Generation mean analysis for fruit yield and its components in bottle gourd [Lagenaria siceraria (Mol.) Standl.]

Introduction

Vegetables constitute an indispensable component of Indian agriculture and nutritional security owing to their short crop duration, high productivity, economic profitability, and rich nutritional composition. India's diverse agro-climatic conditions facilitate year-round cultivation of a wide range of vegetable crops, contributing significantly to agricultural diversification and rural employment generation. During 2024-25, horticultural crops occupied approximately 29.09 million hectares with a production of 367.70 million tonnes, reflecting sustained growth in the sector (Anonymous, 2024) [8] . Vegetables are important dietary sources of vitamins, minerals, proteins, dietary fibre, and bioactive phytochemicals possessing antioxidant and anti-carcinogenic properties. Rapid urbanization, increasing health awareness, changing dietary preferences, and rising per capita income have further accelerated vegetable production and consumption in India. Among vegetable crops, cucurbits occupy a prominent position due to their adaptability, high yield potential, and economic importance, particularly for small and marginal farmers. The family Cucurbitaceae comprises nearly 118 genera and 825 species worldwide, of which India harbours considerable diversity with several endemic species. Bottle gourd [Lagenaria siceraria (Mol.) Standl; 2n = 2x = 22] is one of the oldest domesticated cucurbitaceous vegetables and an important monoecious, cross-pollinated crop cultivated extensively throughout tropical and subtropical regions. It is valued not only for its culinary utility but also for its medicinal, nutritional, and industrial significance. The fruits are easily digestible and rich in carbohydrates, minerals, vitamins, and moisture, while different plant parts possess therapeutic properties against several ailments. Dried fruits are also utilized for preparing utensils, containers, and musical instruments. Despite its economic importance, bottle gourd productivity in many regions of India, including Gujarat, remains constrained due to cultivation of genetically heterogeneous local cultivars with limited yield potential.

Although improved cultivars and hybrids such as Anand Bottle Gourd-1 and Gujarat Anand Bottle Gourd Hybrid-1 have been released, there remains substantial scope for genetic enhancement of yield and quality traits. Bottle gourd exhibits 60-80 percent natural cross-pollination, facilitating extensive variability and providing immense opportunities for exploitation of heterosis in breeding programmes. Owing to its monoecious flowering behaviour, high seed recovery, low seed requirement, and comparatively low inbreeding depression, the crop is highly amenable to hybrid breeding. Improvement of fruit yield in bottle gourd is genetically complex because yield is a quantitative trait governed by polygenic inheritance and influenced by environmental interactions. Therefore, understanding the nature and magnitude of gene effects is essential for devising efficient breeding strategies. Generation mean analysis provides comprehensive information on additive, dominance, and epistatic gene interactions, including additive × additive, additive × dominance, and dominance × dominance effects, thereby enabling precise identification of suitable breeding approaches such as pedigree selection, recurrent selection, or heterosis breeding. Simultaneously, assessment of heterosis and inbreeding depression assists in identifying superior cross combinations and evaluating the extent of non-additive genetic effects governing economically important traits.

Materials and Methods

Plant material

The experimental materials comprised of six basic generations viz., P 1 , P 2 , F 1 , F 2 , BC 1 and BC 2 of four Crosses namely ABGS 11-24 × P. Komal, ABGS 11-17 × NDBG 132, P. Komal × P. Long and Local × P. Long were made between six parents by manual emasculation and pollen transfer. F 1 plants were selfed to obtain seed for the F 2 generation and backcrossed with their respective parents to generate BC 1 and BC 2 generations. Thus, a total of six generations were obtained.

Field Trial

The experimental material consisted of six parental lines, viz., ABGS 11-24, ABGS 11-17, Punjab Komal, NDBG-132, local, Punjab Long and their F 1 , F 2 , and back crosses (B 1 and B 2 ). Each family was made up of six generations: P 1 , P 2 , F 1 , F 2 , B 1 and B 2 . The seed of parents were taken from the Main Vegetable Research Station, Anand. For the generation mean analysis, six basic generations (P 1 , P 2 , F 1 , F 2 , B 1 , and B 2 ) were developed following standard breeding procedures in bottle gourd (Lagenaria siceraria). The parental lines (P 1 and P 2 ) were first self-pollinated to maintain genetic purity, and seeds thus obtained were used for hybridization. Fresh F 1 seeds were produced by crossing P 1 and P 2 during the rabi 2022-23. The F 2 generation was obtained by selfing the F 1 plants. Backcross generations were produced during the kharif 2023, with B 1 derived from crossing F 1 with P 1 , and B 2 from crossing F 1 with P 2 . This set of six generations formed the basis for subsequent generation mean analysis to determine the nature of gene action governing the traits under study. Seventeen different characters were studied viz., days to first male flower appearance, days to first female flower appearance, node number at which first male flower appearance, node number at which first female flower appearance, number of branches per plant, vine length (m), days to first picking, fruit length (cm), fruit girth (cm), number of fruits per plant, average fruit weight (g), yield per plant (kg), chlorophyll-a, chlorophyll-b, total chlorophyll, moisture content (%) and total soluble solids (ºBrix)

Selfing and crossing technique

In bottle gourd, anthesis takes place from 6 am to 11 am. Thus, the best time for bottle gourd pollination, whether natural or hand-assisted, is typically when the flowers are fully open, which usually occurs in the evening hours. The female and male flower buds likely to open on the next day were enclosed during morning hours with butter paper bags just to protect them from stray pollens. During evening, the bags were removed and female buds having receptive stigma were then pollinated by rubbing the dehisced anthers from the desired male parent. The pollinated flowers were identified and covered with white paper bags. When the F 1 was available, backcrosses were created using the same methodology. To obtain the parents' and the F 2 s' seeds, a few closed buds from the F 1 and parents' plants were selfed.

Statistical analysis

For statistical analysis, the data recorded for each character from the observations made on each representative (tagged) plants of each generation of each family in each replication were used. Statistical analysis was performed for the following traits. The analysis of variance among families (Table 1) indicated significant mean square differences among all the three families (crosses) for all the characters studied. Likewise, the mean squares among progenies within each family revealed that the variations among the six generations were significant for all the characters studied in all the three crosses.

Results and Discussion

Analysis of variance and per se performance

Per se performance of different generations in four bottle gourd crosses revealed the predominance of diverse gene actions for yield and its component traits. F₁ hybrids exhibited overdominance for earliness, vine length, branching, fruit length, average fruit weight, yield per plant, moisture content, and total soluble solids in several crosses, indicating strong heterotic effects and the importance of non-additive gene action. Complete dominance was observed for traits such as node number of flowering, vine length, chlorophyll-b, moisture content, and TSS, whereas incomplete dominance governed fruit girth, fruit number, fruit weight, chlorophyll content, and days to fruit picking, suggesting partial dominance effects. Additive gene action was evident where F₁ means remained at par with midparent values for flowering, vegetative, fruit quality, and chlorophyll traits. F₂ generations showed absence of inbreeding depression for several traits including flowering behaviour, branching, fruit number, chlorophyll content, and moisture content, while significant inbreeding depression for fruit length, fruit weight, and yield per plant confirmed the role of dominance effects. Superior F₂ performance over F₁ for certain characters indicated the occurrence of desirable transgressive segregants. The BC₁ and BC₂ generations recorded superior performance for several yield, vegetative, fruit quality, and biochemical traits, demonstrating the accumulation of favourable alleles and suggesting the effectiveness of backcross breeding for improvement of bottle gourd genotypes.

Components of Generation Means Based on Additive-Dominance Model

Generation mean analysis in four bottle gourd crosses, namely Cross I (ABGS 11-24 × P. Komal), Cross II (ABGS 11-17 × NDBG-132), Cross III (P. Komal × P. Long), and Cross IV (Local × Punjab Long), revealed the involvement of both additive and non-additive gene effects for most of the studied traits. Scaling tests and joint χ² analysis indicated that the additive-dominance model was adequate for certain traits and crosses, whereas inadequacy in others suggested the presence of non-allelic interactions. Significant additive effects were predominantly observed for days to first male flower appearance, number of branches per plant, chlorophyll-a, chlorophyll-b, total chlorophyll, and moisture content, indicating the importance of fixable gene action and the effectiveness of selection. Dominance effects were more pronounced for fruit length, average fruit weight, yield per plant, and total soluble solids, reflecting the role of heterosis and non-additive gene action in the inheritance of these traits.

Components of Generation Means Based on Six-Parameter Model (Hayman, 1958) [43] [43]

The six-parameter model of Hayman (1958) [43] revealed the presence of significant epistatic interactions governing several quantitative traits in bottle gourd. Along with significant mean (m), additive (d), and dominance (h) effects, epistatic components such as additive × additive (i), additive × dominance (j), and dominance × dominance (l) interactions were detected in different crosses and traits.

Duplicate and complementary types of epistasis were observed for fruit length, yield per plant, chlorophyll-b, total chlorophyll, and total soluble solids, indicating the complex inheritance pattern of these characters. The predominance of dominance and dominance × dominance interactions for yield and fruit traits suggested the importance of nonadditive gene action, whereas additive and additive × additive effects for chlorophyll and moisture-related traits indicated scope for effective selection in early generations.

Components of Generation Means

Based on Six-Parameter Model (Cavalli, 1952) [18] Cavalli's six-parameter model further confirmed the predominance of digenic interactions with limited higherorder epistasis across the studied bottle gourd crosses. Significant mean (m) effects were recorded for all traits, indicating quantitative inheritance. Additive gene effects played a major role in flowering behaviour, branching, chlorophyll content, and moisture content, whereas dominance effects were important for fruit length, average fruit weight, yield per plant, and total soluble solids. The significance of epistatic components emphasized the contribution of both additive and non-additive gene actions in trait expression. Overall, the results suggested that improvement of economically important traits in bottle gourd can be effectively achieved through suitable breeding approaches such as pedigree selection, biparental mating, recurrent selection, and exploitation of heterosis for accumulation of favourable alleles and development of superior genotypes. (Cavalli, 1952) [18] (m) 45 (Cavalli, 1952) [18] (m) 41 (Cavalli, 1952) [18] (m) (Cavalli, 1952) [18] (m) (Cavalli, 1952) [18] (m) 50 (Cavalli, 1952) [18] (m) (Cavalli, 1952) [18] (m) 83 (Cavalli, 1952) [18] (m) 0.

Conclusion

the present investigation on generation mean analysis in bottle gourd [Lagenaria siceraria (Mol.) Standl.] was undertaken to understand the genetic control of yield, earliness, and fruit quality traits using six generations (P₁, P₂, F₁, F₂, BC₁, and BC₂) derived from four crosses. Significant variability among generations indicated ample genetic diversity for selection. Per se performance of F₁s exhibited substantial heterosis for vine length, branches per plant, fruit length, average fruit weight, and yield per plant, while reduced F₂ performance for several traits confirmed inbreeding depression and the role of non-additive gene action. Scaling tests revealed the inadequacy of the additivedominance model for most traits, indicating the involvement of epistatic interactions. Generation mean analysis demonstrated the importance of additive, dominance, and digenic epistatic effects in the inheritance of flowering, yield, and quality traits. Additive gene action coupled with high heritability and genetic advance was predominant for earliness traits, suggesting the effectiveness of direct selection, whereas dominance and overdominance effects were more important for yield and associated traits, indicating the potential of heterosis breeding. The occurrence of additive × additive and dominance × dominance interactions suggested the usefulness of biparental mating and recurrent selection, while superior BC₁ and BC₂ performance indicated the effectiveness of backcross breeding for accumulation of favourable alleles and genetic improvement in bottle gourd.

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