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Terrestrial ecosystems are increasingly enriched with resources such as atmospheric CO 2 that limit ecosystem processes. The consequences for ecosystem carbon cycling depend on the feedbacks from other limiting resources and plant community change, which remain poorly understood for soil CO 2 efflux, J CO2 , a primary carbon flux from the biosphere to the atmosphere. We applied a unique CO 2 enrichment gradient (250 to 500 µL L -1 ) for eight years to grassland plant communities on soils from different landscape positions. We identified the trajectory of J CO2 responses and feedbacks from other resources, plant diversity [effective species richness, exp(H)], and community change (plant species turnover). We found linear increases in J CO2 on an alluvial sandy loam and a lowland clay soil, and an asymptotic increase on an upland silty clay soil. Structural equation modeling identified CO 2 as the dominant limitation on J CO2 on the clay soil. In contrast with theory predicting limitation from a single limiting factor, the linear J CO2 response on the sandy loam was reinforced by positive feedbacks from aboveground net primary productivity and exp(H), while the asymptotic J CO2 response on the silty clay arose from a net negative feedback among exp(H), species turnover, and soil water potential. These findings support a multiple resource limitation view of the effects of global change drivers on grassland ecosystem carbon cycling and highlight a crucial role for positive or negative feedbacks between limiting resources and plant community structure. Incorporating these feedbacks will improve models of terrestrial carbon sequestration and ecosystem services., Competing Interests: The authors declare no competing interest.

Atmospheric CO 2 enrichment usually increases the aboveground net primary productivity (ANPP) of grassland vegetation, but the magnitude of the ANPP-CO 2 response differs among ecosystems. Soil properties affect ANPP via multiple mechanisms and vary over topographic to geographic gradients, but have received little attention as potential modifiers of the ANPP-CO 2 response. We assessed the effects of three soil types, sandy loam, silty clay and clay, on the ANPP response of perennial C 3 /C 4 grassland communities to a subambient to elevated CO 2 gradient over 10 yr in Texas, USA. We predicted an interactive, rather than additive, effect of CO 2 and soil type on ANPP. Contrary to prediction, CO 2 and soil additively influenced grassland ANPP. Increasing CO 2 by 250 μl l -1 increased ANPP by 170 g m -2 across soil types. Increased clay content from 10% to 50% among soils reduced ANPP by 50 g m -2 . CO 2 enrichment increased ANPP via a predominant direct effect, accompanied by a smaller indirect effect mediated by a successional shift to increased dominance of the C 4 tallgrass Sorghastrum nutans. Our results indicate a large, positive influence of CO 2 enrichment on grassland productivity that resulted from the direct physiological benefits of CO 2 augmented by species succession, and was expressed similarly across soils of differing physical properties., (No claim to original US Government works New Phytologist © 2018 New Phytologist Trust.)

Intraspecific variation in biomass production responses to elevated atmospheric carbon dioxide (eCO2) could influence tree species' ecological and evolutionary responses to climate change. However, the physiological mechanisms underlying genotypic variation in responsiveness to eCO2 remain poorly understood. In this study, we grew 17 Eucalyptus camaldulensis Dehnh. subsp. camaldulensis genotypes (representing provenances from four different climates) under ambient atmospheric CO2 and eCO2. We tested whether genotype leaf-scale photosynthetic and whole-tree carbon (C) allocation responses to eCO2 were predictive of genotype biomass production responses to eCO2. Averaged across genotypes, growth at eCO2 increased in situ leaf net photosynthesis (Anet) (29%) and leaf starch concentrations (37%). Growth at eCO2 reduced the maximum carboxylation capacity of Rubisco (-4%) and leaf nitrogen per unit area (Narea, -6%), but Narea calculated on a total non-structural carbohydrate-free basis was similar between treatments. Growth at eCO2 also increased biomass production and altered C allocation by reducing leaf area ratio (-11%) and stem mass fraction (SMF, -9%), and increasing leaf mass area (18%) and leaf mass fraction (5%). Overall, we found few significant CO2 × provenance or CO2 × genotype (within provenance) interactions. However, genotypes that showed the largest increases in total dry mass at eCO2 had larger increases in root mass fraction (with larger decreases in SMF) and photosynthetic nitrogen-use efficiency (PNUE) with CO2 enrichment. These results indicate that genetic differences in PNUE and carbon sink utilization (in roots) are both important predictors of tree productivity responsiveness to eCO2.

Atmospheric CO2 (ca) has increased since the last glacial period, increasing photosynthetic water use efficiency and improving plant productivity. Evolution of C4 photosynthesis at low ca led to decreased stomatal conductance (gs), which provided an advantage over C3 plants that may be reduced by rising ca. Using controlled environments, we determined how increasing ca affects C4 water use relative to C3 plants. Leaf gas exchange and mass per area (LMA) were measured for four C3 and four C4 annual, crop-related grasses at glacial (200 µmol mol-1), ambient (400 µmol mol-1), and super-ambient (640 µmol mol-1) ca. C4 plants had lower gs, which resulted in a water use efficiency advantage at all ca and was broadly consistent with slower stomatal responses to shade, indicating less pressure on leaf water status. At glacial ca, net CO2 assimilation and LMA were lower for C3 than for C4 leaves, and C3 and C4 grasses decreased leaf hydraulic conductance (Kleaf) similarly, but only C4 leaves decreased osmotic potential at turgor loss. Greater carbon availability in C4 leaves at glacial ca generated a different hydraulic adjustment relative to C3 plants. At current and future ca, C4 grasses have advantages over C3 grasses due to lower gs, lower stomatal sensitivity, and higher absolute water use efficiency.

Continuing enrichment of atmospheric CO 2 may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO 2 enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C 4 -dominated grassland communities on contrasting soils along a CO 2 concentration gradient spanning pre-industrial to expected mid-21st century levels (250-500 μl/L). CO 2 enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO 2 enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R 2  = .34-.74), a measure of species abundance. CO 2 enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species-specific controls on flowering in two general forms: (1) CO 2 effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO 2 effect. Understanding the heterogeneity in species responses to CO 2 enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO 2 effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes., (Published 2017. This article is a U.S. Government work and is in the public domain in the USA.)

Plant gas exchange is regulated by stomata, which coordinate leaf-level water loss with xylem transport. Stomatal opening responds to internal concentrations of CO₂ in the leaf, but changing CO₂ can also lead to changes in stomatal density that influence transpiration. Given that stomatal conductance increases under subambient concentrations of CO₂ and, conversely, that plants lose less water at elevated concentrations, can downstream effects of atmospheric CO₂ be observed in xylem tissue? We approached this problem by evaluating leaf stomatal density, xylem transport, xylem anatomy and resistance to cavitation in Helianthus annuus plants grown under three CO₂ regimes ranging from pre-industrial to elevated concentrations. Xylem transport, conduit size and stomatal density all increased at 290 ppm relative to ambient and elevated CO₂ concentrations. The shoots of the 290-ppm-grown plants were most vulnerable to cavitation, whereas xylem cavitation resistance did not differ in 390- and 480-ppm-grown plants. Our data indicate that, even as an indirect driver of water loss, CO₂ can affect xylem structure and water transport by coupling stomatal and xylem hydraulic functions during plant development. This plastic response has implications for plant water use under variable concentrations of CO₂, as well as the evolution of efficient xylem transport., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Leaf-level gas exchange data support the mechanistic understanding of plant fluxes of carbon and water. These fluxes inform our understanding of ecosystem function, are an important constraint on parameterization of terrestrial biosphere models, are necessary to understand the response of plants to global environmental change, and are integral to efforts to improve crop production. Collection of these data using gas analyzers can be both technically challenging and time consuming, and individual studies generally focus on a small range of species, restricted time periods, or limited geographic regions. The high value of these data is exemplified by the many publications that reuse and synthesize gas exchange data, however the lack of metadata and data reporting conventions make full and efficient use of these data difficult. Here we propose a reporting format for leaf-level gas exchange data and metadata to provide guidance to data contributors on how to store data in repositories to maximize their discoverability, facilitate their efficient reuse, and add value to individual datasets. For data users, the reporting format will better allow data repositories to optimize data search and extraction, and more readily integrate similar data into harmonized synthesis products. The reporting format specifies data table variable naming and unit conventions, as well as metadata characterizing experimental conditions and protocols. For common data types that were the focus of this initial version of the reporting format, i.e., survey measurements, dark respiration, carbon dioxide and light response curves, and parameters derived from those measurements, we took a further step of defining required additional data and metadata that would maximize the potential reuse of those data types. To aid data contributors and the development of data ingest tools by data repositories we provided a translation table comparing the outputs of common gas exchange instruments. Extensive consultation with data collectors, data users, instrument manufacturers, and data scientists was undertaken in order to ensure that the reporting format met community needs. The reporting format presented here is intended to form a foundation for future development that will incorporate additional data types and variables as gas exchange systems and measurement approaches advance in the future. The reporting format is published in the U.S. Department of Energy's ESS-DIVE data repository, with documentation and future development efforts being maintained in a version control system.

The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO2 response curves, including data from 141 C3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common-garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Circadian resonance, whereby a plant's endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C3 plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole-plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian-regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.

Climate change threatens the ability of agriculture and forestry to meet growing global demands for food, fibre and wood products. Information gathered from genotype-by-environment interactions (G × E), which demonstrate intraspecific variation in phenotypic plasticity (the ability of a genotype to alter its phenotype in response to environmental change), may prove important for bolstering agricultural and forest productivity under climate change. Nonetheless, very few studies have explicitly quantified genotype plasticity-productivity relationships in agriculture or forestry. Here, we conceptualize the importance of intraspecific variation in agricultural and forest species plasticity, and discuss the physiological and genetic factors contributing to intraspecific variation in phenotypic plasticity. Our discussion highlights the need for an integrated understanding of the mechanisms of G × E, more extensive assessments of genotypic responses to climate change under field conditions, and explicit testing of genotype plasticity-productivity relationships. Ultimately, further investigation of intraspecific variation in phenotypic plasticity in agriculture and forestry may prove important for identifying genotypes capable of increasing or sustaining productivity under more extreme climatic conditions.

Climate change threatens the ability of agriculture and forestry to meet growing global demands for food, fibre and wood products. Information gathered from genotype-by-environment interactions (G × E), which demonstrate intraspecific variation in phenotypic plasticity (the ability of a genotype to alter its phenotype in response to environmental change), may prove important for bolstering agricultural and forest productivity under climate change. Nonetheless, very few studies have explicitly quantified genotype plasticity-productivity relationships in agriculture or forestry. Here, we conceptualize the importance of intraspecific variation in agricultural and forest species plasticity, and discuss the physiological and genetic factors contributing to intraspecific variation in phenotypic plasticity. Our discussion highlights the need for an integrated understanding of the mechanisms of G × E, more extensive assessments of genotypic responses to climate change under field conditions, and explicit testing of genotype plasticity-productivity relationships. Ultimately, further investigation of intraspecific variation in phenotypic plasticity in agriculture and forestry may prove important for identifying genotypes capable of increasing or sustaining productivity under more extreme climatic conditions.

Abstract: Continuing enrichment of atmospheric CO2 may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO2 enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C4‐dominated grassland communities on contrasting soils along a CO2 concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO2 enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO2 enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R2 = .34–.74), a measure of species abundance. CO2 enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO2 effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO2 effect. Understanding the heterogeneity in species responses to CO2 enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO2 effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes. [ABSTRACT FROM AUTHOR]

Leaf-level gas exchange data support the mechanistic understanding of plant fluxes of carbon and water. These fluxes inform our understanding of ecosystem function, are an important constraint on parameterization of terrestrial biosphere models, are necessary to understand the response of plants to global environmental change, and are integral to efforts to improve crop production. Collection of these data using gas analyzers can be both technically challenging and time consuming, and individual studies generally focus on a small range of species, restricted time periods, or limited geographic regions. The high value of these data is exemplified by the many publications that reuse and synthesize gas exchange data, however the lack of metadata and data reporting conventions make full and efficient use of these data difficult. Here we propose a reporting format for leaf-level gas exchange data and metadata to provide guidance to data contributors on how to store data in repositories to maximize their discoverability, facilitate their efficient reuse, and add value to individual datasets. For data users, the reporting format will better allow data repositories to optimize data search and extraction, and more readily integrate similar data into harmonized synthesis products. The reporting format specifies data table variable naming and unit conventions, as well as metadata characterizing experimental conditions and protocols. For common data types that were the focus of this initial version of the reporting format, i.e., survey measurements, dark respiration, carbon dioxide and light response curves, and parameters derived from those measurements, we took a further step of defining required additional data and metadata that would maximize the potential reuse of those data types. To aid data contributors and the development of data ingest tools by data repositories we provided a translation table comparing the outputs of common gas exchange instruments. Extensive consultation with data collectors, data users, instrument manufacturers, and data scientists was undertaken in order to ensure that the reporting format met community needs. The reporting format presented here is intended to form a foundation for future development that will incorporate additional data types and variables as gas exchange systems and measurement approaches advance in the future. The reporting format is published in the U.S. Department of Energy's ESS-DIVE data repository, with documentation and future development efforts being maintained in a version control system. • The first data reporting format for leaf-level gas exchange data • Strong community consensus from data contributors and users • Key data types identified and explored in more detail • A cornerstone for future development of the format [ABSTRACT FROM AUTHOR]