Shifts and plasticity of plant leaf mass per area and leaf size among slope aspects in a subalpine meadow

The composition of vegetation on a slope frequently changes substantially owing to the different micro‐environments of various slope aspects. To understand how the slope aspect affects the vegetation changes, we examined the variations in leaf mass per area (LMA) and leaf size (LS) within and among populations for 66 species from 14 plots with a variety of slope aspects in a subalpine meadow. LMA is a leaf economic trait that is tightly correlated with plant physiological traits, while the LS shows a tight correlation with leaf temperature, indicating the strategy of plants to self‐adjust in different thermal and hydraulic conditions. In this study, we compared the two leaf traits between slope aspects and between functional types and explored their correlation with soil variables and heat load. Our results showed that high‐LMA, small‐leaved species were favored in south‐facing slopes, while the reverse was true in north‐facing areas. In detail, small dense‐leaved graminoids dominated the south slopes, while large thin‐leaved forbs dominated the north slopes. Soil moisture and the availability of soil P were the two most important soil factors that related to both LMA and LS, and heat load also contributed substantially. Moreover, we disentangled the relative importance of intraspecific trait variation and species turnover in the trait variation among plots and found that the intraspecific variation contributed 98% and 56% to LMA and LS variation among communities, respectively, implying a large contribution of intraspecific trait plasticity. These results indicate that LMA and LS are two essential leaf traits that affect the adaptation or acclimation of plants underlying the vegetation composition changes in different slope aspects in the subalpine meadow.     

The anatomical and compositional basis of leaf mass per area

Leaf dry mass per unit leaf area (LMA) is a central trait in ecology, but its anatomical and compositional basis has been unclear. An explicit mathematical and physical framework for quantifying the cell and tissue determinants of LMA will enable tests of their influence on species, communities and ecosystems. We present an approach to explaining LMA from the numbers, dimensions and mass densities of leaf cells and tissues, which provided unprecedented explanatory power for 11 broadleaved woody angiosperm species diverse in LMA (33–262 g m^?2; R² = 0.94; P < 0.001). Across these diverse species, and in a larger comparison of evergreen vs. deciduous angiosperms, high LMA resulted principally from larger cell sizes, greater major vein allocation, greater numbers of mesophyll cell layers and higher cell mass densities. This explicit approach enables relating leaf anatomy and composition to a wide range of processes in physiological, evolutionary, community and macroecology.     

The leaf economics spectrum’s morning coffee: plant size-dependent changes in leaf traits and reproductive onset in a perennial tree crop

Background and AimsSize-dependent changes in plant traits are an important source of intraspecific trait variation. However, there are few studies that have tested if leaf trait co-variation and/or trade-offs follow a within-genotype leaf economics spectrum (LES) related to plant size and reproductive onset. To our knowledge, there are no studies on any plant species that have tested whether or not the shape of a within-genotype LES that describes how traits covary across whole plant sizes, is the same as the shape of a within-genotype LES that represents environmentally driven trait plasticity.MethodsWe quantified size-dependent variation in eight leaf traits in a single coffee genotype (Coffea arabica var. Caturra) in managed agroecosystems with different environmental conditions (light and fertilization treatments), and evaluated these patterns with respect to reproductive onset. We also evaluated if trait covariation along a within-genotype plant-size LES differed from a within-genotype environmental LES defined with trait data from coffee growing in different environmental conditions.Key ResultsLeaf economics traits related to resource acquisition – maximum photosynthetic rates (A) and mass-based leaf nitrogen (N) concentrations – declined linearly with plant size. Structural traits – leaf mass, leaf thickness, and leaf mass per unit area (LMA) – and leaf area increased with plant size beyond reproductive onset, then declined in larger plants. Three primary LES traits (mass-based A, leaf N and LMA) covaried across a within-genotype plant-size LES, with plants moving towards the ‘resource-conserving’ end of the LES as they grow larger; in coffee these patterns were nearly identical to a within-genotype environmental LES.ConclusionsOur results demonstrate that a plant-size LES exists within a single genotype. Our findings indicate that in managed agroecosystems where resource availability is high the role of reproductive onset in driving within-genotype trait variability, and the strength of covariation and trade-offs among LES traits, are less pronounced compared with plants in natural systems. The consistency in trait covariation in coffee along both plant-size and environmental LES axes indicates strong constraints on leaf form and function that exist within plant genotypes.     

Individual‐level leaf trait variation and correlation across biological and spatial scales

Even with increasing interest in the ecological importance of intraspecific trait variation (ITV) for better understanding ecological processes, few studies have quantified ITV in seedlings and assessed constraints imposed by trade‐offs and correlations among individual‐level leaf traits. Estimating the amount and role of ITV in seedlings is important to understand tree recruitment and long‐term forest dynamics. We measured ten different size, economics, and whole leaf traits (lamina and petiole) for more than 2,800 seedlings (height ≥ 10 cm and diameter at breast height < 1 cm) in 283 seedling plots and then quantified the amount of ITV and trait correlations across two biological (intraspecific and interspecific) and spatial (within and among plots) scales. Finally, we explored the effects of trait variance and sample size on the strength of trait correlations. We found about 40% (6%–63%) variation in leaf‐level traits was explained by ITV across all traits. Lamina and petiole traits were correlated across biological and spatial scales, whereas leaf size traits (e.g., lamina area) were weakly correlated with economics traits (e.g., specific lamina area); lamina mass ratio was strongly related to the petiole length. Trait correlations varied among species, plots, and different scales but there was no evidence that the strength of trait relationships was stronger at broader than finer biological and spatial scales. While larger trait variance increased the strength of correlations, the sample size was the most important factor that was negatively related to the strength of trait correlations. Our results showed that a large amount of trait variation was explained by ITV, which highlighted the importance of considering ITV when using trait‐based approaches in seedling ecology. In addition, sample size was an important factor that influenced the strength of trait correlations, which suggests that comparing trait correlations across studies should consider the differences in sample size.     

The analysis of the causes of variability of the relationship between leaf dry mass and area in plants

The lamina dry mass: area ratio (LMA - Leaf Mass per Area) is a quite variable trait. Leaf dry mass consists of symplast mass (a set of all leaf protoplasts) and apoplast mass (a set of all cell walls in a leaf). The ratio between symplast and apoplast masses is positively related to any functional trait of leaf calculated per unit of dry mass. The value of this ratio is defined by cells size and their number per unit of leaf area, number of mesophyll cells layers and their differentiation between palisade and spongy ones, and also by density of cells packing. The LMA value is defined by leaf thickness and density. The extent and direction of variability in both leaf traits define the extent and direction of variability in LMA. Negative correlation between leaf thickness and density reduces the level of LMA variability. As a consequence of this correlation the following pattern emerges: the thinner a leaf, the denser it is. Changes in the traits that define the LMA value take place both within a species under the influence of environmental factors and between species that differ in leaf structure and functions. Light is the most powerful environmental factor that influences the LMA, increase in illumination leading to increase in LMA. This effect occurs during leaf growth at the expense of structural changes associated with the reduction of symplast/apoplast mass ratio. Under conditions of intense illumination, LMA may increase due to accumulation of starch. With regard to the majority of leaf functions, the mass of starch may be ascribed to apoplast. Starch accumulation in leaves is observed also under conditions of elevated CO2 concentration in the air. Under high illumination, however, LMA increases also due to increased apoplast contribution to leaf dry mass. Scarce mineral nutrition leads to LMA increase due to lowering of growth zones demands for phothosyntates and, therefore, to increase in starch content of leaves. High level of mineral nutrition during leaf growth period leads to LMA increase at the expense of mesophyll thickening where components of photosynthesis system are located. When additional environmental factors are involved, starch accumulation may be partly responsible for increase in LMA. LMA increase at the expense of starch accumulation, unlike that at the expense of mesophyll thickening, is accompanied by increased leaf density. Under conditions of water deficiency LMA increases, which in mature leaf may be caused by starch accumulation. LMA increase during leaf growth period under conditions of water deficiency is associated with decrease in the symplast/apoplast mass ratio.     

Changes of Leaf Morphological, Anatomical Structure and Carbon Isotope Ratio with the Height of the Wangtian Tree （Parashorea chinensis） in Xishuangbanna, China

Leaf morphological and anatomical structure and carbon isotope ratio （δ^13C） change with increasing tree height. To determine how tree height affects leaf characteristics, we measured the leaf area, specific leaf mass （ratio of leaf mass to leaf area [LMA]）, thickness of the total leaf, cuticle, epidermis, palisade and sponge mesophyll, stomata traits and δ^13C at different heights of Parashorea chinensis with methods of light and scanning electron microscopy （SEM） and isotope-ratio mass spectrometry. The correlation and stepwise regression between tree height and leaf structure traits were carried out with SPSS software. The results showed that leaf structures and δ^13C differed significantly along the tree height gradient. The leaf area, thickness of sponge mesophyll and size of stomata decreased with increasing height, whereas the thickness of lamina, palisade mesophyll, epidermis, and cuticle, ratios of palisade to spongy thickness, density of stomata and vascular bundles, LMA and δ^13C increased with tree height. Tree height showed a significant relationship with all leaf indices and the most significant relationship was with epidermis thickness, leaf area, cuticle thickness, δ^13C. The δ^13C value showed a significantly positive relationship with LMA （R = 0.934）. Our results supported the hypothesis that the leaf structures exhibited more xeromorphic characteristics with the increasing gradient of tree height.     

Leaf thickness controls variation in leaf mass per area (LMA) among grazing-adapted grasses in Serengeti

Leaf mass per area (LMA) is a primary plant functional trait that represents the cost of constructing a leaf. Ultimately, plants modify LMA by altering leaf thickness (LT), leaf dry matter content (LDMC), or both. While LMA can be modified through both of these constituents, studies of LMA have found that there is variation in whether LT or LDMC changes are responsible for LMA—and the relationships change depending on the species or functional groups being compared. In this study, we used a phylogenetic framework to determine that evolutionary shifts in LMA are driven by LT, and not LDMC, among 45 Serengeti grass species. We considered two alternative hypotheses that could result in evolutionary correlation of LMA on LT but not LDMC: either (1) LT is more labile than LDMC—and is therefore a less costly means to change LMA or (2) LDMC is tightly coupled to a different dimension of leaf variation (e.g., leaf hydraulics), leaving LT as the source of variation in LMA. LT was not more labile than LDMC, leading us to conclude that the evolution of LMA has been shaped by LT because LDMC is responding to other demands on leaf physiology. We speculate that leaf hydraulics provide this constraint on LDMC. The decoupling of LDMC from LT may allow plants to better optimize resource allocation in ecosystems where gradients in light competition, herbivory, and aridity place competing demands on leaf economics.     

Leaf Mass per Area (LMA) and Its Relationship with Leaf Structure and Anatomy in 34 Mediterranean Woody Species along a Water Availability Gradient

Leaf mass per area (LMA) is a morphological trait widely used as a good indicator of plant functioning (i.e. photosynthetic and respiratory rates, chemical composition, resistance to herbivory, etc.). The LMA can be broken down into the leaf density (LD) and leaf volume to area ratio (LVA or thickness), which in turn are determined by anatomical tissues and chemical composition. The aim of this study is to understand the anatomical and chemical characteristics related to LMA variation in species growing in the field along a water availability gradient. We determined LMA and its components (LD, LVA and anatomical tissues) for 34 Mediterranean (20 evergreen and 14 deciduous) woody species. Variation in LMA was due to variation in both LD and LVA. For both deciduous and evergreen species LVA variation was strongly and positively related with mesophyll volume per area (VA or thickness), but for evergreen species positive relationships of LVA with the VA of epidermis, vascular plus sclerenchyma tissues and air spaces were found as well. The leaf carbon concentration was positively related with mesophyll VA in deciduous species, and with VA of vascular plus sclerenchymatic tissues in evergreens. Species occurring at the sites with lower water availability were generally characterised by a high LMA and LD.     

Size‐dependent and environment‐mediated shifts in leaf traits of a deciduous tree species in a subtropical forest

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.     

兴安落叶松针叶解剖结构变化及其光合能力对气候变化的适应性

叶片易受环境因子影响,其形态解剖结构特征不但与叶片的生理功能密切相关,而且反映树木对环境变化的响应和适应。叶片结构的改变势必会改变树木的生理功能。同一树种长期生长在异质环境条件下,经过自然选择和适应,会在形态和生理特性等方面产生变异,形成特定的地理种群。另外,母体所经受的环境胁迫也会影响到其子代的生长、发育和生理等特征。因此,了解植物叶片形态结构对环境变化的响应与适应是探索植物对环境变化的响应适应机制的基础。兴安落叶松(Larix gmelinii Rupr.)是我国北方森林的优势树种,主要分布在我国东北地区,但日益加剧的气候变化可能会改变其现有的分布区。为了区分叶片对气候变化的可塑性和适应性,本研究采用同质园法比较测定了6个不同气候条件下的兴安落叶松种源的32年生树木的针叶解剖结构和光合生理相关因子,利用石蜡切片方法分析了针叶的解剖结构特征、光合能力（Pmax-a）、水分利用效率（WUE）之间的关系及其对气候变化的适应性。结果表明：表皮细胞厚度、叶肉细胞厚度、传输组织厚度、维管束厚度、内皮层厚度以及叶片总厚度均存在显著的种源间差异（P < 0.05）。叶肉细胞厚度与Pmax-a、气孔导度和WUE之间均存在显著的正相关关系（P < 0.05）。叶肉细胞厚度、表皮细胞厚度、叶片总厚度以及叶肉细胞厚度和表皮细胞厚度在叶片总厚度中所占比例均与种源地的干燥度指数（即年蒸发量与年降水量之比）呈正线性关系。这些结果说明：不同种源兴安落叶松针叶解剖结构因对种源原地气候条件的长期适应而产生显著的差异,从而引起其针叶光合作用、水分利用等生理功能发生相应的变化,从而有利于该树种在气候变化的情景下得以生存和繁衍。     

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

An evolutionary perspective on leaf economics: phylogenetics of leaf mass per area in vascular plants

In plant leaves, resource use follows a trade‐off between rapid resource capture and conservative storage. This “worldwide leaf economics spectrum” consists of a suite of intercorrelated leaf traits, among which leaf mass per area, LMA, is one of the most fundamental as it indicates the cost of leaf construction and light‐interception borne by plants. We conducted a broad‐scale analysis of the evolutionary history of LMA across a large dataset of 5401 vascular plant species. The phylogenetic signal in LMA displayed low but significant conservatism, that is, leaf economics tended to be more similar among close relatives than expected by chance alone. Models of trait evolution indicated that LMA evolved under weak stabilizing selection. Moreover, results suggest that different optimal phenotypes evolved among large clades within which extremes tended to be selected against. Conservatism in LMA was strongly related to growth form, as were selection intensity and phenotypic evolutionary rates: woody plants showed higher conservatism in relation to stronger stabilizing selection and lower evolutionary rates compared to herbaceous taxa. The evolutionary history of LMA thus paints different evolutionary trajectories of vascular plant species across clades, revealing the coordination of leaf trait evolution with growth forms in response to varying selection regimes.     

Patterns in leaf traits of leguminous and non-leguminous dominant trees along a rainfall gradient in Ghana

Aims Both dominance distribution of species and the composition of the dominant species determine the distribution of traits within community. Leaf carbon (C) and nitrogen (N) isotopic composition are important leaf traits, and such traits of dominant species are associated with ecosystem C, water and N cycling. Very little is known how dominant species with distinct traits (e.g. N-fixing leguminous and non-leguminous trees) mediate resource utilization of the ecosystems in stressful environment.Methods Leaves of 81 dominant leguminous and non-leguminous trees were collected in forest (moist semi-deciduous and dry semi-deciduous ecosystems) and savanna (costal savanna, Guinean savanna and west Sudanian savanna ecosystems) areas and the transitional zone (between the forest and the savanna) along the transect from the south to the north of Ghana. We measured leaf traits, i.e. leaf δ 13 C, leaf δ 15 N, leaf water content, leaf mass per area (LMA) and C and N concentration. Correlation analyses were used to examine trait–trait relationships, and relationships of leaf traits with temperature and precipitation. We used analysis of covariance to test the differences in slopes of the linear regressions between legumes and non-legumes.Important findings Leaf δ 13 C, δ 15 N, leaf water content and LMA did not differ between leguminous and non-leguminous trees. Leaf N concentration and C:N ratio differed between the two groups. Moreover, leaf traits varied significantly among the six ecosystems. δ 13 C values were negatively correlated with annual precipitation and positively correlated with mean annual temperature. In contrast, leaf δ 15 N of non-leguminous trees were positively correlated with annual precipitation and negatively correlated with mean annual temperature. For leguminous trees, such correlations were not significant. We also found significant coordination between leaf traits. However, the slopes of the linear relationships were significantly different between leguminous and non-leguminous trees. Our results indicate that shifts in dominant trees with distinct water-use efficiency were corresponded to the rainfall gradient. Moreover, leguminous trees, those characterized with relative high water-use efficiency in the low rainfall ecosystems, were also corresponded to the relative high N use efficiency. The high proportion of leguminous trees in the savannas is crucial to mitigate nutrient stress.     

The interplay among intraspecific leaf trait variation,niche breadth and species abundance along light and soil nutrient gradients

It is assumed that widespread, generalist species have high phenotypic variation, but we know little about how intraspecific trait variation (ITV) relates to species abundance and niche breadth. In the temperate rainforest of southern Chile, we hypothesized that species with wide niche breadth would exhibit 1) high among‐plot ITV, 2) a strong relationship between trait values and the environment, and 3) a close fit between traits and local environment trait optima. We measured leaf functional traits (leaf area, LMA, leaf N and P concentrations) of saplings in woody species, and compared the relative abundance of each species with its niche breadth, measured as the range of light, soil N and P availability. We used the slope of the linear regression of species’ trait–environment relationships to assess the strength and direction of these relationships, and measured the degree to which species’ trait values track the environmental optimum across plots. In some cases, species having wide niche breadth had high ITV in leaf N and also matched traits (LMA and leaf P) to local optima along the light gradient; they also had high ITV in general and matched leaf P to local optima along the soil P gradient. The relationship between species with wide niche breadth and the strength of intraspecific trait–environment relationships was generally weak and varied depending on the niche dimension and trait in question. Species varied considerably in the strength of trait–environment relationships and total magnitude of ITV, and this variation was not generally strongly related to species abundances or niche breadth patterns. In conclusion, trait variation at the community level is not driven by a few abundant, widely distributed species, but depends on the aggregate trait responses of both abundant and rare species. This makes it difficult to scale individual species trait responses up to the community level.     

Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands

Knowledge of the leaf anatomy of grassland plants is crucial for understanding how these plants adapt to the environment. Tibetan alpine grasslands and Inner Mongolian temperate grasslands are two major grassland types in northern China. Tibetan alpine grasslands occur in high-altitude regions where the low temperatures limit plant growth. Inner Mongolian temperate grasslands are found in arid regions where moisture is the limiting factor. Few comparative studies concerning the leaf anatomy of grassland plants of the Tibetan Plateau and Inner Mongolian Plateau have been conducted. We examined leaf characteristics at 71 sites and among 65 species, across the alpine grasslands of the Tibetan Plateau and the temperate grasslands of the Inner Mongolian Plateau. We compared the leaf structures of plants with different life forms and taxonomies, and their adaptation to arid or cold environments. We explored relationships among leaf features and the effects of climatic factors (i.e., growing season temperature and precipitation) on leaf characteristics. Our results showed that (i) there were significant differences in leaf anatomy between Tibetan alpine and Inner Mongolian temperate grasslands. Except for mesophyll cell density, the values obtained for thickness of leaf tissue, surface area and volume of mesophyll cells were larger on the Tibetan Plateau than on the Inner Mongolian Plateau. (ii) Within the same family or genus, leaf anatomy showed significant differences between two regions, and trends were consistent with those of whole species. (iii) Leaf anatomy of woody and herbaceous plants also showed significant differences between the regions. Except for mesophyll cell density, the values obtained for the thickness of leaf tissue, and the surface area and volume of mesophyll cells were larger in herbaceous than in woody plants. (iv) Leaf anatomical traits changed accordingly. Total leaf thickness, thicknesses of lower and upper epidermal cells, and surface area and volume of mesophyll cells were positively correlated, while mesophyll cell density was negatively associated with those traits. (v) Growing season temperature had stronger effects on leaf anatomy than growing season precipitation. Although the communities in Tibetan and Inner Mongolian grasslands were similar in appearance, leaf anatomy differed; this was probably due to the combined effects of evolutionary adaptation of plants to environment and environmental stress induced by climatic factors.     

三峡水库消落带植物叶片光合与营养性状特征

自2007年三峡大坝试运行以来,其独特的人工水位调度节律给当地的水库消落带生态系统带来了巨大的负面影响。植物功能性状可以反映某一特殊生境植物的生理生态过程特殊性,是指示生态系统结构与功能的有效指标。因此,在三峡水库消落带形成2a后,于2009年调查了消落带的42种适生植物以及对照带33种植物的6个叶片功能性状:最大净光合速率(Amax)、叶片气孔导度(Gs)、比叶重(LMA)、叶片全氮含量(Nmass)、全磷含量(Pmass)和全钾含量(Kmass)。运用标准化主轴回归分析对消落带植物叶片各功能性状之间关系进行分析,并对照全球尺度叶片功能性状数据库,旨在说明反季节淹水对消落带植物叶片功能性状之间关系与全球尺度对比发生了哪些变化。同时,运用成对方差t检验的分析方法,对比了在消落带和对照带都存在的33个种的叶片光合与营养性状之间的差异,以阐明消落带植物对消落带特殊生境的生理响应。结果表明:(1)消落带植物叶片各性状关系呈现出与全球尺度基本一致的格局,表现出植物叶性状之间关系的趋同性;(2)消落带植物Amass、Nmass、Pmass和Kmass显著高于全球尺度,而LMA则显著低于全球尺度。处于驯化阶段的消落带植物各叶片性状处在全球叶片经济型谱"低投入-快速回收"的一端。(3)消落带植物叶片Amass与对照带相比,有显著提高。表明三峡水库消落带植物叶片光合能力得到显著提高,这可能是其适应消落带特殊生境的关键生理生态对策之一。     

Is leaf dry matter content a better predictor of soil fertility than specific leaf area?

Background and Aims

Specific leaf area (SLA), a key element of the ‘worldwide leaf economics spectrum’, is the preferred ‘soft’ plant trait for assessing soil fertility. SLA is a function of leaf dry matter content (LDMC) and leaf thickness (LT). The first, LDMC, defines leaf construction costs and can be used instead of SLA. However, LT identifies shade at its lowest extreme and succulence at its highest, and is not related to soil fertility. Why then is SLA more frequently used as a predictor of soil fertility than LDMC?

Methods

SLA, LDMC and LT were measured and leaf density (LD) estimated for almost 2000 species, and the capacity of LD to predict LDMC was examined, as was the relative contribution of LDMC and LT to the expression of SLA. Subsequently, the relationships between SLA, LDMC and LT with respect to soil fertility and shade were described.

Key Results

Although LD is strongly related to LDMC, and LDMC and LT each contribute equally to the expression of SLA, the exact relationships differ between ecological groupings. LDMC predicts leaf nitrogen content and soil fertility but, because LT primarily varies with light intensity, SLA increases in response to both increased shade and increased fertility.

Conclusions

Gradients of soil fertility are frequently also gradients of biomass accumulation with reduced irradiance lower in the canopy. Therefore, SLA, which includes both fertility and shade components, may often discriminate better between communities or treatments than LDMC. However, LDMC should always be the preferred trait for assessing gradients of soil fertility uncoupled from shade. Nevertheless, because leaves multitask, individual leaf traits do not necessarily exhibit exact functional equivalence between species. In consequence, rather than using a single stand-alone predictor, multivariate analyses using several leaf traits is recommended.     

Leaf‐IT: An Android application for measuring leaf area

The use of plant functional traits has become increasingly popular in ecological studies because plant functional traits help to understand key ecological processes in plant species and communities. This also includes changes in diversity, inter‐ and intraspecific interactions, and relationships of species at different spatiotemporal scales. Leaf traits are among the most important traits as they describe key dimensions of a plant's life history strategy. Further, leaf area is a key parameter with relevance for other traits such as specific leaf area, which in turn correlates with leaf chemical composition, photosynthetic rate, leaf longevity, and carbon investment. Measuring leaf area usually involves the use of scanners and commercial software and can be difficult under field conditions. We present Leaf‐IT, a new smartphone application for measuring leaf area and other trait‐related areas. Leaf‐IT is free, designed for scientific purposes, and runs on Android 4 or higher. We tested the precision and accuracy using objects with standardized area and compared the area measurements of real leaves with the well‐established, commercial software WinFOLIA using the Altman–Bland method. Area measurements of standardized objects show that Leaf‐IT measures area with high accuracy and precision. Area measurements with Leaf‐IT of real leaves are comparable to those of WinFOLIA. Leaf‐IT is an easy‐to‐use application running on a wide range of smartphones. That increases the portability and use of Leaf‐IT and makes it possible to measure leaf area under field conditions typical for remote locations. Its high accuracy and precision are similar to WinFOLIA. Currently, its main limitation is margin detection of damaged leaves or complex leaf morphologies.     

Trait coordination and environmental filters shape functional trait distributions of forest understory herbs

1. Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.
2. To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ¹³C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.
3. Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.
4. Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.
5. Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     

Variations of leaf longevity in tropical moist forests predicted by a trait‐driven carbon optimality model

Leaf longevity (LL) varies more than 20‐fold in tropical evergreen forests, but it remains unclear how to capture these variations using predictive models. Current theories of LL that are based on carbon optimisation principles are challenging to quantitatively assess because of uncertainty across species in the ‘ageing rate:’ the rate at which leaf photosynthetic capacity declines with age. Here, we present a meta‐analysis of 49 species across temperate and tropical biomes, demonstrating that the ageing rate of photosynthetic capacity is positively correlated with the mass‐based carboxylation rate of mature leaves. We assess an improved trait‐driven carbon optimality model with in situLL data for 105 species in two Panamanian forests. We show that our model explains over 40% of the cross‐species variation in LL under contrasting light environment. Collectively, our results reveal how variation in LL emerges from carbon optimisation constrained by both leaf structural traits and abiotic environment.