Genome-Wide Association Study Elucidating Key Genetic Determinants of Stalk Water Content in Mature Maize (Zea mays L.)

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Meili Yang, Bangtai Wang, Qiyue Wang, Hua Guo, Guiwen Qin, Jianmei Cheng, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7282731/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Background The Stalk Moisture Content (SMC) is an important measure of stem fullness, disease resistance and lodging resistance. Results In this study, 236 inbred lines were used as the basic materials, and the water content of stalk in the third section was determined using the drying method and calibrated probe method for maize were investigated at the tasseling, grain filling, and maturity stages. 461,053 high-quality SNPs obtained through whole-genome resequencing were used for a genome-wide association study. As a result of mixed linear model (MLM, P < 9.94×10 − 6 ) analysis, 55 significant SNPs related to traits were detected, explaining 9.11%-15.13% phenotypic variation, among which 22 SNPs were found by the drying method (DMSMC) and 33 SNPs by the calibrated probe method(PMSMC). A total of 7 candidate genes were detected repeatedly, The Zm00001d014449 (detected four times) encodes a chromodomain protein involved in vascular bundle formation. The Zm00001d003476 (detected twice) encodes ubiquitin carboxyl-terminal hydrolase. The Zm00001d005351 gene was associated with the DMSMC2 phase and encodes the synthesis of aromatic acid dehydratase. The Zm00001d025134 gene was a chromosome domain protein, which was found to be related to drought resistance of maize. Conclusions This study helps to deepen the understanding of the genetic basis of maize stalk water content traits, and provides theoretical guidance for screening germplasm materials. Maize Stalk traits Lodging resistance Stalk water content Genome-wide association study Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1 INTRODUCTION Maize (Zea mays L.) is an important food and feed crop globally in the world, and playing a significant role in world food security [ 1 ] . Stalk is an important component of the maize plant, which is closely related to plant lodging, material transport, disease resistance stress and grain yield [ 2 – 4 ] . Stalk water content is an important index to measure stalk fullness, stalk maturity and grain dehydration rate, which is affected by the physiological effects of plants, climate and cultivation measures, and is complementary to other chemical components of stalks [ 5 – 8 ] . Stalk water content is directly related to the quality of the stalks, and it is an important criterion for the maturity of maize live stalks. The higher of the stalk water content, the stronger vitality of the stalks, and also the more resistant to external forces. In the maturity period of maize, lodging is easily caused due to the low water content of stalks [ 9 – 11 ] . Brandon evaluated the influence of water content on maize stalk and the tissue properties of peel, developed two compression test methods to obtain the elastic index model, and evaluated the influence of stalk water content on the tissue properties [ 12 ] . Water content was negatively correlated with the elastic modulus of peel. There are direct and indirect methods for evaluating the stalk water content, and the drying method is the most commonly used direct measurement method [ 13 ] . Indirect measurement methods mainly include the electronic moisture meter measurement method, near infrared reflectance spectroscopy analysis method and probe moisture meter method, which are characterized by simple and fast measurement [ 14 – 15 ] . Domestic and foreign scholars research on maize stalks mainly focuses on phenotypic traits such as stalk height, ear height, stalk node number, and stalk thickness [ 16 – 19 ] , while there is relatively little research on measurement traits such as stalk moisture content. In the early stage of maize growth, the stalk moisture content is less affected by density, but in the later stage of growth, it is more affected by density and decreases with increasing density [ 12 ] . Therefore, maintaining a certain stalk moisture content in the physiological later stage of maize can effectively enhance stem strength, increase plant lodging resistance, and meet the production needs of high-density maize planting. The trend of stalk moisture content changes varies among different varieties at different growth stages [ 5 ] . Different types of inbred lines also exhibit diversity in stalk moisture content, but it is relatively stable during the grain filling stage [ 20 ] . In the later stages of growth, as the plant naturally declines, it leads to a decrease in leaf, stem, and root vitality, resulting in a decrease in stalk moisture content [ 21 ] . The moisture content of maize male ear stem has a wide range of phenotypic variations and a high general heritability [ 22 – 23 ] . The hybrid advantage of maize stalk moisture content is mainly manifested by additive effects, with less environmental influence, and can be selected in early generations [ 6 ] . However, Zhou found that different genetic systems of stalk moisture content are significantly influenced by genotype and environmental interaction effects, and should be selected in later generations [ 24 ] . While the genetic basis of moisture content in the maize ear has been extensively studied, the genetics of stalk moisture content-particularly in the third above-ground internode-remains largely unexplored. This study aims to fill this knowledge gap through a genome-wide association study to identify candidate genes controlling this trait. In the present study, we used a wide diversity composed of 236 inbred lines. The water content of the third stem on the ground was determined using the direct drying method and the corrected probe moisture method. The genotype and stalk water content traits were analyzed in the late stage of two years of growth, and significant SNPs related to stalk water content traits were screened. Relevant candidate genes were identified, in order to provide a theoretical reference for cultivating breakthrough maize varieties with strong comprehensive stress resistance. 2 MATERIALS AND METHODS 2.1 Test materials and design 236 materials include 141 inbred lines of regular maize, 34 inbred lines of explosive maize, and 61 inbred lines of glutinous maize. The experimental design refers to Wang [ 25 ] . 2.2 Genotype and genome-wide association analysis A total of 236 samples were randomly selected from the study population. Genomic DNA was extracted using a magnetic bead-based method, with concentration, purity, and integrity subsequently assessed. Qualified DNA samples were resequenced to 5× coverage (BGI, Shenzhen, China; www.bgi.com ). Reads were aligned to the B73 reference genome (RefSeq v4.0) to detect SNPs. After filtering for variants with missing ratios 0.05, 461,053 high-quality SNPs were retained for genome-wide association analysis (Fig. 1 ). Associations between SNPs and stalk thrust resistance were tested at a significance threshold of P < 0.01. The mixed linear model (MLM) of TASSEL V5.0 software [ 26 ] was used to conduct correlation analysis on the stalk correlation at different growth stages. The significance threshold was set according to the method in the reference literature [ 25 , 27 ] , and the threshold for this study is P < 9.94×10 − 6 . 2.3 Phenotypic data measurement method and statistical method During the tasseling, filling, and ripening stages of maize, the moisture content of the third section of the stem on the ground was determined using direct drying method [ 14 ] and calibrated probe puncture method [ 7 ] (In 2021 and 2022, the average moisture content of drying stems at tasseling stage, filling stage and maturity stage were DMSMC1, DMSMC2 and DMSMC3, also by calibrated probe method were PMSMC1, PMSMC2 and PMSMC3). Select three representative plants with consistent growth conditions for each growth period and measure them three times, taking the average. The average of the three replicates represents the phenotype value of the trait at that growth period. Data were analyzed using Excel 2007 and DPS 7.05. Calculate group PCA and Kinship using TASSEL V5.0 software. Use the R "CMplot" software package to draw a correlation graph( https://github.com/YinLiLin/CMplot ). 2.4 Prediction of candidate genes Based on the sequence information of the B73 reference genome (B73-RefGenv4) in the reference database, candidate genes related to stalk water content were identified within a range of 2.5Kb upstream and downstream of significant loci. Annotate and predict the functions of candidate genes in the Uniport protein database( https://www.uniprot.org/uniprotkb/ ). 3 RESULTS AND DISCUSSION 3.1 Phenotypic analysis of moisture content traits in maize stalks The basic description statistics and frequency distribution of stalk moisture content traits in different growth stages of maize are shown in Table 1 and Fig. 2. The variation range of traits is large and the phenotypic diversity is rich. As the growth process progresses, the stalk moisture content gradually decreases. The skewness and kurtosis of each trait, except for the absolute value of PMSMC1 are all less than 1. Combined with the frequency histogram of trait distribution, they basically show a normal distribution trend, which is consistent with typical quantitative trait characteristics. The generalized heritability of stalk moisture content measured by the drying method is 67.63%, mainly influenced by genetic factors, while the generalized heritability of stalk moisture content measured by the calibrated probe method is 37.51%, which is greatly affected by environmental factors. Table 1 Statistical analysis of moisture content traits in maize stalk at different growth stages Trait Range Mean CV (%) Skewness Kurtosis Broad-sense heritability DMSMC1 76.09–88.32 82.81 0.03 -0.03 -0.15 67.63 DMSMC2 69.27–88.80 78.73 0.04 0.16 -0.09 DMSMC3 67.82–86.87 77.48 0.05 0.13 -0.03 PMSMC1 78.25–93.93 88.12 0.03 -0.7 1.11 37.51 PMSMC2 71.66–92.68 82.71 0.05 -0.48 0.11 PMSMC3 59.68–87.35 74.49 0.07 -0.09 -0.2 3.2 Variance analysis of stalk moisture content of the tested inbred lines The variance analysis of stalk moisture content traits in 236 inbred line materials (Table 2 ) showed that the genotype effects of the direct drying method and probe-based measurement of stalk moisture reached a highly significant level (P < 0.01), indicating significant genetic variation in stalk moisture content traits in the constructed natural population. Table 2 Variance analysis of various traits in different maize inbred lines Trait Variation Sum of squares DF SD F DMSMC Environment 3659.667 2 1829.833 580.853** Genotype 5380.766 235 22.8969 7.268** Error 1480.618 470 3.1503 PMSMC Environment 22224.16 2 11112.08 1118.25** Genotype 6539.6 235 27.8281 2.8** Error 4670.404 470 9.937 Note: * or * * indicate a significant correlation at the 0.05 or 0.01 level. 3.3 Genome - wide association study of stalk moisture content at different growth stages of maize The genome-wide association study was conducted on the stalk water content using MLM models of 236 maize inbred lines at different growth stages over two years by Tassel 5.0, and the Manhattan map and QQ map were drawn (Fig. 3a, 3b). The MLM model was used to analyze the correlation between stalk water content at different growth stages over two years. A total of 55 significantly associated SNPs loci were detected on chromosomes of maize (Table 3 ). We detected SNPs distributed across chromosomes 1–10 as follows: 4, 3, 3, 1, 33, 0, 2, 3, 3, and 3 respectively. A total of 22 SNPs (DMSMC1: 1, DMSMC2: 3, DMSMC3: 18) were detected related to DMSMC, as well as 33 SNPs (PMSMC1:14, PMSMC2: 1, PMSMC3: 18) related to PMSMC. The explained phenotypic variation ranged from 9.11–15.13%, with the highest phenotypic interpretation rate being DMSMC3 on chromosome 5_47800595 and the lowest being PMSMC1 on chromosome 4 _13346360. Table 3 Significant locus information on the correlation of maize stalk moisture content Trait Marker Chr Model Pos(bp) p Marker Rsq% DMSMC3 1_206350619 1 MLM 206350619 2.93E-06 10.33 PMSMC1 1_87740598 1 MLM 87740598 4.30E-06 11.42 PMSMC1 1_164402750 1 MLM 164402750 7.36E-06 13.53 PMSMC1 1_76763257 1 MLM 76763257 7.69E-06 11.91 PMSMC1 2_45422750 2 MLM 45422750 1.24E-06 12.34 PMSMC1 2_45418447 2 MLM 45418447 2.06E-06 12.47 DMSMC2 2_170637158 2 MLM 170637158 2.49E-06 9.92 DMSMC3 3_204520247 3 MLM 204520247 1.77E-06 11.95 PMSMC3 3_204520247 3 MLM 204520247 7.42E-06 10.54 PMSMC3 3_122700273 3 MLM 122700273 8.31E-06 10.68 PMSMC1 4_13346360 4 MLM 13346360 9.94E-06 9.11 DMSMC3 5_47800595 5 MLM 47800595 4.40E-08 15.13 PMSMC3 5_47800595 5 MLM 47800595 8.16E-08 14.62 DMSMC3 5_47752006 5 MLM 47752006 3.76E-07 11.84 DMSMC3 5_47727153 5 MLM 47727153 3.92E-07 12.31 PMSMC3 5_47752006 5 MLM 47752006 5.72E-07 11.57 DMSMC3 5_47876150 5 MLM 47876150 5.80E-07 11.96 DMSMC3 5_47645572 5 MLM 47645572 6.35E-07 11.7 PMSMC3 5_47727153 5 MLM 47727153 8.69E-07 11.56 DMSMC3 5_47711262 5 MLM 47711262 9.01E-07 12.63 PMSMC3 5_47645572 5 MLM 47645572 1.02E-06 11.45 DMSMC3 5_47750439 5 MLM 47750439 1.04E-06 12.56 DMSMC3 5_47726081 5 MLM 47726081 1.09E-06 12.51 PMSMC3 5_47711262 5 MLM 47711262 1.17E-06 12.53 PMSMC3 5_47876150 5 MLM 47876150 1.17E-06 11.37 PMSMC3 5_47726081 5 MLM 47726081 1.45E-06 12.35 DMSMC3 5_47925170 5 MLM 47925170 1.53E-06 12.21 PMSMC3 5_47750439 5 MLM 47750439 1.78E-06 12.19 PMSMC3 5_47925170 5 MLM 47925170 1.95E-06 12.05 DMSMC3 5_47735500 5 MLM 47735500 2.54E-06 12.95 DMSMC3 5_47747099 5 MLM 47747099 2.67E-06 11.79 DMSMC3 5_47716318 5 MLM 47716318 2.87E-06 11.52 DMSMC3 5_47880130 5 MLM 47880130 3.11E-06 11.71 PMSMC3 5_47880130 5 MLM 47880130 3.39E-06 11.77 PMSMC3 5_47747099 5 MLM 47747099 3.55E-06 11.59 DMSMC3 5_47744387 5 MLM 47744387 3.85E-06 11.37 PMSMC3 5_47735500 5 MLM 47735500 3.93E-06 12.72 DMSMC2 5_47800595 5 MLM 47800595 4.16E-06 10.77 PMSMC3 5_47716318 5 MLM 47716318 4.21E-06 11.31 PMSMC3 5_47744387 5 MLM 47744387 4.98E-06 11.24 PMSMC1 5_46831646 5 MLM 46831646 5.74E-06 12.88 DMSMC3 5_47718537 5 MLM 47718537 5.76E-06 11.12 PMSMC3 5_47718537 5 MLM 47718537 7.41E-06 10.81 PMSMC1 5_100064005 5 MLM 100064005 7.46E-06 10.7 PMSMC1 7_96409678 7 MLM 96409678 8.02E-07 14.4 PMSMC2 7_114436113 7 MLM 114436113 9.65E-06 11.97 PMSMC1 8_67202484 8 MLM 67202484 6.87E-07 15.02 DMSMC3 8_159862794 8 MLM 159862794 8.42E-06 11.34 PMSMC3 8_159862794 8 MLM 159862794 9.40E-06 11.2 PMSMC1 9_84316697 9 MLM 84316697 1.77E-06 12.37 PMSMC1 9_18935130 9 MLM 18935130 1.92E-06 12.25 PMSMC1 9_70247735 9 MLM 70247735 2.45E-06 14.38 PMSMC1 10_33621054 10 MLM 33621054 2.79E-06 11.63 DMSMC1 10_121739764 10 MLM 121739764 5.32E-06 11.28 DMSMC2 10_105984280 10 MLM 105984280 5.61E-06 11.07 Further analysis revealed that 17 SNPs related to stalk water content traits were detected more than twice by independent association analysis using the MLM models (Table 4 ), suggesting that these loci carry important genes that can stably regulate stalk water content traits in different environments. Among them, 4 SNPs were detected more than twice by independent association analyses at different growth stages with the same stalk moisture content traits. It is speculated that these loci may carry important genes in the growth process of maize stalk moisture content, which are continuously expressed and play a role in multiple growth stages. Different methods for measuring stalk moisture content have identified 16 SNPs detected more than twice by independent association analyses, suggesting that the candidate genes carried by these loci are more likely to be stably expressed in maize stalk moisture traits. These SNPs that can be significantly correlated with stalk-related traits multiple times have high confidence (HC) Among them, HC-SNP locus 5_47800595 was found to be significantly correlated with DMSMC twice and PMSMC3 once in two growth stages. It was also found that 31 SNPs significantly correlated with DMSMC and PMSMC were clustered in the 5_4764557, 2_47925170 range at different growth stages, which is an enriched region of stalk water content-related genes. Significant correlations were found twice with DMSMC and PMSMC at adjacent sites of 2_245418447, 3_204520247, and 8_159862794 respectively. 3.4 Analysis of genes associated with stalk moisture content traits Based on the association of two measurement methods, candidate genes were screened and predicted within a range of 2.5kb upstream and downstream of 55 significant SNP loci related to stalk water content traits. Combined with gene functional annotation, 14 candidate genes were identified (Table 4 ), and 7 candidate genes were detected more than twice. These genes may be highly reliable associated genes for stalk water content traits. Gene ontology (GO)enrichment analysis using the 14 candidate genes, most of the candidate genes discovered are related to Ubiquitin system such as aromatic acid dehydratase, ubiquitin carboxy terminal dehydratase, and threonine phosphatase (Fig. 4). At the same time, this study also found three unknown functional genes, namely LOC103636628 , Zm00001d049035 , and Zm00001d014448 . 3.4 Identification and functional analysis of high-confidence associated genes Using two detection methods, a total of 7 candidate genes were detected more than twice. Among them, Zm00001d014449 was found to be associated with DMSMC and PMSMC 4 times, with a phenotypic variation explanation rate of 11.24%-12.56%. This gene originates from the assembly of CRD101 and is associated with the LHP1 gene. Which is mainly expressed in cells with active cell division and tissue differentiation, such as vascular bundle synthesis, which is of great significance for maintaining cell and tissue structure specificity. Zm00001d014448 was found to be associated with DMSMC and PMSMC 3 times, with a phenotypic variation explanation rate of 10.81%-12.63%. The function of this gene is unknown. Zm00001d003476 was found to be associated with DMSMC and PMSMC 2 times, with a phenotypic variation explanation rate of 12.34%-12.47%, encoding the synthesis of ubiquitin carboxylic acid terminal hydrolase; Zm00001d043592 and Zm00001d043593 were found to be associated with DMSMC and PMSMC 2 times, with a phenotypic variation explanation rate of 10.54%-11.95%, They encode cellulose synthase A4 and participate in the SPA1 expression pathway. Zm00001d014451 was found to be associated with DMSMC and PMSMC traits twice, with a phenotypic variation explanation rate of 12.05%-12.25%, which is involved in glutamic acid receptor synthesis. 3.5 Analysis of haplotype water content traits By analyzing the re-sequencing data and evaluating the phenotypic performance of the population, the association between the above five SNPs and the water content traits of maize stalk was verified. At 2_45418447 locus, the GG genotype was associated with high stem water content of PMSMC1, while the AA genotype was associated with low content, while at 2_45422750 locus, the AA genotype was associated with high stem water content of PMSMC1, and the GG genotype was associated with low content. At the 2_170637158 locus, the TT genotype was associated with high stem water content of DMSMC2, while the CC genotype was associated with low stem water content; At the 5_47750439 locus, the GG genotype was associated with high stem water content of DMSMC3, the AA genotype was associated with low stem water content, and there was no significant difference between the GG genotype and the PMSMC 3 stem water content (Fig. 5). Table 4 Prediction information of the candidate gene Trait Marker Chr Pos p Marker Rsq % Gene accession number Gene annotation PMSMC1 1_76763257 1 76763257 7.69E-06 11.91 LOC103636628 NO PMSMC1 1_87740598 1 87740598 4.30E-06 11.42 Zm00001d029806 NEP1-interacting protein-like 1 DMSMC2 2_170637158 2 170637158 2.49E-06 9.92 Zm00001d005351 Arogenate dehydratase, 4.2.1.91 PMSMC1 2_45418447 2 45418447 2.06E-06 12.47 Zm00001d003476 Ubiquitin carboxyl-terminal hydrolase PMSMC1 2_45422750 2 45422750 1.24E-06 12.34 DMSMC3 3_204520247 3 204520247 1.77E-06 11.95 Zm00001d043592, Zm00001d043593 SPA1-related 3, CESA4 PMSMC3 3_204520247 3 204520247 7.42E-06 10.54 PMSMC1 4_13346360 4 13346360 9.94E-06 9.11 Zm00001d049035 NO DMSMC3 5_47711262 5 47711262 9.01E-07 12.63 Zm00001d014448 NO PMSMC3 5_47711262 5 47711262 1.17E-06 12.53 DMSMC3 5_47716318 5 47716318 2.87E-06 11.52 PMSMC3 5_47716318 5 47716318 4.21E-06 11.31 DMSMC3 5_47718537 5 47718537 5.76E-06 11.12 PMSMC3 5_47718537 5 47718537 7.41E-06 10.81 DMSMC3 5_47744387 5 47744387 3.85E-06 11.37 Zm00001d014449, crd101 Chromo domain-containing protein LHP1, Chrom domain-containing protein CRD101 PMSMC3 5_47744387 5 47744387 4.98E-06 11.24 DMSMC3 5_47747099 5 47747099 2.67E-06 11.79 PMSMC3 5_47747099 5 47747099 3.55E-06 11.59 DMSMC3 5_47750439 5 47750439 1.04E-06 12.56 PMSMC3 5_47750439 5 47750439 1.78E-06 12.19 DMSMC3 5_47752006 5 47752006 3.76E-07 11.84 PMSMC3 5_47752006 5 47752006 5.72E-07 11.57 DMSMC3 5_47925170 5 47925170 1.53E-06 12.21 Zm00001d014451 Glutamate receptor PMSMC3 5_47925170 5 47925170 1.95E-06 12.05 PMSMC1 9_18935130 9 18935130 1.92E-06 12.25 Zm00001d045322, Zm00001d045323 Ubiquinone biosynthesis O-methyltransferase, Double B-box zinc finger protein 11 DMSMC2 10_105984280 10 105984280 5.61E-06 11.07 Zm00001d025134 FPL domain-containing protein 4. Discussion 4.1 Comparative analysis of significant SNPs loci in stalk moisture content traits By comparing and analyzing the genetic mapping results of stalk moisture content traits at home and abroad, it was found that among the many previous studies on moisture content, there were relatively few studies on the localization of the third stalk moisture content in maize fields, only related studies on the localization of male ear stem, cob, bract leaf, and grain moisture content. Shang studied the stem traits of 70 major maize inbred lines and found that the stalk moisture content of maize was relatively stable during the grain filling stage [ 20 ] . Only 4 SNPs were found to be significantly associated with stalk moisture content during the grain filling stage, while 36 SNPs were associated during the maturity stage. Li conducted a GWAS analysis on the moisture content of maize male ear stems, and found that the significant locus on 10_121259591 was located at a distance similar to the significant locus 10_121739764 located during the tasseling stage stalk moisture content in this study [ 22 ] . The water content trait of bract leaves was located at significant loci 1_219374182 and 3_206692486, which are similar in distance to the significant loci 1-206350619 and 3_204520247 for mature stalk water content identified in this study; The water content trait of the spike axis was located at significant loci 2_170297547 and 2_45753838, which are close to the significant loci of stalk water content in the milk ripening stage and in the tasseling stage, which were located at positions 2_170637158 and 2_45418447 respectively. The significant SNPs interval of stalk moisture content trait identified in this study overlaps significantly with the correlation traits studied by previous researchers, indicating high reliability of the localization results and suitability for further exploration of trait-associated genes. In addition, the new significant SNP loci detected in this experiment, which are different from previous studies, especially some highly reliable significant SNP loci and highly efficient significant SNPs with a phenotype explanatory rate of over 15%, and also that great significance for exploring new significant loci of stalk water content traits and subsequently discovering new associated genes. 4.2 Analysis of Related Genes in Maize Stalks Studies demonstrate that post-physiological maturity, natural plant aging causes decrease in stem dry matter and water content, which is the main reason for the decrease in stem puncture strength and the increase in stem folding rate. In the early stage of research on the lodging resistance related traits of maize stems [ 25 – 27 ] , similar significant SNPs and candidate gene information were also discovered in multiple identical intervals, especially the candidate gene Zm00001d014448 and Zm00001d014449 , which were clustered near 5_47747099. Zm00001d014449 is related to the LHP1 gene and participates in vascular bundle synthesis. It is located in the above-ground part of maize and encodes heterochromatin protein 1. It is mainly expressed in cells or tissues with active cell division and tissue differentiation, and is important for maintaining the specificity of cell and tissue structure [ 28 ] . The Zm00001d005351 gene associated with 2_170637158 expresses the product of aromatic acid dehydratase, which is a key enzyme catalyzing the conversion of aromatic acids to phenylalanine in the chloroplasts and plastid matrix of vascular plants. It plays an important role in the biosynthesis of cell wall lignin and photosynthesis, and can be used for improving the dehydration performance of plant stems. Expression in Arabidopsis can reduce lignin content, resulting in a decrease in stem weight and length [ 29 ] . At low nitrogen levels, it is associated with maize leaf type-related traits, and downregulation of expression directly affects the synthesis of tyrosine and alanine [ 30 ] . The expression product of the Zm00001d003476 gene, also associated with the attachment of 2_45418447, is a ubiquitin carboxy terminal hydrolase, this enzyme is an extremely important factor in cell biology that degrades proteins and can coordinate and regulate various biological processes and pathways to maintain cell health. The gene Zm00001d043593 associated with the attachment of 3_204520247 expresses cellulose synthase A4, which has been validated for expression in plants such as tobacco and plays an important role in plant stem cellulose synthesis [ 31 ] . In addition, protein differential expression occurs in the stems of maize under drought stress. Zm00001d029806 regulates the absorption of lead in maize seedling roots [ 32 ] , affects the long-distance transport of nitrogen elements in maize [ 33 ] , and is related to the development of ear length and row grain number [ 34 ] . The expression level of Zm00001d014451 gene decreased significantly after high-temperature treatment [ 35 ] . Zm00001d045322 is expressed and enriched in low-temperature-tolerant maize inbred lines, which may be related to ubiquitin biosynthesis [ 36 ] . The Zm00001d045323 gene is within the range of active recombinant genes in the infiltrated population of large grass [ 37 ] . The Zm00001d025134 gene is associated with maize leaf length traits at low nitrogen levels [ 30 ] . The Zm00001d025134 gene is associated with drought resistance in maize, and ZmVIL2 may inhibit its expression by binding to the promoter, thereby enhancing the drought resistance of maize plants [ 38 ] . 4.3 Correlation analysis of moisture content traits in maize stalks The stem is the bone of maize, mainly responsible for the transportation and supply of water and nutrients. Studies have shown that stem thickness, stem strength, stem tissue water absorption capacity, and transport capacity can all be used as indicators of maize drought resistance [ 39 ] . Anderson's research suggests that the moisture content of maize stems gradually decreases after tasseling, and there is a significant positive correlation between the moisture content of the stem pulp and stem strength. Stem strength decreases with the decrease in moisture content. At the same time, the plant's ability to resist stem rot is related to stalk moisture content and soluble sugar content. Plants with high moisture content in the stem pulp after physiological maturity of maize have a lower incidence of stem rot [ 40 ] . Djordjevic's believe that the moisture content of mature maize stems is significantly negatively correlated with lodging rate [ 41 ] . The impurity rate of maize kernels harvested is significantly positively correlated with the wet weight of maize stems, and significantly negatively correlated with the moisture content of stems [ 42 ] . The stem moisture content is significantly or extremely significantly positively correlated with stem height, number of stem nodes, ear height, and ear nodes, indicating that the height of the plant and ear affect the stalk moisture content. The stalk moisture content of maize inbred lines bred in China in the 1980s and 1990s decreased by 3 percentage points, which may be related to the breeding goal of focusing more on the selection of lodging resistance traits with lower plant height and ear height [ 43 ] . There is a significant positive correlation between the crushing strength of stems and their moisture content and fresh density. Increasing the fresh density and moisture content of stems can improve the mechanical characteristics of maize stems during flowering, thereby enhancing their resistance to stem lodging [ 44 ] . The thickness of the stem bark and the moisture content of the stem internode directly determine the strength of the stem puncture and transverse bending, and are also important criteria for judging the mechanical properties of the stem. In addition, spraying chemical regulators such as phosphatidylcholine can significantly enhance the moisture content of the maize stem in the later stage of maturity, thereby enhancing its ability to resist lodging [ 9 ] . During germplasm improvement for stem lodging resistance and variety breeding, selecting varieties with slow consumption rates of water and soluble sugar content under the ear in the late stage of physiological maturity, or delaying leaf senescence after physiological maturity through cultivation and breeding methods to increase stem soluble sugar content, can effectively reduce the risk of stem lodging and increase stem carbohydrate accumulation [ 45 – 46 ] . The 236 inbred lines used in this study included conventional maize inbred lines, sweet glutinous maize inbred lines, and burst maize inbred lines, with a wide variety of types. The selected germplasm materials with slow stalk moisture reduction in the late maturity stage can be used to improve the stem lodging resistance of core germplasm materials. In addition, the candidate genes associated with the study are undergoing functional validation and the development of stalk water content-related functional markers can further promote the improvement and creation of germplasm materials with high stalk water content in the later stage of physiological maturity. Declarations Ethics approval and consent to participate No ethics approval was required. The authors declare that the experimental methods conducted in this study complied with current Chinese laws and regulations. Availability of data and materials The datasets generated during the current study are available in the [GVM000940] repository, [https://bigd.big.ac.cn/gvm/getProjectDetail?Project=GVM000940]. Competing interests The authors declare no competing interests. Consent for publication Not applicable. Conflict of interest The authors declare that they have no conflicts of interest. Funding This work was supported by Henan Provincial Key Science and Technology Project Creation of Germplasm Resources and New Variety Breeding and Application of High-Yield and Multi-Resistance Maize (2022010202), the open-ended foundation of National Key Laboratory of Wheat and Maize Crop Science, Henan Agricultural University (39990109), Henan Provincial Science and Technology Project Analysis of Genetic Basis and Application of Stalk Quality Traits by GWAs (202102110248). Authors’ contributions † Meili Yang, Bangtai Wang contributed equally to this work. * Corresponding author: Hongwei Lu( [email protected] ), Jiafa Chen( [email protected] ) MLY, BTW, HWL and JFC designed this study. QYW, JMC developed the populations. ZHW recorded the data. BTW analyzed the data. BTW and MLY drafted the manuscript. HWL, JFS, XHZ and JFC revised the manuscript. All authors read and approved the final manuscript. Acknowledgements We thank professor Chunhua Mu of Shandong Academy of Agricultural Sciences for providing the maize population. Author details 1Hebi Academy of Agricultural Sciences, Hebi 458031, Henan, China; 2 Henan Agricultural University College of Life Sciences, Zhengzhou 450002, Henan, China; 3Henan Maize Breeding Engineering Technology Research Center, Hebi 458031, Henan, China; 4Hebi Maize Breeding Key Laboratory, Hebi 458031, Henan, China; 5National Key Laboratory of Wheat and Maize Crop Science; Collaborative Innovation Center of Henan Grain Crops; College of Agronomy; Henan Agricultural University, Zhengzhou 450002, Henan, China. References Chou H G, Zhang S H, Yang J, Jing Y. Development of China's maize industry, challenges in the future and policy suggestions. Journal of Agricultural Science and Technology, 2013(1):20-24. Zhou Y, Gu W R, Zhao M, Tong T, Liu X M, Li C F, Li J, Wei T. Differences of stalk characteristics and grain mechanically harvesting qualities of different maturing type spring maize in Heilongjiang Province. Acta Agriculturae Boreali-Sinica, 2017, 32(S1): 140-146. Feng G, Jing X Q, Li Y Y, Wang L, Huang C L. Correlation and path analysis of lodging resistance with maize stem characters. Acta Agriculturae Boreali-Sinica, 2010, 25(S1): 72-74. Yang Jianping, Li Meng, Yin Yue, Liu Yan, Gan Xinke, Mu Xiaohuan, Li Hanqin, Li Jiankun, Li Haochuan, Zheng Jun, Gou Mingyue. Spatial accumulation of lignin monomers and cellulose underlying stalk strength in maize, Plant Physiology and Biochemistry, 2024, 214:108918. Jin Yingjie. Effects of Stalk Mechanical Characteristics and Chemical Components on Lodging Resistance of Different Maize Cultivars. Master's Thesis of Henan Agricultural University, 2018. (In Chinese) Zhao Yanming, Xiao Jun, Zhang Baoshi, Wang Changtao, Sha Wancun. Genetic Studies on Moisture Content in Maize-stalk. Liaoning Agricultural Sciences, 2002(3): 19-20. Wang Bangtai, Yang Meili, Wang Jing, Wang Zhihong, Guo Hua, Hou Xianjun, Zhang Wenbo,Chang Jianzhi. Correlation Analysis and Rapid Determination of Grain Dehydration Rate of Maize Varieties. Chinese Agricultural Science Bulletin, 2020, 36(15): 18-23. Xiao Yingjie, Liu Haijun, Wu Liuji, Warburton Marilyn, Yan Jianbing. Genome-wide Association Studies in Maize: Praise and Stargaze. 2017, 10(3): 359-374. Pei Zhichao. Effects of Chemistry Regulation on Straw Quality of High-yield Maize Master's Thesis of Chinese Academy of Agricultural Science Dissertation, 2010. (In Chinese) Xue Jun, Ming Bo, Xie Ruizhi, Wang Keru, Hou Peng, Li Shaokun. Evaluation of maize lodging resistance based on the critical wind speed of stalk breaking during the late growth stage, Plant Methods, 2020, 16:148-156. Liu Shenqun, Guo Jinrui, Zhang Weijian, Ren Jun, Song Zhenwei, Yan Xiaogong, Song Fengbin, Zhu Xianchan. Effect of Planting Density on Water Content of Stalk and Root in Spring Maize. Soil and Crop, 2014, 3(3): 93-98. Brandon Sutherland, Kirsten Steele, Joseph Carter, Douglas D. Cook. The influence of water content on the longitudinal modulus of elasticity of maize stalk pith and rind tissues. BMC Plant Methods, 2023, 19(64): 2-11. Xiang Kui. Genetic analysis and measuring method development of kernel fast dry down rate in maize. Doctor's Thesis of Sichuan Agricultural University, 2011. (In Chinese) Guo Hua, Wang Jing, Yang Meili, Wang Zhihong, Wang Bangtai. Building and Utilization of One Quick Water-content Measuring Method in Kernel of Maize Based on Regression Model. Journal of Maize Sciences, 2018, 26(3): 63-68. Wang Guorui, Yuan Zhen, Zhang Pengyu, Liu Zhixue, Wang Tongchao, Wei Li. Genome-wide analysis of NAC transcription factor family in maize under drought stress and rewatering. PHYSIOLOGY AND MOLECULAR BIOLOGY OF PLANTS, 2020, 26(4): 705-717. Ma Yajie, Bao Jianxi, Gao Yuexin, Li Yanan, Qin Wenxuan, Wang Yanbo, Long Yan, Li Jinping, Dong Zhenying, Wan Xiangyuan, Genome-wide association analysis of plant height and ear height related traits in maize. ACTA AGRONOMICA SINICA, 2023, 49(3): 647-661. Sun Qiang, Ren Jiaojiao, Xu Xiaoming, Li Zongze, Huang Bowen, Chen Zhanhui, Wu Penghao. QTL Mapping and Genomic Selection for Plant Height and Ear Height in Maize. Journal of Maize Sciences, 2022, 30(4): 40-47. Wang Xintao, Wei Feng, Dai Ziju, Li Baoye, Hao Junjie. Identification of Candidate Genes Associating with Stem Diameter in Maize (Zea mays L.) Based on Integrating QTL Mapping and RNA-seq. Journal of Plant Genetic Resources, 2022, 23(6): 1737-1745. Wang Jingjing, Li Rongfa, Liu Dongyao, Zhao Wenlong, Miao Zhengyan, Jia Xucun, Dong Pengfei, Wang Qun. Endogenous Hormones Improve Lodging Tolerance of Maize (Zea mays L.) by Regulating Stalk Structure Under Elevated Temperature. JOURNAL OF PLANT GROWTH REGULATION, 2024, 43(2):445-457. Shang Qiqi, Zhang Degui, Wang Kaixin, Wang Guoliang, Pan Jinbao, Li Xinhai, Shi Liyu. Diversity Analysis of Stalk Traits in Maize Inbred Lines in China. Journal of Plant Genetic Resources, 2020, 21(2): 321-329. Xue Jun, Wang Keru, Xie Ruizhi, Gou Ling, Zhang Wangfeng, Ming Bo, Hou Peng, Li Shaokun. Research Progress of Maize Lodging During Late Stage. Scientia Agricultura Sinica, 2018, 51(10): 1845-1854. Li Zhen. Rapid Determination Method and Genome-Wide Association Study of Maize Kernel Moisture Content in Mature Period. Master's Thesis of Hebei Agricultural University, 2019. (In Chinese) Sun Gaoyang, Zhang Xuehai, Duan Haiyang, Gao Jionghao, Li Na, Su Pingping, Xie Huiling, Li, Weihua, Fu Zhiyuan, Huang Yubi, Tang Jihua. Dissection of the genetic architecture of peduncle vascular bundle-related traits in maize by a genome-wide association study. PLANT BIOTECHNOLOGY JOURNAL, 2022, 20(6):1042-1053. Zhou Dongsheng, Zhao Yanming, Hou Hengjun, Li Guangxian. Study on Genetic Main Effects and Genotype Environment Interaction Effects for Moisture Content of Different Parts in Maize. Journal of Maize Sciences, 2011, 19(3): 35-38. Wang Bangtai, Yang Meili, Guo Hua, Wang Jing, Wang Zhihong, Lu Hongwei, Qin Guiwen, Chen Jiafa. Genome-wide association study for stalk lodging resistance related traits in maize (Zea mays L.). BMC Genomics, 2024, 25:19-32. -5. Bradbury PJ, Zhang Z, Kroon DE, Casstevens TM, Ramdoss Y, Buckler ES, Bishop PM. TASSEL: software for association mapping of complex traits in diverse samples. Bioinformatics, 2007,23(19): 2633 Wang M, Yan J, Zhao J, Song W, Zhang X, Xiao Y, Zheng Y. Genome-wide association study (GWAS) of resistance to head smut in maize. Plant Sci, 2012, 196: 125-131. Lv Yuhua. The Evolution Analysis of Plant LHP1 and Expression patterns of LHP1 from three species of Poaceae. Master's Thesis of Guangxi University, 2011. (In Chinese) Oliver R A Corea, CL Cardenas, SE Brewer, AM Patten, LB Davin, NG Lewis. Arogenate dehydratase isoenzymes profoundly and differentially modulate carbon flux into lignins. Journal of Biological Chemistry. 2012, 287(14): 11446-11459. He Bing. QTL Mapping for Controlling Maize leaf type Related traits under different Nitrogen Treatment Conditions. Master's Thesis of Guizhou University, 2021. (In Chinese) Guan Lufan. Genetic Transformation of Cellulose Synthase gene BpCesA4 from Betula Platyphylla. Master's Thesis of Northeast Forestry University, 2009. (In Chinese) Hou Fengxia. Molecular network analysis of ZmbZIP54 and ZmbZIP107 transcription factors regulating lead tolerance in maize seedlings. Doctor’s Thesis of Sichuan Agricultural University, 2023. (In Chinese) Xu Jiahui. Elucidation of the responsive mechanisms of maize roots to long distance nitrogen signals using a split root system. Master's Thesis of Sichuan Agricultural University, 2020. (In Chinese) Ning Qiang. Cloning and Functional analysis of a major QTL qEl7 for Ear length and Kernel Number per row in Maize. Doctor’s Thesis of Huazhong Agricultural University, 2021. (In Chinese) Zhou Shenghui, Zhang Lei, Lv Xinze, Huang Jinguang. Ning Qiang. Identification and Analysis of GLR Family Genes in Maize. Journal of Maize Sciences, 2021. 29(2): 35-42. Meng Aiju. Genome-wide Association Studies and Functional Studies of Candidate Genes for Germination-related Traits of Low Temperature Tolerance in Maize. Doctor’s Thesis of Shandong Agricultural University, 2022. (In Chinese) Fan Feifei. Genetic Analysis of Genomic Expression Variation in An Introgressive Population of Teosinate. Master's Thesis of Huazhong Agricultural University, 2019. (In Chinese) Zhang Dongling. Functional analysis of maize gene ZmVIL2 in response to drought stress. Doctor's Thesis of Henan Agricultural University, 2024. (In Chinese) Tan Guobo, Zhao Liqun, Zhang Lihua, Zhao Hongxiang, Fang Xiangqian, Meng Xiangmeng, Bian Shaofeng, Effects of Different Water Stresses on Plant Character, Photosynthetic Physiology and Yield in Maize Jointing Stage, Journal of Maize Science, 2010, (1): 96-98. Anderson B. Evaluation of Methods for Identification of Corn Genotypes with Stalk Rot and Lodging Resistance. Plant Disease, 1994, 78(6): 590. Djordjevic Jovan S, Ivanovic Mile R. Genetic Analysis for Stalk Lodging Resistance in Narrow-Base Maize Synthetic Population ZPS14. Crop Science, 1996, 36(4): 909-913. Zhu Xueqing. Selection and evaluation of maize varieties suitable for machine harvesting. Master's Thesis of Hunan Agricultural University, 2021. (In Chinese) Wang Kaixin, Cheng Zimeng, Yang Yihan, Zhang Degui, Li Xinhai, Shi Liyu. Evolvement of Stem-related Traits of Maize Inbred Lines Through Different Decades in China. Journal of Plant Genetic Resources, 2021, 22(1): 157-164. Liu Shengqun, Liu Tiedong, Song Fengbin, Wang Yang, Qi Xiaoning, Zhu Xianchan. Effects of Row Orientation and Planting Pattern on Traits Associated with Stem Lodging in Maize. Soils and Crops, 2016, 5(3): 159-165. Chen Xiang. The change of stalk lodging resistance characteristics before and after physiological maturity of maize. Master's Thesis of Sichuan Agricultural University, 2022. (In Chinese) Li Qin, Zhou Shuangzhen, Liu Wenyu, Zhai Zhensheng, Pan Yitian, Liu Changchang, Chen Masheng, Wang Hongwei, Huang Min, Zhang Zuxin, Tang Jihua, Du Hewei. A chlorophyll a oxygenase 1 gene ZmCAO1 contributes to grain yield and waterlogging tolerance in maize. JOURNAL OF EXPERIMENTAL BOTANY, 2021, 72(8): 3155-3167. Additional Declarations No competing interests reported. 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461053 SNP markers\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/be07503cb6adebaa85403f4f.jpg"},{"id":90072573,"identity":"92c72f8a-4fa6-49d2-99f8-764cca8a7cb6","added_by":"auto","created_at":"2025-08-28 07:22:46","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":187063,"visible":true,"origin":"","legend":"\u003cp\u003eFrequency distribution map of moisture content traits in maize stalk\u003c/p\u003e","description":"","filename":"2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/859d8da1ca99a37ec340fd7c.jpg"},{"id":90072162,"identity":"7ed2b147-6a81-4f45-a925-a38382f806e5","added_by":"auto","created_at":"2025-08-28 07:14:46","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":120875,"visible":true,"origin":"","legend":"\u003cp\u003ea. Show that from the inside to the outside, the SNP sites identified in the PMSMC1, PMSMC2, PMSMC3 and DMSMC1, DMSMC2, DMSMC3 of maize in different years are represented respectively; The red highlights represent the significant SNP loci of each trait at 0.01 levels, respectively; Figure 3b shows the QQ chart of stalk related traits of maize in different years.\u003c/p\u003e\n\u003cp\u003eFigure 3b. Genome-wide association study of stalk moisture content resistance under the MLM model Manhattan map and QQ map\u003c/p\u003e","description":"","filename":"3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/a534d741c27cecd6722f7586.jpg"},{"id":90072576,"identity":"565b77cf-a655-4941-bb95-3ad057cba41e","added_by":"auto","created_at":"2025-08-28 07:22:46","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":52457,"visible":true,"origin":"","legend":"\u003cp\u003eThe Enrichment map of 14 associated candidate genes\u003c/p\u003e","description":"","filename":"4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/92e32ee44eee510711b69440.jpg"},{"id":90072165,"identity":"b5f2b9ed-b636-4907-bf7e-4177cddba29a","added_by":"auto","created_at":"2025-08-28 07:14:46","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":72820,"visible":true,"origin":"","legend":"\u003cp\u003eThe box chart of phenotypic differences of genotypes at five significant loci (* * *: significant differences)\u003c/p\u003e","description":"","filename":"5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/2bd18f5610ef5b4e4852e91d.jpg"},{"id":104429931,"identity":"50fda93a-d065-4f86-a2b3-7e653cc0e6dc","added_by":"auto","created_at":"2026-03-11 15:27:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1946650,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7282731/v1/af60d03d-1b1f-48da-b049-f7048f346722.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Genome-Wide Association Study Elucidating Key Genetic Determinants of Stalk Water Content in Mature Maize (Zea mays L.)","fulltext":[{"header":"1 INTRODUCTION","content":"\u003cp\u003eMaize (Zea mays L.) is an important food and feed crop globally in the world, and playing a significant role in world food security \u003csup\u003e[\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]\u003c/sup\u003e. Stalk is an important component of the maize plant, which is closely related to plant lodging, material transport, disease resistance stress and grain yield \u003csup\u003e[\u003cspan additionalcitationids=\"CR3\" citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]\u003c/sup\u003e. Stalk water content is an important index to measure stalk fullness, stalk maturity and grain dehydration rate, which is affected by the physiological effects of plants, climate and cultivation measures, and is complementary to other chemical components of stalks \u003csup\u003e[\u003cspan additionalcitationids=\"CR6 CR7\" citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]\u003c/sup\u003e. Stalk water content is directly related to the quality of the stalks, and it is an important criterion for the maturity of maize live stalks. The higher of the stalk water content, the stronger vitality of the stalks, and also the more resistant to external forces. In the maturity period of maize, lodging is easily caused due to the low water content of stalks \u003csup\u003e[\u003cspan additionalcitationids=\"CR10\" citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]\u003c/sup\u003e. Brandon evaluated the influence of water content on maize stalk and the tissue properties of peel, developed two compression test methods to obtain the elastic index model, and evaluated the influence of stalk water content on the tissue properties \u003csup\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e. Water content was negatively correlated with the elastic modulus of peel. There are direct and indirect methods for evaluating the stalk water content, and the drying method is the most commonly used direct measurement method \u003csup\u003e[\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]\u003c/sup\u003e. Indirect measurement methods mainly include the electronic moisture meter measurement method, near infrared reflectance spectroscopy analysis method and probe moisture meter method, which are characterized by simple and fast measurement \u003csup\u003e[\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eDomestic and foreign scholars research on maize stalks mainly focuses on phenotypic traits such as stalk height, ear height, stalk node number, and stalk thickness \u003csup\u003e[\u003cspan additionalcitationids=\"CR17 CR18\" citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]\u003c/sup\u003e, while there is relatively little research on measurement traits such as stalk moisture content. In the early stage of maize growth, the stalk moisture content is less affected by density, but in the later stage of growth, it is more affected by density and decreases with increasing density \u003csup\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e. Therefore, maintaining a certain stalk moisture content in the physiological later stage of maize can effectively enhance stem strength, increase plant lodging resistance, and meet the production needs of high-density maize planting. The trend of stalk moisture content changes varies among different varieties at different growth stages \u003csup\u003e[\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]\u003c/sup\u003e. Different types of inbred lines also exhibit diversity in stalk moisture content, but it is relatively stable during the grain filling stage \u003csup\u003e[\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]\u003c/sup\u003e. In the later stages of growth, as the plant naturally declines, it leads to a decrease in leaf, stem, and root vitality, resulting in a decrease in stalk moisture content \u003csup\u003e[\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]\u003c/sup\u003e. The moisture content of maize male ear stem has a wide range of phenotypic variations and a high general heritability \u003csup\u003e[\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]\u003c/sup\u003e. The hybrid advantage of maize stalk moisture content is mainly manifested by additive effects, with less environmental influence, and can be selected in early generations \u003csup\u003e[\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]\u003c/sup\u003e. However, Zhou found that different genetic systems of stalk moisture content are significantly influenced by genotype and environmental interaction effects, and should be selected in later generations \u003csup\u003e[\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eWhile the genetic basis of moisture content in the maize ear has been extensively studied, the genetics of stalk moisture content-particularly in the third above-ground internode-remains largely unexplored. This study aims to fill this knowledge gap through a genome-wide association study to identify candidate genes controlling this trait. In the present study, we used a wide diversity composed of 236 inbred lines. The water content of the third stem on the ground was determined using the direct drying method and the corrected probe moisture method. The genotype and stalk water content traits were analyzed in the late stage of two years of growth, and significant SNPs related to stalk water content traits were screened. Relevant candidate genes were identified, in order to provide a theoretical reference for cultivating breakthrough maize varieties with strong comprehensive stress resistance.\u003c/p\u003e"},{"header":"2 MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1 Test materials and design\u003c/h2\u003e\u003cp\u003e236 materials include 141 inbred lines of regular maize, 34 inbred lines of explosive maize, and 61 inbred lines of glutinous maize. The experimental design refers to Wang \u003csup\u003e[\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2 Genotype and genome-wide association analysis\u003c/h2\u003e\u003cp\u003eA total of 236 samples were randomly selected from the study population. Genomic DNA was extracted using a magnetic bead-based method, with concentration, purity, and integrity subsequently assessed. Qualified DNA samples were resequenced to 5\u0026times; coverage (BGI, Shenzhen, China; \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ewww.bgi.com\u003c/span\u003e\u003cspan address=\"http://www.bgi.com\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). Reads were aligned to the B73 reference genome (RefSeq v4.0) to detect SNPs. After filtering for variants with missing ratios\u0026thinsp;\u0026lt;\u0026thinsp;0.40 and minor allele frequencies\u0026thinsp;\u0026gt;\u0026thinsp;0.05, 461,053 high-quality SNPs were retained for genome-wide association analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Associations between SNPs and stalk thrust resistance were tested at a significance threshold of P\u0026thinsp;\u0026lt;\u0026thinsp;0.01.\u003c/p\u003e\u003cp\u003eThe mixed linear model (MLM) of TASSEL V5.0 software \u003csup\u003e[\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]\u003c/sup\u003e was used to conduct correlation analysis on the stalk correlation at different growth stages. The significance threshold was set according to the method in the reference literature \u003csup\u003e[\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]\u003c/sup\u003e, and the threshold for this study is P\u0026thinsp;\u0026lt;\u0026thinsp;9.94\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;6\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\u003ch2\u003e2.3 Phenotypic data measurement method and statistical method\u003c/h2\u003e\u003cp\u003eDuring the tasseling, filling, and ripening stages of maize, the moisture content of the third section of the stem on the ground was determined using direct drying method \u003csup\u003e[\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]\u003c/sup\u003e and calibrated probe puncture method \u003csup\u003e[\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]\u003c/sup\u003e (In 2021 and 2022, the average moisture content of drying stems at tasseling stage, filling stage and maturity stage were DMSMC1, DMSMC2 and DMSMC3, also by calibrated probe method were PMSMC1, PMSMC2 and PMSMC3). Select three representative plants with consistent growth conditions for each growth period and measure them three times, taking the average. The average of the three replicates represents the phenotype value of the trait at that growth period.\u003c/p\u003e\u003cp\u003eData were analyzed using Excel 2007 and DPS 7.05. Calculate group PCA and Kinship using TASSEL V5.0 software. Use the R \"CMplot\" software package to draw a correlation graph(\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://github.com/YinLiLin/CMplot\u003c/span\u003e\u003cspan address=\"https://github.com/YinLiLin/CMplot\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003e2.4 Prediction of candidate genes\u003c/h2\u003e\u003cp\u003eBased on the sequence information of the B73 reference genome (B73-RefGenv4) in the reference database, candidate genes related to stalk water content were identified within a range of 2.5Kb upstream and downstream of significant loci. Annotate and predict the functions of candidate genes in the Uniport protein database(\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.uniprot.org/uniprotkb/\u003c/span\u003e\u003cspan address=\"https://www.uniprot.org/uniprotkb/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e"},{"header":"3 RESULTS AND DISCUSSION","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003e3.1 Phenotypic analysis of moisture content traits in maize stalks\u003c/h2\u003e\u003cp\u003eThe basic description statistics and frequency distribution of stalk moisture content traits in different growth stages of maize are shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and Fig.\u0026nbsp;2. The variation range of traits is large and the phenotypic diversity is rich. As the growth process progresses, the stalk moisture content gradually decreases. The skewness and kurtosis of each trait, except for the absolute value of PMSMC1 are all less than 1. Combined with the frequency histogram of trait distribution, they basically show a normal distribution trend, which is consistent with typical quantitative trait characteristics. The generalized heritability of stalk moisture content measured by the drying method is 67.63%, mainly influenced by genetic factors, while the generalized heritability of stalk moisture content measured by the calibrated probe method is 37.51%, which is greatly affected by environmental factors.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eStatistical analysis of moisture content traits in maize stalk at different growth stages\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTrait\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRange\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMean\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCV (%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSkewness\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eKurtosis\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eBroad-sense heritability\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e76.09\u0026ndash;88.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e82.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e-0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e-0.15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e67.63\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e69.27\u0026ndash;88.80\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e78.73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e-0.09\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e67.82\u0026ndash;86.87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e77.48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e-0.03\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e78.25\u0026ndash;93.93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e88.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e-0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e37.51\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e71.66\u0026ndash;92.68\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e82.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e-0.48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e59.68\u0026ndash;87.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e74.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e-0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e-0.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003e3.2 Variance analysis of stalk moisture content of the tested inbred lines\u003c/h2\u003e\u003cp\u003eThe variance analysis of stalk moisture content traits in 236 inbred line materials (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) showed that the genotype effects of the direct drying method and probe-based measurement of stalk moisture reached a highly significant level (P\u0026thinsp;\u0026lt;\u0026thinsp;0.01), indicating significant genetic variation in stalk moisture content traits in the constructed natural population.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eVariance analysis of various traits in different maize inbred lines\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTrait\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVariation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSum of squares\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eDF\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSD\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eDMSMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEnvironment\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3659.667\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e1829.833\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e580.853**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGenotype\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5380.766\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e235\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e22.8969\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e7.268**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eError\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1480.618\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e470\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e3.1503\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003ePMSMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEnvironment\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e22224.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e11112.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1118.25**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGenotype\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6539.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e235\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e27.8281\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2.8**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eError\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4670.404\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e470\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e9.937\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"6\"\u003eNote: * or * * indicate a significant correlation at the 0.05 or 0.01 level.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\u003ch2\u003e\u003cb\u003e3.3 Genome\u003c/b\u003e-\u003cb\u003ewide association study of stalk moisture content at different growth stages of maize\u003c/b\u003e\u003c/h2\u003e\u003cp\u003eThe genome-wide association study was conducted on the stalk water content using MLM models of 236 maize inbred lines at different growth stages over two years by Tassel 5.0, and the Manhattan map and QQ map were drawn (Fig.\u0026nbsp;3a, 3b). The MLM model was used to analyze the correlation between stalk water content at different growth stages over two years. A total of 55 significantly associated SNPs loci were detected on chromosomes of maize (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). We detected SNPs distributed across chromosomes 1\u0026ndash;10 as follows: 4, 3, 3, 1, 33, 0, 2, 3, 3, and 3 respectively. A total of 22 SNPs (DMSMC1: 1, DMSMC2: 3, DMSMC3: 18) were detected related to DMSMC, as well as 33 SNPs (PMSMC1:14, PMSMC2: 1, PMSMC3: 18) related to PMSMC. The explained phenotypic variation ranged from 9.11\u0026ndash;15.13%, with the highest phenotypic interpretation rate being DMSMC3 on chromosome 5_47800595 and the lowest being PMSMC1 on chromosome 4 _13346360.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSignificant locus information on the correlation of maize stalk moisture content\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTrait\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMarker\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChr\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eModel\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePos(bp)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003ep\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eMarker Rsq%\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1_206350619\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e206350619\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.93E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.33\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1_87740598\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e87740598\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.30E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.42\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1_164402750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e164402750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7.36E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e13.53\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1_76763257\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e76763257\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7.69E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.91\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2_45422750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e45422750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.24E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.34\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2_45418447\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e45418447\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.06E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.47\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2_170637158\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e170637158\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.49E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e9.92\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3_204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.77E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.95\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3_204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7.42E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.54\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3_122700273\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e122700273\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.31E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.68\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4_13346360\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e13346360\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9.94E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e9.11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.40E-08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e15.13\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.16E-08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e14.62\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.76E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.84\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47727153\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47727153\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.92E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.31\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.72E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.57\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47876150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47876150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.80E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.96\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47645572\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47645572\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e6.35E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47727153\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47727153\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.69E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.56\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9.01E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.63\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47645572\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47645572\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.02E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.45\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.04E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.56\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47726081\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47726081\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.09E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.51\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.17E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.53\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47876150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47876150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.17E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47726081\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47726081\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.45E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.35\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.53E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.21\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.78E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.19\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.95E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47735500\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47735500\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.54E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.95\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.67E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.79\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.87E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.52\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47880130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47880130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.11E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.71\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47880130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47880130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.39E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.77\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.55E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.59\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.85E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47735500\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47735500\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.93E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.72\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47800595\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.16E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.77\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.21E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.31\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.98E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.24\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_46831646\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e46831646\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.74E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.88\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.76E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7.41E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.81\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5_100064005\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e100064005\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7.46E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e10.7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7_96409678\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e96409678\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.02E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e14.4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7_114436113\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e114436113\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9.65E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.97\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8_67202484\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e67202484\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e6.87E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e15.02\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8_159862794\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e159862794\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.42E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.34\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8_159862794\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e159862794\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9.40E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e9_84316697\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e84316697\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.77E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e9_18935130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e18935130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.92E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e12.25\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e9_70247735\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e70247735\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.45E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e14.38\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10_33621054\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e33621054\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.79E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.63\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10_121739764\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e121739764\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.32E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.28\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10_105984280\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMLM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e105984280\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.61E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11.07\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eFurther analysis revealed that 17 SNPs related to stalk water content traits were detected more than twice by independent association analysis using the MLM models (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), suggesting that these loci carry important genes that can stably regulate stalk water content traits in different environments. Among them, 4 SNPs were detected more than twice by independent association analyses at different growth stages with the same stalk moisture content traits. It is speculated that these loci may carry important genes in the growth process of maize stalk moisture content, which are continuously expressed and play a role in multiple growth stages. Different methods for measuring stalk moisture content have identified 16 SNPs detected more than twice by independent association analyses, suggesting that the candidate genes carried by these loci are more likely to be stably expressed in maize stalk moisture traits. These SNPs that can be significantly correlated with stalk-related traits multiple times have high confidence (HC) Among them, HC-SNP locus 5_47800595 was found to be significantly correlated with DMSMC twice and PMSMC3 once in two growth stages. It was also found that 31 SNPs significantly correlated with DMSMC and PMSMC were clustered in the 5_4764557, 2_47925170 range at different growth stages, which is an enriched region of stalk water content-related genes. Significant correlations were found twice with DMSMC and PMSMC at adjacent sites of 2_245418447, 3_204520247, and 8_159862794 respectively.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003e3.4 Analysis of genes associated with stalk moisture content traits\u003c/h2\u003e\u003cp\u003eBased on the association of two measurement methods, candidate genes were screened and predicted within a range of 2.5kb upstream and downstream of 55 significant SNP loci related to stalk water content traits. Combined with gene functional annotation, 14 candidate genes were identified (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), and 7 candidate genes were detected more than twice. These genes may be highly reliable associated genes for stalk water content traits. Gene ontology (GO)enrichment analysis using the 14 candidate genes, most of the candidate genes discovered are related to Ubiquitin system such as aromatic acid dehydratase, ubiquitin carboxy terminal dehydratase, and threonine phosphatase (Fig.\u0026nbsp;4). At the same time, this study also found three unknown functional genes, namely \u003cem\u003eLOC103636628\u003c/em\u003e, \u003cem\u003eZm00001d049035\u003c/em\u003e, and \u003cem\u003eZm00001d014448\u003c/em\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003e3.4 Identification and functional analysis of high-confidence associated genes\u003c/h2\u003e\u003cp\u003eUsing two detection methods, a total of 7 candidate genes were detected more than twice. Among them, \u003cem\u003eZm00001d014449\u003c/em\u003e was found to be associated with DMSMC and PMSMC 4 times, with a phenotypic variation explanation rate of 11.24%-12.56%. This gene originates from the assembly of CRD101 and is associated with the LHP1 gene. Which is mainly expressed in cells with active cell division and tissue differentiation, such as vascular bundle synthesis, which is of great significance for maintaining cell and tissue structure specificity. \u003cem\u003eZm00001d014448\u003c/em\u003e was found to be associated with DMSMC and PMSMC 3 times, with a phenotypic variation explanation rate of 10.81%-12.63%. The function of this gene is unknown. \u003cem\u003eZm00001d003476\u003c/em\u003e was found to be associated with DMSMC and PMSMC 2 times, with a phenotypic variation explanation rate of 12.34%-12.47%, encoding the synthesis of ubiquitin carboxylic acid terminal hydrolase; \u003cem\u003eZm00001d043592\u003c/em\u003e and \u003cem\u003eZm00001d043593\u003c/em\u003e were found to be associated with DMSMC and PMSMC 2 times, with a phenotypic variation explanation rate of 10.54%-11.95%, They encode cellulose synthase A4 and participate in the SPA1 expression pathway. \u003cem\u003eZm00001d014451\u003c/em\u003e was found to be associated with DMSMC and PMSMC traits twice, with a phenotypic variation explanation rate of 12.05%-12.25%, which is involved in glutamic acid receptor synthesis.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003e3.5 Analysis of haplotype water content traits\u003c/h2\u003e\u003cp\u003eBy analyzing the re-sequencing data and evaluating the phenotypic performance of the population, the association between the above five SNPs and the water content traits of maize stalk was verified. At 2_45418447 locus, the GG genotype was associated with high stem water content of PMSMC1, while the AA genotype was associated with low content, while at 2_45422750 locus, the AA genotype was associated with high stem water content of PMSMC1, and the GG genotype was associated with low content. At the 2_170637158 locus, the TT genotype was associated with high stem water content of DMSMC2, while the CC genotype was associated with low stem water content; At the 5_47750439 locus, the GG genotype was associated with high stem water content of DMSMC3, the AA genotype was associated with low stem water content, and there was no significant difference between the GG genotype and the PMSMC 3 stem water content (Fig.\u0026nbsp;5).\u003c/p\u003e\u003cp\u003eTable 4 Prediction information of the candidate gene\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"No\" id=\"Taba\" border=\"1\"\u003e\u003ccolgroup cols=\"13\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eTrait\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMarker\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eChr\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003ePos\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003ep\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c11\" namest=\"c9\"\u003e\u003cp\u003eMarker Rsq %\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" colname=\"c12\"\u003e\u003cp\u003eGene accession number\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c13\"\u003e\u003cp\u003eGene annotation\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e1_76763257\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e76763257\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e7.69E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eLOC103636628\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eNO\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e1_87740598\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e87740598\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e4.30E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eZm00001d029806\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eNEP1-interacting protein-like 1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e2_170637158\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e170637158\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e2.49E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e9.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eZm00001d005351\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eArogenate dehydratase, 4.2.1.91\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e2_45418447\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e45418447\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e2.06E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c12\" namest=\"c11\" rowspan=\"2\"\u003e\u003cp\u003e\u003cem\u003eZm00001d003476\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eUbiquitin carboxyl-terminal hydrolase\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e2_45422750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e45422750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.24E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.34\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e3_204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.77E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c12\" namest=\"c11\" rowspan=\"2\"\u003e\u003cp\u003e\u003cem\u003eZm00001d043592, Zm00001d043593\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eSPA1-related 3, CESA4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e3_204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e204520247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e7.42E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e10.54\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e4_13346360\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e13346360\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e9.94E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e9.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eZm00001d049035\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eNO\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e9.01E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.63\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" morerows=\"5\" nameend=\"c12\" namest=\"c11\" rowspan=\"6\"\u003e\u003cp\u003e\u003cem\u003eZm00001d014448\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\" morerows=\"5\" rowspan=\"6\"\u003e\u003cp\u003eNO\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47711262\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.17E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.53\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e2.87E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.52\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47716318\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e4.21E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.31\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e5.76E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47718537\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e7.41E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e10.81\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e3.85E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" morerows=\"7\" nameend=\"c12\" namest=\"c11\" rowspan=\"8\"\u003e\u003cp\u003e\u003cem\u003eZm00001d014449, crd101\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\" morerows=\"7\" rowspan=\"8\"\u003e\u003cp\u003eChromo domain-containing protein LHP1,\u003c/p\u003e\u003cp\u003eChrom domain-containing protein CRD101\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47744387\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e4.98E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.24\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e2.67E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.79\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47747099\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e3.55E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.59\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.04E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.56\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47750439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.78E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.19\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e3.76E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.84\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47752006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e5.72E-07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.57\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.53E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c12\" namest=\"c11\" rowspan=\"2\"\u003e\u003cp\u003e\u003cem\u003eZm00001d014451\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eGlutamate receptor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e5_47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e47925170\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.95E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePMSMC1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e9_18935130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e18935130\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e1.92E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e12.25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eZm00001d045322, Zm00001d045323\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eUbiquinone biosynthesis O-methyltransferase,\u003c/p\u003e\u003cp\u003eDouble B-box zinc finger protein 11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDMSMC2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e10_105984280\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e105984280\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e5.61E-06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e11.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003e\u003cem\u003eZm00001d025134\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003eFPL domain-containing protein\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e\u003ch2\u003e4.1 Comparative analysis of significant SNPs loci in stalk moisture content traits\u003c/h2\u003e\u003cp\u003eBy comparing and analyzing the genetic mapping results of stalk moisture content traits at home and abroad, it was found that among the many previous studies on moisture content, there were relatively few studies on the localization of the third stalk moisture content in maize fields, only related studies on the localization of male ear stem, cob, bract leaf, and grain moisture content. Shang studied the stem traits of 70 major maize inbred lines and found that the stalk moisture content of maize was relatively stable during the grain filling stage \u003csup\u003e[\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]\u003c/sup\u003e. Only 4 SNPs were found to be significantly associated with stalk moisture content during the grain filling stage, while 36 SNPs were associated during the maturity stage. Li conducted a GWAS analysis on the moisture content of maize male ear stems, and found that the significant locus on 10_121259591 was located at a distance similar to the significant locus 10_121739764 located during the tasseling stage stalk moisture content in this study \u003csup\u003e[\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]\u003c/sup\u003e. The water content trait of bract leaves was located at significant loci 1_219374182 and 3_206692486, which are similar in distance to the significant loci 1-206350619 and 3_204520247 for mature stalk water content identified in this study; The water content trait of the spike axis was located at significant loci 2_170297547 and 2_45753838, which are close to the significant loci of stalk water content in the milk ripening stage and in the tasseling stage, which were located at positions 2_170637158 and 2_45418447 respectively. The significant SNPs interval of stalk moisture content trait identified in this study overlaps significantly with the correlation traits studied by previous researchers, indicating high reliability of the localization results and suitability for further exploration of trait-associated genes. In addition, the new significant SNP loci detected in this experiment, which are different from previous studies, especially some highly reliable significant SNP loci and highly efficient significant SNPs with a phenotype explanatory rate of over 15%, and also that great significance for exploring new significant loci of stalk water content traits and subsequently discovering new associated genes.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\u003ch2\u003e4.2 Analysis of Related Genes in Maize Stalks\u003c/h2\u003e\u003cp\u003eStudies demonstrate that post-physiological maturity, natural plant aging causes decrease in stem dry matter and water content, which is the main reason for the decrease in stem puncture strength and the increase in stem folding rate. In the early stage of research on the lodging resistance related traits of maize stems \u003csup\u003e[\u003cspan additionalcitationids=\"CR26\" citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]\u003c/sup\u003e, similar significant SNPs and candidate gene information were also discovered in multiple identical intervals, especially the candidate gene \u003cem\u003eZm00001d014448\u003c/em\u003e and \u003cem\u003eZm00001d014449\u003c/em\u003e, which were clustered near 5_47747099. \u003cem\u003eZm00001d014449\u003c/em\u003e is related to the LHP1 gene and participates in vascular bundle synthesis. It is located in the above-ground part of maize and encodes heterochromatin protein 1. It is mainly expressed in cells or tissues with active cell division and tissue differentiation, and is important for maintaining the specificity of cell and tissue structure \u003csup\u003e[\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]\u003c/sup\u003e. The \u003cem\u003eZm00001d005351\u003c/em\u003e gene associated with 2_170637158 expresses the product of aromatic acid dehydratase, which is a key enzyme catalyzing the conversion of aromatic acids to phenylalanine in the chloroplasts and plastid matrix of vascular plants. It plays an important role in the biosynthesis of cell wall lignin and photosynthesis, and can be used for improving the dehydration performance of plant stems. Expression in Arabidopsis can reduce lignin content, resulting in a decrease in stem weight and length \u003csup\u003e[\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]\u003c/sup\u003e. At low nitrogen levels, it is associated with maize leaf type-related traits, and downregulation of expression directly affects the synthesis of tyrosine and alanine \u003csup\u003e[\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]\u003c/sup\u003e. The expression product of the \u003cem\u003eZm00001d003476\u003c/em\u003e gene, also associated with the attachment of 2_45418447, is a ubiquitin carboxy terminal hydrolase, this enzyme is an extremely important factor in cell biology that degrades proteins and can coordinate and regulate various biological processes and pathways to maintain cell health. The gene \u003cem\u003eZm00001d043593\u003c/em\u003e associated with the attachment of 3_204520247 expresses cellulose synthase A4, which has been validated for expression in plants such as tobacco and plays an important role in plant stem cellulose synthesis \u003csup\u003e[\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]\u003c/sup\u003e. In addition, protein differential expression occurs in the stems of maize under drought stress. \u003cem\u003eZm00001d029806\u003c/em\u003e regulates the absorption of lead in maize seedling roots \u003csup\u003e[\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]\u003c/sup\u003e, affects the long-distance transport of nitrogen elements in maize \u003csup\u003e[\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]\u003c/sup\u003e, and is related to the development of ear length and row grain number \u003csup\u003e[\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]\u003c/sup\u003e. The expression level of \u003cem\u003eZm00001d014451\u003c/em\u003e gene decreased significantly after high-temperature treatment \u003csup\u003e[\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]\u003c/sup\u003e. \u003cem\u003eZm00001d045322\u003c/em\u003e is expressed and enriched in low-temperature-tolerant maize inbred lines, which may be related to ubiquitin biosynthesis \u003csup\u003e[\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]\u003c/sup\u003e. The \u003cem\u003eZm00001d045323\u003c/em\u003e gene is within the range of active recombinant genes in the infiltrated population of large grass \u003csup\u003e[\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e]\u003c/sup\u003e. The \u003cem\u003eZm00001d025134\u003c/em\u003e gene is associated with maize leaf length traits at low nitrogen levels \u003csup\u003e[\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]\u003c/sup\u003e. The \u003cem\u003eZm00001d025134\u003c/em\u003e gene is associated with drought resistance in maize, and ZmVIL2 may inhibit its expression by binding to the promoter, thereby enhancing the drought resistance of maize plants \u003csup\u003e[\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\u003ch2\u003e4.3 Correlation analysis of moisture content traits in maize stalks\u003c/h2\u003e\u003cp\u003eThe stem is the bone of maize, mainly responsible for the transportation and supply of water and nutrients. Studies have shown that stem thickness, stem strength, stem tissue water absorption capacity, and transport capacity can all be used as indicators of maize drought resistance \u003csup\u003e[\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]\u003c/sup\u003e. Anderson's research suggests that the moisture content of maize stems gradually decreases after tasseling, and there is a significant positive correlation between the moisture content of the stem pulp and stem strength. Stem strength decreases with the decrease in moisture content. At the same time, the plant's ability to resist stem rot is related to stalk moisture content and soluble sugar content. Plants with high moisture content in the stem pulp after physiological maturity of maize have a lower incidence of stem rot \u003csup\u003e[\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]\u003c/sup\u003e. Djordjevic's believe that the moisture content of mature maize stems is significantly negatively correlated with lodging rate \u003csup\u003e[\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]\u003c/sup\u003e. The impurity rate of maize kernels harvested is significantly positively correlated with the wet weight of maize stems, and significantly negatively correlated with the moisture content of stems \u003csup\u003e[\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]\u003c/sup\u003e. The stem moisture content is significantly or extremely significantly positively correlated with stem height, number of stem nodes, ear height, and ear nodes, indicating that the height of the plant and ear affect the stalk moisture content. The stalk moisture content of maize inbred lines bred in China in the 1980s and 1990s decreased by 3 percentage points, which may be related to the breeding goal of focusing more on the selection of lodging resistance traits with lower plant height and ear height \u003csup\u003e[\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]\u003c/sup\u003e. There is a significant positive correlation between the crushing strength of stems and their moisture content and fresh density. Increasing the fresh density and moisture content of stems can improve the mechanical characteristics of maize stems during flowering, thereby enhancing their resistance to stem lodging \u003csup\u003e[\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]\u003c/sup\u003e. The thickness of the stem bark and the moisture content of the stem internode directly determine the strength of the stem puncture and transverse bending, and are also important criteria for judging the mechanical properties of the stem. In addition, spraying chemical regulators such as phosphatidylcholine can significantly enhance the moisture content of the maize stem in the later stage of maturity, thereby enhancing its ability to resist lodging \u003csup\u003e[\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eDuring germplasm improvement for stem lodging resistance and variety breeding, selecting varieties with slow consumption rates of water and soluble sugar content under the ear in the late stage of physiological maturity, or delaying leaf senescence after physiological maturity through cultivation and breeding methods to increase stem soluble sugar content, can effectively reduce the risk of stem lodging and increase stem carbohydrate accumulation \u003csup\u003e[\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]\u003c/sup\u003e. The 236 inbred lines used in this study included conventional maize inbred lines, sweet glutinous maize inbred lines, and burst maize inbred lines, with a wide variety of types. The selected germplasm materials with slow stalk moisture reduction in the late maturity stage can be used to improve the stem lodging resistance of core germplasm materials. In addition, the candidate genes associated with the study are undergoing functional validation and the development of stalk water content-related functional markers can further promote the improvement and creation of germplasm materials with high stalk water content in the later stage of physiological maturity.\u003c/p\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNo ethics approval was required. The authors declare that the experimental methods conducted in this study complied with current Chinese laws and regulations.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated during the current study are available in the [GVM000940] repository, [https://bigd.big.ac.cn/gvm/getProjectDetail?Project=GVM000940].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no conflicts of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by Henan Provincial Key Science and Technology Project Creation of Germplasm Resources and New Variety Breeding and Application of High-Yield and Multi-Resistance Maize (2022010202), the open-ended foundation of National Key Laboratory of Wheat and Maize Crop Science, Henan Agricultural University (39990109), Henan Provincial Science and Technology Project Analysis of Genetic Basis and Application of Stalk Quality Traits by GWAs (202102110248).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026rsquo; contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003e\u0026dagger;\u003c/sup\u003eMeili Yang, Bangtai Wang contributed equally to this work.\u003c/p\u003e\n\u003cp\u003e* Corresponding author: Hongwei Lu([email protected]), Jiafa Chen([email protected])\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eMLY, BTW, HWL and JFC designed this study. QYW, JMC developed the populations. ZHW recorded the data. BTW analyzed the data. BTW and MLY drafted the manuscript. HWL, JFS, XHZ and JFC revised the manuscript. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank professor Chunhua Mu of Shandong Academy of Agricultural Sciences for providing the maize population.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor details\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e1Hebi Academy of Agricultural Sciences, Hebi 458031, Henan, China; 2 Henan Agricultural University College of Life Sciences, Zhengzhou 450002, Henan, China; 3Henan Maize Breeding Engineering Technology Research Center, Hebi 458031, Henan, China; 4Hebi Maize Breeding Key Laboratory, Hebi 458031, Henan, China; 5National Key Laboratory of Wheat and Maize Crop Science; Collaborative Innovation Center of Henan Grain Crops; College of Agronomy; Henan Agricultural University, Zhengzhou 450002, Henan, China.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eChou H G, Zhang S H, Yang J, Jing Y. Development of China\u0026apos;s maize industry, challenges in the future and policy suggestions. Journal of Agricultural Science and Technology, 2013(1):20-24.\u003c/li\u003e\n\u003cli\u003eZhou Y, Gu W R, Zhao M, Tong T, Liu X M, Li C F, Li J, Wei T. Differences of stalk characteristics and grain mechanically harvesting qualities of different maturing type spring maize in Heilongjiang Province. Acta Agriculturae Boreali-Sinica, 2017, 32(S1): 140-146.\u003c/li\u003e\n\u003cli\u003eFeng G, Jing X Q, Li Y Y, Wang L, Huang C L. Correlation and path analysis of lodging resistance with maize stem characters. Acta Agriculturae Boreali-Sinica, 2010, 25(S1): 72-74.\u003c/li\u003e\n\u003cli\u003eYang Jianping, Li Meng, Yin Yue, Liu Yan, Gan Xinke, Mu Xiaohuan, Li Hanqin, Li Jiankun, Li Haochuan, Zheng Jun, Gou Mingyue. Spatial accumulation of lignin monomers and cellulose underlying stalk strength in maize, Plant Physiology and Biochemistry, 2024, 214:108918.\u003c/li\u003e\n\u003cli\u003eJin Yingjie. Effects of Stalk Mechanical Characteristics and Chemical Components on Lodging Resistance of Different Maize Cultivars. Master\u0026apos;s Thesis of Henan Agricultural University, 2018. (In Chinese)\u003c/li\u003e\n\u003cli\u003eZhao Yanming, Xiao Jun, Zhang Baoshi, Wang Changtao, Sha Wancun. Genetic Studies on Moisture Content in Maize-stalk. Liaoning Agricultural Sciences, 2002(3): 19-20.\u003c/li\u003e\n\u003cli\u003eWang Bangtai, Yang Meili, Wang Jing, Wang Zhihong, Guo Hua, Hou Xianjun, Zhang Wenbo,Chang Jianzhi. Correlation Analysis and Rapid Determination of Grain Dehydration Rate of Maize Varieties. Chinese Agricultural Science Bulletin, 2020, 36(15): 18-23.\u003c/li\u003e\n\u003cli\u003eXiao Yingjie, Liu Haijun, Wu Liuji, Warburton Marilyn, Yan Jianbing. Genome-wide Association Studies in Maize: Praise and Stargaze. 2017, 10(3): 359-374.\u003c/li\u003e\n\u003cli\u003ePei Zhichao. Effects of Chemistry Regulation on Straw Quality of High-yield Maize Master\u0026apos;s Thesis of Chinese Academy of Agricultural Science Dissertation, 2010. (In Chinese)\u003c/li\u003e\n\u003cli\u003eXue Jun, Ming Bo, Xie Ruizhi, Wang Keru, Hou Peng, Li Shaokun. Evaluation of maize lodging resistance based on the critical wind speed of stalk breaking during the late growth stage, Plant Methods, 2020, 16:148-156.\u003c/li\u003e\n\u003cli\u003eLiu Shenqun, Guo Jinrui, Zhang Weijian, Ren Jun, Song Zhenwei, Yan Xiaogong, Song Fengbin, Zhu Xianchan. Effect of Planting Density on Water Content of Stalk and Root in Spring Maize. Soil and Crop, 2014, 3(3): 93-98.\u003c/li\u003e\n\u003cli\u003eBrandon Sutherland, Kirsten Steele, Joseph Carter, Douglas D. Cook. The influence of water content on the longitudinal modulus of elasticity of maize stalk pith and rind tissues. BMC Plant Methods, 2023, 19(64): 2-11.\u003c/li\u003e\n\u003cli\u003eXiang Kui. Genetic analysis and measuring method development of kernel fast dry down rate in maize. Doctor\u0026apos;s Thesis of Sichuan Agricultural University, 2011. (In Chinese)\u003c/li\u003e\n\u003cli\u003eGuo Hua, Wang Jing, Yang Meili, Wang Zhihong, Wang Bangtai. Building and Utilization of One Quick Water-content Measuring Method in Kernel of Maize Based on Regression Model. Journal of Maize Sciences, 2018, 26(3): 63-68.\u003c/li\u003e\n\u003cli\u003eWang Guorui, Yuan Zhen, Zhang Pengyu, Liu Zhixue, Wang Tongchao, Wei Li. Genome-wide analysis of NAC transcription factor family in maize under drought stress and rewatering. PHYSIOLOGY AND MOLECULAR BIOLOGY OF PLANTS, 2020, 26(4): 705-717.\u003c/li\u003e\n\u003cli\u003eMa Yajie, Bao Jianxi, Gao Yuexin, Li Yanan, Qin Wenxuan, Wang Yanbo, Long Yan, Li Jinping, Dong Zhenying, Wan Xiangyuan, Genome-wide association analysis of plant height and ear height related traits in maize. ACTA AGRONOMICA SINICA, 2023, 49(3): 647-661.\u003c/li\u003e\n\u003cli\u003eSun Qiang, Ren Jiaojiao, Xu Xiaoming, Li Zongze, Huang Bowen, Chen Zhanhui, Wu Penghao. QTL Mapping and Genomic Selection for Plant Height and Ear Height in Maize. Journal of Maize Sciences, 2022, 30(4): 40-47.\u003c/li\u003e\n\u003cli\u003eWang Xintao, Wei Feng, Dai Ziju, Li Baoye, Hao Junjie. Identification of Candidate Genes Associating with Stem Diameter in Maize (Zea mays L.) Based on Integrating QTL Mapping and RNA-seq. Journal of Plant Genetic Resources, 2022, 23(6): 1737-1745.\u003c/li\u003e\n\u003cli\u003eWang Jingjing, Li Rongfa, Liu Dongyao, Zhao Wenlong, Miao Zhengyan, Jia Xucun, Dong Pengfei, Wang Qun. Endogenous Hormones Improve Lodging Tolerance of Maize (Zea mays L.) by Regulating Stalk Structure Under Elevated Temperature. JOURNAL OF PLANT GROWTH REGULATION, 2024, 43(2):445-457.\u003c/li\u003e\n\u003cli\u003eShang Qiqi, Zhang Degui, Wang Kaixin, Wang Guoliang, Pan Jinbao, Li Xinhai, Shi Liyu. Diversity Analysis of Stalk Traits in Maize Inbred Lines in China. Journal of Plant Genetic Resources, 2020, 21(2): 321-329.\u003c/li\u003e\n\u003cli\u003eXue Jun, Wang Keru, Xie Ruizhi, Gou Ling, Zhang Wangfeng, Ming Bo, Hou Peng, Li Shaokun. Research Progress of Maize Lodging During Late Stage. Scientia Agricultura Sinica, 2018, 51(10): 1845-1854.\u003c/li\u003e\n\u003cli\u003eLi Zhen. Rapid Determination Method and Genome-Wide Association Study of Maize Kernel Moisture Content in Mature Period. Master\u0026apos;s Thesis of Hebei Agricultural University, 2019. (In Chinese)\u003c/li\u003e\n\u003cli\u003eSun Gaoyang, Zhang Xuehai, Duan Haiyang, Gao Jionghao, Li Na, Su Pingping, Xie Huiling, Li, Weihua, Fu Zhiyuan, Huang Yubi, Tang Jihua. Dissection of the genetic architecture of peduncle vascular bundle-related traits in maize by a genome-wide association study. PLANT BIOTECHNOLOGY JOURNAL, 2022, 20(6):1042-1053.\u003c/li\u003e\n\u003cli\u003eZhou Dongsheng, Zhao Yanming, Hou Hengjun, Li Guangxian. Study on Genetic Main Effects and Genotype Environment Interaction Effects for Moisture Content of Different Parts in Maize. Journal of Maize Sciences, 2011, 19(3): 35-38.\u003c/li\u003e\n\u003cli\u003eWang Bangtai, Yang Meili, Guo Hua, Wang Jing, Wang Zhihong, Lu Hongwei, Qin Guiwen, Chen Jiafa. Genome-wide association study for stalk lodging resistance related traits in maize (Zea mays L.). BMC Genomics, 2024, 25:19-32. -5.\u003c/li\u003e\n\u003cli\u003eBradbury PJ, Zhang Z, Kroon DE, Casstevens TM, Ramdoss Y, Buckler ES, Bishop PM. TASSEL: software for association mapping of complex traits in diverse samples. Bioinformatics, 2007,23(19): 2633\u003c/li\u003e\n\u003cli\u003eWang M, Yan J, Zhao J, Song W, Zhang X, Xiao Y, Zheng Y. Genome-wide association study (GWAS) of resistance to head smut in maize. Plant Sci, 2012, 196: 125-131.\u003c/li\u003e\n\u003cli\u003eLv Yuhua. The Evolution Analysis of Plant LHP1 and Expression patterns of LHP1 from three species of Poaceae. Master\u0026apos;s Thesis of Guangxi University, 2011. (In Chinese)\u003c/li\u003e\n\u003cli\u003eOliver R A Corea, CL Cardenas, SE Brewer, AM Patten, LB Davin, NG Lewis. Arogenate dehydratase isoenzymes profoundly and differentially modulate carbon flux into lignins. Journal of Biological Chemistry. 2012, 287(14): 11446-11459.\u003c/li\u003e\n\u003cli\u003eHe Bing. QTL Mapping for Controlling Maize leaf type Related traits under different Nitrogen Treatment Conditions. Master\u0026apos;s Thesis of Guizhou University, 2021. (In Chinese)\u003c/li\u003e\n\u003cli\u003eGuan Lufan. Genetic Transformation of Cellulose Synthase gene BpCesA4 from Betula Platyphylla. Master\u0026apos;s Thesis of Northeast Forestry University, 2009. (In Chinese)\u003c/li\u003e\n\u003cli\u003eHou Fengxia. Molecular network analysis of ZmbZIP54 and ZmbZIP107 transcription factors regulating lead tolerance in maize seedlings. Doctor\u0026rsquo;s Thesis of Sichuan Agricultural University, 2023. (In Chinese)\u003c/li\u003e\n\u003cli\u003eXu Jiahui. Elucidation of the responsive mechanisms of maize roots to long distance nitrogen signals using a split root system. Master\u0026apos;s Thesis of Sichuan Agricultural University, 2020. (In Chinese)\u003c/li\u003e\n\u003cli\u003eNing Qiang. Cloning and Functional analysis of a major QTL qEl7 for Ear length and Kernel Number per row in Maize. Doctor\u0026rsquo;s Thesis of Huazhong Agricultural University, 2021. (In Chinese)\u003c/li\u003e\n\u003cli\u003eZhou Shenghui, Zhang Lei, Lv Xinze, Huang Jinguang. Ning Qiang. Identification and Analysis of GLR Family Genes in Maize. Journal of Maize Sciences, 2021. 29(2): 35-42.\u003c/li\u003e\n\u003cli\u003eMeng Aiju. Genome-wide Association Studies and Functional Studies of Candidate Genes for Germination-related Traits of Low Temperature Tolerance in Maize. Doctor\u0026rsquo;s Thesis of Shandong Agricultural University, 2022. (In Chinese)\u003c/li\u003e\n\u003cli\u003eFan Feifei. Genetic Analysis of Genomic Expression Variation in An Introgressive Population of Teosinate. Master\u0026apos;s Thesis of Huazhong Agricultural University, 2019. (In Chinese)\u003c/li\u003e\n\u003cli\u003eZhang Dongling. Functional analysis of maize gene ZmVIL2 in response to drought stress. Doctor\u0026apos;s Thesis of Henan Agricultural University, 2024. (In Chinese)\u003c/li\u003e\n\u003cli\u003eTan Guobo, Zhao Liqun, Zhang Lihua, Zhao Hongxiang, Fang Xiangqian, Meng Xiangmeng, Bian Shaofeng, Effects of Different Water Stresses on Plant Character, Photosynthetic Physiology and Yield in Maize Jointing Stage, Journal of Maize Science, 2010, (1): 96-98.\u003c/li\u003e\n\u003cli\u003eAnderson B. Evaluation of Methods for Identification of Corn Genotypes with Stalk Rot and Lodging Resistance. Plant Disease, 1994, 78(6): 590.\u003c/li\u003e\n\u003cli\u003eDjordjevic Jovan S, Ivanovic Mile R. Genetic Analysis for Stalk Lodging Resistance in Narrow-Base Maize Synthetic Population ZPS14. Crop Science, 1996, 36(4): 909-913.\u003c/li\u003e\n\u003cli\u003eZhu Xueqing. Selection and evaluation of maize varieties suitable for machine harvesting. Master\u0026apos;s Thesis of Hunan Agricultural University, 2021. (In Chinese)\u003c/li\u003e\n\u003cli\u003eWang Kaixin, Cheng Zimeng, Yang Yihan, Zhang Degui, Li Xinhai, Shi Liyu. Evolvement of Stem-related Traits of Maize Inbred Lines Through Different Decades in China. Journal of Plant Genetic Resources, 2021, 22(1): 157-164.\u003c/li\u003e\n\u003cli\u003eLiu Shengqun, Liu Tiedong, Song Fengbin, Wang Yang, Qi Xiaoning, Zhu Xianchan. Effects of Row Orientation and Planting Pattern on Traits Associated with Stem Lodging in Maize. Soils and Crops, 2016, 5(3): 159-165.\u003c/li\u003e\n\u003cli\u003eChen Xiang. The change of stalk lodging resistance characteristics before and after physiological maturity of maize. Master\u0026apos;s Thesis of Sichuan Agricultural University, 2022. (In Chinese)\u003c/li\u003e\n\u003cli\u003eLi Qin, Zhou Shuangzhen, Liu Wenyu, Zhai Zhensheng, Pan Yitian, Liu Changchang, Chen Masheng, Wang Hongwei, Huang Min, Zhang Zuxin, Tang Jihua, Du Hewei. A chlorophyll a oxygenase 1 gene ZmCAO1 contributes to grain yield and waterlogging tolerance in maize. JOURNAL OF EXPERIMENTAL BOTANY, 2021, 72(8): 3155-3167.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Maize, Stalk traits, Lodging resistance, Stalk water content, Genome-wide association study","lastPublishedDoi":"10.21203/rs.3.rs-7282731/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7282731/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground\u003c/h2\u003e\u003cp\u003eThe Stalk Moisture Content (SMC) is an important measure of stem fullness, disease resistance and lodging resistance.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e\u003cp\u003eIn this study, 236 inbred lines were used as the basic materials, and the water content of stalk in the third section was determined using the drying method and calibrated probe method for maize were investigated at the tasseling, grain filling, and maturity stages. 461,053 high-quality SNPs obtained through whole-genome resequencing were used for a genome-wide association study. As a result of mixed linear model (MLM, P\u0026thinsp;\u0026lt;\u0026thinsp;9.94\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;6\u003c/sup\u003e) analysis, 55 significant SNPs related to traits were detected, explaining 9.11%-15.13% phenotypic variation, among which 22 SNPs were found by the drying method (DMSMC) and 33 SNPs by the calibrated probe method(PMSMC). A total of 7 candidate genes were detected repeatedly, The \u003cem\u003eZm00001d014449\u003c/em\u003e (detected four times) encodes a chromodomain protein involved in vascular bundle formation. The \u003cem\u003eZm00001d003476\u003c/em\u003e (detected twice) encodes ubiquitin carboxyl-terminal hydrolase. The \u003cem\u003eZm00001d005351\u003c/em\u003e gene was associated with the DMSMC2 phase and encodes the synthesis of aromatic acid dehydratase. The \u003cem\u003eZm00001d025134\u003c/em\u003e gene was a chromosome domain protein, which was found to be related to drought resistance of maize.\u003c/p\u003e\u003ch2\u003eConclusions\u003c/h2\u003e\u003cp\u003eThis study helps to deepen the understanding of the genetic basis of maize stalk water content traits, and provides theoretical guidance for screening germplasm materials.\u003c/p\u003e","manuscriptTitle":"Genome-Wide Association Study Elucidating Key Genetic Determinants of Stalk Water Content in Mature Maize (Zea mays L.)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-08-28 07:14:41","doi":"10.21203/rs.3.rs-7282731/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"b1743c1b-6cec-4eb9-ba4b-af6cbe65e06c","owner":[],"postedDate":"August 28th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-03-11T15:24:31+00:00","versionOfRecord":[],"versionCreatedAt":"2025-08-28 07:14:41","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7282731","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7282731","identity":"rs-7282731","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}