受地表呼吸过程所控制,季节变化趋势 与林冠上方相反。林冠上方CO₂浓度低于林内近地层CO₂浓度,且差异较大;在日尺度上,各月（除12月外）CO₂浓度的最大差值皆大于80 mg·m ^－3,且出现在傍晚;在季节尺度上,最大值为-62.9 mg·m^－3,出现在10月,最小值为-8.4 mg·m^－3,出现在12月。     

短期CO2加富对凤梨光合作用及生长发育的影响

 研究了CO₂加富对丹尼斯凤梨（Guzmania`Denise’）和吉利凤梨（Guzmania `Cherry’）叶片光合速率、植株生长、开花和光合相关酶活性的 影响。结果表明,处理30 d期间,处理(600±40)、(900±40) μmol CO₂&;#8226;mol^-1的净光合速率分别比同期对照增加了6.24%～31.91%和11.92%～ 41.48%;CO₂加富下促进了叶片中可溶性糖和淀粉的积累, 蒸腾速率和气孔导度下降,Rubisco活性增加,乙醇酸氧化酶活性则明显下降。(600 ±40)μmol CO₂&;#8226;mol^-1处理下的株高、叶面积分别比同期对照下增加了6.94%～14.63%和1.66%～7. 06%,而处理(900±40) μmol CO₂&;#8226;mol^-1下 分别增加了9.71%～20.85%和2.87%～11.62%;CO₂加富下促进了干重和鲜重的积累。此外,CO₂加富提前了吉利凤梨的花期。     

青海省三江源区人工草地生态系统CO2通量

 了解三江源人工草地净生态系统CO₂交换(Net ecosystem CO₂ exchange, NEE)的季节变化规律和主要生物因子及环境因子对这些过程的影响将有助于认识青藏高原人工草地生态系统碳循环、生态价值、功能,以及对三江源区的生态安全的重要意义。该研究利用涡度相关技术,于2005年9月1日至2006年8月31日对位于青海腹地的垂穗披碱草（Elymus nutans）人工草地的NEE及生物和环境因子进行观测, 阐明NEE及其组分的动态变化特征和影响因子。三江源区人工草地生态系统的日最大吸收量为2.38 g C·m^－2·d^－1,出 现在7月30日。日间最大吸收率和最大排放率都出现在8月,分别为-6.82和2.95μmol CO₂·m^－2·s^－1。在生长季, 白天的NEE主要受光合有效辐射(Photosynthe tically active rad iation, PAR)变化控制,同时又与叶面积指数和群落多样性交互作用,共同调节光合速率和光合效率的强度。最大光合同化速率为2.46～10.39μmol CO₂·m^－2·s^－1,表观初始光能利用率为0.013～0.070μmol CO₂·μmol^－1 PAR。 在碳交换日过程中,NEE并不完全随着 PAR的增加而增大,当PAR超过某一值（>1 200μmol ·m^－2·s^－1）时,NEE随PAR的增加而降低。受温度的影响,生长季的生态系统的呼吸商Q₁₀（1.8）小于非生长季节的 2.6）。 生态系统呼吸主要受温度的控制,同时也受到叶面积指数的显著影响。生长季昼夜温差大并不利于生态系统的碳获取。 三江源区人工草地生态系统是一个较强的碳汇,为-49.35 g C·m^－2·a^－1。     

线性和指数回归方法对土壤呼吸CO2扩散速率估算的影响

 理论上,土壤呼吸通量的量值可以通过观测土壤呼吸CO₂扩散速率(&#601;c/&#601;t)计算得到。但是为获得&#601;c/&#601;t,通常须允许土壤呼吸箱内CO₂浓度升高,因此,如何估算外界大气CO₂浓度条件下的&#601;c/&#601;t是土壤呼吸观测技术的关键,关系到观测结果的准确性。通常&#601;c/&#601;t的估算会受土壤表层大气CO₂扩散梯度(即土壤呼吸箱内CO₂扩散梯度和大气CO₂浓度昼夜变化)的影响。目前,线性回归方法是土壤呼吸观测中估算&#601;c/&#601;t的基本方法。然而,常用的线性回 归方法会低估&#601;c/&#601;t,而指数回归方法则可以准确地估算&#601;c/&#601;t。夜间&#601;c/&#601;t的变化与大气CO₂ 浓度之间存在非常明显的负相关关系。夜间土壤表层大气CO₂扩散梯度的减小导致线性回归方法明显低估&#601;c/&#601;t。&#601;c/&#601;t的昼夜变化过程存在明显的非对称性现象,而指数回归方法可以更好地描述&#601;c/&#601;t昼夜变化的非对称性响应。     

Deconvolution of pigment and physiologically related photochemical reflectance index variability at the canopy scale over an entire growing season

The sensitivity of the photochemical reflectance index (PRI) to leaf pigmentation and its impacts on its potential as a proxy for light‐use efficiency (LUE) have recently been shown to be problematic at the leaf scale. Most leaf‐to‐leaf and seasonal variability can be explained by such a confounding effect. This study relies on the analysis of PRI light curves that were generated at the canopy scale under natural conditions to derive a precise deconvolution of pigment‐related and physiologically related variability in the PRI. These sources of variability were explained by measured or estimated physiologically relevant variables, such as soil water content, that can be used as indicators of water availability and canopy chlorophyll content. The PRI mainly reflected the variability in the pigment content of the canopy. However, the corrected PRI, which was obtained by subtracting the pigment‐related seasonal variability from the PRI measurement, was highly correlated with the upscaled LUE measurements. Moreover, the sensitivity of the PRI to the leaf pigment content may mask the PRI versus LUE relationship or result in an artificial relationship that reflects the relationship of chlorophyll versus LUE, depending on the species phenology.     

Relationship between photochemical reflectance index and leaf ecophysiological and biochemical parameters under two different water statuses: towards a rapid and efficient correction method using real‐time measurements

The use of the photochemical reflectance index (PRI) as a promising proxy of light use efficiency (LUE) has been extensively studied, and some issues have been identified, notably the sensitivity of PRI to leaf pigment composition and the variability in PRI response to LUE because of stress. In this study, we introduce a method that enables us to track the short‐term PRI response to LUE changes because of photosynthetically active radiation (PAR) changes. The analysis of these short‐term relationships between PRI and LUE throughout the growing season in two species (Quercus robur L. and Fagus sylvatica L.) under two different soil water statuses showed a clear change in PRI response to LUE, which is related to leaf pigment content. The use of an estimated or approximated PRI0, defined as the PRI of perfectly dark‐adapted leaves, allowed us to separate the PRI variability due to leaf pigment content changes and the physiologically related PRI variability over both daily (PAR‐related) and seasonal (soil water content‐related) scales. The corrected PRI obtained by subtracting PRI0 from the PRI measurements showed a good correlation with the LUE over both of the species, soil water statuses and over the entire growing season.     

Physiology of the seasonal relationship between the photochemical reflectance index and photosynthetic light use efficiency

The photochemical reflectance index (PRI) is regarded as a promising proxy to track the dynamics of photosynthetic light use efficiency (LUE) via remote sensing. The implementation of this approach requires the relationship between PRI and LUE to scale not only in space but also in time. The short-term relationship between PRI and LUE is well known and is based on the regulative process of non-photochemical quenching (NPQ), but at the seasonal timescale the mechanisms behind the relationship remain unclear. We examined to what extent sustained forms of NPQ, photoinhibition of reaction centres, seasonal changes in leaf pigment concentrations, or adjustments in the capacity of alternative energy sinks affect the seasonal relationship between PRI and LUE during the year in needles of boreal Scots pine. PRI and NPQ were highly correlated during most of the year but decoupled in early spring when the foliage was deeply downregulated. This phenomenon was attributed to differences in the physiological mechanisms controlling the seasonal dynamics of PRI and NPQ. Seasonal adjustments in the pool size of the xanthophyll cycle pigments, on a chlorophyll basis, controlled the dynamics of PRI, whereas the xanthophyll de-epoxidation status and other xanthophyll-independent mechanisms controlled the dynamics of NPQ at the seasonal timescale. We conclude that the PRI leads to an underestimation of NPQ, and consequently overestimation of LUE, under conditions of severe stress in overwintering Scots pine, and most likely also in species experiencing severe drought. This severe stress-induced decoupling may challenge the implementation of the PRI approach.     

Constitutive changes in pigment concentrations: implications for estimating isoprene emissions using the photochemical reflectance index

The photochemical reflectance index (PRI), through its relationship with light use efficiency (LUE) and xanthophyll cycle activity, has recently been shown to hold potential for tracking isoprene emissions from vegetation. However, both PRI and isoprene emissions can also be influenced by changes in carotenoid pigment concentrations. Xanthophyll cycle activity and changes in carotenoid concentrations operate over different timescales, but the importance of constitutive changes in pigment concentrations for accurately estimating isoprene emissions using PRI is unknown. To clarify the physiological mechanisms behind the PRI–isoprene relationship, the light environment of potted Salix viminalis (osier willow) trees was modified to induce acclimation in photosynthetic rates, phytopigments, isoprene emissions and PRI. Acclimation resulted in differences in pigment concentrations, isoprene emissions and PRI. Constitutive changes in carotenoid concentration were significantly correlated with both isoprene emissions and PRI, suggesting that the relationship between PRI and isoprene emissions is significantly influenced by constitutive pigment changes. Consequently knowledge regarding how isoprene emissions are affected by both longer term changes in total carotenoid concentrations and shorter term dynamic adjustments of LUE is required to facilitate interpretation of PRI for monitoring isoprene emissions.     

Relationships between the photochemical reflectance index (PRI) and chlorophyll fluorescence parameters and plant pigment indices at different leaf growth stages

This study aimed to evaluate the photochemical reflectance index (PRI) for assessing plant photosynthetic performance throughout the plant life cycle. The relationships between PRI, chlorophyll fluorescence parameters, and leaf pigment indices in Solanum melongena L. (aubergine; eggplant) were studied using photosynthetic induction curves both in short-term (diurnal) and long-term (seasonal) periods under different light intensities. We found good correlations between PRI/non-photochemical quenching (NPQ) and PRI/electron transport rate (ETR) in the short term at the same site of a single leaf but these relationships did not hold throughout the life of the plant. In general, changes in PRI owing to NPQ or ETR variations in the short term were <20?% of those that occurred with leaf aging. Results also showed that PRI was highly correlated to plant pigments, especially chlorophyll indices measured by spectral reflectance. Moreover, relationships of steady-state PRI/ETR and steady-state PRI/photochemical yield of photosystem II (Φ(PSII)) measured at uniform light intensity at different life stages proved that overall photosynthesis capacity and steady-state PRI were better correlated through chlorophyll content than NPQ and xanthophylls. The calibrated PRI index accommodated these pigments effects and gave better correlation with NPQ and ETR than PRI. Further studies of PRI indices based on pigments other than xanthophylls, and studies on PRI mechanisms in different species are recommended.     

Photosynthesis, chlorophyll fluorescence and spectral reflectance in Sphagnum moss at varying water contents

Moss samples from the Fluxnet-Canada western peatland flux station in the Boreal Region of Alberta were measured in the laboratory to obtain the net photosynthesis rate and chlorophyll fluorescence of the moss under controlled environmental conditions, including the regulation of moss water content, simultaneously with measurements of moss spectral reflectance. One objective was to test whether the photochemical reflectance index (PRI) detected changes in moss photosynthetic light-use efficiency that were consistent with short-term (minutes to hours) changes in xanthophyll cycle pigments and associated changes in non-photochemical quenching (NPQ), as recorded by chlorophyll fluorescence. The rate of net photosynthesis was strongly inhibited by water content at values exceeding approximately 9 (fresh weight/dry weight) and declined as the water content fell below values of approximately 8. Chlorophyll fluorescence measurements of maximum photosystem II efficiency generally remained high until the water content was reduced from the maximum of about 20 to values of approximately 10–11, and then declined with further reductions in moss water content. A significant linear decline in NPQ was observed as moss water content was reduced from maximum to low water content values. There was a strong negative correlation between changes in NPQ and PRI. These data suggest that PRI measurements are a good proxy for short-term shifts in photosynthetic activity in Sphagnum moss. A second objective was to test how accurately the water band index (WBI, ratio of reflectance at 900 and 970 nm) recorded changes in moss water content during controlled laboratory studies. Strong linear relationships occurred between changes in moss water content and the WBI, although the slopes of the linear relationships were significantly different among sample replicates. Therefore, WBI appeared to be a useful tool to determine sample-specific water content without destructive measurements.     

Confounding effects of snow cover on remotely sensed vegetation indices of evergreen and deciduous trees: An experimental study

Located at northern latitudes and subject to large seasonal temperature fluctuations, boreal forests are sensitive to the changing climate, with evidence for both increasing and decreasing productivity, depending upon conditions. Optical remote sensing of vegetation indices based on spectral reflectance offers a means of monitoring vegetation photosynthetic activity and provides a powerful tool for observing how boreal forests respond to changing environmental conditions. Reflectance-based remotely sensed optical signals at northern latitude or high-altitude regions are readily confounded by snow coverage, hampering applications of satellite-based vegetation indices in tracking vegetation productivity at large scales. Unraveling the effects of snow can be challenging from satellite data, particularly when validation data are lacking. In this study, we established an experimental system in Alberta, Canada including six boreal tree species, both evergreen and deciduous, to evaluate the confounding effects of snow on three vegetation indices: the normalized difference vegetation index (NDVI), the photochemical reflectance index (PRI), and the chlorophyll/carotenoid index (CCI), all used in tracking vegetation productivity for boreal forests. Our results revealed substantial impacts of snow on canopy reflectance and vegetation indices, expressed as increased albedo, decreased NDVI values and increased PRI and CCI values. These effects varied among species and functional groups (evergreen and deciduous) and different vegetation indices were affected differently, indicating contradictory, confounding effects of snow on these indices. In addition to snow effects, we evaluated the contribution of deciduous trees to vegetation indices in mixed stands of evergreen and deciduous species, which contribute to the observed relationship between greenness-based indices and ecosystem productivity of many evergreen-dominated forests that contain a deciduous component. Our results demonstrate confounding and interacting effects of snow and vegetation type on vegetation indices and illustrate the importance of explicitly considering snow effects in any global-scale photosynthesis monitoring efforts using remotely sensed vegetation indices.     

穿透雨减少下锐齿栎叶片光合色素季节动态及其反射光谱响应

通过林地穿透雨排除的方法模拟降雨减少,测定河南宝天曼自然保护区锐齿栎叶片光合色素含量与反射光谱的季节变化,对减雨处理造成的光合色素变化及其反射光谱的变化进行了定量分析,并探讨了水分控制条件下反射光谱对叶片光合色素变化的响应机制.结果表明: 锐齿栎叶片的光合色素含量和色素比率均呈现明显的季节变化.减雨样地与对照样地叶片的光合色素含量和比率在生长季的各个时期存在差异,其中,叶片叶绿素b(Chl b)含量的差异显著,说明Chl b对减雨处理的敏感性最高,叶片类胡萝卜素(Car)含量的差异较小,说明Car对减雨处理的敏感性相对较弱.550 nm处的光谱反射率对色素季节变化的响应最敏感,以其构造的简单比值指数(SR_750,550)与叶片Chl a、Chl b、总Chl和Car含量的正相关关系显著,光化学反射指数(PRI)与叶片Car/Chl的负相关关系显著.550 nm处的光谱反射率对减雨处理造成的色素变化响应最为敏感.SR_750,550对减雨处理造成的叶片Chl a、Chl b和总Chl的含量变化表现敏感(P<0.01),对Chl a/b的变化不敏感.PRI对减雨处理造成的叶片Car/Chl变化表现敏感(P<0.01).