Enhanced decreases in rice evapotranspiration in response to elevated atmospheric carbon dioxide under warmer environments.

A short period of exposure to elevated CO2 is known to decrease evapotranspiration via stomatal closure. Based on theoretical evaluation of a canopy transpiration model, we hypothesized that this decrease in the evapotranspiration of rice under elevated CO2 was greater under higher temperature conditions due to an increased sensitivity of transpiration to changes in CO2 induced by the greater vapour pressure deficit. In a temperature gradient chamber-based experiment, a 200 ppm increase in CO2 concentration led to 0.4 mm (-7%) and 1.5 mm (-15%) decreases in 12 h evapotranspiration under ambient temperature and high temperature (+3.7°C) conditions, respectively. Model simulations revealed that the greater vapour pressure deficit under higher temperature conditions explained the variations in the reduction of evapotranspiration observed under elevated CO2 levels between the temperature treatments. Our study suggests the utility of a simple modelling framework for mechanistic understanding of evapotranspiration and crop energy balance system under changing environmental conditions.

Alternative method for determining leaf CO2 assimilation without gas exchange measurements: Performance, comparison and sensitivity analysis.

We present an alternative method to determine leaf CO2 assimilation rate (An ), eliminating the need for gas exchange measurements in proximal and remote sensing. This method combines the Farquhar-von Caemmerer-Berry photosynthesis model with mechanistic light reaction (MLR) theory and leaf energy balance (EB) analysis. The MLR theory estimates the actual electron transport rate (J) by leveraging chlorophyll fluorescence via pulse amplitude-modulated fluorometry for proximal sensing or sun-induced chlorophyll fluorescence measurements for remote sensing, along with spectral reflectance. The EB equation is used to directly estimate stomatal conductance from leaf temperature. In wheat and soybean, the MLR-EB model successfully estimated An variations, including midday depression, under various environmental and phenological conditions. Sensitivity analysis revealed that the leaf boundary layer conductance (gb ) played an equal, if not more, crucial role compared to the variables for J. This was primarily caused by the indirect influence of gb through the EB equation rather than its direct impact on convective CO2 exchange on the leaf. Although the MLR-EB model requires an accurate estimation of gb , it can potentially reduce uncertainties and enhance applicability in photosynthesis assessment when gas exchange measurements are unavailable.

High-throughput characterization, correlation, and mapping of leaf photosynthetic and functional traits in the soybean (Glycine max) nested association mapping population.

Photosynthesis is a key target to improve crop production in many species including soybean [Glycine max (L.) Merr.]. A challenge is that phenotyping photosynthetic traits by traditional approaches is slow and destructive. There is proof-of-concept for leaf hyperspectral reflectance as a rapid method to model photosynthetic traits. However, the crucial step of demonstrating that hyperspectral approaches can be used to advance understanding of the genetic architecture of photosynthetic traits is untested. To address this challenge, we used full-range (500-2,400 nm) leaf reflectance spectroscopy to build partial least squares regression models to estimate leaf traits, including the rate-limiting processes of photosynthesis, maximum Rubisco carboxylation rate, and maximum electron transport. In total, 11 models were produced from a diverse population of soybean sampled over multiple field seasons to estimate photosynthetic parameters, chlorophyll content, leaf carbon and leaf nitrogen percentage, and specific leaf area (with R2 from 0.56 to 0.96 and root mean square error approximately <10% of the range of calibration data). We explore the utility of these models by applying them to the soybean nested association mapping population, which showed variability in photosynthetic and leaf traits. Genetic mapping provided insights into the underlying genetic architecture of photosynthetic traits and potential improvement in soybean. Notably, the maximum Rubisco carboxylation rate mapped to a region of chromosome 19 containing genes encoding multiple small subunits of Rubisco. We also mapped the maximum electron transport rate to a region of chromosome 10 containing a fructose 1,6-bisphosphatase gene, encoding an important enzyme in the regeneration of ribulose 1,5-bisphosphate and the sucrose biosynthetic pathway. The estimated rate-limiting steps of photosynthesis were low or negatively correlated with yield suggesting that these traits are not influenced by the same genetic mechanisms and are not limiting yield in the soybean NAM population. Leaf carbon percentage, leaf nitrogen percentage, and specific leaf area showed strong correlations with yield and may be of interest in breeding programs as a proxy for yield. This work is among the first to use hyperspectral reflectance to model and map the genetic architecture of the rate-limiting steps of photosynthesis.

Predicting biochemical acclimation of leaf photosynthesis in soybean under in-field canopy warming using hyperspectral reflectance.

Traditional gas exchange measurements are cumbersome, which makes it difficult to capture variation in biochemical parameters, namely the maximum rate of carboxylation measured at a reference temperature (Vcmax25 ) and the maximum electron transport at a reference temperature (Jmax25 ), in response to growth temperature over time from days to weeks. Hyperspectral reflectance provides reliable measures of Vcmax25 and Jmax25 ; however, the capability of this method to capture biochemical acclimations of the two parameters to high growth temperature over time has not been demonstrated. In this study, Vcmax25 and Jmax25 were measured over multiple growth stages during two growing seasons for field-grown soybeans using both gas exchange techniques and leaf spectral reflectance under ambient and four elevated canopy temperature treatments (ambient+1.5, +3, +4.5, and +6°C). Spectral vegetation indices and machine learning methods were used to build predictive models for Vcmax25 and Jmax25 , based on the leaf reflectance. Results showed that these models yielded an R2 of 0.57-0.65 and 0.48-0.58 for Vcmax25 and Jmax25 , respectively. Hyperspectral reflectance captured biochemical acclimation of leaf photosynthesis to high temperature in the field, improving spatial and temporal resolution in the ability to assess the impact of future warming on crop productivity.

Quantifying high-temperature stress on soybean canopy photosynthesis: The unique role of sun-induced chlorophyll fluorescence.

High temperature and accompanying high vapor pressure deficit often stress plants without causing distinctive changes in plant canopy structure and consequential spectral signatures. Sun-induced chlorophyll fluorescence (SIF), because of its mechanistic link with photosynthesis, may better detect such stress than remote sensing techniques relying on spectral reflectance signatures of canopy structural changes. However, our understanding about physiological mechanisms of SIF and its unique potential for physiological stress detection remains less clear. In this study, we measured SIF at a high-temperature experiment, Temperature Free-Air Controlled Enhancement, to explore the potential of SIF for physiological investigations. The experiment provided a gradient of soybean canopy temperature with 1.5, 3.0, 4.5, and 6.0°C above the ambient canopy temperature in the open field environments. SIF yield, which is normalized by incident radiation and the fraction of absorbed photosynthetically active radiation, showed a high correlation with photosynthetic light use efficiency (r = 0.89) and captured dynamic plant responses to high-temperature conditions. SIF yield was affected by canopy structural and plant physiological changes associated with high-temperature stress (partial correlation r = 0.60 and -0.23). Near-infrared reflectance of vegetation, only affected by canopy structural changes, was used to minimize the canopy structural impact on SIF yield and to retrieve physiological SIF yield (ΦF ) signals. ΦF further excludes the canopy structural impact than SIF yield and indicates plant physiological variability, and we found that ΦF outperformed SIF yield in responding to physiological stress (r = -0.37). Our findings highlight that ΦF sensitively responded to the physiological downregulation of soybean gross primary productivity under high temperature. ΦF , if reliably derived from satellite SIF, can support monitoring regional crop growth and different ecosystems' vegetation productivity under environmental stress and climate change.

Genome-wide association mapping for phenotypic plasticity in rice.

Phenotypic plasticity of plants in response to environmental changes is important for adapting to changing climate. Less attention has been paid to exploring the advantages of phenotypic plasticity in resource-rich environments to enhance the productivity of agricultural crops. Here, we examined genetic variation for phenotypic plasticity in indica rice (Oryza sativa L.) across two diverse panels: (1) a Phenomics of Rice Adaptation and Yield (PRAY) population comprising 301 accessions; and (2) a Multi-parent Advanced Generation Inter-Cross (MAGIC) indica population comprising 151 accessions. Altered planting density was used as a proxy for elevated atmospheric CO2 response. Low planting density significantly increased panicle weight per plant compared with normal density, and the magnitude of the increase ranged from 1.10 to 2.78 times among accessions for the PRAY population and from 1.05 to 2.45 times for the MAGIC population. Genome-wide-association studies validate three Environmental Responsiveness (ER) candidate alleles (qER1-3) that were associated with relative response of panicle weight to low density. Two of these alleles were tested in 13 genotypes to clarify their biomass responses during vegetative growth under elevated CO2 in Japan. Our study provides evidence for polymorphisms that control rice phenotypic plasticity in environments that are rich in resources such as light and CO2 .

Finlay-Wilkinson's regression coefficient as a pre-screening criterion for yield responsiveness to elevated atmospheric CO2 concentration in crops.

The rising atmospheric CO2 concentration ([CO2 ]) can increase crop productivity, but there are likely to be intraspecific variations in the response. To meet future world food demand, screening for genotypes with high [CO2 ] responsiveness will be a useful option, but there is no criterion for high [CO2 ] responsiveness. We hypothesized that the Finlay-Wilkinson regression coefficient (RC) (for the relationship between a genotype's yield versus the mean yield of all genotypes in a specific environment) could serve as a pre-screening criterion for identifying genotypes that respond strongly to elevated [CO2 ]. We collected datasets on the yield of 6 rice and 10 soybean genotypes along environmental gradients and compared their responsiveness to elevated [CO2 ] based on the regression coefficients (i.e. the increases of yield per 100 µmol mol-1 [CO2 ]) identified in previous reports. We found significant positive correlations between the RCs and the responsiveness of yield to elevated [CO2 ] in both rice and soybean. This result raises the possibility that the coefficient of the Finlay-Wilkinson relationship could be used as a pre-screening criterion for [CO2 ] responsiveness.

Phenotypic Plasticity Conditions the Response of Soybean Seed Yield to Elevated Atmospheric CO2 Concentration.

Selection for cultivars with superior responsiveness to elevated atmospheric CO2 concentrations (eCO2) is a powerful option for boosting crop productivity under future eCO2. However, neither criteria for eCO2 responsiveness nor prescreening methods have been established. The purpose of this study was to identify traits responsible for eCO2 responsiveness of soybean (Glycine max). We grew 12 Japanese and U.S. soybean cultivars that differed in their maturity group and determinacy under ambient CO2 and eCO2 for 2 years in temperature gradient chambers. CO2 elevation significantly increased seed yield per plant, and the magnitude varied widely among the cultivars (from 0% to 62%). The yield increase was best explained by increased aboveground biomass and pod number per plant. These results suggest that the plasticity of pod production under eCO2 results from biomass enhancement, and would therefore be a key factor in the yield response to eCO2, a resource-rich environment. To test this hypothesis, we grew the same cultivars at low planting density, a resource-rich environment that improved the light and nutrient supplies by minimizing competition. Low planting density significantly increased seed yield per plant, and the magnitude ranged from 5% to 105% among the cultivars owing to increased biomass and pod number per plant. The yield increase due to low-density planting was significantly positively correlated with the eCO2 response in both years. These results confirm our hypothesis and suggest that high plasticity of biomass and pod production at a low planting density reveals suitable parameters for breeding to maximize soybean yield under eCO2.

Planting geometry as a pre-screening technique for identifying CO2 responsive rice genotypes: a case study of panicle number.

Identifying CO(2) responsive genotypes is a major target for enhancing crop productivity under future global elevated atmospheric CO(2) concentration ([CO(2)]). However, [CO(2)]-fumigation facilities are extremely expensive and are not easily accessible, and are limited in space for large-scale screening. Hence, reliable donors for initiating [CO(2)]-responsive breeding programs are not in place for crops, including rice. We propose a simple and novel phenotyping method for identifying [CO(2)]-responsive genotypes, and quantify the responsiveness to low planting density over 4-year trials across both temperate and tropical conditions. Panicle number per plant is the key determinant of grain yield and hence was the focus trait across all our trials. In temperate climate, a 3-season field screening using 127 diverse rice genotypes and employing two planting densities (normal and low density) was conducted. Two japonica genotypes were selected based on their higher responsiveness to low planting density as candidates for validating the proposed phenotyping protocol as a pre-screen for [CO(2)]-responsiveness. The approach using the two selected candidates and three standard genotypes was confirmed using a free-air CO(2) enrichment facility and temperature gradient chambers under elevated [CO(2)]. In tropical climate, we grew three rice cultivars, previously identified for their [CO(2)]-responsiveness, at two planting densities. The experiments provided confirmation that responsiveness to low planting density was correlated with that of [CO(2)]-responsiveness across both the temperate and tropical conditions. The planting density would be useful pre-screening method for testing large panels of diverse germplasm at low cost complemented by available CO(2) -control facilities for final validation of candidates from the pre-screens.

Dorsoventral asymmetry of photosynthesis and photoinhibition in flag leaves of two rice cultivars that differ in nitrogen response and leaf angle.

Rice is believed to show photosynthetic symmetry between adaxial and abaxial leaf sides. To verify this, we re-examined dorsoventral asymmetry in photosynthesis, chlorophyll fluorescence and anatomical traits in flag leaves of two Oryza sativa cultivars that differ in nitrogen (N) response and in leaf angle: 'Akenohoshi', a cultivar that can adapt to low-N (LN), with low leaf angle (more erect leaves), and 'Shirobeniya', a cultivar that is unable to adapt to LN, with higher leaf angle. Plants were grown under standard-N (SN) and LN conditions. LN leaves of both cultivars became more erect than SN, but LN Akenohoshi still had more erect ones than Shirobeniya. Contrary to results of previous studies, leaves of both cultivars showed an asymmetry in photosynthetic rate between adaxial and abaxial sides (higher on the adaxial side) under SN. SN leaves of both cultivars showed lower susceptibility to photoinhibition on the adaxial side than on the abaxial side. However, leaves of Akenohoshi showed less asymmetry in these traits under LN than under SN, whereas leaves of Shirobeniya had similar degrees of asymmetry in these traits under both SN and LN. Both cultivars also showed dorsoventral asymmetry in anatomical traits of mesophyll tissue regardless of N level, but the degree of asymmetry was lower in LN Akenohoshi. These data reveal that rice leaves exhibit dorsoventral asymmetry in photosynthetic and anatomical features, and that the degree of asymmetry varies with cultivar and N level. It is suggested that lower leaf angles (particularly in Akenohoshi) in the presence of LN represent a light acclimation to prevent photoinhibition.

Ammonia emission from rice leaves in relation to photorespiration and genotypic differences in glutamine synthetase activity.

BACKGROUND AND AIMS: Rice (Oryza sativa) plants lose significant amounts of volatile NH(3) from their leaves, but it has not been shown that this is a consequence of photorespiration. Involvement of photorespiration in NH(3) emission and the role of glutamine synthetase (GS) on NH(3) recycling were investigated using two rice cultivars with different GS activities. METHODS: NH(3) emission (AER), and gross photosynthesis (P(G)), transpiration (Tr) and stomatal conductance (g(S)) were measured on leaves of 'Akenohoshi', a cultivar with high GS activity, and 'Kasalath', a cultivar with low GS activity, under different light intensities (200, 500 and 1000 µmol m(-2) s(-1)), leaf temperatures (27·5, 32·5 and 37·5 °C) and atmospheric O(2) concentrations ([O(2)]: 2, 21 and 40 %, corresponding to 20, 210 and 400 mmol mol(-1)). KEY RESULTS: An increase in [O(2)] increased AER in the two cultivars, accompanied by a decrease in P(G) due to enhanced photorespiration, but did not greatly influence Tr and g(S). There were significant positive correlations between AER and photorespiration in both cultivars. Increasing light intensity increased AER, P(G), Tr and g(S) in both cultivars, whereas increasing leaf temperature increased AER and Tr but slightly decreased P(G) and g(S). 'Kasalath' (low GS activity) showed higher AER than 'Akenohoshi' (high GS activity) at high light intensity, leaf temperature and [O(2)]. CONCLUSIONS: Our results demonstrate that photorespiration is strongly involved in NH(3) emission by rice leaves and suggest that differences in AER between cultivars result from their different GS activities, which would result in different capacities for reassimilation of photorespiratory NH(3). The results also suggest that NH(3) emission in rice leaves is not directly controlled by transpiration and stomatal conductance.