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。     

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     

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。     

不同光强下生长的两种榕树叶片光合能力与比叶重、氮含量及分配的关系

观测了不同光强下生长的两种榕树幼苗叶片的光合能力、比叶重(LMA)、氮含量及在光合机构中的分配.结果表明,两种榕树幼苗的最大净光合速率(Pmax)均随生长光强的升高而升高,这与LMA、单位面积氮含量(NA)和光合氮利用效率随生长光强的升高而升高有关.除36%光强下外,相同光强下生长的喜光的斜叶榕(Ficus tinctoria)的Pmax均显著高于耐荫的假斜叶榕(Ficus subulata),这与其叶片中氮在羧化组份和生物力能学组份中的分配系数、LMA和NA较高有关.     

喜光榕树和耐荫榕树光适应机制的差异

100%和36%光强下生长的喜光的斜叶榕的光合能力高于耐荫的假斜叶榕,而热耗散能力与之相似,说明强光下斜叶榕主要通过光合作用利用光能和热耗散、假斜叶榕主要通过热耗散防御光破坏.100%光强下生长的两种榕树的日间光抑制程度相似,但叶表光强相同情况下各光强下生长的假斜叶榕的光抑制均比斜叶榕严重.100%光强下假斜叶榕叶片悬挂角大于斜叶榕,导致日间叶表光强低于斜叶榕,这可能是两种榕树日间光抑制程度相似的原因,表明叶片悬挂角的适应变化对假斜叶榕有重要的意义.     

氮素对红波罗花光合作用和生长的影响

以三年生红波罗花为试验材料,比较了不同氮处理下植株的叶片性状、气体交换特性、叶氮分配、叶绿素荧光和生长参数,探讨了在高山环境中氮素对红波罗花光合特性和生长的影响.结果表明:不施氮时,虽然红波罗花减少了叶氮在捕光复合体中的分配,但还是出现了光抑制现象,并且由于叶氮含量(N)、叶绿素含量(Chl)和叶氮在光合组分中的分配量是最少的,导致光合能力最低,生长最缓.随外界施氮量的增加,红波罗花的N,Chl,叶肉厚度、叶氮在光合组分中的分配量(NR、NB、NL),光能利用效率均随之增加,光合能力相应增强.三个施氮处理中,红波罗花分配给生物力能学组分的叶氮(NB)多于分配给Rubisco的(NR),从而导致Jmax/Vcmax增加,有利于其适应高山低温和高光强.施氮量增加,植株的相对生长速率(RGR)增加,这对于红波罗花当年的开花繁殖,以及来年的生长发育都非常有利.当施氮量超过0.2g/kg基质时,植株的N,Chl,叶肉组织厚度、光合速率以及RGR都不再显著增加,因此当土壤中纯氮含量为220～230mg/kg时,即能满足红波罗花较佳光合和生长所需.     

不同光强下焕镛木和观光木的光合参数变化

 生长在全日光强下的焕镛木(Woonyoungia septentrionalis)和观光木(Tsoongiodendron lotungensis)幼树叶片的最大光合速率、表观量子产率和光能转换效率均较生长在40%和20%日光强的高。当生长光强从全日光强降低至40%日光强时,焕镛木的表观量子产率和光能转换效率分别降低13.1%和6.3%,而观光木则相应分别降低23.8%和33.4%。生长光强降低至40%日光强时,焕镛木的Rubisco最大羧化速率(Vcmax)未见变化;而最大电子传递速率（Jmax）则降低14.1%,表明Jmax对光强降低的响应较Vcmax敏感。当生长光强从全日光强降低到40%和20%日光强时,观光木的Vcmax分别降低7.7%和31.7%,而Jmax则分别降低9.7%和42%。光强从全日光强降低至40%日光强,焕镛木叶氮在Rubisco和捕光叶绿素蛋白复合体中的分配系数没有明显改变,而叶氮在生物力能学组分中的分配系数降低则较为明显（20.4%）,表明生长光强降低对叶氮在光合电子传递链组分分配的影响较在Rubisco的大。结果表明,焕镛木表现阳生树种特性,在迁地保育中宜选择向阳小生境种植,而观光木较耐荫,可种植在较遮荫的环境。     

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     

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。     

西双版纳两种生态特性不同的外来草本植物对生长环境光强的适应策略

于干热季测定了紫茎泽兰和阳春砂仁叶片比叶重、色素含量、光合能力和叶绿素荧光动力学参数,探讨了它们适应环境光强的策略及其生理生态学机制。36 %光强下砂仁叶片光抑制较明显;10 0 %光强下紫茎泽兰光抑制不严重,光系统 最大光能转换效率和量子效率始终维持在较高水平。随着环境光强的升高,紫茎泽兰最大净光合速率(Pmax)、比叶重、非光化学猝灭系数(N PQ)和单位面积叶片类胡萝卜素含量升高,单位干重叶片叶绿素含量降低。紫茎泽兰能通过形态和生理特性的变化适应大幅度的光强范围,这可能是其表现强入侵性的重要原因之一。强光下紫茎泽兰热耗散并不多,主要通过提高Pmax利用更多的光能来保护光合机构,光系统 反应中心可逆失活也能耗散部分光能。虽然砂仁Pmax也能随生长环境光强的升高而增大,但其值较低,增幅不大,相反其热耗散的增加较多,N PQ较高,白天初始荧光明显低于黎明,表明砂仁主要是通过热耗散来保护光合机构。     

Photosynthetic performance along a light gradient as related to leaf characteristics of a naturally occurring <Emphasis Type="Italic">Cypripedium flavum</Emphasis>

Photosynthesis, leaf structure, nitrogen content and nitrogen allocation in photosynthetic functions of Cypripedium flavum were studied in a naturally varying light regime. Light-saturated leaf net photosynthetic rate (A (max)) was strongly correlated with leaf dry mass per area (LMA), mesophyll conductance (g (m)) and area-based leaf nitrogen content (N(area)), with all variables increasing with increasing irradiance. Such coordinate variation of all these parameters illustrates the plastic response of leaf structure to high light (HL). Leaf N(area) was greater under HL than in low light (LL). The fractions of leaf nitrogen partitioning in carboxylation (P (R)) and bioenergetics (P (B)) were positively related to LMA. In contrast, P (R) and P (B) decreased with increasing mass-based leaf nitrogen content (N(mass)). However, no correlation was found between leaf nitrogen investment in light harvesting (P (L)) and either LMA or N(mass). Like maximum rate of carboxylation (V (cmax)) and electron transport (J (max)), the J (max)/V (cmax) ratio, which was strongly correlated to LMA, also increased significantly with irradiance. Under HL, leaf maximum photosynthetic nitrogen efficiency (ANUE) and intrinsic water use efficiency (WUE) were greater than in LL conditions, despite a small difference in WUE. This suggests that a functional balance in the photosynthetic machinery favors leaf photosynthetic plasticity of C. flavum in response to different light conditions. Given an ample soil nitrogen supply, C. flavum may offset its susceptibility to HL by efficient nitrogen use and higher stomatal and mesophyll conductance against photoinhibition so as to keep leaf photosynthesis positive.     

热带雨林蒲桃属3个树种的幼苗光合作用对生长光强的适应

 测定了生长于3种光强下（100％、42％和14％自然光强） 热带雨林演替早期（思茅蒲桃Syzygium latilimbum）、中期（乌墨S.cumini）和后期（阔叶蒲桃S. szemaoense）出现的蒲桃属 （Syzygium）3个树种幼苗叶片气体交换参数和叶绿素荧光参数的日变化、比叶重和叶绿素含量。发现演替早期树种光合能力和光合可塑性最大、中期树种次之,后期树种光合能力最弱且在强光下受到显著抑制。但是,生长在强光下,3个树种均未发生长期光抑制和光破坏。随生长光强增加,3个树种热耗散速率都升高。不过在强光下,中期和后期树种通过热耗散消耗的过剩光能较多,通过叶黄素循环的热耗散可能对于这两个树种在强光下避免光合机构的光破坏起到了重要作用。14％光强下,3个树种最大净光合速率、光合作用的光饱和点、光补偿点、暗呼吸速率、比叶重降低,叶片单位干重叶绿素含量提高,对低光环境有了较好的形态学和生理学适应。但是,在4％光强下思茅蒲桃和乌墨幼苗全部死亡,只有阔叶蒲桃幼苗仍能存活,说明阔叶蒲桃幼苗适应弱光环境的能力高于思茅蒲桃和乌墨,与它们的演替状态一致。     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

Spatial distribution of leaf nitrogen and photosynthetic capacity within the foliage of individual trees: disentangling the effects of local light quality,leaf irradiance,and transpiration

There is presently no consensus about the factor(s) driving photosynthetic acclimation and the intra-canopy distribution of leaf characteristics under natural conditions. The impact was tested of local (i) light quality (red/far red ratio), (ii) leaf irradiance (PPFD(i)), and (iii) transpiration rate (E) on total non-structural carbohydrates per leaf area (TNC(a)), TNC-free leaf mass-to-area ratio (LMA), total leaf nitrogen per leaf area (N(a)), photosynthetic capacity (maximum carboxylation rate and light-saturated electron transport rate), and leaf N partitioning between carboxylation and bioenergetics within the foliage of young walnut trees grown outdoors. Light environment (quantity and quality) was controlled by placing individual branches under neutral or green screens during spring growth, and air vapour pressure deficit (VPD) was prescribed and leaf transpiration and photosynthesis measured at branch level by a branch bag technique. Under similar levels of leaf irradiance, low air vapour pressure deficit decreased transpiration rate but did not influence leaf characteristics. Close linear relationships were detected between leaf irradiance and leaf N(a), LMA or photosynthetic capacity, and low R/FR ratio decreased leaf N(a), LMA and photosynthetic capacity. Irradiance and R/FR also influenced the partitioning of leaf nitrogen into carboxylation and electron light transport. Thus, local light level and quality are the major factors driving photosynthetic acclimation and intra-canopy distribution of leaf characteristics, whereas local transpiration rate is of less importance.     

Interspecific prediction of photosynthetic light response curves using specific leaf mass and leaf nitrogen content: effects of differences in soil fertility and growth irradiance

Background and Aims

Previous work has shown that the entire photosynthetic light response curve, based on both Mitscherlich and Michaelis–Menten functions, could be predicted in an interspecific context through allometric relations linking the parameters of these functions to two static leaf traits: leaf nitrogen (N) content and leaf mass per area (LMA). This paper describes to what extent these allometric relations are robust to changes in soil fertility and the growth irradiance of the plants.

Methods

Plants of 25 herbaceous species were grown under controlled conditions in factorial combinations of low/high soil fertility and low/high growth irradiance. Net photosynthetic rates per unit dry mass were measured at light intensities ranging from 0 to 700 µmol m⁻² s⁻¹ photosynthetically active radiation (PAR).

Key Results

The differing growth environments induced large changes in N, LMA and in each of the parameter estimates of the Mitscherlich and Michaelis–Menten functions. However, the differing growth environments induced only small (although significant) changes in the allometric relationships linking N and LMA to the parameters of the two functions. As a result, 88 % (Mitcherlich) and 89 % (Michaelis–Menten) of the observed net photosynthetic rates over the full range of light intensities (0–700 µmol m⁻² s⁻¹ PAR) and across all four growth environments could be predicted using only N and LMA using the same allometric relations.

Conclusions

These results suggest the possibility of predicting net photosynthetic rates in nature across species over the full range of light intensities using readily available data.     

Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field

Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.