ZmC2H2-149 negatively regulates drought tolerance by repressing ZmHSD1 in maize.

Drought is a major abiotic stress that limits maize production worldwide. Therefore, it is of great importance to improve drought tolerance in crop plants for sustainable agriculture. In this study, we examined the roles of Cys2 /His2 zinc-finger-proteins (C2H2-ZFPs) in maize's drought tolerance as C2H2-ZFPs have been implicated for plant stress tolerance. By subjecting 150 Ac/Ds mutant lines to drought stress, we successfully identified a Ds-insertion mutant, zmc2h2-149, which shows increased tolerance to drought stress. Overexpression of ZmC2H2-149 in maize led to a decrease in both drought tolerance and crop yield. DAP-Seq, RNA-Seq, Y1H and LUC assays additionally showed that ZmC2H2-149 directly suppresses the expression of a positive drought tolerance regulator, ZmHSD1 (hydroxysteroid dehydrogenase 1). Consistently, the zmhsd1 mutants exhibited decreased drought tolerance and grain yield under water deficit conditions compared to their respective wild-type plants. Our findings thus demonstrated that ZmC2H2-149 can regulate ZmHSD1 for drought stress tolerance in maize, offering valuable theoretical and genetic resources for maize breeding programmes that aim for improving drought tolerance.

Analysis of the molecular mechanisms regulating how ZmEREB24 improves drought tolerance in maize (Zea mays) seedlings.

Drought stress is one of the most limiting factors of maize productivity and can lead to a sharp reduction in the total biomass when it occurs at the seedling stage. Improving drought tolerance at the seedling stage is of great importance for maize breeding. The AP2/ERF transcription factor family plays a critical role in plant response to abiotic stresses. Here, we used a preliminary previously-generated ranscriptomic dataset to identify a highly drought-stress-responsive AP2 gene, i.e., ZmEREB24. Compared to the wild type, the overexpression of ZmEREB24 in maize significantly promotes drought tolerance of transgenic plants at the seedling stage. CRISPR/Cas9-based ZmEREB24-knockout mutants showed a drought-sensitive phenotype. RNA-seq analysis and EMSA assay revealed AATGG.CT and GTG.T.GCC motifs as the main binding sites of ZmEREB24 to the promoters of downstream target genes. DAP-seq identified four novel target genes involved in proline and sugar metabolism and hormone signal transduction of ZmEREB24. Our data indicate that ZmEREB24 plays important biological functions in regulating drought tolerance by binding to the promoters of drought stress genes and modulating their expression. The results further suggest a role of ZmEREB24 in regulating drought adaptation in maize, indicating its potential importance for employing molecular breeding in the development of high-yield drought-tolerant maize cultivars.

ZmGI2 regulates flowering time through multiple flower development pathways in maize.

GIGANTEA (GI) encodes a component of the circadian clock core oscillator and has been identified as a regulatory pathway of the circadian rhythm and photoperiodic flowering in model plants. However, the regulatory pathway of GI affecting flowering time is unknown in maize. Here, we identified that the zmgi2 mutant flowered earlier than the wild type under long day (LD) conditions, whereas the difference in flowering time was not apparent under short day (SD) conditions. The 24 h optimal expression of the gene in the stem apex meristems (SAM) appeared at 9 h after dawn under LD conditions and at 11 h after dawn under SD conditions. DAP-Seq and RNA-Seq further revealed that ZmGI2 delays flowering by directly binding to the upstream regions of ZmVOZs, ZmZCN8 and ZmFPF1 to repress the expression of these genes and by directly binding to the upstream regions of ZmARR11, ZmDOF and ZmUBC11 to promote the expression of these genes. The genetic and biochemical evidence suggests a model for the potential role of ZmGI2 in regulating the flowering time-dependent photoperiodic pathway. This study provides novel insights into the function of ZmGIs in maize and further demonstrates their potential importance for floral transition. These results contribute to a comprehensive understanding of the molecular mechanisms and regulatory networks of GI transcription factors in regulating flowering time in maize.

Functional analysis of a late embryogenesis abundant protein ZmNHL1 in maize under drought stress.

Maize is an important feed and industrial cereal crop and is crucial for global food security. The development of drought-tolerant genotypes is a major aim of breeding programs to fight water scarcity and maintain sustainable maize production. Late embryogenesis abundant (LEA) proteins are a family of proteins related to osmotic regulation that widely exist in organisms. Here, we implemented a previously generated maize transcriptomic dataset to identify a drought-responsive gene designated ZmNHL1. Bioinformatics analysis of ZmNHL1 showed that the protein encoded by ZmNHL1 belongs to the LEA-2 protein family. Tissue specific expression analysis showed that ZmNHL1 is relatively abundant in stems and leaves, highly expressed in tassels and only slightly expressed in roots, pollens and ears. Moreover, the activity of SOD and POD of plants from three 35S::ZmNHL1 transgenic lines under either the induced drought stress conditions (by 20% PEG6000) or the natural water deficit treatment (by water withholding) were higher than that of the WT plants, while the electrolyte leakage of the 35S::ZmNHL1 transgenic plants was lower than that of the WT plants under both drought treatments. Our data further revealed that ZmNHL1 promotes maize tolerance to drought stress in 35S::ZmNHL1 transgenic plants by improving ROS scavenging and maintaining the cell membrane permeability. Overall, our data revealed that ZmNHL1 promotes maize tolerance to drought stress and contributes to provide elite germplasm resources for maize drought tolerance breeding programs.

A Nucleoporin NUP58 modulates responses to drought and salt stress in maize (Zea mays L.).

Nuclear pore complex (NUP) is the main transport channel between cytoplasm and nucleoplasm, which plays an important role in stress response. The function of NUPs was widely reported in yeast and vertebrate but rarely in plants. Here, we identified a nuclear pore complex (ZmNUP58), that is tightly associated with drought and salt tolerance phenotype accompanied with phenotypic and physiological changes under drought and salt stress. The overexpression of ZmNUP58 in maize (Zea mays L.) significantly promotes both chlorophyll content and activities of antioxidant enzymes under drought- and salt-stressed conditions. RNA-Seq analysis showed that ZmNUP58 could regulate the expression of genes related to phytohormone synthesis and signaling, osmotic adjustment substances, antioxidant enzyme system, cell wall biosynthesis, glucose metabolism and aquaporin. The results provide novel insights into the regulatory role of ZmNUP58 in improving drought and salt tolerance through regulating phytohormone and other stress response genes in maize.

Fine Mapping and Cloning of a Major QTL qph12, Which Simultaneously Affects the Plant Height, Panicle Length, Spikelet Number and Yield in Rice (Oryza sativa L.).

Plant height is one of the most important agronomical traits in rice (Oryza sativa L.). Introducing the semidwarf rice in the 1960s significantly enhanced the rice yield potential in Asia. Implementing near-isogenic lines (NILs) is the most powerful tool for the identification and fine mapping of quantitative trait loci (QTLs). In this study, 176 NILs were produced from the crossing and back-crossing of two rice cultivars. Specifically, the indica rice cultivar Jiafuzhan served as a recipient, and the restorer japonica cultivar Hui1586 served as a donor. Using the 176 NILs, we identified a novel major QTL for reduced plant height in the NIL36 line. The qph12 QTL was mapped to a 31 kb genomic region between the indel markers Indel12-29 and Indel12-31. The rice genome annotation indicated the presence of three candidate genes in this genomic region. Through gene prediction and cDNA sequencing, we confirmed that LOC_Os12g40890 (qPH12) is the target gene in the NIL36 line. Further analysis showed that the qph12 QTL is caused by a 1 bp deletion in the first exon that resulted in premature termination of the qPH12. Knockout experiments showed that the qph12 QTL is responsible for the reduced plant height phenotype of the NIL36 line. Although the qph12 gene from the NIL36 line showed a shorter panicle length, fewer spikelets per panicle and a lower plant grain yield, the plant also exhibited a lower plant height. Taken together, our results revealed that the qph12 have good specific application prospects in future rice breeding.

Comparative transcriptomic and epigenetic analyses reveal conserved and divergent regulatory pathways in barley response to temperature stresses.

DNA methylation and histone modification enable plants to rapidly adapt to adverse temperature stresses, including low temperature (LT) and high temperature (HT) stress. In this study, we conducted physiological, epigenetic, and transcriptomic analyses of barley seedlings grown under control (22°C), mild low temperature (MLT, 14°C) and HT (38°C) conditions to elucidate the underlying molecular mechanisms. Compared to MLT, HT implies greater deleterious effects on barley seedlings' growth. The methylation-sensitive amplification polymorphism analysis showed that MLT induced more DNA methylation and HT more DNA demethylation compared to control. Besides, the higher levels of H3K9ac and H3K4me3 under HT compared to MLT stresses might lead to the loosening of chromatin and, subsequently, the activation of gene expression. Consistently, the transcriptome analysis revealed that there were more differentially expressed genes (DEGs) in plants subjected to HT stress than MLT stress compared to control. The common and unique pathways of these DEGs between MLT and HT were also analyzed. Transcription factors, such as ERF, bHLH, NAC, HSF, and MYB, were most involved in MLT and HT stress. The underlying gene regulation networks of epigenetic modulation-related genes were further explored by weight gene co-expression network analysis. Our study provides new insights into the understanding of epigenetic regulation responses to temperature stress in barley, which will lead to improved strategies for the development of cold- and heat-tolerant barley varieties for sustainable barley production in a climate-changing world.

HvbZIP21, a Novel Transcription Factor From Wild Barley Confers Drought Tolerance by Modulating ROS Scavenging.

Drought stress is a common environmental stress, which adversely affects the yield and quality of crops. Due to its excellent drought tolerance, wild barley from the Middle East region is considered a valuable source for barley improvement. Here, we compared the growth rate, stomatal regulation and capacity to metabolize reactive oxygen species (ROS) of two barley cultivars and one wild barley accession. The results indicated the wild barley EC_S1 showed a more significant decline in stomatal aperture and less ROS production. Transcriptomic analysis revealed that EC_S1 has slower transcriptional regulation (5,050 DEGs) in the early stage of drought stress (14 days) than Baudin (7,022 DEGs) and Tadmor (6,090 DEGs). In addition, 30 hub genes, including nine known drought-related genes were identified by WGCNA analysis. Then, we cloned a novel bZIP transcription factor, HvbZIP21, from EC_S1. HvbZIP21 was subcellularly targeted to the nucleus. Overexpression of HvbZIP21 in Arabidopsis enhanced drought tolerance due to increasing activities of superoxide dismutase, peroxidase, and catalase activities as well as glutathione content. Silencing of HvbZIP21 in EC_S1 suppressed drought tolerance in BSMV:HvbZIP21-inoculated plants. Taken together, our findings suggest that HvbZIP21 play a critical role in drought tolerance by manipulating ROS scavenging.

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

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

ZmERF21 directly regulates hormone signaling and stress-responsive gene expression to influence drought tolerance in maize seedlings.

Drought stress adversely impacts crop development and yield. Maize frequently encounters drought stress during its life cycle. Improvement of drought tolerance is a priority of maize breeding programs. Here, we identified a novel transcription factor encoding gene, APETALA2 (AP2)/Ethylene response factor (ERF), which is tightly associated with drought tolerance in maize seedlings. ZmERF21 is mainly expressed in the root and leaf and it can be highly induced by polyethylene glycol treatment. Genetic analysis showed that the zmerf21 mutant plants displayed a reduced drought tolerance phenotype, accompanied by phenotypical and physiological changes that are commonly observed in drought conditions. Overexpression of ZmERF21 in maize significantly increased the chlorophyll content and activities of antioxidant enzymes under drought conditions. RNA-Seq and DNA affinity purification sequencing analysis further revealed that ZmERF21 may directly regulate the expression of genes related to hormone (ethylene, abscisic acid) and Ca signaling as well as other stress-response genes through binding to the promoters of potential target genes. Our results thereby provided molecular evidence of ZmERF21 is involved in the drought stress response of maize.

Aerenchyma formation in the root of leaf-vegetable sweet potato: Programmed cell death initiated by ethylene-mediated H2 O2 accumulation.

Sweet potato, commonly planted in Southeast Asia and South America with abundant rainfall, often suffers from waterlogging. The aerenchyma formation in roots is an effective way for plants to facilitate gas exchange. In the present study, tolerant and sensitive varieties, respectively, designated NC1 and C211, were evaluated under water oxygen content at 2.0 mg·L-1 (hypoxia treatment) and 8.0 mg·L-1 (control). The results showed that NC1 variety has a relatively higher root growth rate under low oxygen condition. In NC1 plants, aerenchyma was observed in the mid-section of the main adventitious root and spread to the proximal and distal ends, forming a complete channel in the cortex. However, in C211 plants, the aerenchyma occurred relatively later and could not turn into a whole channel. Ethylene synthesis-related (ACS1, ACS4, ACS5, etc.) and signal transduction-related (ETR1, ERS1, EIN2, etc.) genes were upregulated in the NC1 plants and led to changes in the reactive oxygen species-related genes (RBOHA, SOD, CAT, etc.) and enzyme activities. It was found that programmed cell death was induced by H2 O2 accumulation. A regulatory model of lysigenous aerenchyma formation in the root of sweet potato was constructed. Our study enriches the understanding of the mechanisms of the aerenchyma formation in plants.

Understanding a Mechanistic Basis of ABA Involvement in Plant Adaptation to Soil Flooding: The Current Standing.

Soil flooding severely impairs agricultural crop production. Plants can cope with flooding conditions by embracing an orchestrated set of morphological adaptations and physiological adjustments that are regulated by the elaborated hormonal signaling network. The most prominent of these hormones is ethylene, which has been firmly established as a critical signal in flooding tolerance. ABA (abscisic acid) is also known as a "stress hormone" that modulates various responses to abiotic stresses; however, its role in flooding tolerance remains much less established. Here, we discuss the progress made in the elucidation of morphological adaptations regulated by ABA and its crosstalk with other phytohormones under flooding conditions in model plants and agriculturally important crops.

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

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

Identification of ZmNF-YC2 and its regulatory network for maize flowering time.

Flowering time is an important agronomic trait that determines the distribution and adaptation of plants. The accurate prediction of flowering time in elite germplasm is critical for maize breeding. However, the molecular mechanisms underlying the photoperiod response remain elusive in maize. Here we cloned the flowering time-controlling gene, ZmNF-YC2, by map-based cloning and confirmed that ZmNF-YC2 is the nuclear transcription factor Y subunit C-2 protein and a positive regulator of flowering time in maize under long-day conditions. Our results show that ZmNF-YC2 promotes the expression of ZmNF-YA3. ZmNF-YA3 negatively regulates the transcription of ZmAP2. ZmAP2 suppresses the expression of ZMM4 to delay flowering time. We then developed a gene regulatory model of flowering time in maize using ZmNF-YC2, ZmNF-YA3, ZmAP2, ZMM4, and other key genes. The cascading regulation by ZmNF-YC2 of maize flowering time has not been reported in other species.

Conservation and divergence of the TaSOS1 gene family in salt stress response in wheat (Triticum aestivum L.).

Salinity is one of the most important problems that adversely affect crops growth, productivity and quality worldwide. Salt Overly Sensitive 1 (SOS1) gene family plays vital roles in plant response to salt stress. Herein, we report the identification of the SOS family in wheat and the exploration of the expression profiles of SOSs under salt stress. Complete genome sequences of T. aestivum were downloaded from Ensembl plant database. Conservation and divergence of TaSOS1 family were conducted by using phylogenetic tree, gene structure and synteny distribution analysis. Expression profiles of TaSOS1s were obtained based on transcriptome and qRT-PCR analysis. Totally, 119 TaSOS1 proteins in wheat were identified at the genome-wide level and classified into three groups. Six motifs were conserved in TaSOS1 gene family. Moreover, 25 TaSOS1 genes had three copies distributing in three sub-genomes (A, B and D). A total of 32, 28 and 29 TaSOS1 genes were located on the sub-genomes A, B and D, respectively. Moreover, there were 19, 12, 6, 7, 28, 5 and 12 genes located on the three homologous of chromosomes 1, 2, 3, 4, 5, 6 and 7, respectively. Two genes were mapped to unattributed scaffolds. The duplication events analysis indicated that tandem repeats contributed to the expansion of the SOS1 family in wheat. Collinearity analysis demonstrated that segmental duplications play an important role in the expansion of SOS1 members. Chromosome 7, 5, 3, and 2 showed collinear relationship. Tissue specific expression pattern analysis revealed that 41 TaSOS1 genes expressed in various tissues, such as root, shoot, leaf, spike and grain. Transcriptomic analysis revealed that 28 and 26 genes were up- and down-regulated under salinity stress, respectively, of which 18 genes were further confirmed by RT-qPCR. The plants with high expression level of these genes displayed higher tolerance to salinity stress, stronger root system, higher Fv/Fm value and water potential. The results could be helpful for further elucidating the molecular mechanism of TaSOS1 related to salt tolerance in wheat and provide a toolkit for improving the salinity tolerance of wheat. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s12298-021-01009-y.

Identification and Validation of Major QTLs, Epistatic Interactions, and Candidate Genes for Soybean Seed Shape and Weight Using Two Related RIL Populations.

Understanding the genetic mechanism underlying seed size, shape, and weight is essential for enhancing soybean cultivars. High-density genetic maps of two recombinant inbred line (RIL) populations, LM6 and ZM6, were evaluated across multiple environments to identify and validate M-QTLs as well as identify candidate genes behind major and stable quantitative trait loci (QTLs). A total of 239 and 43 M-QTLs were mapped by composite interval mapping (CIM) and mixed-model-based composite interval mapping (MCIM) approaches, from which 180 and 18, respectively, are novel QTLs. Twenty-two QTLs including four novel major QTLs were validated in the two RIL populations across multiple environments. Moreover, 18 QTLs showed significant AE effects, and 40 pairwise of the identified QTLs exhibited digenic epistatic effects. Thirty-four QTLs associated with seed flatness index (FI) were identified and reported here for the first time. Seven QTL clusters comprising several QTLs for seed size, shape, and weight on genomic regions of chromosomes 3, 4, 5, 7, 9, 17, and 19 were identified. Gene annotations, gene ontology (GO) enrichment, and RNA-seq analyses of the genomic regions of those seven QTL clusters identified 47 candidate genes for seed-related traits. These genes are highly expressed in seed-related tissues and nodules, which might be deemed as potential candidate genes regulating the seed size, weight, and shape traits in soybean. This study provides detailed information on the genetic basis of the studied traits and candidate genes that could be efficiently implemented by soybean breeders for fine mapping and gene cloning, and for marker-assisted selection (MAS) targeted at improving these traits individually or concurrently.

Recycling of beet sugar byproducts and wastes enhances sugar beet productivity and salt redistribution in saline soils.

Soil salinity adversely affects the growth, yield, and quality parameters of sugar beet, leading to a reduction in root and sugar yields. Improving the physical and chemical properties of salt-affected soils is essential for sustainable cultivation and sugar beet production. A field experiment was conducted at the Delta Sugar Company Research Farm, El-Hamool, Kafr El-Sheikh, Egypt, to evaluate the response of sugar beet to the application of beet sugar filter cake treated with sulfuric and phosphoric acid-treated, phosphogypsum (PG), desaline, humic acid, and molasses under saline soil conditions. The application of treated filter cake enhanced root length, diameter, and leaf area. The application of molasses enhanced root length, diameter, and leaf area as well. Application of molasses increased sugar content and root yield. The application of either treated filter cake or molasses produced the highest recoverable sugar yield. Linear regression analysis revealed that the root yield, quality index, and recoverable sugar yield increased in response to the increased availability of either Ca2+ or K content in the soil which increases in response to the application of soil amendments and molasses. The application of treated beet sugar filter cake and molasses increased the calcium, magnesium, and potassium availability in the soil. Treated filter cake is a promising organic soil amendment that enhanced the yield by 29% and yield-related traits of sugar beet by improving the physical and chemical properties of the soil.

Application of beet sugar byproducts improves sugar beet biofortification in saline soils and reduces sugar losses in beet sugar processing.

Improving the chemical and physical properties of saline soils is crucial for the sustainable production of sugar beet and efficient processing of beet sugar. Here, the impacts of the application of treated filter cake on sugar beet biofortification under saline soil and sugar losses into molasses during beet sugar processing were evaluated for the first time. The application of treated filter cake significantly reduced K%, Na%, and α-amino-N while enhanced sucrose content and quality index of beet root juice. Consequently, sugar loss percentage, sugar loss yield, and relative sugar loss yield were reduced, whereas recoverable sugar yield was enhanced. Linear regression analysis revealed that quality index and sugar loss yield were increased, whereas sugar loss percentage and relative sugar loss yield were reduced in response to the reduction of soil Na+ content accompanied with increasing Ca2+ content in the soil increased. The results provide treated filter cake as a promising amendment for saline soil remediation for improving biofortification of sugar beet and reducing sugar losses during beet sugar processing.

Exploring genetic architecture for pod-related traits in soybean using image-based phenotyping.

Mature pod color (PC) and pod size (PS) served as important characteristics are used in the soybean breeding programs. However, manual phenotyping of such complex traits is time-consuming, laborious, and expensive for breeders. Here, we collected pod images from two different populations, namely, a soybean association panel (SAP) consisting of 187 accessions and an inter-specific recombinant inbred line (RIL) population containing 284 RILs. An image-based phenotyping method was developed and used to extract the pod color- and size-related parameters from images. Genome-wide association study (GWAS) and linkage mapping were performed to decipher the genetic control of pod color- and size-related traits across 2 successive years. Both populations exhibited wide phenotypic variations and continuous distribution in pod color- and size-related traits, indicating quantitative polygenic inheritance of these traits. GWAS and linkage mapping approaches identified the two major quantitative trait loci (QTL) underlying the pod color parameters, i.e., qPC3 and qPC19, located to chromosomes 3 and 19, respectively, and 12 stable QTLs for pod size-related traits across nine chromosomes. Several genes residing within the genomic region of stable QTL were identified as potential candidates underlying these pod-related traits based on the gene annotation and expression profiling data. Our results provide the useful information for fine-mapping/map-based cloning of QTL and marker-assisted selection of elite varieties with desirable pod traits. Supplementary Information: The online version contains supplementary material available at 10.1007/s11032-021-01223-2.

Genotypic variation in the grain photosynthetic contribution to grain filling in rice.

Grain filling in rice, a staple cereal crop worldwide, is a critical determinant of grain yield and quality. However, there is little available information on the relationship between grain filling and grain photosynthetic capacity in rice. This study evaluated the genetic diversity among six rice cultivars for their grain filling rate (GR0) and the relationships with the grain chlorophyll contents and grain net photosynthetic rate (PN). Significant variations in GR0, PN, and the chlorophyll contents (a, b, and total) in the grains of the cultivars were observed. Approximately 90 % of the variation in GR0 was explained by the grain PN. General linear model regression revealed significant positive correlations between PN/GR0 and the chlorophyll contents (a, b, and total) in the grains. There was also a significant positive correlation between PN and GR0. These positive correlations suggest a direct positive relationship between the grain filling rate and grain chlorophyll contents, which is indicative of the high photosynthetic capacity of the grains during the early grain filling period. These results suggest that the grain chlorophyll contents could be used as a molecular marker in marker-assisted breeding programs for rice cultivars with high grain net photosynthetic capacity during the early period of grain filling to improve grain yield.