Assessment of Nitrogen Fixation, Uptake, and Leaching in Maize/Soybean Intercropping System at Varied Soil Depths and under Phosphorus Application in Chinese Agricultural Settings

The study of Nitrogen fixation, uptake, and leaching at different soil depths in the co-cultivation of maize and soybean under phosphorus fertilization is important for sustainable agriculture. This study was conducted in Quzhou, Hebei Province, China, with MC812 maize and Jidou12 soybean varieties. Soil samples were taken from each plot to create a composite sample. The results show that nitrogen concentration varies at different depths and is higher in all treatments between 40 and 100 cm. Incorporating intercropping of maize and soybeans into farming practices can lead to more sustainable and environmentally friendly agriculture in China.

Assessing the Influence of Phosphorus Fertilization on the Growth and Yield of Maize/Soybean Intercrop by Analyzing Nitrogen Uptake

Intercropping, particularly the combination of maize and soybeans, has been widely recognized for its potential to improve nitrogen uptake and promote sustainable agriculture. This study examines the patterns of nitrogen uptake in maize and soybean intercropping systems under different growth stages and phosphorus fertilization levels and investigates the influence of nitrogen uptake on growth parameters such as plant height, leaf area, and biomass accumulation in the maize/soybean intercrop under different phosphorus fertilization regimes. The study also collected chlorophyll samples at different growth stages of maize in monoculture and intercropping with maize or soybean. The results showed that plant height was greater in V10 in both fertilized and unfertilized treatments for intercropped maize and soybean, and chlorophyll concentration was higher in VT intercropped maize. The results also showed a higher accumulation of biomass. Understanding the growth dynamics of these plants in monoculture and intercropping systems and the impact of fertilization practices is crucial for optimizing crop productivity and sustainability in agricultural systems.

Image-based root phenotyping for field-grown crops:An example under maize/soybean intercropping

Root architecture,which determines the water and nutrient uptake ability of crops,is highly plastic in response to soil environmental changes and different cultivation patterns.Root phenotyping for field-grown crops,especially topological trait extraction,is rarely performed.In this study,an image-based semi-automatic root phenotyping method for field-grown crops was developed.The method consisted of image acquisition,image denoising and segmentation,trait extraction and data analysis.Five global traits and 40 local traits were extracted with this method.A good consistency in 1st-order lateral root branching was observed between the visually counted values and the values extracted using the developed method,with R^(2)=0.97.Using the method,we found that the interspecific advantages for maize mainly occurred within 5 cm from the root base in the nodal roots of the 5th-7th nodes,and that the obvious inhibition of soybean was mostly reflected within 20 cm from the root base.Our study provides a novel approach with high-throughput and high-accuracy for field research on root morphology and branching features.It could be applied to the 3D reconstruction of field-grown root system architecture to improve the inputs to data-driven models(e.g.,OpenSimRoot)that simulate root growth,solute transport and water uptake.

Soybean maize strip intercropping:A solution for maintaining food security in China

The practice of intercropping leguminous and gramineous crops is used for promoting sustainable agriculture,optimizing resource utilization,enhancing biodiversity,and reducing reliance on petroleum products.However,promoting conventional intercropping strategies in modern agriculture can prove challenging.The innovative technology of soybean maize strip intercropping(SMSI)has been proposed as a solution.This system has produced remarkable results in improving domestic soybean and maize production for both food security and sustainable agriculture.In this article,we provide an overview of SMSI and explain how it differs from traditional intercropping.We also discuss the core principles that foster higher yields and the prospects for its future development.

Grain yield and N uptake of maize in response to increased plant density under reduced water and nitrogen supply conditions

The development of modern agriculture requires the reduction of water and chemical N fertilizer inputs.Increasing the planting density can maintain higher yields,but also consumes more of these restrictive resources.However,whether an increased maize density can compensate for the negative effects of reduced water and N supply on grain yield and N uptake in the arid irrigated areas remains unknown.This study is part of a long-term positioning trial that started in 2016.A split-split plot field experiment of maize was implemented in the arid irrigated area of northwestern China in 2020 to 2021.The treatments included two irrigation levels:local conventional irrigation reduced by 20%(W1,3,240 m^(3)ha^(-1))and local conventional irrigation(W2,4,050 m^(3)ha^(-1));two N application rates:local conventional N reduced by 25%(N1,270 kg ha^(-1))and local conventional N(360 kg ha^(-1));and three planting densities:local conventional density(D1,75,000 plants ha^(-1)),density increased by 30%(D2,97,500 plants ha-1),and density increased by 60%(D3,120,000 plants ha^(-1)).Our results showed that the grain yield and aboveground N accumulation of maize were lower under the reduced water and N inputs,but increasing the maize density by 30% can compensate for the reductions of grain yield and aboveground N accumulation caused by the reduced water and N supply.When water was reduced while the N application rate remained unchanged,increasing the planting density by 30% enhanced grain yield by 13.9% and aboveground N accumulation by 15.3%.Under reduced water and N inputs,increasing the maize density by 30% enhanced N uptake efficiency and N partial factor productivity,and it also compensated for the N harvest index and N metabolic related enzyme activities.Compared with W2N2D1,the N uptake efficiency and N partial factor productivity increased by 28.6 and 17.6%under W1N1D2.W1N2D2 had 8.4% higher N uptake efficiency and 13.9% higher N partial factor productivity than W2N2D1.W1N2D2 improved urease activity and nitrate reductase activity by 5.4% at the R2(blister)stage and 19.6% at the V6(6th leaf)stage,and increased net income and the benefit:cost ratio by 22.1 and 16.7%,respectively.W1N1D2 and W1N2D2 reduced the nitrate nitrogen and ammoniacal nitrogen contents at the R6 stage in the 40-100 cm soil layer,compared with W2N2D1.In summary,increasing the planting density by 30% can compensate for the loss of grain yield and aboveground N accumulation under reduced water and N inputs.Meanwhile,increasing the maize density by 30% improved grain yield and aboveground N accumulation when water was reduced by 20% while the N application rate remained constant in arid irrigation areas.

Manure substitution improves maize yield by promoting soil fertility and mediating the microbial community in lime concretion black soil

Synthetic nitrogen(N)fertilizer has made a great contribution to the improvement of soil fertility and productivity,but excessive application of synthetic N fertilizer may cause agroecosystem risks,such as soil acidification,groundwater contamination and biodiversity reduction.Meanwhile,organic substitution has received increasing attention for its ecologically and environmentally friendly and productivity benefits.However,the linkages between manure substitution,crop yield and the underlying microbial mechanisms remain uncertain.To bridge this gap,a three-year field experiment was conducted with five fertilization regimes:i)Control,non-fertilization;CF,conventional synthetic fertilizer application;CF_(1/2)M_(1/2),1/2 N input via synthetic fertilizer and 1/2 N input via manure;CF_(1/4)M_(3/4),1/4 N input synthetic fertilizer and 3/4 N input via manure;M,manure application.All fertilization treatments were designed to have equal N input.Our results showed that all manure substituted treatments achieved high soil fertility indexes(SFI)and productivities by increasing the soil organic carbon(SOC),total N(TN)and available phosphorus(AP)concentrations,and by altering the bacterial community diversity and composition compared with CF.SOC,AP,and the soil C:N ratio were mainly responsible for microbial community variations.The co-occurrence network revealed that SOC and AP had strong positive associations with Rhodospirillales and Burkholderiales,while TN and C:N ratio had positive and negative associations with Micromonosporaceae,respectively.These specific taxa are implicated in soil macroelement turnover.Random Forest analysis predicted that both biotic(bacterial composition and Micromonosporaceae)and abiotic(AP,SOC,SFI,and TN)factors had significant effects on crop yield.The present work strengthens our understanding of the effects of manure substitution on crop yield and provides theoretical support for optimizing fertilization strategies.

Identification, pathogenicity, and fungicide sensitivity of Eutiarosporella dactylidis associated with leaf blight on maize in China

Maize(Zea mays L.) is an economically vital grain crop that is cultivated worldwide. In 2011, a maize foliar disease was detected in Lingtai and Lintao counties in Gansu Province, China. The characteristic signs and symptoms of this disease include irregular chlorotic lesions on the tips and edges of infected leaves and black punctate fruiting bodies in dead leaf tissues. Given favourable environmental conditions, this disease spread to areas surrounding Gansu. In this study, infected leaves were collected from Gansu and Ningxia Hui Autonomous Region between 2018and 2020 to identify the disease-causing pathogen. Based on morphological features, pathogenicity tests, and multilocus phylogenetic analysis involving internal transcribed spacer(ITS), 18S small subunit rDNA(SSU), 28S large subunit rDNA(LSU), translation elongation factor 1-alpha(TEF), and β-tubulin(TUB) sequences, Eutiarosporella dactylidis was identified as the causative pathogen of this newly discovered leaf blight. Furthermore, an in vitro bioassay was conducted on representative strains using six fungicides, and both fludioxonil and carbendazim were found to significantly inhibit the mycelial growth of E. dactylidis. The results of this study provide a reference for the detection and management of Eutiarosporella leaf blight.

Dynamics and genetic regulation of macronutrient concentrations during grain development in maize

Nitrogen(N), phosphorus(P), and potassium(K) are essential macronutrients that are crucial not only for maize growth and development, but also for crop yield and quality. The genetic basis of macronutrient dynamics and accumulation during grain filling in maize remains largely unknown. In this study, we evaluated grain N, P, and K concentrations in 206 recombinant inbred lines generated from a cross of DH1M and T877 at six time points after pollination. We then calculated conditional phenotypic values at different time intervals to explore the dynamic characteristics of the N, P, and K concentrations. Abundant phenotypic variations were observed in the concentrations and net changes of these nutrients. Unconditional quantitative trait locus(QTL) mapping revealed 41 non-redundant QTLs, including 17, 16, and 14 for the N, P, and K concentrations, respectively. Conditional QTL mapping uncovered 39 non-redundant QTLs related to net changes in the N, P, and K concentrations. By combining QTL, gene expression, co-expression analysis, and comparative genomic data, we identified 44, 36, and 44 candidate genes for the N, P, and K concentrations, respectively, including GRMZM2G371058 encoding a Doftype zinc finger DNA-binding family protein, which was associated with the N concentration, and GRMZM2G113967encoding a CBL-interacting protein kinase, which was related to the K concentration. The results deepen our understanding of the genetic factors controlling N, P, and K accumulation during maize grain development and provide valuable genes for the genetic improvement of nutrient concentrations in maize.

Effects of drip and flood irrigation on carbon dioxide exchange and crop growth in the maize ecosystem in the Hetao Irrigation District,China

Drip irrigation and flood irrigation are major irrigation methods for maize crops in the Hetao Irrigation District,Inner Mongolia Autonomous Region,China.This research delves into the effects of these irrigation methods on carbon dioxide(CO_(2))exchange and crop growth in this region.The experimental site was divided into drip and flood irrigation zones.The irrigation schedules of this study aligned with the local commonly used irrigation schedule.We employed a developed chamber system to measure the diurnal CO_(2)exchange of maize plants during various growth stages under both drip and flood irrigation methods.From May to September in 2020 and 2021,two sets of repeated experiments were conducted.In each experiment,a total of nine measurements of CO_(2)exchange were performed to obtain carbon exchange data at different growth stages of maize crop.During each CO_(2)exchange measurement event,CO_(2)flux data were collected every two hours over a day-long period to capture the diurnal variations in CO_(2)exchange.During each CO_(2)exchange measurement event,the biological parameters(aboveground biomass and crop growth rate)of maize and environmental parameters(including air humidity,air temperature,precipitation,soil water content,and photosynthetically active radiation)were measured.The results indicated a V-shaped trend in net ecosystem CO_(2)exchange in daytime,reducing slowly at night,while the net assimilation rate(net primary productivity)exhibited a contrasting trend.Notably,compared with flood irrigation,drip irrigation demonstrated significantly higher average daily soil CO_(2)emission and greater average daily CO_(2)absorption by maize plants.Consequently,within the maize ecosystem,drip irrigation appeared more conducive to absorbing atmospheric CO_(2).Furthermore,drip irrigation demonstrated a faster crop growth rate and increased aboveground biomass compared with flood irrigation.A strong linear relationship existed between leaf area index and light utilization efficiency,irrespective of the irrigation method.Notably,drip irrigation displayed superior light use efficiency compared with flood irrigation.The final yield results corroborated these findings,indicating that drip irrigation yielded higher harvest index and overall yield than flood irrigation.The results of this study provide a basis for the selection of optimal irrigation methods commonly used in the Hetao Irrigation District.This research also serves as a reference for future irrigation studies that consider measurements of both carbon emissions and yield simultaneously.

Combining field data and modeling to better understand maize growth response to phosphorus(P) fertilizer application and soil P dynamics in calcareous soils

We used field experimental data to evaluate the ability of the agricultural production system model (APSIM) to simulate soil P availability,maize biomass and grain yield in response to P fertilizer applications on a fluvo-aquic soil in the North China Plain.Crop and soil data from a 2-year experiment with three P fertilizer application rates(0,75 and 300 kg P_(2)O_(5) ha^(–1)) were used to calibrate the model.Sensitivity analysis was carried out to investigate the influence of APSIM SoilP parameters on the simulated P availability in soil and maize growth.Crop and soil P parameters were then derived by matching or relating the simulation results to observed crop biomass,yield,P uptake and Olsen-P in soil.The re-parameterized model was further validated against 2 years of independent data at the same sites.The re-parameterized model enabled good simulation of the maize leaf area index (LAI),biomass,grain yield,P uptake,and grain P content in response to different levels of P additions against both the calibration and validation datasets.Our results showed that APSIM needs to be re-parameterized for simulation of maize LAI dynamics through modification of leaf size curve and a reduction in the rate of leaf senescence for modern staygreen maize cultivars in China.The P concentration limits (maximum and minimum P concentrations in organs)at different stages also need to be adjusted.Our results further showed a curvilinear relationship between the measured Olsen-P concentration and simulated labile P content,which could facilitate the initialization of APSIM P pools in the NCP with Olsen-P measurements in future studies.It remains difficult to parameterize the APSIM SoilP module due to the conceptual nature of the pools and simplified conceptualization of key P transformation processes.A fundamental understanding still needs to be developed for modelling and predicting the fate of applied P fertilizers in soils with contrasting physical and chemical characteristics.

Salicylic acid positively regulates maize defenses against lepidopteran insects

In response to insect attack,plants use intricate signaling pathways,including phytohormones,such as jasmonate(JA),ethylene(ET),and salicylic acid(SA),to activate defenses.Maize(Zea mays)is one of the most important staple food crops around the world.Previous studies have shown that the JA and ET signaling play important roles in maize defense against insects,but little is known about whether and how SA regulates maize resistance to insect herbivores.In this study,we ectopically expressed the NahG(salicylate hydroxylase)gene in maize plants(NahG maize)to block the accumulation of SA.It was found that compared with the wild-type(WT)maize,the NahG-maize exhibited decreased resistance to the generalist insects Spodoptera litura and Spodoptera frugiperda and the specialist Mythimna separata,and the compromised resistance in the NahG maize was associated with decreased levels of defensive metabolites benzoxazinoids(Bxs)and chlorogenic acid(CA).Quantification of simulated S.litura feedinginduced JA,JA-isoleucine conjugate(JA-Ile),and ET in the WT and NahG maize indicated that SA does not regulate JA or JA-Ile,but positively controls ET.We provide evidence suggesting that the SA pathway does not crosstalk with the JA or the ET signaling in regulating the accumulation of Bxs and CA.Transcriptome analysis revealed that the bHLH,ERF,and WRKY transcription factors might be involved in SAregulated defenses.This study uncovers a novel and important phytohormone pathway in maize defense against lepidopterous larvae.

The cytosolic isoform of triosephosphate isomerase,ZmTPI4,is required for kernel development and starch synthesis in maize(Zea mays L.)

Triosephosphate isomerase(TPI)is an enzyme that functions in plant energy production,accumulation,and conversion.To understand its function in maize,we characterized a maize TPI mutant,zmtpi4.In comparison to the wild type,zmtpi4 mutants showed altered ear development,reduced kernel weight and starch content,modified starch granule morphology,and altered amylose and amylopectin content.Protein,ATP,and pyruvate contents were reduced,indicating ZmTPI4 was involved in glycolysis.Although subcellular localization confirmed ZmTPI4 as a cytosolic rather than a plastid isoform of TPI,the zmtpi4 mutant showed reduced leaf size and chlorophyll content.Overexpression of ZmTPI4 in Arabidopsis led to enlarged leaves and increased seed weight,suggesting a positive regulatory role of ZmTPI4 in kernel weight and starch content.We conclude that ZmTPI4 functions in maize kernel development,starch synthesis,glycolysis,and photosynthesis.

ZmbZIP27 regulates nitrogen-mediated leaf angle by modulating lignin deposition in maize

In grain crops such as maize(Zea mays),leaf angle(LA)is a key agronomic trait affecting light interception and thus planting density and yield.Nitrogen(N)affects LA in plants,but we lack a good understanding of how N regulates LA.Here,we report that N deficiency enhanced lignin deposition in the ligular region of maize seedlings.In situ hybridization showed that the bZIP transcription factor gene ZmbZIP27 is mainly expressed in the phloem of maize vascular bundles.Under N-sufficient conditions,transgenic maize overexpressing ZmbZIP27 showed significantly smaller LA compared with wild type(WT).By contrast,zmbzip27_(ems)mutant showed larger LA under both N-deficient and N-sufficient conditions compared with WT.Overexpression of ZmbZIP27 enhanced lignin deposition in the ligular region of maize in the field.We further demonstrated that ZmbZIP27 could directly bind the promoters of the microRNA genes ZmMIR528a and ZmMIR528b and negatively regulate the expression levels of ZmmiR528.ZmmiR528 knockdown transgenic maize displayed erect architecture in the field by increasing lignin content in the ligular region of maize.Taken together,these results indicate that ZmbZIP27 regulates N-mediated LA size by regulating the expression of ZmmiR528 and modulating lignin deposition in maize.

Timing effect of high temperature exposure on the plasticity of internode and plant architecture in maize

The occurrence of high temperature(HT)in crop production is becoming more frequent and unpredictable with global warming,severely threatening food security.The state of an organ’s growth and development is largely determined by the temperature conditions it is exposed to over time.Maize is the main cereal crop,and its stem growth and plant architecture are closely related to lodging resistance,and especially sensitive to temperature.However,systematic research on the timing effect of HT on the sequentially developing internode and stem is currently lacking.To identify the timing effect of HT on the morphology and plasticity of the stem in maize,two hybrids(Zhengdan 958(ZD958),Xianyu 335(XY335))characterized by distinct morphological traits in the stem were exposed to a 7-day HT treatment from the V6 to V17 stages(Vn presents the vegetative stage with n leaves fully expanded)in 2019-2020.The results demonstrated that exposure to HT during V6-V12 accelerated the rapid elongation of stems.For instance,HT occurring at V7 and V12 specifically promoted the lengths and weights of the 3rd-5th and 9th-11th internodes,respectively.Meanwhile,HT slowed the growth of internodes adjacent to the promoted internodes.Interestingly,compared with control,the plant height was significantly increased soon after HT treatment,but the promotion effect became narrower at the subsequent flowering stage,demonstrating a self-adjusting mechanism in the maize plant in response to HT.Importantly,HT altered the plant architectures,including a rising of the ear position and increase in the ear position coefficient.XY335 exhibited greater sensitivity in stem development than ZD958 under HT treatment.These findings improve our systematic understanding of the plasticity of internode and plant architecture in response to the timing of HT exposure.

Application of Polygonum minus Extract in Enhancing Drought Tolerance in Maize by Regulating Osmotic and Antioxidant System

Drought stress is a major factor affecting plant growth and crop yield production.Plant extracts as natural biostimulants hold great potential to strengthen plants to overcome drought impacts.To explore the effect of Polygonum minus extract(PME)in enhancing drought tolerance in plants,a study was set up in a glasshouse environment using 10 different treatment combinations.PME foliar application were designed in CRD and effects were closely observed related to the growth,physiology,and antioxidant system changes in maize(Zea mays L.)under well-watered and drought conditions.The seaweed extract(SWE)was used as a comparison.Plants subjected to drought stress exhibited a significant reduction in fresh weight,dry weight,relative water content(RWC),and soluble sugar,but they stimulated the phenolic,flavonoid,proline,glutathione(GSH),malondialdehyde(MDA)and antioxidant enzyme(catalase,CAT;peroxidase,POD;superoxide dismutase,SOD)activities.Foliar application of PME improved fresh and dry weight(FW:33.1%~41.4%;DW:48.0%~43.1%),chlorophyll content(Chl b:87.9%~100.76%),soluble sugar(23.6%~49.3%),and soluble protein(48.6%~56.9%)as well as antioxidant enzyme activities(CAT and POD)compared to CK under drought conditions.while decreasing the level of MDA.Notably,the mitigating effect of PME application with high concentration was more effective than those of SWE.Our study reveals that PME could alleviate drought stress by regulating osmoprotectant content and antioxidant defense system and can be used as an economical and environmentally friendly biostimulants for promoting maize growth under drought stress.

Effects of drying methods on the drying kinetics and quality of maize

At present, maize is mostly dried by hot air, and the quality of maize after drying in this way is poor. So it is particularly important to explore the influence of new drying methods on the drying characteristics and quality of maize. Five drying methods, including hot air drying (HAD), vacuum drying (VD), infrared drying (IRD), variable temperature drying (VTD), and vacuum IR drying (VID), were used to analyze the drying rate (DR), moisture ratio (MR), effective moisture diffusion coefficient (Deff), hardness, nutrient composition, color, and microstructure of maize to investigate the effects of different drying methods on the drying kinetics and quality of maize kernels. The results showed that among the five drying methods, the Modified Page drying model could most reflect moisture changes. VTD was better than the other methods in terms of DR, cracking rate, hardness, and crude fat. The highest lysine content in maize was obtained using HAD. The protein content was higher in IRD (p<0.05). The dough characteristics were better in VID and VTD than in IRD. The IRD, VTD, and VID-treated maize had a better color appearance. Microstructure analysis showed that the starch granules of VID, IRD, and VD-treated maize were oval, but large gaps could be found between the granules. The granules were also densely stacked, with most of them relatively intact. Correlation and clustering analysis showed different degrees of correlation between the physicochemical indicators. The overall quality in VTD was the best, followed by VID, whereas HAD and VD showed poorer quality. In terms of economic value and product quality, VTD was the most suitable drying method for maize. This study can provide a theoretical basis for the application of maize drying in the processing industry.

Overexpression of the peroxidase gene ZmPRX1 increases maize seedling drought tolerance by promoting root development and lignification

Drought is a main abiotic stress factor hindering plant growth,development,and crop productivity.Therefore,it is crucial to understand the mechanisms by which plants cope with drought stress.Here,the function of the maize peroxidase gene ZmPRX1 in drought stress tolerance was investigated by measurement of its expression in response to drought treatment both in a ZmPRX1 overexpression line and a mutant line.The higher root lignin accumulation and seedling survival rate of the overexpression line than that of the wild type or mutant support a role for ZmPRX1 in maize drought tolerance by regulating root development and lignification.Additionally,yeast one-hybrid,Dule luciferase and ChIP-qPCR assays showed that ZmPRX1 is negatively regulated by a nuclear-localized ZmWRKY86 transcription factor.The gene could potentially be used for breeding of drought-tolerant cultivars.

Phenotypic Characterization and QTL/Gene Identification for Internode Number and Length Related Traits in Maize

Internode number and length are the foundation to constitute plant height, ear height and the above-ground spatial structure of maize plant. In this study, segregating populations were constructed between EHel with extremely low ear height and B73. Through the SNP-based genotyping and phenotypic characterization, 13 QTL distributed on the chromosomes (Chrs) of Chr1, Chr2, Chr5-Chr8 were detected for four traits of internode no. above ear (INa), average internode length above ear (ILaa), internode no. below ear (INb), and average internode length below ear (ILab). Phenotypic variation explained (PVE) by a single QTL ranged from 6.82% (qILab2-2) to 12.99% (qILaa5). Zm00001d016823 within the physical region of qILaa5, the major QTL for ILaa with the largest PVE was determined as the candidate through the genomic annotation and sequence alignment between EHel and B73. Product of Zm00001d016823 was annotated as a WEB family protein homogenous to At1g75720. qRT-PCR assay showed that Zm00001d016823 highly expressed within the tissue of internode, exhibiting statistically higher expression levels among internodes of IN4 to IN7 in EHel than those in B73 (P Zm00001d016823 might provide novel insight into molecular mechanism beyond phytohormones controlling internode development in maize.

Genetic analysis and candidate gene identification of salt tolerancerelated traits in maize

Soil salinization poses a threat to maize production worldwide,but the genetic mechanism of salt tolerance in maize is not well understood.Therefore,identifying the genetic components underlying salt tolerance in maize is of great importance.In the current study,a teosinte-maize BC2F7 population was used to investigate the genetic basis of 21 salt tolerance-related traits.In total,125 QTLs were detected using a high-density genetic bin map,with one to five QTLs explaining 6.05–32.02%of the phenotypic variation for each trait.The total phenotypic variation explained(PVE)by all detected QTLs ranged from 6.84 to 63.88%for each trait.Of all 125 QTLs,only three were major QTLs distributed in two genomic regions on chromosome 6,which were involved in three salt tolerance-related traits.In addition,10 pairs of epistatic QTLs with additive effects were detected for eight traits,explaining 0.9 to 4.44%of the phenotypic variation.Furthermore,18 QTL hotspots affecting 3–7 traits were identified.In one hotspot(L5),a gene cluster consisting of four genes(ZmNSA1,SAG6,ZmCLCg,and ZmHKT1;2)was found,suggesting the involvement of multiple pleiotropic genes.Finally,two important candidate genes,Zm00001d002090 and Zm00001d002391,were found to be associated with salt tolerance-related traits by a combination of linkage and marker-trait association analyses.Zm00001d002090 encodes a calcium-dependent lipid-binding(CaLB domain)family protein,which may function as a Ca^(2+)sensor for transmitting the salt stress signal downstream,while Zm00001d002391 encodes a ubiquitin-specific protease belonging to the C19-related subfamily.Our findings provide valuable insights into the genetic basis of salt tolerance-related traits in maize and a theoretical foundation for breeders to develop enhanced salt-tolerant maize varieties.

The transcription factor ZmNAC84 increases maize salt tolerance by regulating ZmCAT1 expression

Salt stress severely affects plant growth and yield.The transcription factor NAC plays a variety of important roles in plant abiotic stress,but we know relatively little about the specific molecular mechanisms of NAC in antioxidant defense.Here,our genetic studies reveal the positive regulation of salt tolerance in maize by the transcription factor ZmNAC84.Under salt stress,overexpression of ZmNAC84 in maize increased the expression of ZmCAT1,enhanced CAT activity,and consequently reduced H_(2)O_(2) accumulation,thereby improving salt stress tolerance in maize.Whereas RNA interference-mediated knockdown of ZmNAC84 produced the opposite effect.Subsequently,we found that ZmNAC84 directly binds to and regulates the expression of the ZmCAT1 promoter,and the hybridized material also demonstrated that ZmCAT1 is a downstream target gene of ZmNAC84.In addition,phenotypic and biochemical analyses indicated that ZmCAT1 positively regulated salt tolerance by regulating H_(2)O_(2) accumulation under salt stress.Taken together,these results reveal the function of ZmNAC84 in regulating ZmCAT1-mediated antioxidant defense in response to salt stress in plants.