Variation In Specific Leaf Area Research Articles

Specific leaf area and its within-individual variation in understory evergreen and deciduous woody species in New Zealand

Specific leaf area (SLA) plays a critical role in carbon assimilation and nutrient cycling. While leaf habit (deciduous vs. evergreen) has often been recognized as a reliable predictor of SLA—with deciduous species typically having higher mean SLA values due to lower concentration of structural components compared to evergreens—high variation in SLA among evergreen species suggests further investigation of variation for species with this leaf habit could improve predictions of SLA effects on community and ecosystem processes. Furthermore, the presence of leaves of different ages in evergreen plants, emerging over multiple years, could amplify the within-individual variation in SLA, which remains underexplored. Here we report variations of SLA measured from c. 2000 leaves collected from 36 individuals across 19 woody species in an understory environment in New Zealand (NZ). We found that while most deciduous species, predominantly non-native, clustered towards higher SLA values, evergreen species presented a wide SLA spectrum. Moreover, we found that while being deciduous, having a smaller leaf size, and younger leaves, collected from lateral branches, correlated with elevated SLA values, the leaf habit did not primarily drive the within-individual SLA variation. Instead, smaller leaf size emerged as a significant predictor of within-individual SLA variation. The branch-order effect on SLA underscores a methodological consideration: accurate estimation of total leaf area in evergreen trees requires representative sampling across all branch orders. Our study also highlights the need for integrating leaf traits such as leaf size and branch order into functional trait analyses. Further research is vital to understand the underlying mechanisms of these trait variations and their impacts on ecosystem functioning.

Climate factors dominate the elevational variation in grassland plant resource utilization strategies.

Specific leaf area (SLA) and leaf dry matter content (LDMC) are key leaf functional traits often used to reflect plant resource utilization strategies and predict plant responses to environmental changes. In general, grassland plants at different elevations exhibit varying survival strategies. However, it remains unclear how grassland plants adapt to changes in elevation and their driving factors. To address this issue, we utilized SLA and LDMC data of grassland plants from 223 study sites at different elevations in China, along with climate and soil data, to investigate variations in resource utilization strategies of grassland plants along different elevational gradients and their dominant influencing factors employing linear mixed-effects models, variance partitioning method, piecewise Structural Equation Modeling, etc. The results show that with increasing elevation, SLA significantly decreases, and LDMC significantly increases (P < 0.001). This indicates different resource utilization strategies of grassland plants across elevation gradients, transitioning from a "faster investment-return" at lower elevations to a "slower investment-return" at higher elevations. Across different elevation gradients, climatic factors are the main factors affecting grassland plant resource utilization strategies, with soil nutrient factors also playing a non-negligible coordinating role. Among these, mean annual precipitation and hottest month mean temperature are key climatic factors influencing SLA of grassland plants, explaining 28.94% and 23.88% of SLA variation, respectively. The key factors affecting LDMC of grassland plants are mainly hottest month mean temperature and soil phosphorus content, with relative importance of 24.24% and 20.27%, respectively. Additionally, the direct effect of elevation on grassland plant resource utilization strategies is greater than its indirect effect (through influencing climatic and soil nutrient factors). These findings emphasize the substantive impact of elevation on grassland plant resource utilization strategies and have important ecological value for grassland management and protection under global change.

Evaluation of the Community Land Model-Simulated Specific Leaf Area with Observations over China: Impacts on Modeled Gross Primary Productivity

Specific leaf area (SLA) is a key leaf functional trait associated with the ability to acquire light. Substantial variations in SLA have not been well described in the community land model (CLM) and similar terrestrial biosphere models. How these SLA variations influence the simulation of gross primary productivity (GPP) remains unclear. Here, we evaluated the mismatch in SLA between the CLM4.5 and observed data collected from China and quantified the impacts of SLA variation calculated from both observations and the default values across seven terrestrial biosphere models on modeled GPP using CLM4.5. The results showed that CLM4.5 tended to overestimate SLA values at the top and gradient of the canopy. The higher default SLA values could cause an underestimation of the modeled GPP by 5–161 g C m−2 yr−1 (1%–7%) for temperate needleleaf evergreen tree (NET), temperate broadleaf deciduous tree (BDT), and C3 grass and an overestimation by 50 g C m−2 yr−1 (2%) for temperate broadleaf evergreen tree (BET). Moreover, the observed SLA variation among species ranged from 21% to 59% for 14 plant functional types (PFTs), which was similar to the variation in default SLA values across models (9%–60%). These SLA variations would lead to greater changes in modeled GPP by 7%–19% for temperate NET and temperate BET than temperate BDT and C3 grass (2%–9%). Our study suggested that the interspecific variation in SLA and its responses to environmental factors should be involved in terrestrial biosphere models; otherwise, it would cause substantial bias in the prediction of ecosystem productivity.

极端干旱区多枝柽柳叶片功能性状及其与土壤理化因子的关系

PDF HTML阅读 XML下载 导出引用 引用提醒 极端干旱区多枝柽柳叶片功能性状及其与土壤理化因子的关系 DOI: 10.5846/stxb202204070904 作者: 作者单位: 作者简介: 通讯作者: 中图分类号: 基金项目: 新疆维吾尔自治区自然科学基金(2020D01C053);国家自然科学基金(32001145);新疆维吾尔自治区教育厅"天池博士计划"(TCBS202054) Leaf functional traits of Tamarix ramosissima in extremely arid region and their relationship with soil physicochemical factors Author: Affiliation: Fund Project: 摘要 | 图/表 | 访问统计 | 参考文献 | 相似文献 | 引证文献 | 资源附件 | 文章评论 摘要:叶片是植物获取资源的重要器官,研究其功能性状与环境因子的关系,有助于更好地了解植物对环境要素变化的生态适应性。以塔里木盆地北缘不同生境下(绿洲、过渡带、荒漠)典型荒漠植物多枝柽柳(Tamarix ramosissima)为研究对象,分析其叶片结构和生理性状在不同生境间的差异,建立叶片性状与土壤理化因子的相关关系,解析荒漠植物的生态适应性并揭示影响荒漠植物叶片功能性状的关键环境因子。结果表明:(1)多枝柽柳叶片功能性状具有不同程度的变异,其中叶片面积变异幅度最大(40%),比叶面积、可溶性蛋白含量等变异幅度最小(均为15%)。(2)多枝柽柳叶片厚度和叶片面积等结构性状以及叶片可溶性蛋白、脯氨酸、淀粉含量等生理性状在不同生境中均具有显著差异(P<0.05),其中可溶性蛋白、脯氨酸含量均在荒漠生境中达到最高。(3)多枝柽柳叶片部分功能性状之间存在显著的相关关系,其中叶片厚度、淀粉含量等与叶组织密度呈显著负相关关系(P<0.05),而叶片厚度与淀粉含量之间、可溶性糖含量与非结构性碳水化合物含量之间均呈极显著正相关关系(P<0.01)。(4)通过RDA排序分析发现,土壤速效磷含量、土壤含水量、pH、容重等土壤环境因子影响较大,对多枝柽柳叶片功能性状有较好的解释。研究表明多枝柽柳通过叶片功能性状变化以及性状之间协同-权衡的生态对策来适应极端干旱环境,土壤水分和速效磷含量是影响其叶片功能性状最关键的土壤环境因子,研究可为极端干旱区荒漠植被的科学管理、保护和恢复提供理论依据。 Abstract:Leaf is an important organ for plants to obtain resources. Studying the relationships between its functional traits and environmental factors helps to better understand the ecological adaptability of plants to environmental changes. In this study, with the aim of determining the ecological adaptations of desert plants and revealing the environmental factors that influencing the functional traits of them, we investigated a typical desert plant, Tamarix ramosissima, in different habitats (oasis, transition zone and desert) at the northern edge of the Tarim Basin, analyzed the differences in its leaf structure and physiological traits, and established the correlation between them with soil physicochemical factors. The results showed that: (1) the functional traits of Tamarix ramosissima leaves exhibited different degrees of variation, with the greatest variation in leaf area (40%) and the least variation in specific leaf area and soluble protein content (both 15%). (2) The structural traits including leaf thickness and leaf area, as well as physiological traits including leaf soluble protein, proline, and starch contents of Tamarix ramosissima were significantly different among the three habitats (P<0.05), with soluble protein and proline contents reaching the highest levels in desert habitats. (3) There were significant correlations between several functional traits of Tamarix ramosissima leaves. Leaf thickness and starch content were significantly negatively correlated with leaf tissue density (P<0.05), while the correlations between leaf thickness and starch content, soluble sugar content and non-structural carbohydrate content were significantly positive (P<0.01). (4) The RDA analysis revealed that soil environmental factors including soil available phosphorus content, soil water content, pH and bulk density had the greatest influence, and could explain the leaf functional trait variations of Tamarix ramosissima better. This study indicates that Tamarix ramosissima adapts to extreme arid environment by developing ecological responses that varying in leaf functional traits and forming synergistic trade-offs between them. Soil available phosphorus content and soil water content are the most critical soil environmental factors affecting leaf functional traits of Tamarix ramosissima. The findings of this study provide a theoretical basis for the scientific management, conservation, and restoration of desert vegetation in extreme arid region. 参考文献 相似文献 引证文献

Specific leaf area is lower on ultramafic than on neighbouring non-ultramafic soils

ABSTRACT Background Specific leaf area (SLA) is a core trait within the leaf economic spectrum that describes differences in plant performance and productivity. Research on the sources of variation in the leaf economic spectrum and SLA has primarily focused on climate. Much less is known about SLA variation across unusual edaphic environments, such as on ultramafic soils. Aims To determine the role of ultramafic soils as a driver of SLA variation. Methods We measured SLA for dominant species on paired ultramafic and non-ultramafic soils in five biogeographically distinct regions around the globe and compared mean SLA values to globally reported values. Results SLA was lower on ultramafic than on non-ultramafic soils in all regions, except Puerto Rico, and both climate and soil were important drivers of SLA. For three of the five regions, SLA values on ultramafic soils were lower than the global average. Conclusions Soils can be a major driver of SLA along with climate. Low SLA on ultramafic soil points to selection for stress resistance strategies. Furthermore, in some bioregions, SLA values on ultramafic soils were among the lowest on the planet and thus represent globally rare phenotypes that should be conserved within these unique edaphic habitats.

Divergent response and adaptation of specific leaf area to environmental change at different spatio-temporal scales jointly improve plant survival.

Specific leaf area (SLA) is one of the most important plant functional traits. It integrates multiple functions and reflects strategies of plants to obtain resources. How plants employ different strategies (e.g., through SLA) to respond to dynamic environmental conditions remains poorly understood. This study aimed to explore the spatial variation in SLA and its divergent adaptation through the lens of biogeographic patterns, evolutionary history, and short-term responses. SLA data for 5424 plant species from 76 natural communities in China were systematically measured and integrated with meta-analysis of field experiments (i.e., global warming, drought, and nitrogen addition). The mean value of SLA across all species was 21.8m2 kg-1 , ranging from 0.9 to 110.2m2 kg-1 . SLA differed among different ecosystems, temperature zones, vegetation types, and functional groups. Phylogeny had a weak effect on SLA, but plant species evolved toward higher SLA. Furthermore, SLA responded nonlinearly to environmental change. Unexpectedly, radiation was one of the main factors determining the spatial variation in SLA on a large scale. Conversely, short-term manipulative experiments showed that SLA increased with increased resource availability and tended to stabilize with treatment duration. However, different species exhibited varying response patterns. Overall, variation in long-term adaptation of SLA to environmental gradients and its short-term response to resource pulses jointly improve plant adaptability to a changing environment. Overall SLA-environment relationships should be emphasized as a multidimensional strategy for elucidating environmental change in future research.

Effects of Precipitation Change and Nitrogen and Phosphorus Additions on Traits and Abundance of Potentilla anserina in an Alpine Meadow

Changes in precipitation patterns and eutrophication can cause changes in plant traits and abundance, potentially affecting plant community structure and functions. Here, we studied responses of traits and abundance of Potentilla anserina to precipitation change and nitrogen (N) and phosphorus (P) additions, and the effect of traits on its abundance in an alpine meadow of the Qinghai-Tibet Plateau. We found that precipitation change and N and P additions significantly affected the mean value of traits such as specific leaf area (SLA), leaf dry matter content (LDMC), single leaf area, plant height and individual size, while only P addition significantly affected intraspecific variation of SLA and individual size. Increased precipitation and N and P additions shifted plant traits to more resource acquisitive, and increased plant abundance. Responses of plant traits to P addition were larger than that of N addition. Plant abundance was mainly affected by precipitation, and was limited by N or P dependent on precipitation conditions. In conclusions, our research shows that P. anserina can respond to environmental changes by changing its traits to improve its adaptability, potentially affecting community structure and ecosystem functions.

Leaf Traits Predict Water‐Use Efficiency in U.S. Pacific Northwest Grasslands Under Rain Exclusion Treatment

AbstractDoes drought stress in temperate grasslands alter the relationship between plant structure and function? Here we report data from an experiment focusing on growth form and species traits that affect the critical functions of water‐ and nutrient‐use efficiency in prairie and pasture plant communities. A total of 139 individuals of 12 species (11 genera and four families) were sampled in replicated plots maintained for three years across a 520 km latitudinal gradient in the Pacific Northwest, USA. Rain exclusion did not alter the interspecific relationship between foliar traits and stoichiometry or intrinsic water‐use efficiency (iWUE). Rain exclusion reduced iWUE in grasses, an effect was primarily species‐specific, although leaf morphology, life history strategy, and phylogenetic distance predicted iWUE for all 12 species when analyzed together. Variation in specific leaf area explained most of the variation in iWUE between different functional groups, with annual forbs and annual grasses at opposite ends of the resource‐use spectrum. Our findings are consistent with expected trait‐driven tradeoffs between productivity and resource‐use efficiency, and provide insight into strategies for the sustainable use and conservation of temperate grasslands.

Variation in leaf traits among and within dominant shrubs of contrasting Pine Barrens habitats

Leaf anatomical and physiological traits influence rates of individual plant and ecosystem function. Although leaf trait variation among different species can be largely attributed to broad climatic gradients, variation at finer spatial scales and within species is less known for many habitats and taxa. Pine barrens are particularly understudied despite regional and ecological significance in northeastern North America. We assessed variation in specific leaf area (SLA), leaf dry matter content (LDMC) and leaf thickness (Lth) for three dominant understory shrub species in the New Jersey Pine Barrens. We hypothesized that traits would vary among and within species along an environmental gradient from upland forest to lowland bog. We further hypothesized that SLA would be larger and LDMC and Lth would be smaller in the upland forest than in the wetter, more nutrient-poor bog, reflective of stress tolerant plant strategies. Unexpectedly, all three species had smaller SLA and larger Lth in the forest than in the bog. LDMC was larger in the forest for Black Huckleberry (Gaylussacia baccata) and Sweetgale (Myrica gale) but smaller in the forest for Lowbush Blueberry (Vaccinium angustifolium). A trade-off between SLA and Lth appears constrained and potentially adaptive in multiple ecological contexts. However, trait-trait correlations involving LDMC were inconsistent across scales. These shrub species may be responding in similar ways to dominant environmental variables in the bog, while multiple environmental variables in the forest may promote species-specific responses and divergent trait correlations. Differences in light and water availability may be influencing leaf trait variation more than soil nutrient availability, while soil pH may be an additional driver of variation in leaf functional traits for these understory temperate shrubs.

Intra‐ and interspecific variability of specific leaf area mitigate the reduction of community stability in response to warming and nitrogen addition

Global environmental changes are reducing the diversity and affecting the functioning of natural ecosystems as well as their ability to reliably provide ecosystem functions and services to mankind. Many studies have shown that a greater plant diversity can stabilize community productivity against environmental fluctuations. However, most of these studies focused on plant species richness, thus overlooking the potential role of functional traits in stabilizing community productivity against environmental fluctuations. Whether and how functional trait mean and variability influence community stability in response to environmental changes and their relative contributions to community stability are largely unknown. Here, we used a 10‐year experiment to investigate the role of species richness, as well as functional mean and intra‐ and interspecific variability of specific leaf area (SLA) of plants within‐ and among communities in driving community stability in response to nitrogen (N) addition and warming. We found that both N addition and warming reduced the temporal stability of community productivity by reducing species richness and its contribution to species asynchrony and species stability. In contrast, changes in the mean and variability of SLA in response to N addition and warming mitigated the reduction of community stability. Specifically, N addition reduced variation in SLA both by reducing interspecific differences in SLA within communities and differences in mean values of SLA among communities. Warming increased intraspecific differences in SLA among communities, leading to higher species stability that partly buffered the reduction of community stability. Our study demonstrates the role of trait mean and variability in mitigating the reduction of community stability in response to two pervasive global environmental changes. Gaining a deeper understanding of the processes linking global changes and the stability of our ecosystems requires integrating both trait mean values and trait variability.

Variations in leaf and stem traits across two elevations in subtropical forests.

Understanding the variations in plant traits across elevations may provide valuable insights into the species structure and function of forests and their responses to climate change. To explore the patterns of trait variation across elevations, we analysed 14 leaf and stem traits associated with resource acquisition and stress tolerance in Schima superba Gardner & Champion, Castanopsis chinensis (Sprengel) Hance, and Pinus massoniana Lambert trees at two elevations in a subtropical forest in southern China. Wood density increased, whereas crown width, leaf water potential at 0700 hours (Ψ L-0700 ), and leaf δ 18 O decreased in high-elevation plants. Vessel diameter, daily maximum sap flux density, leaf δ 13 C, and leaf C and N concentrations per unit mass were comparable across elevations. We found species-specific variations in specific leaf area, midday leaf water potential, and leaf P concentration across elevations. Decreasing crown width with increasing elevation was associated with decreasing leaf δ 18 O and Ψ L-0700 , suggesting that higher stomatal conductance may moderate the loss of carbon assimilation. We elucidated the adaptive strategies of plants in response to environmental change, and showed that physiological traits varied in coordination with structural traits. Future studies incorporating multi-dimensional trait analyses can improve our understanding of the responses of forest ecosystems to climate change and global warming.

中国温带典型森林植物比叶面积的空间格局及其影响因素

比叶面积（SLA）能够反映植物自身生长对策、对光能的捕获能力、对外界环境变化的适应和对可利用资源的分配策略，对植物的生境适应状况和群落的自然演替程度也起到重要的指示作用。为了深入了解温带森林植物SLA随空间变化的变异特征及其影响因素，跨越1200 km选取中国东北部12个典型温带森林（寒温带兴安落叶松林、温带红松针阔混交林和暖温带落叶阔叶栎林），通过对样地内物种进行系统性的调查，分析了植物SLA在空间上的变化规律及环境因素的影响效应。结果显示：中国温带森林植物SLA范围为2.02-99.65 m²/kg（均值为34.18 m²/kg），其中乔木SLA范围为2.02-58.74 m²/kg（均值为21.32 m²/kg），灌木SLA范围为2.88-99.65 m²/kg（均值为31.60 m²/kg），草本SLA范围为4.66-98.53 m²/kg（均值为38.75 m²/kg）。SLA在不同森林类型之间差异显著，具体表现为：温带红松针阔混交林 > 暖温带落叶阔叶栎林 > 寒温带兴安落叶松林。SLA受到气候因素和土壤因素的影响，其中随着年均降水、土壤碳氮含量的增加显著增加；随着年均温度、紫外辐射、光合有效辐射，呈先增加后降低的趋势；随着土壤碳氮比的增加显著降低。以上结果揭示了温带森林SLA的空间变异规律，可以为将来区域乃至全球尺度下植物功能性状研究提供理论支撑。;Specific leaf area (SLA) can reflect plant growth strategies, the ability to capture light energy, adaptation to changing environments, and allocation strategies of available resources. SLA critically plays an important role in indicating plant adaptation to their habitat and the degree of community succession as well. SLA is a complex of plant leaf area and weight, and is defined as the ratio of single leaf area to its dry weight. The current research on SLA in temperate forests mostly focused on the spatial variation of dominant species or just a certain tree, and the vertical variation among forests. However, the systematic investigation of the entire community was rarely carried out. This research was conducted in the northeast of China in which covered three forest types. These types were respectively cold-temperate Larix gmelinii forests, temperate broad-leaved Korean pine mixed forests and warm-temperate deciduous broad-leaved Oak forests. This study selected 12 typical temperate forests spanning 1200 km, and performed a systematic survey to explore the spatial patterns and influencing factors of plant SLA. The results showed that the SLA of China's temperate forest plants ranged from 2.02 to 99.65 m²/kg (mean, 34.18 m²/kg). Specifically, the SLA range was 2.02-58.74 m²/kg for trees (mean, 21.32 m²/kg), 2.88-99.65 m²/kg for shrubs (mean, 31.60 m²/kg), and 4.66-98.53 m²/kg for herbs (mean, 38.75 m²/kg). Plant SLA significantly differed among different forest types, with the greatest SLA in temperate broad-leaved Korean pine mixed forest, followed by warm-temperate deciduous broad-leaved Oak forest and cold-temperate Larix gmelinii forest. Significant latitudinal pattern of plant SLA was observed. As latitude increased, SLA initially increased and then decreased. SLA was found to be affected by both climatic and soil factors. SLA increased significantly with the increasing mean annual precipitation, soil carbon content, and soil nitrogen content. However, with the increase of mean annual temperature, ultraviolet radiation, and photosynthetically active radiation, SLA first increased, followed by a decline. Moreover, it decreased significantly as the soil carbon to nitrogen ratio increased. In summary, this study reveals the spatial variation of SLA in temperate forests, and it can not only provide a new perspective for the study on the response of different forest ecosystems to global change, but also provide an important theoretical basis for the development of the adaptive management of natural forests in China. Moreover, it also provides a theoretical support for future studies on plant functional traits at the regional and global scales.

Variations in Leaf Traits Modulate Plant Vegetative and Reproductive Phenological Sequencing Across Arid Mediterranean Shrublands.

Structural and nutrient traits of a leaf are important for understanding plant ecological strategies (e.g., drought avoidance). We studied the specific leaf area (SLA), leaf carbon content (LCC), leaf nitrogen content (LNC), leaf phosphorous content (LPC), and the phenophase sequence index (PSI) in 126 Mediterranean perennial species from predesert (SMS) and semiarid (SaMS) to subalpine (SAS), alpine cushion (AcS), and oro-Mediterranean (AjS) shrublands, which represent eight functional groups (evergreen and deciduous trees, evergreen large and half shrubs, deciduous large and half shrubs, succulents and perennial herbs). We analyzed the variation and relationships between leaf traits and PSI among shrublands, functional groups, and within species with drought-avoidance mechanisms. SLA variation of 20–60% could be ascribed to differences between functional groups and only 38–48% to different shrublands increasing from the predesert to the alpine. Alpine species display low PSI and N:P and high SLA, LNC, LPC, LCC, and C:N. On the contrary, predesert and semiarid showed high PSI and low SLA. SLA mediates the vegetative and reproductive phenological plant sequencing, high SLA is often associated with the overlapping in growth and reproductive phenophases with a seasonal reduction of vegetative growth, whereas low SLA is associated with vegetative and reproductive sequencing and a seasonal extension of vegetative growth. Species with drought-avoidance mechanisms (e.g., semideciduous species) contribute to an increase in the mean values of the SLA and LNC because these species show similar leaf and phenological patterns as the deciduous (high SLA and LNC and low PSI). The N:P indicates that only the alpine shrublands could present P limitations. The positive correlations between SLA and LPC and LNC and LPC (leaf economic spectrum) and the negative correlation between SLA and C:N were consistently maintained in the studied arid Mediterranean shrublands.

Functional diversity and identity of plant genotypes regulate rhizodeposition and soil microbial activity.

Our understanding of the linkages between plant diversity and soil carbon and nutrient cycling is primarily derived from studies at the species level, while the importance and mechanisms of diversity effects at the genotype level are poorly understood. Here we examine how genotypic diversity and identity, and associated variation in functional traits, within a common grass species, Anthoxanthum odoratum, modified rhizodeposition, soil microbial activity and litter decomposition. Root litter quality was not significantly affected by plant genotypic diversity, but decomposition was enhanced in soils with the legacy of higher genotypic diversity. Plant genotypic diversity and identity modified rhizodeposition and associated microbial activity via two independent pathways. Plant genotypic diversity enhanced soil functioning via positive effects on variation in specific leaf area and total rhizodeposition. Genotype identity affected both rhizodeposit quantity and quality, and these effects were mediated by differences in mean specific leaf area, shoot mass and plant height. Rhizodeposition was more strongly predicted by aboveground than belowground traits, suggesting strong linkages between photosynthesis and root exudation. Our study demonstrates that functional diversity and identity of plant genotypes modulates belowground carbon supply and quality, representing an important but overlooked pathway by which biodiversity affects ecosystem functioning.

Divergent long- and short-term responses to environmental gradients in specific leaf area of grassland species

Specific leaf area (SLA) is one of the most widely researched plant traits, integrating multiple functions and reflecting plant resource acquisition strategies. However, the underlying mechanisms that drive temporal and spatial variation in SLA are still not clear. We used a combination of transect investigation and short-term manipulative experiments to identify patterns of variation in SLA over naturally occurring and simulated environmental gradients. We measured SLA of 631 species from the Tibetan (TP), Loess (LP), and Mongolian Plateaus (MP), where plant growth is limited by temperature, soil nutrient availability, and precipitation, respectively. In each plateau, we established an east–west transect of 10 sites, including meadow steppe (meadow), typical steppe (steppe), and desert steppe (desert) habitats. We further measured SLA of 15 species from the TP and MP, using manipulative experiments that altered temperature, precipitation and soil nitrogen levels. SLA ranged from 0.9 to 58.90 m2 kg−1 on average, and was significantly larger in desert than in meadow and steppe, except in the LP. Plants from the TP had significantly lower SLA than those from the LP and MP, but variation in SLA across the environmental gradients was small and consistent across the three plateaus. Conversely, the short-term manipulative experiments revealed that SLA increased with resource availability. Shifts in SLA across the broad environmental gradients was relatively small, and SLA had high sensitivity to short-term resource enrichment, suggesting that grassland species in these plateaus express divergent short- (acclimation) and long-term responses (adaptation) to the environment, which may be an important strategy for coping with ongoing shifts in climate.

The Effects of Tree and Stand Traits on the Specific Leaf Area in Managed Scots Pine Forests of Different Ages

The purpose of this study was to understand the relationships between stand structure (tree size, volume, biomass, social position, stand density) and the variability of specific leaf area (SLA) at the stand level, which could improve forest management modeling. The study was carried out on 100 trees selected from 10 stands of Scots pine located in northwestern Poland. The stands had been established in a similar way and were similarly managed. Five mid-aged (51–60 years) and five mature (81–90 years) pure Scots pine stands were selected. To obtain the SLA index, we used the direct method, which involves scanning ca. 50 needles from each part of the tree crown. The average SLA was from 4.65 to 6.62 m2·kg−1 and differed significantly according to the part of the crown measured (p &lt; 0.0001) and the tree age (p &lt; 0.0001). The smallest SLA was in the upper part of the crown and the largest in the lower part of the crown, which is in line with the known relation to the light exposure of needles. Mid-aged stands of Scots pine have higher SLA values than mature ones. Dominant trees in mid-aged stands have a lower SLA than more shaded intermediate ones, which is probably due to the different lighting conditions within the canopy. No clear relationship is observed between the stand density and the SLA.

Reprint of: Drivers of within-tree leaf trait variation in a tropical planted forest varying in tree species richness

In plant ecology, community-weighted trait means are often used as predictors for ecosystem functions. More recently, also within-species trait variation has been confirmed to contribute to ecosystem functioning. We here go even further and assess within-individual trait variation, assuming that every leaf in a plant individually adjusts to its micro-environment. Using forest plots varying in tree species richness (Sardinilla experiment, Panama), we analysed how leaf traits within individual trees vary along the vertical crown gradient. Furthermore, we tested whether niche partitioning in mixed stands results in a decrease of within-species leaf trait variation and whether niche partitioning can be also observed at the level of individual trees. We focused on leaf traits that describe the growth strategy along the conservative-acquisitive spectrum of growth. We found a decrease in within-species variation of specific leaf area (SLA) with increasing neighbourhood species richness. Both sampling height and local neighbourhood richness contributed to explaining within-species leaf trait variation, which however, varied in importance among different species and traits. With increasing sampling height, leaf dry matter content (LDMC), carbon to nitrogen ratio and lignin content increased, while leaf nitrogen concentration (leaf N), SLA, cellulose and hemicellulose decreased. Variation in leaf N decreased with increasing neighbourhood species richness, while the magnitude of within-individual variation of most traits was unaffected by neighbourhood species richness. Our results suggest an increased niche partitioning with increasing species richness both in a plant community and at the level of individual plants. Our findings highlight the importance of including within-individual trait variation to understand biodiversity-ecosystem functioning relationships.

Active layer depth and soil properties impact specific leaf area variation and ecosystem productivity in a boreal forest.

Specific leaf area (SLA, leaf area per unit dry mass) is a key canopy structural characteristic, a measure of photosynthetic capacity, and an important input into many terrestrial process models. Although many studies have examined SLA variation, relatively few data exist from high latitude, climate-sensitive permafrost regions. We measured SLA and soil and topographic properties across a boreal forest permafrost transition, in which dominant tree species changed as permafrost deepened from 54 to >150 cm over 75 m hillslope transects in Caribou-Poker Creeks Research Watershed, Alaska. We characterized both linear and threshold relationships between topographic and edaphic variables and SLA and developed a conceptual model of these relationships. We found that the depth of the soil active layer above permafrost was significantly and positively correlated with SLA for both coniferous and deciduous boreal tree species. Intraspecific SLA variation was associated with a fivefold increase in net primary production, suggesting that changes in active layer depth due to permafrost thaw could strongly influence ecosystem productivity. While this is an exploratory study to begin understanding SLA variation in a non-contiguous permafrost system, our results indicate the need for more extensive evaluation across larger spatial domains. These empirical relationships and associated uncertainty can be incorporated into ecosystem models that use dynamic traits, improving our ability to predict ecosystem-level carbon cycling responses to ongoing climate change.

Sampling protocols of specific leaf area for improving accuracy of the estimation of forest leaf area index

Litterfall collection is a non-destructive direct method to estimate forest leaf area index (LAI) and validate indirect LAI products. However, the potential errors associated with the variation in specific leaf area (SLA) are rarely explored. Here, we measured the SLA of leaf litter for each tree species in a temperate deciduous forest using the litterfall collection method from 2012 to 2018, and assessed the spatial and temporal variation in SLA and its consequence on LAI estimates. The results showed that the spatial and temporal variation in SLA for the 10 major species across the seven years ranged from 0.8% to 24.3%, with the variation across the nine permanent plots (20 m × 30 m) being higher than that within plot (five 1-m2 traps), and interannual higher than seasonal. Across the 63 plot-years, the 10 simplified SLA sampling protocols introduced the errors in LAI by −12.4% to 22.2% relative to the reference protocol (sampling leaves from each trap at each collection date for the major species). Applying the SLA obtained from one trap in each plot, one plot, at the leaf fall peak for each year, or for a single year induced the errors in LAI by –3.7% to 2.9%, −6.4% to 14.7%, −9.2% to 4.7%, and −8.1% to 8.6%, respectively. Considering the trade-off between inter-plot and interannual variation, we recommend sampling from one-trap for each plot at the leaf-fall peak for measuring SLA and quantifying the spatial pattern of LAI, and sampling each species in the typical plot for measuring SLA and monitoring the temporal fluctuation in LAI. The tight relationship between the MODIS and ground reference LAI across the seven years (R2 = 0.77) validated the use of MODIS LAI to study the long-term change in forest LAI. These findings help to establish a standardized protocol for long-term accurately measuring forest LAI with the litterfall collection.

Drivers of within-tree leaf trait variation in a tropical planted forest varying in tree species richness

In plant ecology, community-weighted trait means are often used as predictors for ecosystem functions. More recently, also within-species trait variation has been confirmed to contribute to ecosystem functioning. We here go even further and assess within-individual trait variation, assuming that every leaf in a plant individually adjusts to its micro-environment. Using forest plots varying in tree species richness (Sardinilla experiment, Panama), we analysed how leaf traits within individual trees vary along the vertical crown gradient. Furthermore, we tested whether niche partitioning in mixed stands results in a decrease of within-species leaf trait variation and whether niche partitioning can be also observed at the level of individual trees. We focused on leaf traits that describe the growth strategy along the conservative-acquisitive spectrum of growth. We found a decrease in within-species variation of specific leaf area (SLA) with increasing neighbourhood species richness. Both sampling height and local neighbourhood richness contributed to explaining within-species leaf trait variation, which however, varied in importance among different species and traits. With increasing sampling height, leaf dry matter content (LDMC), carbon to nitrogen ratio and lignin content increased, while leaf nitrogen concentration (leaf N), SLA, cellulose and hemicellulose decreased. Variation in leaf N decreased with increasing neighbourhood species richness, while the magnitude of within-individual variation of most traits was unaffected by neighbourhood species richness. Our results suggest an increased niche partitioning with increasing species richness both in a plant community and at the level of individual plants. Our findings highlight the importance of including within-individual trait variation to understand biodiversity-ecosystem functioning relationships.