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Growing chickpea (Cicer arietinum L.) faces significant challenges due to rising temperatures and drought stress, particularly during the reproductive and seed-filling phases. This study investigated the single and joint impacts of drought and heat stress on seed development, focusing on the responses of drought-tolerant (DT) and drought-sensitive (DS) chickpea genotypes. Initially raised in an outdoor environment (mean day and night temperature of 27 and 16±1 °C, respectively, light intensity of 1230–1440 µmol m−2 s−1, relative humidity of 70/43 %) until seed filling (around 110–113 days after sowing) commenced. The plants were subsequently exposed to single or combined heat and drought stress under controlled conditions until maturity. Control pots were maintained at day and night temperature of 25 and 15 °C, respectively with 500 µmol m−2 s−1 light, 60–65 % RH, and regular irrigation, and drought-stressed pots were kept at 50 % field capacity under the same conditions of light and humidity. Heat stress in pots was gradually increased to 32(day)/20 °C (night) under regular irrigation, while combined stress pots experienced both drought (50 % field capacity) and heat stress conditions 32(day)/20 °C (night) under the same light and humidity conditions with irrigation. All stress treatments adversely affected cell membranes, photosynthesis, and water regulation, with more pronounced effects under combined stress. While heat stress increased stomatal conductance, drought and combined stress significantly reduced it. Seed filling rate and duration decreased under all stress conditions, especially combined stress. The stresses in combination severely reduced seed weight and pod numbers compared to individual stresses. Enzyme activities involved in starch and sucrose synthesis and hydrolysis substantially decreased under the combined stress. Seed composition elements (starch, storage proteins, sugars, fat, crude fiber, and ash) exhibited significant reductions across all stress treatments, particularly for the combined stress. Thus, under combined stresses, starch, proteins, and soulube sugars were markedly decreased to 13–20 %, 6.4–12.4 %, and 3–5 % in seeds, compared to 37–39 %, 21–24 %, and 6 % in control seeds. The DT genotype outperformed the DS genotype for all traits under individual and combined stress conditions. Principal component analysis revealed a complex interplay among various physiological responses (membrane damage, chlorophyll, chlorophyll fluorescence, relative leaf water content, and stomatal conductance), seed yield, and seed composition under the combined stress. This study highlighted that combined heat and drought stress severely impacted chickpea yield and nutritional traits, such as seed starch and protein content, compared to individual stresses underscoring the need to develop cultivars tolerant to this stress combination.

Abstract Background Chickpea is prone to many abiotic stresses such as heat, drought, salinity, etc. which cause severe loss in yield. Tolerance towards these stresses is quantitative in nature and many studies have been done to map the loci influencing these traits in different populations using different markers. This study is an attempt to meta-analyse those reported loci projected over a high-density consensus map to provide a more accurate information on the regions influencing heat, drought, cold and salinity tolerance in chickpea. Results A meta-analysis of QTL reported to be responsible for tolerance to drought, heat, cold and salinity stress tolerance in chickpeas was done. A total of 1512 QTL responsible for the concerned abiotic stress tolerance were collected from literature, of which 1189 were projected on a chickpea consensus genetic map. The QTL meta-analysis predicted 59 MQTL spread over all 8 chromosomes, responsible for these 4 kinds of abiotic stress tolerance in chickpea. The physical locations of 23 MQTL were validated by various marker-trait associations and genome-wide association studies. Out of these reported MQTL, CaMQAST1.1, CaMQAST4.1, CaMQAST4.4, CaMQAST7.8, and CaMQAST8.2 were suggested to be useful for different breeding approaches as they were responsible for high per cent variance explained (PVE), had small intervals and encompassed a large number of originally reported QTL. Many putative candidate genes that might be responsible for directly or indirectly conferring abiotic stress tolerance were identified in the region covered by 4 major MQTL- CaMQAST1.1, CaMQAST4.4, CaMQAST7.7, and CaMQAST6.4, such as heat shock proteins, auxin and gibberellin response factors, etc. Conclusion The results of this study should be useful for the breeders and researchers to develop new chickpea varieties which are tolerant to drought, heat, cold, and salinity stresses.

Abstract Increasing temperature affects all food crops, thereby reducing their yield potential. Chickpea is a cool-season food legume vital for its nutritive value, but it is sensitive to high temperatures (> 32/20 °C maximum/minimum) during its reproductive and seed-filling stages. This study evaluated the effects of heat stress on yield and qualitative traits of chickpea seeds in a controlled environment. Chickpea genotypes differing in heat sensitivity [two heat-tolerant (HT) and two heat-sensitive (HS)] were raised in pots, initially in an outdoor environment (average 23.5/9.9 °C maximum/minimum), until the beginning of pod set (107–110 days after sowing). At this stage, the plants were moved to a controlled environment in the growth chamber to impose heat stress (32/20 °C) at the seed-filling stage, while maintaining a set of control plants at 25/15 °C. The leaves of heat-stressed plants of the HT and HS genotypes showed considerable membrane damage, altered stomatal conductance, and reduced leaf water content, chlorophyll content, chlorophyll fluorescence, and photosynthetic ability (RuBisCo, sucrose phosphate synthase, and sucrose activities) relative to their corresponding controls. Seed filling duration and seed rate drastically decreased in heat-stressed plants of the HT and HS genotypes, severely reducing seed weight plant–1 and single seed weight, especially in the HS genotypes. Yield-related traits, such as pod number, seed number, and harvest index, noticeably decreased in heat-stressed plants and more so in the HS genotypes. Seed components, such as starch, proteins, fats, minerals (Ca, P, and Fe), and storage proteins (albumin, globulins, glutelin, and prolamins), drastically declined, resulting in poor-quality seeds, particularly in the HS genotypes. These findings revealed that heat stress significantly reduced leaf sucrose production, affecting the accumulation of various seed constituents, and leading to poor nutritional quality. The HT genotypes were less affected than the HS genotypes because of the greater stability of their leaf water status and photosynthetic ability, contributing to better yield and seed quality traits in a heat-stressed environment.

Abstract Terminal drought is one of the major constraints to crop production in chickpea (Cicer arietinum L.). In order to map drought tolerance related traits at high resolution, we sequenced multi‐parent advanced generation intercross (MAGIC) population using whole genome resequencing approach and phenotyped it under drought stress environments for two consecutive years (2013–14 and 2014–15). A total of 52.02 billion clean reads containing 4.67 TB clean data were generated on the 1136 MAGIC lines and eight parental lines. Alignment of clean data on to the reference genome enabled identification of a total, 932,172 of SNPs, 35,973 insertions, and 35,726 deletions among the parental lines. A high‐density genetic map was constructed using 57,180 SNPs spanning a map distance of 1606.69 cM. Using compressed mixed linear model, genome‐wide association study (GWAS) enabled us to identify 737 markers significantly associated with days to 50% flowering, days to maturity, plant height, 100 seed weight, biomass, and harvest index. In addition to the GWAS approach, an identity‐by‐descent (IBD)‐based mixed model approach was used to map quantitative trait loci (QTLs). The IBD‐based mixed model approach detected major QTLs that were comparable to those from the GWAS analysis as well as some exclusive QTLs with smaller effects. The candidate genes like FRIGIDA and CaTIFY4b can be used for enhancing drought tolerance in chickpea. The genomic resources, genetic map, marker‐trait associations, and QTLs identified in the study are valuable resources for the chickpea community for developing climate resilient chickpeas.

Abstract Soil salinity affects various crop cultivation but legumes are the most sensitive to salinity. Osmotic stress is the first stage of salinity stress caused by excess salts in the soil on plants which adversely affects the growth instantly. The Trehalose-6-phosphate synthase (TPS) genes play a key role in the regulation of abiotic stresses resistance from the high expression of different isoform. Selected genotypes were evaluated to estimate for salt tolerance as well as genetic variability at morphological and molecular level. Allelic variations were identified in some of the selected genotypes for the TPS gene. A comprehensive analysis of the TPS gene from selected genotypes was conducted. Presence of significant genetic variability among the genotypes was found for salinity tolerance. This is the first report of allelic variation of TPS gene from chickpea and results indicates that the SNPs present in these conserved regions may contribute largely to functional distinction. The nucleotide sequence analysis suggests that the TPS gene sequences were found to be conserved among the genotypes. Some selected genotypes were evaluated to estimate for salt tolerance as well as for comparative analysis of physiological, molecular and allelic variability for salt responsive gene Trehalose-6-Phosphate Synthase through sequence similarity. Allelic variations were identified in some selected genotypes for the TPS gene. It is found that Pusa362, Pusa1103, and IG5856 are the most salt-tolerant lines and the results indicates that the identified genotypes can be used as a reliable donor for the chickpea improvement programs for salinity tolerance.

The development of genomic selection (GS) methods has allowed plant breeding programs to select favorable lines using genomic data before performing field trials. Improvements in genotyping technology have yielded high-dimensional genomic marker data which can be difficult to incorporate into statistical models. In this paper, we investigated the utility of applying dimensionality reduction (DR) methods as a pre-processing step for GS methods. We compared five DR methods and studied the trend in the prediction accuracies of each method as a function of the number of features retained. The effect of DR methods was studied using three models that involved the main effects of line, environment, marker, and the genotype by environment interactions. The methods were applied on a real data set containing 315 lines phenotyped in nine environments with 26,817 markers each. Regardless of the DR method and prediction model used, only a fraction of features was sufficient to achieve maximum correlation. Our results underline the usefulness of DR methods as a key pre-processing step in GS models to improve computational efficiency in the face of ever-increasing size of genomic data.

Pusa 391, a mega desi chickpea variety with medium maturity duration is extensively cultivated in the Central Zone of India. Of late, this variety has become susceptible to Fusarium wilt (FW), which has drastic impact on its yield. Presence of variability in the wilt causing pathogen, Fusarium oxysporum f.sp. ciceri (foc) across geographical locations necessitates the role of pyramiding for FW resistance for different races (foc 1,2,3,4 and 5). Subsequently, the introgression lines developed in Pusa 391 genetic background were subjected to foreground selection using three SSR markers (GA16, TA 27 and TA 96) while 48 SSR markers uniformly distributed on all chromosomes, were used for background selection to observe the recovery of recurrent parent genome (RPG). BC1F1 lines with 75–85% RPG recovery were used to generate BC2F1. The plants that showed more than 90% RPG recovery in BC2F1 were used for generating BC3F1. The plants that showed more than 96% RPG recovery were selected and selfed to generate BC3F3. Multi-location evaluation of advanced introgression lines (BC2F3) in six locations for grain yield (kg/ha), days to fifty percent flowering, days to maturity, 100 seed weight and disease incidence was done. In case of disease incidence, the genotype IL1 (BGM 20211) was highly resistant to FW in Junagarh, Indore, New Delhi, Badnapur and moderately resistant at Sehore and Nandyal. GGE biplot analysis revealed that IL1(BGM20211) was the most stable genotype at Junagadh, Sehore and Nandyal. GGE biplot analysis revealed that IL1(BGM 20211) and IL4(BGM 20212) were the top performers in yield and highly stable across six environments and were nominated for Advanced Varietal Trials (AVT) of AICRP (All India Coordinated Research Project on Chickpea) in 2018–19. BGM20211 and BGM 20212 recorded 29 and 28.5% average yield gain over the recurrent parent Pusa 391, in the AVT-1 and AVT-2 over five environments. Thus, BGM20211 was identified for release and notified as Pusa Manav/Pusa Chickpea 20211 for Madhya Pradesh, Gujarat and Maharashtra, Southern Rajasthan, Bundhelkhand region of Uttar Pradesh states by the Central Sub-Committees on Crop Standards, Notification and Release of Varieties of Agricultural Crops, Ministry of Agriculture and Farmers Welfare, Government of India, for commercial cultivation in India (Gazette notification number S.O.500 (E) dt. 29-1-2021).Such pyramided lines give resilience to multiple races of fusarium wilt with added yield advantage.

Micronutrient malnutrition is a serious concern in many parts of the world; therefore, enhancing crop nutrient content is an important challenge. Chickpea (Cicer arietinum L.), a major food legume crop worldwide, is a vital source of protein and minerals in the vegetarian diet. This study evaluated a diverse set of 258 chickpea germplasm accessions for 12 key nutritional traits. A significant variation was observed for several nutritional traits, including crude protein (16.56–24.64/100 g), β-Carotene (0.003–0.104 mg/100 g), calcium (60.69–176.55 mg/100 g), and folate (0.413–6.537 mg/kg). These data, combined with the available whole-genome sequencing data for 318,644 SNPs, were used in genome-wide association studies comprising single-locus and multi-locus models. We also explored the effect of varying the minor allele frequency (MAF) levels and heterozygosity. We identified 62 significant marker-trait associations (MTAs) explaining up to 28.63% of the phenotypic variance (PV), of which nine were localized within genes regulating G protein-coupled receptor signaling pathway, proteasome assembly, intracellular signal transduction, and oxidation–reduction process, among others. The significant effect MTAs were located primarily on Ca1, Ca3, Ca4, and Ca6. Importantly, varying the level of heterozygosity was found to significantly affect the detection of associations contributing to traits of interest. We further identified seven promising accessions (ICC10399, ICC1392, ICC1710, ICC2263, ICC1431, ICC4182, and ICC16915) with superior agronomic performance and high nutritional content as potential donors for developing nutrient-rich, high-yielding chickpea varieties. Validation of the significant MTAs with higher PV could identify factors controlling the nutrient acquisition and facilitate the design of biofortified chickpeas for the future.

Chickpea yield is severely affected by drought stress, which is a complex quantitative trait regulated by multiple small-effect genes. Identifying genomic regions associated with drought tolerance component traits may increase our understanding of drought tolerance mechanisms and assist in the development of drought-tolerant varieties. Here, a total of 187 F8 recombinant inbred lines (RILs) developed from an interspecific cross between drought-tolerant genotype GPF 2 (Cicer arietinum) and drought-sensitive accession ILWC 292 (C. reticulatum) were evaluated to identify quantitative trait loci (QTLs) associated with drought tolerance component traits. A total of 21 traits, including 12 morpho-physiological traits and nine root-related traits, were studied under rainfed and irrigated conditions. Composite interval mapping identified 31 QTLs at Ludhiana and 23 QTLs at Faridkot locations for morphological and physiological traits, and seven QTLs were identified for root-related traits. QTL analysis identified eight consensus QTLs for six traits and five QTL clusters containing QTLs for multiple traits on linkage groups CaLG04 and CaLG06. The identified major QTLs and genomic regions associated with drought tolerance component traits can be introgressed into elite cultivars using genomics-assisted breeding to enhance drought tolerance in chickpea.

Measurement(s) genetic variation • SNP • Linkage Disequilibrium Technology Type(s) DNA sequencing • bioinformatics analysis • computational phylogenetic analysis Sample Characteristic - Organism Glycine soja • Glycine max Machine-accessible metadata file describing the reported data: https://doi.org/10.6084/m9.figshare.13568552

An association mapping panel consisting of 380 genotypes of chickpea was evaluated for three different years, including 2014–2015, 2015–2016 and 2016–2017, for yield-contributing parameters, including the seed number and seed weight. The AMMI analysis presented mainly concentrated on the seed weight and seed number, which are the two most important yield-contributing traits. The genotypes contributed 93.08% of the total variance, while the interaction effect was comparatively low, with 4.1% for the two traits. AMMI biplot analysis identified IG5986, IG5982, ILC6025 and ICCV14307 as desirable genotypes for the seed weight and IG5893, ILC6891 and IG5856 for the seed number. Identifying stable genotypes would help in strategic planning for yield improvement through component trait breeding.

Abstract The current pace of crop improvement is inadequate to feed the burgeoning human population by 2050. Higher, more stable, and sustainable crop production is required against a backdrop of drought stress, which causes significant losses in crop yields. Tailoring crops for drought adaptation may hold the key to address these challenges and provide resilient production systems for future harvests. Understanding the genetic and molecular landscape of the functionality of alleles associated with adaptive traits will make designer crop breeding the prospective approach for crop improvement. Here, we highlight the potential of genomics technologies combined with crop physiology for high‐throughput identification of the genetic architecture of key drought‐adaptive traits and explore innovative genomic breeding strategies for designing future crops.

The rich genetic diversity existing within exotic, indigenous, and diverse germplasm lays the foundation for the continuous improvement of crop cultivars. The composite collection has been suggested as a gateway to identifying superior germplasm for use in crop improvement programs. Here, a chickpea global composite collection was evaluated at five locations in India over two years for five agronomic traits to identify agronomically superior accessions. The desi, kabuli, and intermediate types of chickpea accessions differed significantly for plant height (PLHT) and 100-seed weight (100 SW). In contrast, the intermediate type differed substantially from kabuli for days to maturity (DM). Several highly significant trait correlations were detected across different locations. The most stable and promising accessions from each of the five locations were prioritised based on their superior performance over the best-performing check cultivar. Accordingly, the selected germplasm accessions of desi type showed up to 176% higher seed yield (SY), 29% lower flowering time, 21% fewer maturity days, 64% increase in PLHT, and 183% larger seeds than the check cultivar JG11 or Annigeri. The prioritised kabuli accessions displayed up to 270% more yield, 13% less flowering time, 8% fewer maturity days, 111% increase in PLHT, and 41% larger seeds over the check cultivar KAK2. While the intermediate type accessions had up to 169% better yield, 1% early flowering, 3% early maturity, 54% taller plants, and 25% bigger seeds over the check cultivar JG 11 or KAK2. These accessions can be utilised in chickpea improvement programs to develop high-yielding, early flowering, short duration, taller, and large-seeded varieties with a broad genetic base.

The world faces a grave situation of nutrient deficiency as a consequence of increased uptake of calorie-rich food that threaten nutritional security. More than half the world’s population is affected by different forms of malnutrition. Unhealthy diets associated with poor nutrition carry a significant risk of developing non-communicable diseases, leading to a high mortality rate. Although considerable efforts have been made in agriculture to increase nutrient content in cereals, the successes are insufficient. The number of people affected by different forms of malnutrition has not decreased much in the recent past. While legumes are an integral part of the food system and widely grown in sub-Saharan Africa and South Asia, only limited efforts have been made to increase their nutrient content in these regions. Genetic variation for a majority of nutritional traits that ensure nutritional security in adverse conditions exists in the germplasm pool of legume crops. This diversity can be utilized by selective breeding for increased nutrients in seeds. The targeted identification of precise factors related to nutritional traits and their utilization in a breeding program can help mitigate malnutrition. The principal objective of this review is to present the molecular mechanisms of nutrient acquisition, transport and metabolism to support a biofortification strategy in legume crops to contribute to addressing malnutrition.

Chickpea—the second most important grain legume worldwide—is cultivated mainly on marginal soils. Phosphorus (P) deficiency often restricts chickpea yields. Understanding the genetics of traits encoding P-acquisition efficiency and P-use efficiency will help develop strategies to reduce P-fertilizer application. A genome-wide association mapping approach was used to determine loci and genes associated with root architecture, root traits associated with P-acquisition efficiency and P-use efficiency, and any associated proxy traits. Using three statistical models—a generalized linear model (GLM), a mixed linear model (MLM), and a fixed and random model circulating probability unification (FarmCPU) —10, 51, and 40 marker-trait associations (MTAs), respectively were identified. A single nucleotide polymorphism (SNP) locus (Ca1_12310101) on Ca1 associated with three traits, i.e., physiological P-use efficiency, shoot dry weight, and shoot P content was identified. Genes related to shoot P concentration (NAD kinase 2, dynamin-related protein 1C), physiological P-use efficiency (fasciclin-like arabinogalactan protein), specific root length (4-coumarate–CoA ligase 1) and manganese concentration in mature leaves (ABC1 family protein) were identified. The MTAs and novel genes identified in this study can be used to improve P-use efficiency in chickpea.

Abstract Genetic enhancement for resistance against the pod borer, Helicoverpa armigera is crucial for enhancing production and productivity of chickpea. Here we provide some novel insights into the genetic architecture of natural variation in H. armigera resistance in chickpea, an important legume, which plays a major role in food and nutritional security. An interspecific recombinant inbred line (RIL) population developed from a cross between H. armigera susceptible accession ICC 4958 (Cicer arietinum) and resistant accession PI 489777 (Cicer reticulatum) was evaluated for H. armigera resistance component traits using detached leaf assay and under field conditions. A high‐throughput AxiomCicerSNP array was utilized to construct a dense linkage map comprising of 3,873 loci and spanning a distance of 949.27 cM. Comprehensive analyses of extensive genotyping and phenotyping data identified nine main‐effect QTLs and 955 epistatic QTLs explaining up to 42.49% and 38.05% phenotypic variance, respectively, for H. armigera resistance component traits. The main‐effect QTLs identified in this RIL population were linked with previously described genes, known to modulate resistance against lepidopteran insects in crop plants. One QTL cluster harbouring main‐effect QTLs for three H. armigera resistance component traits and explaining up to 42.49% of the phenotypic variance, was identified on CaLG03. This genomic region, after validation, may be useful to improve H. armigera resistance component traits in elite chickpea cultivars.

Abstract With an aim of enhancing drought tolerance using a marker‐assisted backcrossing (MABC) approach, we introgressed the “QTL‐hotspot” region from ICC 4958 accession that harbors quantitative trait loci (QTLs) for several drought‐tolerance related traits into three elite Indian chickpea (Cicer arietinum L.) cultivars: Pusa 372, Pusa 362, and DCP 92‐3. Of eight simple sequence repeat (SSR) markers in the QTL‐hotspot region, two to three polymorphic markers were used for foreground selection with respective cross‐combinations. A total of 47, 53, and 46 SSRs were used for background selection in case of introgression lines (ILs) developed in genetic backgrounds of Pusa 372, Pusa 362, and DCP 92‐3, respectively. In total, 61 ILs (20 BC3F3 in Pusa 372; 20 BC2F3 in Pusa 362, and 21 BC3F3 in DCP 92‐3), with >90% recurrent parent genome recovery were developed. Six improved lines in different genetic backgrounds (e.g. BGM 10216 in Pusa 372; BG 3097 and BG 4005 in Pusa 362; IPC(L4‐14), IPC(L4‐16), and IPC(L19‐1) in DCP 92‐3) showed better performance than their respective recurrent parents. BGM 10216, with 16% yield gain over Pusa 372, has been released as Pusa Chickpea 10216 by the Central Sub‐Committees on Crop Standards, Notification and Release of Varieties of Agricultural Crops, Ministry of Agriculture and Farmers Welfare, Government of India, for commercial cultivation in India. In summary, this study reports introgression of the QTL‐hotspot for enhancing yield under rainfed conditions, development of several introgression lines, and release of Pusa Chickpea 10216 developed through molecular breeding in India.

Unravelling the genetic architecture underlying yield components and agronomic traits is important for enhancing crop productivity. Here, a recombinant inbred line (RIL) population, developed from ICC 4958 and DCP 92-3 cross, was used for constructing linkage map and QTL mapping analysis. The RIL population was genotyped using a high-throughput Axiom®CicerSNP array, which enabled the development of a high-density genetic map consisting of 3,818 SNP markers and spanning a distance of 1064.14 cM. Analysis of phenotyping data for yield, yield components and agronomic traits measured across three years together with genetic mapping data led to the identification of 10 major-effect QTLs and six minor-effect QTLs explaining up to 59.70% phenotypic variance. The major-effect QTLs identified for 100-seed weight, and plant height possessed key genes, such as C3HC4 RING finger protein, pentatricopeptide repeat (PPR) protein, sugar transporter, leucine zipper protein and NADH dehydrogenase, amongst others. The gene ontology studies highlighted the role of these genes in regulating seed weight and plant height in crop plants. The identified genomic regions for yield, yield components, and agronomic traits, and the closely linked markers will help advance genetics research and breeding programs in chickpea.

After drought salinity is the major abiotic stress that severely affects agricultural productivity globally. Chickpea (Cicer arietinum L.) is the important grain legume which suffers approximately 8-10% of total global yield loss due to salinity. Screening for salt stress is difficult and traits that correlate salinity tolerance are least understood. The present study was carried out at ICAR-IARI, New Delhi 2017-18, deals with the important morphological and physiological traits like RWC (Relative water content), EL (Electrolyte Leakage), Na/K (sodium and potassium ratio) to characterize the salt tolerant genotypes under hydroponic condition which is a quick and easy method to screen large number of chickpea genotypes at initial stage under salt stress condition. Genotypes showing high RWC, low EL and Na/K ratio were tolerant like ICCV 10, JG 11, JG 62 and CSG-8962 whereas genotypes like ICC4958 and Pusa362 fall under moderately tolerant genotypes and DCP 93-3, Pusa 256, Phule G5 and SBD 377 were classified as susceptible genotypes. This study also attempts to understand the candidate genes responsible for salt-stress related pathways in chickpea genotypes based on sequence similarity approach exploiting known salt-stress responsive genes from model crops or other crop species.

Much of the world’s population growth will occur in regions where food insecurity is prevalent, with large increases in food demand projected in regions of Africa and South Asia. While improving food security in these regions will require a multi-faceted approach, improved performance of crop varieties in these regions will play a critical role. Current rates of genetic gain in breeding programs serving Africa and South Asia fall below rates achieved in other regions of the world. Given resource constraints, increased genetic gain in these regions cannot be achieved by simply expanding the size of breeding programs. New approaches to breeding are required. The Genomic Open-source Breeding informatics initiative (GOBii) and Excellence in Breeding Platform (EiB) are working with public sector breeding programs to build capacity, develop breeding strategies, and build breeding informatics capabilities to enable routine use of new technologies that can improve the efficiency of breeding programs and increase genetic gains. Simulations evaluating breeding strategies indicate cost-effective implementations of genomic selection (GS) are feasible using relatively small training sets, and proof-of-concept implementations have been validated in the International Maize and Wheat Improvement Center (CIMMYT) maize breeding program. Progress on GOBii, EiB, and implementation of GS in CIMMYT and International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) breeding programs are discussed, as well as strategies for routine implementation of GS in breeding programs serving Africa and South Asia.

Susceptibility to drought stress has restrained chickpea productivity at a global level, and the development of drought-tolerant varieties is essential to maintain its productivity. Therefore, the present study was conducted to evaluate genetic divergence in selected genotypes of chickpea and their morpho-physiological responses under irrigated and stressed conditions to identify the traits that account for the better performance of these genotypes under stressed conditions, as well as genotypes with improved drought tolerance. The genotypes were evaluated for two years under irrigated and drought stressed conditions, and significant variation was found amongst the genotypes for different morpho-physiological and yield traits. The maximum reduction was observed for plant yield (33.23%) under stressed conditions. Principle component analysis (PCA)-based biplots and correlation studies established its strong positive correlation with relative water content (RWC), membrane stability index (MSI), chlorophyll index (CI), secondary branches (SB) and yield traits and negative correlations with drought susceptibility index (DSI), days to maturity (DM) and 100 seed weight (100 SW) under drought stress, suggesting their use in selecting drought-tolerant germplasm. Ten genotypes with high values of RWC, MSI, CI, SB, yield traits and lower DSI were identified as drought-tolerant and might serve as ideal donors in the forthcoming breeding of elite chickpea cultivars. The seed-filling stage began earlier in these genotypes, with significantly reduced days to maturity under stressed conditions. Our results indicate selection for earliness offers a promising strategy for the development of drought-tolerant chickpea cultivars.

Globally, soil salinity has been on the rise owing to various factors that are both human and environmental. The abiotic stress caused by soil salinity has become one of the most damaging abiotic stresses faced by crop plants, resulting in significant yield losses. Salt stress induces physiological and morphological modifications in plants as a result of significant changes in gene expression patterns and signal transduction cascades. In this comprehensive review, with a major focus on recent advances in the field of plant molecular biology, we discuss several approaches to enhance salinity tolerance in plants comprising various classical and advanced genetic and genetic engineering approaches, genomics and genome editing technologies, and plant growth-promoting rhizobacteria (PGPR)-based approaches. Furthermore, based on recent advances in the field of epigenetics, we propose novel approaches to create and exploit heritable genome-wide epigenetic variation in crop plants to enhance salinity tolerance. Specifically, we describe the concepts and the underlying principles of epigenetic recombinant inbred lines (epiRILs) and other epigenetic variants and methods to generate them. The proposed epigenetic approaches also have the potential to create additional genetic variation by modulating meiotic crossover frequency.

The Translational Chickpea Genomics Consortium (TCGC) was set up to increase the production and productivity of chickpea (Cicer arietinum L.). It represents research institutes from six major chickpea growing states (Madhya Pradesh, Maharashtra, Andhra Pradesh, Telangana, Karnataka and Uttar Pradesh) of India. The TCGC team has been engaged in deploying modern genomics approaches in breeding and popularizing improved varieties in farmers’ fields across the states. Using marker-assisted backcrossing, introgression lines with enhanced drought tolerance and fusarium wilt resistance have been developed in the genetic background of 10 elite varieties of chickpea. Multi-location evaluation of 100 improved lines (70 desi and 30 kabuli) during 2016–2017 and 2018–2019 enabled the identification of top performing desi and kabuli lines. In total, 909 Farmer Participatory Varietal Selection trials were conducted in 158 villages in 16 districts of the five states, during 2017–2018, 2018–2019, and 2019–2020, involving 16 improved varieties. New molecular breeding lines developed in different genetic backgrounds are potential candidates for national trials under the ICAR-All India Coordinated Research Project on Chickpea. The comprehensive efforts of TCGC resulted in the development and adoption of high-yielding varieties that will increase chickpea productivity and the profitability of chickpea growing farmers.

Chickpea has a profound nutritional and economic value in vegetarian society. Continuous decline in chickpea productivity is attributed to insufficient genetic variability and different environmental stresses. Chickpea like several other legumes is highly susceptible to terminal drought stress. Multiple genes control drought tolerance and ASR gene plays a key role in regulating different plant stresses. The present study describes the molecular characterization and functional role of Abscissic acid and stress ripening (ASR) gene from chickpea (Cicer arietinum) and the gene sequence identified was submitted to NCBI Genbank (MK937569). Molecular analysis using MUSCLE software proved that the ASR nucleotide sequences in different legumes show variations at various positions though ASR genes are conserved in chickpea with only few variations. Sequence similarity of ASR gene to chickpea putative ABA/WDS induced protein mRNA clearly indicated its potential involvement in drought tolerance. Physiological screening and qRT-PCR results demonstrated increased ASR gene expression under drought stress possibly enabled genotypes to perform better under stress. Conserved domain search, protein structure analysis, prediction and validation, network analysis using Phyre2, Swiss-PDB viewer, ProSA and STRING analysis established the role of hypothetical ASR protein NP_001351739.1 in mediating drought responses. NP_001351739.1 might have enhanced the ASR gene activity as a transcription factor regulating drought stress tolerance in chickpea. This study could be useful in identification of new ASR genes that play a major role in drought tolerance and also develop functional markers for chickpea improvement.

Core Ideas Quantitative trait locus (QTL) analyses for carotenoids in chickpea were completed for three F2 populations. A moderate number of QTLs and candidate genes associated with carotenoid concentration in chickpea seeds were identified. Green cotyledon color is positively associated with provitamin A carotenoids. Three F2 populations derived from crosses between cultivars with green and yellow cotyledon colors were used to identify quantitative trait loci (QTLs) associated with carotenoid components in chickpea (Cicer arietinum L.) seeds developed by the Crop Development Centre (CDC). Carotenoids including violaxanthin, lutein, zeaxanthin, β‐cryptoxanthin, and β‐carotene were assessed in the F2:3 seeds via high‐performance liquid chromatography (HPLC). In the ‘CDC Jade’ × ‘CDC Frontier’ population, 1068 bin markers derived from the 50K Axiom CicerSNP array were mapped onto eight linkage groups (LGs). Eight QTLs, including two each for β‐carotene and zeaxanthin and one each for total carotenoids, β‐cryptoxanthin, β‐carotene, and violaxanthin were identified in this population. In the ‘CDC Cory’ × ‘CDC Jade’ population, 694 bin markers were mapped onto eight LGs and one partial LG. Quantitative trait loci for β‐cryptoxanthin, β‐carotene, violaxanthin, lutein, and total carotenoids were identified on LG8. A map with eight LGs was developed from 581 bin markers in the third population derived from the ‘ICC4475’ × ‘CDC Jade’ cross. One QTL for β‐carotene and four QTLs, one each for β‐cryptoxanthin, β‐carotene, lutein, and total carotenoids, were identified in this population. The highest phenotypic variation explained by the QTLs was for β‐carotene, which ranged from 58 to 70% in all three populations. A major gene for cotyledon color was mapped on LG8 in each population. A significant positive correlation between cotyledon color and carotenoid concentration was observed. Potential candidate genes associated with carotenoid components were obtained and their locations on the chickpea genome are presented.

Genetic diversity among 40 chickpea (Cicer arietinum L.) genotypes was investigated using 125 microsatellite (SSR, simple sequence repeat) markers. Twenty five polymorphic markers with average genetic diversity and PIC (Polymorphic Information Content) value of 0.489 and 0.437, respectively, generated a total of 90 alleles. High PIC and gene diversity (HE) values indicated good variability amongst the chickpea genotypes. Sequential Agglomerative Hierarchical Non-overlapping (SAHN) grouping revealed two main clusters with 29 genotypes in cluster I and 11 genotypes in cluster II. The Cluster analysis did not follow geographical diversity rather it was in agreement for genetic diversity with respect to seed type and parentage/pedigree. Grouping clearly delineated the diverse kabuli and desi genotypes. Molecular variance analysis also indicated 97% variation within the populations and 3% variation among the populations. Principal coordinate analysis (PCoA) divided all the 40 genotypes into three populations based on their seed type and pedigree. The 2D plot largely supported the dendrogram with similar pattern of clustering. It also indicated that the material used was diverse. Thus, the study proved that SSR markers are informative tools for assessing genetic diversity and can be recommended for characterization studies in chickpea.

Chickpea is one of the most important food legumes that holds the key to meet rising global food and nutritional demand. In order to deploy molecular breeding approaches in crop improvement programs, user friendly and cost effective marker resources remain prerequisite. The advent of next generation sequencing (NGS) technology has resulted in the generation of several thousands of markers as part of several large scale genome sequencing and re-sequencing initiatives. Very recently, PCR based Insertion-deletions (InDels) are becoming a popular gel based genotyping solution because of their co-dominant, inexpensive, and highly polymorphic nature. With an objective to expand marker resources for genomics assisted breeding (GAB) in chickpea, whole genome re-sequencing data generated on five parental lines of one interspecific (ICC 4958 × PI 489777) and two intra-specific (ICC 283 × ICC 8261 and ICC 4958 × ICC 1882) mapping populations, were used for identification of InDels. A total of 231,658 InDels were identified using Dindel software with default parameters. Further, a total of 8,307 InDels with ≥20 bp size were selected for development of gel based markers, of which primers could be designed for 7,523 (90.56%) markers. On average, markers appeared at a frequency of 1,038 InDels/LG with a maximum number of markers on CaLG04 (1,952 InDels) and minimum on CaLG08 (360 InDels). In order to validate these InDels, a total of 423 primer pairs were randomly selected and tested on the selected parental lines. A high amplification rate of 80% was observed ranging from 46.06 to 58.01% polymorphism rate across parents on 3% agarose gel. This study clearly reflects the usefulness of available sequence data for the development of genome-wide InDels in chickpea that can further contribute and accelerate a wide range of genetic and molecular breeding activities in chickpea.

[This corrects the article DOI: 10.1371/journal.pone.0096758.].

Genomic selection (GS) unlike marker-assisted backcrossing (MABC) predicts breeding values of lines using genome-wide marker profiling and allows selection of lines prior to field-phenotyping, thereby shortening the breeding cycle. A collection of 320 elite breeding lines was selected and phenotyped extensively for yield and yield related traits at two different locations (Delhi and Patancheru, India) during the crop seasons 2011-12 and 2012-13 under rainfed and irrigated conditions. In parallel, these lines were also genotyped using DArTseq platform to generate data on 3,000 polymorphic markers. Phenotypic and genotypic data were used with six statistical GS models to estimate the prediction accuracies. GS models were tested for four yield related traits viz. seed yield, 100 seed weight, days to 50% flowering and days to maturity. Prediction accuracy for the models tested varied from 0.138 (seed yield) to 0.912 (100 seed weight), whereas performance of models did not show any significant difference for estimating prediction accuracy within traits. Kinship matrix calculated using genotyping data reaffirmed existence of two different groups within selected lines. There was not much effect of population structure on prediction accuracy. In brief, present study establishes the necessary resources for deployment of GS in chickpea breeding.

To understand the genetic basis of tolerance to drought and heat stresses in chickpea, a comprehensive association mapping approach has been undertaken. Phenotypic data were generated on the reference set (300 accessions, including 211 mini-core collection accessions) for drought tolerance related root traits, heat tolerance, yield and yield component traits from 1-7 seasons and 1-3 locations in India (Patancheru, Kanpur, Bangalore) and three locations in Africa (Nairobi, Egerton in Kenya and Debre Zeit in Ethiopia). Diversity Array Technology (DArT) markers equally distributed across chickpea genome were used to determine population structure and three sub-populations were identified using admixture model in STRUCTURE. The pairwise linkage disequilibrium (LD) estimated using the squared-allele frequency correlations (r2; when r2

To estimate genetic diversity within and between 10 interfertile Cicer species (94 genotypes) from the primary, secondary and tertiary gene pool, we analysed 5,257 DArT markers and 651 KASPar SNP markers. Based on successful allele calling in the tertiary gene pool, 2,763 DArT and 624 SNP markers that are polymorphic between genotypes from the gene pools were analyzed further. STRUCTURE analyses were consistent with 3 cultivated populations, representing kabuli, desi and pea-shaped seed types, with substantial admixture among these groups, while two wild populations were observed using DArT markers. AMOVA was used to partition variance among hierarchical sets of landraces and wild species at both the geographical and species level, with 61% of the variation found between species, and 39% within species. Molecular variance among the wild species was high (39%) compared to the variation present in cultivated material (10%). Observed heterozygosity was higher in wild species than the cultivated species for each linkage group. Our results support the Fertile Crescent both as the center of domestication and diversification of chickpea. The collection used in the present study covers all the three regions of historical chickpea cultivation, with the highest diversity in the Fertile Crescent region. Shared alleles between different gene pools suggest the possibility of gene flow among these species or incomplete lineage sorting and could indicate complicated patterns of divergence and fusion of wild chickpea taxa in the past.

Kompetitive allele-specific polymerase chain reaction (KASP) assays have emerged as cost-effective marker assays especially for molecular breeding applications. Therefore, a set of 96 informative single nucleotide polymorphisms (SNPs) was used to develop KASP assays in groundnut or peanut ( spp.). Developed assays were designated as groundnut KASP assay markers (GKAMs) and screened on 94 genotypes (validation set) that included parental lines of 27 mapping populations, seven synthetic autotetraploid and amphidiploid lines, and 19 wild species accessions. As a result, 90 GKAMs could be validated and 73 GKAMs showed polymorphism in the validation set. Validated GKAMs were screened on 280 diverse genotypes of the reference set for estimating diversity features and elucidating genetic relationships. Cluster analysis of marker allelic data grouped accessions according to their genome type, subspecies, and botanical variety. The subspecies L. subsp. Waldron and subsp. formed distinct cluster; however, some overlaps were found indicating their frequent intercrossing during the course of evolution. The wild species, having diploid genomes, were grouped into a single cluster. The average polymorphism information content value for polymorphic GKAMs was 0.32 in the validation set and 0.31 in the reference set. These validated and highly informative GKAMs may be useful for genetics and breeding applications in species.

Single nucleotide polymorphisms (SNPs) are ideal molecular markers due to their higher abundance. Although several types of genotyping platforms for assaying large number of SNPs are available, in cases such as marker-assisted selection, where few markers are required for genotyping a set of potential lines, high-throughput SNP genotyping platforms (e.g., iScan or Infinium) may not be cost effective. In this scenario, GoldenGate assays based on VeraCode technology using Illumina BeadXpress seems to be the most cost-effective platform. The objective of this study was to develop cost-effective SNP genotyping platforms in chickpea ( L.) and pigeonpea ( L.). Two sets of SNPs, one each for chickpea (96 SNPs) and pigeonpea (48 SNPs), were developed and tested by genotyping 288 diverse genotypes from respective reference sets. The SNPs selected for the oligo pool assays had high transferability to crop wild relative species. The mean polymorphism information content value of assayed SNP markers was 0.31 and 0.32 in chickpea and pigeonpea, respectively. No unique pattern was observed in the chickpea reference set whereas two major groups were observed in the case of the pigeonpea reference set. The Illumina BeadXpress platform assays developed for chickpea and pigeonpea are highly informative and cost effective for undertaking genetic studies in these legume species.