Drought Tolerance In Maize Research Articles

Dissection of Maize Drought Tolerance at the Flowering Stage Using Genome-Wide Association Studies.

Drought is one of the most critical environmental factors constraining maize production. When it occurs at the flowering stage, serious yield losses are caused, and often, the damage is irretrievable. In this study, anthesis to silk interval (ASI), plant height (PH), and ear biomass at the silking date (EBM) of 279 inbred lines were studied under both water-stress (WS) and well-water (WW) field conditions, for three consecutive years. Averagely, ASI was extended by 25.96%, EBM was decreased by 17.54%, and the PH was reduced by 12.47% under drought stress. Genome-wide association studies were carried out using phenotypic values under WS, WW, and drought-tolerance index (WS-WW or WS/WW) and applying a mixed linear model that controls both population structure and relative kinship. In total, 71, 159, and 21 SNPs, located in 32, 59, and 12 genes, were significantly (P < 10−5) associated with ASI, EBM, and PH, respectively. Only a few overlapped candidate genes were found to be associated with the same drought-related traits under different environments, for example, ARABIDILLO 1, glycoprotein, Tic22-like, and zinc-finger family protein for ASI; 26S proteasome non-ATPase and pyridoxal phosphate transferase for EBM; 11-ß-hydroxysteroid dehydrogenase, uncharacterised, Leu-rich repeat protein kinase, and SF16 protein for PH. Furthermore, most candidate genes were revealed to be drought-responsive in an association panel. Meanwhile, the favourable alleles/key variations were identified with a haplotype analysis. These candidate genes and their key variations provide insight into the genetic basis of drought tolerance, especially for the female inflorescence, and will facilitate drought-tolerant maize breeding.

Transcriptome expression profiles reveal response mechanisms to drought and drought-stress mitigation mechanisms by exogenous glycine betaine in maize.

Drought stress is one of the major abiotic stresses that limit growth, development and yield of maize crops. To better understand the responses of maize inbred lines with different levels of drought resistance and the molecular mechanism of exogenous glycine betaine (GB) in alleviating drought stress, the responses of two maize inbred lines to drought stress and to the stress-mitigating effects of exogenous GB were investigated. Seedling morphology, physiological and biochemical indexes, root cell morphology and root transcriptome expression profiles were compared between a drought-tolerant inbred line Chang 7-2 and drought-sensitive inbred line TS141. Plants of both lines were subjected to treatments of drought stress alone and drought stress with application of exogenous GB. The results showed that with the increase of drought treatment time, the growth and development of TS141 were inhibited, while those of Chang 7-2 were not significantly different from those of the control (no drought stress and GB). Compared with the corresponding data of the drought-stress group, every index measured from the two inbred lines indicated mitigating effects from exogenous GB, and TS141 produced stronger mitigating responses due to the GB. Transcriptome analysis showed that 562 differentially expressed genes (DEGs) were up-regulated and 824 DEGs were down-regulated in both inbred lines under drought stress. Due to the exogenous GB, 1061 DEGs were up-regulated and 424 DEGs were down-regulated in both lines. In addition, quantitative real-time polymerase chain reaction (qRT-PCR) was used to verify 10 DEGs screened from the different treatments. These results showed that the expression of 9 DEGs were basically consistent with their respective transcriptome expression profiles. The results of this study provide models of potential mechanisms of drought tolerance in maize as well as potential mechanisms of how exogenous GB may regulate drought tolerance.

Genome-wide identification of HD-ZIP transcription factors in maize and their regulatory roles in promoting drought tolerance.

Drought is the main limiting factor of maize productivity, therefore improving drought tolerance in maize has potential practical importance. Cloning and functional verification of drought-tolerant genes is of great importance to understand molecular mechanisms under drought stress. Here, we employed a bioinformatic pipeline to identify 42 ZmHDZ drought responsive genes using previously reported maize transcriptomic datasets. The coding sequences, exon-intron structure and domain organization of all the 42 genes were identified. Phylogenetic analysis revealed evolutionary conservation of members of the ZmHDZ genes in maize. Several regulatory elements associated with drought tolerance were identified in the promoter regions of ZmHDZ genes, indicating the implication of these genes in plant response to drought stress. 42 ZmHDZ genes were distributed unevenly on 10 chromosomes, and 24 pairs of gene duplications were the segmental duplication. The expression of several ZmHDZ genes was upregulated under drought stress, and ZmHDZ9 overexpressing transgenic plants exhibited higher SOD and POD activities and higher accumulation of soluble proteins under drought stress which resulted in enhanced developed phenotype and improved resistance. The present study provides evidence for the evolutionary conservation of HD-ZIP transcription factors homologs in maize. The results further provide a comprehensive insight into the roles of ZmHDZ genes in regulating drought stress tolerance in maize.

Impact of water deficit stress on traits influencing the drought tolerance and yield of maize (Zea mays L.) genotypes

Impact of water deficit stress on traits influencing the drought tolerance and yield of maize (Zea mays L.) genotypes

Poly-\u03b3-glutamic acid enhanced the drought resistance of maize by improving photosynthesis and affecting the rhizosphere microbial community

BackgroundCompared with other abiotic stresses, drought stress causes serious crop yield reductions. Poly-γ-glutamic acid (γ-PGA), as an environmentally friendly biomacromolecule, plays an important role in plant growth and regulation.ResultsIn this project, the effect of exogenous application of γ-PGA on drought tolerance of maize (Zea mays. L) and its mechanism were studied. Drought dramatically inhibited the growth and development of maize, but the exogenous application of γ-PGA significantly increased the dry weight of maize, the contents of ABA, soluble sugar, proline, and chlorophyll, and the photosynthetic rate under severe drought stress. RNA-seq data showed that γ-PGA may enhance drought resistance in maize by affecting the expression of ABA biosynthesis, signal transduction, and photosynthesis-related genes and other stress-responsive genes, which was also confirmed by RT–PCR and promoter motif analysis. In addition, diversity and structure analysis of the rhizosphere soil bacterial community demonstrated that γ-PGA enriched plant growth promoting bacteria such as Actinobacteria, Chloroflexi, Firmicutes, Alphaproteobacteria and Deltaproteobacteria. Moreover, γ-PGA significantly improved root development, urease activity and the ABA contents of maize rhizospheric soil under drought stress. This study emphasized the possibility of using γ-PGA to improve crop drought resistance and the soil environment under drought conditions and revealed its preliminary mechanism.ConclusionsExogenous application of poly-γ-glutamic acid could significantly enhance the drought resistance of maize by improving photosynthesis, and root development and affecting the rhizosphere microbial community.

Physiological and molecular level understanding of advanced carbon dots to enhance maize drought tolerance: modulation of photosynthesis and signaling molecules

Foliar application of polyacrylic acid modified and nitrogen doped carbon dots (PNDs) could enhance maize drought tolerance through scavenging ROS, improving photosynthesis, and modulating plant signals.

Overexpression of the homeobox-leucine zipper protein ATHB-6 improves the drought tolerance of maize (Zea mays L.)

Homeo-Leucine Zipper (HD-Zip) proteins are a class of transcription factors unique to higher plants and are involved in plant stress responses and regulation of growth and development. However, the function of maize HD-Zip genes in enhancing drought tolerance is unknown. Here, Sub-Cellular Localization results showed that ATHB-6 fusion proteins were only localized in the nucleus. The malondialdehyde content was lower than the wild type under drought tolerance, proving that the introduction of the ATHB-6 gene can improve the drought tolerance of plants. Follow-up analysis showed that ATHB-6 could promote root growth and activities of a series of ROS-scavenging enzymes in maize. Moreover, overexpression of ATHB-6 in maize activated the expression of critical genes in the ROS signals pathway and ABA-dependent pathway under drought tolerance.Our results provides a significant advancement in undestanding the functions of HD-Zip transcription factors in maize.

Is Photoprotection of PSII One of the Key Mechanisms for Drought Tolerance in Maize?

Drought is one of the most important abiotic stress factors limiting maize production worldwide. The objective of this study was to investigate whether photoprotection of PSII was associated with the degree of drought tolerance and yield in three maize hybrids (30Y87, 31R88, P3939). To do this, three maize hybrids were subjected to three cycles of drought, and we measured the activities of photosystem II (PSII) and photosystem I (PSI). In a second field experiment, three maize hybrids were subjected to drought by withholding irrigation, and plant water status, yield and yield attributes were measured. Drought stress decreased leaf water potential (ΨL) in three maize hybrids, and this reduction was more pronounced in hybrid P3939 (−40%) compared to that of 30Y87 (−30%). Yield and yield attributes of three maize hybrids were adversely affected by drought. The number of kernels and 100-kernel weight was the highest in maize hybrid 30Y87 (−56%, −6%), whereas these were lowest in hybrid P3939 (−88%, −23%). Drought stress reduced the quantum yield of PSII [Y(II)], photochemical quenching (qP), electron transport rate through PSII [ETR(II)] and NPQ, except in P3939. Among the components of NPQ, drought increased the Y(NPQ) with concomitant decrease in Y(NO) only in P3939, whereas Y(NO) increased in drought-stressed plants of hybrid 30Y87 and 31R88. However, an increase in cyclic electron flow (CEF) around PSI and Y(NPQ) in P3939 might have protected the photosynthetic machinery but it did not translate in yield. However, drought-stressed plants of 30Y87 might have sufficiently downregulated PSII to match the energy consumption in downstream biochemical processes. Thus, changes in PSII and PSI activity and development of NPQ through CEF are physiological mechanisms to protect the photosynthetic apparatus, but an appropriate balance between these physiological processes is required, without which plant productivity may decline.

Melatonin Enhances Drought Tolerance by Regulating Leaf Stomatal Behavior, Carbon and Nitrogen Metabolism, and Related Gene Expression in Maize Plants

It is commonly known that exogenously applied melatonin can alleviate the impact of drought stress, but the mechanism used by melatonin to regulate stomatal behavior and carbon (C) and nitrogen (N) metabolism to increase drought resistance remains elusive. Herein, our aim was to investigate the influence of exogenous melatonin on the regulation of C and N metabolism in maize plants under water deficit. In this study, we analyzed stomatal behavior, the key components of C and N metabolism, and the gene expression and activity of enzymes involved in the C and N metabolism in maize plants. The results showed that the application of melatonin (100 μM) significantly increased maize growth and sustained the opening of stomata, and secondarily increased the photosynthetic capacity in maize. Under drought stress, foliar application of melatonin induced the gene transcription and activities of sucrose phosphate synthetase, ADP-glucose pyrophosphorylase, phosphoenolpyruvate carboxylase, and citrate synthase, resulting in the enhancement of sucrose and starch synthesis and the tricarboxylic acid (TCA) cycle. This enhancement in sugar biosynthesis and the TCA cycle might lead to stronger N assimilation. As anticipated, NO3– reduction and NH4+ assimilation were also strengthened after melatonin treatment under drought stress. An increase was observed in some key enzymatic activities and transcription involved in nitrogen metabolism, such as that of nitrate reductase, nitrite reductase, glutamate synthase, and glutamine synthetase, in melatonin-treated, drought-stressed maize. Moreover, melatonin attenuated the drought-induced damage by reducing protein degradation and increasing the level of proline. Conclusively, our results indicate that exogenous melatonin enhances drought tolerance in maize via promoting stomatal opening and regulating C and N metabolism and related gene expression.

A vegetative storage protein improves drought tolerance in maize.

SummaryVegetative storage proteins (VSPs) are known to serve as nitrogen reserves in many dicot plants but remain undiscovered in grasses, most widely grown group of crops globally. We identified and characterized a VSP in maize and demonstrated that its overexpression improved drought tolerance. Nitrogen supplementation selectively induced a mesophyll lipoxygenase (ZmLOX6), which was targeted to chloroplasts by a novel N‐terminal transit peptide of 62 amino acids. When ectopically expressed under the control of various tissue‐specific promoters, it accumulated to a fivefold higher level upon expression in the mesophyll cells than the wild‐type plants. Constitutive expression or targeted expression specifically to the bundle sheath cells increased its accumulation by less than twofold. The overexpressed ZmLOX6 was remobilized from the leaves like other major proteins during grain development. Evaluated in the field over locations and years, transgenic hybrids overexpressing ZmLOX6 in the mesophyll cells significantly outyielded nontransgenic sibs under managed drought stress imposed at flowering. Additional storage of nitrogen as a VSP in maize leaves ameliorated the effect of drought on grain yield.

Silicon Seed Priming Combined with Foliar Spray of Sulfur Regulates Photosynthetic and Antioxidant Systems to Confer Drought Tolerance in Maize (Zea mays L.)

The present study evaluated the effect of silicon (Si) seed priming and sulfur (S) foliar spray on drought tolerance of two contrasting maize hybrids viz. drought tolerant Hi-Corn 11 and susceptible P-1574. The maize seeds were primed with (3 mM Na2SiO3) or without Si (hydropriming) and later sown in pots filled with sandy loam soil. Drought stress (25–30% water holding capacity or WHC) was initiated at cob development stage (V5) for two weeks, whereas the well-watered plants were grown at 65–70% WHC. On appearance of drought symptoms, foliar spray of S was done using 0.5% and 1.0% (NH4)2SO4, whereas water spray was used as a control. The drought-stressed plants were grown for further two weeks at 25–30% WHC before the final harvest. The results showed a marked effect of Si seed priming and foliar S spray on biomass, physiological and enzymatic processes as well as macronutrient concentrations of maize. In comparison to control, the highest increase in leaf relative water content (25%), chlorophyll a content (56%), carotenoids (26%), photosynthetic rate (64%), stomatal conductance (56%) and intercellular CO2 concentration (48%) was observed by Si seed priming + S foliar spray (Si + S) under water deficit conditions. Also, Si + S application stimulated the activity of catalase (45%), guaiacol peroxidase (38%) and superoxide dismutase (55%), and improved NPK concentrations (40–63%) under water limitations. Our results suggest that Si seed priming + foliar spray of S is more effective than the individual application of these nutrients to enhance drought tolerance in maize.

Foliar carbon dot amendment modulates carbohydrate metabolism, rhizospheric properties and drought tolerance in maize seedling

Improving maize drought tolerance is of great importance for scaling up production due to food security and population growth. Carbon dots (CDs) were synthesized by hydrothermal method with citric acid and ethylenediamine as carbon sources. Then, CDs (5 ml, 5 mg‧L−1) were sprayed on 25th day-old maize (Zea mays L., drought-stress, 35% soil moisture) for seven consecutive days (spraying ultra-pure water as control), after which the physiological parameters and rhizospheric properties of maize under drought were evaluated. Foliar sprayed CDs (5 mg‧L−1) could increase root exudates (e.g., succinic acid (14.5 folds), pyruvic acid (10.0 folds), and betaine (11.8 folds)), and modify microbial community. Particularly, the relative abundance of Pseudomonas, Sphingomonas, Nitrospira, and Conocybe were significantly increased by 344.4%, 233.3%, 126.2%, and 122.6%, respectively. The altered microbial abundance could improve soil available nitrogen and phosphorus by 33.5% and 16.8%, respectively, and increase plant water uptake by 37.2%. The change of exudate synthesis and microbial abundance could be driven by the significantly increased in net photosynthesis rate by 122.9%, and carbohydrate content by 35.4% in shoots and 113.6% in roots, respectively upon foliar application of CDs. Meanwhile, fresh weight of shoots and roots were increased by 62.1% and 50.6%, and dry weight of shoots and roots were increased by 29.2% and 37.5%, respectively. These results demonstrated that foliar application of CDs could improve the rhizosphere environment to enhance maize drought tolerance and even growth. Therefore, foliar application of CDs would be a promising strategy for sustainable nano-agriculture in response to drought stress.

ZmCCT regulates photoperiod-dependent flowering and response to stresses in maize

BackgroundAppropriate flowering time is very important to the success of modern agriculture. Maize (Zea mays L.) is a major cereal crop, originated in tropical areas, with photoperiod sensitivity. Which is an important obstacle to the utilization of tropical/subtropical germplasm resources in temperate regions. However, the study on the regulation mechanism of photoperiod sensitivity of maize is still in the early stage. Although it has been previously reported that ZmCCT is involved in the photoperiod response and delays maize flowering time under long-day conditions, the underlying mechanism remains unclear.ResultsHere, we showed that ZmCCT overexpression delays flowering time and confers maize drought tolerance under LD conditions. Implementing the Gal4-LexA/UAS system identified that ZmCCT has a transcriptional inhibitory activity, while the yeast system showed that ZmCCT has a transcriptional activation activity. DAP-Seq analysis and EMSA indicated that ZmCCT mainly binds to promoters containing the novel motifs CAAAAATC and AAATGGTC. DAP-Seq and RNA-Seq analysis showed that ZmCCT could directly repress the expression of ZmPRR5 and ZmCOL9, and promote the expression of ZmRVE6 to delay flowering under long-day conditions. Moreover, we also demonstrated that ZmCCT directly binds to the promoters of ZmHY5, ZmMPK3, ZmVOZ1 and ZmARR16 and promotes the expression of ZmHY5 and ZmMPK3, but represses ZmVOZ1 and ZmARR16 to enhance stress resistance. Additionally, ZmCCT regulates a set of genes associated with plant development.ConclusionsZmCCT has dual functions in regulating maize flowering time and stress response under LD conditions. ZmCCT negatively regulates flowering time and enhances maize drought tolerance under LD conditions. ZmCCT represses most flowering time genes to delay flowering while promotes most stress response genes to enhance stress tolerance. Our data contribute to a comprehensive understanding of the regulatory mechanism of ZmCCT in controlling maize flowering time and stress response.

Physiological and transcriptomic analyses of the effects of exogenous melatonin on drought tolerance in maize (Zea mays L.)

Water deficit inhibits maize (Zea mays L.) seedling growth and yield. Application of exogenous melatonin can improve drought tolerance of corn, but little is known regarding the transcriptional mechanisms of melatonin-mediated drought tolerance in maize. Increased understanding of the effects of melatonin on maize plants under drought stress is vital to alleviate the adverse effects of drought on food production in the future. The aim of this investigation was to use physiological and transcriptome analyses for exploring the possible mechanisms of exogenous melatonin against drought stress in maize. In this study, maize seedlings were subjected to drought stress and some were treated with exogenous melatonin. The physiological results showed that melatonin inhibited H2O2 accumulation and promoted the scavenging of excessive reactive oxygen species to reduce oxidative damage in maize leaves. Transcriptomic analysis identified 957 differentially expressed genes between melatonin and non-melatonin treatment groups. Further detailed analyses suggested that melatonin-regulated genes are mainly related to glutathione metabolism, calcium signaling transduction, and jasmonic acid biosynthesis. Some transcription factor families, such as WRKY, AP2/ERF-ERF, MYB, NAC, and bZIP, were also activated by exogenous melatonin. Moreover, crosstalk between melatonin and other hormones that mediate drought tolerance was observed. In conclusion, the combination of physiological and transcriptome analyses revealed some mechanisms underlying the role of melatonin in alleviating drought; knowledge of these mechanisms may assist in successful maize cultivation under drought stress.

Two calcium-dependent protein kinases enhance maize drought tolerance by activating anion channel ZmSLAC1 in guard cells.

SummaryStomatal closure is an important process to prevent water loss in plants response to drought stress, which is finely modulated by ion channels together with their regulators in guard cells, especially the S‐type anion channel AtSLAC1 in Arabidopsis. However, the functional characterization and regulation analyses of anion channels in gramineous crops, such as in maize guard cells are still limited. In this study, we identified an S‐type anion channel ZmSLAC1 that was preferentially expressed in maize guard cells and involved in stomatal closure under drought stress. We found that two Ca2+‐dependent protein kinases ZmCPK35 and ZmCPK37 were expressed in maize guard cells and localized on the plasma membrane. Lesion of ZmCPK37 resulted in drought‐sensitive phenotypes. Mutation of ZmSLAC1 and ZmCPK37 impaired ABA‐activated S‐type anion currents in maize guard cells, while the S‐type anion currents were increased in the guard cells of ZmCPK35‐ and ZmCPK37‐overexpression lines. Electrophysiological characterization in maize guard cells and Xenopus oocytes indicated that ZmCPK35 and ZmCPK37 could activate ZmSLAC1‐mediated Cl‐ and NO3 ‐ currents. The maize inbred and hybrid lines overexpressing ZmCPK35 and ZmCPK37 exhibited enhanced tolerance and increased yield under drought conditions. In conclusion, our results demonstrate that ZmSLAC1 plays crucial roles in stomatal closure in maize, whose activity is regulated by ZmCPK35 and ZmCPK37. Elevation of ZmCPK35 and ZmCPK37 expression levels is a feasible way to improve maize drought tolerance as well as reduce yield loss under drought stress.

Comparative Effects of Individual and Consortia Plant Growth Promoting Bacteria on Physiological and Enzymatic Mechanisms to Confer Drought Tolerance in Maize (Zea mays L.)

Mitigation strategies based on plant–microbe interactions to increase the performance of plants under water-deficit conditions are well documented. However, little is known about a suitable consortium of bacterial inoculants and underlying physiological and enzymatic events to improve drought tolerance in maize. We performed laboratory and pot experiments to understand the synergistic interactions among plant growth-promoting bacteria to alleviate the drought-induced damages in maize. Initially, ten bacterial strains were evaluated for their osmotic stress tolerance capacity by growing them in a media containing 0, 10, 20, and 30% polyethylene glycol (PEG-6000). Also, the seeds of a drought tolerant (NK-6654) and sensitive (SD-626) maize cultivar were inoculated with these bacterial strains in the first pot experiment to determine their effects on the growth and physiological processes. Later, in the second pot experiment, the best performing inoculants were selected to study the individual and synergistic effects of bacterial inoculation to confer drought tolerance in maize. Our findings showed that the inoculation with tolerant strains resulted in higher photosynthetic activity (25–39%), maintenance of leaf water status (14–18%) and pigments (27–32%), and stimulation of antioxidant machinery (28–38%) than no inoculation in water-stressed maize seedlings. Moreover, the treatment with bacteria consortia further stimulated the drought protective mechanisms and resulted in higher efficiency of photosynthetic (47–61%) and antioxidant systems (42–62%) than the individual inoculants under water-deficit conditions. We conclude that the inoculation with microbial consortia regulates water uptake, photosynthetic performance, and stress metabolites to minimize drought-induced damages in maize.

Exogenously Applied Proline Modulates Drought Tolerance in Maize (Zea mays L)

Improving drought stress tolerance in maize is essential to increase its production and yield worldwide. Thus, the present study was conducted to investigate the improvement of drought tolerance in maize (Zea mays L.) by exogenous application of proline (25 and 50mM) on two maize varieties. Maize plants were subjected to drought stress at various phases of plant growth under pot culture conditions and proline was applied as foliar spray. Water deficit stress caused a significant decrease (by approximately 25%) in growth and yield of both maize varieties by decreasing plant height, cob length, dry root weight, grains per cob and 100-grain weight. Water deficit stress also decreased chlorophyll and intercellular proline contents, and antioxidant enzyme activities viz. catalase (CAT), guaiacol peroxidase (POX) and ascorbate peroxidase (APX). Exogenous application of proline (50 mM) was found to be more effective in increasing growth and yield of both varieties. These increases were positively associated with increased levels (by at least 15%) of chlorophyll and intracellular proline, and enhanced activities of CAT, POX and APX enzymes in both varieties. Interaction effects of exogenous proline and water deficit stress were significant in aspects of higher growth and yields and enhanced levels of chlorophyll, intracellular proline and antioxidant enzyme activities. Therefore, it is concluded that foliar application of proline improves drought tolerance by modulating chlorophyll and intracellular proline contents, and antioxidant enzyme activities.

ZmWRKY104 Transcription Factor Phosphorylated by ZmMPK6 Functioning in ABA-Induced Antioxidant Defense and Enhance Drought Tolerance in Maize.

Simple SummaryCurrent knowledge about the downstream substrate proteins of MAPKs is still limited. Our study screened a new WRKY IIa transcription factor as the substrate protein of ZmMPK6, and its phosphorylation at Thr-59 is critical to the role of ZmWRKY104 in ABA-induced antioxidant defense. Moreover, overexpression ZmWRKY104 in maize enhances the drought tolerance of transgenic plants. These findings define a mechanism for the function of ZmWRKY104 phosphorylated by ZmMPK6 in ABA-induced antioxidant defense and drought tolerance.Mitogen-activated protein kinase (MAPK) cascades are primary signaling pathways involved in various signaling pathways triggered by abiotic and biotic stresses in plants. The downstream substrate proteins of MAPKs in maize, however, are still limited. Here, we screened a WRKY IIa transcription factor (TF) in maize (Zea mays L.), ZmWRKY104, and found that it is a substrate of ZmMPK6. ZmWRKY104 physically interacts with ZmMPK6 in vitro and in vivo. Liquid chromatography–tandem mass spectrometry (LC-MS/MS) analysis results showed that threonine-59 (Thr-59, T59) was the major phosphorylation site of ZmWRKY104 by ZmMPK6. Subcellular localization analysis suggested that ZmWRKY104 acts in the nucleus and that ZmMPK6 acts in the nucleus and cytoplasmic membrane in the cytosol. Functional analysis revealed that the role of ZmWRKY104 in ABA-induced antioxidant defense depends on ZmMPK6. Moreover, overexpression of ZmWRKY104 in maize can enhance drought tolerance and relieve drought-induced oxidative damage in transgenic lines. The above results help define the mechanism of the function of ZmWRKY104 phosphorylated by ZmMPK6 in ABA-induced antioxidant defense and drought tolerance in maize.

Genomic basis underlying the metabolome-mediated drought adaptation of maize

BackgroundDrought is a major environmental disaster that causes crop yield loss worldwide. Metabolites are involved in various environmental stress responses of plants. However, the genetic control of metabolomes underlying crop environmental stress adaptation remains elusive.ResultsHere, we perform non-targeted metabolic profiling of leaves for 385 maize natural inbred lines grown under well-watered as well as drought-stressed conditions. A total of 3890 metabolites are identified and 1035 of these are differentially produced between well-watered and drought-stressed conditions, representing effective indicators of maize drought response and tolerance. Genetic dissections reveal the associations between these metabolites and thousands of single-nucleotide polymorphisms (SNPs), which represented 3415 metabolite quantitative trait loci (mQTLs) and 2589 candidate genes. 78.6% of mQTLs (2684/3415) are novel drought-responsive QTLs. The regulatory variants that control the expression of the candidate genes are revealed by expression QTL (eQTL) analysis of the transcriptomes of leaves from 197 maize natural inbred lines. Integrated metabolic and transcriptomic assays identify dozens of environment-specific hub genes and their gene-metabolite regulatory networks. Comprehensive genetic and molecular studies reveal the roles and mechanisms of two hub genes, Bx12 and ZmGLK44, in regulating maize metabolite biosynthesis and drought tolerance.ConclusionOur studies reveal the first population-level metabolomes in crop drought response and uncover the natural variations and genetic control of these metabolomes underlying crop drought adaptation, demonstrating that multi-omics is a powerful strategy to dissect the genetic mechanisms of crop complex traits.

Efficient Regulation of CO2 Assimilation Enables Greater Resilience to High Temperature and Drought in Maize.

Increasing temperatures and extended drought episodes are among the major constraints affecting food production. Maize has a relatively high temperature optimum for photosynthesis compared to C3 crops, however, the response of this important C4 crop to the combination of heat and drought stress is poorly understood. Here, we hypothesized that resilience to high temperature combined with water deficit (WD) would require efficient regulation of the photosynthetic traits of maize, including the C4–CO2 concentrating mechanism (CCM). Two genotypes of maize with contrasting levels of drought and heat tolerance, B73 and P0023, were acclimatized at high temperature (38°C versus 25°C) under well-watered (WW) or WD conditions. The photosynthetic performance was evaluated by gas exchange and chlorophyll a fluorescence, and in vitro activities of key enzymes for carboxylation (phosphoenolpyruvate carboxylase), decarboxylation (NADP-malic enzyme), and carbon fixation (Rubisco). Both genotypes successfully acclimatized to the high temperature, although with different mechanisms: while B73 maintained the photosynthetic rates by increasing stomatal conductance (gs), P0023 maintained gs and showed limited transpiration. When WD was experienced in combination with high temperatures, limited transpiration allowed water-savings and acted as a drought stress avoidance mechanism. The photosynthetic efficiency in P0023 was sustained by higher phosphorylated PEPC and electron transport rate (ETR) near vascular tissues, supplying chemical energy for an effective CCM. These results suggest that the key traits for drought and heat tolerance in maize are limited transpiration rate, allied with a synchronized regulation of the carbon assimilation metabolism. These findings can be exploited in future breeding efforts aimed at improving maize resilience to climate change.