Acclimation of photosynthetic characteristics of the moss Pleurozium schreberi to among‐habitat and within‐canopy light gradients

Light availability varies strongly among moss habitats and within the moss canopy, and vertical variation in light within the canopy further interacts with the age gradient. The interacting controls by habitat and canopy light gradient and senescence have not been studied extensively. We measured light profiles, chlorophyll (Chl), carotenoid (Car) and nitrogen (N) concentrations, and photosynthetic electron transport capacity (J_max) along habitat and canopy light gradients in the widespread, temperate moss Pleurozium schreberi to separate sources of variation in moss chemical and physiological traits. We hypothesised that this species, like typical feather mosses with both apical and lateral growth, exhibits greater plasticity in the canopy than between habitats due to deeper within‐canopy light gradients. For the among‐habitat light gradient, Chl, Chl/N and Chl/Car ratio increased with decreasing light availability, indicating enhanced light harvesting in lower light and higher capacity for photoprotection in higher light. N and J_max were independent of habitat light availability. Within the upper canopy, until 50–60% above‐canopy light, changes in moss chemistry and photosynthetic characteristics were analogous to patterns observed for the between‐habitat light gradient. In contrast, deeper canopy layers reflected senescence of moss shoots, with pigment and nitrogen concentrations and photosynthetic capacity decreasing with light availability. Thus, variation in chemical and physiological traits within the moss canopy is a balance between acclimation and senescence. This study demonstrates extensive light‐dependent variation in moss photosynthetic traits, but also that between‐habitat and within‐canopy light gradient affects moss physiology and chemistry differently.     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

太岳山典型阔叶乔木冠层叶片性状的分布格局

以太岳山4种阔叶乔木不同冠层高度的叶片为研究对象,用LI-3000A叶面积仪和Li-6400便携式光合作用测定系统分别测定了这4种乔木不同冠层高度叶片的叶面积大小和单位面积的叶光饱和速率(Aarea);同时测定了其叶氮含量;计算了其比叶面积(SLA)、单位面积叶氮含量(Narea)、单位重量叶氮含量(Nmass)、单位重量的叶光饱和速率(Amass)和光合氮素利用效率(PNUE),对植株不同冠层高度叶片的SLA、叶氮和光合特性的空间分布格局进行了比较研究,结果表明:Aarea、Amass、Nmass、PNUE、SLA和Narea在树冠上层、中层和下层的差异均达到了极显著水平(P<0.001),表明树冠不同高度的叶片性状参数差异较大;在相同SLA下,Nmass和Narea在冠层中的分布均表现为中层>上层>下层,并出现平行位移现象;Aarea和Nmass都以中层值最大,表明冠层光合能力分布格局以中层相对较高。     

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.     

Divergent patterns of photosynthetic phosphorus-use efficiency versus nitrogen-use efficiency of tree leaves along nutrient-availability gradients

1.  Nitrogen (N) and phosphorus (P) are essential nutrients for photosynthetic carbon assimilation and most frequently limit primary productivity in terrestrial ecosystems. Efficient use of those nutrients is important for plants growing in nutrient-poor environments.
2.  We investigated the pattern of photosynthetic phosphorus-use efficiency (PPUE) in comparison with photosynthetic nitrogen-use efficiency (PNUE) along gradients of P and N availability across biomes with 340 tree and shrub species. We used both total soil N and P concentration and foliar N/P ratios for indicating nutrient-availability gradients.
3.  Photosynthetic phosphorus-use efficiency increased with greater leaf mass per area (LMA) toward decreasing P availability. By contrast, PNUE decreased with greater LMA towards decreasing N and P availability.
4.  The increase in PPUE with decreasing P availability was caused by the net effects of a relatively greater reduction in foliar P concentration and a relatively constant photosynthetic carbon assimilation rate. The decrease in PNUE with decreasing N availability was caused by the effects of a reduction in photosynthetic carbon assimilation rate with greater LMA.
5. Synthesis . Our results suggest that higher PPUE may be an effective leaf-level adaptation to P-poor soils, especially in tropical tree species. Future research should focus on the difference between PPUE and PNUE in relation to leaf economics, physiology and strategy.     

Acclimation of plants to light gradients in leaf canopies: evidence for a possible role for cytokinins transported in the transpiration stream.

The mechanism of response of plants to vertical light intensity gradients in leaf canopies was investigated. Since shaded leaves transpire less than leaves in high light, it was hypothesized that cytokinins (CKs) carried by mass transport in the transpiration stream would be distributed over the leaf area of partially shaded plants parallel to the gradient in light intensity. It was also hypothesized that this causes the distribution of leaf growth, leaf N and photosynthetic capacity, and possibly chloroplast acclimation as observed in plants growing in leaf canopies. In a field experiment, the distribution of Ca, N and CKs in a bean leaf canopy of a dense and an open stand supported the concept of a role for CKs in the response of N allocation to the light gradient when a decreasing sensitivity for CKs with increasing leaf age is assumed. Both shading of one leaf of the pair of primary bean leaves and independent reduction of its transpiration rate in a growth cabinet experiment caused lower dry mass, N and Ca per unit leaf area in comparison to the opposite not treated leaf. Shading caused a parallel reduction in CK concentration, which supports the hypothesis, but independent reduction of transpiration rate failed to do the same. Application of benzylaminopurine (BA) counteracted the reduction caused by shade of leaf N, photosynthetic capacity and leaf area growth. The experiments show an important role for the transpiration stream in the response of plants to light gradients. Evidence is presented here that CKs carried in the transpiration stream may be important mediators for the acclimation of plants to leaf canopy density.     

Vertical gradients in photosynthetic physiology diverge at the latitudinal range extremes of white spruce

Light availability drives vertical canopy gradients in photosynthetic functioning and carbon (C) balance, yet patterns of variability in these gradients remain unclear. We measured light availability, photosynthetic CO₂ and light response curves, foliar C, nitrogen (N) and pigment concentrations, and the photochemical reflectance index (PRI) on upper and lower canopy needles of white spruce trees (Picea glauca) at the species' northern and southern range extremes. We combined our photosynthetic data with previously published respiratory data to compare and contrast canopy C balance between latitudinal extremes. We found steep canopy gradients in irradiance, photosynthesis and leaf traits at the southern range limit, but a lack of variation across canopy positions at the northern range limit. Thus, unlike many tree species from tropical to mid-latitude forests, high latitude trees may not require vertical gradients of metabolic activity to optimize photosynthetic C gain. Consequently, accounting for self-shading is less critical for predicting gross primary productivity at northern relative to southern latitudes. Northern trees also had a significantly smaller net positive leaf C balance than southern trees suggesting that, regardless of canopy position, low photosynthetic rates coupled with high respiratory costs may ultimately constrain the northern range limit of this widely distributed boreal species.     

栲树冠层光合生理特性的空间异质性

森林冠层在能量传输、光合有效辐射和微气象等方面的差异可导致冠层光合生产力空间分布的变化.叶片光合生理特性在空间上的差异对精确估算森林冠层的初级生产力十分重要.本文以亚热带常绿阔叶林优势种---栲树(Castanopsis fargesii)为对象,研究叶片光合生理特性在冠层空间上的变化.结果表明:1)在垂直方向上,冠层北向叶的饱和光合速率(Amax)、光饱和点(LSP)和CO2羧化效率(CCE)均表现为上部>中部>底部,且依次平均降低19.4%、18.1%和37.1%;光补偿点(LCP)、光下暗呼吸(Rd)以及冠层南向叶的饱和光合速率、光饱和点和CO2羧化效率均表现为上部>底部>中部,上部比中部和底部高出12.3%~71.4%;表观量子效率(AQY)表现为底部>上部>中部,底部分别是顶部和中部叶的1.2和1.3倍;2)在水平方向上,冠层上部和底部南向叶的饱和光合速率、光饱和点和CO2羧化效率比北向叶高0.9%~31.5%;冠层中部北向叶的饱和光合速率等6个参数比南向叶高9.6%~63.2%.因此,在冠层水平上模拟和估算植物生产力时,必须考虑冠层光合生理特性的空间差异.     

黄土高原油松和刺槐叶片光合生理适应性比较

以黄土高原地区由南向北分布的杨凌、永寿、富县、安塞、米脂、神木等县为研究地点，研究不同地区油松和刺槐的光合特性与叶结构性状间的关系.结果表明，不同地区油松针叶和刺槐叶片的净光合速率（P_n）、光合氮利用效率（PNUE）、水分利用效率（WUE）、比叶质量（LMA）、氮含量（N_mass）和叶绿素相对含量（Chl）差异均达极显著水平（P＜0.001）,说明不同地区油松和刺槐的光合能力和叶结构性状参数差异很大.由南向北，油松的P_n、WUE和PNUE呈略微增加趋势，而刺槐则呈显著降低趋势，表明油松在干旱生境下仍能维持较高的光合能力，而刺槐光合能力明显受到抑制；油松和刺槐的LMA均呈略微上升趋势，而N_mass和Chl均呈略微下降趋势，且刺槐的变化幅度高于油松，说明油松从生理代谢和叶结构性状上对干旱环境的适应能力均强于刺槐.相关分析表明，不同地区油松和刺槐的LMA与N_mass整体上呈极显著负相关；P_n、PNUE与LMA、N_mass相关不显著，与Chl呈极显著正相关；WUE与LMA呈显著负相关（P＜0.05），与N_mass呈显著正相关.     

Plant biodiversity in a larch plantation from the view point of photosynthetic nitrogen use efficiency in northeast China

Forest floor of larch species often provides growth habitat for many kinds of understory species because of relatively sparse structure in a larch canopy. A rich flora of forest understory species may play an essential role in maintaining fertility of a larch stand. An attempt was made to evaluate photosynthetic nitrogen use efficiency (PNUE) of many understory and overstory species according to their Raunkiaer lifeform. By studying 72 perennial deciduous species in a larch plantation in northeast China, marked photosynthetic differences between phanerophytes (Ph) and other three lifeforms of chamaephytes (Ch), hemicryptophytes (He), and cryptophytes (Cr) were found, with marginal differences found among Ch, He, and Cr. Ph species had much lower PNUE, and much lower values of rate of nitrogen allocation to chlorophyll (Chl./N) and nitrogen allocation to carboxylation processes (V _cmax/N) were concurrently observed in Ph compared with the other three lifeforms. Ph had much lower leaf nitrogen per unit of projection area (N _area) and specific leaf area (SLA, cm² g^–1). At lower SLA, for Ph species the change of PNUE with SLA was small, but these changes became very large at higher SLA for Ch, He, and Cr species. Our findings indicate that leaf morphological change is important for clarifying photosynthesis differences among species with different lifeform.     

Effects of altitudinal gradient on Salix atopantha foliar delta13C

In 2010, measurements were conducted on the foliar delta13C, photosynthesis, CO2 diffusive conductivity, nitrogen content, photosynthetic nitrogen use efficiency (PNUE), and special leaf area (SLA) of Salix atopantha at different altitudes (2350 m, 2700 m, 3150 m, and 3530 m) in Wolong Natural Reserve. With the increase of altitude, the foliar nitrogen content (especially the nitrogen content per unit leaf area, N(area)) and the PNUE increased, and the foliar delta13C had a significant increase, with an increment of 1.4 per thousand per 1000 m altitude. The stomatal and mesophyll CO2 diffusion conductance also increased with increasing altitude, which had definite negative effect on the increase of foliar delta13C, but the effect was not strong enough. Comparing with CO2 diffusion conductance, carboxylation capacity was a more important factor limiting the P(c)/P(a), and even, the foliar delta13C. At altitude 2350-2700 m, air temperature was the main factor affecting the allocation of nitrogen in S. atopantha photosynthetic system, whereas at altitude 2700-3530 m, light could be the main affecting factor. No significant difference was observed in the SLA at different altitudes.     

Comparative ecophysiology of leaf and canopy photosynthesis

Leaves and herbaceous leaf canopies photosynthesize efficiently although the distribution of light, the ultimate resource of photosynthesis, is very biased in these systems. As has been suggested in theoretical studies, if a photosynthetic system is organized such that every photosynthetic apparatus photosynthesizes in concert, the system as a whole has the sharpest light response curve and is most adaptive. This condition can be approached by (i) homogenization of the light environment and (ii) acclimation of the photosynthetic properties of leaves or chloroplasts to their local light environments. This review examines these two factors in the herbaceous leaf canopy and in the leaf. Changes in the inclination of leaves in the canopy and differentiation of mesophyll into palisade and spongy tissue contribute to the moderation of the light gradient. Leaf and chloroplast movements in the upper parts of these systems under high irradiances also moderate light gradients. Moreover, acclimation of leaves and chloroplasts to the local light environment is substantial. These factors increase the efficiency of photosynthesis considerably. However, the systems appear to be less efficient than the theoretical optimum. When the systems are optically dense, the light gradients may be too great for leaves or chloroplasts to acclimate. The loss of photosynthetic production attributed to the imperfect adjustment of photosynthetic apparatus to the local light environment is most apparent when the photosynthesis of the system is in the transition between the light-limited and light-saturated phases. Although acclimation of the photosynthetic apparatus and moderation of light gradients are imperfect, these markedly raise the efficiency of photosynthesis. Thus more mechanistic studies on these adaptive attributes are needed. The causes and consequences of imperfect adjustment should also be investigated.     

Canopy Light Gradient Perception by Cytokinin

We have recently identified cytokinin as an important xylem-carried signal involved in the photosynthetic acclimation of plants to light gradients in dense canopies. Lower leaves become shaded in a dense canopy and consequently have reduced transpiration rates. our measurements have shown that this results in a reduced delivery of cytokinins carried in the transpiration stream to shaded leaves, as compared to light-exposed leaves. Cytokinins are involved in the regulation of photosynthetic acclimation to the light gradient by stimulating the expression of photosynthetic enzymes in light-exposed leaves. In shaded leaves, the low delivery rate of cytokinin leads to reduced photosynthetic capacity and ultimately senescence. We show evidence for this role of cytokinin, as part of a complex of signaling pathways where other regulatory mechanisms are also involved. A model is presented depicting the regulation of photosynthetic acclimation by cytokinin delivery to leaves dependent on the irradiance they receive.Key Words: canopy light gradient, transpiration, photosynthetic acclimation, cytokinin, nitrate, systemic signaling     

Photosynthetic acclimation to simultaneous and interacting environmental stresses along natural light gradients: optimality and constraints

There is a strong natural light gradient from the top to the bottom in plant canopies and along gap-understorey continua. Leaf structure and photosynthetic capacities change close to proportionally along these gradients, leading to maximisation of whole canopy photosynthesis. However, other environmental factors also vary within the light gradients in a correlative manner. Specifically, the leaves exposed to higher irradiance suffer from more severe heat, water, and photoinhibition stresses. Research in tree canopies and across gap-understorey gradients demonstrates that plants have a large potential to acclimate to interacting environmental limitations. The optimum temperature for photosynthetic electron transport increases with increasing growth irradiance in the canopy, improving the resistance of photosynthetic apparatus to heat stress. Stomatal constraints on photosynthesis are also larger at higher irradiance because the leaves at greater evaporative demands regulate water use more efficiently. Furthermore, upper canopy leaves are more rigid and have lower leaf osmotic potentials to improve water extraction from drying soil. The current review highlights that such an array of complex interactions significantly modifies the potential and realized whole canopy photosynthetic productivity, but also that the interactive effects cannot be simply predicted as composites of additive partial environmental stresses. We hypothesize that plant photosynthetic capacities deviate from the theoretical optimum values because of the interacting stresses in plant canopies and evolutionary trade-offs between leaf- and canopy-level plastic adjustments in light capture and use.     

Ecophysiological significance of leaf traits in Cypripedium and Paphiopedilum

There is a close phylogenetic relationship between Paphiopedilum and Cypripedium, but these two genera diverge considerably in terms of their leaf traits. To understand the evolution and the ecophysiological significance of leaf traits, we investigated the leaf traits of three Paphiopedilum species and three Cypripedium species in southwestern China. Cypripedium tibeticum and C. flavum showed a significantly higher light-saturated photosynthetic rate (P(max) ), stomatal conductance (g(s) ), photosynthetic nitrogen utilization efficiency (PNUE) and specific leaf area (SLA), but lower ratio of leaf carbon to nitrogen content (C/N) and leaf construction cost (CC) than Paphiopedilum. These leaf traits of Cypripedium suggest its high resource use efficiency and high growth rate reflecting adaptation to a short growing period and abundant soil nutrients and water in alpine habitats. Conversely, the low P(max) , g(s) , PNUE, SLA and the ratio of chlorophyll a to chlorophyll b (Chl a/b), but high leaf nitrogen investment in light-harvesting component (P(L) ), CC and C/N in Paphiopedilum indicate its adaptation to a low light, nutrient-poor and limited soil water habitats in karst areas. As a sympatric species of Paphiopedilum, although C. lentiginosum retained the phylogenetic leaf traits of Cypripedium, such as high mass-based light-saturated photosynthetic rate (P(max-M) ), g(s) and PNUE, it had some similar leaf traits to those of Paphiopedilum, such as low mesophyll conductance (g(m) ) and Chl a/b, and high P(L) , which reflected an adaptation to the same habitat. Our results show that the evolution of the leaf traits of Paphiopedilum and Cypripedium are shaped by both phylogeny and environment.     

Community succession and photosynthetic physiological characteristics of pasture plants in a sub-alpine meadow in Gannan,China

 为了解甘南亚高山草甸围封地恢复演替动态, 探究围封恢复进程中植物光合生理特征的变化规律及其影响因子, 对围封试验地内5个典型群落样地进行样方调查, 测定了各群落优势种及3个共有种的光合参数和叶性状参数, 并测定了群落表层土壤(0–20 cm)的水分含量及全氮含量。结果显示: 该围封地内形成一个以草本植物→半灌木→灌木的演替序列, 群落表层土壤含水量及全氮含量随着演替的进行逐渐增加; 在演替的时间尺度上, 各演替阶段优势种光合生理特征间存在明显差异, 随着演替的进行, 群落优势种的净光合速率(A_area)、光合水分利用效率(WUE)、相对叶绿素含量(SPAD)呈降低趋势, 其叶片氮含量(N_mass)、光合氮利用效率(PNUE)、比叶面积(SLA)随演替变化没有严格一致的规律, 而更多地表现为不同植物功能型之间的差异; 从演替前期到后期, 同种植物的A_area、SPAD值逐渐降低, 非豆科植物披碱草(Elymus dahuricus)、老鹳草(Geranium wilfordii)的PNUE、WUE随演替进行呈降低趋势, 其N_mass、SLA随演替进行却呈增加趋势, 然而豆科植物紫苜蓿(Medicago sativa)由于具有固氮能力, 受养分限制不明显, 这些光合生理特征值没有随演替发生明显的变化。这些结果表明, 在恢复演替过程中, 该围封地由一个物质获得能力强的群落向物质保持能力强的群落过渡, 土壤水分含量及全氮含量是推动这种过渡发生的主要因子。掌握围封地群落演替过程中的光合生理动态对于亚高寒退化草甸恢复具有一定的理论指导意义。     

温带针阔混交林叶片性状随树冠垂直高度的变化规律

自然界中,森林植物叶片的生长随树冠高度呈现明显的垂直分布现象;然而,有关叶片性状随着树冠垂直高度增加的变化规律仍不清楚。为了更好地揭示植物叶片对光环境变化的适应策略以及对资源的利用能力,有必要深入探讨叶片性状与冠层高度的定量关系及其内在调控机制。以中国广泛分布的温带针阔混交林为对象,选取8种主要树种为研究对象(白桦、蒙古栎、水曲柳、大青杨、色木槭、千金榆、核桃楸和红松),通过测定这些物种9个冠层高度的叶片比叶面积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(N)、叶片磷含量(P)、氮磷比(N∶P)和叶绿素含量(Chl)等属性,探讨了针阔混交林叶片性状的差异以及各性状之间的相关关系,进而揭示叶片性状随树冠垂直高度的变化规律。实验结果表明:1)温带针阔混交林内优势树种的部分叶片性状在不同冠层高度之间差异显著。2)随着树冠垂直高度的增加,SLA、LDMC、N、P、N∶P和Chl呈现不同的变化趋势。其中,阔叶树种SLA随着树冠垂直高度的增加而减小;所有树种的LDMC随着树冠垂直高度的增加而增加;不同树种的N、P、N∶P和Chl随着树冠垂直高度的变化规律存在差异。3)对于温带针阔混交林冠层中,SLA与N、P、N∶P均存在显著的正相关关系,高SLA伴随着高的N、P、N∶P,表明植物通过SLA与N、P等性状的协同来提高叶片的光合作用(或对光热资源的利用效率)。本研究通过定量分析探讨温带针阔混交林叶片性状随冠层高度的变化规律,一定程度地揭示了树木对光、热和水资源竞争的适应机制,以及植物叶片的资源利用和分配策略,不仅拓展了传统性状研究的范畴,其相关研究结论也有助于树木生长模型的构建和优化。     

Increases of chlorophyll a/b ratios during acclimation of tropical woody seedlings to nitrogen limitation and high light

According to the theory of optimal nitrogen partitioning within a leaf, the chlorophyll (Chl) a/b ratio is expected to increase when leaf N content decreases. Here, we report the first empirical support for this prediction. The Chl a/b ratio increased while Chl content decreased in response to N limitation in photosynthetic cotyledons and leaves of seedlings of four tropical woody species in the Bignoniaceae. The responses of all four species were in the same direction, but differed in magnitude. For Tabebuia rosea, the species that exhibited the greatest increase in Chl a/b ratios (up to values of 5.9), detailed photosynthetic characteristics were also examined. Light and N availability were positively correlated with the light- and CO2-saturated photosynthetic O2 evolution rate, as well as with leaf carboxylation capacity (Vcmax) and electron transport rate (Vj). Severe N limitation and high light did not cause chronic photo-inhibition (i.e. no change in quantum yield or in dark-acclimated Fv/Fm). The observed change in the ratio of Vcmax to leaf N in response to N availability was consistent with likely functional reasons for change in the Chl a/b ratio. Adjustment of the Chl a/b ratio was apparently an integral feature of acclimation to high light conditions and low N availability.