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Chromium (Cr)-contaminated in irrigation water poses a significant threat to the safety of wheat (Triticum aestivum L.) production safety. Recent studies suggest that melatonin (MT) could enhance crop tolerance to Cr pollution. This study aimed to investigate the effects of foliar spraying MT on alleviating Cr toxicity and accumulation in wheat irrigated with K2Cr2O7 solution at concentrations of 5, 10, and 20 mg/kg Cr in the soil. Our results showed that Cr-contaminated water irrigation significantly reduced dry weight, grain numbers, grain weight, yield, harvest index, net photosynthetic rate (Pn), maximum and actual photochemical efficiency of photosystem II (Fv/Fm and ΦPSII), chlorophyll contents, and the a/b ratio. It also increased PSII photodamage and oxidative stress in wheat leaves, resulting in high Cr accumulation in roots, leaves, and grains. Foliar spraying of MT alleviated Cr toxicity by improving Pn, Fv/Fm, and ΦPSII, enhancing chlorophyll content, promoting dry matter accumulation and yield, and reducing oxidative stress and Cr translocation. Furthermore, MT application enhanced transcriptional regulation, alleviated oxidative stress by boosting antioxidant enzyme activities, and restricted Cr translocation from roots to leaves and grains by increasing the accumulation of secondary metabolites, such as lignin and metallothionein. These findings suggest that MT application could serve as a viable strategy for reducing Cr contamination in cereals and supporting phytoremediation efforts.

Abstract Wheat grain starch content displays large variations within different pearling fractions, which affecting the processing quality of corresponding flour, while the underlying mechanism on starch gradient formation is unclear. Here, we show that wheat caryopses acquire sugar through the transfer of cells (TCs), inner endosperm (IE), outer endosperm (OE), and finally aleurone (AL) via micro positron emission tomography-computed tomography (PET-CT). To obtain integrated information on spatial transcript distributions, developing caryopses are laser microdissected into AL, OE, IE, and TC. Most genes encoding carbohydrate transporters are upregulated or specifically expressed, and sugar metabolites are more highly enriched in the TC group than in the AL group, in line with the PET-CT results. Genes encoding enzymes in sucrose metabolism, such as sucrose synthase, beta-fructofuranosidase, glucose-1-phosphate adenylyltransferase show significantly lower expression in AL than in OE and IE, indicating that substrate supply is crucial for the formation of starch gradients. Furthermore, the low expressions of gene encoding starch synthase contribute to low starch content in AL. Our results imply that transcriptional regulation represents an important means of impacting starch distribution in wheat grains and suggests breeding targets for enhancing specially pearled wheat with higher quality.

The use of slow-release fertilizers and seed-fertilizers cause localized high-ammonium (NH4+) environments in agricultural fields, adversely affecting wheat growth and development and delaying its yield. Thus, it is important to investigate the physiological responses of wheat and its tolerance to NH4+ stress to improve the adaptation of wheat to high NH4+ environments. In this study, the physiological mechanisms of ammonium tolerance in wheat (Triticum aestivum) were investigated in depth by comparative analysis of two cultivars: NH4+-tolerant Xumai25 and NH4+-sensitive Yangmai20. Cultivation under hydroponic conditions with high NH4+ (5 mM NH4+, AN) and nitrate (5 mM NO3-, NN), as control, provided insights into the nuanced responses of both cultivars. Compared to Yangmai20, Xumai25 displayed a comparatively lesser sensitivity to NH4+ stress, as evident by a less pronounced reduction in dry plant biomass and a milder adverse impact on root morphology. Despite similarities in NH4+ efflux and the expression levels of TaAMT1.1 and TaAMT1.2 between the two cultivars, Xumai25 exhibited higher NH4+ influx, while maintaining a lower free NH4+ concentration in the roots. Furthermore, Xumai25 showed a more pronounced increase in the levels of free amino acids, including asparagine, glutamine, and aspartate, suggesting a superior NH4+ assimilation capacity under NH4+ stress compared to Yangmai20. Additionally, the enhanced transcriptional regulation of vacuolar glucose transporter and glucose metabolism under NH4+ stress in Xumai25 contributed to an enhanced carbon skeleton supply, particularly of 2-oxoglutarate and pyruvate. Taken together, our results demonstrate that the NH4+ tolerance of Xumai25 is intricately linked to enhanced glucose metabolism and optimized glucose transport, which contributes to the robust NH4+ assimilation capacity.

The impact of ammonium (NH4+) stress on plant growth varies across species and cultivars, necessitating an in-depth exploration of the underlying response mechanisms. This study delves into elucidating the photosynthetic responses and differences in tolerance to NH4+ stress by investigating the effects on two wheat (Triticum aestivum L.) cultivars, Xumai25 (NH4+-less sensitive) and Yangmai20 (NH4+-sensitive). The cultivars were grown under hydroponic conditions with either sole ammonium nitrogen (NH4+, AN) or nitrate nitrogen (NO3−, NN) as the nitrogen source. NH4+ stress exerted a profound inhibitory effect on seedling growth and photosynthesis in wheat. However, these effects were less pronounced in Xumai25 than in Yangmai20. Dynamic photosynthetic analysis revealed that the suppression in photosynthesis was primarily attributed to stomatal limitation associated with a decrease in leaf water status and osmotic potential. Compared to Yangmai20, Xumai25 exhibited a significantly higher leaf K+ concentration and TaAKT1 upregulation, leading to a stronger stomatal opening and, consequently, a better photosynthetic performance under NH4+ stress. In conclusion, our study suggested stomatal limitation as the primary factor restricting photosynthesis under NH4+ stress. Furthermore, we demonstrated that improved regulation of osmotic substances contributed to higher stomatal conductance and enhanced photosynthetic performance in Xumai25.

Shoot branching is inhibited by a low red/far-red ratio (R/FR). Prior studies have shown that the R/FR suppressed Arabidopsis thaliana branching by promotes bud abscisic acid (ABA) accumulation directly. Given that wheat tiller buds are wrapped in leaf sheaths and may not respond rapidly to a R/FR, systemic cytokinin (CTK) may be more critical. Here, systemic hormonal signals including indole-3-acetic acid (IAA), gibberellins (GA) and CTK and bud ABA signals in wheat were tested under a low R/FR. The results showed that a low R/FR reduced the percentage of tiller occurrence of tiller IV and the tiller number per plant. The low R/FR did not rapidly induced ABA accumulation in the tiller IV because of the protection of the leaf sheath and had little effect on IAA content and signaling in the tiller nodes. The significant change in the CTK levels was observed earlier than those of other hormone (ABA, IAA and GA) and exogenous cytokinin restored the CTK levels and tiller number per plant under low R/FR conditions. Further analysis revealed that the decrease in cytokinin levels was mainly associated with upregulation of cytokinin degradation genes (TaCKX5, TaCKX11) in tiller nodes. In addition, exposure to a decreased R/FR upregulated the expression of GA biosynthesis genes (TaGA20ox1, TaGA3ox2), resulting in elevated GA levels, which might further promote CTK degradation in tiller nodes and inhibit tillering. Therefore, our results provide evidence that the enhancement of cytokinin degradation is a novel mechanism underlying the wheat tillering response to a low R/FR.

Water and nitrogen (N) deficiencies are the major limitations to crop production, particularly when they occur simultaneously. By supporting metabolism, even when tissue water capacity is lower, nitrogen and priming may reduce drought pressure on plants. Therefore, the current study investigates the impact of nitrogen and priming on wheat to minimize post-anthesis drought stress. Plant morphology, physiology, and biochemical changes were observed before, during, and after stress at the post-anthesis stage. The plants were exposed to three water levels, i.e., well watering (WW), water deficit (WD), and priming at jointing and water deficit (PJWD) at the post-anthesis stage, and two different nitrogen levels, i.e., N180 (N1) and N300 (N2). Nitrogen was applied in three splits, namely, sowing, jointing, and booting stages. The results showed that the photosynthesis of plants with N1 was significantly reduced under drought stress. Moreover, drought stress affected chlorophyll (Chl) fluorescence and water-related parameters (osmotic potential, leaf water potential, and relative water content), grain filling duration (GFD), and grain yield. In contrast, PJWD couple with high nitrogen treatment (N300 kg ha–1) induced the antioxidant activity of peroxidase (37.5%), superoxide dismutase (29.64%), and catalase (65.66%) in flag leaves, whereas the levels of hydrogen peroxide (H2O2) and superoxide anion radical (O2–) declined by 58.56 and 66.64%, respectively. However, during the drought period, the primed plants under high nitrogen treatment (N300 kg ha–1) maintained higher Chl content, leaf water potential, and lowered lipid peroxidation (61%) (related to higher activities of ascorbate peroxidase and superoxide dismutase). Plants under high nitrogen treatment (N300 kg ha–1) showed deferred senescence, improved GFD, and grain yield. Consequently, the research showed that high nitrogen dose (N300 kg ha–1) played a synergistic role in enhancing the drought tolerance effects of priming under post-anthesis drought stress in wheat.

Drought stress is a limiting factor for wheat production and food security. Drought priming has been shown to increase drought tolerance in wheat. However, the underlying mechanisms are unknown. In the present study, the genes encoding the biosynthesis and metabolism of abscisic acid (ABA) and jasmonic acid (JA), as well as genes involved in the ABA and JA signaling pathways were up-regulated by drought priming. Endogenous concentrations of JA and ABA increased following drought priming. The interplay between JA and ABA in plant responses to drought priming was further investigated using inhibitors of ABA and JA biosynthesis. Application of fluridone (FLU) or nordihydroguaiaretic acid (NDGA) to primed plants resulted in lower chlorophyll-fluorescence parameters and activities of superoxide dismutase and glutathione reductase, and higher cell membrane damage, compared to primed plants (PD) under drought stress. NDGA + ABA, but not FLU + JA, restored priming-induced tolerance, as indicated by a finding of no significant difference from PD under drought stress. Under drought priming, NDGA induced the suppression of ABA accumulation, while FLU did not affect JA accumulation. These results were consistent with the expression of genes involved in the biosynthesis of ABA and JA. They suggest that ABA and JA are required for priming-induced drought tolerance in wheat, with JA acting upstream of ABA.

Starch plays an important role in food industry. In this study, three wheat cultivars with different protein contents were used to investigate the different ratios of starch addition on starch pasting properties, starch thermal performance, dough rheology, biscuit quality, and their relationships. Results showed that with the increase in starch content, gluten, protein and glutenin macropolymer (GMP), lactic acid solvent retention capacity (SRC), sucrose SRC, and onset temperature (To) decreased, while most pasting parameters and gelatinization enthalpy (ΔH) increased. Viscosity parameters were significantly negatively correlated with dough stability time, farinograph quality number (FQN), and sucrose SRC. Biscuit quality was improved by starch addition, indicated by lower thickness and hardness, higher diameter, spread ratio, and sensory score. Viscosity parameters were positively correlated to diameter, spread ratio, and sensory score of biscuit, while negatively correlated to hardness and thickness of biscuit. Image analysis showed that the crumbs of biscuit were improved as shown by bigger pores in the bottom side. The results provide useful information for the clarification of the role of starch in determining biscuit quality and the inter-relationships of flour, dough, and biscuit.

Nitrogen, one of the essential elements, is a key determinant for improving peanut growth and yield. GATA zinc finger transcription factors have been found to be involved in regulation of nitrogen metabolism. However, a systematic characterization of the GATA gene family and patterns of their expression under different nitrogen levels remains elusive. In this study, a total of 45 GATA genes distributed among 17 chromosomes were identified in the peanut genome and classified into three subfamilies I, II and III with 26, 13 and 6 members, respectively, whose physicochemical characteristics, gene structures and conserved motifs were also analyzed. Furthermore, the optimal level of nitrogen fertilizer on the growth of peanut cultivar Yuhua 23 was determined by pod yield and value cost ratio from 2020 to 2022, and the results revealed that 150 kg hm−2 nitrogen was the best for cultivation of peanut Yuhua 23 because of its highest pod yield and relatively higher VCR of more than four. In addition, expression patterns of peanut GATA genes under different nitrogen levels were detected by real-time quantitative PCR and several GATA genes were significantly changed under a nitrogen level of 150 kg hm−2. Overall, the above results would be helpful for further understanding biological functions of the GATA gene family in cultivated peanut.

The asymmetric warming in diurnal and seasonal temperature patterns plays an important role in crop distribution and productivity. Asymmetric warming during the early growth periods of winter wheat profoundly affects its vegetative growth and post-anthesis grain productivity. Field experiments were conducted on winter wheat to explore the impact of night warming treatment in winter (Winter warming treatment, WT) or spring (Spring warming treatment, ST) on the senescence of flag leaves and yield of wheat plants later treated with night warming during grain filling (Warming treatment during grain filling, FT). The results showed that FT decreased wheat yield by reducing the number of grains per panicle and per 1,000-grain weight and that the yield of wheat plants treated with FT declined to a greater extent than that of wheat plants treated with WT + FT or ST + FT. The net photosynthetic rate, chlorophyll content, and chlorophyll fluorescence parameters of the flag leaves of wheat plants treated with WT + FT or ST + FT were higher than those under the control treatment from 0 to 7 days after anthesis (DAA) but were lower than those under the control treatment and higher than those of wheat plants treated with FT alone from 14 to 28 DAA. The soluble protein and Rubisco contents in the flag leaves of wheat plants treated with WT + FT or ST + FT were high in the early grain-filling period and then gradually decreased to below those of the control treatment. These contents were greater in wheat plants treated with WT + FT than in wheat plants treated with ST + FT from 0 to 14 DAA, whereas the opposite was true from 21 to 28 DAA. Furthermore, WT + FT and ST + FT inhibited membrane lipid peroxidation by increasing superoxide dismutase and peroxidase activities and lowering phospholipase D (PLD), phosphatidic acid (PA), lipoxygenase (LOX), and free fatty acid levels in the early grain-filling period, but their inhibitory effects on membrane lipid peroxidation gradually weakened during the late grain-filling period. Night-warming priming alleviated the adverse effect of post-anthesis warming on yield by delaying the post-anthesis senescence of flag leaves.

The aim of this study was to test the significant effects of inorganic sulfur and cysteine on grain protein and flour quality in wheat and to provide a theoretical basis of wheat cultivation techniques with high yield and quality. In the field experiment, a winter wheat cultivar, Yangmai 16, was used, and five treatments were established, i.e., S0 (no sulfur fertilizer application during the whole wheat growth period), S(B)60 (60 kg ha−1 inorganic sulfur fertilizer was applied as the basal fertilizer), Cys(B)60 (60 kg ha−1 cysteine sulfur fertilizer was applied as the basal fertilizer), S(J)60 (60 kg ha−1 inorganic sulfur fertilizer was applied as the jointing fertilizer), and Cys(J)60 (60 kg ha−1 cysteine sulfur fertilizer was applied as the jointing fertilizer). The fertilizer application at jointing stage showed a better influence than basal fertilizer application on protein quality; for the content of albumin, gliadin, and high molecular weight glutenin (HMW-GS), Cys(J)60 was the best among these treatments. An increase of 7.9%, 24.4%, 43.5%, 22.7% and 36.4% was found in grain yield, glutenin content, glutenin macro-polymer (GMP), low molecular weight glutenin (LMW-GS), and S content under Cys(J)60, in relation to the control, respectively. A similar trend was found in the end-use quality, as exemplified by an increase of 38.6%, 10.9%, 60.5%, and 109.8% in wet gluten content, dry gluten content, sedimentation volume, and bread-specific volume, respectively; a decrease of 69.3% and 69.1% in bread hardness and bread chewiness was found under Cys(J)60. In terms of application period, topdressing at jointing stage is compared with base fertilizer, the sulfur fertilizer application at jointing stage showed larger effects on grain protein and flour quality, from the different types of sulfur fertilizer, the application of cysteine performed better than the use of inorganic sulfur. The Cys(J)60 exhibited the best effects on protein and flour quality. It was suggested that sufficient sulfur application at jointing stage has the potential to enhance the grain protein and flour quality.

Chromium (Cr) pollution has serious harm to crop growth, while little is known on the role of melatonin (MT) on seed germination and physiology in Cr-stressed wheat. The effects of seed soaking with MT on growth, reserve mobilization, osmotic regulation and antioxidant capacity of wheat seeds during germination under hexavalent chromium (100 μM) stress were investigated. The results indicated that Cr toxicity decreased the seed germination rate by 16% and suppressed the growth of germinated seeds compared to unstressed seeds. MT in the concentration-dependent manner increased germination rate and promoted subsequent growth when seeds were exposed to Cr stress, but the effect could be counteracted at high concentration. Seed soaking with MT (100 μM) markedly decreased Cr accumulation in seeds, radicals and coleoptiles by 15%, 6% and 15%, respectively, and enhanced α-amylase activity and soluble sugar and free amino acids content in seeds to improve reserve mobilization under Cr stress, compared with Cr treatment. Furthermore, decreasing the level of osmotic regulators (soluble sugar and soluble protein) in radicles under MT combined with Cr treatment confirmed the reduction of osmotic stress caused by Cr stress. Importantly, MT pretreatment reduced H2O2 content by 19% and O2·− release rate by 45% in radicles under Cr toxicity compared with Cr-stressed wheat, in terms of promoting scavenging ability and decreasing production ability, which was to upregulate the activities and encoding genes expression levels of superoxide dismutase (SOD), catalase (CAT), ascorbic acid peroxidase (APX) and peroxidase (POD) and to downregulate plasma membrane-bound NADPH oxidase (NOX) encoding genes (TaRbohD, TaRbohF) expression, respectively. In all, these results provided evidence that seed soaking with MT could be a potentially method to protect wheat seeds from Cr toxicity, which effectively ameliorated germination under Cr stress by enhancing reserve mobilization and antioxidant metabolism in wheat.

Inhibited photosynthesis caused by post-anthesis high-temperature stress (HTS) leads to decreased wheat grain yield. Magnesium (Mg) plays critical roles in photosynthesis; however, its function under HTS during wheat grain filling remains poorly understood. Therefore, in this study, we investigated the effects of Mg on the impact of HTS on photosynthesis during wheat grain filling by conducting pot experiments in controlled-climate chambers. Plants were subjected to a day/night temperature cycle of 32°C/22°C for 5 days during post-anthesis; the control temperature was set at 26°C/16°C. Mg was applied at the booting stage, with untreated plants used as a control. HTS reduced the yield and net photosynthetic rate (Pn) of wheat plants. The maximum carboxylation rate (VCmax), which is limited by Rubisco activity, decreased earlier than the light-saturated potential electron transport rate. This decrease in VCmax was caused by decreased Rubisco activation state under HTS. Mg application reduced yield loss by stabilizing Pn. Rubisco activation was enhanced by increasing Rubisco activase activity following Mg application, thereby stabilizing Pn. We conclude that Mg maintains Rubisco activation, thereby helping to stabilize Pn under HTS.

With global climate change, waterlogging stress is becoming more frequent. Waterlogging stress inhibits root growth and physiological metabolism, which ultimately leads to yield loss in wheat. Waterlogging priming has been proven to effectively enhance waterlogging tolerance in wheat. However, it is not known whether waterlogging priming can improve the offspring’s waterlogging resistance. Here, wheat seeds that applied waterlogging priming for one generation, two generations and three generations are separately used to test the hypoxia stress tolerance in wheat, and the physiological mechanisms are evaluated. Results found that progeny of primed plants showed higher plant biomass by enhancing the net photosynthetic rate and antioxidant enzyme activity. Consequently, more sugars are transported to roots, providing a metabolic substrate for anaerobic respiration and producing more ATP to maintain the root growth in the progeny of primed plants compared with non-primed plants. Furthermore, primed plants’ offspring promote ethylene biosynthesis and further induce the formation of a higher rate of aerenchyma in roots. This study provides a theoretical basis for improving the waterlogging tolerance of wheat.

The intensive use of groundwater in agriculture under the current climate conditions leads to acceleration of soil salinization. Given that almond is a salt-sensitive crop, selection of salt-tolerant rootstocks can help maintain productivity under salinity stress. Selection for tolerant rootstocks at an early growth stage can reduce the investment of time and resources. However, salinity-sensitive markers and salinity tolerance mechanisms of almond species to assist this selection process are largely unknown. We established a microscopy-based approach to investigate mechanisms of stress tolerance in and identified cellular, root anatomical, and molecular traits associated with rootstocks exhibiting salt tolerance. We characterized three almond rootstocks: Empyrean-1 (E1), Controller-5 (C5), and Krymsk-86 (K86). Based on cellular and molecular evidence, our results show that E1 has a higher capacity for salt exclusion by a combination of upregulating ion transporter expression and enhanced deposition of suberin and lignin in the root apoplastic barriers, exodermis, and endodermis, in response to salt stress. Expression analyses revealed differential regulation of cation transporters, stress signaling, and biopolymer synthesis genes in the different rootstocks. This foundational study reveals the mechanisms of salinity tolerance in almond rootstocks from cellular and structural perspectives across a root developmental gradient and provides insights for future screens targeting stress response.

Nitrogen (N) supply and seed rate (SR) are two essential factors that affect the accumulation and partitioning of N and dry matter (DM) and, therefore, grain yield (GY) and N use efficiency (NUE). The objective of this experiment was to optimize N application and SR to regulate wheat growth and increase both GY and NUE. The results revealed that net photosynthetic rate (Pn), stomatal conductance (Gs), chlorophyll content, and activities of metabolic enzymes (NR and GS) significantly increased with increasing of N levels while decreasing SR. Plant tillers, GY, DM before anthesis, and N translocation, N agronomic efficiency (NAE), N recovery efficiency (NRE), and N uptake efficiency (NUPE) were highest in a combined treatment of N235 and SR180. However, N levels beyond 235 kg ha−1 significantly decreased NAE, NRE, and NUPE. By increasing SR from 135 to 180 kg ha−1 an increase of 12.9 % and 9.1% GY and NUPE, respectively, was observed. Based on this result, we estimate that 1 kg N ha−1 might be replaced by an increase of approximately 0.6 kg ha−1 SR. Our study suggested that using a combination of N and SR (N235 + SR180) could attain maximum GY and improve NUE parameters.

Abstract Background Nitrogen is one basic element of amino acids and grain protein in wheat. In field experiments, wheat plants were subjected to different timing of nitrogen topdressing treatments: at the stages of emergence of the top fifth leaf (TL5), top third leaf (TL3) and top first leaf (TL1) to test the regulatory effects of nitrogen topdressing timing on grain protein quality. The underlying mechanisms were elucidated by clarifying the relationship between proteolysis in vegetative organs and accumulation of amino acids in the endosperm cavity, conversion of amino acids, and storage protein synthesis in endosperm of wheat grain. Results Delayed nitrogen topdressing up-regulated gene expression related to nitrogen metabolism and protease synthesis in the flag leaf, followed by more free amino acids being transported to both the cavity and the endosperm from 7 days after anthesis (DAA) to 13 DAA in TL1. TL1 enhanced the conversion between free amino acids in endosperm and upregulated the expression of genes encoding high molecular weight (HMW) and low molecular weight (LMW) subunits and protein disulfide isomerases-like (PDIL) proteins, indicating that the synthesis and folding of glutenin were enhanched by delayed nitrogen topdressing. As a consequense, the content of glutenin macropolymers (GMP) and glutenin increased with delaying nitrogen topdressing. Conclusions The results highlight the relationship between nitrogen remobilization and final grain protein production and suggest that the nitrogen remobilization processes could be a potential target for improving the quality of wheat grain. Additionally, specific gene expression related to nitrogen topdressing was identified, which conferred more detailed insights into underlying mechanism on the modification protein quality.

To develop efficient N management strategies for high wheat NUE and minimizing the environmental impact of N losses under asymmetric warming, 15N micro-plot experiments were conducted to investigate the effects of night-warming during winter (warming by 1.47–1.56°C from tillering to jointing), spring (warming by 1.68–1.82°C from jointing to booting), and winter + spring (warming by 1.53–1.64°C from tillering to booting) on root growth and distribution of winter wheat, the fates of 15N-labeled fertilizer, and their relationships in 2015–2017. The results showed that night-warming increased the recovery of basal 15N and top-dressed 15N, while reduced the residual and loss of basal 15N and top-dressed 15N. The losses decreases of top-dressed 15N were higher than those of basal 15N, indicating that night-warming reduced losses of fertilizer 15N mainly by reducing losses of top–dressed 15N. Moreover, pre-anthesis root dry matter accumulation rate in 0–60 cm soil layer were promoted, resulted in improved root biomass and root/shoot ratio, which favored increasing recovery of fertilizer 15N and reducing losses of fertilizer 15N. Furthermore, residual fertilizer 15N content in 0–100 cm soil layer was reduced, which was associated with improved root weight density in 0–60 cm soil layer, resulted in reduced leaching losses of fertilizer 15N. The path analysis showed that root dry matter distribution in 0–20 cm soil layer was the most important in contributing to reducing losses of total fertilizer 15N compared with other soil layers. Two years data showed that winter and spring night-warming gave better root growth and distribution in 0–20 cm soil layer, resulted in reduced the losses of fertilizer 15N and improved the recovery of fertilizer 15N, while maximizing grain yield of winter wheat, and winter + spring night-warming resulted in higher advantages than winter night-warming and spring night-warming.

Drought is among the main environmental stressors that reduces wheat production. Nitrogen (N) availability affects plant adaptation to abiotic stress, but the effect of low N (LN) on drought tolerance is unclear. To identify the effect of LN priming on water-deficit stress tolerance in wheat seedlings, we primed cultivar Yangmai158 with 0.25 mM N for 7 days, and then added 20% polyethylene glycol 6000 as a water-deficit treatment for 5 days. The net photosynthetic rate (Pn), plant biomass, and plant growth rate (GR) were significantly reduced under water-deficit conditions; such decreases were less severe in LN-primed (LND) plants than non-primed (CKD) plants. The leaf relative water content (LRWC) decreased under water-deficit conditions, which in turn led to a reduced transpiration rate, stomatal conductance, and intercellular CO2 concentration (Ci), causing a stomatal limitation on photosynthesis. LN priming also enhanced root growth, resulting in a higher LRWC and less stomatal limitation in LND plants than CKD plants. PSII quantum efficiency, photochemical quenching, and maximum PSII quantum efficiency were reduced under water-deficit conditions, indicating photoinhibition. However, LN priming increased the electron flux to photorespiration and the Mehler pathway, reducing photoinhibition. In conclusion, LN priming improved the leaf water status and increased alternative electron flux to attenuate photoinhibition, thus alleviating the inhibition of photosynthesis, and growth due to water deficiency.

Salicylic acid (SA) can induce plant resistance to biotic and abiotic stresses through cross talk with other signaling molecules, whereas the interaction between hydrogen peroxide (H2O2) and abscisic acid (ABA) in response to SA signal is far from clear. Here, we focused on the roles and interactions of H2O2 and ABA in SA-induced freezing tolerance in wheat plants. Exogenous SA pretreatment significantly induced freezing tolerance of wheat via maintaining relatively higher dark-adapted maximum photosystem II quantum yield, electron transport rates, less cell membrane damage. Exogenous SA induced the accumulation of endogenous H2O2 and ABA. Endogenous H2O2 accumulation in the apoplast was triggered by both cell wall peroxidase and membrane-linked NADPH oxidase. The pharmacological study indicated that pretreatment with dimethylthiourea (H2O2 scavenger) completely abolished SA-induced freezing tolerance and ABA synthesis, while pretreatment with fluridone (ABA biosynthesis inhibitor) reduced H2O2 accumulation by inhibiting NADPH oxidase encoding genes expression and partially counteracted SA-induced freezing tolerance. These findings demonstrate that endogenous H2O2 and ABA signaling may form a positive feedback loop to mediate SA-induced freezing tolerance in wheat.

Heat stress during grain filling substantially decreases wheat productivity; thus, to ensure food security, heat tolerance in wheat needs to be developed. In this study, we evaluated the effect of heat priming applied during the stem-elongation stage, booting and anthesis, followed by 5 days of severe heat stress (a 7.86°C rise in temperature) during the grain-filling stage on physiological activities and grain yield of winter wheat in pot experiments during the 2015-2017 growing seasons using the winter wheat cultivars Yangmai 18 (a vernal type) and Yannong 19 (a facultative type). Compared with the damage observed in non-primed plants, heat priming during the stem-elongation stage and booting significantly prevented the grain-yield damage caused by heat stress during grain filling. Heat-primed plants displayed higher sucrose contents and sucrose-phosphate activity in leaves and greater above-ground dry matter than non-primed plants. Priming during stem elongation and booting led to increased photosynthetic capacity, stomatal conductance and chlorophyll contents in comparison with non-priming. Improved tolerance to heat stress due to the enhanced activities of antioxidant enzymes superoxide dismutase and peroxidase and reductions in reactive oxygen species and malondialdehyde production was observed in primed plants compared with non-primed plants of both cultivars. The positive effect of heat priming on the response to heat stress during grain filling was more pronounced in plants primed at the booting stage than in those primed at the stem-elongation or anthesis stage. Moreover, the vernal-type Yangmai 18 benefited more from heat priming than did Yannong 19, as evidenced by its higher productivity. We conclude that heat priming during early reproductive-stage growth can improve post-anthesis heat tolerance in winter wheat.

Drought is the major abiotic stress that decreases plant water status, inhibits photosynthesis, induces oxidative stress, restricts growth and finally lead to the reduction of wheat yield. It has been proven that drought priming during vegetative growth stage could enhance tolerance to drought stress at grain filling in wheat. However, whether drought priming imposed at grain filling in parental plants could induce drought tolerance in the offspring is not known. In this study, drought priming was successively applied in the first, the second and the third generation of wheat to obtain the plants of T1 (primed for one generation), T2 (primed for two generations), T3 (primed for three generations). The differently primed plants were then subjected to drought stress during grain filling in the fourth generation. Under drought stress, the parentally primed (T1D, T2D, T3D) plants, disregarding the number of generations, showed higher grain yield, leaf photosynthetic rate and antioxidant capacity as well as lower O2•− release rate and contents of H2O2 and MDA than the non-primed (T0D) plants, suggesting that drought priming induced the transgenerational stress tolerance to drought stress. Moreover, the parentally primed plants showed higher leaf water status, which may result from the higher contents of proline and glycine betaine, and higher activities of Δ1-pyrroline-5-carboxylate synthetase (P5CS) and betaine aldehyde dehydrogenase (BADH), compared with the non-primed plants under drought stress. In addition, there was no significant difference among three generations under drought, and the drought priming in parental generations did not affect the grain yield of the offspring plants under control condition. Collectively, the enhanced accumulation of proline and glycine betaine in the parentally primed plants could have played critical roles in parental priming induced tolerance to drought stress. This research provided a potential approach to improve drought tolerance of offspring plants by priming parental plants.

Systemic wound response (SWR), a well-characterized systemic signaling response, plays crucial roles in plant defense responses. Progress in understanding of the SWR in abiotic stress has also been aided by the researchers. However, the function of SWR in freezing stress remains elusive. In this study, we showed that local mild mechanical wounding enhanced freezing tolerance in newly occurred systemic leaves of wheat plants (Triticum aestivum L.). Wounding significantly increased the maximal photochemical efficiency of photosystem II, net photosynthetic rate, and the activities of the antioxidant enzymes under freezing stress. Wounding also alleviated freezing-induced chlorophyll decomposition, electrolyte leakage, water lose, and membrane peroxidation. In addition, wounding-induced freezing stress mitigation was closely associated with the ratio between reduced glutathione (GSH) and oxidized glutathione (GSSG), and the ratio between ascorbate (AsA) and dehydroascorbate (DHA), as well as the contents of total soluble sugars and free amino acids. Importantly, pharmacological study showed that wounding-induced freezing tolerance was substantially arrested by pretreatment of wheat leaves with the scavenger of hydrogen peroxide (H2O2) or the inhibitor of NADPH oxidase (RBOH). These results support the hypothesis that local mechanical wounding-induced SWR in newly occurred leaves is largely attributed to RBOH-dependent H2O2 production, which may subsequently induce freezing tolerance in wheat plants. This mechanism may have a potential application to reduce the yield losses of wheat under late spring freezing conditions.Highlights:In our previous research, we found that local mechanical wounding could induce freezing tolerance in the upper systemic leaves of wheat plants. Surprisingly, in this paper, we further demonstrated that local mechanical wounding could also increase freezing resistance in newly occurred leaves of wheat plants. RBOH mediated H2O2 and ascorbate–glutathione cycle participate in this systemic wound response.

Mechanical wounding is a common stress caused by herbivores or manual and natural manipulations, whereas its roles in acclimation response to a wide spectrum of abiotic stresses remain unclear. The present work showed that local mechanical wounding enhanced freezing tolerance in untreated systemic leaves of wheat plants (Triticum aestivum L.), and meanwhile the signal molecules hydrogen peroxide (H2O2) and nitric oxide (NO) were accumulated systemically. Pharmacological study showed that wounding-induced NO synthesis was substantially arrested by pretreatment with scavengers of reactive oxygen species and an inhibitor of NADPH oxidase (respiratory burst oxidase homolog, RBOH). On the contrary, wounding-induced H2O2 accumulation was not sensitive to NO synthetic inhibitors or scavenger, indicating that H2O2 acts upstream of NO in wounding signal transduction pathways. Cytochemical and vascular tissues localizations approved that RBOH-dependent H2O2 acts as long-distance signal in wounding response. Transcriptome analysis revealed that 279 genes were up-regulated in plants treated with wounding and freezing, but not in plants treated with freezing alone. Importantly, freezing- and wounding-induced genes were significantly enriched in the categories of “photosynthesis” and “signaling.” These results strongly supported that primary mechanical wounding can induce freezing tolerance in wheat through the systemic accumulation of NO and H2O2, and further modifications in photosystem and antioxidant system.

The diurnal and seasonal temperature rising patterns “asymmetric warming,” plays an important role in crop distribution and productivity. Asymmetric warming during the early growth periods of winter wheat (Triticum aestivum L.) profoundly affects vegetative growth and post-anthesis grain productivity, but the underlying physiological mechanism is still unclear. We conducted field experiments from 2012 to 2014 using two wheat cultivars, namely, Yangmai-13 (vernal type) and Yannong-19 (semi-winter type), to investigate the influences of night-warming during the winter (warming by 1.56–1.67°C from tillering to jointing) or during the spring (warming by 1.78–1.92°C from jointing to booting) on post-anthesis physiological activities and grain-filling processes. Both night-warming treatments enhanced the source activity by increasing flag leaf area, chlorophyll content, and photosynthetic capability in both cultivars compared with those of the control. The night-warming treatments caused an increase in the antioxidant activities of superoxide dismutase (SOD), peroxidase, and catalase (CAT) in the flag leaves of both cultivars, while ROS contents such as superoxide anion radical (O2•−) and hydrogen peroxide (H2O2) decreased. Moreover, the expression levels of Rubisco activase B (RcaB), major chlorophyll a/b-binding protein (Cab), chloroplast Cu/Zn superoxide dismutase (Cu/Zn-SOD), mitochondrial manganese superoxide dismutase (Mn-SOD), and CAT genes were upregulated at anthesis and were associated with higher photosynthetic capacity and antioxidant activities. Furthermore, night-warming improved sink activities by increasing the concentrations of grain indole-3-acetic acid and cytokinins as well as the sucrose synthase activity for both cultivars. Winter night-warming showed greater potential for improving source strength and grain filling, with consistent performance in both cultivars compared with that of spring night-warming. We concluded form these results that night-warming can improve source and sink capacities in winter wheat, and winter night-warming has greater advantages in this respect than does spring warming.

Most of the studies about NH4+ stress mechanism simply address the effects of free NH4+, failing to recognize the changed nitrogen assimilation products. The objective of this study was to elucidate the effects of glutamate on root growth under high ammonium (NH4+) conditions in winter wheat (Triticum aestivum L.). Hydroponic experiments were conducted using two wheat cultivars, AK58 (NH4+-sensitive) and Xumai25 (NH4+-tolerant) with either 5 mM NH4+ nitrogen (AN) as stress treatment or 5 mM nitrate (NO3-) nitrogen as control. To evaluate the effects of NH4+-assimilation products on plant growth, 1 μM L-methionine sulfoximine (MSO) (an inhibitor of glutamine synthetase (GS)) and 1 mM glutamates (a primary N assimilation product) were added to the solutions, respectively. The AN significantly reduced plant biomass, total root length, surface area and root volume in both cultivars, but less effect was observed in Xumai25. The inhibition effects were alleviated by the application of MSO but strengthened by the application of glutamate. The AN increased the activities of GS, glutamate dehydrogenase (GDH) in both cultivars, resulting in higher glutamate contents. However, its contents were decreased by the application of MSO. Compared to AK58, Xumai25 showed lower glutamate contents due to its higher activities of glutamic-oxaloacetic transaminase (GOT) and glutamic-pyruvic transaminase (GPT). With the indole-3-acetic acid (IAA) contents decreasing in roots, the ratio of shoot to root in IAA was increased, and further increased by the application of glutamate, and reduced by the application of MSO, but the ratio was lower in Xumai25. Meanwhile, the total soluble sugar contents and its root to shoot ratio also showed similar trends. These results indicate that the NH4+-tolerant cultivar has a greater transamination ability to prevent glutamate over-accumulation to maintain higher IAA transport ability, and consequently promoted soluble sugar transport to roots, further maintaining root growth.

Exploring Approaches To Optimizing Spatial Distributions of Leaf Area index (Lai) and Leaf Nitrogen Content (Lnc) Should Be Useful For increasing Grain Yield in Rice (Oryza Sativa). THe Primary Objective of This Study Was ToCharacterize The Variation of Lai and Lnc Distributions Within The Canopy in Relation To Grain Yield and N Uptake in Rice.Two Experiments With Different N Fertilization Rates Under Conventional and intermittent Irrigation (Ci and Ii, Respectively) Were Conducted in 2002 and 2003, Using Japonica Rice Cultivar Wuyujing9. The Results Showed That Grain Yield and N Uptake Were Significantly Different Among Application Rates of N (N Rates), But Did Not Differ Between Ci and Ii. Lai Distribution At Full Heading Was Affected Significantly By N Rate But Hardly By The Irrigation Method. individual Lai increased With The N Rate. To Achieve A High Yield, The Proper Distribution of Lai in The Canopy Can Be Designed As The Largest in The 2Nd Leaf From The Top, Followed By The 3Rd and 4Th Leaves, and The Smallest in The Top Leaf. Lnc On The Base of Both Area and Dry Matter At 15 Days After Full Heading Decreased From The Top To Lower Leaves in The Canopy, and Significantly increased With The N Rate. Grain Yield Was Enhanced Linearly With The increasing N Content of The Upper Two Leaves, But Hindered By The High N Content of Lower Leaves. These Results indicate That The Spatial Distributions of Both Lnc and Lai Could Be Optimized To Achieve Maximum Canopy Photosynthesis and Grain Yield in Rice.

Meteorological conditions including temperature, sunshine and precipitation during grain growth are the primary factors determining the variation of the protein content of grain (PC) in wheat. On the basis of field experiments, a simplified regression model was developed for predicting the PC in winter wheat. From stepwise regression analysis, it was found that the PC of high-protein cultivars was correlated with the difference between daily maximum and minimum temperatures (∆T) when ∆ T variation under the environment was significant, but with the interaction of mean temperature (Tmean) × total sunshine hours from anthesis to maturity (TSUN) under the environment with ∆ T variation less than 5%. In medium-protein cultivars, the PC was correlated with TSUN, and in low-protein cultivars, with the combination of Tmean, total rainfall from anthesis to maturity (TR) and TSUN. The climatic factors influencing PC were further quantified incorporating five genetic parameters. The ∆ T and TSUN were linearly correlated with PC, and Tmean was quadratically correlated with PC. The precipitation was linearly correlated with PC if it was less than 50mm, otherwise quadratically. The average root mean square error (RMSE) values of the estimated PC relative to the observed value were less than 7 percent, indicating a good fit between the estimated and observed PC. Thus, it is concluded that the present model can predict the PC of different winter wheat cultivars under various climate environments.

Source-sink relationships influence photosynthesis. So far, the limiting factors for photosynthesis of wheat cultivars with different source-sink relationships have not been determined. We aimed to determine the variation patterns of photosynthetic characteristics of wheat cultivars with different source-sink relationships. In this study, two wheat cultivars with different source-sink relationships were selected for photosynthetic physiological analyses. The results showed that YM25 (source-limited cultivar) had higher photosynthetic efficiency compared to YM1 (sink-limited cultivar). This is mainly due to a stronger photochemical efficiency, electron transfer capacity, and Rubisco carboxylation capacity of YM25. YM25 accumulated less soluble carbohydrates in flag leaves than YM1. This is mainly due to the stronger sucrose synthesis and transport capacity of YM25 by presenting higher sucrose-related enzyme activities and gene expression. A PCA analysis showed that Rubisco was the main factor limiting the photosynthetic capacity of YM25. The soluble sugar accumulation in flag leaves and sink limitation decreased the photosynthetic activity of YM1. Increased N application improved source-sink relationships and increased grain yield and source leaf photosynthetic capacity in both two wheat cultivars. Taken together, our findings suggest that Rubisco and sucrose synthesis and translocation are involved in the regulation of photosynthesis of wheat cultivars with different sourcesink relationships and that source and sink limitation effects should be considered in photosynthesis. [ABSTRACT FROM AUTHOR]

The use of slow-release fertilizers and seed-fertilizers cause localized highammonium (NH4+) environments in agricultural fields, adversely affecting wheat growth and development and delaying its yield. Thus, it is important to investigate the physiological responses of wheat and its tolerance to NH4+ stress to improve the adaptation of wheat to high NH4+ environments. In this study, the physiological mechanisms of ammonium tolerance in wheat (Triticum aestivum) were investigated in depth by comparative analysis of two cultivars: NH4+-tolerant Xumai25 and NH4+-sensitive Yangmai20. Cultivation under hydroponic conditions with high NH4+ (5 mM NH4+, AN) and nitrate (5 mM NO3-, NN), as control, provided insights into the nuanced responses of both cultivars. Compared to Yangmai20, Xumai25 displayed a comparatively lesser sensitivity to NH4+ stress, as evident by a less pronounced reduction in dry plant biomass and a milder adverse impact on root morphology. Despite similarities in NH4+ efflux and the expression levels of TaAMT1.1 and TaAMT1.2 between the two cultivars, Xumai25 exhibited higher NH4+ influx, while maintaining a lower free NH4+ concentration in the roots. Furthermore, Xumai25 showed a more pronounced increase in the levels of free amino acids, including asparagine, glutamine, and aspartate, suggesting a superior NH4+ assimilation capacity under NH4+ stress compared to Yangmai20. Additionally, the enhanced transcriptional regulation of vacuolar glucose transporter and glucose metabolism under NH4+ stress in Xumai25 contributed to an enhanced carbon skeleton supply, particularly of 2-oxoglutarate and pyruvate. Taken together, our results demonstrate that the NH4+ tolerance of Xumai25 is intricately linked to enhanced glucose metabolism and optimized glucose transport, which contributes to the robust NH4+ assimilation capacity. [ABSTRACT FROM AUTHOR]