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The triose phosphate utilization (TPU) rate has been identified as one of the processes that can limit terrestrial plant photosynthesis. However, we lack a robust quantitative assessment of TPU limitation of photosynthesis at the global scale. As a result, TPU, and its potential limitation of photosynthesis, is poorly represented in terrestrial biosphere models (TBMs). In this study, we utilized a global data set of photosynthetic CO2 response curves representing 141 species from tropical rainforests to Arctic tundra. We quantified TPU by fitting the standard biochemical model of C3 photosynthesis to measured photosynthetic CO2 response curves and characterized its instantaneous temperature response. Our results demonstrate that TPU does not limit leaf photosynthesis at the current ambient atmospheric CO2 concentration. Furthermore, our results showed that the light-saturated photosynthetic rates of plants growing in cold environments are not more often limited by TPU than those of plants growing in warmer environments. In addition, our study showed that the instantaneous temperature response of TPU is distinct from temperature response of the maximum rate of Rubisco carboxylation. The new formulations of the temperature response of TPU derived in this study may prove useful in quantifying the biochemical limits to terrestrial plant photosynthesis and improve the representation of plant photosynthesis in TBMs.

Plant hydraulic traits capture the impacts of drought stress on plant function, yet vegetation models lack sufficient information regarding trait coordination and variation with climate-of-origin across species. Here, we investigated key hydraulic and carbon economy traits of 12 woody species in Australia from a broad climatic gradient, with the aim of identifying the coordination among these traits and the role of climate in shaping cross-species trait variation. The influence of environmental variation was minimized by a common garden approach, allowing us to factor out the influence of environment on phenotypic variation across species. We found that hydraulic traits (leaf turgor loss point, stomatal sensitivity to drought [Pgs ], xylem vulnerability to cavitation [Px ], and branch capacitance [Cbranch ]) were highly coordinated across species and strongly related to rainfall and aridity in the species native distributional range. In addition, trade-offs between drought tolerance and plant growth rate were observed across species. Collectively, these results provide critical insight into the coordination among hydraulic traits in modulating drought adaptation and will significantly advance our ability to predict drought vulnerability in these dominant trees species.

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.

Models of stomatal conductance (g(s)) are based on coupling between g(s) and CO(2) assimilation (A(net)), and it is often assumed that the slope of this relationship ('g(1) ') is constant across species. However, if different plant species have adapted to different access costs of water, then there will be differences in g(1) among species. We hypothesized that g(1) should vary among species adapted to different climates, and tested the theory and its linkage to plant hydraulics using four Eucalyptus species from different climatic origins in a common garden. Optimal stomatal theory predicts that species from sub-humid zones have a lower marginal water cost of C gain, hence lower g(1) than humid-zone species. In agreement with the theory that g(1) is related to tissue carbon costs for water supply, we found a relationship between wood density and g(1) across Eucalyptus species of contrasting climatic origins. There were significant reductions in the parameter g(1) during drought in humid but not sub-humid species, with the latter group maintaining g(1) in drought. There are strong differences in stomatal behaviour among related tree species in agreement with optimal stomatal theory, and these differences are consistent with the economics involved in water uptake and transport for carbon gain.

Under elevated atmospheric CO2 concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO2 effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta-analysis to test the hypotheses that: (1) elevated atmospheric CO2 stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO2 induces a C allocation shift towards below-ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO2. Soil N concentration strongly interacted with CO2 fumigation: the effect of elevated CO2 on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO2 are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.

The temperature dependence of C 3 photosynthesis is known to vary with growth environment and with species. In an attempt to quantify this variability, a commonly used biochemically based photosynthesis model was parameterized from 19 gas exchange studies on tree and crop species. The parameter values obtained described the shape and amplitude of the temperature responses of the maximum rate of Rubisco activity ( V cmax ) and the potential rate of electron transport ( J max ). Original data sets were used for this review, as it is shown that derived values of V cmax and its temperature response depend strongly on assumptions made in derivation. Values of J max and V cmax at 25 ° C varied considerably among species but were strongly correlated, with an average J max : V cmax ratio of 1·67. Two species grown in cold climates, however, had lower ratios. In all studies, the J max : V cmax ratio declined strongly with measurement temperature. The relative temperature responses of J max and V cmax were relatively constant among tree species. Activation energies averaged 50 kJ mol − 1 for J max and 65 kJ mol − − − 1 for V cmax , and for most species temperature optima averaged 33 ° ° ° C for J max and 40 ° C for V cmax . However, the cold climate tree species had low temperature optima for both J max ( 19 ° C) and V cmax (29 ° ° ° C), suggesting acclimation of both processes to growth temperature. Crop species had somewhat different temperature responses, with higher activation energies for both J max and V cmax , implying narrower peaks in the temperature response for these species. The results thus suggest that both growth environment and

Responses of plant processes to temperature may vary according to the time scale on which they are measured. In this study, both short-term and seasonal responses of photosynthesis to temperature were examined. A field study of seasonal changes in the temperature response of photosynthesis was conducted on two provenances, French and Moroccan, of mature maritime pine ( Pinus pinaster Ait.). Measurements were made every 2 months over a 1-year period and used to parameterize a mechanistic model of photosynthesis. Temperature responses of maximum Rubisco activity, V cmax , and potential electron transport rate, J max , were obtained for each measurement period, as was the response of stomatal conductance, g s , to water vapour pressure deficit (VPD). Absolute values of V cmax and J max at 25 ° C were related to needle nitrogen content, N area. N area , and thus V cmax and J max , were negatively correlated with the mean minimum temperature in the month preceding measurements. The ratio of J max : V cmax at 25 ° C varied between 1 and 1·7 but did not show any seasonal trend. Nor was there any seasonal trend in the relative temperature response of V cmax , which had an activation energy H a of approximately 57 kJ mol − 1 throughout the experiment. The activation energy of J max was also close to constant throughout the experiment, averaging 39 kJ mol − 1 . For the French provenance, the optimal temperature of J max was positively correlated with the maximum temperature of the previous day, but no such correlation was found for the Moroccan provenance. The response of g s to VPD also varied seasonally, with much stronger stomatal closure in winter months. Taken together, these results implied a translational shift downwards of the photosynthetic temperature response curve with increasing T prev , and a shift in the temperature optimum of photosynthesis of 5‐10 ° C between summer and winter. These results illustrate that the short-term temperature response of photosynthesis varies significantly on a seasonal basis.

The effects of elevated atmospheric CO2 concentration on growth of forest tree species are difficult to predict because practical limitations restrict experiments to much shorter than the average life-span of a tree. Long-term, processbased computer models must be used to extrapolate from shorter-term experiments. A key problem is to ensure a strong flow of information between experiments and models. In this study, meta-analysis techniques were used to summarize a suite of photosynthetic model parameters obtained from 15 field-based elevated [CO2] experiments on European forest tree species. The parameters studied are commonly used in modelling photosynthesis, and include observed light-saturated photosynthetic rates (Amax), the potential electron transport rate (Jmax), the maximum Rubisco activity (Vcmax) and leaf nitrogen concentration on mass (Nm) and area (Na) bases. Across all experiments, light-saturated photosynthesis was strongly stimulated by growth in elevated [CO2]. However, significant down-regulation of photosynthesis was also observed; when measured at the same CO2 concentration, photosynthesis was reduced by 10‐20%. The underlying biochemistry of photosynthesis was affected, as shown by a down-regulation of the parameters Jmax and Vcmax of the order of 10%. This reduction in Jmax and Vcmax was linked to the effects of elevated [CO2] on leaf nitrogen concentration. It was concluded that the current model is adequate to model photosynthesis in elevated [CO2]. Tables of model parameter values for different European forest species are given.