Effects of ozone and aerosol pollution on photosynthesis of poplar leaves

由于经济的快速发展, 中国大部分地区正面临着严峻的复合型大气污染, 其中臭氧和气溶胶是两种主要污染物。已有的研究表明臭氧对叶片的氧化性伤害能够抑制光合作用, 而气溶胶可通过增加散射辐射比例或缓解高温抑制促进光合作用。但复合污染下, 臭氧和气溶胶如何共同调控叶片光合作用, 仍缺乏研究。该研究利用北京及周边地区之间的污染梯度, 选择加杨(Populus × canadensis)作为实验对象, 于2012-2013年生长季期间对叶片光合速率进行连续观测, 并同时监测臭氧浓度(AOT40)、气溶胶光学厚度(AOD)、空气温度和冠层内外光合有效辐射(PAR)等环境因子, 以期探讨大气复合污染下臭氧和气溶胶变化对植物叶片光合作用的影响及相关机制。结果表明: (1)臭氧浓度与空气温度、气溶胶浓度之间均呈显著正相关关系, 但气溶胶浓度与空气温度没有显著相关关系; (2)臭氧浓度增加显著抑制了阳生叶片的光合作用, 但气溶胶浓度上升促进了阳生叶片的光合作用; 臭氧浓度升高对阴生叶片光合作用的影响较小, 但气溶胶浓度上升促进了阴生叶片的光合作用; (3)标准化后的结果显示, 臭氧对阳生叶片光合作用的影响最大, 此时气溶胶的促进作用一定程度上补偿了臭氧浓度上升所带来的抑制效应。对于阴生叶片光合作用而言, 气溶胶则是最重要的影响因素。该研究发现复合污染下阴生叶和阳生叶光合响应不同, 这表明冠层结构可能通过影响阴生叶和阳生叶的比例, 从而对植物生长产生不同影响。该研究对理解大气复合污染如何影响光合作用提供了的机理支持, 同时也表明, 为了维持生态系统生产力及功能, 需要同时控制气溶胶和臭氧污染。     

黄河小浪底栓皮栎、刺槐叶片电子传递速率-光响应的模拟

为了比较直角双曲线模型、非直角双曲线模型与叶子飘模型的优缺点, 研究阴生叶和阳生叶电子传递速率的差异, 探讨环境/生物因素对电子传递速率等参数的影响, 该文采用LI-6400XT荧光测定系统对黄河小浪底栓皮栎(Quercus variabilis)、刺槐(Robinia pseudoacacia)叶片电子传递速率-光响应(J-I)曲线进行了测定, 利用直角双曲线模型、非直角双曲线模型和叶子飘模型对J-I曲线进行了拟合。结果表明, 3种模型对叶片J-I曲线拟合的决定系数(R ²)在0.96以上, 叶子飘模型的R ²最高(> 0.99)。直角双曲线模型和非直角双曲线模型无法模拟植物叶片光系统II动力学下调现象, 且不能得出饱和光强(I_sat); 直角双曲线模型对最大电子传递速率(J_max)的模拟明显大于实测值; 叶子飘模型能很好地模拟光系统II动力学下调现象, 得出的J_max和I_sat均最接近实测值。对阴生叶和阳生叶J-I曲线研究发现, 栓皮栎、刺槐阴生叶的J_max分别低于阳生叶25.0%和18.0%, 阳生叶的I_sat分别高于阴生叶26.0%和10.1%。栓皮栎和刺槐J_max与气温显著正相关; 刺槐I_sat与气温、土壤水分含量和净光合速率具有显著的正相关关系; 栓皮栎和刺槐J-I曲线初始斜率α均与净光合速率呈显著负相关关系。     

Simulation on the light-response curves of electron transport rate of Quercus variabilis and Robinia pseudoacacia leaves in the Xiaolangdi area,China

为了比较直角双曲线模型、非直角双曲线模型与叶子飘模型的优缺点, 研究阴生叶和阳生叶电子传递速率的差异, 探讨环境/生物因素对电子传递速率等参数的影响, 该文采用LI-6400XT荧光测定系统对黄河小浪底栓皮栎(Quercus variabilis)、刺槐(Robinia pseudoacacia)叶片电子传递速率-光响应(J-I)曲线进行了测定, 利用直角双曲线模型、非直角双曲线模型和叶子飘模型对J-I曲线进行了拟合。结果表明, 3种模型对叶片J-I曲线拟合的决定系数(R ²)在0.96以上, 叶子飘模型的R ²最高(> 0.99)。直角双曲线模型和非直角双曲线模型无法模拟植物叶片光系统II动力学下调现象, 且不能得出饱和光强(I_sat); 直角双曲线模型对最大电子传递速率(J_max)的模拟明显大于实测值; 叶子飘模型能很好地模拟光系统II动力学下调现象, 得出的J_max和I_sat均最接近实测值。对阴生叶和阳生叶J-I曲线研究发现, 栓皮栎、刺槐阴生叶的J_max分别低于阳生叶25.0%和18.0%, 阳生叶的I_sat分别高于阴生叶26.0%和10.1%。栓皮栎和刺槐J_max与气温显著正相关; 刺槐I_sat与气温、土壤水分含量和净光合速率具有显著的正相关关系; 栓皮栎和刺槐J-I曲线初始斜率α均与净光合速率呈显著负相关关系。     

基于叶绿素荧光成像及荧光参数分布特征的叶片光合异质性定量分析

植物叶片光合作用具有高度的空间异质性,叶绿素荧光成像技术为叶片光合异质性的研究提供了便利,但叶片光合异质性的定量分析并没有得到广泛应用。本文利用ImagingPAM叶绿素荧光成像系统,获得 中亚热带地区越冬期小叶榕（Ficus microcarpa）阳生叶和阴生叶的叶绿素荧光参数图像,并利用仪器的分析软件对其进行分析,定量比较了阳生叶和阴生叶的光合异质性特征。研究发现：越冬期小叶榕阳生叶的光合异质性和光抑制程度明显高于阴生叶,变异系数可作为光合异质性的定量指标。低温强光导致阳生叶坏死率（P_LN）达4.30%,并有53.30%的区域处于严重光抑制（0<F_v/F_m<0.627）,但仍有42.27%的区域仅为轻度光抑制（0.627≤ F_v/F_m<0.800）。而低温弱光并未造成阴生叶坏死和严重光抑制。通过对光系统Ⅱ（PSⅡ）的实际光合效率 （Y（Ⅱ））、调节性能量耗散的量子产额（Y（NPQ））和非调节性能量耗散的量子产额（Y（NO））荧光参数异质性的定量分析表明,阳生叶具有相对较高的光化学能力,阴生叶则具有相对较高的热耗散能力;冬季强光虽然会导致小叶榕阳生叶PSⅡ严重激发压积累,存在严重光抑制的潜在风险,但其致死面积并不大,叶片中仍存在一定面积低激发压的低风险区,而低温弱光下的阴生叶则主要以低风险区域为主。     

适度高温胁迫对亚热带森林3种建群树种幼树光合作用的影响

利用Licor-6400光合作用测定系统和叶室荧光仪（Licor-6400LCF）测定适度高温（42℃）胁迫下阳生树种荷木（Schima superba）、耐荫树种黄果厚壳桂（Cryptocarya concinna）和中生性树种红锥（Castanopsis hystrix）在全日光和遮阴（20％全日光）生长下的叶片光合速率和叶绿素a荧光。适度高温胁迫引起全日光和遮阴叶片PSII原初最大光化量子产率（Fv/Tm）降低，反映适度高温胁迫引起PSII功能的部分抑制。其中适度高温对阴生树种黄果厚壳桂和遮阴下生长叶片的PSII抑制较阳生树种荷木在全日光下生长的叶片大。除在全日光下生长的黄果壳桂外，适度高温胁迫能增高全日光或遮阴下生长的荷木和红锥叶片的光合速率。同时亦表现较高的耐高光强抑制的能力。适度高温胁迫降低全日光下生长荷木和红锥叶片的PSII量子产率（ФPSII），但对具有低西ФPSII的阴生树种黄果厚壳桂或在遮阴下生长的阳生树种荷木或中生性树种红锥叶片则影响较小。适度高温胁迫引起生长在全日光下的阳生树种荷木或中生性树种红锥叶片的CO2同化量子需要量降低，但甚少影响阴生树种黄果厚壳桂或遮阴下生长叶片CO2同化量子需要量。适度高温对亚热带森林建群种幼树光合作用的影响依赖于植物种类和叶类型（阳生和阴生叶）。     

珊瑚树阳生和阴生叶片光合特性和状态转换的比较

珊瑚树阳生和阴生叶片是在不同光照环境中长期生长的,它们的光合特性有一些明显的差异.与阳生叶片相比,阴生叶片单位干重的叶绿素含量较多,类囊体膜垛叠程度较高(即每个基粒的类囊体膜垛叠层数较多,基粒类囊体的直径较大),而叶绿素a/b比值、光合作用的饱和光强和最大净光合速率等较低.用弱红光诱导阳生和阴生叶片向状态2转换时,叶绿素荧光Fm/Fo和F685/F735先迅速下降再逐渐回升,这表明两种叶片都先后通过满溢和LHCⅡ转移调节激发能在PSⅡ和PSⅠ之间的分配,改善光能利用,但阳生叶片Fm/Fo和F685/F735下降的幅度较大.     

刺栲（Castanopsis hystrix）的气体交换和叶绿素a荧光特征

用LI-6400便携式光合测定系统和PAM-2100便携式植物荧光分析仪分别在初夏和秋季测定了广州龙眼洞林场刺栲(Castanopsishystrix)幼树叶片的气体交换和叶绿素a荧光参数。结果表明:1)刺栲冠层阳生叶的光饱和点为400μmolphotonsm-2s-1,最大光合速率在4-6μmolm-2s-1之间,说明刺栲幼树保持阳生植物的特征。阴生叶与阳生叶的叶绿素a/b值相似,均约为2.4,而阴生叶叶绿素b含量高于阳生叶,表明阴生叶具有潜在利用弱光的能力;2)刺栲幼树冠层叶片的PSⅡ的最大光化学效率Fv/Fm和光化学量子产额(Yield)的日动态变化在中午略有下降,说明其在强光下能发生可逆性的光抑制。刺栲冠层阳生叶的最大光合能力比较低,整个冠层叶能高效而稳定地利用散射和透射光,这些特征使得常绿的刺栲幼树能在林边或林下良好生长,较好地适应不同的光环境并成功渡过幼苗期,最终成为群落中的上层优势种。     

淮北相山恢复演替群落优势树种叶片的生态解剖

对淮北相山混交林5个优势种叶片的生态解剖学观察表明,其叶片结构具有一定的旱生特征:表皮具发达的表皮毛或角质层,全栅等面叶或具发达的栅栏组织,维管组织发达。牡荆(Vitex negundo var. cannabifolia)和酸枣(Ziziphus jujuba var. spinosa)作为两个广布优势树种,叶片结构表现出很大的可塑性:1)同一群落环境(混交林)中,叶片结构随着季节的变化表现出发育可塑性(5月初的叶片比9月中旬更具有阳生叶的特点);2)不同恢复演替阶段的群落中,叶片结构随着群落环境的变化表现出环境可塑性,其变化趋势为:灌草丛(旱生/阳生)-灌丛(旱生/阳生)-落叶疏林(中生/阳生)-人工侧柏(Platycladus orientalis)林(中生/阴生)。这种可塑性既是植物适应其异质生境的一种重要机制,同时又是不同群落环境的反映。非参数相关分析表明,牡荆和酸枣的叶片结构受多个生态因子综合影响,其中水分和风速是影响叶片结构的主导因子。叶片的上表皮角质层厚度、气孔器密度、栅栏组织厚度、叶片厚度、木质部厚度、韧皮部厚度、维管束厚度等性状均与土壤含水量和空气相对湿度呈显著负相关,与风速呈显著正相关。     

光强对砂仁叶片光合作用光抑制及热耗散的影响

通过测定不同光照条件下砂仁 (AmomumvillosumLour.)叶片气体交换和叶绿素荧光参数 ,探讨了光对其光合机构及其光破坏防御的影响。试验期间 ,上午 11:0 0之前有雾 ,光强较弱。上午砂仁阳生叶净光合速率 (Pn)与下午 (6 .5 3μmol·m-2 ·s-1)相似 ,高于阴生叶 (5 .94μmol·m-2 ·s-1) ,下午阴生叶Pn 高于上午 ,与阳生叶相似。下午砂仁叶片表观量子效率低于上午。其初始荧光 (Fo)、最大荧光 (Fm)、光系统Ⅱ (PSⅡ )最大光能转换效率 (Fv/Fm)、Fm/Fo 及PSⅡ的潜在效率 (Fv/Fo)随日光增强而降低 ,15 :0 0降至最低 ,表明光抑制逐渐加剧。之后随光强减弱这些叶绿素荧光参数升高 ,光抑制得到缓解。与此相反 ,非光化学猝灭系数 (qN)随光强的增加而升高 ,并一直维持在较高水平 ,表明依赖叶黄素循环的保护性反应逐渐增强。阳生叶的光抑制比阴生叶强烈 ,当日遮荫处理使光抑制缓解 ,但各处理间qN 差异不大 ,表明热耗散未受显著影响。结论 :弱光下砂仁叶片即发生光抑制 ,在不同光照下其光抑制的普遍发生 ,是依赖叶黄素循环的保护性反应 ,而非光破坏的结果 ;砂仁叶片叶黄素循环的启动不需过剩光能 ,不同光处理对其影响不大 ;砂仁对光的适应能力较强。     

阳生和阴生地木耳生理生化特性的比较

以生长于不同光环境下的地木耳为材料,对其Fv/Fm的日变化、光合作用特性、叶绿素和类胡萝卜素的含量进行了研究,以了解其光适应的生理生化基础。同阴生地木耳相比,阳生地木耳的光饱和点、类胡萝卜素含量、类胡萝卜素和叶绿素的比值均比较高,但其P-I曲线光限制部分的斜率、暗呼吸速率、Fv/Fm、叶绿素、MAAs含量和单位面积干重较低。二者最大光合速率和光补偿点无明显差异,二者均无明显的光呼吸。同等条件的光抑制后,阳生地木耳在暗处能更快、更大程度地恢复其Fv/Fm活性。原位研究表明,阴生和阳生地木耳在雨后强光下均有不同程度的光抑制发生,但在弱光下或夜晚时会及时恢复。阳生和阴生地木耳的光合特性及色素含量显著不同,以此来适应不同环境中的光因子。     

Interaction of ozone pollution and light effects on photosynthesis in a forest canopy experiment

Ozone pollution may reduce net carbon gain in forests, yet data from mature trees are rare and the effects of irradiance on the response of photosynthesis to ozone remain untested. We used an open-air system to expose 10 branches within the upper canopy of an 18-m-tall stand of sugar maple (Acer saccharum Marsh.) to twice-ambient concentrations of ozone (95nmol mol^?1, 0900 to 1700, 1 h mean) relative to 10 paired, untreated controls (45nmol mol^?1) over 3 months. The branch pairs were selected along a gradient from relatively high irradiance (PPFD 14.5 mol m^?2 d^?1) to deep shade (0.7mol m^?2 d^?1). Ozone reduced light-saturated rates of net photosynthesis (A_sat) and increased dark respiration by as much as 56 and 40%, respectively. Compared to sun leaves, shade leaves exhibited greater proportional reductions in A_sat and had lower chlorophyll concentrations, quantum efficiencies, and leaf absorptances when treated with ozone relative to controls. With increasing ozone dose over time, A_sat became uncoupled from stomatal conductance as ratios of internal to external concentrations of carbon dioxide increased, reducing water-use efficiency. Ozone reduced net photosynthesis and impaired stomatal function, with these effects depending on the irradiance environment of the canopy leaves. Increased ozone sensitivity of shade leaves compared to sun leaves has consequences for net carbon gain in canopies.     

Field evidences for the positive effects of aerosols on tree growth

Theoretical and eddy covariance studies demonstrate that aerosol‐loading stimulates canopy photosynthesis, but field evidence for the aerosol effect on tree growth is limited. Here, we measured in situ daily stem growth rates of aspen trees under a wide range of aerosol‐loading in China. The results showed that daily stem growth rates were positively correlated with aerosol‐loading, even at exceptionally high aerosol levels. Using structural equation modeling analysis, we showed that variations in stem growth rates can be largely attributed to two environmental variables covarying with aerosol loading: diffuse fraction of radiation and vapor pressure deficit (VPD). Furthermore, we found that these two factors influence stem growth by influencing photosynthesis from different parts of canopy. Using field observations and a mechanistic photosynthesis model, we demonstrate that photosynthetic rates of both sun and shade leaves increased under high aerosol‐loading conditions but for different reasons. For sun leaves, the photosynthetic increase was primarily attributed to the concurrent lower VPD; for shade leaves, the positive aerosol effect was tightly connected with increased diffuse light. Overall, our study provides the first field evidence of increased tree growth under high aerosol loading. We highlight the importance of understanding biophysical mechanisms of aerosol‐meteorology interactions, and incorporating the different pathways of aerosol effects into earth system models to improve the prediction of large‐scale aerosol impacts, and the associated vegetation‐mediated climate feedbacks.     

Instantaneous canopy photosynthesis: analytical expressions for sun and shade leaves based on exponential light decay down the canopy and an acclimated non-rectangular hyperbola for leaf photosynthesis

Analytical expressions for the contributions of sun and shade leaves to instantaneous canopy photosynthesis are derived. The analysis is based on four assumptions. First, that the canopy is closed in the sense that it is horizontally uniform. Secondly, that there is an exponential profile of light down the canopy with the same decay constant for light from different parts of the sky. Thirdly, that the leaf photosynthetic response to incident irradiance can be described by a three-parameter non-rectangular hyperbola (NRH). And lastly, that light acclimation at the leaf level occurs in only one parameter of the NRH, that describing the light-saturated photosynthetic rate, which is assumed to be proportional to the local averaged leaf irradiance. These assumptions have been extensively researched empirically and theoretically and their limitations are quite well understood. They have been widely used when appropriate. Combining these four assumptions permits the derivation of algebraic expressions for instantaneous canopy photosynthesis which are computationally efficient because they avoid the necessity for numerical integration down the canopy. These are valuable for modelling plant and crop ecosystems, for which canopy photosynthesis is the primary driver. Ignoring the sun/shade dichotomy can result in overestimates of canopy photosynthesis of up to 20 %, but using a rectangular hyperbola instead of a non-rectangular hyperbola to estimate canopy photosynthesis taking account of sun and shade leaves can lead to a similarly sized underestimate.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. I. Leaf model parametrization

In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m^?2 s^?1) than in sun leaves (11–5μmol m^?2 s^?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).     

The phenology of leaf quality and its within‐canopy variation is essential for accurate modeling of photosynthesis in tropical evergreen forests

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per‐area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO₂ assimilation. We developed a two‐fraction leaf (sun and shade), two‐layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within‐canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground‐based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two‐fraction leaf, two‐layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance‐derived CO₂ assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO₂ assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests.     

Effect of local irradiance on CO(2) transfer conductance of mesophyll in walnut

The acclimation responses of walnut leaf photosynthesis to the irradiance microclimate were investigated by characterizing the photosynthetic properties of the leaves sampled on young trees (Juglans nigraxregia) grown in simulated sun and shade environments, and within a mature walnut tree crown (Juglans regia) in the field. In the young trees, the CO(2) compensation point in the absence of mitochondrial respiration (Gamma*), which probes the CO(2) versus O(2) specificity of Rubisco, was not significantly different in sun and shade leaves. The maximal net assimilation rates and stomatal and mesophyll conductances to CO(2) transfer were markedly lower in shade than in sun leaves. Dark respiration rates were also lower in shade leaves. However, the percentage inhibition of respiration by light during photosynthesis was similar in both sun and shade leaves. The extent of the changes in photosynthetic capacity and mesophyll conductance between sun and shade leaves under simulated conditions was similar to that observed between sun and shade leaves collected within the mature tree crown. Moreover, mesophyll conductance was strongly correlated with maximal net assimilation and the relationships were not significantly different between the two experiments, despite marked differences in leaf anatomy. These results suggest that photosynthetic capacity is a valuable parameter for modelling within-canopies variations of mesophyll conductance due to leaf acclimation to light.     

Rate limiting factors in leaf photosynthesis. II. Electron transport

Increased scattering of a weak 535 nm measuring beam which indicates the light-dependent formation of a transthylakoid proton gradient in leaves was used to examine the role of the electron-transport chain in limiting photosynthetic carbon assimilation. The proton gradient is supported by electron flux and indicates thylakoid energization. In CO₂-free air, half saturation of thylakoid energization was observed at intensities of red light ranging from 2 to 50 W·m⁻² in different plant species. The differences were attributed to different carbohydrate availability for energy-consuming photorespiratory processes when external CO₂ was absent. Thylakoid energization of shade leaves (Asarum, Fagus) was saturated at lower light intensities than that of sun leaves (Phaseolus, Fagus). When photorespiratory carbohydrate oxidation was suppressed by decreasing the O₂ concentration from 21 to 2% in the absence of CO₂, thylakoid energization saturated at lower light intensities than in CO₂-free air. CO₂ decreased thylakoid energization particularly at low light intensities. Under high intensity illumination, however, thylakoid energization was remarkably high even in the presence of saturating CO₂. Apparently, electron transport was capable of maintaining the energy status of the photosynthetic apparatus at a high level even when photosynthetic carbon fluxes were maximal. This suggests that electron transport is less important in limiting photosynthesis than previously thought.     

Beech leaf colonization by the endophyte Apiognomonia errabunda dramatically depends on light exposure and climatic conditions

Ozone and light effects on endophytic colonization by Apiognomonia errabunda of adult beech trees (Fagus sylvatica) and their putative mediation by internal defence compounds were studied at the Kranzberg Forest free-air ozone fumigation site. A. errabunda colonization was quantified by "real-time PCR" (QPCR). A. errabunda-specific primers allowed detection without interference by DNA from European beech and several species of common genera of plant pathogenic fungi, such as Mycosphaerella, Alternaria, Botrytis, and Fusarium. Colonization levels of sun and shade leaves of European beech trees exposed either to ambient or twice ambient ozone regimes were determined. Colonization was significantly higher in shade compared to sun leaves. Ozone exhibited a marginally inhibitory effect on fungal colonization only in young leaves in 2002. The hot and dry summer of 2003 reduced fungal colonization dramatically, being more pronounced than ozone treatment or sun exposure. Levels of soluble and cell wall-bound phenolic compounds were approximately twice as high in sun than in shade leaves. Acylated flavonol 3- O-glycosides with putatively high UV-B shielding effect were very low in shade canopy leaves. Ozone had only a minor influence on secondary metabolites in sun leaves. It slightly increased kaempferol 3- O-glucoside levels exclusively in shade leaves. The frequently prominent hydroxycinnamic acid derivative, chlorogenic acid, was tested for its growth inhibiting activity against Apiognomonia and showed an IC50 of approximately 8 mM. Appearance of Apiognomonia-related necroses strongly correlated with the occurrence of the stress metabolite, 3,3',4,4'-tetramethoxybiphenyl. Infection success of Apiognomonia was highly dependent on light exposure, presumably affected by the endogenous levels of constitutive phenolic compounds. Ozone exerted only minor modulating effects, whereas climatic factors, such as pronounced heat periods and drought, were dramatically overriding.