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The leaf economics spectrum (LES) represents a suite of intercorrelated leaf traits concerning construction costs per unit leaf area, nutrient concentrations, and rates of carbon fixation and tissue turnover. Although broad trade-offs among leaf structural and physiological traits have been demonstrated, we still do not have a comprehensive view of the fundamental constraints underlying the LES trade-offs. Here, we investigated physiological and structural mechanisms underpinning the LES by analysing a novel data compilation incorporating rarely considered traits such as the dry mass fraction in cell walls, nitrogen allocation, mesophyll CO2 diffusion and associated anatomical traits for hundreds of species covering major growth forms. The analysis demonstrates that cell wall constituents are major components of leaf dry mass (18–70%), especially in leaves with high leaf mass per unit area (LMA) and long lifespan. A greater fraction of leaf mass in cell walls is typically associated with a lower fraction of leaf nitrogen (N) invested in photosynthetic proteins; and lower within-leaf CO2 diffusion rates, as a result of thicker mesophyll cell walls. The costs associated with greater investments in cell walls underpin the LES: long leaf lifespans are achieved via higher LMA and in turn by higher cell wall mass fraction, but this inevitably reduces the efficiency of photosynthesis.

Just as people with the same weight can have different body builds, woods with the same wood density can have different anatomies. Here, our aim was to assess the magnitude of anatomical variation within a restricted range of wood density and explore its potential ecological implications. Twig wood of 69 angiosperm tree and shrub species was analyzed. Species were selected so that wood density varied within a relatively narrow range (0.38-0.62 g cm-3). Anatomical traits quantified included wood tissue fractions (fibres, axial parenchyma, ray parenchyma, vessels, and conduits with maximum lumen diameter below 15 μm), vessel properties, and pith area. To search for potential ecological correlates of anatomical variation the species were sampled across rainfall and temperature contrasts, and several other ecologically-relevant traits were measured (plant height, leaf area to sapwood area ratio, and modulus of elasticity). Despite the limited range in wood density, substantial anatomical variation was observed. Total parenchyma fraction varied from 0.12 to 0.66 and fibre fraction from 0.20 to 0.74, and these two traits were strongly inversely correlated (r = -0.86, P < 0.001). Parenchyma was weakly (0.24 ≤|r|≤ 0.35, P < 0.05) or not associated with vessel properties nor with height, leaf area to sapwood area ratio, and modulus of elasticity (0.24 ≤|r|≤ 0.41, P < 0.05). However, vessel traits were fairly well correlated with height and leaf area to sapwood area ratio (0.47 ≤|r|≤ 0.65, all P < 0.001). Modulus of elasticity was mainly driven by fibre wall plus vessel wall fraction rather than by the parenchyma component. Overall, there seem to be at least three axes of variation in xylem, substantially independent of each other: a wood density spectrum, a fibre-parenchyma spectrum, and a vessel area spectrum. The fibre-parenchyma spectrum does not yet have any clear or convincing ecological interpretation.

• Leaf area (LA), mass per area (LMA), nitrogen per unit area (Narea) and the leaf-internal to ambient CO₂ ratio (χ) are fundamental traits for plant functional ecology and vegetation modelling. Here we aimed to assess how their variation, within and between species, tracks environmental gradients. • Measurements were made on 705 species from 116 sites within a broad north–south transect from tropical to temperate Australia. Trait responses to environment were quantified using multiple regression; within- and between-species responses were compared using analysis of covariance and trait-gradient analysis. • Leaf area, the leaf economics spectrum (indexed by LMA and Narea) and χ (from stable carbon isotope ratios) varied almost independently among species. Across sites, however, χ and LA increased with mean growing-season temperature (mGDD₀) and decreased with vapour pressure deficit (mVPD₀) and soil pH. LMA and Narea showed the reverse pattern. Climate responses agreed with expectations based on optimality principles. Within-species variability contributed < 10% to geographical variation in LA but > 90% for χ, with LMA and Narea intermediate. • These findings support the hypothesis that acclimation within individuals, adaptation within species and selection among species combine to create predictable relationships between traits and environment. However, the contribution of acclimation/adaptation vs species selection differs among traits.

Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves, the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot, sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich “next-generation” vegetation models in which leaf temperature and water use during photosynthesis play key roles.

Leaf functional traits are important indicators of plant growth and ecosystem dynamics. Despite a wealth of knowledge about leaf trait relationships, a mechanistic understanding of how biotic and abiotic factors quantitatively influence leaf trait variation and scaling is still incomplete. We propose that leaf water content (LWC) inherently affects other leaf traits, although its role has been largely neglected. Here, we present a modification of a previously validated model based on metabolic theory and use an extensive global leaf trait dataset to test it. Analyses show that mass-based photosynthetic capacity and specific leaf area increase nonlinearly with LWC, as predicted by the model. When the effects of temperature and LWC are controlled, the numerical values for the leaf area-mass scaling exponents converge onto 1.0 across plant functional groups, ecosystem types, and latitudinal zones. The data also indicate that leaf water mass is a better predictor of whole-leaf photosynthesis and leaf area than whole-leaf nitrogen and phosphorus masses. Our findings highlight a comprehensive theory that can quantitatively predict some global patterns from the leaf economics spectrum. Leaf functional traits are increasingly used as proxies for plant functions. Here, the authors show that leaf water affects other leaf traits and is a better predictor of whole-leaf photosynthesis and leaf area than leaf nitrogen or phosphorus content.

Summary Generalised dose–response curves are essential to understand how plants acclimate to atmospheric CO2. We carried out a meta‐analysis of 630 experiments in which C3 plants were experimentally grown at different [CO2] under relatively benign conditions, and derived dose–response curves for 85 phenotypic traits. These curves were characterised by form, plasticity, consistency and reliability. Considered over a range of 200–1200 µmol mol−1 CO2, some traits more than doubled (e.g. area‐based photosynthesis; intrinsic water‐use efficiency), whereas others more than halved (area‐based transpiration). At current atmospheric [CO2], 64% of the total stimulation in biomass over the 200–1200 µmol mol−1 range has already been realised. We also mapped the trait responses of plants to [CO2] against those we have quantified before for light intensity. For most traits, CO2 and light responses were of similar direction. However, some traits (such as reproductive effort) only responded to light, others (such as plant height) only to [CO2], and some traits (such as area‐based transpiration) responded in opposite directions. This synthesis provides a comprehensive picture of plant responses to [CO2] at different integration levels and offers the quantitative dose–response curves that can be used to improve global change simulation models.

Summary 565 I. LMA in perspective 566 II. LMA in the field 567 III. Inherent differences 568 IV. Relation with anatomy and chemical composition 570 V. Environmental effects 572 VI. Differences in space and time 577 VII. Molecular regulation and physiology 579 VIII. Ecological consequences 580 IX. Conclusions and perspectives 582 Acknowledgements 582 References 582 Appendices 587

Background and AimsThe leaf economic spectrum (LES) is related to dry mass and nutrient investments towards photosynthetic processes and leaf structures, and to the duration of returns on those investments (leaf lifespan, LL). Phosphorus (P) is a key limiting nutrient for plant growth, yet it is unclear how the allocation of leaf P among different functions is coordinated with the LES. We addressed this question among 12 evergreen woody species co-occurring on P-impoverished soils in south-eastern Australia.MethodsLeaf ‘economic’ traits, including LL, leaf mass per area (LMA), light-saturated net photosynthetic rate per mass (Amass), dark respiration rate, P concentration ([Ptotal]), nitrogen concentration, and P resorption, were measured for three pioneer and nine non-pioneer species. Leaf P was separated into five functional fractions: orthophosphate P (Pi), metabolite P (PM), nucleic acid P (PN), lipid P (PL), and residual P (PR; phosphorylated proteins and unidentified compounds that contain P).ResultsLL was negatively correlated with Amass and positively correlated with LMA, representing the LES. Pioneers occurred towards the short-LL end of the spectrum and exhibited higher [Ptotal] than non-pioneer species, primarily associated with higher concentrations of Pi, PN and PL. There were no significant correlations between leaf P fractions and LL or LMA, while Amass was positively correlated with the concentration of PR.ConclusionsAllocation of leaf P to different fractions varied substantially among species. This variation was partially associated with the LES, which may provide a mechanism underlying co-occurrence of species with different ecological strategies under P limitation.

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals ( r 2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates ( r 2 = 0.46), and much weaker for large trees ( r 2 ≤ 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height ( H max ) are key traits among forest trees. Seed mass, LMA, WD, and H max are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak ( r 2 < 0.1 for three and r 2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for >80% of the explained variation and, after WD was included, LMA and H max made insignificant contributions. Virtually the full range of values of SM, LMA, and H max occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.

1. A prevalent question in the study of plant invasions has been whether or not invasions can be explained on the basis of traits. Despite many attempts, a synthetic view of multi‐trait differences between alien and native species is not yet available. 2. We compiled a database of three ecologically important traits (specific leaf area, typical maximum canopy height, individual seed mass) for 4473 species sampled over 95 communities (3784 species measured in their native range, 689 species in their introduced range, 207 in both ranges). 3. Considering each trait separately, co‐occurring native and alien species significantly differed in their traits. These differences, although modest, were expressed in a combined 15% higher specific leaf area, 16% lower canopy height and 26% smaller seeds. 4. Using three novel multi‐trait metrics of functional diversity, aliens showed significantly smaller trait ranges, larger divergences and a consistent differentiation from the median trait combination of co‐occurring natives. 5. We conclude that the simultaneous evaluation of multiple traits is an important novel direction in understanding invasion success. Our results support the phenotypic divergence hypothesis that predicts functional trait differences contribute to the success of alien species.