冠层部位和叶龄对红松光合蒸腾特性的影响

利用Li-6400便携式CO2/H2O红外气体分析仪测定了红松不同冠层部位和叶龄针叶的光合蒸腾特性及其环境影响因子.结果表明:冠层部位和叶龄显著地影响最大净光合速率(Pmax)、光饱和点(LSP)、光补偿点(LCP)、表观最大量子效率(α)、蒸腾速率(Tr)和比叶面积(SLA),但对水分利用效率(WUE)影响不显著.随着冠层部位的下降和叶龄的增加,红松针叶的Pmax逐渐下降,其平均值变动在6·55～9·05μmol·m-2·s-1之间.不同冠层部位和叶龄针叶的LSP和LCP的差异很大,以树冠中部针叶对弱光和强光的利用能力最大.Tr随着冠层部位的下降而降低;不同叶龄针叶的Tr在1·37～1·59mmol·m-2·s-1之间变化.不同部位和叶龄红松针叶的Tr和光合有效辐射存在极显著正相关关系(R2=0·967).红松的WUE与净光合速率紧密相关(R2=0·860).随冠层部位的上升和叶龄的增大,红松针叶的SLA递减,分别在6·61～8·41m2·kg-1和6·65～8·38m2·kg-1之间波动.     

白马雪山阴坡林线长苞冷杉(Abies georgei)种群结构特征

长苞冷杉(Abies georgei)是青藏高原特有林线树种.对白马雪山阴坡海拔4400 m以上林线0.8 hm2样地长苞冷杉种群进行每木调查,分析其种群结构、数量特征及空间分布格局.结果表明:(1)结构呈典型金字塔型,幼苗和幼树在种群中所占比重大,表现为增长型种群,种群个体数随径级的增加而减少,密度为幼苗》幼树》成年树;(2)存活曲线接近Deevy-Ⅲ型,高径级种群趋于稳定,种群具两个死亡高峰,低径级种群尤其幼苗死亡率高达90%,这是林线区的气候条件如低温、强光照、积雪及冬季冻害等综合作用的结果,种群另一死亡高峰出现于V～VI龄级,种内和种间对空间、光照和养分等生存因子的激烈竞争引起自疏,导致死亡率再度上升;(3)长苞冷杉种群各龄级空间点格局在不同尺度上表现为聚集、随机和均匀分布,以聚集分布为主,由于幼苗来源于种子库且依赖成年树的微生境,幼苗聚集强度和尺度都最大;各龄级关系密切,在不同尺度上表现出显著的相关性.     

干扰对高山林线再形成过程的影响

高山林线是一类典型的生态交错带,因其特殊的结构和功能以及对外界环境的高度敏感性而成为全球气候变化研究的热点之一.本文简要介绍了高山林线的相关概念及其界定,从高山林线海拔位置波动、植被格局变化、生态交错带物种组成变化及其生理生态特征变化等几个方面阐述了干扰对高山林线再形成过程的不同影响,总结了高山林线物种对干扰的两种基本响应方式,即退行和入侵.认为人为干扰在一定程度上弱化了当前气候变暖对高山林线波动的影响,因而在不同地区必须紧密结合当地可能的干扰来讨论高山林线的波动,否则结果有可能因误差较大而失去应有的价值.指出该研究在高海拔地区进行植被恢复的指导意义.     

Development of acute frost drought in Rhododendron ferrugineum at the alpine timberline

Summary Acute winter frost drought in Rhododendron ferrugineum at the alpine timberline was provoked by removing the snow that had covered the shrubs and by irradiating and rewarming the twigs in situ in a climatized chamber simulating snow melt conditions. Water potential decreased to -2.3 MPa within three days. After activation of photosynthesis on the 1st day, the COr-uptake decreased during the 2nd day and approached zero at the 3rd day. With advanced desiccation, the internal COr concentration of the leaves increased indicating an impairment of chloroplast functions. The procedure of the disturbances appears to be similar to that of other acute drought events.     

The effects of wind and temperature on cuticular transpiration of Picea abies and Pinus cembra and their significance in dessication damage at the alpine treeline

Summary The importance of high winter winds and plant temperatures as causes of winter desiccation damage at the alpine treeline were studied in the Austrian Alps. Samples of 1- and 2-year twigs of Picea abies and Pinus cembra were collected from the valley bottom (1,000 m a.s.l.), forestline (1,940 m a.s.l.), kampfzone (2.090 m a.s.l.), wind-protected treeline (2,140 m a.s.l.), and wind-exposed treeline (2,140 m a.s.l.). Cuticular transpiration was measured at three different levels of wind speed (4, 10, and 15 ms^-1) and temperature (15°, 20°, and 25° C). At elevated wind speeds slight increases in water loss were observed, whereas at higher temperatures much greater increases occurred. Studies on winter water relations show a significant decline in the actual moisture content and osmotic potentials of twigs, especially in the kampfzone and at treeline. The roles of high winds and temperatures in depleting the winter water economy and causing desiccation damage in the alpine treeline environment are discussed.     

Changes in radial tree growth for Picea abies, Larix decidua, Pinus cembra and Pinus uncinata near the alpine timberline since 1750

Changes in radial growth of the four coniferous species growing in the French Alps near the upper treeline are investigated. Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.], European larch (Larix decidua Mill.), Swiss stone pine (Pinus cembra L.) and mountain pine (Pinus uncinata Mill. ex Mirb.) were sampled by taking 1320 cores and analysing tree-ring widths. Sites were chosen in various climatic conditions (macroclimate and aspect) and on two kinds of bedrock in order to take into account the ecological behaviour of these species. Belledonne, Moyenne-Tarentaise, Haute-Maurienne and Briançonnais areas were sampled along increasing gradients of summer aridity and winter continentality. The calculation of time series after removing the age trend brings strong evidence for an increase in radial growth during the two last centuries, but with different stages and fluctuations for each species. This growth trend is significantly enhanced since 1860 for the spruce, and since 1920 for the two pine species. Furthermore, it also appears on Larix decidua with the same pattern despite periodical growth reduction due to attacks of the larch bud moth (Zeiraphera diniana Gn.). The analysis of ring-widths at a given cambial age reveals that this enhanced phenomenon is observed especially during the tree’s early years (25–75 years). The analysis of four regional climatic series, and three longer series of temperature (in farther single sites) reveals synchronous decadal fluctuations and an evident secular increase in minimum temperatures (especially in January and from July to October), that may be involved in tree-growth enhancement. Thermic amplitudes are significantly reduced during the whole growing period, what is more pronounced in Belledonne, the most oceanic region. Long term growth changes are well described by stepwise regression models, especially for the pine species. These models involved both a linear trend (CO₂ concentration or N-deposition) and low frequency of Turin monthly temperatures. However, they show different patterns than those observed from response functions at a yearly scale.     

Winter at the alpine timberline causes complex within-tree patterns of water potential and embolism in Picea abies

Winter temperatures at the alpine timberline cause ice formation in the xylem of conifers blocking water uptake as well as water shifts within the axes system. This amplifies drought stress that, in combination with freeze–thaw events, causes embolism. This study focussed on within-tree patterns of water potential (ψ) and embolism in Norway spruce [ Picea abies (L.) Karst.]. At five sampling dates in midwinter, ψ was determined at numerous positions in the crown of three trees, and at the end of March, the extent of embolism in representative sections of the axes system was analysed. Until 14 March, mean ψ decreased to −3.77 ± 0.11 MPa with less negative ψ in exposed crown parts. On 30 March, ψ was −1.60 ± 0.06 MPa, while loss of conductivity reached up to 100%. Conductivity losses increased with exposition and were highest in the smallest tree. The observed complex within-tree patterns of ψ and embolism were caused by ice blockages and differences in stress intensities within the xylem. High conductivity losses despite moderate ψ in exposed crown parts indicated freeze–thaw events to be a major inducer of winter embolism. Tree size may play a critical role for winter water relations as trees profit from water stored in the stem and in crown parts below the snow cover.     

Direct and indirect climate change effects on photosynthesis and transpiration

Climate change affects plants in many different ways. Increasing CO(2) concentration can increase photosynthetic rates. This is especially pronounced for C(3) plants, at high temperatures and under water-limited conditions. Increasing temperature also affects photosynthesis, but plants have a considerable ability to adapt to their growth conditions and can function even at extremely high temperatures, provided adequate water is available. Temperature optima differ between species and growth conditions, and are higher in elevated atmospheric CO(2). With increasing temperature, vapour pressure deficits of the air may increase, with a concomitant increase in the transpiration rate from plant canopies. However, if stomata close in response to increasing CO(2) concentration, or if there is a reduction in the diurnal temperature range, then transpiration rates may even decrease. Soil organic matter decomposition rates are likely to be stimulated by higher temperatures, so that nutrients can be more readily mineralised and made available to plants. This is likely to increase photosynthetic carbon gain in nutrient-limited systems. All the factors listed above interact strongly so that, for different combinations of increases in temperature and CO(2) concentration, and for systems in different climatic regions and primarily affected by water or nutrient limitations, photosynthesis must be expected to respond differently to the same climatic changes.     

Hydraulic efficiency and safety of leader shoots and twigs in Norway spruce growing at the alpine timberline

Xylem within trees varies in its hydraulic efficiency and safety. Trees at the alpine timberline were expected to exhibit a hydraulic architecture protecting the leader shoot from winter embolism. Hydraulic and related anatomical parameters were compared as well as seasonal courses of winter embolism in leader shoots and twigs of Norway spruce trees growing at 2000 m. Leader shoots had a 1.4-fold higher specific hydraulic conductivity (ks) as well as a 4.9-fold higher leaf specific conductivity (kl) than side twigs. Vulnerability to drought-induced embolism was lower in leader shoots with a 50% loss of conductivity occurring at a water potential (Psi 50) 0.7 MPa lower than in twigs. Higher ks and kl were related to 1.2-fold wider tracheid diameters in leader shoots. Lower vulnerability corresponded to smaller pit dimensions but not to wood density. High ks and kl reflect the hydraulic dominance of the leader shoot, which is important for its water supply during summer. Low vulnerability protects the leader shoot from embolism during the winter season. In field measurements at the timberline during the winter of 2001/2002, conductivity losses of up to 56% were observed only in twigs while leader shoots showed little or no embolism. Results demonstrate that leader shoot xylem is both hydraulically efficient and safe.     

高寒草地不同坡向披针叶黄华蒸腾速率与叶性状的关系

植物蒸腾速率(Tr)与叶性状间的协同变异关系,对理解异质性生境下植物叶片形态构建模式及其生态适应性具有重要意义。利用ArcGIS建立研究区域的数字高程模型(DEM),并提取样地坡度数据,研究了祁连山高寒草地不同坡向披针叶黄华叶性状与Tr的关系。结果表明:随着坡向由北坡向东坡、西坡、南坡转变,草地群落的密度、高度和土壤含水量逐渐减小,披针叶黄华叶面积(LA)与Tr呈减小趋势,而叶厚度呈增大趋势;在南坡和北坡披针叶黄华的Tr与LA之间存在极显著正相关关系(P0.01),与叶厚度之间存在极显著负相关关系(P0.01),在东坡和西坡披针叶黄华的Tr与LA之间存在显著正相关关系(P0.05),与叶厚度之间存在显著负相关关系(P0.05)。生长在南坡的披针叶黄华选择了小而厚的叶片和低的Tr,生长在北坡的披针叶黄华选择了大而薄的叶片和高的Tr,体现了异质生境中植物通过叶片生物量分配机制实现资源有效利用的生存策略。     

高山林线形成机理及植物相关生理生态学特性研究进展

高山林线作为极端环境条件下树木生存的界限,由于其对气候变化的敏感性,在全球变化研究中得到了广泛关注。研究高山林线形成机理以及林线地带植物相关生理生态学特性成为预测未来气候变化条件下植被动态变化的出发点。对于高山林线形成机理研究主要关注两方面问题：（1）林线地带外界环境如何限制乔木生长和分布,其内在机理如何;（2）灌木及草本相对于乔木在林线地区有哪些生存优势,从乔木到灌木及草本生活型演变的功能及意义如何。综述了当前高山林线形成机理及相关生理生态特性的国内外最新研究成果,指出尽管温度（尤其是生长季低温）在全球尺度上能解释大部分高山区域林线的分布,但树木生长和生存受限的内在机理并没有弄清楚,目前主要存在“碳受限”以及“生长受限”假说两大争论焦点。另外,理论上受温度控制的高山林线对气候变化的响应表现出不同的模式,表明全球变化对林线分布和植被生长影响的复杂性和不确定性。因此,未来的研究应该关注影响林线地区植被生长的多种生理生态学过程,比如水分及养分利用过程,以及从乔木到灌木及草本生活型演变的功能意义,从而为林线形成机理以及对气候变化的响应提供更好的解释。     

环境因子对辣椒光合与蒸腾特性的影响

采用灰色关联分析方法研究环境因子对辣椒(Capsicum annuum Linn.)光合和蒸腾特性的影响.结果表明,在强光照下影响净光合速率的环境因子大小顺序为相对湿度、CO2浓度、温度、光照强度、气孔导度;净光合速率与相对湿度、叶温呈显著的负相关,与CO2浓度呈极显著的正相关;蒸腾速率与气温呈极显著的正相关,与相对湿度呈极显著的负相关;水分利用效率与相对湿度呈极显著的正相关,与温度呈极显著的负相关.弱光照时影响净光合速率的环境因子大小顺序为光照强度、温度、相对湿度、CO2浓度;净光合速率与光照强度、CO2浓度呈显著的正相关,与相对湿度呈显著的负相关.在强光照下适当遮阳、灌水、增施CO2肥,在弱光照下减少荫蔽、通风透气、降低相对湿度、增加温度可促进辣椒光合作用,提高辣椒产量.     

Response of transpiration and photosynthesis to a transient change in illuminated foliage area for a Pinus radiata D. Don tree

Sudden but transient changes in the fraction or illuminated foliage area in a well-watered 7-year-old Pinus radiata D. Don tree were imposed by completely covering either the upper 22% or the lower 78% of the foliage for periods of up to 36 h. Measurements of transpiration flux density (E), tree conductance (g_t), stomatal conductance (g_s) and net photosynthesis (A) were made to test the hypothesis that compensatory responses would occur in the remaining illuminated foliage when the cover was installed. When the lower foliage was covered there was an immediate decrease in g_t. However, when tree conductance was normalized with respect to the illuminated leaf area (g_t'), it increased between 50 and 75%, depending on the value of air saturation deficit (D). The effect was also apparent from concurrent measurements of increases in g_s and A up to 59 and 24%, respectively, for needles in the top third of (he crown. When the cover was removed these effects were reversed. The changes in the lower foliage when the upper foliage was covered were much smaller. Changes in bulk needle water potential were small. It is suggested that the observed responses occurred because of a perturbation to the hydraulic pathway in the xylem that could have triggered the action of a chemical signal to regulate stomatal conductance and photosynthesis.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Mesophyll conductance in land surface models: effects on photosynthesis and transpiration

The CO₂ transfer conductance within plant leaves (mesophyll conductance, g_m) is currently not considered explicitly in most land surface models (LSMs), but instead treated implicitly as an intrinsic property of the photosynthetic machinery. Here, we review approaches to overcome this model deficiency by explicitly accounting for g_m, which comprises the re‐adjustment of photosynthetic parameters and a model describing the variation of g_m in dependence of environmental conditions. An explicit representation of g_m causes changes in the response of photosynthesis to environmental factors, foremost leaf temperature, and ambient CO₂ concentration, which are most pronounced when g_m is small. These changes in leaf‐level photosynthesis translate into a stronger climate and CO₂ response of gross primary productivity (GPP) and transpiration at the global scale. The results from two independent studies show consistent latitudinal patterns of these effects with biggest differences in GPP in the boreal zone (up to ~15%). Transpiration and evapotranspiration show spatially similar, but attenuated, changes compared with GPP. These changes are indirect effects of g_m caused by the assumed strong coupling between stomatal conductance and photosynthesis in current LSMs. Key uncertainties in these simulations are the variation of g_m with light and the robustness of its temperature response across plant types and growth conditions. Future research activities focusing on the response of g_m to environmental factors and its relation to other plant traits have the potential to improve the representation of photosynthesis in LSMs and to better understand its present and future role in the Earth system.     

Effects of guttation prevention on photosynthesis and transpiration in leaves of Alchemilla mollis

The ecophysiological function(s) and consequences of guttation, a phenomenon by which water is exuded by and accumulated as droplets along the leaf margins under high humidity in many plants that grow in wet soil, has been poorly studied and remains largely unknown. Thus, leaf gas exchange and chlorophyll fluorescence were examined, using two experimental approaches, in Alchemilla mollis plants under conditions that promoted guttation and those that prevented this phenomenon. Although results were variable, depending on the experimental approach, prevention of guttation effected reductions in photosynthesis and transpiration, as well as photochemical activity measured with fluorescence techniques. These findings lend partial support for a previously hypothesized function of guttation: prevention of excess water in leaves, yet they contradict those of several other studies. More work is required in order to adequately understand the function of guttation.     

作物光合、蒸腾与水分高效利用的试验研究

通过田间试验,对作物光合、蒸腾、气孔行为及其影响因素进行了研究。结果表明,光合与蒸腾的非线性关系可以用抛物线方程表述,其中光合速率最高时的蒸腾速率为临界值,超出该值即为奢侈蒸腾,干旱处理的临界值较低,通过合适的调控措施,抑制奢侈蒸腾并不影响光合生产,综合分析光合速率、蒸腾速率与气孔导度的关系,气孑L导度大于0．12mol·m^-2·s^-1,实施提高气孔阻力并抑制蒸腾的措施,既节约水分又促进光合作用,增加产量．光合速率基本上随光合有效辐射的增加而提高,并有光饱和点存在,水分条件影响叶片光合作用达到饱和的早晚,干旱处理的光饱和点远远低于湿润处理,强光需要水分充足相耦合,才能充分发挥光能利用率,蒸腾与辐射的线性关系十分显著。从光合有效辐射入手,在光合有效辐射大于1000μmol·m^-2·s^-1时实施措施,既可大大降低蒸腾,又可改善光合,节水增产效果不言而喻。