Links between leaf anatomy and leaf mass per area of herbaceous species across slope aspects in an eastern Tibetan subalpine meadow

Leaf anatomy varies with abiotic factors and is an important trait for understanding plant adaptive responses to environmental conditions. Leaf mass per area (LMA) is a key morphological trait and is related to leaf performance, such as light‐saturated photosynthetic rate per leaf mass, leaf mechanical strength, and leaf lifespan. LMA is the multiplicative product of leaf thickness (LT) and leaf density (LD), both of which vary with leaf anatomy. Nevertheless, how LMA, LT, and LD covary with leaf anatomy is largely unexplored along natural environmental gradients. Slope aspect is a topographic factor that underlies variations in solar irradiation, air temperature, humidity, and soil fertility. In the present study, we examined (1) how leaf anatomy varies with different slope aspects and (2) how leaf anatomy is related to LMA, LD, and LT. Leaf anatomy was measured for 30 herbaceous species across three slope aspects (south‐, west‐, and north‐facing slopes; hereafter, SFS, WFS, and NFS, respectively) in an eastern Tibetan subalpine meadow. For 18 of the 30 species, LMA data were available from previous studies. LD was calculated as LMA divided by LT. Among the slope aspects, the dominant species on the SFS exhibited the highest LTs with the thickest spongy mesophyll layers. The thicker spongy mesophyll layer was related to a lower LD via larger intercellular airspaces. In contrast, LD was the highest on NFS among the slope aspects. LMA was not significantly different among the slope aspects because higher LTs on SFS were effectively offset by lower LDs. These results suggest that the relationships between leaf anatomy and LMA were different among the slope aspects. Mechanisms underlying the variations in leaf anatomy may include different solar radiation, air temperatures, soil water, and nutrient availabilities among the slope aspects.     

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.     

内蒙古荒漠草原植物叶片功能性状关系及其经济谱分析

气候变化和人为干扰导致草原荒漠化加剧, 引发了严重的环境问题。因此, 对荒漠草原植物与环境变化关系的研究愈加迫切, 分析比较荒漠草原不同功能型物种叶片经济谱具有重要意义。该研究通过测定内蒙古荒漠草原生态系统不同功能型植物叶片的光合及叶绿素荧光参数、比叶面积和叶片氮素含量, 验证了荒漠草原植物叶片经济谱的存在, 明确了各功能型植物叶片性状间的关系及其在叶片经济谱中的位置。荒漠草原不同功能型植物叶片性状差异明显, 草本植物的比叶面积(SLA)、单位质量叶氮含量(N_mass)分别是灌木的2.39倍和1.20倍; 一年生植物单位面积最大净光合速率(A_area)、SLA、光合氮利用效率(PNUE)分别是多年生植物的1.93倍、2.13倍和4.24倍; C₄植物的A_area、SLA、PNUE分别是C₃植物的2.25倍、1.73倍和3.61倍。除A_area与单位面积叶氮含量(N_area)、PSII的实际光化学效率(Φ_PSII)与SLA之间不存在显著相关关系外, 叶片性状间存在广泛的相关关系, 且均达到极显著水平。这验证了叶片经济谱在内蒙古荒漠草原植物中也同样存在。进一步分析表明, 一年生植物、草本植物、C₄植物叶片在叶片经济谱中位于靠近薄叶、光合能力强、寿命短的一端; 而多年生植物、灌木、C₃植物叶片靠近厚叶、光合能力弱、寿命长的一端。这说明荒漠草原中不同功能型植物可通过权衡其经济性状间的关系而采取不同的适应策略, 对于荒漠草原生态系统管理具有重要的理论指导意义。     

Ozone effects on wheat in relation to CO2: modelling short‐term and long‐term responses of leaf photosynthesis and leaf duration

A combined stomatal–photosynthesis model was extended to simulate the effects of ozone exposure on leaf photosynthesis and leaf duration in relation to CO₂. We assume that ozone has a short‐term and a long‐term effect on the Rubisco‐limited rate of photosynthesis, A_c. Elevated CO₂ counteracts ozone damage via stomatal closure. Ozone is detoxified at uptake rates below a threshold value above which A_c decreases linearly with the rate of ozone uptake. Reduction in A_c is transient and depends on leaf age. Leaf duration decreases depending on accumulated ozone uptake. This approach is introduced into the mechanistic crop simulation model AFRCWHEAT2. The derived model, AFRCWHEAT2‐O3, is used to test the capability of these assumptions to explain responses at the plant and crop level. Simulations of short‐term and long‐term responses of leaf photosynthesis, leaf duration and plant and crop growth to ozone exposure in response to CO₂ are analysed and compared with experimental data derived from the literature. The model successfully reproduced published responses of leaf photosynthesis, leaf duration, radiation use efficiency and final biomass of wheat to elevated ozone and CO₂. However, simulations were unsatisfactory for cumulative radiation interception which had some impact on the accuracy of predictions of final biomass. There were responses of leaf‐area index to CO₂ and ozone as a result of effects on tillering which were not accounted for in the present model. We suggest that some model assumptions need to be tested, or analysed further to improve the mechanistic understanding of the combined effects of changes in ozone and CO₂ concentrations on leaf photosynthesis and senescence. We conclude that research is particularly needed to improve the understanding of leaf‐area dynamics in response to ozone exposure and elevated CO₂.     

Variation in ecophysiology and carbon economy of invasive and native woody vines of riparian zones in south‐eastern Queensland

Exotic and invasive woody vines are major environmental weeds of riparian areas, rainforest communities and remnant natural vegetation in coastal eastern Australia, where they smother standing vegetation, including large trees, and cause canopy collapse. We investigated, through glasshouse resource manipulative experiments, the ecophysiological traits that might facilitate faster growth, better resource acquisition and/or utilization and thus dominance of four exotic and invasive vines of South East Queensland, Australia, compared with their native counterparts. Relative growth rate was not significantly different between the two groups but water use efficiency (WUE) was higher in the native species while the converse was observed for light use efficiency (quantum efficiency, AQE) and maximum photosynthesis on a mass basis (). The invasive species, as a group, also exhibited higher respiration load, higher light compensation point and higher specific leaf area. There were stronger correlations of leaf traits and greater structural (but not physiological) plasticity in invasive species than in their native counterparts. The scaling coefficients of resource use efficiencies (WUE, AQE and respiration efficiency) as well as those of fitness (biomass accumulated) versus many of the performance traits examined did not differ between the two species‐origin groups, but there were indications of significant shifts in elevation (intercept values) and shifts along common slopes in many of these relationships – signalling differences in carbon economy (revenue returned per unit energy invested) and/or resource usage. Using ordination and based on 14 ecophysiological attributes, a fair level of separation between the two groups was achieved (51.5% explanatory power), with AQE, light compensation point, respiration load, WUE, specific leaf area and leaf area ratio, in decreasing order, being the main drivers. This study suggests similarity in trait plasticity, especially for physiological traits, but there appear to be fundamental differences in carbon economy and resource conservation between native and invasive vine species.     

Elevated carbon dioxide is predicted to promote coexistence among competing species in a trait‐based model

Differential species responses to atmospheric CO₂ concentration (C_a) could lead to quantitative changes in competition among species and community composition, with flow‐on effects for ecosystem function. However, there has been little theoretical analysis of how elevated C_a (eC_a) will affect plant competition, or how composition of plant communities might change. Such theoretical analysis is needed for developing testable hypotheses to frame experimental research. Here, we investigated theoretically how plant competition might change under eC_a by implementing two alternative competition theories, resource use theory and resource capture theory, in a plant carbon and nitrogen cycling model. The model makes several novel predictions for the impact of eC_a on plant community composition. Using resource use theory, the model predicts that eC_a is unlikely to change species dominance in competition, but is likely to increase coexistence among species. Using resource capture theory, the model predicts that eC_a may increase community evenness. Collectively, both theories suggest that eC_a will favor coexistence and hence that species diversity should increase with eC_a. Our theoretical analysis leads to a novel hypothesis for the impact of eC_a on plant community composition. This hypothesis has potential to help guide the design and interpretation of eC_a experiments.     

Roles of non‐native species in large‐scale regeneration of moist tropical forests on anthropogenic grassland

This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.     

Trait‐abundance relation in response to nutrient addition in a Tibetan alpine meadow: The importance of species trade‐off in resource conservation and acquisition

In competition‐dominated communities, traits promoting resource conservation and competitive ability are expected to have an important influence on species relative abundance (SRA). Yet, few studies have tested the trait‐abundance relations in the line of species trade‐off in resource conservation versus acquisition, indicating by multiple traits coordination. We measured SRA and key functional traits involving leaf economic spectrum (SLA, specific leaf area; LDMC, leaf dry matter content; LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LPC, leaf phosphorus concentration; Hs, mature height) for ten common species in all plots subjected to addition of nitrogen fertilizer (N), phosphorus fertilizer (P), or both of them (NP) in a Tibetan alpine meadow. We test whether SRA is positively related with traits promoting plant resource conservation, while negatively correlated with traits promoting plant growth and resource acquisition. We found that species were primarily differentiated along a trade‐off axis involving traits promoting nutrient acquisition and fast growth (e.g., LPC and SLA) versus traits promoting resource conservation and competition ability (e.g., large LDMC). We further found that SRA was positively correlated with plant height, LDMC, and LCC, but negatively associated with SLA and leaf nutrient concentration irrespective of fertilization. A stronger positive height‐SRA was found in NP‐fertilized plots than in other plots, while negative correlations between SRA and SLA and LPC were found in N or P fertilized plots. The results indicate that species trade‐off in nutrient acquisition and resource conservation was a key driver of SRA in competition‐dominated communities following fertilization, with the linkage between SRA and traits depending on plant competition for specific soil nutrient and/or light availability. The results highlight the importance of competitive exclusion in plant community assembly following fertilization and suggest that abundant species in local communities become dominated at expense of growth while infrequent species hold an advantage in fast growth and dispersals to neighbor meta‐communities.     

The acquisitive–conservative axis of leaf trait variation emerges even in homogeneous environments

Background and AimsThe acquisitive–conservative axis of plant ecological strategies results in a pattern of leaf trait covariation that captures the balance between leaf construction costs and plant growth potential. Studies evaluating trait covariation within species are scarcer, and have mostly dealt with variation in response to environmental gradients. Little work has been published on intraspecific patterns of leaf trait covariation in the absence of strong environmental variation.MethodsWe analysed covariation of four leaf functional traits [specific leaf area (SLA) leaf dry matter content (LDMC), force to tear (F_t) and leaf nitrogen content (N_m)] in six Poaceae and four Fabaceae species common in the dry Chaco forest of Central Argentina, growing in the field and in a common garden. We compared intraspecific covariation patterns (slopes, correlation and effect size) of leaf functional traits with global interspecific covariation patterns. Additionally, we checked for possible climatic and edaphic factors that could affect the intraspecific covariation pattern.Key ResultsWe found negative correlations for the LDMC–SLA, F_t–SLA, LDMC–N_m and F_t–N_m trait pairs. This intraspecific covariation pattern found both in the field and in the common garden and not explained by climatic or edaphic variation in the field follows the expected acquisitive–conservative axis. At the same time, we found quantitative differences in slopes among different species, and between these intraspecific patterns and the interspecific ones. Many of these differences seem to be idiosyncratic, but some appear consistent among species (e.g. all the intraspecific LDMC–SLA and LDMC–N_m slopes tend to be shallower than the global pattern).ConclusionsOur study indicates that the acquisitive–conservative leaf functional trait covariation pattern occurs at the intraspecific level even in the absence of relevant environmental variation in the field. This suggests a high degree of variation–covariation in leaf functional traits not driven by environmental variables.     

Climate‐ and successional‐related changes in functional composition of European forests are strongly driven by tree mortality

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late‐successional short‐statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm‐like strategies at low water availability in forests formerly dominated by broad‐leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm‐like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.     

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.     

柯-青冈常绿阔叶林优势树种叶片性状变异及适应策略

厘清叶片功能性状的变异及性状间的权衡关系,对揭示植物对环境变化的响应及适应策略具有重要意义。以中亚热带柯(Lithocarpus glaber)-青冈(Cyclobalanopsis glauca)常绿阔叶林为对象,测量了1 hm²固定监测样地内6个优势树种的叶面积(LA)、比叶面积(SLA)、干物质含量(LDMC)、叶片厚度(LT)、叶片碳(LC)、叶片氮(LN)、叶片磷(LP)含量和叶片碳氮比(LC ∶ LN)8个功能性状,采用多重比较、主成分分析(PCA)法分析了群落叶片功能性状的生活型、种内和种间变异及性状间关系。结果表明：(1)群落内叶片功能性状种内、种间差异显著,变异中等(CV： 0.02-0.59),其结构性状的可塑性较化学性状保守,变异格局符合"性状空间变异分割假说";针叶树种的LA、SLA显著低于阔叶树种,常绿树种的LC和LDMC最小,而落叶树种的SLA、LN和LP最大以及LT和LC ∶ LN最小。(2)群落叶片功能性状变异主要来源于生活型和种间变异,种内变异亦有显著贡献;生活型对多数性状的贡献率较大,其对LDMC、LC ∶ LN的贡献分别高达93.11%和91.76%;种间变异(LDMC除外)对结构性状的贡献率均高于化学性状;种内变异对LP的贡献率(23.66%)较种间变异高。(3)叶片性状之间多表现出显著相关关系,针叶树和阔叶树在PCA排序图中聚集于不同区域,叶经济型谱在柯-青冈群落中具有适用性。不同树种通过叶片结构、化学性状之间的权衡策略来适应环境变化,从而实现群落物种共存。结果可为理解森林群落物种的环境适应策略、预测群落动态变化和制定植被恢复措施提供科学依据。     

New aspects of grassland recovery in old‐fields revealed by trait‐based analyses of perennial‐crop‐mediated succession

Classical old‐field succession studies focused on vegetation changes after the abandonment of annual croplands or on succession after the elimination of cultivated crops. Perennial‐crop‐mediated succession, where fields are initially covered by perennial crops, reveals alternative aspects of old‐field succession theories. We tested the validity of classical theories of old‐field succession for perennial‐crop‐mediated succession. We formulated the following hypotheses: (1) functional diversity increases with increasing field age; (2) resource acquisition versus conservation trade‐off shifts toward conservation at community level during the succession; (3) the importance of spatial and temporal seed dispersal decreases during the succession; and (4) competitiveness and stress‐tolerance increases and ruderality decreases at community level during the succession. We studied functional diversity, trait distributions and plant strategies in differently aged old‐fields using chronosequence method. We found increasing functional richness and functional divergence, but also unchanged or decreasing functional evenness. We detected a shift from resource acquisition to resource conservation strategy of communities during the succession. The role of spatial and temporal seed dispersal was found to be important not only at the initial but also at latter successional stages. We found an increasing stress‐tolerance and a decreasing ruderality during succession, while the competitiveness remained unchanged at the community level. Despite the markedly different starting conditions, we found that classical and perennial‐crop‐mediated old‐field successions have some similarities regarding the changes of functional diversity, resource acquisition versus conservation trade‐off, and seed dispersal strategies. However, we revealed also the subsequent differences. The competitive character of communities remained stable during the succession; hence, the initial stages of perennial‐crop‐mediated succession can be similar to the middle stages of classical old‐field succession. Moreover, the occupied functional niche space and differentiation were larger in the older stages, but resources were not effectively utilized within this space, suggesting that the stabilization of the vegetation requires more time.     

Functional diversity of photosynthesis during drought in a model tropical rainforest – the contributions of leaf area, photosynthetic electron transport and stomatal conductance to reduction in net ecosystem carbon exchange

The tropical rainforest mesocosm within the Biosphere 2 Laboratory, a model system of some 110 species developed over 12 years under controlled environmental conditions, has been subjected to a series of comparable drought experiments during 2000–2002. In each study, the mesocosm was subjected to a 4–6 week drought, with well‐defined rainfall events before and after the treatment. Ecosystem CO₂ uptake rate (A_eco) declined 32% in response to the drought, with changes occurring within days and being reversible within weeks, even though the deeper soil layers did not become significantly drier and leaf‐level water status of most large trees was not greatly affected. The reduced A_eco during the drought reflected both morphological and physiological responses. It is estimated that the drought‐induced 32% reduction of A_eco has three principal components: (1) leaf fall increased two‐fold whereas leaf expansion growth of some canopy dominants declined to 60%, leading to a 10% decrease in foliage coverage of the canopy. This might be the main reason for the persistent reduction of A_eco after rewatering. (2) The maximum photosynthetic electron transport rate at high light intensities in remaining leaves was reduced to 71% for three of the four species measured, even though no chronic photo‐inhibition occurred. (3) Stomata closed, leading to a reduced ecosystem water conductance to water vapour (33% of pre‐drought values), which not only reduced ecosystem carbon uptake rate, but may also have implications for water and energy budgets of tropical ecosystems. Additionally, individual rainforest trees responded differently, expressing different levels of stress and stress avoiding mechanisms. This functional diversity renders the individual response heterogeneous and has fundamental implications to scale leaf level responses to ecosystem dynamics.