干旱和湿润生境中主要优势树种叶片功能性状的比较

以云南南部的沟谷雨林(湿润生境)和干热河谷萨瓦纳(savanna)稀树灌丛(干旱生境)中共31种主要优势木本植物为材料,研究了叶片主要功能性状单位面积叶干重(LMA)、单位干重氮含量(Nmass)和单位干重最大净光合速率(Amass)在2个生境中的差异及各性状间的关系。结果表明:在物种水平上,干旱和湿润生境中各树种的LMA值分别为46.88～178.63和45.35～93.16g·m-2;Amass值分别为48.12～176.65和71.38～265.76nmol·g-1·s-1;Nmass值分别为11.8～36.71和12.98～31.78mg·g-1;在群落水平上,干旱生境中LMA显著高于湿润生境,而Amass显著低于湿润生境,Nmass在2个生境中差异不显著;在群落尺度上,干旱生境中LMA与Nmass和Amass均呈显著负相关;在2生境中Nmass与Amass均呈显著正相关。同一生境下不同植物对环境的适应策略有差异,但各性状间的相关关系具有趋同性;从湿润到干旱生境,植物由快速生长转向以提高资源利用效率为主的适应策略。     

Interactions and competition processes among tree species in young experimental mixed forests,assessed with chlorophyll fluorescence and leaf morphology

Chlorophyll a fluorescence (ChlF) and leaf morphology were assessed in two sites in Europe (Kaltenborn, Germany, and Satakunta, Finland) within a forest diversity experiment. Trees at Satakunta, planted in 1999, form a stratified canopy, while in Kaltenborn the trees are 7 years old, with no apparent canopy connection among broadleaf species. The following ChlF parameters from measured OJIP transient curves were examined: F_V/F_M (a proxy for maximum quantum yield); Ψ_Eo (a proxy for efficiency in transferring an electron from reduced QA to the electron transport chain); I‐P phase (a proxy for efficiency of reducing final acceptors beyond PSI); and PI_tot (total performance index for potential energy conservation from photons absorbed by PSII to reduction of PSI end acceptors). At Satakunta F_V/F_M and Ψ_Eo in Betula pendula were higher in monocultures and lower in mixed plots, perhaps due to increasing light availability in mixed plots, which can induce photoinhibition. The opposite trend was observed in Picea abies, which was shaded in mixed plots. At Kaltenborn F_V/F_M decreased in Fagus sylvatica and P. abies in mixed plots due to competition both above‐ and belowground. At Satakunta LMA increased in B. pendula leaves with increasing species richness. Leaf area of ten leaves was reduced in F. sylvatica in mixed plots at Kaltenborn. By up‐scaling the overall fluorescence response to plot level (PI_{tot_plot}), a significant positive correlation with tree diversity was found at Kaltenborn, but not at Satakunta. This could suggest that competition/facilitation processes in mixed stands play a significant role in the early stages of forest establishment, but then tend to be compensated in more mature stands.     

Linking vein properties to leaf biomechanics across 58 woody species from a subtropical forest

• Leaf venations have elements with relatively lower elasticity than other leaf tissue components, which are thought to contribute to leaf biomechanics. A better mechanistic understanding of relationships between vein traits and leaf mechanical properties is essential for ecologically relevant interpretation of leaf structural variations.
• We investigated 13 major (first to third order) and minor (>third order) vein traits, six leaf mechanical properties and other structural traits across 58 woody species from a subtropical forest to elucidate how vein traits contribute to leaf biomechanics.
• Across species, vein dry mass density (ρ_v), total vein dry mass per leaf area (VMA) and minor vein diameter (VD_min), but not the lower‐order vein density (VLA_1?2), were positively correlated with leaf force to punch (F_p) and force to tear (F_t). Structural equation models showed that ρ_v and VD_min not only contribute to leaf mechanical properties directly (direct pathway), but also had impacts on leaf biomechanics by influencing leaf thickness and leaf dry mass per area (indirect pathway).
• Our study demonstrated that vein dry mass density and minor vein diameter are the key vein properties for leaf biomechanics. We also suggest that the mechanical characteristics of venations are potential factors influencing leaf mechanical resistance, structure and leaf economics spectrum.

     

Specific leaf area, leaf nitrogen content, and photosynthetic acclimation of Trifolium repens L. seedlings grown at different irradiances and nitrogen concentrations

Clover seedlings were grown at different nitrogen concentrations (5, 10, 15, 20, 25 mM NO₃ ⁻, i.e. N₅ to N₂₅) and two irradiances, I (200 and 400 μmol m⁻² s⁻¹ of photon flux density, i.e. I ₂₀₀ and I ₄₀₀). Net photosynthetic rate (P _N), photosynthetic nitrogen use efficiency (PNUE), leaf chlorophyll (Chl) content, maximum photochemical efficiency (F_v/F_m), and actual photochemical efficiency of photosystem 2 (PS2) (Φ_PS2) increased from N₅ to N₁₅ and decreased with N₁₅ to N₂₅. P _N, PNUE, and Φ_PS2 were higher at I ₄₀₀ than at I ₂₀₀, but F_v/F_m and leaf Chl contents at I ₄₀₀ were lower than at I ₂₀₀. The effects of the N and I on specific leaf area (SLA) and N contents per unit dry mass (N_m) were similar, the SLA and N_m increased from N₅ to N₂₅ and they were higher at I ₂₀₀ than at I ₄₀₀. The nitrogen contents per unit area (N_a) increased from N₅ to N₂₀, but decreased from N₂₀ to N₂₅. The N_a was higher at I ₂₀₀ than at I ₄₀₀ when Trifolium repens grew at N₅ and N₁₀, but it was higher at I ₄₀₀ than at I ₂₀₀ at N₁₅ to N₂₅.     

Chemical and mechanical changes during leaf expansion of four woody species of dry Restinga woodland

Young leaves are preferential targets for herbivores, and plants have developed different strategies to protect them. This study aimed to evaluate different leaf attributes of presumed relevance in protection against herbivory in four woody species (Erythroxylum argentinum, Lithrea brasiliensis, Myrciaria cuspidata, and Myrsine umbellata), growing in a dry restinga woodland in southern Brazil. Evaluation of leaf parameters was made through single-point sampling of leaves (leaf mass per area and leaf contents of nitrogen, carbon, and pigments) at three developmental stages and through time-course sampling of expanding leaves (area and strength). Leaves of M. umbellata showed the highest leaf mass per area (LMA), the largest area, and the longest expansion period. On the other extreme, Myrc. cuspidata had the smallest LMA and leaf size, and the shortest expansion period. Similarly to L. brasiliensis, it displayed red young leaves. None of the species showed delayed-greening, which might be related to the high-irradiance growth conditions. Nitrogen contents reduced with leaf maturity and reached the highest values in the young leaves of E. argentinum and Myrc. cuspidata and the lowest in M. umbellata. Each species seems to present a different set of protective attributes during leaf expansion. Myrciaria cuspidata appears to rely mostly on chemical defences to protect its soft leaves, and anthocyanins might play this role at leaf youth, while M. umbellata seems to invest more on mechanical defences, even at early stages of leaf growth, as well as on a low allocation of nitrogen to the leaves. The other species display intermediate characteristics.     

Effects of leaf age,elevation and light conditions on photosynthesis and leaf traits in saplings of two evergreen conifers,Abies veitchii and A. mariesii

亚高山的针叶树物种分布在广泛的海拔范围内,因此它们必须具备抵抗严酷自然条件(如高海拔和长期的黑暗)的能力。 在日本中部的亚高山地带,两种常绿的针叶树冷杉(Abies veitchii)和马尾冷杉(Abies mariesii)分别在低海拔和高海拔地区占据优势。本研究的目的是探讨叶龄、海拔和光照条件对两个物种叶片形态和生理性状的影响。我们以两种针叶树的幼树为样本,分别在两种针叶树下部 (1600米)和上部(2300米)的林下层和冠层间隙中取样,并分析叶龄、海拔和光照条件对冷杉和马尾冷杉的光合作用和叶片性状的影响。研究结果表明,两种针叶树的光合作用和叶片性状对叶龄呈现相似的响应,但其对海拔和光照条件则表现出不同的响应。冷杉最大光合速率与单位叶面积质量和非结构性碳水化合物浓度呈负相关。两种树木的单位叶面积质量在高海拔处均增加,而非结构性碳水化合物浓度仅在冷杉中增加。因此,冷杉的最大光合速率在高海拔时降低。此外,两个物种的最大光合速率均与氮浓度呈正相关。在林下层,叶片氮浓度在冷杉中下降,而在马尾冷杉中上升。在林下条件下,只有马尾冷杉的单位叶面积质量下降、叶绿素-氮比增加,表明在黑暗条件下,马尾冷杉比冷杉有更高的捕光效率。本研究表明,与冷杉相比,马尾冷杉的光合和叶片特性具有更强的耐阴性,光合速率受海拔的影响较小,从而使马尾冷杉在林下植被中生存,并在高海拔地区占据优势。     

不同光环境下亚热带常绿阔叶树种和落叶阔叶树种幼苗的叶形态和光合生理特征

由于地球环境的演变,亚热带常绿阔叶林中常有落叶阔叶成分的存在,但从生态角度,其存在的机理尚不清楚.通过比较研究常绿阔叶优势树种大头茶与落叶阔叶树种枫香幼苗的叶形态和光合生理特征对不同光环境(旷地、林窗、林下)的响应,尝试解释落叶成分遗留的基础.结果表明:①枫香具有较小的比叶重(LMA)和较高的光合氮利用效率(PNUE),在高光环境中,并未受到光抑制,具有较高的光合可塑性,尤其在林窗,表现出较大头茶高3倍的光合能力(Pnmax),保证了其在较短的生长季节积累更多的光合产物,增强与常绿树种的竞争力;②常绿阔叶树种大头茶具有较大的LMA,在旷地受到严重的光抑制,将更多的氮用于化学防御中.在林窗和林下具有相对高的Pnmax,生长幅度较广,但在林下将更多的氮投入到比生长更为重要的生存消耗中.而枫香在林下响应表现为Rubisco活性降低,光合受阻;③两树种在林窗均表现出较高的光合适应性,具有最大的光合能力(Pnmax)及合理的氮在光合机构中的分配系数,说明林窗是它们更新的最佳环境.但枫香的光合作用更依赖于光照,而大头茶则更依赖于CO2浓度;④总之,落叶阔叶树种以其较高的形态可塑性和对高光的光合生理可塑性能够在常绿阔叶林种生存、生长,并成为常绿阔叶林中固有成分的主要原因之一.     

Comparison of chlorophyll fluorescence emission characteristics of wheat leaf tissue and isolated thylakoids as a function of excitation wavelength

Abstract. Chlorophyll fluorescence emission spectra and the kinetics of 685 mm fluorescence emission from wheat leaf tissue and thylakoids isolated from such tissue were examined as a function of excitation wavelength. A considerable enhancement of fluorescence emission above 700 nm relative to that at 685 nm was observed from leaf tissue when it was excited with 550 nm rather than 450 nm radiation. Such excitation wavelength dependent changes in the emission spectrum occurred over an excitation spectral range of 440–660 nm and appeared to be directly related to the total quantity of radiation absorbed at a given excitation wavelength. Experiments with isolated thylakoid preparations demonstrated that changes in the fluorescence emission spectrum of the leaf were attributable to the optical properties of the leaf and were not due to the intrinsic characteristies of the thylakoid photochemical apparatus. This was not the case for the observed excitation wavelength dependent changes in the 685 nm fluorescence induction curve obtained from leaf tissue infiltrated with DCMU. Excitation wavelength dependent changes in the ratio of the variable to maximal fluorescence emission and the shape of the variable fluorescence induction were observed for leaf tissue. Isolated thylakoid studies showed that such changes in the leaf fluorescence kinetics were representative of the way in which the photochemical apparatus in vivo was processing the absorbed radiation at the different excitation wavelengths. The results are considered in the context of the use of fluorescence emission characteristics of leaves as non-destructive probes of the photochemical apparatus in vivo.     

The influence of chlorophyll fluorescence on the light gradients and the phytochrome state in a green model leaf under natural conditions

Abstract. The effect of chlorophyll fluoresence on the spectral light gradients within a model green leaf was examined under different light qualities (day-light, sunset, canopy) and different quantum efficiencies. Light fluxes within the leaf tissue are nearly doubled in the emission domain of fluorescence but the effect on the phytochrome photoequilibrium is very small.     

Functional relationship between chlorophyll content and leaf reflectance, and light-capturing efficiency of Japanese forest species

We examined the functional relationship between chlorophyll concentrations and light spectral absorption in 16 species of woody, vine and herbaceous plants in northern Japan. Leaves of each species from under forest shade and in more open sites were measured for chlorophyll, specific leaf area (SLA) and spectral absorption. In all species, SLA increased and the Chl a : b ratio declined in shade- vs open-grown leaves indicating an adaptive adjustment to forest shade in these leaf characters. However, the expected increase in the ratio of 680 to 700 nm absorption in shade leaves did not occur in all species. Light absorption at 680 relative to 700 nm was lower in the shade leaves of Acer japonicum. Kalopanax pictus, Panax japonicus and Petasites japonicus even with a reduced Chl a : b , a commonly accepted indicator of shade adaptation. Therefore, spectral measurements in these species failed to support Chl concentrations that were expected to confer an improvement in the absorption of red light (<680nm) deficient relative to far-red light (>700 nm) in the forest shade. Compared with other species, the absorption pattern of these four 'non-conforming' species is associated with a higher ratio of shade:open leaves in reflectance spectra in the 600–750 nm range. This suggests an increased reflectance in shade leaves caused by changes in leaf surface properties which are not immediately apparent. We conclude that adaptive spectral absorption cannot always be inferred from changes in specific leaf area and chlorophyll a and b concentrations.     

Photosynthetic characteristics of three varieties of Lilium “Oriental Hybrids” in the central areas of Yunnan Province, China

To understand the ecophysiological adapta-tion of Lilium "Oriental Hybrids", which are grown for their commercial bulbs, the gas exchange, leaf N and chlorophyll content of the three varieties were investigated in the central areas of the Yunnan Province. Among the three varieties, light-saturated photosynthetic rate at ambient CO2 (Amax) of Tiber was the highest, while that of Siberia was the lowest. The difference in the Amax was related to the carboxylation efficiency (CE), leaf mass per unit area and leaf N content per mass, which indicated that their photosynthetic capacity was influenced by the activity and/or the quantity of Rubisco. The three vari-eties had lower photosynthetic saturation points and pho-tosynthetic compensation points, but the photosynthetic rates were not decreased up to 2000 μmol.m-2s-1 of the light intensity. This indicates that the three varieties had broad adaptability to light intensity. There were signifi-cant differences in the photosynthetic optimum temper-ature among the three varieties. Siberia had the highest photosynthetic optimum temperature (25.5℃-34.9℃), and is likely to grow well in warm areas. Sorbonne had the lowest photosynthetic optimum temperature (19.3℃-25.6℃), and its growth is favored in cool areas. Tiber can maintain a high photosynthetic rate within a wide range of temperature. Therefore, Tiber is the most suitable variety for the climate in the central areas of the Yunnan Province, China.     

Evaluation of a technique for the measurement of chlorophyll fluorescence from leaves exposed to continuous white light

Abstract An instrument for the generation and measurement of modulated chlorophyll fluorescence signals from leaves exposed to continuous, highintensity white light is described. Modulated fluorescence is generated in the leaf by pulsed diodes emitting low-intensity yellow radiation and is detected with a photodiode whose output is fed to an amplifier locked in to the frequency of the lightemitting diodes. Comparisons are made between the modulated fluorescence signals measured with this instrument and the continuous fluorescence signals emitted from dark-adapted leaf tissue and isolated thylakoids when photosynthetic activity is induced by exposure to a range of intensities of continuous broad-band, blue-green light. The modulated fluorescence signals were similar to the continuous fluorescence signals, but they were not always identical. The small differences between the two signals are mainly attributable to differences in the populations of chloroplasts being monitored in the two measurements as a result of differential penetration of the modulated and actinic light sources into the sample.     

Vertical variation and economic strategy of leaf trait of major tree species in a typical mixed broadleaved-Korean pine forest

分析不同树种叶片性状的变化有助于了解植物群落结构。该文通过对典型阔叶红松(Pinus koraiensis)林15种阔叶树种的比叶质量、叶片厚度、叶干物质含量、叶绿素含量指数、叶片碳、氮、磷含量的测定, 分析了冠层高度对叶性状及叶性状间相关关系的影响。结果表明, 水曲柳(Fraxinus mandshurica)和大青杨(Populus ussuriensis)上层的比叶质量显著大于下层, 而其他树种冠层间的比叶质量无显著变化; 叶绿素含量指数在白桦(Betula platyphylla)和春榆(Ulmus japonica)冠层间的分布分别为上层显著大于下层和上层显著大于中层; 单位质量氮含量在水曲柳的中层显著大于上层。叶片性状间存在着广泛的相关性, 比叶质量与叶片厚度、干物质含量在三层间均呈显著正相关关系, 而有些性状, 只在一或二个冠层中存在一定的相关性。山杨(Populus davidiana)和大青杨的叶片倾向于选择光合能力较低、营养浓度较低、呼吸速率较慢的一端, 而黄檗(Phellodendron amurense)和山槐(Maackia amurensis)叶片更倾向于光合能力强、营养物质浓度高的一端。不同树种对光照响应的差异可能会改变不同冠层中叶片的形态和化学性状, 从而有助于群落构建和物种共存。     

典型城区与郊区环境大叶黄杨气体交换及叶绿素荧光特性比较

采用Li-6400便携式光合作用测定系统对夏秋季典型城区与郊区环境下大叶黄杨的气体交换和叶绿素荧光特性进行了现场实验比较研究.研究显示,叶片净光合速率的大小由总光合速率(光合能力)和呼吸速率共同决定,城区环境温度较高、相对湿度较低、大气CO2浓度较高, 不同月份城区和郊区样点大叶黄杨的净光合速率差异显著性存在不同.城区环境下大叶黄杨的胞间CO2浓度、叶面水气压亏缺、蒸腾速率高于郊区环境.城区环境中温度、大气CO2浓度等的变化会影响叶片呼吸作用,造成呼吸速率升高或是降低,城区环境中污染物浓度变化也会损伤叶片光合结构从而导致总光合能力降低,这两者都会引起净光合速率的变化.通过大叶黄杨叶片叶绿素荧光指标的进一步对比分析发现,城区大叶黄杨叶片叶绿素总量、叶绿素a/b、Fv/Fm、Fv/Fo、qP、ΦPSⅡ、ETR降低,但qN升高.表明叶片叶绿体PSⅡ的功能受到负面影响.城区大叶黄杨叶片荧光参数的变化,从微观机制上表明城区环境中污染物浓度的上升导致叶绿素及叶绿体光合结构受损的确是叶片光合能力下降的主要原因之一.     

Underestimate of PS2 efficiency in the field due to high leaf temperature resulting from leaf clipping and its amendment

Chlorophyll fluorescence parameter F_v/F_m, an indicator of the maximum efficiency of PS2, is routinely measured in the field with plant leaves darkened by leaf clips. I found that on a sunny day of subtropical summer, the F_v/F_m ratio was often underestimated because of a large F₀ value resulted from a high leaf temperature caused by clipping the leaf under high irradiance, especially for long (e.g. 20 min) duration. This phenomenon may overestimate the down-regulation of PS2 efficiency under high irradiance. When leaf temperature was lower than 40 °C, the F₀ level of rice leaves under clipping remained practically unchanged. However, F₀ increased drastically with leaf temperature rising over 40 °C. In most measurements, no significant difference in F_m was found between rice leaves dark-adapted by leaf clips for 10 min and for 20 min. Therefore, shading leaf clips to prevent a drastic increase of leaf temperature, using F₀ measured immediately after the leaf being darkened to calculate F_v/F_m, as well as shortening the duration of leaf clipping are useful means to avoid an underestimate of F_v/F_m.     

典型阔叶红松林主要树种叶性状的垂直变异及经济策略

分析不同树种叶片性状的变化有助于了解植物群落结构。该文通过对典型阔叶红松(Pinus koraiensis)林15种阔叶树种的比叶质量、叶片厚度、叶干物质含量、叶绿素含量指数、叶片碳、氮、磷含量的测定, 分析了冠层高度对叶性状及叶性状间相关关系的影响。结果表明, 水曲柳(Fraxinus mandshurica)和大青杨(Populus ussuriensis)上层的比叶质量显著大于下层, 而其他树种冠层间的比叶质量无显著变化; 叶绿素含量指数在白桦(Betula platyphylla)和春榆(Ulmus japonica)冠层间的分布分别为上层显著大于下层和上层显著大于中层; 单位质量氮含量在水曲柳的中层显著大于上层。叶片性状间存在着广泛的相关性, 比叶质量与叶片厚度、干物质含量在三层间均呈显著正相关关系, 而有些性状, 只在一或二个冠层中存在一定的相关性。山杨(Populus davidiana)和大青杨的叶片倾向于选择光合能力较低、营养浓度较低、呼吸速率较慢的一端, 而黄檗(Phellodendron amurense)和山槐(Maackia amurensis)叶片更倾向于光合能力强、营养物质浓度高的一端。不同树种对光照响应的差异可能会改变不同冠层中叶片的形态和化学性状, 从而有助于群落构建和物种共存。     

A test of the generality of leaf trait relationships on the Tibetan Plateau

Leaf mass per area (LMA), nitrogen concentration (on mass and area bases, N(mass) and N(area), respectively), photosynthetic capacity (A(mass) and A(area)) and photosynthetic nitrogen use efficiency (PNUE) are key foliar traits, but few data are available from cold, high-altitude environments. Here, we systematically measured these leaf traits in 74 species at 49 research sites on the Tibetan Plateau to examine how these traits, measured near the extremes of plant tolerance, compare with global patterns. Overall, Tibetan species had higher leaf nitrogen concentrations and photosynthetic capacities compared with a global dataset, but they had a slightly lower A(mass) at a given N(mass). These leaf trait relationships were consistent with those reported from the global dataset, with slopes of the standardized major axes A(mass)-LMA, N(mass)-LMA and A(mass)-N(mass) identical to those from the global dataset. Climate only weakly modulated leaf traits. Our data indicate that covarying sets of leaf traits are consistent across environments and biogeographic regions. Our results demonstrate functional convergence of leaf trait relationships in an extreme environment.     

Divergent patterns of foliar δ13C and δ15N in Quercus aquifolioides with an altitudinal transect on the Tibetan Plateau: an integrated study based on multiple key leaf functional traits

Aims With a close association with plant water availability, foliar δ 13 C had been investigated extensively in alpine regions; however, foliar δ 15 N has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar δ 13 C and δ 15 N found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient.Methods To test our hypothesis, we measured foliar δ 13 C and δ 15 N in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m.Important findings (i) Contrary to our hypothesis, foliar δ 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar δ 15 N initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar δ 13 C, while differences in foliar δ 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between δ 13 C and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with δ 15 N and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.