Temperature responses of leaf respiration in light and darkness are similar and modulated by leaf development.

Our ability to predict temperature responses of leaf respiration in light and darkness (RL and RDk ) is essential to models of global carbon dynamics. While many models rely on constant thermal sensitivity (characterized by Q10 ), uncertainty remains as to whether Q10 of RL and RDk are actually similar. We measured short-term temperature responses of RL and RDk in immature and mature leaves of two evergreen tree species, Castanopsis carlesii and Ormosia henry in an open field. RL was estimated by the Kok method, the Yin method and a newly developed Kok-iterCc method. When estimated by the Yin and Kok-iterCc methods, RL and RDk had similar Q10 (c. 2.5). The Kok method overestimated both Q10 and the light inhibition of respiration. RL /RDk was not affected by leaf temperature. Acclimation of respiration in summer was associated with a decline in basal respiration but not in Q10 in both species, which was related to changes in leaf nitrogen content between seasons. Q10 of RL and RDk in mature leaves were 40% higher than in immature leaves. Our results suggest similar Q10 values can be used to model RL and RDk while leaf development-associated changes in Q10 require special consideration in future respiration models.

Short- and long-term responses of leaf day respiration to elevated atmospheric CO2.

Evaluating leaf day respiration rate (RL), which is believed to differ from that in the dark (RDk), is essential for predicting global carbon cycles under climate change. Several studies have suggested that atmospheric CO2 impacts RL. However, the magnitude of such an impact and associated mechanisms remain uncertain. To explore the CO2 effect on RL, wheat (Triticum aestivum) and sunflower (Helianthus annuus) plants were grown under ambient (410 ppm) and elevated (820 ppm) CO2 mole fraction ([CO2]). RL was estimated from combined gas exchange and chlorophyll fluorescence measurements using the Kok method, the Kok-Phi method, and a revised Kok method (Kok-Cc method). We found that elevated growth [CO2] led to an 8.4% reduction in RL and a 16.2% reduction in RDk in both species, in parallel to decreased leaf N and chlorophyll contents at elevated growth [CO2]. We also looked at short-term CO2 effects during gas exchange experiments. Increased RL or RL/RDk at elevated measurement [CO2] were found using the Kok and Kok-Phi methods, but not with the Kok-Cc method. This discrepancy was attributed to the unaccounted changes in Cc in the former methods. We found that the Kok and Kok-Phi methods underestimate RL and overestimate the inhibition of respiration under low irradiance conditions of the Kok curve, and the inhibition of RL was only 6%, representing 26% of the apparent Kok effect. We found no significant long-term CO2 effect on RL/RDk, originating from a concurrent reduction in RL and RDk at elevated growth [CO2], and likely mediated by acclimation of nitrogen metabolism.

Overestimated gains in water-use efficiency by global forests.

Increases in terrestrial water-use efficiency (WUE) have been reported in many studies, pointing to potential changes in physiological forcing of global carbon and hydrological cycles. However, gains in WUE are of uncertain magnitude over longer (i.e. >10 years) periods of time largely owing to difficulties in accounting for structural and physiological acclimation. 13 C signatures (i.e. δ13 C) of plant organic matter have long been used to estimate WUE at temporal scales ranging from days to centuries. Mesophyll conductance is a key uncertainty in estimated WUE owing to its influence on diffusion of CO2 to sites of carboxylation. Here we apply new knowledge of mesophyll conductance to 464 δ13 C chronologies in tree-rings of 143 species spanning global biomes. Adjusted for mesophyll conductance, gains in WUE during the 20th century (0.15 ppm year-1 ) were considerably smaller than those estimated from conventional modelling (0.26 ppm year-1 ). Across the globe, mean sensitivity of WUE to atmospheric CO2 was 0.15 ppm ppm-1 . Ratios of internal-to-atmospheric CO2 (on a mole fraction basis; ci /ca ) in leaves were mostly constant over time but differed among biomes and plant taxa-highlighting the significance of both plant structure and physiology. Together with synchronized responses in stomatal and mesophyll conductance, our results suggest that ratios of chloroplastic-to-atmospheric CO2 (cc /ca ) are constrained over time. We conclude that forest WUE may have not increased as much as previously suggested and that projections of future climate forcing via CO2 fertilization may need to be adjusted accordingly.

Estimation of intrinsic water-use efficiency from δ13C signature of C3 leaves: Assumptions and uncertainty.

Carbon isotope composition (δ13C) has been widely used to estimate the intrinsic water-use efficiency (iWUE) of plants in ecosystems around the world, providing an ultimate record of the functional response of plants to climate change. This approach relies on established relationships between leaf gas exchange and isotopic discrimination, which are reflected in different formulations of 13C-based iWUE models. In the current literature, most studies have utilized the simple, linear equation of photosynthetic discrimination to estimate iWUE. However, recent studies demonstrated that using this linear model for quantitative studies of iWUE could be problematic. Despite these advances, there is a scarcity of review papers that have comprehensively reviewed the theoretical basis, assumptions, and uncertainty of 13C-based iWUE models. Here, we 1) present the theoretical basis of 13C-based iWUE models: the classical model (iWUEsim), the comprehensive model (iWUEcom), and the model incorporating mesophyll conductance (iWUEmes); 2) discuss the limitations of the widely used iWUEsim model; 3) and make suggestions on the application of the iWUEmes model. Finally, we suggest that a mechanistic understanding of mesophyll conductance associated effects and post-photosynthetic fractionation are the bottlenecks for improving the 13C-based estimation of iWUE.

Accounting for mesophyll conductance substantially improves 13 C-based estimates of intrinsic water-use efficiency.

Carbon isotope discrimination (Δ) has been used widely to infer intrinsic water-use efficiency (iWUE) of C3 plants, a key parameter linking carbon and water fluxes. Despite the essential role of mesophyll conductance (gm ) in photosynthesis and Δ, its effect on Δ-based predictions of iWUE has generally been neglected. Here, we derive a mathematical expression of iWUE as a function of Δ that includes gm (iWUEmes ) and exploits the gm -stomatal conductance (gsc ) relationship across drought-stress levels and plant functional groups (deciduous or semideciduous woody, evergreen woody and herbaceous species) in a global database. iWUEmes was further validated with an independent dataset of online-Δ and CO2 and H2 O gas exchange measurements with seven species. Drought stress reduced gsc and gm by nearly one-half across all plant functional groups, but had no significant effect on the gsc  : gm ratio, with a well supported value of 0.79 ± 0.07 (95% CI, n = 198). gm was negatively correlated to iWUE. Incorporating the gsc  : gm ratio greatly improved estimates of iWUE, compared with calculations that assumed infinite gm . The inclusion of the gsc  : gm ratio, fixed at 0.79 when gm was unknown, proved desirable to eliminate significant errors in estimating iWUE from Δ across various C3 vegetation types.

[Mesophyll conductance to CO2: Methods and current knowledge].

Mesophyll conductance (gm), the total conductance of CO2 diffusion from substomatal cavity to the site of carboxylation within chloroplast, is a major limiting factor for photosynthesis and a key parameter for improving photosynthetic resource use efficiency of crops. Online 13C discrimination method is an important method for plant eco-physiological studies and a well-established method for measuring gm of C3 plants, although it has not been widely used due to challenges in methodology and high demands on experimental facilities. In this review, we summarized the characteristics of commonly used methods for gm, introduced the basic theory of the online 13C discrimination method, namely Farquhar's photosynthetic 13C discrimination model; systematically introduced the practical measurements, equations and the components of facilities; and reviewed the drivers for variation in gm of C3 plants. At the last part, we discussed the outlook of the development of methodology, new experimental protocols, and applications in measurement scenarios.