Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.Green leaves are fundamental for the functioning of terrestrial ecosystems. Their pigments are the predominant signal seen from space. Nitrogen uptake and carbon assimilation by plants and the decomposability of leaves drive biogeochemical cycles. Animals, fungi and other heterotrophs in ecosystems are fuelled by photosynthate, and their habitats are structured by the stems on which leaves are deployed. Plants invest photosynthate and mineral nutrients in the construction of leaves, which in turn return a revenue stream of photosynthate over their lifetimes. The photosynthate is used to acquire mineral nutrients, to support metabolism and to re-invest in leaves, their supporting stems and other plant parts.There are more than 250,000 vascular plant species, all engaging in the same processes of investment and reinvestment of carbon and mineral nutrients, and all making enough surplus to ensure continuity to future generations. These processes of investment and re-investment are inherently economic in nature [1][2][3] . Understanding how these processes vary between species, plant functional types and the vegetation of different biomes is a major goal for plant ecology and crucial for modelling how nutrient fluxes and vegetation boundaries will shift with land-use and climate change. Data set and parametersWe formed a global plant trait network (Glopnet) to quantify leaf economics across the world's plant species. The Glopnet data set spans 2,548 species from 219 families at 175 sites (approximately 1% of the extant vascular plant species). The coverage of traits, species and sites is at least tenfold greater than previous data compilations [4][5][6][7][8][9][10][11] , extends to all vegetated continents, and represents a wide range of vegetation types, from arctic tundra to tropical rainforest, from hot to cold deserts, from boreal forest to grasslands. Site elevation ranges from below sea level (Death Valley, USA) to 4,800 m. Mean annual temperature (MAT) ranges from 216.5 8C to 27.5 8C; mean annual rainfall (MAR) ranges from 133 to 5,300 mm per year. This cove...

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

AimOur aim was to quantify climatic influences on key leaf traits and relationships at the global scale. This knowledge provides insight into how plants have adapted to different environmental pressures, and will lead to better calibration of future vegetation-climate models.Location The data set represents vegetation from 175 sites around the world.Methods For more than 2500 vascular plant species, we compiled data on leaf mass per area (LMA), leaf life span (LL), nitrogen concentration (N mass ) and photosynthetic capacity (A mass ). Site climate was described with several standard indices. Correlation and regression analyses were used for quantifying relationships between single leaf traits and climate. Standardized major axis (SMA) analyses were used for assessing the effect of climate on bivariate relationships between leaf traits. Principal components analysis (PCA) was used to summarize multidimensional trait variation.Results At hotter, drier and higher irradiance sites, (1) mean LMA and leaf N per area were higher; (2) average LL was shorter at a given LMA, or the increase in LL was less for a given increase in LMA (LL-LMA relationships became less positive); and (3) A mass was lower at a given N mass , or the increase in A mass was less for a given increase in N mass . Considering all traits simultaneously, 18% of variation along the principal multivariate trait axis was explained by climate.Main conclusions Trait-shifts with climate were of sufficient magnitude to have major implications for plant dry mass and nutrient economics, and represent substantial selective pressures associated with adaptation to different climatic regimes.

Summary• Global-scale quantification of relationships between plant traits gives insight into the evolution of the world's vegetation, and is crucial for parameterizing vegetationclimate models.• A database was compiled, comprising data for hundreds to thousands of species for the core 'leaf economics' traits leaf lifespan, leaf mass per area, photosynthetic capacity, dark respiration, and leaf nitrogen and phosphorus concentrations, as well as leaf potassium, photosynthetic N-use efficiency (PNUE), and leaf N : P ratio.• While mean trait values differed between plant functional types, the range found within groups was often larger than differences among them. Future vegetationclimate models could incorporate this knowledge.• The core leaf traits were intercorrelated, both globally and within plant functional types, forming a 'leaf economics spectrum'. While these relationships are very general, they are not universal, as significant heterogeneity exists between relationships fitted to individual sites. Much, but not all, heterogeneity can be explained by variation in sample size alone. PNUE can also be considered as part of this trait spectrum, whereas leaf K and N : P ratios are only loosely related.

Most plants show considerable capacity to adjust their photosynthetic characteristics to their growth temperatures (temperature acclimation). The most typical case is a shift in the optimum temperature for photosynthesis, which can maximize the photosynthetic rate at the growth temperature. These plastic adjustments can allow plants to photosynthesize more efficiently at their new growth temperatures. In this review article, we summarize the basic differences in photosynthetic reactions in C3, C4, and CAM plants. We review the current understanding of the temperature responses of C3, C4, and CAM photosynthesis, and then discuss the underlying physiological and biochemical mechanisms for temperature acclimation of photosynthesis in each photosynthetic type. Finally, we use the published data to evaluate the extent of photosynthetic temperature acclimation in higher plants, and analyze which plant groups (i.e., photosynthetic types and functional types) have a greater inherent ability for photosynthetic acclimation to temperature than others, since there have been reported interspecific variations in this ability. We found that the inherent ability for temperature acclimation of photosynthesis was different: (1) among C3, C4, and CAM species; and (2) among functional types within C3 plants. C3 plants generally had a greater ability for temperature acclimation of photosynthesis across a broad temperature range, CAM plants acclimated day and night photosynthetic process differentially to temperature, and C4 plants was adapted to warm environments. Moreover, within C3 species, evergreen woody plants and perennial herbaceous plants showed greater temperature homeostasis of photosynthesis (i.e., the photosynthetic rate at high-growth temperature divided by that at low-growth temperature was close to 1.0) than deciduous woody plants and annual herbaceous plants, indicating that photosynthetic acclimation would be particularly important in perennial, long-lived species that would experience a rise in growing season temperatures over their lifespan. Interestingly, across growth temperatures, the extent of temperature homeostasis of photosynthesis was maintained irrespective of the extent of the change in the optimum temperature for photosynthesis (T opt), indicating that some plants achieve greater photosynthesis at the growth temperature by shifting T opt, whereas others can also achieve greater photosynthesis at the growth temperature by changing the shape of the photosynthesis-temperature curve without shifting T opt. It is considered that these differences in the inherent stability of temperature acclimation of photosynthesis would be reflected by differences in the limiting steps of photosynthetic rate.

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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 CO 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 CO 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.

Growth temperature alters temperature dependence of the photosynthetic rate (temperature acclimation). In many species, the optimal temperature that maximizes the photosynthetic rate increases with increasing growth temperature. In this minireview, mechanisms involved in changes in the photosynthesis-temperature curve are discussed. Based on the biochemical model of photosynthesis, change in the photosynthesis-temperature curve is attributable to four factors: intercellular CO2 concentration, activation energy of the maximum rate of RuBP (ribulose-1,5-bisphosphate) carboxylation (Vc max), activation energy of the rate of RuBP regeneration (Jmax), and the ratio of Jmax to Vc max. In the survey, every species increased the activation energy of Vc max with increasing growth temperature. Other factors changed with growth temperature, but their responses were different among species. Among these factors, activation energy of Vc max may be the most important for the shift of optimal temperature of photosynthesis at ambient CO2 concentrations. Physiological and biochemical causes for the change in these parameters are discussed.