Genetic variability, heritability and genetic advance studies for seed cotton yield and yield contributing characters in upland cotton

Introduction

Cotton, popularly known as the "white gold," is one of the most important commercial fibre crops in the world. Among the cultivated cotton species, Gossypium hirsutum (L.), commonly referred to as upland or upland cotton, occupies a predominant position in global cotton production scenario owing to its high yield potential, wide adaptability and superior fibre characteristics. It contributes more than 90 per cent of the world's cotton production (Morales et al., 2022) [1] . India ranks first in the world in area under cotton cultivation, whereas second in total production and consumption. The area under cotton cultivation during 2025-26 was about 114.82 lakh hectares with production of 290.91 lakh bales, with productivity of 428-451 kg per hectare. However, domestic consumption during the same period resulted in a supply gap met partly through imports, underscoring the continuing need to raise productivity through genetic improvement. In recognition of this, the Government of India has approved the five-year Mission for Cotton Productivity (2026-27 to 2030-31) to enhance yield and fibre quality across major cotton-growing states (Telangana Today, 2026). Seed cotton yield is a complex quantitative trait governed by polygenes and strongly influenced by environmental factors, and it is the end product of several interacting yield components such as number of bolls per plant, boll weight, plant height, number of sympodial and monopodial branches, and the duration to flowering and maturity (Fang et al., 2014) [2] . Gossypium hirsutum is a tetraploid (2𝑛 = 4𝑥 = 52) species with a broad genetic base; however, intensive breeding and continuous selection for specific traits have narrowed the variability available upland cotton (Zhang et al., 2016) [3] . Systematic evaluation of available genotypes for seed cotton yield and its component traits is essential for identifying promising parental material for future breeding programme. Several workers have reported considerable genetic variability for seed cotton yield and its component characters in hirsutum cotton (Sahar et al., 2021a; Chapara et al., 2022; Divya et al., 2023) [5,4,6] , yet there remains a continuous need to evaluate locally adapted genotypes under specific agro-climatic conditions to identify superior lines which can be utilized for cotton improvement programme to develop promising hybrids. Statistical tools such as analysis of variance, estimation of genotypic and phenotypic coefficients of variation, heritability and genetic advance are widely employed to quantify the nature and extent of genetic variability present in a set of genotypes and to determine the scope for improvement through selection. The magnitude of genotypic coefficient of variation together with heritability and genetic advance as per cent of mean indicates whether a character is governed predominantly by additive gene action, rendering it amenable to direct phenotypic selection, or by non-additive gene action. In such scenario hybridization could be useful to exploit heterosis. In view of this the present investigation was undertaken to assess the seed cotton yield performance and the extent of genetic variability for seed cotton yield and yield-contributing traits among a set of hirsutum cotton genotypes evaluated at Cotton Research Station, Nanded, Maharashtra.

Materials and Methods

Experimental site, material and design

The with spacing of 60 × 30 cm. All recommended agronomic package of practices and need-based plant protection measures were followed for healthy crop growth.

Observations recorded

Observations were recorded on five competitive and randomly selected plants in each replication of every genotype for eight yield and yield-contributing characters. The mean values per plant were worked out. Days to 50 per cent flowering was recorded as the number of days from sowing to the date on which at least 50 per cent of plants had flowered. Plant height (cm) was measured from the ground level to the growing point of the main stem at final picking. Days to maturity was recorded as the number of days from sowing to maturity of 80 per cent of plants in a plot. Number of sympodia per plant and number of monopodia per plant were recorded by counting the reproductive (fruiting) and vegetative (non-fruiting) branches, respectively, on the main stem at maturity. Number of bolls per plant was recorded by counting the fully matured, open bolls on each observational plant. Boll weight (g) was determined from the average weight of seed cotton of ten bolls per treatment after final picking. Seed cotton yield per plant (g) was recorded as the seed cotton obtained from each of the five randomly selected plants, averaged per treatment.

Statistical analysis

Replication-wise mean values for all characters were subjected to analysis of variance (ANOVA) as per the standard method of Panse et al. (1985) [7] to test the significance of genotypic differences. Genotypic and phenotypic variances were estimated from the respective mean sums of squares following Burton (1952) [8] , and the genotypic and phenotypic coefficients of variation (GCV and PCV) were calculated by the method of Burton (1952) [8] and categorized as low (< 10%), moderate (10-20%) or high (≥ 20%) following Sivasubramanian et al. (1973) [9] . Broad-sense heritability (h² b.s.) was estimated according to Allard (1960) [10] and classified as low (< 30%), moderate (30-60%) or high (≥ 60%) as suggested by Robinson et al. (1949) [11] . Genetic advance (GA) at 5 per cent selection intensity (K = 2.06) and genetic advance as per cent of mean (GAM) were computed as per method given by Johnson et al. (1955) [12] and classified as low (< 10%), moderate (10-20%) or high (≥ 20%). The standard error of difference between genotypic means and the critical difference (CD) at 5 per cent and 1 per cent levels of significance were also worked out for each character.

Results and Discussion

The present investigation was undertaken to evaluate seventeen upland cotton (Gossypium hirsutum L.) genotypes, comprising seventeen elite genotypes and three checks (NH 677, NH 545 and NH 615), for their seed cotton yield performance and associated yield-contributing characters. Results of the statistical analyses carried out on the data recorded for the seventeen genotypes for eight yield and yield-contributing traits are presented and discussed below, and are compared with relevant earlier investigations.

Analysis of variance

The mean sum of squares due to replication, treatment (genotypes) and error for the eight yield and yield-contributing characters are presented in Table 1. The analysis of variance revealed highly significant differences (P ≤ 0.01) among the seventeen genotypes for all the eight characters studied, viz., days to 50 per cent flowering, days to maturity, plant height, number of sympodia per plant, number of monopodia per plant, number of bolls per plant, boll weight and seed cotton yield per plant. This indicated the presence of substantial amount of genetic variability present in seventeen genotypes screened for seed cotton yield and its component traits. This confirms that selection could be effective for improvement of these characters. The mean sum of squares due to replication was comparatively small for all characters, indicating a reasonably uniform experimental environment across the three replications. Similar highly significant genotypic differences for yield and its component traits in upland cotton were earlier reported by Anjani et al. (2020) [13] and Chapara et al. (2024) [14] .

Mean performance of the genotypes for yield and yieldcontributing traits

Mean performance of the seventeen G. hirsutum genotypes for the eight yield and yield-contributing characters is furnished in Table 2, along with the general mean, standard error (SE ± ), coefficient of variation (CV%) and critical difference (CD) at 5 and 1 per cent levels of significance.

Days to 50 per cent flowering

Mean days to 50 per cent flowering ranged from 66.00 (NH 825) to 76.00 (NH 1136), with a general mean of 72.27 days; NH 1136 was followed by NH 817 and NH 1187 (75.00 days each) for the highest values. The spread between the highest and lowest performing genotypes exceeded the critical difference (CD @ 5% = 4.64), confirming statistically significant genotypic differences for this trait, consistent with the highly significant treatment mean sum of squares in Table 1 and with the findings of Ishaq et al. (2025) [15] and Kaleri et al. (2025) [16] in upland cotton.

Days to maturity

The maximum mean for days to maturity were recorded by NH 814 (145.67), followed by NH 1212 (144.67) and NH 824 (144.33). Days to maturity ranged from 131.33 to 145.67 days with a general mean of 141.97 days, the spread exceeding the CD @ 5% of 5.56 and indicating appreciable genetic variability for maturity duration. The results are in accordance with the reports of Sendekie et al. (2024) [17] and Hugar et al. (2024b) [18] .

Highest value of plant height was recorded by NH 1187 (206.67 cm), followed by NH 1222 (206.33 cm) and NH 864 (204.80 cm). Whereas, genotype NH 1226 recorded lowest value for plant height (163.33 cm). The general mean was 183.57 cm, with the range (163.33-206.67 cm) exceeding the CD @ 5% value of 9.77 cm, confirming significant genotypic differences. Similar kind of results were reported by Patil et al. (2023) [19] and Damtew et al. (2022) [20] .

Number of sympodia per plant

Mean values for number of sympodia per plant ranged from 25 [21] and Aslan et al. (2022) [22] .

Number of monopodia per plant

Number of monopodia per plant ranged from 2.80 (NH 1187) to 6.70 (NH 1201), with a general mean of 5.38. The genotype NH 1201 (5.38) recorded maximum monopodia per plant followed by NH 825 (6.67) and NH 820 (6.63). The spread exceeded the CD @ 5% of 0.57, confirming significant genotypic differences. Similar results were reported by Kamran et al. (2020) [23] and Manonmani et al. (2019) [24] .

Number of bolls per plant

The [25] and Bakhsh et al. (2019) [26] .

Maximum boll weight was recorded by NH 821 (3.50 g), followed by NH 824 (3.27 g) and NH 1136 (3.20 g). Whereas, genotype NH 860 recorded the least boll weight (2.22 g). The general mean for boll weight was 2.82 g, with the range (2.22-3.50 g) exceeding the CD @ 5% of 0.41 g, confirming significant genotypic differences. The results are in agreement with results of Karademir et al. (2015) [27] and Karademir et al. (2012) [28] .

Seed cotton yield per plant is the most important and complex character governed by many genes. It is chief objective to improve seed cotton yield in any cotton improvement programme. Mean values for seed cotton yield ranged from 49.86 g (NH 860) to 89.96 g (NH 824), against a general mean of 72.27 g. Genotype NH 824 recorded the highest mean seed cotton yield, followed by the NH 615 (85.90 g) and NH 821 (84.52 g). The difference between the highest and lowest yielding genotypes exceeded the CD @ 5% of 13.32 g, confirming a statistically significant genotypic difference. The results are in agreement with the reports of Aslan et al. (2022) [22] and Karademir et al. (2015) [27] . The promising genotypes for seed cotton yield per plant were NH 824, NH 615 (check) and NH 821. These genotypes could be utilized as potential parents or donor lines for seed cotton yield per plant in future hybridization aimed at seed cotton yield improvement in G. hirsutum cotton. Considering seed cotton yield and its contributing traits as a whole, NH 824 and NH 821 identified as the most promising entries, exhibiting above-average performance for seed cotton yield per plant coupled with a associated seed cotton yield component traits such as number of bolls per plant and boll weight. These results are in agreement with the reports of Karademir et al. (2015) [27] ,

Bakhsh et al. (2019) [26] , Aslan et al. (2022) [22] , Damtew et al. (2022) [20] and Sendekie et al. (2024) [17] in G. hirsutum genotypes. These genotypes could be used for improvement of earliness, plant stature and seed cotton yield attributes.

Genetic variability, heritability and genetic advance

Estimates of range, mean, genotypic and phenotypic variance, genotypic and phenotypic coefficients of variation (GCV and PCV), broad-sense heritability (h²) and genetic advance as per cent of mean (GAM) for the eight yield and yield-contributing characters are presented in Table 3. The phenotypic coefficient of variation was higher in magnitude than the corresponding genotypic coefficient of variation for all eight characters, indicating some influence of environment on trait expression; however, the comparatively narrow gap between GCV and PCV for most characters suggested that a considerable proportion of the observed variability was under genetic control.

Genotypic and phenotypic coefficients of variation

Genotypic and phenotypic coefficients of variation were estimated as per classification given by Sivasubramanian et al. (1973) [9] . High GCV (≥ 20%) was recorded for number of monopodia per plant (23.80%). Moderate GCV (10-20%) was observed for seed cotton yield per plant (13.65%) and boll weight (12.06%), while the remaining characters, viz; Days to 50 per cent flowering (2.86%), days to maturity (1.69%), plant height (8.01%), number of sympodia per plant (4.81%) and number of bolls per plant (4.54%), exhibited low GCV. The phenotypic coefficient of variation followed a similar trend but was slightly higher in magnitude for every character. High PCV (24.64%) was recorded only for number of monopodia per plant while moderate PCV for seed cotton yield per plant (17.62%) and boll weight (14.95%). High GCV and PCV for number of monopodia per plant and seed cotton yield per plant indicated substantial genetic variability for these traits, providing considerable scope for improvement through direct selection. The results are in agreement with reports of Sakthi et al. (2007) [29] and Devi et al. (2020) [30] .

Heritability (broad sense)

Heritability was estimated as per classification of Robinson et al.

(1949) [11] . High heritability (≥ 60%) was recorded for number of monopodia per plant (93.23%), plant height (86.09%) and boll weight (65.09%). While moderate heritability (30-60%) was observed for seed cotton yield per plant (59.97%), number of sympodia per plant (37.86%), number of bolls per plant (36.72%), days to 50 per cent flowering (35.14%) and days to maturity (33.73%). The high heritability for number of monopodia per plant indicated that this character was least influenced by environment and direct selection could be effective for improvement of it. These results are consistent with the reports of Sakthi et al. (2007) [29] and Meena et al. (2001) [31] .

Genetic advance as per cent of mean

Genetic advance as per cent of mean was estimated as per classification of Johnson et al. (1955) [12] . High genetic advance as per cent of mean (≥ 20%) was recorded for number of monopodia per plant (47.33%), seed cotton yield per plant (21.77%) and boll weight (20.04%). While the remaining characters showed low to moderate GAM. Plant height recorded a moderate value of 15.31 per cent. Characters coupled high heritability and high GAM are viz; Number of monopodia per plant and boll weight, indicated that heritability in these traits was largely attributable to additive gene action, so that selection based on phenotypic expression would be effective for their improvement. Seed cotton yield per plant, despite only moderate heritability, combined this with a high genetic advance as per cent of mean (21.77%), suggesting that a meaningful part of its heritable variation is additive in nature and amenable to selection gains, notwithstanding some environmental influence on its expression. In contrast, plant height exhibited high heritability (86.09%) but only moderate GAM (15.31%), suggesting the possible involvement of non-additive gene action (dominance and epistasis) in its expression, in which case recurrent selection would be effective (Selection after in termating of individuals). These findings are in accidence with the findngs of Desalegn et al. (2009) [32] , Sakthi et al. (2007) [29] and Meena et al. (2001) [31] for yield and its component traits in G. hirsutum. Taken together, the results of the present investigation demonstrated the presence of appreciable genetic variability for seed cotton yield and its contributing characters among the seventeen hirsutum cotton genotypes evaluated. Substancial amount of genetic variability was noticed for number of monopodia per plant, boll weight and seed cotton yield per plant. These characters combining reasonably high heritability with high genetic advance, and therefore amenable to improvement through direct selection. Genotypes NH 824, NH 821 and the check NH 615, which expressed high seed cotton yield with a desirable magnitude of yield components, emerged as promising genetic material for use as parents/donors in hybridization programmes directed at seed cotton yield improvement in upland cotton.

Conclusion

The present investigation demonstrated highly significant genotypic differences among seventeen upland cotton genotypes for all eight yield and yield-contributing traits studied. Which revealed presence of substantial amount of genetic variability that is largely under genetic control. Whereas, environmental influence is negligible. Seed cotton yield per plant ranged from 49.86 g to 89.96 g, and genotypes NH 824, NH 821 and the check NH 615 were identified as the most promising entries, combining superior yield with favourable expression of number of bolls per plant and boll weight. Number of monopodia per plant, boll weight and seed cotton yield per plant, which combined moderate-to-high heritability with high genetic advance as per cent of mean, appeared to be governed predominantly by additive gene action and are therefore amenable to improvement through direct phenotypic selection, whereas plant height, which showed high heritability but only moderate genetic advance, would benefit from hybridization-based breeding approaches. These findings provide valuable genetic information to cotton breeders to select genetically diverse superior parental genotypes to be utilized in cotton improvement programmes in upland cotton. Furthermore these genotypes must be tested at multi-locations in multi-season trials to confirm the stable performance prior to their use in breeding or release.

Acknowledgement

The authors acknowledge to Cotton Specialist, Cotton Resarch Station, Bafna T Point, Nanded under jurisdiction of Vasantrao Naik Marathwada Krishi Vidyapeeth, Parbhani for providing all the facilities for smooth conduct of experiment. traits in seventeen upland cotton genotypes

References

1. Morales KY, Bridgeland AH, Hake KD, Udall JA, Thomson MJ, Yu JZ. Homology-based identification of candidate genes for male sterility editing in upland cotton (Gossypium hirsutum L.). Frontiers in Plant Science. 2022;13:1006264.
2. Fang DD, Jenkins JN, Deng DD, McCarty JC, Li P, Wu J. Quantitative trait loci analysis of fiber quality traits using a random-mated recombinant inbred population in upland cotton (Gossypium hirsutum L.). BMC Genomics. 2014;15:397.
3. Zhang J, Wu M, Yu J, Li X, Pei W. Breeding potential of introgression lines developed from interspecific crossing between upland cotton (Gossypium hirsutum) and Gossypium barbadense: Heterosis, combining ability and genetic effects. PLOS ONE. 2016;11(1):e0143646.
4. Chapara R, Siva Reddy KV, Sudha Rani M, Sree Lakshmi B, Roja V, Pranaya J. Variability studies and genetic divergence in cotton (Gossypium hirsutum L.) germplasm using multivariate analysis. Electronic Journal of Plant Breeding. 2022;13(4):13051311.
5. Sahar A, Majeed S, Rana IA, Atif RM. Genetic variability for yield and fiber related traits in genetically modified cotton (Gossypium hirsutum L.). Journal of Natural Fibers. 2021;19(13):6439-6450.
6. Divya M, Sabitha N, Mohan Vishnuvardhan K. Studies on genetic variability for yield and fibre quality traits in upland cotton (Gossypium hirsutum L.). The Journal of Research ANGRAU. 2023;51(3):59-64.
7. Panse VG, Sukhatme PV. Statistical methods for agricultural workers (4th ed.). Indian Council of Agricultural Research. 1985.
8. Burton GW. Quantitative inheritance in grasses. In Proceedings of the 6th International Grassland Congress. 1952;1:277-283.
9. Sivasubramanian S, Menon M. Genotypic and phenotypic variability in rice. Madras Agricultural Journal. 1973;60:1093-1096.
10. Allard RW. Principles of plant breeding. John Wiley & Sons; 1960.
11. Robinson HF, Comstock RE, Harvey PH. Estimates of heritability and the degree of dominance in corn. Agronomy Journal. 1949;41(8):353-359.
12. Johnson HW, Robinson HF, Comstock RE. Estimates of genetic and environmental variability in soybeans. Agronomy Journal. 1955;47(7):314-318.
13. Anjani A, Padma V, Ramana JV, Satish Y. Evaluation of genetic parameters of agro-morpho-quality traits in upland cotton (Gossypium hirsutum L.). Electronic Journal of Plant Breeding. 2020;11(1):279-282.
14. Chapara R, Reddy KVS, Rani MS, Sudhamani K. Association and divergence studies for yield and its attributing traits in upland cotton. The Journal of Research ANGRAU. 2024;52(1):1-8.
15. Ishaq M, Hussain A, Ali M, Ahmad S. Yield performance and genetic variability of advanced upland cotton lines under field conditions. Journal of Plant Breeding and Crop Science. 2025;17(1):12-20.
16. Kaleri AA, Panhwar SA, Keerio AA, Soomro ZA. Multivariate analysis for genetic diversity in upland cotton crosses for yield and fiber traits. Pakistan Journal of Botany. 2025;57(2):421-430.
17. Sendekie Y, Habite DN. Performance Evaluation of Cotton (Gossypium hirsutum L.) Varieties for Yield and Related Traits in North-Western Ethiopia. EAS Journal of Biotechnology and Genetics. 2024;6(1):7-11.
18. Hugar AY, Katageri IS, Jadhav MP, Vamadevaiah HM. Molecular diversity analysis and heterosis prediction in upland cotton (Gossypium hirsutum L.) for yield and fibre traits. Indian Journal of Genetics and Plant Breeding. 2024;84(1):45-52.
19. Patil SS, Deshmukh SB, Patil VN, Wadikar PB. Stability analysis of seed cotton yield in upland cotton (Gossypium hirsutum L.) using AMMI and GGE biplot methods. Journal of Cotton Research. 2023;6(18):1-11.
20. Damtew Z, Mohammed W, Tefera H. Genetic variability and performance evaluation of upland cotton genotypes for yield and yield components. Cogent Food and Agriculture. 2022;8(1):2039874.
21. Li Y, Wang Y, Chen Q, Zhang Z, Liu R, Ma X. Genome-wide association study of yield and fiber quality traits in upland cotton (Gossypium hirsutum L.). Frontiers in Plant Science. 2022;13:835612.
22. Aslan R, Aygün YZ, Mert M. Yield and fiber quality characteristics of some cotton (Gossypium hirsutum L.) cultivars grown in the Southeastern Anatolian conditions. Turkish Journal of Field Crops. 2022;27(2):285-292.
23. Kamran M, Afzal I, Basra SMA, Mahmood A, Sarwar G. Harvesting and post-harvest management for improving seed quality and subsequent crop yield of cotton. Crop and Pasture Science. 2020;71(12):1041-1049.
24. Manonmani S, Manickam S, Raveendran TS. Genetic diversity analysis in cotton (Gossypium hirsutum L.) using SSR markers. Electronic Journal of Plant Breeding. 2019;10(3):1054-1061.
25. Shahzad K, Zhang X, Guo L, Qi T, Bao L, Zhang M. Evaluation of genotype x environment interaction for seed cotton yield and fiber quality traits in upland cotton (Gossypium hirsutum L.). Field Crops Research. 2019;236:70-79.
26. Bakhsh A, Rehman M, Salman S, Ullah R. Evaluation of Cotton Genotypes for Seed Cotton Yield and Fiber Quality Traits Under Water Stress and Non-Stress Conditions. Sarhad Journal of Agriculture. 2019;35(1):161-170.
27. Karademir C, Karademir E, Gencer O. Determination of yield and quality characters of some cotton (Gossypium hirsutum L.) genotypes. In Proceedings of the Plenary Meeting of the InterRegional Cooperative Research Network on Cotton. 2015:92-95.
28. Karademir C, Karademir E, Akışcan C. Effect of heat stress on leaf area in cotton (Gossypium hirsutum L.). Journal of Food, Agriculture & Environment. 2012;10(2):522-525.
29. Sakthi AR, Kumar M, Ravikesavan R. Variability and association analysis using morphological and quality traits in cotton (Gossypium hirsutum L.). Journal of Cotton Research and Development. 2007;21(2):148-152.
30. Devi GG, Nidagundi JM, Diwan JR, Mahanthashivyogayya K, Hanchinal SG. Evaluation of brown coloured cotton genotypes by genetic studies (Gossypium hirsutum L.). International Journal of Current Microbiology and Applied Sciences. 2020;9(10):14051412.
31. Meena RA, Mishra MN, Dani RG. Genetic variability and correlation for seed-quality parameters upland cotton (Gossypium hirsutum). The Indian Journal of Agricultural Sciences. 2001;71(6):417-420.
32. Desalegn Z, Ratanadilok N, Kaveeta R. Correlation and heritability for yield and fiber quality parameters of Ethiopian cotton (Gossypium hirsutum L.) estimated from 15 (diallel) crosses. Kasetsart Journal (Natural Science). 2009;43(1):1-11.