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Item Trans-QTL alliance of HKT1 and PHL7 modulate salinity stress tolerance and enhance crop yield endurance(John Wiley & Sons, 2026) Mohanty, Jitendra K.; Yadav, Antima; Narnoliya, Laxmi; Thakro, Virevol; Rathore, Deepanshi; Tripathi, Shailesh; Sinharoy, Senjuti; Agarwal, Pinky; Parida, Swarup K.Salinity stress can cause significant yield losses in crops because of its major impact on reproductive success. The complexity of salinity stress responses, particularly their tissue- and cell-specific regulation, continues to challenge the translation of molecular insights into tangible crop yield improvements. In the present study, the authors deployed a genomic strategy combining a genome-wide association study, regional association analysis, QTL mapping, fine mapping, and map-based cloning to delineate a pair of novel CaPHL7 and CaHKT1 alleles that regulate yield under salinity stress. The selected contrasting accessions, developed near-isogenic lines (NILs), overexpressed chickpea lines and complemented Arabidopsis lines collectively underscore the functional significance of the identified alleles in relaying yield endurance under salinity stress conditions. Functional characterisation of the genes revealed the intricate transcriptional regulation of CaHKT1 by CaPHL7, which influences the degree of salinity stress tolerance. Furthermore, in our efforts to enhance yield endurance, we discovered a novel regulatory role for the phosphorus (P) starvation-responsive gene (PHL7) in legumes, facilitating salinity stress adaptation. This study provides the first functional validation of a trans-QTL regulatory model in chickpea, where CaPHL7, located on one chromosome, transcriptionally activates CaHKT1 on a separate chromosome. The regulatory mechanism plays a key role in excluding sodium from the transpiration stream, thereby protecting reproductive processes from salinity-induced damage and mitigating yield penalties. This inter-locus regulation explains yield stability and offers useful insights that may be considered in future efforts to enhance salt resilience in chickpea.Item Method for the dissection of genomic loci associated with chickpea root penetration traits in compact soil(Springer Nature Publishing AG, 2026) Ganotra, Jahanvi; Pandey, Mandavi; Donde, Ravindra; Giri, JitenderMechanical impedance in agricultural land is a significant constraint in modern agriculture. It dramatically affects seed germination, plant growth, development, and grain yield. Soil compaction hinders root growth and the ability to access deeper nutrients and water resources, impacting climate resilience, crop productivity, and global food security. Crops display variations in root system architecture (RSA) traits when grown in compacted soils. We can better understand the mechanisms behind soil compaction by examining root-related traits and their associated genes. Our recently published study investigated RSA traits across different soil compaction levels and identified significant genomic associations in chickpeas. We developed reliable methods for creating soils with varying bulk densities (i.e., compaction levels), growing chickpea seedlings, and harvesting the roots. We also conducted high-throughput phenotyping and screening of root-related traits using winRHIZO software. By integrating these phenotypic data with available genotypic data through Genome-Wide Association Studies (GWAS), we could identify genetic loci influencing root penetration in response to increasing soil compaction. These methods will help us identify key architectural traits of roots that can be targeted in crop breeding efforts to enhance resilience and productivity in compacted soils. By improving the root system and understanding the genes involved, we aim to develop plants more responsive to root penetration.Item Genome-wide association analysis using multi model approach identified novel associations for plant architectural traits in chickpea germplasm collection(Springer Nature Publishing AG, 2026) Raiya, Rahul; Yadav, Hemant Kumar; Kumar, Kuldeep; Prakash, Nitish Ranjan; Parida, Swarup K.; Bharadwaj, Chellapilla; Hegde, Venkatraman; Tripathi, ShaileshCrop plant architecture dictates plant performance under different ecological conditions and is responsible for its establishment, development, and morphology. It plays an important role in plant breeding for yield optimization, regulating photosynthetic rate and efficiency, utilization of resources, occurrence of pests and diseases, effective mechanical harvesting, space optimization, and improving the quality of plant produce. The subtle changes in the plant’s architecture could help the plant adapt to different ecological niches and are very important, keeping in view the challenges posed due to climate change. A set of 280 diverse genotypes, which included the core collection of chickpea was evaluated during the rabi season of 2021 and 2022. A total of 10 plant architecture related traits, including plant height, first pod height, canopy width, inter-nodal length, and days to flowering were studied and significant variability was observed as per the analysis of variance (ANOVA) and phenotypic descriptors. Diversity based on π and θ estimates suggests the presence of substantial diversity, while Tajima’s D reflects balancing selection due to the abundance of shared alleles. Significant marker trait associations (MTAs) for traits like plant height (PH), first pod height (FDPH) and days to flowering were observed using trait based or BLUP estimates. In total, 97 and 51 MTAs were identified using trait based and BLUP based on multi model GWAS analysis, respectively. Among these 17 were consistent MTAs being present either across the year or were identified using more than one GWAS model. Likewise, 9 consistent MTAs were identified using the BLUP estimates. Interestingly, two genomic regions present on chromosome 5 and 7 were found to harbor multiple MTAs for PH and FPDH. The linkage disequilibrium (LD) block analysis reflects the prevalence of multiple LD blocks in these regions. The allelic effects of the MTAs reflect their additive nature in determining the phenotype. Overall, the MTAs identified in the current study are highly useful for the chickpea breeder in modulating the plant architecture, mainly PH and FPDH.Item A next-generation combinatorial genomic strategy scans genomic loci governing heat stress tolerance in chickpea(John Wiley & Sons, 2025) Mohanty, Jitendra K.; Yadav, Antima; Narnoliya, Laxmi; Thakro, Virevol; Nayyar, Harsh; Dixit, Girish P.; Jha, Uday Chand; Prasad, P. V. Vara; Agarwal, Pinky; Parida, Swarup K.In the wake of rising earth temperature, chickpea crop production is haunted by the productivity crisis. Chickpea, a cool season legume manifests tolerance in several agro-physiological level, which is complex quantitative in nature, and regulated by multiple genes and genetic networks. Understanding the molecular genetic basis of this tolerance and identifying key regulators can leverage chickpea breeding against heat stress. This study employed a genomics-assisted breeding strategy utilizing multi-locus GWAS to identify 10 key genomic regions linked to traits contributing to heat stress tolerance in chickpea. These loci subsequently delineated few key candidates and hub regulatory genes, such as RAD23b, CIPK25, AAE19, CK1 and WRKY40, through integrated genomics, transcriptomics and interactive analyses. The differential transcript accumulation of these identified candidates in contrasting chickpea accessions suggests their potential role in heat stress tolerance. Differential ROS accumulation along with their scavengers' transcript abundance aligning with the expression of identified candidates in the contrasting chickpea accessions persuade their regulatory significance. Additionally, their functional significance is ascertained by heterologous expression and subsequent heat stress screening. The high confidence genomic loci and the superior genes and natural alleles delineated here has great potential for swift genomic interventions to enhance heat resilience and yield stability in chickpea.Item Transcriptome-wide association mapping provides insights into the genetic basis and candidate genes governing flowering, maturity and seed weight in rice bean (Vigna umbellata)(BioMed Central Ltd, 2024) Sahu, Tanmaya Kumar; Verma, Sachin Kumar; Gayacharan; Singh, Nagendra Pratap; Joshi, Dinesh Chandra; Wankhede, D. P.; Singh, Mohar; Bhardwaj, Rakesh; Singh, Badal; Parida, Swarup K.; Chattopadhyay, Debasis; Singh, Gyanendra Pratap; Singh, Amit KumarBackground: Rice bean (Vigna umbellata), an underrated legume, adapts to diverse climatic conditions with the potential to support food and nutritional security worldwide. It is used as a vegetable, minor food crop and a fodder crop, being a rich source of proteins, minerals, and essential fatty acids. However, little effort has been made to decipher the genetic and molecular basis of various useful traits in this crop. Therefore, we considered three economically important traits i.e., flowering, maturity and seed weight of rice bean and identified the associated candidate genes employing an associative transcriptomics approach on 100 diverse genotypes out of 1800 evaluated rice bean accessions from the Indian National Genebank. Results: The transcriptomics-based genotyping of one-hundred diverse rice bean cultivars followed by pre-processing of genotypic data resulted in 49,271 filtered markers. The STRUCTURE, PCA and Neighbor-Joining clustering of 100 genotypes revealed three putative sub-populations. The marker-trait association analysis involving various genome-wide association study (GWAS) models revealed significant association of 82 markers on 48 transcripts for flowering, 26 markers on 22 transcripts for maturity and 22 markers on 21 transcripts for seed weight. The transcript annotation provided information on the putative candidate genes for the considered traits. The candidate genes identified for flowering include HSC80, P-II PsbX, phospholipid-transporting-ATPase-9, pectin-acetylesterase-8 and E3-ubiquitin-protein-ligase-RHG1A. Further, the WRKY1 and DEAD-box-RH27 were found to be associated with seed weight. Furthermore, the associations of PIF3 and pentatricopeptide-repeat-containing-gene with maturity and seed weight, and aldo–keto-reductase with flowering and maturity were revealed. Conclusion: This study offers insights into the genetic basis of key agronomic traits in rice bean, including flowering, maturity, and seed weight. The identified markers and associated candidate genes provide valuable resources for future exploration and targeted breeding, aiming to enhance the agronomic performance of rice bean cultivars. Notably, this research represents the first transcriptome-wide association study in pulse crop, uncovering the candidate genes for agronomically useful traits.Item Dissecting chickpea genomic loci associated with the root penetration responsive traits in compacted soil(Springer Nature Publishing AG, 2024) Donde, Ravindra; Kohli, Pawandeep Singh; Pandey, Mandavi; Sirohi, Ujjwal; Singh, Bhagat; Giri, JitenderSoil compaction is a major concern for modern agriculture, as it constrains plant root growth, leading to reduced resource acquisition. Phenotypic variation for root system architecture (RSA) traits in compacted soils is present for various crops; however, studies on genetic associations with these traits are lacking. Therefore, we investigated RSA traits in diferent soil compaction levels and identifed signifcant genomic associations in chickpea. We conducted a Genome-Wide Association Study (GWAS) of 210 chickpea accessions for 13 RSA traits under three bulk densities (BD) (1.1BD, 1.6BD, and 1.8BD). Soil compaction decreases root exploration by reducing 12 RSA traits, except average diameter (AD). Further, AD is negatively correlated with lateral root traits, and this correlation increases in 1.8BD, suggesting the negative efect of AD on lateral root traits. Interestingly, we identifed probable candidate genes such as GLP3 and LRX for lateral root traits and CRF1-like for total length (TL) in 1.6BD soil. In heavy soil compaction, DGK2 is associated with lateral root traits. Reduction in laterals during soil compaction is mainly due to delayed seedling establishment, thus making lateral root number a critical trait. Interestingly, we also found a higher contribution of the GxE component of the number of root tips (Tips) to the total variation than the other lateral traits. We also identifed a pectin esterase, PPE8B, associated with Tips in high soil compaction and a signifcantly associated SNP with the relative change in Tips depicting a trade-of between Tips and AD. Identifed genes and loci would help develop soil-compaction-resistant chickpea varieties.Item Natural alleles of Mediator subunit genes modulate plant height in chickpea(John Wiley & Sons, 2023) Malik, Naveen; Basu, Udita; Srivastava, Rishi; Daware, Anurag; Ranjan, Rajeev; Sharma, Akash; Thakro, Virevol; Mohanty, Jitendra K.; Jha, Uday Chand; Tripathi, Shailesh; Tyagi, Akhilesh K.; Parida, Swarup K.Plant height (PH) is an important plant architectural trait targeted during Green Revolution to enhance crop yields. Identification of genes and natural alleles governing plant height without compromising agronomic performance can fill the lacuna of knowledge connecting ideal plant architecture with maximum achievable yield in chickpea. Through coherent strategy involving genome-wide association study, QTL/fine mapping, map-based cloning, molecular haplotyping, and downstream functional genomics, the current study identified two Mediator subunit genes namely, CaMED23 and CaMED5b and their derived natural alleles/haplotypes underlying the major QTLs and trans-acting eQTLs regulating plant height in chickpea. Differential accumulation of haplotype-specific transcripts of these two Mediator genes in corresponding haplotype-introgressed near-isogenic lines (NILs) correlates negatively with the plant height trait. Quantitative as well as qualitative estimation based on histology, scanning electron microscopy, and histochemical assay unraveled the reduced lengths and cell sizes of internodes along with compromised lignin levels in dwarf/semi-dwarf chickpea NILs introgressed with superior CaMED23 and CaMED5b gene haplotypes. This observation, supported by global transcriptome profiling-based diminished expression of various phenylpropanoid pathway genes upstream of lignin biosynthesis in dwarf/semi-dwarf NILs, essentially links plant height with lignin accumulation. The identified molecular signatures in the Mediator subunit genes can be efficiently utilized to develop desirable dwarf/semi-dwarf-type chickpea cultivars without affecting their yield per plant via modulating lignin/phenylpropanoid biosynthesis.Item A superior gene allele involved in abscisic acid signaling enhances drought tolerance and yield in chickpea(Oxford University Press, 2023) Thakro, Virevol; Malik, Naveen; Basu, Udita; Srivastava, Rishi; Narnoliya, Laxmi; Daware, Anurag; Varshney, Nidhi; Mohanty, Jitendra K; Bajaj, Deepak; Dwivedi, Vikas; Tripathi, Shailesh; Jha, Uday Chand; Dixit, Girish Prasad; Singh, Ashok K; Tyagi, Akhilesh K.; Upadhyaya, Hari D; Parida, Swarup K.Identifying potential molecular tags for drought tolerance is essential for achieving higher crop productivity under drought stress. We employed an integrated genomics-assisted breeding and functional genomics strategy involving association mapping, fine mapping, map-based cloning, molecular haplotyping and transcript profiling in the introgression lines (ILs)- and near isogenic lines (NILs)-based association panel and mapping population of chickpea (Cicer arietinum). This combinatorial approach delineated a bHLH (basic helix-loop-helix) transcription factor, CabHLH10 (Cicer arietinum bHLH10) underlying a major QTL, along with its derived natural alleles/haplotypes governing yield traits under drought stress in chickpea. CabHLH10 binds to a cis-regulatory G-box promoter element to modulate the expression of RD22 (responsive to desiccation 22), a drought/ABA-responsive gene (via a trans-expression QTL), and two strong yield-enhancement photosynthetic efficiency (PE) genes. This, in turn, upregulates other downstream drought-responsive and abscisic acid signaling genes, as well as yield-enhancing PE genes, thus increasing plant adaptation to drought with reduced yield penalty. We showed that a superior allele of CabHLH10 introgressed into the NILs improved root and shoot biomass and PE, thereby enhancing yield and productivity during drought without compromising agronomic performance. Furthermore, overexpression of CabHLH10 in chickpea and Arabidopsis (Arabidopsis thaliana) conferred enhanced drought tolerance by improving root and shoot agro-morphological traits. These findings facilitate translational genomics for crop improvement and the development of genetically-tailored, climate-resilient, high-yielding chickpea cultivars.Item Rice Pangenome Genotyping Array: an efficient genotyping solution for pangenome-based accelerated genetic improvement in rice(John Wiley & Sons, 2022) Daware, Anurag; Malik, Ankit; Srivastava, Rishi; Das, Durdam; Ellur, Ranjith K; Singh, Ashok K; Tyagi, Akhilesh K.; Parida, Swarup K.The advent of the pangenome era has unraveled previously unknown genetic variation existing within diverse crop plants, including rice. This untapped genetic variation is believed to account for a major portion of phenotypic variation existing in crop plants. However, the use of conventional single reference-guided genotyping often fails to capture large portion of this genetic variation leading to a reference bias. This makes it difficult to identify and utilize novel population/cultivar-specific genes for crop improvement. Thus, we developed a rice pangenome genotyping array (RPGA) harboring probes assaying 80K single nucleotide polymorphisms (SNPs) and presence-absence variants (PAVs) spanning the entire 3K rice pangenome. This array provides a simple, user-friendly and cost-effective (60 to 80 USD per sample) solution for rapid pangenome-based genotyping in rice. The GWAS conducted using RPGA-SNP genotyping data of a rice diversity panel detected a total of 42 loci, including previously known as well as novel genomic loci regulating grain size/weight traits in rice. Eight of these identified trait-associated loci (dispensable loci) could not be detected with conventional single reference genome-based GWAS. A WD repeat-containing PROTEIN 12 gene underlying one of such dispensable locus on chromosome 7 (qLWR7) along with other non-dispensable loci were subsequently detected using high-resolution QTL mapping confirming authenticity of RPGA-led GWAS. This demonstrates the potential of RPGA-based genotyping to overcome reference bias. The application of RPGA-based genotyping for population structure analysis, hybridity testing, ultra-high-density genetic map construction and chromosome-level genome assembly, and marker-assisted selection was also demonstrated. A web application (http://www.rpgaweb.com) was further developed to provide easy to use platform for the imputation of RPGA-based genotyping data using 3K Rice Reference Panel and subsequent GWAS.Item GWAS identifies genetic loci underlying nitrogen responsiveness in the climate resilient C4 model Setaria italica (L.)(Elsevier B.V., 2022) Bandyopadhyay, Tirthankar; Swarbreck, Stéphanie M; Jaiswal, Vandana; Maurya, Jyoti; Gupta, Rajeev; Bentley, Alison R.; Griffiths, Howard; Prasad, ManojIntroduction N responsiveness is the capacity to perceive and induce morpho-physiological adaptation to external and internal Nitrogen (N). Crop productivity is propelled by N fertilizer and requires the breeding/selection of cultivars with intrinsically high N responsiveness. This trait has many advantages in being more meaningful in commercial/environmental context, facilitating in-season N management and not being inversely correlated with N availability over processes regulating NUE. Current lack of its understanding at the physio-genetic basis is an impediment to select for cultivars with a predictably high N response. Objectives To dissect physio-genetic basis of N responsiveness in 142 diverse population of foxtail millet, Setaria italica (L.) by employing contrasting N fertilizer nutrition regimes. Methods We phenotyped S. italica accessions for major yield related traits under low (N10, N25) and optimal (N100) growth conditions and genotyped them to subsequently perform a genome-wide association study to identify genetic loci associated with nitrogen responsiveness trait. Groups of accessions showing contrasting trait performance and allelic forms of specific linked genetic loci (showing haplotypes) were further accessed for N dependent transcript abundances of their proximal genes. Results Our study show that N dependent yield rise in S. italica is driven by grain number whose responsiveness to N availability is genetically underlined. We identify 22 unique SNP loci strongly associated with this trait out of which six exhibit haplotypes and consistent allelic variation between lines with contrasting N dependent grain number response and panicle architectures. Furthermore, differential transcript abundances of specific genes proximally linked to these SNPs in same lines is indicative of their N dependence in a genotype specific manner. Conclusion The study demonstrates the value/ potential of N responsiveness as a selection trait and identifies key genetic components underlying the trait in S. italica. This has major implications for improving crop N sustainability and food security.