Critical Leaf Water Content for Maize Photosynthesis under Drought Stress and Its Response to Rewatering

Crop photosynthesis is closely related to leaf water content (LWC), and clarifying the LWC conditions at critical points in crop photosynthesis has great theoretical and practical value for accurately monitoring drought and providing early drought warnings. This experiment was conducted to study the response of LWC to drought and rewatering and to determine the LWC at which maize photosynthesis reaches a maximum and minimum and thus changes from a state of stomatal limitation (SL) to non-stomatal limitation (NSL). The effects of rehydration were different after different levels of drought stress intensity at different growth stages, and the maize LWC recovered after rewatering following different drought stresses at the jointing stage; however, the maize LWC recovered more slowly after rewatering following 43 days and 36 days of drought stress at the tasselling and silking stages, respectively. The LWC when maize photosynthesis changed from SL to NSL was 75.4% ± 0.38%, implying that the maize became rehydrated under physiologically impaired conditions. The LWCs at which the maize Vcmax25 reached maximum values and zero differed between the drought and rewatering periods. After exposure to drought stress, the maize exhibited enhanced drought stress tolerance, an obviously reduced suitable water range, and significantly weakened photosynthetic capacity. These results provide profound insight into the turning points in maize photosynthesis and their responses to drought and rewatering. They may also help to improve crop water management, which will be useful in coping with the increased frequency of drought and extreme weather events expected under global climate change.

Lima Bean Growth, Leaf Stomatal and Nonstomatal Limitations to Photosynthesis, and 13C Discrimination in Response to Saline Irrigation

Soil salinization is a widespread problem severely impacting crop production. Understanding how salt stress affects growth-controlling photosynthetic performance is essential for improving crop salt tolerance and alleviating the salt impact. Lima bean (Phaseolus lunatus) is an important crop, but little information is available on its growth and leaf gas exchange in relation to a wide range of salinity. In this study, the responses of leaf gas exchange and whole plant growth of lima bean (cv. Fordhook 242) to six salinities with electrical conductivity (EC) of 2.9 (control), 5.7, 7.8, 10.0, 13.0, and 15.5 dS·m−1 in irrigation waters were assessed. Significant linear reduction by increasing salinity was observed on plant biomass, bean yield, and leaf net carbon assimilation rate (A). As EC increased from the control to 15.5 dS·m−1, plant biomass and A decreased by 87% and 69%, respectively, at the vegetative growth stage, and by 96% and 83%, respectively, at the pod growth stage, and bean yield decreased by 98%. Judged by the linear relations, the reduction in A accounted for a large portion of the growth reduction and bean yield loss. Salinity also had a significantly negative and linear effect on leaf stomatal conductance (gS). Leaf intercellular CO2 concentration (Ci) and leaf C13 isotope discrimination (Δ13) declined in parallel significantly with increasing salinity. The A-Ci curve analysis revealed that stomatal limitation [Lg (percent)] to A increased significantly and linearly, from 18% to 78% and from 22% to 87% at the vegetative and pod-filling stages, respectively, as EC increased from the control to the highest level. Thus, relatively nonstomatal or biochemical limitation [Lm (percent), Lm = 100 − Lg] to A responded negatively to increasing salinity. This result is coincident with the observed Δ13 salt-response trend. Furthermore, leaf carboxylation efficiency and CO2-saturated photosynthetic capacity [maximum A (Amax)] were unaffected by increasing salinity. Our results strongly indicate that the reduction in lima bean A by salt stress was mainly due to stomatal limitation and biochemical properties for photosynthesis might not be impaired. Because stomatal limitation reduces A exactly from lowering CO2 availability to leaves, increasing CO2 supply with an elevated CO2 concentration may raise A of the salt-stressed lima bean leaves and alleviate the salt impact. This is supported by our finding that the external CO2 concentration for 50% of Amax increased significantly and linearly with increasing salinity at the both growth stages. Leaf water use efficiency showed an increasing trend and no evident decline in leaf chlorophyll soil plant analysis development (SPAD) readings was observed as salinity increased.

Leaf Gas Exchange of Tomato Depends on Abscisic Acid and Jasmonic Acid in Response to Neighboring Plants under Different Soil Nitrogen Regimes

High planting density and nitrogen shortage are two important limiting factors for crop yield. Phytohormones, abscisic acid (ABA), and jasmonic acid (JA), play important roles in plant growth. A pot experiment was conducted to reveal the role of ABA and JA in regulating leaf gas exchange and growth in response to the neighborhood of plants under different nitrogen regimes. The experiment included two factors: two planting densities per pot (a single plant or four competing plants) and two N application levels per pot (1 and 15 mmol·L−1). Compared to when a single plant was grown per pot, neighboring competition decreased stomatal conductance (gs), transpiration (Tr) and net photosynthesis (Pn). Shoot ABA and JA and the shoot-to-root ratio increased in response to neighbors. Both gs and Pn were negatively related to shoot ABA and JA. In addition, N shortage stimulated the accumulation of ABA in roots, especially for competing plants, whereas root JA in competing plants did not increase in N15. Pearson’s correlation coefficient (R2) of gs to ABA and gs to JA was higher in N1 than in N15. As compared to the absolute value of slope of gs to shoot ABA in N15, it increased in N1. Furthermore, the stomatal limitation and non-stomatal limitation of competing plants in N1 were much higher than in other treatments. It was concluded that the accumulations of ABA and JA in shoots play a coordinating role in regulating gs and Pn in response to neighbors; N shortage could intensify the impact of competition on limiting carbon fixation and plant growth directly.

Drought stress strengthens the link between chlorophyll fluorescence parameters and photosynthetic traits

Chlorophyll fluorescence (ChlF) has been used to understand photosynthesis and its response to climate change, particularly with satellite-based data. However, it remains unclear how the ChlF ratio and photosynthesis are linked at the leaf level under drought stress. Here, we examined the link between ChlF ratio and photosynthesis at the leaf level by measuring photosynthetic traits, such as net CO2 assimilation rate (An), the maximum carboxylation rate of Rubisco (Vcmax), the maximum rate of electron transport (Jmax), stomatal conductance (gs) and total chlorophyll content (Chlt). The ChlF ratio of the leaf level such as maximum quantum efficiency of PSII (Fv/Fm) is based on fluorescence kinetics. ChlF intensity ratio (LD685/LD740) based on spectrum analysis was obtained. We found that a combination of the stomatal limitation, non-stomatal limitation, and Chlt regulated leaf photosynthesis under drought stress, while Jmax and Chlt governed the ChlF ratio. A significant link between the ChlF ratio and An was found under drought stress while no significant correlation in the control, which indicated that drought stress strengthens the link between the ChlF ratio and photosynthetic traits. These results suggest that the ChlF ratio can be a powerful tool to track photosynthetic traits of terrestrial ecosystems under drought stress.

Incorporating non-stomatal limitation improves the performance of leaf and canopy models at high vapour pressure deficit

Vapour pressure deficit (D) is projected to increase in the future as temperatures rise. In response to increased D, stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water, and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A-gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A-gs models (Leuning 1995 and Medlyn et al. 2011) were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs, two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. While the model incorporating hydraulic limitation captured the observed A and gs, it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model.

Stomatal and non-stomatal limitations to photosynthesis of Populus euphratica Oliv. leaves in an extremely arid area of northwestern China

On the basis of successive measurements of leaf gas exchange during the main growing seasons of Populus euphratica Oliv. in 2013 and 2014, respectively, we analyzed the stomatal and non-stomatal limitations to photosynthesis under natural conditions in an extremely arid region of northwestern China. Our results showed that (1) the distribution patterns of net photosynthesis (Pn) and stomatal conductance (gs) were similar, both of which increased in the morning, peaked at around noon, and then decreased. This contrasted with the observed changes in sub-stomatal CO2 concentrations (Ci). (2) The phenomenon of midday depression of photosynthesis (MDP) was obvious from July to September during the two years. At the beginning of MDP, the stomatal limitation to photosynthesis (Ls) peaked, where its predominance was supported by Ci being at a minimum. Thereafter, Ls decreased and Ci/gs increased sharply, indicating that the non-stomatal limitation to photosynthesis predominated. (3) Both the Ls and relative stomatal limitation to photosynthesis (RLs) increased in the morning, and then decreased, whereas Ci/gs presented contrary changes. (4) The RLs values were greater than the Ls values, which was mainly due to the nonlinearity of the Pn/Ci curve, which often leads to large overestimations. (5) The Ls values in our study were much greater than those from other studies under natural conditions. The most probable reason was that the extremely high temperature and scarce water resource caused the stomata to close to reduce transpiration, resulting in the stomatal limitation to photosynthesis being more intense.