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Studies on the inheritance of fruit bursting in Indian Snapmelon (Cucumis melo L. Var momordica)

Published online by Cambridge University Press:  07 November 2025

Rashmi K. Naik ,
Shivapriya Manchali Open the ORCID record for Shivapriya Manchali [Opens in a new window] ,
Ravishankar V. Kundapura  and
Vishnuvardhana
Rashmi K. Naik
Affiliation:
Department of Biotechnology and Crop Improvement, University of Horticultural Sciences, Bagalkot, India
Shivapriya Manchali*
Affiliation:
Department of Biotechnology and Crop Improvement, University of Horticultural Sciences, Bagalkot, India
Ravishankar V. Kundapura
Affiliation:
ICAR–Indian Institute of Horticulture Research, Hesaragatta, Bengaluru, India
Vishnuvardhana
Affiliation:
Department of Biotechnology and Crop Improvement, University of Horticultural Sciences, Bagalkot, India
*
Corresponding author: Shivapriya Manchali; Email: priyachidambara@gmail.com
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Abstract

Snapmelon is an indigenous crop of Indian origin that has contributed resistance genes against major melon diseases. Despite its importance, Snapmelon is an under-utilized crop, and its cultivation may be limited due to its very short post-harvest life, fruit-bursting on maturity and very thin pericarp that is unsuitable for long-distance transportation. Studies on the inheritance of melon fruit bursting are scanty, and according to the available literature, it is due to complementary gene action. The inheritance of fruit bursting in Snapmelon in our work was studied through an inter-botanical cross between Snapmelon (IC632170, bursting fruit type) and Chandalak melon (Kashi Madhu, non-bursting fruit type). F2 and backcross populations were developed, and all three populations were evaluated for fruit bursting under polyhouse conditions for two seasons. Chi-square analysis indicated non-significant deviations from the expected 3:1 Mendelian ratio for non-bursting: bursting type fruits in the F2 population, suggesting the monogenic control of fruit bursting. The backcross with parent Kashi Madhu exhibited 1:0 (non-bursting: bursting) ratios, while the backcross with parent IC632170 revealed 1:1 (non-bursting: bursting) ratio. Selective genotyping in the F2:3 population of Kashi Madhu x IC632170 using 86 polymorphic Simple sequence repeat (SSR) markers revealed the association of the marker CMCT505 for fruit bursting at maturity. Knowledge on the genetic basis of fruit bursting is crucial for designing effective breeding strategies aimed at developing non-bursting Snapmelon varieties suitable for commercial cultivation and improved post-harvest handling.

Keywords

Cucumis melointer-botanical crossmomordicaselective genotypingSnapmelon

Information

Type
Research Article
Information
Plant Genetic Resources , Volume 24 , Issue 2 , April 2026 , pp. 108 - 114
Copyright
© The Author(s), 2025. Published by Cambridge University Press on behalf of National Institute of Agricultural Botany.

Introduction

Snapmelon (Cucumis melo L. var momordica, 2n = 24) is one among the 19 botanical groups of the melons of the Cucurbitaceae family as described by Pitrat (Reference Pitrat, Grumet, Katzir and Garcia-Mas2016). It is native to India, and some of its accessions are known to possess and contribute multiple disease-resistant genes for melon breeding (Perchepied et al. Reference Perchepied, Bardin, Dogimont and Pitrat2005; Pitrat Reference Pitrat, Prohens and Nuez2008; Pitrat and Besombes Reference Pitrat, Besombes and Pitrat2008; Liu et al. Reference Liu, Chen, Su, Zhang and Zhu2010; Dhillon et al. Reference Dhillon, Monforte, Pirate, Pandey and Singh2012; Manchali et al. Reference Manchali, Kavya and Sudhakara2019; Hemanthkumar 2024).

Snapmelon is allogamous (cross-pollinated) due to the monoecious flower type (Pitrat et al. Reference Pitrat, Hanelt and Hammer2000; Manchali et al., Reference Manchali, Mamatha, Umesha, Lingaiah and Mohankumar2021a). Our previous studies involving melons belonging to 11 botanical classes have shown cross-compatibility of melons (including Snapmelon) and the successful fruit set (Chaitra 2019). The cross-compatibility between various botanical groups of melon (Luan et al. Reference Luan, Sheng, Wang and Staub2010; Pandey et al. Reference Pandey, Dhillon, Sureja, Singh and Malik2010; Singh et al. Reference Singh, Pandey, Raghuwanshi, Singh, Jha and Singh2011; John et al. Reference John, Scariah, Nissar, Latha, Gopalakrishnan, Yadav and Bhat2013; Chaitra 2019; Rashmi Reference Rashmi2023) helps in the easy transfer of genes across the botanical groups. Melons are nutritious and vitamin-rich, wholesome fruits (Lester Reference Lester1997; Manchali and Murthy Reference Manchali, Murthy and Jaiswal2020). Snapmelons are low in total soluble solids (Manchali et al. Reference Manchali, Murthy, Vishnuvardana and Patil2021b) and usually consumed with jaggery or sugar. The fruit pulp is mealy and suitable for making juice.

Fruit cracking determines the shelf life of a fruit, and it is very common in Snapmelon. A very thin exocarp of the fruits results in peeling off and splitting of the fruit at maturity (Pitrat Reference Pitrat, Grumet, Katzir and Garcia-Mas2016). The very short post-harvest life due to the fruit-bursting nature of Snapmelon hinders its large-scale cultivation and long-distance transportation, but despite its short shelf life, in Kerala state of Southern India, it is being cultivated at a commercial scale for the juice industry (Pareek et al. Reference Pareek, Vashistha and Samadia1999). Bursting of fruits and skin peeling on maturity are unique characteristics of Snapmelon and various bursting patterns of fruit rind and wide variability for fruit traits are reported in Snapmelon of India (Dhillon et al. Reference Dhillon, Ranjana, Singh, Eduardo, Monforte, Pitrat, Dhillon and Singh2007; Fergany et al. Reference Fergany, Kaur, Monforte, Pitrat, Rys, Lecoq, Dhillon and Dhaliwal2011; Singh et al. Reference Singh, Pandey, Raghuwanshi, Singh, Jha and Singh2011; Pitrat Reference Pitrat, Grumet, Katzir and Garcia-Mas2016; Manchali et al. Reference Manchali, Mamatha, Umesha, Lingaiah and Mohankumar2021a; Hemanthkumar 2024) as well as Brazilian States (Valadares et al. Reference Valadares, Melo, Sarinho, Oliveira, Rocha, Silva and Menezes2018).

Inheritance of melon fruit bursting/cracking is still not very clear, and one of the studies (Singh et al. Reference Singh, Pandey, Raghuwanshi, Singh, Jha and Singh2011) using the F2 population of Snapmelon with a non-bursting fruit type line noticed a complementary gene action (the digenic control of a character with a 9:7 ratio in the F2 population). Considering the importance of post-harvest life of Snapmelon, in the present work, we made an attempt to study the inheritance of fruit bursting and also to find out its association with the molecular markers. Snapmelon accession IC632170 (fruit bursting type) is a landrace collected from a farmer’s field and has resistance to powdery and downy mildew diseases (Manchali et al. Reference Manchali, Kavya and Sudhakara2019; Manchali et al. Reference Manchali, Mamatha, Umesha, Lingaiah and Mohankumar2021a), and the other parent, Kashi Madhu (non-bursting fruit type), is a popular released variety developed through selection from landrace ‘Mau’ melon (Pandey et al. Reference Pandey, Rai, Prasanna and Kalloo2008). These two parents were selected for crossing as they were in contrast with respect to many characters, including the fruit-bursting nature.

In Selective genotyping, DNA markers are tested on the most informative individuals in the higher and lower tails of a mapping population that exhibit extreme phenotypic values. The best (higher 17%) and worst (lower 17%) of the individuals in a mapping population for a particular phenotypic character can account for about 81% of the total linkage information (Lander and Botstein Reference Lander and Botstein1989). It is an alternative productive strategy for understanding the association of a trait and linked markers (Sun et al. Reference Sun, Wang, Crouch and Xu2010). This method of selective genotyping can reduce genotyping expenses without compromising information, but it can be done only for one trait at a time (Lebowitz et al. Reference Lebowitz, Soller and Beckmann1987). Thus, this particular approach is advantageous over the Quantitative trait loci (QTL) mapping, when the time and expense of assaying molecular markers far exceeds phenotyping (Merk et al. Reference Merk, Ashrafi and Foolad2012). Thus, in order to comprehend the inheritance of fruit bursting in Indian Snapmelon, the present study was conducted using the bursting and non-bursting type fruit-bearing parents, their hybrid, F2 and backcross populations from the inter-botanical cross between momordica and chandalak.

Materials and methods

Snapmelon (IC632170) (var. momordica) and Kashi Madhu (var. chandalak) were used to develop the inter-botanical cross F1. Snapmelon accession IC632170 is a landrace sourced from farmer’s field in Karnataka, India (Manchali et al. Reference Manchali, Mamatha, Umesha, Lingaiah and Mohankumar2021a). It has shown resistance to powdery and downy mildew diseases in our previous disease screening experiment (Manchali et al. Reference Manchali, Kavya and Sudhakara2019). It is cultivated as a mixed crop with vegetables. Kashi Madhu is a selection from a local landrace ‘Mau’ melon of Uttar Pradesh, India (Pandey et al. Reference Pandey, Rai, Prasanna and Kalloo2008) and is known for its sweet and attractive orange pulp. The parents were very contrasting with respect to the fruit traits. The Snapmelon (IC632170) fruits were oval in shape, possessed smooth, thin rind without any netting or suture, light-orange colored, thick, mealy flesh that was not sweet. The fruits burst/crack on maturity. The ‘Kashi Madhu’ fruits were oblate in shape with a heavily netted and sutured rind. Pulp was dark orange-colored, consistent and sweet. The fruits were of non-bursting/cracking on maturity.

Development of segregating population

The present study included F2 and backcrosses to study the inheritance of fruit bursting. Parents Snapmelon (IC632170) and Kashi Madhu were crossed to obtain the hybrid (F1). Seeds extracted from the crossed fruits of Snapmelon were planted in the next season. About 30 F1 plants were grown. Selfing of flowers on 10 plants was performed to obtain the F2 individuals. Ten F1 plants were used as a female parent to cross with IC632170 and another 10 to cross with Kashi Madhu to create backcrosses BC1 (P1) and BC1 (P2), respectively. All the crossing and selfing were done under insect-proof polyhouse to avoid contamination (Rashmi Reference Rashmi2023).

Evaluation of segregating populations for fruit bursting:

The two parents and their F1, F2, BC1 (P1) and BC1 (P2) lines were cultivated under the polyhouse condition at the Regional Horticultural Research and Extension Centre, University of Horticultural Sciences, Bagalkot campus, Bengaluru, during the winter season of 2020 and 2021. Parents and F1 were replicated three times, and segregating lines were un-replicated. Spacing of 1.8 m between rows and 0.6 m plant to plant was maintained. In 2020, 201 F2, 50 BC1 (P1) and 87 BC1 (P2) lines; in 2021, 300 F2, 150 BC1 (P1) and 100 BC1 (P2) lines were evaluated for fruit cracking/bursting on maturity. Fruits were harvested when the abscission layer developed and the fruits slipped from the plant. The observation on the individual plant for fruit bursting was recorded. The fruits were visually observed for bursting nature and categorized as bursting or non-bursting.

Chi-square test was performed according to Gomez and Gomez, 1984.

\begin{equation*}\chi^2\ =\ \frac{{\sum {{\left( {O - {\text{ }}E} \right)}^2}}}{E}\end{equation*}

O = observed value; E = expected value.

Identification of the SSR markers linked to fruit bursting using selective genotyping:

Seeds were isolated from the F2 genotypes, and 10 seedlings from each of the selected genotypes (F2:3) were raised in pots during the Kharif season of 2021. Leaf samples from 10 plants of each genotype were bulked to form a single family. A total of 160 F2:3 families were generated for selective genotyping based on bursting on maturity. Genomic DNA was isolated using modified Cetyl Tri methyl Ammonium Bromide extraction buffer method (Ravishankar et al. Reference Ravishankar, Anand and Dinesh2000). The parental lines and F1 were initially screened with 125 SSR markers (Supplemental Table S1) with M13 tail primers (Schuelke Reference Schuelke2000) to assess polymorphism of these; 86 markers were polymorphic and subsequently used to genotype the extreme phenotypes for fruit bursting at maturity. Allelic frequency was calculated using the t-test.

Statistical analysis

A two-sample t-test between non-bursting fruits (best) and bursting fruits (worst) was conducted by using the formula (Lebowitz et al. Reference Lebowitz, Soller and Beckmann1987);

\begin{equation*}{\text{t = }}\frac{{{\text{PBest - PWorst}}}}{{\sqrt {\frac{{{\text{p}}\left( {{\text{1 - p}}} \right)}}{{{\text{2}}{{\text{N}}_{\text{1}}}}}{\text{ + }}\frac{{{\text{p}}\left( {{\text{1 - p}}} \right)}}{{{\text{2}}{{\text{N}}_{\text{2}}}}}} }}\end{equation*}

where P Best is the marker allele frequency of best type, P Worst is the marker allele frequency of worst type, p is the marker allele frequency at 0.50, N 1 is the total number of best type plants and N 2 is the total number of worst-type plants.

Results

The parent Snapmelon (IC632170) (P1) produced bursting fruits (Fig. 1), parent Kashi Madhu (P2), as well as the F1 produced non-bursting fruits in both the seasons (Figs. 2 and 3). In the winter season of 2020, the F2s showed 152 lines with non-bursting matured fruits and 49 lines with bursting types of fruits. In winter season of 2021, among 150 F2s, 105 plants produced non-bursting fruits and remaining plants produced bursting fruits (Table 1 and Fig. 4). The calculated chi-square values were found to be non-significant in both the seasons (χ2 = 0.041 and 2; P = > 0.05) and fitted with the expected Mendelian ratio 3:1 (non-bursting: bursting) depicting a monogenic inheritance of fruit bursting.

Figure 1.

Fruit bursting in Snapmelon (IC632170).

Figure 2.

Fruit of Kashi Madhu.

Figure 3.

Fruit of Snapmelon (IC632170) x Kashi Madhu hybrid (F1).

Figure 4.

Variation for fruit bursting trait in F2 population of Snapmelon (IC632170) x Kashi Madhu.

Table 1.

The genetic variation in F2 and backcross melon populations derived from the inter-botanical cross IC632170 x Kashi Madhu for fruit bursting at maturity

BC1 (P1): F1 back crossed with bursting fruit type parent IC632170; BC1 (P2): F1 back crossed with non-bursting fruit type parent Kashi Madhu; E: expected value; F2: F2 generation; O: observed value.

*P: probability at 0.05%.

In back cross with bursting fruit type parent IC632170, the BC1 (P1) revealed 1:1 (non-bursting: bursting fruit type) ratio in both the seasons (41:46 in 2020 with χ2 = 0.287 and 81:69 with χ2 = 0.96 in 2021) (Fig. 5). The BC1 (P2) (back cross with non-bursting fruit type parent Kashi Madhu) exhibited1 (non-bursting): 0 (bursting) fruit type in both the seasons (50:0 and 100:0 in 2020 and 2021, respectively) (Fig. 6). The probability was more than 0.05 in both the cases.

Figure 5.

Variation for fruit bursting trait in backcross of F1 with Snapmelon (IC632170).

Figure 6.

Variation for fruits in backcross of F1 with Kashi Madhu.

Selective genotyping for fruit bursting

Selective genotyping to find the association of SSR markers with fruit bursting at maturity was performed by selecting the extreme individuals in the F2:3 population of IC632170 x Kashi Madhu. The SSR marker CMCT505 exhibited an allelic percentage of 0.73 for non-bursting fruit nature and 0.26 for bursting nature of the fruit, with the significant t-statistic value (3.640) at P = 0.05 (Table 2). This was the only marker found associated with fruit bursting among the 86 polymorphic SSRs used to genotype/screen the parents and extreme genotypes.

Table 2.

Identification of SSR markers associated with fruit bursting at maturity in the inter-botanical cross IC632170 x Kashi Madhu

Probablity at 0.05%.

Discussion

Snapmelon has been cultivated for a long time in India for its good flavour and mealy-textured pulp that suits juice making. Though the shelf life is very short, fruits are wrapped in an arecanut spathe/betel nut spathe or cloth and sold in the local market. The cross compatibility of Snapmelon with other melon botanical groups (Chaitra 2019) makes it possible to create genetic variability and select for non-bursting fruit type genotypes among the segregating lines.

In the present evaluation of the segregating populations along with the parents and F1 for fruit bursting at maturity, the observed numbers of plants with non-bursting and bursting type fruits at maturity were close to the 3:1 ratio for the F2 in both seasons and exhibited monogenic complete dominance. This was supported by the ratio obtained for backcrosses. In backcross of F1 with Kashi Madhu (non-bursting fruit type parent), no fruit bursting was observed in the progenies. A 1:1 ratio was noticed in a backcross of F1 with Snapmelon (bursting fruit parent). The chi-square values were non-significant (5% level of significance) and indicated that the difference between observed and expected values was due to chance. Singh et al. (Reference Singh, Pandey, Raghuwanshi, Singh, Jha and Singh2011) noticed a complementary gene action for the inheritance of fruit bursting in their study with an F2 population developed for IVMM-3 and B-159 (momordica) parents.

The selective genotyping methodology has been successfully employed to identify molecular markers associated with various agronomic traits across multiple crops. Notable examples are salt-tolerance in tomato (Foolad et al. Reference Foolad, Stoltz, Dervinis, Rodriguez and Jones1997), early blight resistance in wild tomato (Zhang et al. Reference Zhang, Lin, Nino-Liu and Foolad2003), fusarium head blight resistance in barley (Wingbermuehle et al. Reference Wingbermuehle, Gustus and Smith2004), sclerotiniamid-stalkrot resistance in sunflower (Micic et al. Reference Micic, Hahn, Bauer, Melchinger, Knapp, Tang and Schon2005), genomicregions involved in nitrogen use efficiency in maize (Coque and Gallais, Reference Coque and Gallais2006), late blight disease resistance in tomato (Merk et al. Reference Merk, Ashrafi and Foolad2012), resistance to Fusarium oxysporum race1inwatermelon (Lambel et al. Reference Lambel, Lanini, Vivoda, Fauve, Wechter, Harris-Shultz, Massey and Levi2014) and seed protein and oil in soybean (Phansak et al. Reference Phansak, Soonsuwon, Hyten, Song, Cregan, Graef and Specht2016). Despite its widespread application, no studies have yet reported the use of selective genotyping in melon.

In the present study, based on the phenotypic characterization of the F2, extreme genotypes for the fruit bursting were grouped for selective genotyping in F2:3 lines. Our study identified a significant association between the SSR marker CMCT505 and fruit bursting at maturity (t-statistic = 3.640, P = 0.05). The marker showed distinct allelic frequencies between phenotypes: 0.73 in non-bursting fruits versus 0.26 in extreme bursters. In one of the previous studies, Singh et al. (Reference Singh, Sanjeev, Singh, Rai, Singh and Pandey2015) used IVMM-3 and a different accession of Snapmelon (B159) to develop F2:3 because of their contrasting nature and identified five QTLs for fruit characters such as fruit length, fruit weight, number of fruits per plant and ascorbic acid content, but loci associated with fruit cracking were not identified.

The marker CMCT505, which is linked to fruit bursting in our study, is associated with varying fruit characters in different genetic backgrounds. For example, marker CMCT505 is associated with fruit diameter and seed cell diameter on LG 1 from a cross between USDA-846-1 and Top Mark (Paris et al. Reference Paris, Zalapa, McCreight and Staub2008) and flesh thickness on LG 1 for F2 mapping population of cantaloupe and snake melon (Ramamurthy and Waters Reference Ramamurthy and Waters2015). The same marker was mapped on LG 6 in a study involving Piel de Sapo cultivar (PS) and PI 161375 (Monforte et al. Reference Monforte, Oliver, Gonzalo, Alvarez, Sanjuan and Arus2004); LG 3 (Zalapa et al. Reference Zalapa, Staub, McCreight, Chung and Cuevas2007), but there was no association of this marker with any of the fruit traits in both the studies. Different accessions of Snapmelon of Indian origin have been used as one of the parents in mapping QTLs related to powdery mildew (Cui et al. Reference Cui, Siskos, Wang, Schouten, Visser and Bai2022). To date, there are no studies that have successfully linked molecular markers to its fruit-cracking trait. Similarly, although momordica accessions have been employed to map fruit firmness QTLs (Nimmakayala et al. Reference Nimmakayala, Tomason, Abburi, Alvarado, Saminathan, Vajja, Salazar, Panicker, Levi, Wechter and McCreight2016) and other studies have identified QTLs for ethylene regulation and other traits (PérinC et al., Reference PérinC, Hagen, Dogimont, Pech and Latché2002; Moreno et al. Reference Moreno, Obando, Dos-Santos, Fernández-Trujillo and Monforte2008), the genetic basis of fruit cracking in Snapmelon remains unexplored.

Information regarding the inheritance of fruit bursting in Snapmelon is scanty, and our study helps in understanding the inheritance of fruit bursting at maturity in Snapmelon, as well as the possible association of SSR marker CMCT505 with this trait. Further, it aids in planning for breeding programme to develop Snapmelon lines with non-bursting fruits. Our study also resulted in the creation of wide genetic variability in the segregating populations for various plant and fruit morphological characters. Further selection of lines with non-bursting fruit type in the advanced recombinant lines is being done. Validation of molecular marker CMCT505 for its association with fruit bursting is also necessary, as it is reported to be mapped to various loci on chromosomes in different studies.

Supplementary material

The supplementary material for this article can be found at https://doi.org/10.1017/S1479262125100361.

Acknowledgements

None.

Ethical standards

This study was partially funded by the student funds from the University of Horticultural Sciences, Bagalkot, Karnataka state, India.

Conflict of interest

Nil.

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Figure 0

Figure 1. Fruit bursting in Snapmelon (IC632170).

Figure 1

Figure 2. Fruit of Kashi Madhu.

Figure 2

Figure 3. Fruit of Snapmelon (IC632170) x Kashi Madhu hybrid (F1).

Figure 3

Figure 4. Variation for fruit bursting trait in F2 population of Snapmelon (IC632170) x Kashi Madhu.

Figure 4

Table 1. The genetic variation in F2 and backcross melon populations derived from the inter-botanical cross IC632170 x Kashi Madhu for fruit bursting at maturity

Figure 5

Figure 5. Variation for fruit bursting trait in backcross of F1 with Snapmelon (IC632170).

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Figure 6. Variation for fruits in backcross of F1 with Kashi Madhu.

Figure 7

Table 2. Identification of SSR markers associated with fruit bursting at maturity in the inter-botanical cross IC632170 x Kashi Madhu

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