Browsing by Author "Wright, Ian J"
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Item ‘‘Diminishing returns’’ in the scaling of functional leaf traits across and within species groups(National Academy of Sciences, 2007) Niklas, Karl J; Cobb, Edward D; Niinemets, Ulo; Reich, Peter B; Sellin, Arne; Shipley, Bill; Wright, Ian JMore than 5,000 measurements from 1,943 plant species were used to explore the scaling relationships among the foliar surface area and the dry, water, and nitrogen/phosphorus mass of mature individual leaves. Although they differed statistically, the exponents for the relationships among these variables were numerically similar among six species groups (ferns, graminoids, forbs, shrubs, trees, and vines) and within 19 individual species. In general, at least one among the many scaling exponents was <1.0, such that increases in one or more features influencing foliar function (e.g., surface area or living leaf mass) failed to keep pace with increases in mature leaf size. Thus, a general set of scaling relationships exists that negatively affects increases in leaf size. We argue that this set reflects a fundamental property of all plants and helps to explain why annual growth fails to keep pace with increases in total body mass across species.Item An evolutionary perspective on leaf economics: Phylogenetics of leaf mass per area in vascular plants(2014) Flores, Olivier; Garnier, Eric; Wright, Ian J; Reich, Peter B; Pierce, Simon; D ıaz, Sandra; Pakeman, Robin J; Rusch, Graciela M; Bernard-Verdier, Maud; Testi, Baptiste; Bakker1, Jan P; Bekker, Renee M; Cerabolini, Bruno E L; Ceriani, Roberta M; Cornu, Guillaume; Cruz, Pablo; Delcamp, Matthieu; Dolezal, Jiri; Eriksson, Ove; Fayolle, Adeline; Freitas, Helena; Golodets, Carly; Gourlet-Fleury, Sylvie; Hodgson, John G; Brusa, Guido; Kleyer, Michael; Kunzmann, Dieter; Lavorel, Sandra; Papanastasis, Vasilios P; Perez-Harguindeguy, Natalia; Vendramini, Fernanda; Weiher, EvanIn plant leaves, resource use follows a trade-off between rapid resource capture and conservative storage. This “worldwide leaf economics spectrum” consists of a suite of intercorrelated leaf traits, among which leaf mass per area, LMA, is one of the most fundamental as it indicates the cost of leaf construction and light-interception borne by plants. We conducted a broad-scale analysis of the evolutionary history of LMA across a large dataset of 5401 vascular plant species. The phylogenetic signal in LMA displayed low but significant conservatism, that is, leaf economics tended to be more similar among close relatives than expected by chance alone. Models of trait evolution indicated that LMA evolved under weak stabilizing selection. Moreover, results suggest that different optimal phenotypes evolved among large clades within which extremes tended to be selected against. Conservatism in LMA was strongly related to growth form, as were selection intensity and phenotypic evolutionary rates: woody plants showed higher conservatism in relation to stronger stabilizing selection and lower evolutionary rates compared to herbaceous taxa. The evolutionary history of LMA thus paints different evolutionary trajectories of vascular plant species across clades, revealing the coordination of leaf trait evolution with growth forms in response to varying selection regimes.Item Functional traits and the growth-mortality tradeoff in tropical trees(Ecological Society of America, 2010) Wright, S Joseph; Kitajima, Kaoru; Kraft, Nathan J B; Reich, Peter B; Wright, Ian J; Bunker, Daniel E; Condit, Richard; Dalling, James W; Davies, Stuart J; Diaz, Sandra; Engelbrecht, Bettina M J; Harms, Kyle E; Hubbell, Stephen P; Marks, Christian O; Ruiz-Jaen, Maria C; Salvador, Cristina M; Zanne, Amy EA 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 (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 ≤ 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 (Hmax) are key traits among forest trees. Seed mass, LMA, WD, and Hmax 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 (r2 < 0.1 for three and r2 < 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 Hmax made insignificant contributions. Virtually the full range of values of SM, LMA, and Hmax 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.Item Global effects of soil and climate on leaf photosynthetic traits and rates(Wiley, 2015) Maire, Vincent; Wright, Ian J; Prentice, I. Colin; Batjes, Niels H; Bhaskar, Radika; Bodegom, Peter M; Cornwell, Will K; Ellsworth, David; Niinemets, Ülo; Ordonez, Alejandro; Reich, Peter B; Santiago, Louis SAim The influence of soil properties on photosynthetic traits in higher plants is poorly quantified in comparison with that of climate. We address this situation by quantifying the unique and joint contributions to global leaf-trait variation from soils and climate. Location Terrestrial ecosystems world-wide. Methods Using a trait dataset comprising 1509 species from 288 sites, with climate and soil data derived from global datasets, we quantified the effects of 20 soil and 26 climate variables on light-saturated photosynthetic rate (Aarea), stomatal conductance (gs), leaf nitrogen and phosphorus (Narea and Parea) and specific leaf area (SLA) using mixed regression models and multivariate analyses. Results Soil variables were stronger predictors of leaf traits than climatic variables, except for SLA. On average, Narea, Parea and Aarea increased and SLA decreased with increasing soil pH and with increasing site aridity. gs declined and Parea increased with soil available P (Pavail). Narea was unrelated to total soil N. Joint effects of soil and climate dominated over their unique effects on Narea and Parea, while unique effects of soils dominated for Aarea and gs. Path analysis indicated that variation in Aarea reflected the combined independent influences of Narea and gs, the former promoted by high pH and aridity and the latter by low Pavail. Main conclusions Three environmental variables were key for explaining variation in leaf traits: soil pH and Pavail, and the climatic moisture index (the ratio of precipitation to potential evapotranspiration). Although the reliability of global soil datasets lags behind that of climate datasets, our results nonetheless provide compelling evidence that both can be jointly used in broad-scale analyses, and that effects uniquely attributable to soil properties are important determinants of leaf photosynthetic traits and rates. A significant future challenge is to better disentangle the covarying physiological, ecological and evolutionary mechanisms that underpin trait–environment relationships.Item The global spectrum of plant form and function(2016) Díaz, Sandra; Kattge, Jens; Cornelissen, Johannes H C; Wright, Ian J; Lavorel, Sandra; Dray, Stéphane; Reu, Björn; Kleyer, Michael; Wirth, Christian; Prentice, I. Colin; Garnier, Eric; Bönisch, Gerhard; Westoby, Mark; Poorter, Hendrik; Reich, Peter B; Moles, Angela T; Dickie, John; Gillison, Andrew N; Zanne, Amy E; Chave, Jérôme; Wright, S. Joseph; Sheremet’ev, Serge N; Jactel, Hervé; Baraloto, Christopher; Cerabolini, Bruno; Pierce, Simon; Shipley, Bill; Kirkup, Donald; Casanoves, Fernando; Joswig, Julia S; Günther, Angela; Falczuk, Valeria; Rüger, Nadja; Mahecha, Miguel D; Gorné, Lucas DEarth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today’s terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.Item Incorporation of plant traits in a land surface model helps explain the global biogeographical distribution of major forest functional types(Wiley, 2016) Lu, Xingjie; Wang, Ying‐Ping; Wright, Ian J; Reich, Peter B; Shi, Zheng; Dai, YongjiuContrasting leaf types with different leaf life spans represent different adaptive strategies in plants. Previous studies explained the adaptive advantages of different strategies on the basis of environmental climatic limits, but could not account for the observed co-dominance of multiple plant functional types (PFTs) in many regions. Here we used a process-based model to explore whether observed inter- and intra-PFT variation in plant traits can explain global biogeographical variation in the dominance and co-dominance of major forest types.Item Predicting leaf functional traits from simple plant and climate attributers using the GLOPNET global data set(2007) Reich, Peter B; Wright, Ian J; Lusk, Christopher HKnowledge of leaf chemistry, physiology, and life span is essential for global vegetation modeling, but such data are scarce or lacking for some regions, especially in developing countries. Here we use data from 2021 species at 175 sites around the world from the GLOPNET compilation to show that key physiological traits that are difficult to measure (such as photosynthetic capacity) can be predicted from simple qualitative plant characteristics, climate information, easily measured (“soft”) leaf traits, or all of these in combination. The qualitative plant functional type (PFT) attributes examined are phylogeny (angiosperm or gymnosperm), growth form (grass, herb, shrub, or tree), and leaf phenology (deciduous vs. evergreen). These three PFT attributes explain between one-third and two-thirds of the variation in each of five quantitative leaf ecophysiological traits: specific leaf area (SLA), leaf life span, mass-based net photosynthetic capacity (Amass), nitrogen content (Nmass), and phosphorus content (Pmass). Alternatively, the combination of four simple, widely available climate metrics (mean annual temperature, mean annual precipitation, mean vapor pressure deficit, and solar irradiance) explain only 5–20% of the variation in those same five leaf traits. Adding the climate metrics to the qualitative PFTs as independent factors in the model increases explanatory power by 3–11% for the five traits. If a single easily measured leaf trait (SLA) is also included in the model along with qualitative plant traits and climate metrics, an additional 5–25% of the variation in the other four other leaf traits is explained, with the models accounting for 62%, 65%, 66%, and 73% of global variation in Nmass, Pmass, Amass, and leaf life span, respectively. Given the wide availability of the summary climate data and qualitative PFT data used in these analyses, they could be used to explain roughly half of global variation in the less accessible leaf traits (Amass, leaf life span, Nmass, Pmass); this can be augmented to two-thirds of all variation if climatic and PFT data are used in combination with the readily measured trait SLA. This shows encouraging possibilities of progress in developing general predictive equations for macro-ecology, global scaling, and global modeling.Item Three keys to the radiation of angiosperms into freezing environments(Nature Publishing Group, 2014) Zanne, Amy E; Tank, David C; Cornwell, William K; Eastman, Jonathan M; Smith, Stephen A; FitzJohn, Richard G; McGlinn, Daniel J; O’Meara, Brian C; Moles, Angela T; Reich, Peter B; Royer, Dana L; Soltis, Douglas E; Stevens, Peter F; Westoby, Mark; Wright, Ian J; Aarssen, Lonnie; Bertin, Robert I; Calaminus, Andre; Govaerts, Rafaël; Hemmings, Frank; Leishman, Michelle R; Oleksyn, Jacek; Soltis, Pamela S; Swenson, Nathan G; Warman, Laura; Beaulieu, Jeremy MEarly flowering plants are thought to have been woody species restricted to warm habitats1, 2, 3. This lineage has since radiated into almost every climate, with manifold growth forms4. As angiosperms spread and climate changed, they evolved mechanisms to cope with episodic freezing. To explore the evolution of traits underpinning the ability to persist in freezing conditions, we assembled a large species-level database of growth habit (woody or herbaceous; 49,064 species), as well as leaf phenology (evergreen or deciduous), diameter of hydraulic conduits (that is, xylem vessels and tracheids) and climate occupancies (exposure to freezing). To model the evolution of species’ traits and climate occupancies, we combined these data with an unparalleled dated molecular phylogeny (32,223 species) for land plants. Here we show that woody clades successfully moved into freezing-prone environments by either possessing transport networks of small safe conduits5 and/or shutting down hydraulic function by dropping leaves during freezing. Herbaceous species largely avoided freezing periods by senescing cheaply constructed aboveground tissue. Growth habit has long been considered labile6, but we find that growth habit was less labile than climate occupancy. Additionally, freezing environments were largely filled by lineages that had already become herbs or, when remaining woody, already had small conduits (that is, the trait evolved before the climate occupancy). By contrast, most deciduous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occupancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold.Item Why are non-photosynthetic tissues generally 13C enriched compared with leaves in C3 plants? Review and synthesis of current hypotheses(CSIRO, 2009) Cernusak, Lucas A; Tcherkez, Guillaume; Keitel, Claudia; Cornwell, William K; Santiago, Louis S; Knohl, Alexander; Barbour, Margaret M; Williams, David G; Reich, Peter B; Ellsworth, David S; Dawson, Todd E; Griffiths, Howard G; Farquhar, Graham D; Wright, Ian JNon-photosynthetic, or heterotrophic, tissues in C3 plants tend to be enriched in 13C compared with the leaves that supply them with photosynthate. This isotopic pattern has been observed for woody stems, roots, seeds and fruits, emerging leaves, and parasitic plants incapable of net CO2 fixation. Unlike in C3 plants, roots of herbaceous C4 plants are generally not 13C-enriched compared with leaves. We review six hypotheses aimed at explaining this isotopic pattern in C3 plants: (1) variation in biochemical composition of heterotrophic tissues compared with leaves; (2) seasonal separation of growth of leaves and heterotrophic tissues, with corresponding variation in photosynthetic discrimination against 13C; (3) differential use of day v. night sucrose between leaves and sink tissues, with day sucrose being relatively 13C-depleted and night sucrose 13C-enriched; (4) isotopic fractionation during dark respiration; (5) carbon fixation by PEP carboxylase; and (6) developmental variation in photosynthetic discrimination against 13C during leaf expansion. Although hypotheses (1) and (2) may contribute to the general pattern, they cannot explain all observations. Some evidence exists in support of hypotheses (3) through to (6), although for hypothesis (6) it is largely circumstantial. Hypothesis (3) provides a promising avenue for future research. Direct tests of these hypotheses should be carried out to provide insight into the mechanisms causing within-plant variation in carbon isotope composition.Item The worldwide leaf economics spectrum(Nature Publishing Group, 2004) Wright, Ian J; Reich, Peter B; Westoby, Mark; Ackerly, David D; Baruch, Zdravko; Bongers, Frans; Cavender-Bares, Jeannine; Chapin, Terry; Cornelissen, Johannes H C; Diemer, Matthias; Flexas, Jaume; Garnier, Eric; Groom, Philip K; Gulias, Javier; Hikosaka, Kouki; Lamont, Byron B; Lee, Tali; Lee, William; Lusk, Christopher; Midgley, Jeremy J; Navas, Marie-Laure; Niinemets, Ülo; Oleksyn, Jacek; Osada, Noriyuki; Poorter, Hendrik; Poot, Pieter; Prior, Lynda; Pyankov, Vladimir I; Roumet, Catherine; Thomas, Sean C; Tjoelker, Mark G; Veneklaas, Erik J; Villar, RafaelBringing 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.