四种热带雨林树种幼苗比叶重,光合特性和暗呼吸对生长光环境的适应

雾凉季研究了西双版纳热带雨林4种植物幼苗对生长光环境的适应,其中两个树种幼苗喜光（团花和滇南插柚紫）,两个树种幼苗耐荫（滇南红厚壳和玉蕊）发现弱光环境中生长的4种植物比叶重、光合能力、光饱和点、光补偿点暗呼吸速度、叶绿素a/b比较低,叶绿素含量较高。玉蕊和滇南红厚壳幼苗的光合能力和呼吸速度 于团花树和滇南插柚紫。团花树和滇南插机紫的比叶重和光合作用的可塑性大于玉蕊和滇南红厚壳。高光强下生长的团花树和滇南插机紫增加叶氮分配给羧化酶的比较。减少分配给叶绿素的比例。滇南红厚壳和玉蕊适应弱光环境的能力略强于团花树和滇南插机紫,但适应强光的能力较差。研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说,单位干重叶的光合能力和呼吸速率并未表现出利于光适应的可塑性,表明4种植物生理适应能力较差,形态学上的适应在4种热带雨林树种幼苗光适应方面起到了重要的作用,叶氮分配也是它们光适应的策略之一。     

生长光强对4种热带雨林树苗光合机构的影响

于雾凉季研究了西双版纳热带雨林 4种植物幼苗对光抑制及光破坏的防御。发现不同光强下生长的玉蕊和滇南红厚壳日间光抑制均较团花树和滇南插柚紫重 ,在中光强下还发生了长期光抑制。中光强下生长的团花树和滇南插柚紫叶绿素含量降低 ,光合能力升高 ,同时增加热耗散量 ,维持光能平衡 ,避免了光破坏的发生。中光强下玉蕊和滇南红厚壳减少光能吸收、提高热耗散的同时 ,发生了光系统II光化学量子产量长期下调。中光强下生长的 4种植物叶片的类胡罗卜素的绝对含量并不高 ,但单位叶绿素的类胡罗卜素含量显著升高 ,相对保护作用增强。先锋树种团花树类胡罗卜素含量低于另 3种植物 ,这与其较低的热耗散速率相一致 ,光合能力的提高对其光破坏的防御有重要的作用     

不同生态习性热带雨林树种的幼苗对光能的利用与耗散

研究了生长于100％、25％和8％光照条件下的热带雨林先锋树种团花、演替顶极阶段的冠层树种绒毛番龙眼和中下层树种滇南风吹楠幼苗的光合能力及光能分配特性对光强的响应。与绒毛番龙眼和滇南风吹楠相比,团花具有较高的最大光合速率和最大电子传递速率,从光能分配对光强的响应曲线可以看出,随着光强的增加,3个树种幼苗叶片吸收的光能分配到光化学反应的比例减少,分配到热耗散的比例增加,光能在光化学反应与热耗散之间的分配呈显著负相关,与其它两个种相比,100％光下的团花幼苗将较多的光能分配到光化学反应中,热耗散较弱且未达到饱和。过剩光能少,没有引起长期光抑制,绒毛番龙眼和滇南风吹楠将叶片吸收的较多光能分配到热耗散中,但生长于100％光下的幼苗过剩光能仍然较多,导致幼苗遭受长期光抑制,结果表明,不同生态习性热带雨林树种幼苗更新对光环境的要求与这些幼苗对光能的利用和耗散特性密切相关。     

光照对两种热带雨林树种幼苗光合能力、热耗散和抗氧化系统的影响

 研究了生长于不同光照条件下（100%、25%和8%光强）热带雨林冠层树种绒毛番龙眼（Pometia tomentosa）和中层树种滇南风吹楠（Horsfieldia tetratepala）幼苗的光合能力、热耗散、活性氧和保护性酶的活性。结果表明,绒毛番龙眼的最大光合速率随着生长光强的增加而提高,而滇南风吹楠在全光条件下的最大光合速率反比25%光照条件下的低。全光条件下两个树种光系统II的最大光化学效率（Fv/Fm）都显著降低,表明发生了长期光抑制。当把生长于遮荫条件下的幼苗移到全光下,从凌晨到中午随着光强的增加光抑制加剧,日落时生长于8%光照条件下的绒毛番龙眼及生长于8%和25%光照条件下的滇南风吹楠的光抑制不能完全恢复。非光化学猝灭对光强的响应曲线表明,随着生长光强的增加滇南风吹楠的热耗散能力增强,而生长在全光和25%光照条件下的绒毛番龙眼的热耗散能力都比滇南风吹楠的弱。两个树种叶片中O-[]·2、H2O2含量、SOD和CAT活性均随着生长光强的增加而提高;在同一光照条件下,绒毛番龙眼叶片中O-[]·2、H2O2含量、SOD和CAT活性显著高于滇南风吹楠。上述结果表明,在光抑制条件下,冠层树种绒毛番龙眼较大程度通过提高保护性酶的活性来保护光合机构免受损伤,而中层树种滇南风吹楠却较大程度通过增强非光化学猝灭来耗散过量光能;滇南风吹楠对强光的适应性差。     

6种热带雨林木本植物幼苗光合诱导的研究

 在晴天上午适宜条件下,测定了生长在模拟林下光环境中的6种热带雨林木本植物幼苗的光合特性和光合诱导特征。6种植物分别为先锋树种大穗野桐(Mallotus macrostachys),冠层树种绒毛番龙眼(Pometia tomentosa)、玉蕊(Barringtonia pendala)、望天树(Shorea chinensis)、滇南插柚紫(Linociera insignis)和林下灌木睫毛粗叶木(Lasianthus hookeri)。研究结果表明：暗处理3 h的叶片经连续饱和强光照射后,6种植物的净光合速率呈s形到双曲线形。6种植物达到90%最大净光合速率的时间为4.4～12.5 min,这与所报道的其它热带雨林中一些阴生植物的诱导速率相近。大穗野桐和睫毛粗叶木的诱导速率最快,达到50%和90%最大净光合速率的时间为其它4种冠层植物幼苗的1/2至1/3。诱导过程中,最大气孔导度对强光的响应明显滞后于净光合速率。充分诱导的叶片在黑暗中20 min后,6种植物的诱导状态都较高。其中,大穗野桐的诱导状态消失相对较快,这可能与其气孔导度和羧化能力的快速降低有关。玉蕊诱导状态的消失主要与生化限制有关,因为此时它的气孔导度仍维持相对较高的值。而睫毛粗叶木较高的气孔导度和羧化能力的维持导致了很高的诱导状态。林下植物这种对强光的快速反应和黑暗中高的维持状态对有效利用光斑具有重要的意义,这与其一生中在林下生长和更新的特点是一致的。     

热带雨林冠层树种绒毛番龙眼两种发育阶段叶片的光抑制

 通过测定西双版纳热带雨林冠层树种绒毛番龙眼(Pometia tomentosa)完全伸展嫩叶和成熟叶的叶片解剖、生理特征和雨季晴天自然条件下叶绿素a荧光以及午间强光对部分保护酶活性和膜脂过氧化作用的影响,探讨了两种不同发育阶段叶片光合作用的光抑制与强光和温度的关系。结果表明：绒毛番龙眼全展嫩叶和成熟叶表现出明显的解剖和生理特征差异。与全展嫩叶相比,成熟叶的叶片较厚、叶绿素含量高、气孔导度大、羧化效率高、最大净光合速率和光饱和点高,而气孔密度和保卫细胞长度没有显著差别。在雨季晴天自然条件下,午间最高光强可达2 200 μmol·m-2·s-1以上,最高叶温比气温高7～8 ℃,而成熟叶片的最高温度比全展嫩叶高1.5～2 ℃。上午随光强的增大,两种叶片的非光化学猝灭系数（NPQ）增大,PSⅡ原初光化学效率(Fv/Fm)、实际光化学效率［(Fm′_Fs)/Fm′］逐渐减小,在15∶30左右达最小。下午随着光强的减弱,Fv/Fm逐渐恢复,在傍晚基本恢复到清晨值。初始荧光(F0)在一天中变化很小。这表明绒毛番龙眼叶片光抑制是非辐射能量耗散增加引起的保护光合机构免受光破坏的保护性反应,而非光破坏。全展嫩叶比成熟叶有较低的光化学效率和非辐射耗散能力,对强光和高温处理的敏感性也较强,但在自然条件下一天中的光抑制程度与成熟叶没有显著差别。田间午间强光导致两种叶片的保护酶活性(超氧化物歧化酶,SOD;抗坏血酸过氧化物酶,APX)升高,而H2O2含量变化较小。其中,全展嫩叶的保护酶活性高,丙二醛(MDA)含量低。这表明自然条件下,与成熟叶相比,绒毛番龙眼全展嫩叶通过较低的光能利用效率、较低的叶温和高的保护酶活性减轻了强光高温的光抑制程度。     

夜间低温对不同光强下生长的两种热带树苗光合作用的影响

于雾凉季测定了叶片叶绿素荧光和气体交换参数、H2 O2 和丙二醛 (MDA)含量 ,探讨了 4～ 6℃夜间低温对 3种相对光强 (太阳光的 8%、2 5 %和 5 0 % )下生长的 2种山地雨林树苗光合作用的影响。夜间低温处理导致 3种光强下生长的 2种植物叶片日间和胁迫诱导的光抑制加剧 ,净光合速率 (Pn)和光呼吸速率降低 ,H2 O2 和MDA含量升高。生长环境光强升高可加剧夜间低温的效应 ,弱光下 2种植物受夜间低温影响较小 ,间接表明雾使光强减弱可缓解自然夜温降低对西双版纳热带植物的危害。处理期间滇南插柚紫Pn 主要受气孔限制 ,滇南红厚壳Pn 降低与活性氧增加引起的羧化效率降低有关 ,两种植物Pn 的降低均与光抑制关系不大。夜间低温后滇南红厚壳气孔导度降低较少 ,H2 O2 和MDA含量较高 ,光抑制较重 ,5 0 %相对光强下其未做夜间低温处理的对照植株发生了胁迫诱导的光抑制 ,表明它对夜间低温较敏感。     

夜间低温对2种热带雨林树种幼苗叶绿素荧光的影响

研究了夜间低温(5℃±1℃,12h,连续3个晚上)对生长于3种光照条件下(100%、25%和8%光照)的热带雨林冠层树种绒毛番龙眼(Pometiatomentosa)和中层树种滇南风吹楠(Horsfieldiatetratepala)幼苗叶绿素荧光的影响。结果表明:低温处理使生长于全光下的绒毛番龙眼幼苗的Fv/Fm(PSII最大光化学量子产量)急剧降低,中午发生了强烈的光抑制,但随着低温胁迫的解除Fv/Fm能很快恢复,表明没有发生不可逆的光氧化损伤。低温使生长于全光和25%光照条件下2个树种幼苗的NPQ(非光化学猝灭,热耗散)受到抑制,但没有引起2个树种幼苗Fo(初始荧光)的升高,不会导致幼苗PSII反应中心的失活。2种热带雨林植物对低温的抗性可能与这些植物的地理分布和历史渊源有关。     

生长环境光强对两种热带雨林树种幼苗光合作用的影响

以西双版纳热带雨林中演替后期种绒毛番龙眼和先锋树种山黄麻为材料 ,于雾凉季测定了不同光强下生长的 2种树苗叶片最大净光合速率 (Pmax)、叶绿素荧光参数以及光合色素含量和比叶重 (L MA) ,探讨了不同生态习性热带雨林树种幼苗对光强的适应及光保护机制。发现在一定光强范围内随生长环境光强的增加 ,2种树苗 L MA、荧光的非化学猝灭 (N PQ)、类胡萝卜素(Car)含量、Car与叶绿素 (Chl)之比升高 ,光饱和点和光补偿点也有随生长环境光强的增大而升高的趋势 ,Chl含量降低 ,2种树苗均能通过形态和生理特性的变化适应不同的光强环境。相同的生长光强下 ,绒毛番龙眼光抑制明显比山黄麻重 ,山黄麻适应强光的能力强。随生长环境光强的增加 ,山黄麻 N PQ增加不显著 ,热耗散较少 ,相同光强下其 Pmax显著高于绒毛番龙眼。绒毛番龙眼则相反 ,其热耗散随生长环境光强的升高显著增多 ,但 Pmax差异不显著。表明先锋种山黄麻主要通过提高 Pmax利用光能防止光合机构光破坏 ,而演替后期种绒毛番龙眼却较大程度通过增强非光化学猝灭来耗散过量光能。上午人为降低光强度对先锋种山黄麻影响不大 ,但可以明显缓解绒毛番龙眼的光抑制 ,表明上午一定程度的遮光 (如有雾 )可减缓绒毛番龙眼光抑制     

绒毛番龙眼对生长光强的形态和生理适应

在100％、50％、25％和8％自然光强下栽培绒毛番龙眼幼苗并研究了其对光环境的适应。100％生长光强下绒毛番龙眼通过增大叶片悬挂角(midrib angle,MA)和比叶重(lamina mass per unit area,LMA),减少叶氮在捕光组分中的分配等降低光能捕获;通过增加类胡萝卜素含量增加热耗散。虽如此,还是发生了比较严重的光抑制,加之叶氮在光合机构中的分配最少,导致光合能力最低,长势最差。8％生长光强下绒毛番龙眼通过降低MA、LMA以及叶片技转,增加叶氮在捕光组分中的分配等提高光能捕获能力,光能转换及利用效率较高,热耗散水平较低,但由于环境光较弱,限制了光合碳同化,植株生长也较慢。50％和25％生长光强下绒毛番龙眼有较强的光能捕获、利用和耗散能力,在几种光处理中长势最好。     

Stomatal frequency of Quercus myrsinaefolia grown under different irradiances

Stomatal and epidermal cell frequencies and leaf area were measured in leaves of Quercus myrsinaefolia grown in the field under different relative photon flux density (PFD), which was the ratio of integrated PFD at the leaf surface to that at an open site. Leaf area showed a linear relationship with the relative PFD. Stomatal and epidermal cell frequencies increased with increasing relative PFD. Numbers of stomata and epidermal cells per leaf, and stomatal index (ratio of stomatal number to epidermal cell number) increased with increasing relative PFD.     

Stomatal frequency of Quercus myrsinaefolia grown under different irradiances

Stomatal and epidermal cell frequencies and leaf area were measured in leaves of Quercus myrsinaefolia grown in the field under different relative photon flux density (PFD), which was the ratio of integrated PFD at the leaf surface to that at an open site. Leaf area showed a linear relationship with the relative PFD. Stomatal and epidermal cell frequencies increased with increasing relative PFD. Numbers of stomata and epidermal cells per leaf, and stomatal index (ratio of stomatal number to epidermal cell number) increased with increasing relative PFD. This revised version was published online in June 2006 with corrections to the Cover Date.     

Stomatal control of transpiration from a developing sugarcane canopy

Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.     

Changes in stomatal frequency, stomatal conductance and cuticle thickness during leaf expansion in the broad-leaved evergreen species, Eucalyptus regnans

Aspects of leaf anatomical and physiological development were investigated in the broad-leaved evergreen species, Eucalyptus regnans F.Muell. Newly emergent leaves were tagged in the field and measured for stomatal conductance while a subset was collected every 14 days for the measurement of stomata and cuticle over a 113-day period. Cuticle thickness increased during leaf expansion, the increase following a sigmoid curve. Stomatal frequency (no. mm⁻²) decreased from 56 to 113 days after leaf emergence. The frequency of both immature and intermediate developmental stages of stomata also decreased over this time, but the total number of stomata per leaf remained relatively constant. Stomatal conductance (g _s) of young expanding leaves increased during expansion, and was significantly linearly correlated with stomatal frequency (excluding immature stomata), and with cuticle thickness. The progressive increase in g _s in young developing leaves was contrary to the observed changes in structural characteristics (increased cuticle thickness and decreased stomatal frequency). This increase in g _s with development may be related to the progressive increase in number of mature stomata with larger apertures and, therefore, a higher total pore area in fully expanded leaves.     

Effects of humidity on light-induced stomatal opening: evidence for hydraulic coupling among stomata

A mechanism for co-ordinating behaviour of stomata within an areole during patchy stomatal conductance has recently been proposed. This mechanism depends on hydraulic interactions among stomata that are mediated by transpiration-induced changes in epidermal turgor. One testable prediction that arises from this proposed mechanism is that the strength of hydraulic coupling among stomata should be proportional to evaporative demand and, therefore, inversely proportional to humidity. When a leaf is illuminated following a period of darkness, there is typically a period of time, termed the Spannungsphase, during which guard cell osmotic and turgor pressure are increasing, but the pore remains closed. If hydraulic coupling is proportional to evaporative demand, then variation among stomata in the duration of the Spannungsphase should be lower for leaves at low humidity than for leaves at high humidity. A similar prediction emerged from a computer model based on the proposed hydraulic mechanisms. These predictions were tested by measuring individual stomatal apertures on intact transpiring leaves at low and high humidity and on vacuum-infiltrated leaf pieces (to eliminate transpiration) as PFD was increased to high values from either darkness or a low value. Results showed that the range of Spannungsphasenamong stomata was reduced at low humidity compared to high humidities. Experiments that began at low PFD, rather than at darkness, showed no delay in stomatal opening. These results are discussed in the context of the proposed hydraulic coupling mechanisms.     

A New Porometer Based upon the Electrical Current Produced by Guard Cells

Stomatal guard cells extrude protons when the stomata open.This gives rise to an electrical current which is proportionalto the degree of stomatal opening. An instrument has been developedto measure this leaf surface current which is, in effect, anew type of porometer. The performance of the new porometerhas been compared with that of a commercially available diffusionporometer and a close relationship between leaf surface currentand stomatal conductance was observed for all the species investigated.It is concluded that the instrument has several advantages overthe diffusion porometer, in particular, its small size and simplicityof operation, making it especially suitable for use in the field. Key words: Leaves, stomata, electrical currents, porometry     

Elevated carbon dioxide affects the patterning of subsidiary cells in Tradescantia stomatal complexes

The influence of elevated CO₂ concentration (670 ppm) on thestructure, distribution, and patterning of stomata in Tradescantialeaves was studied by making comparisons with plants grown atambient CO₂. Extra subsidiary cells, beyond the normal complementof four per stoma, were associated with nearly half the stomatalcomplexes on leaves grown in elevated CO₂. The extra cells sharedcharacteristics, such as pigmentation and expansion, with thetypical subsidiary cells. The position and shape of the extrasubsidiary cells in face view differed in the green and purplevarieties of Tradescantia. Substomatal cavities of complexeswith extra subsidiary cells appeared larger than those foundin control leaves. Stomatal frequency expressed on the basisof leaf area did not differ from the control. Stomatal frequencybased on cell counts (stomatal index) was greater in leavesgrown in CO₂-enriched air when all subsidiary cells were countedas part of the stomatal complex. This difference was eliminatedwhen subsidiary cells were included in the count of epidermalcells, thereby evaluating the frequency of guard cell pairs.The extra subsidiary cells were, therefore, recruited from theepidermal cell population during development. Stomatal frequencyin plants grown at elevated temperature (29 C) was not significantlydifferent from that of the control (24 C). The linear aggregationsof stomata were similar in plants grown in ambient and elevatedCO₂. Since enriched CO₂ had no effect on the structure or patterningof guard cells, but resulted in the formation of additionalsubsidiary cells, it is likely that separate and independentevents pattern the two cell types. Plants grown at enrichedCO₂ levels had significantly greater internode lengths, butleaf area and the time interval between the appearance of successiveleaves were similar to that of control plants. Porometric measurementsrevealed that stomatal conductance of plants grown under elevatedCO₂ was lower than that of control leaves and those grown atelevated temperature. Tradescantia was capable of regulatingstomatal conductance in response to elevated CO₂ without changingthe relative number of stomata present on the leaf. Key words: Elevated CO₂, stomata, subsidiary cells, patterning     

How strong is intracanopy leaf plasticity in temperate deciduous trees?

Intracanopy plasticity in tree leaf form is a major determinant of whole-plant function and potentially of forest understory ecology. However, there exists little systematic information for the full extent of intracanopy plasticity, whether it is linked with height and exposure, or its variation across species. For arboretum-grown trees of six temperate deciduous species averaging 13-18 m in height, we quantified intracanopy plasticity for 11 leaf traits across three canopy locations (basal-interior, basal-exterior, and top). Plasticity was pronounced across the canopy, and maximum likelihood analyses indicated that plasticity was primarily linked with irradiance, regardless of height. Intracanopy plasticity (the quotient of values for top and basal-interior leaves) was often similar across species and statistically indistinguishable across species for several key traits. At canopy tops, the area of individual leaves was on average 0.5-0.6 times that at basal-interior, stomatal density 1.1-1.5 times higher, sapwood cross-sectional area up to 1.7 times higher, and leaf mass per area 1.5-2.2 times higher; guard cell and stomatal pore lengths were invariant across the canopy. Species differed in intracanopy plasticity for the mass of individual leaves, leaf margin dissection, ratio of leaf to sapwood areas, and stomatal pore area per leaf area; plasticity quotients ranged only up to ≈2. Across the six species, trait plasticities were uncorrelated and independent of the magnitude of the canopy gradient in irradiance or height and of the species' light requirements for regeneration. This convergence across species indicates general optimization or constraints in development, resulting in a bounded plasticity that improves canopy performance.