Within‐species patterns challenge our understanding of the leaf economics spectrum

The utility of plant functional traits for predictive ecology relies on our ability to interpret trait variation across multiple taxonomic and ecological scales. Using extensive data sets of trait variation within species, across species and across communities, we analysed whether and at what scales leaf economics spectrum (LES) traits show predicted trait–trait covariation. We found that most variation in LES traits is often, but not universally, at high taxonomic levels (between families or genera in a family). However, we found that trait covariation shows distinct taxonomic scale dependence, with some trait correlations showing opposite signs within vs. across species. LES traits responded independently to environmental gradients within species, with few shared environmental responses across traits or across scales. We conclude that, at small taxonomic scales, plasticity may obscure or reverse the broad evolutionary linkages between leaf traits, meaning that variation in LES traits cannot always be interpreted as differences in resource use strategy.     

Irradiance, temperature and rainfall influence leaf dark respiration in woody plants: evidence from comparisons across 20 sites

Leaf dark respiration (R) is one of the most fundamental physiological processes in plants and is a major component of terrestrial CO2 input to the atmosphere. Still, it is unclear how predictably species vary in R along broad climate gradients. Data for R and other key leaf traits were compiled for 208 woody species from 20 sites around the world. We quantified relationships between R and site climate, and climate-related variation in relationships between R and other leaf traits. Species at higher-irradiance sites had higher mean R at a given leaf N concentration, specific leaf area (SLA), photosynthetic capacity (Amass) or leaf lifespan than species at lower-irradiance sites. Species at lower-rainfall sites had higher mean R at a given SLA or Amass than species at higher-rainfall sites. On average, estimated field rates of R were higher at warmer sites, while no trend with site temperature was seen when R was adjusted to a standard measurement temperature. Our findings should prove useful for modelling plant nutrient and carbon budgets, and for modelling vegetation shifts with climate change.     

Global meta-analysis shows that relationships of leaf mass per area with species shade tolerance depend on leaf habit and ontogeny

It was predicted that relationships of leaf mass per area (LMA) with juvenile shade tolerance will depend on leaf habit, and on whether species are compared at a common age as young seedlings, or at a common size as saplings. A meta-analysis of 47 comparative studies (372 species) was used to test predictions, and the effect of light environment on this relationship. The LMA of evergreens was positively correlated with shade tolerance, irrespective of ontogeny or light environment. The LMA of young seedlings (相似文献   

江西九连山常绿阔叶林主要树种叶建成消耗的比较

叶建成消耗（或叶构建消耗）是指构建单位质量（面积）叶片所需要的葡萄糖当量,是植物碳收获过程中必要的成本投资．反映植物在叶片水平上的能量和碳分配策略。基于热值、灰份含量、叶氮含量和比叶面积的测定,估算了江西九连山常绿阔叶林16个主要树种的叶建成消耗,分析比较了不同树种、不同叶寿命、比叶面积和冠层不同部位对叶建成消耗的影响。结果表明,热值组分的变化对单位质量叶建成消耗的影响最大;树种间单位质量叶建成消耗的差异不大（变化率仅3％～6％）,但其单位面积叶建成消耗则存在显著差异（变化率达27％～28％）;热值和单位质量叶建成消耗与比叶面积密切相关,而比叶面积随取样高度呈显著的降低趋势,最终导致热值及单位质量和单位面积的叶建成消耗均随取样高度呈显著的增加趋势。研究结果指出：（1）较高的叶建成消耗是由于植物叶片内含有较高能量投资的化学组分;（2）叶建成消耗和比叶面积随取样高度的变化综合反映了植物对光照和水分资源垂直梯度变化的可塑性适应。     

Linking leaf and root trait syndromes among 39 grassland and savannah species

Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.     

东北次生林主要树种比叶面积对光照强度的响应

通过测定不同光照条件下（15％透光、30％透光、60％透光、全光）东北次生林主要树种（红松、红皮云杉、胡桃楸、水曲柳和黄波罗）的比叶面积,探讨了不同树种对光照变化的响应与适应。结果表明:不同树种比叶面积存在明显的差异,尤其是常绿针叶树种与落叶树种之间差异显著（P<0.001）,指示不同功能型树种存在不同的生存策略;随着光照强度降低,所有树种比叶面积呈现增加的趋势,尤其是弱光照（15％）和低光照（30％）相对于全光和中光照（60％）差异显著（P<0.05）;不同树种比叶面积随光照强度的变化幅度明显不同,且变化最显著的光照区域也有所不同;不同年龄红松比叶面积的变化趋势一致,但变幅不同,4年生幼苗对光照响应则更为明显。本研究结果表明,所有树种均能通过改变叶形态来适应光照条件的变化,在低光条件下通过增加单位质量叶面积,提高叶片光捕获能力;但各树种对光照强度变化的不同响应则表明树种之间不同的光适应机制,这对森林生态系统内部树种的共生具有重要意义;年龄较小的幼苗对光照的强响应指示其在植物更新过程中叶片水平上生理生态学机理研究中的重要性。     

Plasticity in above‐ and belowground resource acquisition traits in response to single and multiple environmental factors in three tree species

Functional trait plasticity is a major component of plant adjustment to environmental stresses. Here, we explore how multiple local environmental gradients in resources required by plants (light, water, and nutrients) and soil disturbance together influence the direction and amplitude of intraspecific changes in leaf and fine root traits that facilitate capture of these resources. We measured population‐level analogous above‐ and belowground traits related to resource acquisition, i.e. “specific leaf area”–“specific root length” (SLA–SRL), and leaf and root N, P, and dry matter content (DMC), on three dominant understory tree species with contrasting carbon and nutrient economics across 15 plots in a temperate forest influenced by burrowing seabirds. We observed similar responses of the three species to the same single environmental influences, but partially species‐specific responses to combinations of influences. The strength of intraspecific above‐ and belowground trait responses appeared unrelated to species resource acquisition strategy. Finally, most analogous leaf and root traits (SLA vs. SRL, and leaf versus root P and DMC) were controlled by contrasting environmental influences. The decoupled responses of above‐ and belowground traits to these multiple environmental factors together with partially species‐specific adjustments suggest complex responses of plant communities to environmental changes, and potentially contrasting feedbacks of plant traits with ecosystem properties. We demonstrate that despite the growing evidence for broadly consistent resource‐acquisition strategies at the whole plant level among species, plants also show partially decoupled, finely tuned strategies between above‐ and belowground parts at the intraspecific level in response to their environment. This decoupling within species suggests a need for many species‐centred ecological theories on how plants respond to their environments (e.g. competitive/stress‐tolerant/ruderal and response‐effect trait frameworks) to be adapted to account for distinct plant‐environment interactions among distinct individuals of the same species and parts of the same individual.     

利用叶片荧光参数估算油蒿灌丛群落生态系统生产力

生态系统生产力（GEP）在全球碳循环中具有重要意义,但其准确估算仍然是一个挑战。近年来,叶绿素荧光和冠层GEP的关联成为生态学的研究热点,关系尚不清楚且存在广泛争议。于2015年对宁夏盐池毛乌素沙地荒漠灌木油蒿（Artemisia ordosica）灌丛生态系统碳交换（NEE）特征进行连续观测,使用多通道荧光监测仪对通量贡献区内油蒿叶片的实时荧光（F_s）和光下荧光（F_m''）进行原位连续监测,叶面积指数（LAI）、归一化植被指数（NDVI）以及环境因子同步观测。利用光合有效辐射（PAR）、LAI和实际光化学效率（Φ_PSII）等参数构建基于叶绿素荧光的生态系统生产力（GEP_ChlF）模型,探究叶片和冠层尺度不同参数对环境因子波动的响应,比较分析GEP_ChlF和基于涡度相关法监测生态系统生产力（GEP_EC）相关性及GEP_ChlF的适用性。研究发现,Φ_PSII和NEE日变化规律一致,Φ_PSII、GEP_ChlF和GEP_EC由PAR控制,受空气温度（T_a）和饱和水汽压差（VPD）调控,土壤含水量（SWC）和Φ_PSII呈显著正相关（P<0.01）。PAR处于400-800 μmol m^-2 s^-1时,GEP_ChlF与GEP_EC线性关系最优,斜率为0.627（R²=0.67,P<0.01）;弱光下GEP_ChlF的低估可能是由于冠层实际光能拦截率高造成;强光下GEP_ChlF显著高于GEP_EC,呈非线性关系,可能是GEP_EC基于夜间温度敏感性所拟合的生态系统呼吸（R_e）无法预测光呼吸部分所导致。GEP_ChlF具有荧光参数的特性,对环境波动更加敏感,和环境因子相关性优于GEP_EC。结果表明,高辐射、极端温度、高蒸腾和干旱是限制叶片和冠层尺度下油蒿光合过程的主要胁迫因素。本研究构建的荒漠生态系统光合过程模型GEP_ChlF能够替代GEP_EC作为一个良好的冠层尺度生态参数,所提出的空间尺度上推方法可为促进区域可持续发展提供数据支撑和决策参考。     

Biogeographic constraints on the world‐wide leaf economics spectrum

Aim The world‐wide leaf economic spectrum (LES) describes tight coordination of leaf traits across global floras, reported to date as being largely independent of phylogeny and biogeography. Here, we present and test an alternative, historical perspective that predicts that biogeography places significant constraints on global trait evolution. These hypothesized constraints could lead to important deviations in leaf trait relationships between isolated floras that were influenced by different magnitudes of genetic constraint and selection. Location Global, including floristic regions of the Northern and Southern Hemispheres, eastern North America, East Asia (EAS), the Hawaiian Islands and tropical mainland floras. Methods We use a large leaf‐trait database (GLOPNET) and species native distribution data to test for variation in leaf trait relationships modulated by floristic region, controlling for climatic differences. Standardized major axis analyses were used to evaluate biogeographic effects on bivariate relationships between LES traits, including relationships of photosynthetic capacity and dark respiration rate (A_mass–R_d‐mass), leaf lifespan and mass per area ratio (LL–LMA), and photosynthetic capacity and nitrogen content (A_mass–N_mass). Results Independent of climate or biome, floras of different evolutionary histories exhibited different leaf trait allometries. Floras of the Northern Hemisphere exhibited greater rates of return on resource investment (steeper slopes for the trait relationships analysed), and the more diverse temperate EAS flora exhibited greater slopes or intercepts in leaf trait relationships, with the exception of the A_mass–N_mass relationship. In contrast to our hypothesis, plants of the floristically isolated Hawaiian Islands exhibited a similar A_mass–N_mass relationship to those of mainland tropical regions. Main conclusions Differences in leaf trait allometries among global floristic regions support a historical perspective in understanding leaf trait relationships and suggest that independent floras can exhibit different tradeoffs in resource capture strategies.     

Simulation of leaf area development based on dry matter partitioning and specific leaf area for cut chrysanthemum

This work aims to predict time courses of leaf area index (LAI) based on dry matter partitioning into the leaves and on specific leaf area of newly formed leaf biomass (SLA(n)) for year-round cut chrysanthemum crops. In five glasshouse experiments, each consisting of several plant densities and planted throughout the year, periodic destructive measurements were conducted to develop empirical models for partitioning and for SLA(n). Dry matter partitioning into leaves, calculated as incremental leaf dry mass divided by incremental shoot dry mass between two destructive harvests, could be described accurately (R(2 )= 0.93) by a Gompertz function of relative time, R(t). R(t) is 0 at planting date, 1 at the start of short-days, and 2 at final harvest. SLA(n), calculated as the slope of a linear regression between periodic measurements of leaf dry mass (LDM) and LAI, showed a significant linear increase with the inverse of the daily incident photosynthetically active radiation (incident PAR, MJ m(-2 )d(-1)), averaged over the whole growing period, the average glasshouse temperature and plant density (R(2 )= 0.74). The models were validated by two independent experiments and with data from three commercial growers, each with four planting dates. Measured shoot dry mass increase, initial LAI and LDM, plant density, daily temperature and incident PAR were input into the model. Dynamics of LDM and LAI were predicted accurately by the model, although in the last part of the cultivation LAI was often overestimated. The slope of the linear regression of simulated against measured LDM varied between 0.95 and 1.09. For LAI this slope varied between 1.01 and 1.12. The models presented in this study are important for the development of a photosynthesis-driven crop growth model for cut chrysanthemum crops.     

Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.     

Variability in leaf optical properties among 26 species from a broad range of habitats

Leaves from 26 species with growth forms from annual herbs to trees were collected from open, intermediate, and shaded understory habitats in Mississippi and Kansas, USA. Leaf optical properties including reflectance, transmittance, and absorptance in visible and near infrared (NIR) wavelengths were measured along with leaf thickness and specific leaf mass (SLM). These leaf properties and internal light scattering have been reported to vary with light availability in studies that have focused on a limited number of species. Our objective was to determine whether these patterns in leaf optics and light availability were consistent when a greater number of species were evaluated. Leaf thickness and SLM varied by tenfold among species sampled, but within-habitat variance was high. Although there was a strong trend toward thicker leaves in open habitats, only SLM was significantly greater in open vs. understory habitats. In contrast, leaf optical properties were strikingly similar among habitats. Reflectance and reflectance/transmittance in the NIR were used to estimate internal light scattering and there were strong relationships (r1 > 0.65) between these optical properties and leaf thickness. We concluded that leaf thickness, which did not vary consistently among habitats, was the best predictor of NIR reflectance and internal light scattering. However, because carbon allocation to leaves was lower in understory species (low SLM) yet gross optical properties were similar among all habitats, the energy investment by shade leaves required to achieve optical equivalence with sun leaves was lower. Differences in leaf longevity and growth form within a habitat may help explain the lack of consistent patterns in leaf optics as the number of species sampled increases.     

Effects of hydraulic architecture and spatial variation in light on mean stomatal conductance of tree branches and crowns

In a Pinus taeda L. (loblolly pine) plantation, we investigated whether the response to vapour pressure deficit (D) of canopy average stomatal conductance (G(S)) calculated from sap flux measured in upper and lower branches and main stems follows a hydraulically modelled response based on homeostasis of minimum leaf water potential (Psi(L)). We tested our approach over a twofold range of leaf area index (L; 2-4 m(2) m(-2)) created by irrigation, fertilization, and a combination of irrigation and fertilization relative to untreated control. We found that G(S) scaled well from leaf-level porometery [porometry-based stomatal conductance (g(s))] to branch-estimated and main stem-estimated G(S). The scaling from branch- to main stem-estimated G(S) required using a 45 min moving average window to extract the diurnal signal from the large high-frequency variation, and utilized a light attenuation model to weigh the contribution of upper and lower branch-estimated G(S). Our analysis further indicated that, regardless of L, lower branch-estimated G(S) represented most of the main stem-estimated G(S) in this stand. We quantified the variability in both upper and lower branch-estimated G(S) by calculating the SD of the residuals from a moving average smoothed diurnal. A light model, which incorporated penumbral effects on vertical distribution of direct light, was employed to estimate the variability in light intensity at each canopy level in order to explain the increasing SD of both upper and lower branch-estimated G(S) with light. The results from the light model showed that the upper limit of the variability in individual branch-estimated G(S) could be attributed to incoming light, but not the variation below that upper limit. A porous medium model of water flow in trees produced a pattern of variation below the upper limit that was consistent with the observed variability in branch-estimated G(S). Our results indicated that stems acted to buffer leaf- and branch-level variation and might transmit a less-variable water potential signal to the roots.     

Relationships between sclerophylly,leaf biomechanical properties and leaf anatomy in some Australian heath and forest species

ABSTRACT

A previous study of 19 south-east Australian heath and forest species with a range of leaf textures showed that they varied considerably in leaf biomechanical properties. By using an index of sclerophylly derived from botanists' rankings (botanists' sclerophylly index, BSI) we determined that leaves considered by botanists to be sclerophyllous generally had both high strength and work to fracture (particularly in punching and tearing tests), both at the level of leaf and per unit leaf thickness. In the current study we have shown that leaves from the same species also varied considerably in leaf specific mass (46–251 g m^-2), neutral detergent fibre concentration (20–59% on a dry weight basis) and in leaf anatomy. Multiple regression indicated a very strong correlation between BSI and the first two components of a principal components analysis (PCA) of leaf anatomy (R ² = 0.91). In addition, there was strong correlation between the first component of a PCA of the mechanical properties (correlated with BSI) and the two axes derived from anatomical characteristics (R ² = 0.66). The anatomical properties contributing most to the significant component axes were thickness of palisade mesophyll and upper cuticle (axis 1) and percentage fibre (neutral detergent fibre) and lower epidermis thickness (axis 2). However, whether these relationships are causal, or reflect correlations with characteristics not measured in this study, such as vascularization and sclerification, is not clear. At a finer scale, however, there is evidence that there are various ways to be sclerophyllous, both in terms of anatomical and mechanical properties. This is illustrated by comparison of two of the sclerophyllous species, Eucalyptus baxteri and Banksia marginata.     

不同沙丘生境主要植物比叶面积和叶干物质含量的比较

研究了生长在不同沙丘生境中 (流动沙丘 ,半固定沙丘和固定沙丘 ) 2 0个植物种 (10个 1年生植物种和 10个多年生植物种 )的比叶面积 (SL A)和叶干物质含量 (L DMC)的变化 ,并且分析了各个沙丘生境的土壤养分特征。结果表明 ,各个植物种的平均 SL A和 L DMC在植物种之间差异显著 ;多数在两种或 3种沙丘生境均有分布的植物其 SL A在不同沙丘生境之间差异显著 ,但是仅有 6个植物种的 L DMC在不同沙丘生境之间表现出差异 (p<0 .0 5 )。与许多研究结果类似 ,1年生植物的 SL A显著大于多年生植物的 SL A,而且两者之间 L DMC存在一定的差异。 1年生植物 SL A和 L DMC之间相关性不显著 ,但多年生植物SL A和 L DMC之间呈显著负相关。综合所有 2 0个植物种可以发现 ,SL A增大时 ,L DMC有下降的趋势