Heterosis of okra [Abelmoschus esculentus (L.) Moench] for fruit yield and its components traits

Introduction

Okra [Abelmoschus esculentus (L.) Moench] is a major vegetable crop cultivated widely in tropical and subtropical areas and is a member of the Malvaceae family with India accounting for a substantial proportion of global production. It is a warm-climate annual crop propagated by seed with an indeterminate growth habit and a chromosome number is 2n = 130. (Dutta and Naug, 1968) [6] . The crop grows well under hot and humid climatic conditions (Lamont, 1999) [14] and is widely cultivated for its tender edible pods, which are rich in dietary fibre, vitamins and minerals and contain a well-balanced amino acid composition comparable to that of soybean (Kumar et al., 2013) [13] . Okra has diverse uses and can adapt to different soil conditions, although well-drained and fertile soils with an appropriate pH are considered most suitable for its cultivation (Jain et al., 2012) [10] . The roots and stems of okra are also utilized in the clarification of gur, khand and raw sugar. Owing to its hardiness, dietary fibre content and favourable seed protein composition, particularly the essential amino acids lysine and tryptophan. In India, major okra-producing states include Gujarat, Maharashtra, Uttar Pradesh, Tamil Nadu, Andhra Pradesh, Karnataka and Punjab. The crop is cultivated during both the Kharif and summer seasons. In Gujarat, Vadodara, Junagadh, Surat, Anand, Surendranagar and Banaskantha is the major okragrowing districts.

Materials and Methods

The investigational material consisted of seven female lines viz., JOL-24-108, JOL-24-113, JOL-21-106, JOL-12-13, AOL-22-16, SKNOL-23-03 and JOL-22-17, along with four male testers of okra viz., GJO-3, Kashi Lalima, GAO-8 and KS-404 (Table 1). A total of 28 F1 hybrids were developed through a line × tester mating design along with one standard check hybrid (GJOH-4). The material was evaluated during kharif 2025 at the Vegetable Research Station, Junagadh Agricultural University, Junagadh using a randomized block design (RBD) with three replications. A spacing of 60 cm between rows and 30 cm between plants was maintained. The crop was raised following the recommended agronomic practices of the region to ensure proper crop growth and development. Observations were recorded on five randomly selected competitive plants from each replication for each genotype. Twelve characters were studied, namely nodal position for first flowering, days to first flowering, days to first picking, plant height (cm), number of primary branches per plant, number of fruits per plant, fruit length (cm), fruit girth (cm), average fruit weight (g), days to last picking, number of pickings and total fruit yield per plant (g). The mean values obtained from these observations were used for statistical analysis. Analysis of variance for each character was carried out following the procedure outlined by Panse and Sukhatme (1995) [22] . Heterosis was assessed using two parameters: heterobeltiosis representing heterosis over the better parent, as proposed by Fonseca and Patterson (1968) [7] and standard heterosis representing heterosis over the standard check as suggested by Meredith and Bridge (1972) [18] .

Results and Discussion

The analysis of variance revealed highly significant differences among the genotypes for all the characters studied. The total genotypic variance was further divided into components comprising parents (lines, testers and lines vs. testers), hybrids and parents vs hybrids. Significant differences were observed among the parents and hybrids for all the traits. The comparison between parents and hybrids was also highly significant for nodal position for first flowering, plant height (cm), number of primary branches per plant, number of fruits per plant, fruit length (cm), days to last picking, number of pickings and total fruit yield per plant (g). The mean squares attributable to lines were significant for all the characters, while those due to testers were also significant for all the traits, demonstrating the presence of substantial genetic variability within the experimental material. The lines vs testers component exhibited significant mean squares for all the characters, indicating differences between the female and male parents with respect to the traits studied. The contribution of testers to the total variance was comparatively greater for nodal position for first flowering, days to first flowering, days to first picking, plant height (cm), number of primary branches per plant, number of fruits per plant, fruit length (cm), average fruit weight (g), days to last picking, number of pickings and total fruit yield per plant (g) indicating a greater influence of testers on the variation observed for these characters. In contrast, lines contributed a larger proportion of the total variance for the remaining traits under investigation (Table 2). The findings were in agreement with the results reported by Patel and Patel (2016) [24] , Punia et al. (2017) [29] , Pithiya et al. (2019) [28] , Das et al. (2020) [5] , Patel et al. (2020) [23] , Samindre et al. (2022) [36] , Chaudhary et al. (2023) [4] , Sakriya et al. (2023) [35] , Pathania et al. (2024) [27] , Ranga et al. (2024) [33] , Vani et al. (2024) [41] and Vekariya et al. (2025) [42] for nodal position for first flowering, days to first flowering, days to first picking, plant height (cm), number of primary branches per plant, number of fruits per plant, fruit length (cm), fruit girth (cm), average fruit weight (g), days to last picking, number of pickings and total fruit yield per plant (g). Several cross combinations exhibited significant heterosis in the desirable direction over both the better parent and the standard check for various traits. The number of crosses showing favourable heterosis over the better parent and standard check was 13 and 9 for nodal position of first flowering, 7 and 7 for days to first flowering, 7 and 6 for days to first picking, 7 and 10 for plant height (cm), 6 and 12 for number of primary branches per plant, 5 and 8 for number of fruits per plant, 4 and 8 for fruit length (cm), 7 and 6 for fruit girth (cm), 4 and 6 for average fruit weight (g), 6 and 11 for days to last picking, 7 and 12 for number of pickings and 6 and 7 for total fruit yield per plant (g), respectively. Evaluation of the crosses for heterosis over the better parent and the standard check (GJOH-4) indicated considerable heterotic effects in the desirable direction for nodal position for first flowering, plant height (cm), number of primary branches per plant, number of fruits per plant, fruit length (cm), number of pickings and total fruit yield per plant (g). In contrast, comparatively lower levels of heterobeltiosis and standard heterosis were observed for days to first flowering, days to first picking, average fruit weight (g), fruit girth (cm) and days to last picking. Comparable findings have also been reported by Punia et al. (2017) [29] , Pithiya et al. (2019) [28] , Das et al. (2020) [5] , Patel et al. (2020) [23] , Samindre et al. (2022) [36] , Chaudhary et al. (2023) [4] , Sakriya et al. (2023) [35] , Pathania et al. (2024) [27] , Ranga et al. (2024) [33] , Vani et al. (2024) [41] and Vekariya et al. (2025) [42] . In okra, negative heterosis is considered desirable for nodal position for first flowering as it indicates earlier flowering. The heterobeltiosis values ranged from-26.85 percent (AOL-22-16 × GAO-8) to 4.46 percent (JOL-22-17 × Kashi Lalima), whereas standard heterosis ranged from-32.32 percent (AOL-22-16 × KS-404) to 18.18 percent (JOL-22-17 × Kashi Lalima) (Table 3). The present findings are in accordance with those reported by Nagesh et al. (2014) [19] , Padadalli et al. (2019) [21] , Pithiya et al. (2019) [28] , Poluru et al. (2023) [30] and Vani et al. (2024) [41] . Early flowering is an important breeding objective in okra, and days to first flowering serves as a key component for determining earliness. The heterobeltiosis estimates for days to first flowering ranged from-9.88 percent (JOL-12-13 × GJO-3) to 5.3 percent (JOL-24-113 × GAO-8), while the standard heterosis values varied from-11.11% (JOL-12-13 × KS-404) to 6.54% (SKNOL-23-03 × GJO-3). The results obtained in the present study were consistent with the findings of Patel et al. (2015) [26] , Tiwari et al. (2016) [40] , Lyngdoh et al. (2017) [15] , Rameshkumar et al. (2017) [31] , Makdoomi et al. (2018) [17] , Chaudhary et al. (2023) [4] and Ranga et al. (2024) [33] . Earliness for days to first picking is an important desirable attribute in okra. For this character heterobeltiosis ranged from-9.64 percent (AOL-22-16 × Kashi Lalima) to 5.56 percent (JOL-24-113 × GAO-8), whereas standard heterosis varied from-11.59% (JOL-12-13 × KS-404) to 6.71% (SKNOL-23-03 × GJO-3). The findings of the present investigation were comparable with those reported by Nagesh et al. (2014) [19] , Pithiya et al. (2019) [28] , Patel and Patel (2016) [24] , Tiwari et al. (2016) [40] and Vani et al. (2024) [41] . Greater plant height is generally associated with higher yield in okra. Therefore, positive heterotic effects for this trait are considered desirable. The heterobeltiosis estimates for plant height varied from-24.98 percent (AOL-22-16 × GJO-3) to 15.83 percent (AOL-22-16 × KS-404). In comparison, heterosis over the standard check ranged from-21.67 percent (SKNOL-23-03 × KS-404) to 20.74 percent (JOL-12-13 × Kashi Lalima). The present findings were in agreement with the observations reported by Patel and Patel (2016) [24] , Lyngdoh et al. (2017) [15] , Rameshkumar et al. (2017) [31] , Das et al. (2020) [5] , Ranga et al. (2024) [33] and Sood (2025) [37] . For the number of primary branches per plant, heterobeltiosis estimates varied from-46.51 percent (JOL-12-13 × KS-404) to 33.33 percent (SKNOL-23-03 × KS-404 and AOL-22-16 × KS-404). The standard heterosis values ranged from-40.54 percent (AOL-22-16 × GAO-8) to 45.95 percent (JOL-22-17 × Kashi Lalima). The results obtained were comparable with those reported by Nagesh et al. (2014) [19] , Mahajan et al. (2017) [16] , Punia and Garg (2017) [29] , Padadalli et al. (2019) [21] , Samindre et al. (2022) [36] , Gawai and Patel (2023) [9] and Abdelkader et al. (2024) [1] . For the number of fruits per plant, heterobeltiosis showed a range from-22.01 percent (JOL-12-13 × GAO-8) to 21.33 percent (JOL-22-17 × GJO-3). The estimates of standard heterosis ranged from-27.42 percent (AOL-22-16 × GAO-8) to 37.63 percent (JOL-22-17 × GJO-3). The findings were consistent with those reported by Patel and Patel (2016) [24] , Pithiya et al. (2019) [28] , Patel et al. (2020) [23] , Subhakar et al. (2022) [39] , Poluru et al. (2023) [30] , Abdelkader et al. (2024) [1] and Vekariya et al. (2025) [42] . Greater fruit length is generally considered favourable in okra because of its potential contribution to higher yield. Hence, positive heterotic effects are desirable for this character. The heterobeltiosis estimates ranged from-20.62 percent (JOL-22-17 × KS-404) to 13.58 percent (JOL-24-113 × GAO-8), whereas standard heterosis varied from-14.66% (JOL-24-113 × KS-404) to 20.7% (JOL-22-17 × GJO-3). None of the crosses exhibited positive and significant heterosis over the standard check (GJOH-4) for fruit length. The present findings were in accordance with those reported by Sabesan et al. (2016) [34] , Punia and Garg (2017) [29] , Samindre et al. (2022) [36] , Chaudhary et al. (2023) [4] and Ranga et al. (2024) [33] . For fruit girth in okra, negative heterosis is considered desirable. The heterobeltiosis values ranged from-5.8 (JOL-22-17 × KS-404) to 6.36 percent (SKNOL-23-03 × Kashi Lalima), while standard heterosis ranged from-7.36 (JOL-22-17 × GAO-8) to 6.86 percent (JOL-12-13 × GJO-3). The findings were in agreement with the results reported by Ram et al. (2016) [32] , Joshi et al. (2019) [11] , Abinaya et al. (2020) [2] , Chaudhary et al. (2023) [4] and Pathania et al. (2024) [27] . For average fruit weight, heterobeltiosis estimates varied from-14.96 (AOL-22-16 × GJO-3) to 11.94 percent (JOL-22-17 × GJO-3), whereas standard heterosis ranged from-17.31 (SKNOL-23-03 × Kashi Lalima) to 23.95 percent (JOL-22-17 × GJO-3). Positive heterotic effects for this trait have also been reported by Sabesan et al. (2016) [34] , Mahajan et al. (2017) [16] , Joshi et al. (2019) [11] , Poluru et al. (2023) [30] and Pathania et al. (2024) [27] . For days to last picking, the heterobeltiosis estimates varied from-5.57 percent (JOL-12-13 × KS-404) to 4.98 percent (SKNOL-23-03 × GAO-8), whereas standard heterosis ranged from-9.97 percent (JOL-12-13 × KS-404) to 10.63 percent (JOL-22-17 × GJO-3). The present results were comparable with the findings reported by Nagesh et al. (2014) [19] , Tiwari et al. (2016) [40] , Punia and Garg (2017) [29] , Chaudhary et al. (2023) [4] and Vekariya et al. (2025) [42] . For number of pickings, heterobeltiosis estimates ranged from-22.58 percent (SKNOL-23-03 × Kashi Lalima) to 15.63 percent (SKNOL-23-03 × GAO-8). Standard heterosis showed a range of-25.00 percent (SKNOL-23-03 × Kashi Lalima) to 40.63 percent (JOL-21-106 × GJO-3). The results obtained in the present investigation were in agreement with the observations reported by Nagesh et al. (2014) [19] , Pithiya et al. (2019) [28] , Patel et al. (2020) [23] and Ranga et al. (2024) [33] . For fruit yield per plant, heterobeltiosis estimates ranged from-19.64 (AOL-22-16 × GJO-3) to 20.07 percent (JOL-22-17 × GJO-3). The highest significant and desirable heterobeltiosis was observed in JOL-22-17 × GJO-3 (20.07%), followed by JOL-24-113 × GAO-8 (18.04%) and JOL-12-13 × GJO-3 (15.33%). Standard heterosis ranged from-23.62 (AOL-22-16 × GAO-8) to 45.33 percent (JOL-22-17 × GJO-3). Among the crosses, JOL-22-17 × GJO-3 recorded the highest significant standard heterosis in the desirable direction (45.33%), followed by JOL-12-13 × GJO-3 (39.59%) and JOL-21-106 × GJO-3 (35.10%). The present findings were in agreement with the results reported by Nagesh et al. (2014) [19] , Neetu et al. (2015) [20] , Bhatt et al. (2016) [3] , Mahajan et al. (2017) [16] , Gavint et al. (2018) [8] , Pithiya et al. (2019) [28] , Sapavadiya et al. (2019) [38] , Samindre et al. (2022) [36] , Chaudhary et al. (2023) [4] , Gawai and Patel (2023) [9] , Pathania et al. (2024) [27] , Ranga et al. (2024) [33] and Vani et al. (2024) [41] . The cross JOL-22-17 × GJO-3 emerged as a superior hybrid and expressed significant standard heterosis in the desirable direction for nodal position for first flowering, number of primary branches per plant, number of fruits per plant, fruit length (cm), average fruit weight (g), days to last picking, number of pickings and total fruit yield per plant (g). This performance indicated the important contribution of these traits to fruit yield per plant. The same cross also exhibited significant heterobeltiosis in the desirable direction for nodal position for first flowering, number of primary branches per plant, number of fruits per plant, fruit length (cm), average fruit weight (g), days to last picking, number of pickings and total fruit yield per plant (g). (Table 4) The second most heterotic cross JOL-12-13 × GJO-3 exhibited significant standard heterosis in the desirable direction for plant height (cm), number of primary branches per plant, number of fruits per plant, average fruit weight (g), fruit girth (cm), average fruit weight (g)), days to last picking, number of pickings and total fruit yield per plant (g). The cross also showed significant heterobeltiosis in the desirable direction for nodal position for first flowering, days to first flowering, days to first picking, plant height (cm), number of fruits per plant, average fruit weight (g), number of pickings and total fruit yield per plant (g). (Table 4) The third highest heterotic cross JOL-21-106 × GJO-3 demonstrated significant standard heterosis in the desirable direction over the standard check for nodal position for first flowering, days to first flowering, days to first picking, number of fruits per plant, days to last picking, number of pickings and total fruit yield per plant (g). The same cross also expressed significant heterobeltiosis in the desirable direction for nodal position for first flowering, number of fruits per plant, days to last picking, number of pickings and total fruit yield per plant (g). (Table 4) It is interesting to note that the crosses exhibiting high heterosis for fruit yield per plant did not necessarily express high heterosis for all the associated yield components (Table 4). Nevertheless, the present investigation revealed that significant and positive heterosis for fruit yield was generally accompanied by desirable and significant heterotic effects for two or more yield contributing characters. Such a combined influence of heterosis in multiple yield components contributing to fruit yield per plant has also been reported by Nagesh et al. (2014) [19] , Patel and Patel (2016) [24] , Pithiya et al. (2019) [28] , Kharat et al. (2022) [12] , Poluru et al. (2023) [30] , Sakriya et al. (2023) [35] and Ranga et al. (2024) [33] .

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