A better understanding of the extent of natural genotypic variation existing in the germplasm of cultivated groundnut is helpful in identification of genotypes likely with greater yield potential and nutritional values that can be utilized as parental material for breeding of improved cultivars. Characterization of 371 germplasm originating from more than fourteen countries, resulted in the identification of trait specific and multi-traits germplasm from the global composite collection.
Genotypic Variability
The prerequisite for successful breeding program is to find sufficient amount of variability, in which desired lines are to be selected for further manipulation to achieve the target. Genetic variability is of paramount importance in selecting the best genotypes for making rapid improvement in yield and desirable characters as well as for selection of the most promising parents for further breeding program. Study of genetic variability reveals variation in different quantitative traits. The success of breeder in selecting suitable quality parameters lies largely on existence and exploitation of genetic variability to the fullest extent (Allard, 1960). The coefficient of variability for hundred kernel weight (g) was 17.43%; 7.45% at two locations, 48.97; 30.96% for pod yield (g/m2) and 8.91; 15.34% for shelling percentage. So, the traits showed significant variation among accessions, with the coefficient of variation of various parameters had a range value (7.45–48.97%), indicating that the trial and data recording was carried out with sufficient precision. In other studies, low values of coefficient of variability were recorded, suggesting better improvement scope for these traits. The coefficients of variations measures magnitude of variability present in the population. In other studies, high values of coefficient of variability were recorded, suggesting better improvement scope for these traits by selection of parents with desirable attributes for breeding program. The frequency distribution of germplasm for yield and related attributes in different groups indicated the presence of high level of genotypic diversity among the germplasm evaluated at the two locations.
Germplasm identification involves identifying accessions with consistent and extraordinary agronomic traits, and biochemical traits such as high oil, sugar, and protein levels. This study found that most biochemical parameters were within normal ranges when compared to past research. Both places tested showed variability in germplasm for biochemical traits. Junagadh has more diversified germplasm for the traits under research than Anantapur. The total variable range for oil, protein, and sugar concentrations in Junagadh was 40.49 to 52.40%, 23.54 to 35.924%, and 3.73 to 9.04%, respectively. At Anantapur, oil, protein, and sugar ranged from 41.36 to 47.75%, 28.33 to 35.96%, and 4.84 to 8.39%, respectively. Oil and protein content values in the population showed continuity, indicating a quantitative inheritance pattern. Genetic and environmental factors cause this metabolic characteristic to fluctuate significantly. The study found that 58.49% of Anantapur germplasm and 36.65% of Junagadh germplasm had oil content between 44.1–46% and 46.1–48%. 53.9% and 37.46% of germplasm from Anantapur and Junagadh had seed protein values between 32.1–34%. Researchers worldwide have found similar seed oil and protein concentrations. Jambunathan et al. 1985 studied 6840 germplasm accessions at ICRISAT in tiny batches over several years. The results showed that accessions had 32–55% oil and 16–34% protein. Upadhyaya et al. 2003 found 45–55% oil content heterogeneity in the groundnut minicore collection, wild Arachis accessions in the ICRISAT gene bank (Upadhyaya et al. 2011), and USDA groundnut germplasm repository (Wang et al. 2012). According to unpublished data from S.N. Nigam of ICRISAT in 2006, genotypes with high oil or protein content in one environment were unsustainable when systematically assessed in multiple environments.
Heba et al. (2021) and Meena et al. (2022) found that increased moisture stress can reduce nutrient uptake, affecting growth and development of groundnut. Breeding program needs reliable sources with high oil concentration, and superior for agronomic traits. The coefficient of variation compares quantitative trait genetic variability. At two locations, oil and protein levels had low coefficient of variability (CV) values of 2.67% and 3.76% and 5.23% and 6.05%, respectively. Asibuo et al. (2008) analysed 20 Ghanaian landraces and found protein (1.50%) and oil (0.34%) with very low CVs. According to Lynch and Walsh (2001), the oil and protein contents exhibited low values, which is indicative of a significant degree of genotypic variability in these traits. Oil, was lower in Anantapur than Junagadh. The mean values of protein, sugar, and FAA at Anantapur were higher than those at Junagadh, indicating more diversity across germplasm accessions. Mid-season drought decreased groundnut oil content (Conkerton et al. 1989), and according to Vaidya et al. (2015) and Yadav et al. (2013) drought stress increased genetic variability, FAA, and total soluble protein as observed at Anantapur condition than that was found at Junagadh. A higher kernel protein content and stress-induced protein breakdown may explain the observed increase in free amino acids. Stress heterogeneously affected total phenol content. Genetic variability makes it easy to identify genotypes with drought-stress-related features for breeding. Small-molecular-mass proteins accumulate quicker than fatty acids in drought-stressed plants. De novo synthesis or amino acid degradation enhances total soluble protein content. Protein increases may help plants tolerate drought by balancing osmotic pressure (Yadav et al. 2013). The current study confirmed previous findings that Anantapur, a location that endures repeated droughts during the rainy season, accumulated less oil but more proteins, carbohydrates, and free amino acids. Chakraborty et al. (2016) found that oil content decreased during drought stress as observed under Anantapur as well.
Mean performance (Y), stability variance (σi 2 ), and rankings (𝘒R)
Anantapur and Junagadh's groundnut germplasm's pod yield, hundred kernel weight (HKW), and shelling percentage (SHP) mean values were 223.66 and 134.47 g/m2, 34.75 and 36.38 g, and 72.11 and 71.72%, respectively. Junagadh has lower pod yield and SHP than Anantapur. Table 2S shows that precipitation, sunshine exposure, and temperature may explain the differences between the two experimental sites. During the rainy season, Anantapur received 630 mm and Junagadh 1305 mm. The rainy season lowest and maximum temperatures at Anantapur and Junagadh were 17.8°C and 35.0°C, and 16.2°C and 38.2°C, respectively. Anantapur had 39.87% greater pod yield and 49.55% higher SHP than Junagadh. In Anantapur and Junagadh, seasonal solar exposure was explored on photosynthesis, photosynthate accumulation and translocation, yield, and related traits. Anantapur's continuous daily sunlight hours above 5 hours may have optimised photosynthetic processes and yields. Figure 1b shows that Junagadh's erratic rainfall and lack of sunshine during pod setting may have reduced yields. Solar radiation interception increases Radiation Use Efficiency (RUE), dry matter production, and yield in several Arachis sp. cultivars (Oluwasemire and Odugbenra, 2014). Junagadh's typical daylight hours of 1–3 hours/day have affected yield traits throughout critical crop growth stages including flowering and pod formation. During kharif season, high moisture stress, disease pressure, and insufficient sunshine in Junagadh impair crop performance. HKW in Junagadh was 51.54% greater than Anantapur's. The observed variation may be attributed to distinct climatic and edaphic factors associated with the cultivation of groundnut. A breeding program aims to generate cultivars with high yield potential and adaptability to various agro-ecological zones. The mean performance and stability statistic of NRCG-14027, NRCG-10272, NRCG-11546, and NRCG-12229 indicated the best cultivars for specific outturn (SHP) and NRCG-14027, NRCG-10366 and NRCG-13960 were best cultivars for pod yield.
The mean values for oil content, protein content, sugar content, phenol and FAA of germplasm over two environments, ANT and JND was 44.62; 46.59%, 32.40; 31.37%, 6.43; 5.97%, 0.25; 0.27%, and 0.66; 0.41% respectively. The biochemical data indicate that the genotypes exhibited different behavior for different traits in response to the two different environments. However, mean values of biochemical traits were comparable from two locations that could help in identifying stable performing germplasm. For oil content, NRCGs-11270, 11515, 11732, 12380, 12465, 12470 and 13945 were most desirable. While for protein content, germplasm viz., NRCGs-11562, 11732, 13959, 14067, 14072, 14073 and 14545 were identified as best performing. Available reports indicate that water stress reduces oil yield (Hashim et al. 1993 and Chakraborty et al. 2016) whereas protein (Dwivedi et al. 1996) and sugar content (Chakraborty et al. 2016) increases. Findings of present study are in agreement with previous reports wherein Anantapur location which faces frequent dry spells during rainy season had reduced accumulation of oil but with increased accumulation of proteins and sugars. This indicates that drought stress at optimum or higher air temperature (Fig. 1a) can significantly affect groundnut oil content. The lack of adequate C-supply from the source tissue (both due to reduced photosynthesis and conversion of assimilate for biosynthesis of organic osmo-protectants) resulted in reduction in kernel oil content, but a relative increase in protein content (Chakraborty et al. 2016).
Inter-Relationship among various traits and Identification of superior accessions for multiple traits
Understanding inter-character correlation is essential to select best genotypes from the population. However, selection for a trait may reduce other attributes. Figure 5 shows groundnut physicochemical and agronomic connections. Trait relationships were examined in two contexts i.e Junagadh and Anantapur conditions seperately. Junagadh data suggests no negative association between pod yield and protein content. Thus, both factors can be improved simultaneously. High oil yield per unit area is difficult due to the negative association between pod yield and oil content. Chiow and Wynne (1983) found a negative association between these variables. In both the environments, oil content and 100-kernel weight had no significant negative connection (0.05 & 0.17**). Two factors were negatively correlated as reported by Janila et al. (2016). When they examined 33 genotypes, Dwivedi et al. (1990) found a positive correlation between kernel oil content and seed mass. Ajay et al. (2012) found that kernel protein and oil content affect confectionery groundnut cultivar preferences. Groundnut seed contains oil, crude protein and carbohydrates (Francisco and Resurreccion 2008). An increase in any of these ingredients should decrease one or more of the others. Assimilates are distributed across several seed components, therefore protein concentration and other important constituents are intrinsically linked. Thus, changing one component affects the others. The correlation coefficients of + 0.93 (ATP) and + 0.54 (JND) show that protein and sugar are positively correlated. NRCGs-10366, 10480, 10485, and 10844 were superior for PY, HKW, and SHP; NRCG-11390 for PY, protein, and sugar; and NRCGs-12469, 14089, and 16492 for important biochemical traits (oil, protein, and sugar content); NRCG-11101 for HKW, protein, and oil; and NRCGs-11154, 11164 for HKW, protein, and sugar. Table 4S describes the morphology of exceptional germplasm with numerous desirable features. After adaptive trials, stable genotypes with better mean performance can be grown and employed as breeding parents.