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.     

植物比叶质量研究进展

比叶质量(LMA)是叶片经济型谱中最基础的叶功能性状, 也是重要的复合型结构参数。LMA不仅与植物的许多生理反应密切相关, 而且能定量测定光合产物在单位叶面积的投入, 因而被认为是反映植物生态策略的重要指标。目前, 有关LMA的深入研究已在植物生态学、农学、林学、植物生理学领域全面展开。该文系统阐述了LMA在叶片整体、组织、细胞三个水平的结构解析和计算方法; 重点分析了LMA对光合作用的影响; 讨论了LMA的内在遗传差异以及外部的环境胁迫因子(温度、水分、光照)对LMA的影响, 以期梳理比叶质量研究的思路、策略和方法, 为今后的研究提供借鉴和参考。     

Ontogenetic and temporal trajectories of chemical defence in a cyanogenic eucalypt

Many studies have shown that similarly aged plants within a species or population can vary markedly in the concentration of defence compounds they deploy to protect themselves from herbivores. Some studies have also shown that the concentration of these compounds can change with development, but no empirical research has mapped such an ontogenetic trajectory in detail. To do this, we grew cyanogenic Eucalyptus yarraensis seedlings from three half-sibling families under constant glasshouse conditions, and followed their foliar cyanogenic glycoside (prunasin) concentration over time for 338 days after sowing (DAS). Plants in all families followed a similar temporal pattern. Plants increased in foliar prunasin concentration from a very low level (10 μg cyanide (CN) equivalents g⁻¹) in their first leaves, to a maximum of, on average, 1.2 mg CN g⁻¹ at about 240 DAS. From 240 to 338 DAS, prunasin concentration gradually decreased to around 0.7 mg CN g⁻¹. Significant differences between families in maximum prunasin concentration were detected, but none were detected in the time at which this maximum occurred. In parallel with these changes in prunasin concentration, we detected an approximately linear increase in leaf mass per unit leaf area (LMA) with time, which reflected a change from juvenile to adult-like leaf anatomy. When ontogenetic trajectories of prunasin against LMA were constructed, we failed to detect a significant difference between families in the LMA at which maximum prunasin concentration occurred. This remarkable similarity in the temporal and ontogenetic trajectories between individuals, even from geographically remote families, is discussed in relation to a theoretical model for ontogenetic changes in plant defence. Our results show that ontogeny can constrain the expression of plant chemical defense and that chemical defense changes in a nonlinear fashion with ontogeny.     

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。     

A model explaining genotypic and ontogenetic variation of leaf photosynthetic rate in rice (Oryza sativa) based on leaf nitrogen content and stomatal conductance

BACKGROUNDS AND AIMS: Identification of physiological traits associated with leaf photosynthetic rate (Pn) is important for improving potential productivity of rice (Oryza sativa). The objectives of this study were to develop a model which can explain genotypic variation and ontogenetic change of Pn in rice under optimal conditions as a function of leaf nitrogen content per unit area (N) and stomatal conductance (g(s)), and to quantify the effects of interaction between N and g(s) on the variation of Pn. METHODS: Pn, N and g(s) were measured at different developmental stages for the topmost fully expanded leaves in ten rice genotypes with diverse backgrounds grown in pots (2002) and in the field (2001 and 2002). A model of Pn that accounts for carboxylation and CO diffusion processes, and assumes that the ratio of internal conductance to g(s) is constant, was constructed, and its goodness of fit was examined. KEY RESULTS: Considerable genotypic differences in Pn were evident for rice throughout development in both the pot and field experiments. The genotypic variation of Pn was correlated with that of g(s) at a given stage, and the change of Pn with plant development was closely related to the change of N. The variation of g(s) among genotypes was independent of that of N. The model explained well the variation in Pn of the ten genotypes grown under different conditions at different developmental stages. Conclusions The response of Pn to increased N differs with g(s), and the increase in Pn of genotypes with low g(s) is smaller than that of genotypes with high g(s). Therefore, simultaneous improvements of these two traits are essential for an effective breeding of rice genotypes with increased Pn.     

植物叶片性状对气候变化的响应研究进展

叶片性状反映了植物对环境的高度适应能力及其在复杂生境下的自我调控能力。叶片性状如何响应和适应气候变化是植物适应性研究的重点内容。该文系统综述了叶片大小、比叶质量、叶片氮含量、碳同位素等指标对气候变化响应的最新研究结果。不同叶片性状对气候变化的响应结果存在差异,所指示的生态学含义也有所不同。单一叶片性状不能全面地反映植物对气候变化的响应;不同尺度的研究(如环境的修饰或筛选作用的研究)还存在很多不确定性。高寒地区的研究工作相对缺乏。该文有助于理解植物与气候之间的相互关系、植物对气候变化的响应与适应对策,对了解植物演化、预测植物在未来气候变化条件下的变化特征具有一定意义。     

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.     

Changes in traits of shrub canopies across an aridity gradient in northern Patagonia, Argentina

Low and highly variable precipitation pulses exert a strong selective pressure on plant traits and this might provide axes of ecological differentiation among plant species in arid ecosystems. We asked whether aridity contributes to maintain high diversity of species and morphotypes in shrub canopies. We selected eleven study sites evenly distributed across a 400-km transect in northern Patagonia, Argentina. Precipitation is low and highly variable within and between years but almost homogeneous across the transect (125–150 mm). Mean annual temperature varied, however, ranging from 8 °C (west) to 13.5 °C (east) creating a west–east gradient of aridity (aridity index from 3.7 to 7.3, respectively). Sheep grazing commenced in the early 1900s at a similar intensity across the transect. We recorded the richness and cover of shrubs by species and by morphotypes (drought deciduous tall shrubs, evergreen tall shrubs, medium shrubs, and dwarf shrubs), and further calculated the species and morphotype Shannon diversity index at each site. We assessed the presence of spiny leaves, leaf pubescence, thorny stems, and photosynthetic stems in shrub species of all morphotypes and collected green leaves of the dominant shrub species (more than 80% of the total shrub cover) to assess the leaf area, leaf mass per unit area, N-, lignin- and soluble phenolic-concentrations per species at each site. Richness and diversity of shrub species and morphotypes were positively associated with aridity. The richness and diversity of shrub species with pubescent leaves and thorny stems, and nitrogen concentration in green leaves of dominant shrubs increased with increasing aridity. We conclude that our findings on increased diversification in life history traits, species and morhotypes in shrub canopies with increasing aridity support the hypothesis that variability in aridity provides axes of ecological differentiation among shrub species facilitating their coexistence.     

神农架海拔梯度上4种典型森林的乔木叶片功能性状特征

研究植物群落叶片功能性状沿环境梯度的变化可以更好地认识群落对环境的适应机制,海拔梯度在较小的空间范围内浓缩了不同的群落类型和环境梯度,是研究和理解群落与环境关系的理想场所。以神农架海拔梯度上4种典型森林为研究对象,分析了4种典型森林优势乔木树种的LMA、LDMC、N_area、N_mass、叶片厚度及叶片叶绿素含量等主要叶片功能性状随海拔的变化规律及各功能性状之间的相关性,以期从叶片功能性状角度揭示群落适应环境的机制。研究结果表明:在780-1970 m范围内,神农架海拔梯度上典型森林乔木优势种叶片的、N_area和叶片厚度随海拔升高而下降,而海拔2570 m的亚高山针叶林的LMA、N_area和叶片厚度最高,分别为187.37 g/cm²、1.92 g/m²和385.31 um,且针叶林的LMA是常绿阔叶林的2倍、落叶阔叶林的3.7倍;除常绿落叶阔叶混交林的N_mass小于落叶阔叶林外,N_mass随海拔升高(780-2570 m)而下降;海拔最高(2570 m)的针叶林和海拔最低(780 m)的常绿阔叶林的LDMC和叶绿素含量最高,分别为454.93 g/kg、407.64 g/kg和48.84、44.12;而海拔介于二者之间的常绿落叶阔叶混交林和落叶阔叶林的LDMC和叶绿素含量则最低,分别为338.93 g/kg、337.3 g/kg和38.73、38.19。叶片各功能性状之间存在着显著的相关性:叶片厚度与LMA/LDMC及LMA显著正相关。叶片叶绿素含量与N_area显著正相关。LMA与LDMC、N_area、叶片厚度、叶绿素含量显著正相关,LDMC与N_area、叶绿素含量、叶片厚度显著正相关,N_area与叶片厚度、叶绿素含量显著正相关,叶片厚度、叶绿素含量显著正相关。本研究进一步表明,随着海拔梯度上升环境条件的变化,森林群落不同物种功能性状间的相互作用共同决定着森林群落对环境的适应性。     

Nitrogen in cell walls of sclerophyllous leaves accounts for little of the variation in photosynthetic nitrogen-use efficiency

Photosynthetic rate per unit nitrogen generally declines as leaf mass per unit area (LMA) increases. To determine how much of this decline was associated with allocating a greater proportion of leaf nitrogen into cell wall material, we compared two groups of plants. The first group consisted of two species from each of eight genera, all of which were perennial evergreens growing in the Australian National Botanic Gardens (ANBG). The second group consisted of seven Eucalyptus species growing in a greenhouse. The percentage of leaf biomass in cell walls was independent of variation in LMA within any genus, but varied from 25 to 65% between genera. The nitrogen concentration of cell wall material was 0.4 times leaf nitrogen concentration for all species apart from Eucalyptus , which was 0.6 times leaf nitrogen concentration. Between 10 and 30% of leaf nitrogen was recovered in the cell wall fraction, but this was independent of LMA. No trade-off was observed between nitrogen associated with cell walls and the nitrogen allocated to ribulose 1·5-bisphosphate carboxylase/oxygenase (Rubisco). Variation in photosynthetic rate per unit nitrogen could not be explained by variation in cell wall nitrogen.     

Regulation of oil accumulation in single glands of Eucalyptus polybractea

The accumulation of terpenoid oil was examined in the leaves of Eucalyptus polybractea at scales ranging from individual oil glands to the whole plant. Variations in oil composition and concentration of oil were measured and related to both morphological and physiological parameters. Within a plant, all glands produced oil of broadly similar composition that was not regulated by leaf age or the position of the gland within the leaf. There were, however, distinct differences between plants, suggesting that composition is controlled primarily at the whole-plant level. Oil concentration, too, was regulated primarily at the whole-plant level and was limited by gland capacity. Gland capacity was linked to leaf area and thickness, the final products of leaf expansion. Leaf and plant oil composition is determined not by a mosaic of glands specializing in producing a single or a small group of compounds, but rather by glands with remarkably similar capacities for terpenoid biosynthesis, although oil concentration, limited by gland capacity, may be linked to leaf expansion rather than biosynthetic capacity.     

Variations in photosynthetic rate and associated parameters with age of oil palm leaves under irrigation

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in an adult oil palm (Elaeis guineensis) canopy were highest in the 9^th leaf and progressively declined with leaf age. Larger leaf area (LA) and leaf dry mass (LDM) were recorded in middle leaves. P _N showed a significant positive correlation with g _s and a negative relationship with leaf mass per area (ALM). The oil palm leaf remains photosynthetically active for a longer time in the canopy which contributes significantly to larger dry matter production in general and greater fresh fruit bunch yields in particular.     

Photosynthetic and Transpiration Rates of Soybean as Affected by Different Irradiances During Growth

The diurnal variation of net photosynthetic (PN) and transpiration (E) rates in soybean [Glycine max (L.) Merr. cv. Fukuyutaka] plants grown under 100, 50, or 25 % of full sun irradiance (I100, I50, I25 plants) were compared. In the morning, activities of the plants were measured at irradiances under which they grew. However, during the afternoon, all the plants were tested under full irradiance. The lower the growth irradiance, the lower PN, E, and mesophyll conductance values were found. Stomatal conductance was considerably lower in I25 plants only. Both the increase in specific leaf area (SLA) and the decrease in nitrogen content per leaf area unit contributed to the PN reduction of soybean plants grown under low irradiances. Though E of the plants grown under different irradiances differed less markedly than PN, the water use efficiency declined from I100 to I25.     

贡嘎山地区不同海拔冷杉比叶质量和非结构性碳水化合物含量变化

以贡嘎山地区磨西山谷东北坡和康定山谷西南坡为样地,研究了高山林线和低海拔冷杉休眠期和生长旺盛期针叶比叶质量和组织非结构性碳水化合物含量.结果表明:生长在温暖湿润的磨西山谷东北坡的冷杉针叶比叶质量和组织非结构性碳水化合物含量均高于炎热干燥的康定山谷西南坡;高山林线冷杉比叶质量大于低海拔冷杉;高山林线冷杉组织非结构性碳水化合物含量总体高于低海拔冷杉,且生长期比休眠期更为显著.研究结果不支持\"碳供应不足导致高山林线形成\"的假说.