Elevated CO2 concentration increases maize growth under water deficit or soil salinity but with a higher risk of hydraulic failure.

Climate change presents a challenge for plants to acclimate their water relations under changing environmental conditions, and may increase the risks of hydraulic failure under stress. In this study, maize plants were acclimated to two different CO2 concentrations ([CO2]; 400 ppm and 700 ppm) while under either water stress (WS) or soil salinity (SS) treatments, and their growth and hydraulic traits were examined in detail. Both WS and SS inhibited growth and had significant impacts on hydraulic traits. In particular, the water potential at 50% loss of stem hydraulic conductance (P50) decreased by 1 MPa in both treatments at 400 ppm. When subjected to elevated [CO2], the plants under both WS and SS showed improved growth by 7-23%. Elevated [CO2] also significantly increased xylem vulnerability (measured as loss of conductivity with decreasing xylem pressure), resulting in smaller hydraulic safety margins. According to the plant desiccation model, the critical desiccation degree (time×vapor pressure deficit) that the plants could tolerate under drought was reduced by 43-64% under elevated [CO2]. In addition, sensitivity analysis showed that P50 was the most important trait in determining the critical desiccation degree. Thus, our results demonstrated that whilst elevated [CO2] benefited plant growth under WS or SS, it also interfered with hydraulic acclimation, thereby potentially placing the plants at a higher risk of hydraulic failure and increased mortality.

A meta-analysis highlights the cross-resistance of plants to drought and salt stresses from physiological, biochemical, and growth levels.

In nature, drought and salt stresses often occur simultaneously and affect plant growth at multiple levels. However, the mechanisms underlying plant responses to drought and salt stresses and their interactions are still not fully understood. We performed a meta-analysis to compare the effects of drought, salt, and combined stresses on plant physiological, biochemical, morphological and growth traits, analyze the different responses of C3 and C4 plants, as well as halophytes and non-halophytes, and identify the interactive effects on plants. There were numerous similarities in plant responses to drought, salt, and combined stresses. C4 plants had a more effective antioxidant defense system, and could better maintain above-ground growth. Halophytes could better maintain photosynthetic rate (Pn) and relative water content (RWC), and reduce growth as an adaptation strategy. The responses of most traits (Pn, RWC, chlorophyll content, soluble sugar content, H2O2 content, plant dry weight, etc.) to combined stress were less-than-additive, indicating cross-resistance rather than cross-sensitivity of plants to drought and salt stresses. These results are important to improve our understanding of drought and salt cross-resistance mechanisms and further induce resistance or screen-resistant varieties under stress combination.

Response of soil respiration and carbon budget to irrigation quantity/quality and biochar addition in a mulched maize field under drip irrigation.

BACKGROUND: Biochar addition strongly alters net carbon (C) balance in agroecosystems. However, the effects of biochar addition on net C balance of maize field under various irrigation water quantities and qualities remains unclear. Thus, a field experiment combining two irrigation levels of full (W1) and deficit irrigation (W2 = 1/2 W1), two water salinity levels of fresh (S0, 0.71 g L-1 ) and brackish water (S1, 4 g L-1 ), and two biochar addition rates of 0 t ha-1 (B0) and 60 t ha-1 (B1) was conducted to investigate soil carbon dioxide (CO2 ) emissions, maize C sequestration and C budget. RESULTS: Compared with W1, W2 reduced average cumulative CO2 emissions by 6.5% and 19.9% for 2020 and 2021, respectively. The average cumulative CO2 emissions under W1S1 treatments were 5.4% and 22.3% lower than W1S0 for 2020 and 2021, respectively, whereas W2S0 and W2S1 had similar cumulative CO2 emissions in both years. Biochar addition significantly increased cumulative CO2 emissions by 17.8-23.5% for all water and salt treatments in 2020, and reduced average cumulative CO2 emissions by 11.9% for W1 but enhanced it by 8.0% for W2 in 2021. Except for W2S1, biochar addition effectively increased total maize C sequestration by 6.9-14.8% for the other three treatments through ameliorating water and salt stress over the 2 years. Compared with W1S0, W1S1 did not affect net C sequestration, but W2 treatments significantly decreased it. Biochar addition increased net C sequestration by 39.47-43.65 t C ha-1 for four water and salt treatments for the 2 years. CONCLUSION: These findings demonstrate that biochar addition is an effective strategy to increase both crop C sequestration and soil C storage under suitable water-saving irrigation methods in arid regions with limited freshwater resources. © 2023 Society of Chemical Industry.

Combination of suitable planting density and nitrogen rate for high yield maize and their source-sink relationship in Northwest China.

BACKGROUND: Increasing crop yield per unit area by increasing planting density is essential to ensure food security. However, the optimal combination of planting density and nitrogen (N) application for high-yielding maize and its source-sink characteristics need to be more clearly understood. RESULTS: A 2-year field experiment was conducted combining three planting densities (D1: 70 000 plants ha-1 ; D2: 100 000 plants ha-1 ; D3: 130 000 plants ha-1 ) and three nitrogen rates (N1: 150 kg hm-2 ; N2: 350 kg hm-2 ; N3: 450 kg hm-2 ). The results showed that increasing planting density significantly increased leaf area index and grain yield but negatively affected ear traits. The Richards model was used to fit the dynamic changes of dry matter accumulation of maize under different treatments, and the fitting results were good. Increasing planting density increased population yield while limiting the development of individual plants, bringing the period of rapid dry matter accumulation to an early end and accelerating leaf senescence. An appropriate nitrogen rate could prolong the period of rapid accumulation of dry matter in maize, and increase the 100-kernel weight. Increasing planting density enhanced post-silking dry matter accumulation to a lesser extent, and the source-sink relationship of the maize population gradually developed from sink limitation to source limitation with increasing planting density. CONCLUSION: The decrease in yield due to the insufficient source strength to meet the sink demand at too high densities was the reason that limited further improvement of the optimal planting density. An appropriate nitrogen rate facilitated the realization of yield potential at high density. © 2023 Society of Chemical Industry.

Plasticity in stomatal behaviour across a gradient of water supply is consistent among field-grown maize inbred lines with varying stomatal patterning.

Stomata regulate leaf CO2 assimilation (A) and water loss. The Ball-Berry and Medlyn models predict stomatal conductance (gs ) with a slope parameter (m or g1 ) that reflects the sensitivity of gs to A, atmospheric CO2  and humidity, and is inversely related to water use efficiency (WUE). This study addressed knowledge gaps about what the values of m and g1 are in C4 crops under field conditions, as well as how they vary among genotypes and with drought stress. Four inbred maize genotypes were unexpectedly consistent in how m and g1 decreased as water supply decreased. This was despite genotypic variation in stomatal patterning, A and gs . m and g1 were strongly correlated with soil water content, moderately correlated with predawn leaf water potential (Ψpd ), but not correlated with midday leaf water potential (Ψmd ). This implied that m and g1 respond to long-term water supply more than short-term drought stress. The conserved nature of m and g1 across anatomically diverse genotypes and water supplies suggests there is flexibility in structure-function relationships underpinning WUE. This evidence can guide the simulation of maize gs across a range of water supply in the primary maize growing region and inform efforts to improve WUE.

Stomatal conductance of tomato leaves is regulated by both abscisic acid and leaf water potential under combined water and salt stress.

Stomatal conductance (gs ) affects water use efficiency (WUE) through coordinating photosynthesis and transpiration and is regulated by chemical and/or hydraulic signals. However, the regulation mechanism of gs of tomato leaves has not been fully explored under combined water and salt stress. Here, we set up four salt treatments and two water treatments in a climate greenhouse and measured stomatal morphologies and conductance and other photosynthesis parameters. Water and salt stress reduced stomatal length (SL), width, perimeter, area (amax ), density (SD), and the maximum stomatal conductance (gsmax ). Water and salt stress had a separate weakening effect on net photosynthetic rate (A) and transpiration rate but interactively reduced gs . The contents of abscisic acid (ABA) and Na+ in tomato leaves increased with the NaCl concentration, while leaf water potential (Ψl ) and chlorophyll content decreased. Under full irrigation, gsmax was coordinated by SD and amax , and gs by ABA content under salt stress. Under water and salt combined stress, gsmax was affected by amax , and gs was coordinated with ABA and Ψl . The decrease of A was caused by both a reduction of chlorophyll content and gs under water and salt stress. Intrinsic WUE did not reduce under full irrigation or mild to moderate salt stress but decreased under a combination of water and severe salt stress, indicating that the leaves of the tested tomato cultivar performed better under moderate salt stress. Collectively, these results can provide useful insights for the efficient management of water and salt to adapt to drought and high salt environments.

Elevated [CO2 ] alleviates the impacts of water deficit on xylem anatomy and hydraulic properties of maize stems.

Plants can modify xylem anatomy and hydraulic properties to adjust to water status. Elevated [CO2 ] can increase plant water potential via reduced stomatal conductance and water loss. This raises the question of whether elevated [CO2 ], which thus improves plant water status, will reduce the impacts of soil water deficit on xylem anatomy and hydraulic properties of plants. To analyse the impacts of water and [CO2 ] on maize stem xylem anatomy and hydraulic properties, we exposed potted maize plants to varying [CO2 ] levels (400, 700, 900, and 1,200 ppm) and water levels (full irrigation and deficit irrigation). Results showed that at current [CO2 ], vessel diameter, vessel roundness, stem cross-section area, specific hydraulic conductivity, and vulnerability to embolism decreased under deficit irrigation; yet, these impacts of deficit irrigation were reduced at elevated [CO2 ]. Across all treatments, midday stem water potential was tightly correlated with xylem traits and displayed similar responses. A distinct trade-off between efficiency and safety in stem xylem water transportation in response to water deficit was observed at current [CO2 ] but not observed at elevated [CO2 ]. The results of this study enhance our knowledge of plant hydraulic acclimation under future climate environments and provide insights into trade-offs in xylem structure and function.

Salinity-specific stomatal conductance model parameters are reduced by stomatal saturation conductance and area via leaf nitrogen.

Modeling stomatal behavior is necessary for accurate stomatal simulation and predicting the terrestrial water­carbon cycle. Although the Ball-Berry and Medlyn stomatal conductance (gs) models have been widely used, variations and the drivers of their key slope parameters (m and g1) remain poorly understood under salinity stress. We measured leaf gas exchange, physiological and biochemical traits, soil water content and electrical conductivity of saturation extract (ECe), and fitted slope parameters of two genotypes of maize growing in two water and two salinity levels. We found m was different between the genotypes, but no difference in g1. Salinity stress reduced m and g1, saturated stomatal conductance (gsat), the fraction of leaf epidermis area allocation to stomata (fs), and leaf nitrogen (N) content, and increased ECe, but no marked decrease in slope parameters under drought. Both m and g1 were positively correlated with gsat, fs, and leaf N content, and negatively correlated with ECe in the same fashion among the two genotypes. Salinity stress altered m and g1 by modulating gsat and fs via leaf N content. The prediction accuracy of gs was improved using salinity-specific slope parameters, with root mean square error (RMSE) being decreased from 0.056 to 0.046 and 0.066 to 0.025 mol m-2 s-1 for the Ball-Berry and Medlyn models, respectively. This study provides a modeling approach to improving the simulation of stomatal conductance under salinity.

The trade-offs between resistance and resilience of forage stay robust with varied growth potentials under different soil water and salt stress.

Water shortage and soil salinization are important factors restricting crop production worldwide. To conduct accurate yield prediction and reasonable crop layout, more attention should be paid to the performances of crop resistance and resilience under water and salt stress and their trade-off relationships. Here, we set different water (full irrigation, W0; moderate deficit irrigation, W1; and severe deficit irrigation, W2) and salt (S0, S1, S2, S3, S4, S5, and S6, representing 0 ‰, 1 ‰, 2 ‰, 3 ‰, 4 ‰, 5 ‰, and 6 ‰ salt in soil) treatments. Together with relevant studies, we analyzed the performances of forage resistance (Rt) and resilience (Rs) and their relationships under varied water and salt stress. The results indicated that logarithmic Rt (lg(Rt), the same as lg(Rs)) and the distribution of lg(Rs) were affected by water and salt stress, however, the relationships of lg(Rs)-lg(Rt) stayed stable with the constant slopes (k) and declined intercepts (m) as stress intensified. The physiological mechanisms and trade-offs for fixed species remained robust while the growth potentials varied under stress, which were closely related to stomatal regulations. Forage with larger |k| was suitable for fully irrigated regions to achieve higher yields, while regions with detrimental water and salt conditions should select cultivars with smaller |k| to ensure production. This study laid the groundwork for the estimation of the perennial forage adaptation and stability, and the method of long-term yield prediction and cultivar management under soil water and salt stress.

Mild water and salt stress improve water use efficiency by decreasing stomatal conductance via osmotic adjustment in field maize.

Water and salt stress often occur simultaneously in heavily irrigated arid agricultural areas, yet they are usually studied in isolation. To understand the physiological bases of water use efficiency (WUE) of field-grown maize (Zea mays) at multi-scales under combined water and salt stress, we investigated the joint effects of water and salt stress on physiology, growth, yield, and WUE of two genotypes (XY335 and ZD958). We measured leaf stomatal conductance (gs), net photosynthesis rate (A) and hydraulic traits, whole-plant growth and water use (ET), and final biomass and grain yield. Leaf osmotic adjustment was a key trait of the physiological differences between XY335 and ZD958 under water and salt stress. Although the responses of the two genotypes were different, mild water and salt stress improved intrinsic water use efficiency (iWUE = A/gs) by (i) decreasing gsvia increasing osmotic adjustment and hydraulic resistance, and (ii) declining A via increasing stomatal limitations rather than reducing photosynthetic capacity. Joint water and salt stress had a synergistic effect on reproductive growth and grain formation of maize. Mild water and salt stress reduced ET, stabilized grain yield, and improved grain WUE via declining gs, maintaining photosynthetic capacity, and improving harvest index. Collectively, our study provides a novel insight into the physiological mechanisms of WUE and demonstrates an approach for the efficient management of water and salt by using a growth stage-based deficit irrigation strategy or/and selecting genotypes with strong osmotic adjustment capacity and high harvest index.

Irrigation water productivity is more influenced by agronomic practice factors than by climatic factors in Hexi Corridor, Northwest China.

Quantifying the influence of driving factors on irrigation water productivity (IWP) is vital for efficient agricultural water use. This study analyzed contributions of agronomic practice and climatic factors to the changes of IWP, based on the data from 1981 to 2012 in Hexi Corridor, Northwest China. Cobb-Douglas production functions were developed by the partial least squares method and contribution rates of the driving factors were calculated. Results showed that IWP and its driving factors increased during the study period, with different changing patterns. IWP was significantly correlated with the agronomic practice factors, daily mean temperature and solar radiation of the crop growing period. The agronomic practice factors including irrigation, fertilization, agricultural film, and agricultural pesticide contributed 20.6%, 32.8%, 42.3% and 11.1% respectively to the increase of IWP; and the contribution rates of the climatic factors, i.e. daily mean temperature and solar radiation, are -0.9% and 0.9%. And the contributions of these factors changed in different sub-periods. It is concluded that agronomic practice factors influenced IWP much more than climatic factors. The improvement of IWP should rely on advanced water-saving technology and application of optimum (need-based) fertilizer, agricultural film and pesticide, ensuring efficient use of agronomic inputs in the study area.

Scaling up stomatal conductance from leaf to canopy using a dual-leaf model for estimating crop evapotranspiration.

The dual-source Shuttleworth-Wallace model has been widely used to estimate and partition crop evapotranspiration (λET). Canopy stomatal conductance (Gsc), an essential parameter of the model, is often calculated by scaling up leaf stomatal conductance, considering the canopy as one single leaf in a so-called "big-leaf" model. However, Gsc can be overestimated or underestimated depending on leaf area index level in the big-leaf model, due to a non-linear stomatal response to light. A dual-leaf model, scaling up Gsc from leaf to canopy, was developed in this study. The non-linear stomata-light relationship was incorporated by dividing the canopy into sunlit and shaded fractions and calculating each fraction separately according to absorbed irradiances. The model includes: (1) the absorbed irradiance, determined by separately integrating the sunlit and shaded leaves with consideration of both beam and diffuse radiation; (2) leaf area for the sunlit and shaded fractions; and (3) a leaf conductance model that accounts for the response of stomata to PAR, vapor pressure deficit and available soil water. In contrast to the significant errors of Gsc in the big-leaf model, the predicted Gsc using the dual-leaf model had a high degree of data-model agreement; the slope of the linear regression between daytime predictions and measurements was 1.01 (R2 = 0.98), with RMSE of 0.6120 mm s-1 for four clear-sky days in different growth stages. The estimates of half-hourly λET using the dual-source dual-leaf model (DSDL) agreed well with measurements and the error was within 5% during two growing seasons of maize with differing hydrometeorological and management strategies. Moreover, the estimates of soil evaporation using the DSDL model closely matched actual measurements. Our results indicate that the DSDL model can produce more accurate estimation of Gsc and λET, compared to the big-leaf model, and thus is an effective alternative approach for estimating and partitioning λET.