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Leaf dark respiration (R d ) acclimates to environmental changes. However, the magnitude, controls and time scales of acclimation remain unclear and are inconsistently treated in ecosystem models. We hypothesized that R d and Rubisco carboxylation capacity (V cmax ) at 25°C (R d,25 , V cmax,25 ) are coordinated so that R d,25 variations support V cmax,25 at a level allowing full light use, with V cmax,25 reflecting daytime conditions (for photosynthesis), and R d,25 /V cmax,25 reflecting night-time conditions (for starch degradation and sucrose export). We tested this hypothesis temporally using a 5-yr warming experiment, and spatially using an extensive field-measurement data set. We compared the results to three published alternatives: R d,25 declines linearly with daily average prior temperature; R d at average prior night temperatures tends towards a constant value; and R d,25 /V cmax,25 is constant. Our hypothesis accounted for more variation in observed R d,25 over time (R 2  = 0.74) and space (R 2  = 0.68) than the alternatives. Night-time temperature dominated the seasonal time-course of R d , with an apparent response time scale of c. 2 wk. V cmax dominated the spatial patterns. Our acclimation hypothesis results in a smaller increase in global R d in response to rising CO 2 and warming than is projected by the two of three alternative hypotheses, and by current models., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

C 4 photosynthesis involves a series of biochemical and anatomical traits that significantly improve plant productivity under conditions that reduce the efficiency of C 3 photosynthesis. We explore how evolution of the three classical biochemical types of C 4 photosynthesis (NADP-ME, NAD-ME and PCK types) has affected the functions and properties of mitochondria. Mitochondria in C 4 NAD-ME and PCK types play a direct role in decarboxylation of metabolites for C 4 photosynthesis. Mitochondria in C 4 PCK type also provide ATP for C 4 metabolism, although this role for ATP provision is not seen in NAD-ME type. Such involvement has increased mitochondrial abundance/size and associated enzymatic capacity, led to changes in mitochondrial location and ultrastructure, and altered the role of mitochondria in cellular carbon metabolism in the NAD-ME and PCK types. By contrast, these changes in mitochondrial properties are absent in the C 4 NADP-ME type and C 3 leaves, where mitochondria play no direct role in photosynthesis. From an eco-physiological perspective, rates of leaf respiration in darkness vary considerably among C 4 species but does not differ systematically among the three C 4 types. This review outlines further mitochondrial research in key areas central to the engineering of the C 4 pathway into C 3 plants and to the understanding of variation in rates of C 4 dark respiration., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Climate change and future warming will significantly affect crop yield. The capacity of crops to dynamically adjust physiological processes (i.e., acclimate) to warming might improve overall performance. Understanding and quantifying the degree of acclimation in field crops could ensure better parameterization of crop and Earth System models and predictions of crop performance. We hypothesized that for field-grown wheat, when measured at a common temperature (25°C), crops grown under warmer conditions would exhibit acclimation, leading to enhanced crop performance and yield. Acclimation was defined as (a) decreased rates of net photosynthesis at 25°C (A 25 ) coupled with lower maximum carboxylation capacity (V cmax 25 ), (b) reduced leaf dark respiration at 25°C (both in terms of O 2 consumption R dark _O 2 25 and CO 2 efflux R dark _CO 2 25 ) and (c) lower R dark _CO 2 25 to V cmax 25 ratio. Field experiments were conducted over two seasons with 20 wheat genotypes, sown at three different planting dates, to test these hypotheses. Leaf-level CO 2 -based traits (A 25 , R dark _CO 2 25 and V cmax 25 ) did not show the classic acclimation responses that we hypothesized; by contrast, the hypothesized changes in R dark_ O 2 were observed. These findings have implications for predictive crop models that assume similar temperature response among these physiological processes and for predictions of crop performance in a future warmer world., (© 2020 John Wiley & Sons Ltd.)

To further our understanding of how sustained changes in temperature affect the carbon economy of rice (Oryza sativa), hydroponically grown plants of the IR64 cultivar were developed at 30°C/25°C (day/night) before being shifted to 25/20°C or 40/35°C. Leaf messenger RNA and protein abundance, sugar and starch concentrations, and gas-exchange and elongation rates were measured on preexisting leaves (PE) already developed at 30/25°C or leaves newly developed (ND) subsequent to temperature transfer. Following a shift in growth temperature, there was a transient adjustment in metabolic gene transcript abundance of PE leaves before homoeostasis was reached within 24 hr, aligning with R dark (leaf dark respiratory CO 2 release) and A n (net CO 2 assimilation) changes. With longer exposure, the central respiratory protein cytochrome c oxidase (COX) declined in abundance at 40/35°C. In contrast to R dark , A n was maintained across the three growth temperatures in ND leaves. Soluble sugars did not differ significantly with growth temperature, and growth was fastest with extended exposure at 40/35°C. The results highlight that acclimation of photosynthesis and respiration is asynchronous in rice, with heat-acclimated plants exhibiting a striking ability to maintain net carbon gain and growth when exposed to heat-wave temperatures, even while reducing investment in energy-conserving respiratory pathways., (© 2019 John Wiley & Sons Ltd.)

High temperatures account for major wheat yield losses annually and, as the climate continues to warm, these losses will probably increase. Both photosynthesis and respiration are the main determinants of carbon balance and growth in wheat, and both are sensitive to high temperature. Wheat is able to acclimate photosynthesis and respiration to high temperature, and thus reduce the negative affects on growth. The capacity to adjust these processes to better suit warmer conditions stands as a potential avenue toward reducing heat-induced yield losses in the future. However, much remains to be learnt about such phenomena. Here, we review what is known of high temperature tolerance in wheat, focusing predominantly on the high temperature responses of photosynthesis and respiration. We also identify the many unknowns that surround this area, particularly with respect to the high temperature response of wheat respiration and the consequences of this for growth and yield. It is concluded that further investigation into the response of photosynthesis and respiration to high temperature could present several methods of improving wheat high temperature tolerance. Extending our knowledge in this area could also lead to more immediate benefits, such as the enhancement of current crop models., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

In many biomes, plants are subject to heatwaves, potentially causing irreversible damage to the photosynthetic apparatus. Field surveys have documented global, temperature-dependent patterns in photosynthetic heat tolerance (P HT ); however, it remains unclear if these patterns reflect acclimation in P HT or inherent differences among species adapted to contrasting habitats. To address these unknowns, we quantified seasonal variations in T crit (high temperature where minimal chlorophyll-a fluorescence rises rapidly, reflecting disruption to photosystem II) in 62 species native to 6 sites from 5 thermally contrasting biomes across Australia. T crit and leaf fatty acid (FA) composition (important for membrane stability) were quantified in three temperature-controlled glasshouses in 20 of those species. T crit was greatest at hot field sites and acclimated seasonally (summer > winter, increasing on average 0.34 °C per °C increase in growth temperature). The glasshouse study showed that T crit was inherently higher in species from warmer habitats (increasing 0.16 °C per °C increase in origin annual mean maximum temperature) and acclimated to increasing growth temperature (0.24 °C °C -1 ). Variations in T crit were positively correlated with the relative abundance of saturated FAs, with FAs accounting for 40% of T crit variation. These results highlight the importance of both plastic adjustments and inherent differences determining contemporary continent-wide patterns in P HT ., (© 2018 John Wiley & Sons Ltd.)

Globally, trees originating from high-rainfall tropical regions typically exhibit lower rates of light-saturated net CO 2 assimilation (A) compared with those from high-rainfall temperate environments, when measured at a common temperature. One factor that has been suggested to contribute towards lower rates of A is lower mesophyll conductance. Using a combination of leaf gas exchange and carbon isotope discrimination measurements, we estimated mesophyll conductance (g m ) of several Australian tropical and temperate wet-forest trees, grown in a common environment. Maximum Rubisco carboxylation capacity, V cmax , was obtained from CO 2 response curves. g m and the drawdown of CO 2 across the mesophyll were both relatively constant. V cmax estimated on the basis of intercellular CO 2 partial pressure, C i , was equivalent to that estimated using chloroplastic CO 2 partial pressure, C c , using 'apparent' and 'true' Rubisco Michaelis-Menten constants, respectively Having ruled out g m as a possible factor in distorting variations in A between these tropical and temperate trees, attention now needs to be focused on obtaining more detailed information about Rubisco in these species., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Understanding of the extent of acclimation of light-saturated net photosynthesis (A n ) to temperature (T), and associated underlying mechanisms, remains limited. This is a key knowledge gap given the importance of thermal acclimation for plant functioning, both under current and future higher temperatures, limiting the accuracy and realism of Earth system model (ESM) predictions. Given this, we analysed and modelled T-dependent changes in photosynthetic capacity in 10 wet-forest tree species: six from temperate forests and four from tropical forests. Temperate and tropical species were each acclimated to three daytime growth temperatures (T growth ): temperate - 15, 20 and 25 °C; tropical - 25, 30 and 35 °C. CO 2 response curves of A n were used to model maximal rates of RuBP (ribulose-1,5-bisphosphate) carboxylation (V cmax ) and electron transport (J max ) at each treatment's respective T growth and at a common measurement T (25 °C). SDS-PAGE gels were used to determine abundance of the CO 2 -fixing enzyme, Rubisco. Leaf chlorophyll, nitrogen (N) and mass per unit leaf area (LMA) were also determined. For all species and T growth , A n at current atmospheric CO 2 partial pressure was Rubisco-limited. Across all species, LMA decreased with increasing T growth . Similarly, area-based rates of V cmax at a measurement T of 25 °C (V cmax 25 ) linearly declined with increasing T growth , linked to a concomitant decline in total leaf protein per unit leaf area and Rubisco as a percentage of leaf N. The decline in Rubisco constrained V cmax and A n and resulted in poor predictions of photosynthesis by currently widely used models that do not account for T growth -mediated changes in Rubisco abundance that underpin the thermal acclimation response of photosynthesis in wet-forest tree species. A new model is proposed that accounts for the effect of T growth -mediated changes in Rubisco abundance that underpin the thermal acclimation response of photosynthesis in wet-forest tree species. A new model is proposed that accounts for the effect of T growth -mediated declines in V cmax 25 on A n , complementing current photosynthetic thermal acclimation models that do not account for T sensitivity of V cmax 25 ., (© 2017 John Wiley & Sons Ltd.)

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (V cmax ), and the maximum rate of electron transport (J max )), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (M a , N a and P a , respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO 2 -fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf P a were key explanatory factors for models of area-based V cmax and J max but did not account for variations in photosynthetic N-use efficiency. At any given N a and P a , the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Simulations of photosynthesis by terrestrial biosphere models typically need a specification of the maximum carboxylation rate (Vcmax ). Estimating this parameter using A-Ci curves (net photosynthesis, A, vs intercellular CO2 concentration, Ci ) is laborious, which limits availability of Vcmax data. However, many multispecies field datasets include net photosynthetic rate at saturating irradiance and at ambient atmospheric CO2 concentration (Asat ) measurements, from which Vcmax can be extracted using a 'one-point method'. We used a global dataset of A-Ci curves (564 species from 46 field sites, covering a range of plant functional types) to test the validity of an alternative approach to estimate Vcmax from Asat via this 'one-point method'. If leaf respiration during the day (Rday ) is known exactly, Vcmax can be estimated with an r(2) value of 0.98 and a root-mean-squared error (RMSE) of 8.19 μmol m(-2) s(-1) . However, Rday typically must be estimated. Estimating Rday as 1.5% of Vcmax, we found that Vcmax could be estimated with an r(2) of 0.95 and an RMSE of 17.1 μmol m(-2) s(-1) . The one-point method provides a robust means to expand current databases of field-measured Vcmax , giving new potential to improve vegetation models and quantify the environmental drivers of Vcmax variation., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

In intact leaves, mitochondrial populations are highly heterogeneous among contrasting cell types; how such contrasting populations respond to sustained changes in the environment remains, however, unclear. Here, we examined respiratory rates, mitochondrial protein composition and response to growth temperature in photosynthetic (mesophyll) and non-photosynthetic (epidermal) cells from fully expanded leaves of warm-developed (WD) and cold-developed (CD) broad bean (Vicia faba L.). Rates of respiration were significantly higher in mesophyll cell protoplasts (MCPs) than epidermal cell protoplasts (ECPs), with both protoplast types exhibiting capacity for cytochrome and alternative oxidase activity. Compared with ECPs, MCPs contained greater relative quantities of porin, suggesting higher mitochondrial surface area in mesophyll cells. Nevertheless, the relative quantities of respiratory proteins (normalized to porin) were similar in MCPs and ECPs, suggesting that ECPs have lower numbers of mitochondria yet similar protein complement to MCP mitochondria (albeit with lower abundance serine hydroxymethyltransferase). Several mitochondrial proteins (both non-photorespiratory and photorespiratory) exhibited an increased abundance in response to cold in both protoplast types. Based on estimates of individual protoplast respiration rates, combined with leaf cell abundance data, epidermal cells make a small but significant (2%) contribution to overall leaf respiration which increases twofold in the cold. Taken together, our data highlight the heterogeneous nature of mitochondrial populations in leaves, both among contrasting cell types and in how those populations respond to growth temperature., (© 2015 John Wiley & Sons Ltd.)

We investigated whether the degree of light inhibition of leaf respiration (R) differs among large Eucalyptus saligna grown in whole-tree chambers and exposed to present and future atmospheric [CO(2) ] and summer drought. Associated with month-to-month changes in temperature were concomitant changes in R in the light (R(light) ) and darkness (R(dark) ), with both processes being more temperature dependent in well-watered trees than under drought. Overall rates of R(light) and R(dark) were not significantly affected by [CO(2) ]. By contrast, overall rates of R(dark) (averaged across both [CO(2) ]) were ca. 25% lower under drought than in well-watered trees. During summer, the degree of light inhibition of leaf R was greater in droughted (ca. 80% inhibition) than well-watered trees (ca. 50% inhibition). Notwithstanding these treatment differences, an overall positive relationship was observed between R(light) and R(dark) when data from all months/treatments were combined (R(2)  = 0.8). Variations in R(light) were also positively correlated with rates of Rubisco activity and nitrogen concentration. Light inhibition resulted in a marked decrease in the proportion of light-saturated photosynthesis respired (i.e. reduced R/A(sat) ). Collectively, these results highlight the need to account for light inhibition when assessing impacts of global change drivers on the carbon economy of tree canopies., (© 2011 Blackwell Publishing Ltd.)

Background and aimsTropical forests exchange more carbon dioxide (CO2) with the atmosphere than any other terrestrial biome. Yet, uncertainty in the projected carbon balance over the next century is roughly three-times greater for the tropics than other ecosystems. Our limited knowledge of tropical plant physiological responses, including photosynthetic, to climate change is a substantial source of uncertainty in our ability to forecast the global terrestrial carbon sink.MethodsWe used a meta-analytic approach, focusing on tropical photosynthetic temperature responses, to address this knowledge gap. Our dataset, gleaned from 18 independent studies, included leaf-level light saturated photosynthetic (Asat) temperature responses from 108 woody species, with additional temperature parameters (35 species) and rates (250 species) of both maximum rates of electron transport (Jmax) and Rubisco carboxylation (Vcmax). We investigated how these parameters responded to mean annual temperature (MAT), temperature variability, aridity, and elevation, as well as also how responses differed among successional strategy, leaf habit, and light environment.Key resultsOptimum temperatures for Asat (ToptA) and Jmax (ToptJ) increased with MAT but not for Vcmax (ToptV). Although photosynthetic rates were higher for "light" than "shaded" leaves, light conditions did not generate differences in temperature response parameters. ToptA did not differ with successional strategy, but early successional species had ~4 °C wider thermal niches than mid/late species. Semi-deciduous species had ~1 °C higher ToptA than broadleaf evergreen. Most global modeling efforts consider all tropical forests as a single "broadleaf evergreen" functional type, but our data show that tropical species with different leaf habits display distinct temperature responses that should be included in modeling efforts.ConclusionsThis novel research will inform modeling efforts to quantify tropical ecosystem carbon cycling and provide more accurate representations of how these key ecosystems will respond to altered temperature patterns in the face of climate warming.

We examined photosynthetic traits of pre-existing and newly developed flag leaves of four wheat genotypes grown in controlled-environment experiments. In newly developed leaves, acclimation of the maximum rate of net CO2 assimilation (A n) to warm nights (i.e. increased A n) was associated with increased capacity of Rubisco carboxylation and photosynthetic electron transport, with Rubisco activation state probably contributing to increased Rubisco activity. Metabolite profiling linked acclimation of A n to greater accumulation of monosaccharides and saturated fatty acids in leaves; these changes suggest roles for osmotic adjustment of leaf turgor pressure and maintenance of cell membrane integrity. By contrast, where A n decreased under warm nights, the decline was related to lower stomatal conductance and rates of photosynthetic electron transport. Decreases in A n occurred despite higher basal PSII thermal stability in all genotypes exposed to warm nights: T crit of 45–46.5 °C in non-acclimated versus 43.8–45 °C in acclimated leaves. Pre-existing leaves showed no change in A n–temperature response curves, except for an elite heat-tolerant genotype. These findings illustrate the impact of night-time warming on the ability of wheat plants to photosynthesize during the day, thereby contributing to explain the impact of global warming on crop productivity. [ABSTRACT FROM AUTHOR]

Summary: The rate with which crop yields per hectare increase each year is plateauing at the same time that human population growth and other factors increase food demand. Increasing yield potential (Yp) of crops is vital to address these challenges. In this review, we explore a component of Yp that has yet to be optimised – that being improvements in the efficiency with which light energy is converted into biomass (εc) via modifications to CO2 fixed per unit quantum of light (α), efficiency of respiratory ATP production (εprod) and efficiency of ATP use (εuse). For α, targets include changes in photoprotective machinery, ribulose bisphosphate carboxylase/oxygenase kinetics and photorespiratory pathways. There is also potential for εprod to be increased via targeted changes to the expression of the alternative oxidase and mitochondrial uncoupling pathways. Similarly, there are possibilities to improve εuse via changes to the ATP costs of phloem loading, nutrient uptake, futile cycles and/or protein/membrane turnover. Recently developed high‐throughput measurements of respiration can serve as a proxy for the cumulative energy cost of these processes. There are thus exciting opportunities to use our growing knowledge of factors influencing the efficiency of photosynthesis and respiration to create a step‐change in yield potential of globally important crops. [ABSTRACT FROM AUTHOR]

SUMMARY: High temperature stress inhibits photosynthesis and threatens wheat production. One measure of photosynthetic heat tolerance is Tcrit – the critical temperature at which incipient damage to photosystem II (PSII) occurs. This trait could be improved in wheat by exploiting genetic variation and genotype‐by‐environment interactions (GEI). Flag leaf Tcrit of 54 wheat genotypes was evaluated in 12 thermal environments over 3 years in Australia, and analysed using linear mixed models to assess GEI effects. Nine of the 12 environments had significant genetic effects and highly variable broad‐sense heritability (H2 ranged from 0.15 to 0.75). Tcrit GEI was variable, with 55.6% of the genetic variance across environments accounted for by the factor analytic model. Mean daily growth temperature in the month preceding anthesis was the most influential environmental driver of Tcrit GEI, suggesting biochemical, physiological and structural adjustments to temperature requiring different durations to manifest. These changes help protect or repair PSII upon exposure to heat stress, and may improve carbon assimilation under high temperature. To support breeding efforts to improve wheat performance under high temperature, we identified genotypes superior to commercial cultivars commonly grown by farmers, and demonstrated potential for developing genotypes with greater photosynthetic heat tolerance. Significance Statement: Here we examined intraspecific variation in photosynthetic heat tolerance (the critical temperature at which damage to PSII occurs ‐ Tcrit); effects of genotype‐by‐environment interactions on Tcrit; and environmental drivers of Tcrit in wheat – the most widely cultivated crop. Our study provides a sound framework for identifying germplasm with superior Tcrit relative to well‐adapted commercial cultivars. This framework can be applied in agricultural and ecological settings in selection of more heat tolerant species. [ABSTRACT FROM AUTHOR]

Wheat photosynthetic heat tolerance can be characterized using minimal chlorophyll fluorescence to quantify the critical temperature (T crit) above which incipient damage to the photosynthetic machinery occurs. We investigated intraspecies variation and plasticity of wheat T crit under elevated temperature in field and controlled-environment experiments, and assessed whether intraspecies variation mirrored interspecific patterns of global heat tolerance. In the field, wheat T crit varied diurnally—declining from noon through to sunrise—and increased with phenological development. Under controlled conditions, heat stress (36 °C) drove a rapid (within 2 h) rise in T crit that peaked after 3–4 d. The peak in T crit indicated an upper limit to PSII heat tolerance. A global dataset [comprising 183 Triticum and wild wheat (Aegilops) species] generated from the current study and a systematic literature review showed that wheat leaf T crit varied by up to 20 °C (roughly two-thirds of reported global plant interspecies variation). However, unlike global patterns of interspecies T crit variation that have been linked to latitude of genotype origin, intraspecific variation in wheat T crit was unrelated to that. Overall, the observed genotypic variation and plasticity of wheat T crit suggest that this trait could be useful in high-throughput phenotyping of wheat photosynthetic heat tolerance. [ABSTRACT FROM AUTHOR]

Summary: C4 photosynthesis involves a series of biochemical and anatomical traits that significantly improve plant productivity under conditions that reduce the efficiency of C3 photosynthesis. We explore how evolution of the three classical biochemical types of C4 photosynthesis (NADP‐ME, NAD‐ME and PCK types) has affected the functions and properties of mitochondria. Mitochondria in C4 NAD‐ME and PCK types play a direct role in decarboxylation of metabolites for C4 photosynthesis. Mitochondria in C4 PCK type also provide ATP for C4 metabolism, although this role for ATP provision is not seen in NAD‐ME type. Such involvement has increased mitochondrial abundance/size and associated enzymatic capacity, led to changes in mitochondrial location and ultrastructure, and altered the role of mitochondria in cellular carbon metabolism in the NAD‐ME and PCK types. By contrast, these changes in mitochondrial properties are absent in the C4 NADP‐ME type and C3 leaves, where mitochondria play no direct role in photosynthesis. From an eco‐physiological perspective, rates of leaf respiration in darkness vary considerably among C4 species but does not differ systematically among the three C4 types. This review outlines further mitochondrial research in key areas central to the engineering of the C4 pathway into C3 plants and to the understanding of variation in rates of C4 dark respiration. [ABSTRACT FROM AUTHOR]

Plant respiration is an important contributor to the proposed positive global carbon‐cycle feedback to climate change. However, as a major component, leaf mitochondrial ('dark') respiration (Rd) differs among species adapted to contrasting environments and is known to acclimate to sustained changes in temperature. No accepted theory explains these phenomena or predicts its magnitude. Here we propose that the acclimation of Rd follows an optimal behaviour related to the need to maintain long‐term average photosynthetic capacity (Vcmax) so that available environmental resources can be most efficiently used for photosynthesis. To test this hypothesis, we extend photosynthetic co‐ordination theory to predict the acclimation of Rd to growth temperature via a link to Vcmax, and compare predictions to a global set of measurements from 112 sites spanning all terrestrial biomes. This extended co‐ordination theory predicts that field‐measured Rd and Vcmax accessed at growth temperature (Rd,tg and Vcmax,tg) should increase by 3.7% and 5.5% per degree increase in growth temperature. These acclimated responses to growth temperature are less steep than the corresponding instantaneous responses, which increase 8.1% and 9.9% per degree of measurement temperature for Rd and Vcmax respectively. Data‐fitted responses proof indistinguishable from the values predicted by our theory, and smaller than the instantaneous responses. Theory and data are also shown to agree that the basal rates of both Rd and Vcmax assessed at 25°C (Rd,25 and Vcmax,25) decline by ~4.4% per degree increase in growth temperature. These results provide a parsimonious general theory for Rd acclimation to temperature that is simpler—and potentially more reliable—than the plant functional type‐based leaf respiration schemes currently employed in most ecosystem and land‐surface models. [ABSTRACT FROM AUTHOR]

The Jurien Bay dune chronosequence in south‐western Australia's biodiversity hotspot comprises sites differing in nutrient availability, with phosphorus (P) availability declining strongly with increasing soil age. We have explored the exceptionally high photosynthetic P‐use efficiency (PPUE) of Proteaceae in this region, triggering the question what the PPUE of co‐occurring species in other families might be along the Jurien Bay chronosequence.We explored how traits associated with PPUE, photosynthetic nitrogen (N)‐use efficiency (PNUE) and leaf respiration might converge along the chronosequence, and whether Proteaceae and non‐Proteaceae species differ in leaf traits associated with nutrient use.Seven to 10 species were sampled at three sites differing in nutrient availability (ranging from N‐ to P‐limited). Measurements of leaf light‐saturated photosynthesis and dark respiration were integrated with measurements of total N and P concentration in both mature and senesced leaves, and leaf mass per unit area (LMA).Contrary to what is known for other chronosequences, rates of photosynthesis and respiration did not decrease with increasing soil age and LMA along the Jurien Bay chronosequence. However, they increased when expressed per unit leaf P. Both N and P were used much more efficiently for photosynthesis on nutrient‐poor sites, in both Proteaceae and non‐Proteaceae species. Proteaceae had the fastest rate of photosynthesis per unit leaf P, followed by species that preferentially allocate P to mesophyll cells, rather than epidermal cells.Synthesis. Our results show that with declining soil P availability, photosynthetic P‐use efficiency of all investigated species from different families increased. Plants growing on the oldest, most nutrient‐impoverished soils exhibited similar rates of CO2 exchange as plants growing on more nutrient‐rich younger soils, and extraordinarily high photosynthetic P‐use efficiency. This indicates convergence in leaf traits related to photosynthetic nutrient use on severely P‐impoverished sites. [ABSTRACT FROM AUTHOR]

Understanding forest tree responses to climate warming and heatwaves is important for predicting changes in tree species diversity, forest C uptake, and vegetation–climate interactions. Yet, tree species differences in heatwave tolerance and their plasticity to growth temperature remain poorly understood. In this study, populations of four Eucalyptus species, two with large range sizes and two with comparatively small range sizes, were grown under two temperature treatments (cool and warm) before being exposed to an equivalent experimental heatwave. We tested whether the species with large and small range sizes differed in heatwave tolerance, and whether trees grown under warmer temperatures were more tolerant of heatwave conditions than trees grown under cooler temperatures. Visible heatwave damage was more common and severe in the species with small rather than large range sizes. In general, species that showed less tissue damage maintained higher stomatal conductance, lower leaf temperatures, larger increases in isoprene emissions, and less photosynthetic inhibition than species that showed more damage. Species exhibiting more severe visible damage had larger increases in heat shock proteins (HSPs) and respiratory thermotolerance (Tmax). Thus, across species, increases in HSPs and Tmax were positively correlated, but inversely related to increases in isoprene emissions. Integration of leaf gas‐exchange, isoprene emissions, proteomics, and respiratory thermotolerance measurements provided new insight into mechanisms underlying variability in tree species heatwave tolerance. Importantly, warm‐grown seedlings were, surprisingly, more susceptible to heatwave damage than cool‐grown seedlings, which could be associated with reduced enzyme concentrations in leaves. We conclude that species with restricted range sizes, along with trees growing under climate warming, may be more vulnerable to heatwaves of the future. [ABSTRACT FROM AUTHOR]

Highlights • Leastcost and coordination hypotheses support species strategy and ecosystem function. • An alternative framework of classifying dynamic roles based in plant traits. • Trait variation was linked to climate within a narrow tropical climatic range. • Optimality concepts and forest dynamics may represent PFTs more faithfully in DGVMs. Abstract The conventional representation of Plant Functional Types (PFTs) in Dynamic Global Vegetation Models (DGVMs) is increasingly recognized as simplistic and lacking in predictive power. Key ecophysiological traits, including photosynthetic parameters, are typically assigned single values for each PFT while the substantial trait variation within PFTs is neglected. This includes continuous variation in response to environmental factors, and differences linked to spatial and temporal niche differentiation within communities. A much stronger empirical basis is required for the treatment of continuous plant functional trait variation in DGVMs. We analyse 431 sets of measurements of leaf and plant traits, including photosynthetic measurements, on evergreen angiosperm trees in tropical moist forests of Australia and China. Confining attention to tropical moist forests, our analysis identifies trait differences that are linked to vegetation dynamic roles. Coordination theory predicts that Rubisco- and electron-transport limited rates of photosynthesis are co-limiting under field conditions. The least-cost hypothesis predicts that air-to-leaf CO 2 drawdown minimizes the combined costs per unit carbon assimilation of maintaining carboxylation and transpiration capacities. Aspects of these predictions are supported for within-community trait variation linked to canopy position, just as they are for variation along spatial environmental gradients. Trait differences among plant species occupying different structural and temporal niches may provide a basis for the ecophysiological representation of vegetation dynamics in next-generation DGVMs. [ABSTRACT FROM AUTHOR]

Phosphorus (P) limitation is known to have substantial impacts on leaf metabolism. However, uncertainty remains around whether P deficiency alters scaling functions linking leaf metabolism to associated traits.Weinvestigated the effect of P deficiency on leaf gas exchange and related leaf traits in 17 contrasting Eucalyptus species that exhibit inherent differences in leaf traits. Saplings were grown under controlled-environment conditions in a glasshouse, where they were subjected to minus and plus P treatments for 15 weeks. P deficiency decreased P concentrations and increased leaf mass per area (LMA) of newly-developed leaves. Rates of photosynthesis (A) and respiration (R) were also reduced in P-deficient plants compared with P-fertilised plants. By contrast, P deficiency had little effect on the temperature sensitivity of R. Irrespective of P treatment, on a log-log basis A and R scaled positively with increasing leaf nitrogen concentration [N] and negatively with increasing LMA. Although P deficiency had limited impact on A-R-LMA relationships, rates of CO2 exchange per unit N were consistently lower in P-deficient plants. Our results highlight the importance of P supply for leaf carbon metabolism and show howP deficiencies (i.e. when excluding confounding genotypic and environmental effects) can have a direct effect on commonly used leaf trait scaling relationships. [ABSTRACT FROM AUTHOR]

Heatwaves are likely to increase in frequency and intensity with climate change, which may impair tree function and forest C uptake. However, we have little information regarding the impact of extreme heatwaves on the physiological performance of large trees in the field. Here, we grew Eucalyptus parramattensis trees for 1 year with experimental warming (+3°C) in a field setting, until they were greater than 6 m tall. We withheld irrigation for 1 month to dry the surface soils and then implemented an extreme heatwave treatment of 4 consecutive days with air temperatures exceeding 43°C, while monitoring whole-canopy exchange of CO2 and H2O, leaf temperatures, leaf thermal tolerance, and leaf and branch hydraulic status. The heatwave reduced midday canopy photosynthesis to near zero but transpiration persisted, maintaining canopy cooling. A standard photosynthetic model was unable to capture the observed decoupling between photosynthesis and transpiration at high temperatures, suggesting that climate models may underestimate a moderating feedback of vegetation on heatwave intensity. The heatwave also triggered a rapid increase in leaf thermal tolerance, such that leaf temperatures observed during the heatwave were maintained within the thermal limits of leaf function. All responses were equivalent for trees with a prior history of ambient and warmed (+3°C) temperatures, indicating that climate warming conferred no added tolerance of heatwaves expected in the future. This coordinated physiological response utilizing latent cooling and adjustment of thermal thresholds has implications for tree tolerance of future climate extremes as well as model predictions of future heatwave intensity at landscape and global scales. [ABSTRACT FROM AUTHOR]

Ecosystem models commonly assume that key photosynthetic traits, such as carboxylation capacity measured at a standard temperature, are constant in time. The temperature responses of modelled photosynthetic or respiratory rates then depend entirely on enzyme kinetics. Optimality considerations, however, suggest this assumption may be incorrect. The "coordination hypothesis" (that Rubisco- and electron-transport-limited rates of photosynthesis are co-limiting under typical daytime conditions) predicts, instead, that carboxylation (Vcmax/ capacity should acclimate so that it increases somewhat with growth temperature but less steeply than its instantaneous response, implying that Vcmax when normalized to a standard temperature (e.g. 25 °C) should decline with growth temperature. With additional assumptions, similar predictions can be made for electron-transport capacity (Jmax/ and mitochondrial respiration in the dark (Rdark/. To explore these hypotheses, photosynthetic measurements were carried out on woody species during the warm and the cool seasons in the semi-arid Great Western Woodlands, Australia, under broadly similar light environments. A consistent proportionality between Vcmax and Jmax was found across species. Vcmax, Jmax and Rdark increased with temperature in most species, but their values standardized to 25 °C declined. The ci V ca ratio increased slightly with temperature. The leaf NV P ratio was lower in the warm season. The slopes of the relationships between log-transformed Vcmax and Jmax and temperature were close to values predicted by the coordination hypothesis but shallower than those predicted by enzyme kinetics. [ABSTRACT FROM AUTHOR]

Drought‐induced reduction of leaf gas exchange entails a complex regulation of the plant leaf metabolism. We used a combined molecular and physiological approach to understand leaf photosynthetic and respiratory responses of 2‐year‐old Quercus ilex seedlings to drought. Mild drought stress resulted in glucose accumulation while net photosynthetic CO2 uptake (Pn) remained unchanged, suggesting a role of glucose in stress signaling and/or osmoregulation. Simple sugars and sugar alcohols increased throughout moderate‐to‐very severe drought stress conditions, in parallel to a progressive decline in Pn and the quantum efficiency of photosystem II; by contrast, minor changes occurred in respiration rates until drought stress was very severe. At very severe drought stress, 2‐oxoglutarate dehydrogenase complex gene expression significantly decreased, and the abundance of most amino acids dramatically increased, especially that of proline and γ‐aminobutyric acid (GABA) suggesting enhanced protection against oxidative damage and a reorganization of the tricarboxylic cycle acid cycle via the GABA shunt. Altogether, our results point to Q. ilex drought tolerance being linked to signaling and osmoregulation by hexoses during early stages of drought stress, and enhanced protection against oxidative damage by polyols and amino acids under severe drought stress. [ABSTRACT FROM AUTHOR]

Contents986I.987II.987III.988IV.991V.992VI.995VII.997VIII.998References998 Summary: It has been 75 yr since leaf respiratory metabolism in the light (day respiration) was identified as a low‐flux metabolic pathway that accompanies photosynthesis. In principle, it provides carbon backbones for nitrogen assimilation and evolves CO2 and thus impacts on plant carbon and nitrogen balances. However, for a long time, uncertainties have remained as to whether techniques used to measure day respiratory efflux were valid and whether day respiration responded to environmental gaseous conditions. In the past few years, significant advances have been made using carbon isotopes, ‘omics’ analyses and surveys of respiration rates in mesocosms or ecosystems. There is substantial evidence that day respiration should be viewed as a highly dynamic metabolic pathway that interacts with photosynthesis and photorespiration and responds to atmospheric CO2 mole fraction. The view of leaf day respiration as a constant and/or negligible parameter of net carbon exchange is now outdated and it should now be regarded as a central actor of plant carbon‐use efficiency. [ABSTRACT FROM AUTHOR]

In the present study we investigated variations in leaf respiration in darkness (RD) and light (RL), and associated traits in response to season, and along a gradient of soil moisture, in Mediterranean woodland dominated by holm oak (Quercus ilex L.) in central and north-eastern Spain respectively. On seven occasions during the year in the central Spain site, and along the soil moisture gradient in north-eastern Spain, we measured rates of leaf RD, RL (using the Kok method), light-saturated photosynthesis (A) and related light response characteristics, leaf mass per unit area (MA) and leaf nitrogen (N) content. At the central Spain site, significant seasonal changes in soil water content and ambient temperature (T) were associated with changes in MA, foliar N, A and stomatal conductance. RD measured at the prevailing daily T and in instantaneous R-T responses, displayed signs of partial acclimation and was not significantly affected by time of year. RL was always less than, and strongly related to, RD, and RL/RD did not vary significantly or systematically with seasonal changes in T or soil water content. Averaged over the year, RL/RD was 0.66±0.05 s.e. (n = 14) at the central Spain site. At the north-eastern Spain site, the soil moisture gradient was characterised by increasing MA and RD, and reduced foliar N, A, and stomatal conductance as soil water availability decreased. Light inhibition of R occurred across all sites (mean RL/RD= 0.69±0.01 s.e. (n = 18)), resulting in ratios of RL/A being lower than for RD/A. Importantly, the degree of light inhibition was largely insensitive to changes in soil water content. Our findings provide evidence for a relatively constrained degree of light inhibition of R (RL/RD ~ 0.7, or inhibition of ~30%) across gradients of water availability, although the combined impacts of seasonal changes in both T and soil water content increase the range of values expressed. The findings thus have implications in terms of the assumptions made by predictive models that seek to account for light inhibition of R, and for our understanding of how environmental gradients impact on leaf trait relationships in Mediterranean plant communities. [ABSTRACT FROM AUTHOR]

Ecosystem models commonly assume that key photosynthetic traits, such as carboxylation-capacity measured at a standard temperature, are constant in time. The temperature responses of modelled photosynthetic/respiratory rates then depend entirely on enzyme kinetics. Optimality considerations suggest this assumption may be incorrect. The 'co-ordination hypothesis' (that Rubisco- and electron-transport limited rates of photosynthesis are co-limiting under typical daytime conditions) predicts instead that carboxylation (Vcmax) and light-harvesting (Jmax) capacities, and mitochondrial respiration in the dark (Rdark), should acclimate so that they increase with growth temperature - but less steeply than their instantaneous response rates. To explore this hypothesis, photosynthetic measurements were carried out on woody species during the warm and the cool seasons in the semi-arid Great Western Woodlands, Australia, under broadly similar light environments. A consistent linear relationship between Vcmax and Jmax was found across species. Vcmax, Jmax and Rdark increased with temperature, but values standardized to 25 °C declined. The ci:ca ratio increased slightly with temperature. The leaf N:P ratio was lower in the warm season. The slopes of the relationships of log-transformed Vcmax and Jmax to temperature were close to values predicted by the co-ordination hypothesis, but shallower than those predicted by enzyme kinetics. [ABSTRACT FROM AUTHOR]

Nitrogen (N) and phosphorus (P) have key roles in leaf metabolism, resulting in a strong coupling of chemical composition traits to metabolic rates in field-based studies. However, in such studies, it is difficult to disentangle the effects of nutrient supply per se on trait-trait relationships., Our study assessed how high and low N (5 mM and 0.4 mM, respectively) and P (1 mM and 2 μM, respectively) supply in 37 species from six plant functional types (PTFs) affected photosynthesis ( A) and respiration ( R) (in darkness and light) in a controlled environment., Low P supply increased scaling exponents (slopes) of area-based log-log A-N or R-N relationships when N supply was not limiting, whereas there was no P effect under low N supply. By contrast, scaling exponents of A-P and R-P relationships were altered by P and N supply. Neither R : A nor light inhibition of leaf R was affected by nutrient supply. Light inhibition was 26% across nutrient treatments; herbaceous species exhibited a lower degree of light inhibition than woody species., Because N and P supply modulates leaf trait−trait relationships, the next generation of terrestrial biosphere models may need to consider how limitations in N and P availability affect trait−trait relationships when predicting carbon exchange. [ABSTRACT FROM AUTHOR]

Leaf dark respiration ( Rdark) represents an important component controlling the carbon balance in tropical forests. Here, we test how nitrogen (N) and phosphorus (P) affect Rdark and its relationship with photosynthesis using three widely separated tropical forests which differ in soil fertility., Rdark was measured on 431 rainforest canopy trees, from 182 species, in French Guiana, Peru and Australia. The variation in Rdark was examined in relation to leaf N and P content, leaf structure and maximum photosynthetic rates at ambient and saturating atmospheric CO2 concentration., We found that the site with the lowest fertility (French Guiana) exhibited greater rates of Rdark per unit leaf N, P and photosynthesis. The data from Australia, for which there were no phylogenetic overlaps with the samples from the South American sites, yielded the most distinct relationships of Rdark with the measured leaf traits., Our data indicate that no single universal scaling relationship accounts for variation in Rdark across this large biogeographical space. Variability between sites in the absolute rates of Rdark and the Rdark : photosynthesis ratio were driven by variations in N- and P-use efficiency, which were related to both taxonomic and environmental variability. [ABSTRACT FROM AUTHOR]

High-temperature tolerance in plants is important in a warming world, with extreme heat waves predicted to increase in frequency and duration, potentially leading to lethal heating of leaves. Global patterns of high-temperature tolerance are documented in animals, but generally not in plants, limiting our ability to assess risks associated with climate warming. To assess whether there are global patterns in high-temperature tolerance of leaf metabolism, we quantified Tcrit (high temperature where minimal chlorophyll a fluorescence rises rapidly and thus photosystem II is disrupted) and Tmax (temperature where leaf respiration in darkness is maximal, beyond which respiratory function rapidly declines) in upper canopy leaves of 218 plant species spanning seven biomes. Mean site-based Tcrit values ranged from 41.5 °C in the Alaskan arctic to 50.8 °C in lowland tropical rainforests of Peruvian Amazon. For Tmax, the equivalent values were 51.0 and 60.6 °C in the Arctic and Amazon, respectively. Tcrit and Tmax followed similar biogeographic patterns, increasing linearly ( ˜8 °C) from polar to equatorial regions. Such increases in high-temperature tolerance are much less than expected based on the 20 °C span in high-temperature extremes across the globe. Moreover, with only modest high-temperature tolerance despite high summer temperature extremes, species in mid-latitude (~20-50°) regions have the narrowest thermal safety margins in upper canopy leaves; these regions are at the greatest risk of damage due to extreme heat-wave events, especially under conditions when leaf temperatures are further elevated by a lack of transpirational cooling. Using predicted heat-wave events for 2050 and accounting for possible thermal acclimation of Tcrit and Tmax, we also found that these safety margins could shrink in a warmer world, as rising temperatures are likely to exceed thermal tolerance limits. Thus, increasing numbers of species in many biomes may be at risk as heat-wave events become more severe with climate change. [ABSTRACT FROM AUTHOR]

We measured a diverse range of foliar characteristics in shrub and tree species in temperate rainforest communities along a soil chronosequence (six sites from 8 to 120 000 years) and used multilevel model analysis to attribute the proportion of variance for each trait into genetic (G, here meaning species-level), environmental (E) and residual error components. We hypothesised that differences in leaf traits would be driven primarily by changes in soil nutrient availability during ecosystem progression and retrogression. Several leaf structural, chemical and gas-exchange traits were more strongly driven by G than E effects. For leaf mass per unit area (MA), foliar [N], net CO2 assimilation and dark respiration rates and foliar carbohydrate concentration, the G component accounted for 60-87% of the total variance, with the variability associated with plot, the E effect, much less important. Other traits, such as foliar [P] and N: P, displayed strong E and residual effects. Analyses revealed significant reductions in the slopes of G-only bivariate relationships when compared with raw relationships, indicating that a large proportion of trait-trait relationships is species based, and not a response to environment per se. This should be accounted for when assessing the mechanistic basis for using such relationships in order to make predictions of responses of plants to short-term environmental change. [ABSTRACT FROM AUTHOR]

In intact leaves, mitochondrial populations are highly heterogeneous among contrasting cell types; how such contrasting populations respond to sustained changes in the environment remains, however, unclear. Here, we examined respiratory rates, mitochondrial protein composition and response to growth temperature in photosynthetic (mesophyll) and non-photosynthetic (epidermal) cells from fully expanded leaves of warm-developed ( WD) and cold-developed ( CD) broad bean ( V icia faba L.). Rates of respiration were significantly higher in mesophyll cell protoplasts ( MCPs) than epidermal cell protoplasts ( ECPs), with both protoplast types exhibiting capacity for cytochrome and alternative oxidase activity. Compared with ECPs, MCPs contained greater relative quantities of porin, suggesting higher mitochondrial surface area in mesophyll cells. Nevertheless, the relative quantities of respiratory proteins (normalized to porin) were similar in MCPs and ECPs, suggesting that ECPs have lower numbers of mitochondria yet similar protein complement to MCP mitochondria (albeit with lower abundance serine hydroxymethyltransferase). Several mitochondrial proteins (both non-photorespiratory and photorespiratory) exhibited an increased abundance in response to cold in both protoplast types. Based on estimates of individual protoplast respiration rates, combined with leaf cell abundance data, epidermal cells make a small but significant (2%) contribution to overall leaf respiration which increases twofold in the cold. Taken together, our data highlight the heterogeneous nature of mitochondrial populations in leaves, both among contrasting cell types and in how those populations respond to growth temperature. [ABSTRACT FROM AUTHOR]