Maize requires more water than most other crops; therefore, the water use efficiency of this crop must be improved for maize production under undesirable land and changing environmental conditions. To elucidate the genetic control of drought in maize, we evaluated approximately 5000 inbred lines from 30 linkage-association joint mapping populations under two contrasting water regimes for seven drought-related traits, including yield and anthesis-silking interval (ASI). The joint linkage analysis was conducted to identify 220 quantitative trait loci (QTLs) under well-watered conditions and 169 QTLs under water-stressed conditions. The genome-wide association analysis identified 365 single nucleotide polymorphisms (SNPs) associated with drought-related traits, and these SNPs were located in 354 candidate genes. Fifty-two of these genes showed significant differential expression in the inbred line B73 under the well-watered and water-stressed conditions. In addition, genomic predictions suggested that the moderate-density SNPs obtained through genotyping-by-sequencing were able to make accurate predictions in the nested association mapping population for drought-related traits with moderate-to-high heritability under the water-stressed conditions. The results of the present study provide important information that can be used to understand the genetic basis of drought stress responses and facilitate the use of beneficial alleles for the improvement of drought tolerance in maize.

The pentatricopeptide repeat (PPR) gene family is one of the largest gene families in land plants (450 PPR genes in Arabidopsis, 477 PPR genes in rice and 486 PPR genes in foxtail millet) and is important for plant development and growth. Most PPR genes are encoded by plastid and mitochondrial genomes, and the gene products regulate the expression of the related genes in higher plants. However, the functions remain largely unknown, and systematic analysis and comparison of the PPR gene family in different maize genomes have not been performed. In this study, systematic identification and comparison of PPR genes from two elite maize inbred lines, B73 and PH207, were performed. A total of 491 and 456 PPR genes were identified in the B73 and PH207 genomes, respectively. Basic bioinformatics analyses, including of the classification, gene structure, chromosomal location and conserved motifs, were conducted. Examination of PPR gene duplication showed that 12 and 15 segmental duplication gene pairs exist in the B73 and PH207 genomes, respectively, with eight duplication events being shared between the two genomes. Expression analysis suggested that 53 PPR genes exhibit qualitative variations in the different genetic backgrounds. Based on analysis of the correlation between PPR gene expression in kernels and kernel-related traits, four PPR genes are significantly negatively correlated with hundred kernel weight, 12 are significantly negatively correlated with kernel width, and eight are significantly correlated with kernel number. Eight of the 24 PPR genes are also located in metaQTL regions associated with yield and kernel-related traits in maize. Two important PPR genes (GRMZM2G353195 and GRMZM2G141202) might be regarded as important candidate genes associated with maize kernel-related traits. Our results provide a more comprehensive understanding of PPR genes in different maize inbred lines and identify important candidate genes related to kernel development for subsequent functional validation in maize.

Drought is a major abiotic stress affecting maize development and growth. Unravelling the molecular mechanisms underlying maize drought tolerance and enhancing the drought tolerance of maize is of great importance. However, due to the complexity of the maize genome and the multiplicity of drought tolerance mechanisms, identifying the genetic effects of drought tolerance remains great challenging. Using a mixed linear model (MLM) based on 362 maize inbred lines, we identified 40 associated loci and 150 candidate genes associated with survival rates. Concurrently, transcriptome analysis was conducted for five drought - tolerant and five drought - sensitive lines under Well-Watered (WW) and Water-Stressed (WS) conditions. Additionally, through co-expression network analysis (WGCNA), we identified five modules significantly associated with the leaf relative water content (RWC) under drought treatment. By integrating the results of GWAS, DEGs, and WGCNA, four candidate genes (Zm00001d006947, Zm00001d038753, Zm00001d003429 and Zm00001d003553) significantly associated with survival rate were successfully identified. Among them, ZmGRAS15 (Zm00001d003553), a GRAS transcription factor considered as a key hub gene, was selected for further functional validation. The overexpression of ZmGRAS15 in maize could significantly enhance drought tolerance through regulating primary root length at the seedling stage. This study provides valuable information for understanding the genetic basis of drought tolerance and gene resources for maize drought tolerance breeding.