高山嵩草气孔导度对环境因子的响应模拟

以青藏高原地区高寒草甸常见建群种高山嵩草(Kobresia pygmaea)为研究对象,利用光合测定仪于2018年7—9月测定高山嵩草气体交换参数与环境因子的日变化值,利用土壤水分测量仪及环刀法、透膜法测得0—10 cm土壤水分数据作为模型变量。首先,采用3种气孔导度模型对高山嵩草气孔导度进行拟合和检验,其次,用Jarvis模型、Leuning模型和Gao模型对高山嵩草气孔导度在不同月份(7,8,9月)日变化的模拟结果进行分析,最后,总结并讨论了气孔导度对3个主要环境因子(光合有效辐射(Photosynthetically active radiation, PAR)、气温(T_air)、饱和水汽压差(Vapor pressure deficit, VPD))的响应特征。主要研究结果:(1)3个气孔导度模型都可以较好地模拟高山嵩草的气孔导度变化,Leuning模型表现最好(R²=0.726),Jarvis模型次之(R²=0.659),Gao模型准确度最低(R²=0.624)。(2)高山嵩草叶片气孔导度...     

半干旱地区3种植物叶片水平的抗旱耐旱特性分析——两个气孔导度模型的应用和比较

在对半干旱区3种植物进行生理生态特性测定的基础上,应用两种气孔导度模型进行参数的非线性拟合,BBL模型平均可以解释77.6%的结果,Gao模型平均可以解释59.3%的结果。但Gao模型作为一个机理性的模型,其参数具有明确的物理意义。模型的行为和敏感性分析结果说明,用BBL计算的气孔导度一般大于Gao模型。BBL模型对于干旱胁迫下的土壤水分亏缺没有响应, 因而不适合用作干旱半干旱区的植物生理生态学分析和生态系统模拟。而Gao模型可以描述在各种水分条件下植物气孔导度的响应。Gao模型的结果说明,与油松 (Pinus tabulaeformis) 和中间锦鸡儿 (Caragana intermedia) 比较,小叶杨 (Populus simonii) 具有最小的抗旱和耐旱能力,油松具有最好的叶片水平的耐旱和抗旱特性,但其气孔导度对土壤水分的不敏感意味着在干旱条件下维持光合作用的同时,也可能会导致过多的水分损失。中间锦鸡儿具有很强的耐旱性,且其气孔导度对土壤水分的变化敏感,二者相结合,中间锦鸡儿可以在土壤水分条件较好的情况下,维持较大的气孔导度以满足光合作用的需要,但在土壤水分胁迫严重的时候能迅速降低气孔导度以保持土壤水分。     

额济纳荒漠河岸胡杨林叶片气孔导度与微环境因子关系的模拟研究

以额济纳荒漠河岸胡杨(Populus euphratica)为研究对象,利用LI-6400光合测定仪于2005年5~9月份观测了胡杨叶片气体交换数据,研究了胡杨叶片气孔导度与光合速率、光合有效辐射与光合速率之间的关系.结果表明:(1)胡杨叶片净光合速率随气孔导度的增大而升高,但当气孔导度增加到一定值后,光合速率的增加变缓慢直至平稳,并主要是非气孔限制因素造成的;Ball-Berry模型(B-B模型)能够很好地描述气孔导度与光合速率之间的关系(R2=0.92).(2)叶片净光合速率随着有效辐射的变化符合非直角双曲线规律(R2=0.99).(3)B-B模型和非直角双曲线光合模型耦合后模拟值与观测值之间存在很好的正相关性(r=0.93),但耦合模型的模拟值还是较实测值偏大.因此,在干旱区还必须考虑水分限制因素对气孔开闭的控制作用,进一步构建适合干旱区生态系统特点的水-碳耦合循环机理模型.     

最优气孔行为理论和气孔导度模拟

气孔调节功能是陆地生态系统碳-水耦合过程中最重要的环节。与即时的气孔导度测量相比, 气孔导度斜率能有效地反映气孔导度对CO₂浓度、饱和水汽压亏缺和光合作用的敏感性, 包含了环境因子对光合作用和临界水分利用效率等的综合影响, 为研究全球变化下陆地生态系统碳-水耦合关系提供了一个简明且综合的框架。气孔导度模型从经验模型、半经验模型发展到机理模型, 经过很多学者的改进, 但是模型参数的生物学意义和变化规律还不明确。鉴于气孔导度斜率方面研究的重要性和研究的不足, 为了加强对气孔导度调节规律的认识, 并减少气孔导度模拟的不确定性, 该文主要综述了长期以来国内外关于最优气孔行为理论和气孔导度模拟方面的研究成果, 其中包括广泛使用的气孔导度模型及参数意义, 讨论影响气孔导度斜率的主要因素以及气孔导度机理模型的应用, 并对最优气孔行为理论和气孔导度模拟方面的研究做了简单展望。     

气孔导度对CO2浓度变化的模拟及其生理机制

基于气孔运动的生理生化机制重点进行了气孔导度(gs)对CO2浓度变化的响应机制分析,并推导得到气孔导度(gs)对CO2浓度变化响应模型,并以9种植物进行了模型验证。结果表明:随着CO2浓度的升高,气孔导度会逐渐降低,且下降的幅度会随着CO2浓度的升高而逐渐减弱。气孔导度对CO2浓度(Cs)变化的响应模型可以表达为gs=gmax/(1+Cs/Cs0),其中式中gmax是最大气孔导度和Cs0是实验常数。该模型较好地模拟了气孔导度随CO2浓度变化的规律,模型参数具有明确的生理意义,与Jarvis模型和Ball-Berry模型相比,该模型如何实现多种环境因子的耦合有待进一步突破。另外,模型是在短期改变叶片CO2浓度的条件下得出的,在CO2浓度长期胁迫下的适用性也有待进一步确认。     

羊草叶片气孔导度特征及数值模拟

对松嫩平原草地羊草叶片气孔导特征及与环境因子关系的研究结果表明,羊草叶片气孔导度日变化与环境因子密切相关,晴天表现为双峰曲线,阴天为单峰曲线,同时叶片气孔导度（gs)对瞬时光合有效辐射（PAR）,叶片与空气间的水汽压亏损（VPD）,空气温度（Ta）反应十分明显,依据野外实测资料,在对国际上两类代表性气孔导度模型验证比较的基础上,建立了适用于羊草草原的羊草叶片气孔导度对环境因子的响应模型gs=PAR(2.01Ta^2 147.74Ta-2321.11)/(444.62 PAR)(-538.04 VPD).     

华北落叶松冠层平均气孔导度模拟及其对环境因子的响应

冠层气孔导度是生态水文学研究中的一个重要参数,研究其对环境因子的响应,能为建立机理性的森林蒸腾模型提供理论依据.本文利用热扩散探针,于2005年5-9月,测定了六盘山叠叠沟小流域华北落叶松人工林树干液流及其同步的环境因子,计算了林分冠层平均气孔导度(gc)并构建了Jarvis形式的冠层平均气孔导度模型,分析了gc对光合有效辐射(PAR)、空气水汽压亏缺(DVP)和土壤相对有效含水率(REW)的响应.结果表明:该模型能有效地计算gc的日变化特征,计算与观测的gc决定系数R2为0.76 (n=952).gc对各环境因子有不同响应关系,并表现为非线性特征.其中PAR是gc的驱动因子,当PAR＜0.35 mmol·m-2·s-1时驱动作用明显,大于此值则驱动作用变小;DVP是gc的限制因子,随着DVP的增加gc降低;REW=41%是gc对土壤水分响应的一个关键阈值,当REW＞41%时,土壤水分对gc的影响较小,当REW＜41%时,土壤水分则成为gc的限制因子.     

不同R∶FR值对菊花叶片气孔特征和气孔导度的影响

以切花菊品种"神马(Jinba)"为试材,2010年10月至2011年2月间在南京信息工程大学试验温室采用不同Red (660±10) nm: Far-red (730±10) nm值的LED光源短日处理,研究了温室切花菊叶片气孔特征和气孔导度对不同R∶FR值的响应。结果表明:不同R∶FR值短日处理35d菊花叶片的上表皮和下表皮的气孔直径分别以R∶FR值4.5和6.5处理最大,均以R∶FR值2.5处理最小,气孔密度和气孔开度均以R∶FR=2.5处理最高,以R∶FR值6.5处理最低,下表皮的气孔密度、气孔开度明显高于上表皮;不同R∶FR值处理叶片的气孔开张比和气孔指数差异不显著;在相同光强下,菊花叶片气孔导度和光合速率由大到小的R∶FR值顺序依次为2.5、4.5、0.5、6.5。叶片气孔导度与气孔指数、气孔密度、气孔开张比和气孔开度成正相关,与气孔长度和气孔宽度呈负相关;R∶FR值在2.5-6.5范围内,随光质中红光成分增加,叶片气孔密度、气孔指数、气孔开度、气孔开张比和气孔导度显著降低。     

胡杨叶片气孔导度特征及其对环境因子的响应

依据2005年对极端干旱区荒漠河岸林胡杨的观测资料,对胡杨气孔运动进行了分析研究以揭示胡杨的水分利用特征与抗旱机理。结果表明:(1)胡杨叶片气孔导度日变化呈现为周期波动曲线,其波动周期为2 h,傍晚(20:00)波动消失;净光合速率和蒸腾速率与气孔导度的波动相对应而呈现同步周期波动。(2)胡杨的阳生叶气孔导度高于阴生叶,且不同季节气孔导度值不同,阳生叶气孔导度的季节变幅大于阴生叶。(3)胡杨气孔导度与气温、相对湿度和叶水势有显著相关关系,当CO2浓度较小时,胡杨气孔导度随CO2浓度的增加而增加,当CO2浓度达到一定值后气孔导度不再增加,反而随CO2浓度的增加大幅度降低。(4)胡杨适应极端干旱区生境的气孔调节机制为反馈式反应,即由于叶水势降低导致气孔导度减小,从而减少蒸腾耗水,达到节约用水、适应干旱的目的,表明胡杨的水分利用效率随气孔限制值的增大而减小,二者呈显著负相关。     

基于叶片尺度观测数据的气孔导度模型评价

气孔导度(g)是控制冠层与大气之间能量和水分交换的重要因素。空气湿度是控制植物叶片气孔导度的一个关键环境因子。在过去的十几年中,普遍得到应用的是Ball-Woodrow-Berry(BWB)模型和Leuning模型中气孔导度与湿度的关系。本研究使用一个诊断变量f(H),基于农田叶片水平的光合-气孔导度观测数据,对BWB模型、Leuning模型以及新发展的power-h模型和power-D模型进行了气孔导度模拟效果的比较和评价。结果表明:BWB模型描述的是g和相对湿度(hs)之间的一种线性关系,当空气较为湿润时,模拟结果存在较大的低估;Leuning模型中反映的是g与饱和水汽压差(Ds)的非线性函数,降低了模拟结果的误差,但仍然不能很好地描述g在较湿状况下的显著升高;相比之下,两个新的模型,即Ds的指数函数和(1-hs)的指数函数形式模型能提高模拟结果的精度。这个研究结果也表明基于Ds的模型模拟效果要好于基于hs的模型。     

幼龄胡杨气孔行为对土壤质地和地下水埋深变化的响应

气孔调节是植物适应水分条件变化的关键途径,研究多变生境中植物气孔行为对认识植物的适应具有重要意义。洪水漫溢新形成的河漫滩是胡杨更新的自然生境,其土壤质地和地下水埋深具高度时空异质性。已有研究主要集中于胡杨对地下水埋深变化的生理生态响应,而对土壤质地与地下水变化交互作用影响植物水分关系的认识不足。通过设置土壤质地(砂土(S₁)、砂壤土(S₂)、黏壤土(S₃)与地下水埋深(W₁(30 cm)、W₂(60 cm)、W₃(90 cm))交互试验模拟幼龄胡杨自然生境,观测分析了不同条件下胡杨气孔导度(G_s)、气孔导度斜率(g₁)、光合的气孔限制(L_s)的变化。研究结果表明：(1)胡杨气孔行为对地下水变化的响应受土壤质地影响;(2)相同地下水埋深时不同土质间G_s具显著差异,W₁时S₂与S₃的G_s...     

Stomatal responses to CO2 during a diel Crassulacean acid metabolism cycle in Kalanchoe daigremontiana and Kalanchoe pinnata

To investigate the diurnal variation of stomatal sensitivity to CO₂, stomatal response to a 30 min pulse of low CO₂ was measured four times during a 24 h time-course in two Crassulacean acid metabolism (CAM) species Kalanchoe daigremontiana and Kalanchoe pinnata , which vary in the degree of succulence, and hence, expression and commitment to CAM. In both species, stomata opened in response to a reduction in p CO₂ in the dark and in the latter half of the light period, and thus in CAM species, chloroplast photosynthesis is not required for the stomatal response to low p CO₂. Stomata did not respond to a decreased p CO₂ in K. daigremontiana in the light when stomata were closed, even when the supply of internal CO₂ was experimentally reduced. We conclude that stomatal closure during phase III is not solely mediated by high internal p CO₂, and suggest that in CAM species the diurnal variability in the responsiveness of stomata to p CO₂ could be explained by hypothesizing the existence of a single CO₂ sensor which interacts with other signalling pathways. When not perturbed by low p CO₂, CO₂ assimilation rate and stomatal conductance were correlated both in the light and in the dark in both species.     

Photosynthesis affects following night leaf conductance in Vicia faba

Night-time stomatal opening in C₃ plants may result in significant water loss when no carbon gain is possible. The objective of this study was to determine if endogenous patterns of night-time stomatal opening, as reflected in leaf conductance, in Vicia faba are affected by photosynthetic conditions the previous day. Reducing photosynthesis with low light or low CO₂ resulted in reduced night-time stomatal opening the following night, irrespective of the effects on daytime stomatal conductance. Likewise, increasing photosynthesis with enriched CO₂ levels resulted in increased night-time stomatal opening the following night. Reduced night-time stomatal opening was not the result of an inability to regulate stomatal aperture as leaves with reduced night-time stomatal opening were capable of greater night-time opening when exposed to low CO₂. After acclimating plants to long or short days, it was found that night-time leaf conductance was greater in plants acclimated to short days, and associated with greater leaf starch and nitrate accumulation, both of which may affect night-time guard cell osmotic potential. Direct measurement of guard cell contents during endogenous night-time stomatal opening will help identify the mechanism of the effect of daytime photosynthesis on subsequent night-time stomatal regulation.     

麦田冠层气孔导度的分层研究

小麦灌浆期和乳熟期冠层各层叶片上、下表面的气孔导度之间呈正相关关系;冠层不同层的叶片气孔导度从早到傍晚有平行变化的趋势,数值上存在较大的差异,一般从冠层上到下递减。经分析,这主要与冠层叶片接受的光强自上而下递减有关,且这时所对应的叶片水势自冠层上到下递增的幅度大。测算结果表明,冠层气孔导度白天亦呈明显的日变化,灌浆期的值大于乳熟期的值。     

几个气孔模型在自然条件下的适用性

在自然条件下,用气孔下腔与叶面间的水汽压差（ＶＰＤｓ）取代原有气孔模型中的大气湿度因子,可以明显提高气孔模型在自然条件下的适用性。理论分析指出,在气孔模型中,用ＶＰＤｓ表达气孔导度对湿度的响应与用蒸腾速率表达气孔导度对蒸腾失水的响应是等价的。     

Photosynthetic CO2 uptake byZea mays leaves as influenced by unilateral irradiation of adaxial and abaxial leaf surfaces

A study was made on the effect of increasing photon fluence rate (I) at a unilateral irradiation of adaxial (normal leaf position) and abaxial (inverse leaf position) blade surface of maize leaves of various insertion levels on net photosynthetic CO₂ uptake (P _n ) by the leaves, as well as the contribution of individual surfaces toP _n of the leaves, and the significance of, or relationship between the stomatal (g _s ) and intracellular (gm) conductances at the CO₂ transport.P _n of leaves of various age according to their insertion level was unaffected by the direction of incident irradiation. Upon irradiation of the leaves in normal and inverse position the contribution of the adaxial and abaxial surfaces toP _n ,g _s and g_m was different. On irradiating the leaves in normal position, the contribution of the irradiated adaxial surface to the characteristics mentioned made on the average 55% of total values, the contribution of the abaxial surface irradiated in inverse position made on the average 70% inP _n andg _m , and 80% ing _s . At lowerI’s g _m was higher thang _s both in irradiated and non-irradiated surfaces. The ratio ofg _s to g_m gradually got square with increasingI. In the irradiated adaxial surface the equilibrium (g _s /g _m = 1.0) took place at the highestI’s, in the irradiated abaxial surface between 500 to 1000 μmol m⁻² s⁻¹. The significance of the ratiog _m in the CO₂ transport through the individual surfaces is discussed.     

黄土高原苹果园蒸腾导度大气驱动规律比较

植物蒸腾导度是表征土壤-植物-大气连续体（SPAC）中植物-大气间水汽传导过程、反映植物水分调控能力的一类重要变量,常见有冠层导度（G_c）、冠层气孔导度（G_s）与叶片气孔导度（g_s）,明确三者在反映冠层蒸腾过程时的异同或关联性对于理解植物水分利用机制具有重要意义。本研究基于对黄土高原果园苹果树生长季内树干液流（J_s）及环境因子的连续观测,计算了G_c、G_s及脱耦联系数（Ω）等变量,并与短期连续观测的叶片气孔导度（g_s）比较,分析了G_c、G_s和g_s在反映冠层蒸腾特征方面的异同及其关系。结果表明,日变化过程中G_s、g_s呈"单峰"型曲线,而G_c则呈"先增后减,午后抬升"的"双峰"型曲线。g_s与G_s存在较紧密的线性关系（R²=0.80）,但与G_c的线性关系较弱（R²=0.02）。G_c、G_s均随大气水汽压亏缺（VPD）的变化呈现确定的规律,其中,上边界函数呈递减的对数函数关系,平均值则符合先增后减的Log-Normal函数关系（R²>0.95）,拐点对应的VPD值分别为1.33和1.16 kPa。在一日内,G_s对VPD变化的响应过程与g_s对VPDL （基于叶片温度计算的水汽压亏缺）变化的响应过程总体一致,其一致性高于G_c对VPD变化的响应。整个生长季（4-10月）中果树的Ω平均值为0.12,随着Ω递减,G_c与G_s的线性相关性愈趋紧密,其斜率呈递增趋势,G_c越来越趋近于G_s。研究结果表明,在北方地区,基于树干液流的监测能较准确的推导整株并估算林分的冠层蒸腾导度。与实测g_s的变化过程比较,G_s比G_c具有更高的一致性,G_s可以作为描述苹果树水分利用过程响应大气驱动的更为恰当的变量。     

Acclimation to humidity modifies the link between leaf size and the density of veins and stomata

The coordination of veins and stomata during leaf acclimation to sun and shade can be facilitated by differential epidermal cell expansion so large leaves with low vein and stomatal densities grow in shade, effectively balancing liquid‐ and vapour‐phase conductances. As the difference in vapour pressure between leaf and atmosphere (VPD) determines transpiration at any given stomatal density, we predict that plants grown under high VPD will modify the balance between veins and stomata to accommodate greater maximum transpiration. Thus, we examined the developmental responses of these traits to contrasting VPD in a woody angiosperm (Toona ciliata M. Roem.) and tested whether the relationship between them was altered. High VPD leaves were one‐third the size of low VPD leaves with only marginally greater vein and stomatal density. Transpirational homeostasis was thus maintained by reducing stomatal conductance. VPD acclimation changed leaf size by modifying cell number. Hence, plasticity in vein and stomatal density appears to be generated by plasticity in cell size rather than cell number. Thus, VPD affects cell number and leaf size without changing the relationship between liquid‐ and vapour‐phase conductances. This results in inefficient acclimation to VPD as stomata remain partially closed under high VPD.     

Stomatal response of hybrid poplar to incident light, sudden darkening and leaf excision

Responses of abaxial and adaxial stomata of Populus trichocarpa Torr. & Gray. × P. deltoides Bartr. (ex Marsh.) cv. Unal to incident light, sudden darkening and leaf excision in the light and in the dark were studied on 5-year-old trees in the field using diffusion porometry. Stomatal closure in the dark was found to be incomplete in most cases studies. Stomata closed after leaf excision in the dark within 90 min. Stomatal closure after darkening of an entire tree or an entire branch (white the rest of the tree was in the light) was slower, and complete stomatal closure was noticed only for adaxial stomata after 3 h. Adaxial stomata were more reactive and sensitive than abaxial stomata to sudden darkening and leaf excision in the light and the dark. In all treatments, stomatal response was more responsive in mature leaves than in young, still expanding leaves.     

Internal conductance does not scale with photosynthetic capacity: implications for carbon isotope discrimination and the economics of water and nitrogen use in photosynthesis

Central paradigms of ecophysiology are that there are recognizable and even explicit and predictable patterns among species, genera, and life forms in the economics of water and nitrogen use in photosynthesis and in carbon isotope discrimination (delta). However most previous examinations have implicitly assumed an infinite internal conductance (gi) and/or that internal conductance scales with the biochemical capacity for photosynthesis. Examination of published data for 54 species and a detailed examination for three well-characterized species--Eucalyptus globulus, Pseudotsuga menziesii and Phaseolus vulgaris--show these assumptions to be incorrect. The reduction in concentration of CO2 between the substomatal cavity (Ci) and the site of carbon fixation (Cc) varies greatly among species. Photosynthesis does not scale perfectly with gi and there is a general trend for plants with low gi to have a larger draw-down from Ci to Cc, further confounding efforts to scale photosynthesis and other attributes with gi. Variation in the gi-photosynthesis relationship contributes to variation in photosynthetic 'use' efficiency of N (PNUE) and water (WUE). Delta is an information-rich signal, but for many species only about two-thirds of this information relates to A/gs with the remaining one-third related to A/gi. Using data for three well-studied species we demonstrate that at common WUE, delta may vary by up to 3 per thousand. This is as large or larger than is commonly reported in many interspecific comparisons of delta, and adds to previous warnings about simplistic interpretations of WUE based on delta. A priority for future research should be elucidation of relationships between gi and gs and how these vary in response to environmental conditions (e.g. soil water, leaf-to-air vapour pressure deficit, temperature) and among species.