Review On Advance Breeding and Biotechnological Approaches For Muskmelon Improvement
Review On Advance Breeding and Biotechnological Approaches For Muskmelon Improvement
Review On Advance Breeding and Biotechnological Approaches For Muskmelon Improvement
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Dr. Priyanka Solanki Dr. Priyanka Solanki, Kishorkumar GK, Ajeet Kumar, Rajat Rajput,
Assistant Professor, Department of Ravi Shankar, Rishabh Kumar Singh, Salma Roshni and Jaydeep Panda
Social Science, College of
Horticulture and Forestry,
Jhalawar, Agricultural University, DOI: https://doi.org/10.33545/26174693.2024.v8.i9Si.2218
Kota, Rajasthan, India
Kishorkumar GK Abstract
Ph.D., Scholar, Department of Melon (Cucumis melo L.) is a warm season old-world cucurbit species which a member of family
Vegetable Science, University of Cucurbitaceae is extensively cultivated for its fleshy fruits. Melons exhibit considerable variation in
Horticultural Sciences Bagalkot,
their phenotypic and biochemical traits, depending on the agroclimatic zones of cultivation and regional
Karnataka, India
preferences. The following aims to assess the progress and potential outcomes of melon breeding
Ajeet Kumar programs utilizing the latest techniques. Significant milestones have been reached in melon breeding
Ph.D., Scholar, Department of over the past century, and this progress is expected to continue. However, further research is needed to
Horticulture (Vegetable Science), uncover new genetic insights related to challenges posed by climate change. The identification of
School of Agricultural Sciences,
Nagaland University, Medziphema valuable hereditary and also metabolic variability in the form of landraces and melon wild relatives will
Campus, Nagaland, India be useful for harvest diversification and also for the broadening of the cultivated melon genetic base.
Whereas, considerable information on genomics, and melon metabolomics, is beneficial for dissecting
Rajat Rajput the basis of the inheritance of important traits. Overall, we hope the manuscript is going to serve as a
Ph.D., Scholar, Department of
Horticulture (Vegetable Science),
crucial resource for the melon breeders. The development of plant biotechnological tools for melons
School of Agricultural Sciences, offers the prospect to develop new varieties, more rapidly, avoiding natural genetic barriers. The use of
Nagaland University, Medziphema these methods has extended to increase the genetic diversity by somatic hybridisation or gene transfer
Campus, Nagaland, India and to optimise conventional breeding programmes.
Ravi Shankar
Ph.D., Scholar, Department of Keywords: Breeding, diversity, genetic engineering, genomics, male sterility, melon, QTLS
Horticulture (Plantation, Spices,
Medicinal and Aromatic Crops),
School of Agricultural Sciences,
Introduction
Nagaland University, Medziphema Muskmelon, also known as cantaloupe (Bailey, 1976) [5], is a hot-season crop belonging to
Campus, Nagaland, India cucurbitaceous family. It can be widely grown for its edible flesh (Kesh and Kaushik, 2020)
[43]
Rishabh Kumar Singh
. The plant has a size of genome 454 Mega basepair (Garcia-Mas et al., 2012) [27] and has
Ph.D., Scholar, Department of grown globally in temperate, subtropical, and tropical regions. Major producers include
Horticulture (Vegetable Science), China, the USA, Spain, Turkey, and Iran. Due to variations in climate and local preferences,
School of Agricultural Sciences,
Nagaland University, Medziphema muskmelons exhibit significant diversity in their substantial, physiological, and
Campus, Nagaland, India morphological traits. (Silberstein et al., 2003) [78]. A closest relative of the muskmelon,
Cucumis picrocarpus F. von Mueller, is found within Aussie Land, with their ancestor
Salma Roshni
M.Sc. Scholar, Department of originating from Bharat. The primary centre of origin for cultivated melon are believed to be
Vegetable Science, College of Southwest Asia and Central Asia. Secondary centre of origin include China, Korea, and the
Horticulture and Forestry, Central
Agricultural University, Pasighat,
Iberian Peninsula (Esquinas-Alcazar and Gulick, 1983) [20]. Muskmelon has undergone
Arunachal Pradesh, India human selection and plant breeding since ancient times, leading to significant varietal
improvements through ancient breeding approaches. However, highly significant sexual
Jaydeep Panda
M.Sc. Scholar, Department of incompatibility challenges during the interspecific and intergeneric methods have limited this
Silviculture and agroforestry, genetic probable in respect of developing noval and improved muskmelon varieties.
College of Horticulture and Forest, Enhancing muskmelon varieties through old hybridization is a cpmparatively moderate
Acharya Narendra Dev University
of Agriculture Technology, process and other is limited to a narrow genice pool. The advancement associated with plant
Ayodhya, Uttar Pradesh, India biotechnological tools regarding muskmelons presents opportunities to create noval species
extra quickly, bypassing natural genetic carrier. Techniques such as somatic hybridization
Corresponding Author: and gene transfer were been developed to enhance genetic diversity and optimize
Rajat Rajput conventional breeding programs through the construction of genetic maps.
Ph.D., Scholar, Department of
Horticulture (Vegetable Science),
The primary aim of cucurbitaceous vegetable research is to enhance long-term productivity
School of Agricultural Sciences, by developing high-quality varieties and hybrids those resistant to biotic and abiotic stresses.
Nagaland University, Medziphema Using superior with inherent inbred and hybrid vigour, combined with modern vegetable
Campus, Nagaland, India
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technology and effective government policies, can lead to a unaffected by environmental conditions and show a greater
steady increase in output. Like other vegetables, cucurbits degree of polymorphism. This enables more effective
are susceptible to various biotic stresses such as diseases, utilization of plant traits in breeding varieties with enhanced
insect pests, nematodes, and weeds. Developing cultivars yield, quality, and resistance to biotic and abiotic stresses.
resistant to these stresses through conventional breeding Additionally, A genetic map using morphological markers
methods is difficult and time-intensive, as strains, races, and has been developed for C. melo. (Garcia et al., 1998;
pathotypes quickly mutate to bypass resistance. Silberstein et al., 1999) [26]. Initial muskmelon mapping
Throughout the extensive background regarding plant efforts with classical markers were restricted regarding
manipulation, 4 primary for methods for plant breeding have varietal surveys (Esquinas-Alcazar, 1977; Perl-Treves et al.,
been innovated along with utilized: 1985) [19]. A initial muskmelon gene map has created with
1) Traditional breeding through crossing with selection the use of 23 markers across 8 linkage groups, focusing on
2) Plant breeding derived from mutations disease resistance, reproductive biology, and vegetative
3) Crop breeding using transgenes traits by Pitrat (1991) [92]. Updated information on mapped
4) Plant breeding utilizing genome editing (Hickey et al. genes for melon was provided. Relying solely on a
2019) [35]. phenotypic marker is inadequate for building a
comprehensive map because of its low occurrence in the
Traditional breeding methods, Hybridization as well as genome and variable expression. In 1996, Baudarcco-Arnas
mutation-based breeding methods typically necessitate long and Pitrat created the initial genetic map of melon using
periods to yield results besides involve extensive labor Random Amplified Polymorphic DNA (RAPD) and
(Chen et al. 2019) [14]. Genetically modified plant breeding Restricted Fragment Length Polymorphism (RFLP)
has advanced Acceleratedly over the past century, emerging molecular markers. The second map, featuring 188
like a highly promising method to combine multiple predominantly Amplified Fragment Length Polymorphism
desirable traits into one variety. However, this technology (AFLP) markers, was developed by Oliver et al. (2001) [59].
has sparked considerable controversy early on, primarily The melon genome was mapped using 400 markers, leading
due to concerns over safety and ethical considerations to the creation of a comprehensive composite map featuring
(Prado et al. 2014) [68]. 12 linkage groups. This high-density map was generated
using a recombinant inbred line (RIL) population and a total
Botanical groups of muskmelon of 668 markers, incorporating AFLP, inter-microsatellite
Cucumis melo is a broad and varied taxonomic group, amplification, and phenotypic markers. Gonzalo et al.
comprising numerous Botanical and horticultural (2005) [31] Another composite melon map was also
classifications. The cultivated varieties muskmelon was established using Simple Sequence Repeats (SSR) markers.
classified within 9 Clades and single natural form by naudin The mapping populations in these studies usually segregate
in 1859, whose work served as the foundation for all with respect to disease resistance genes and various other
subsequent studies on the subject. Munger and Robinson horticultural characteristics.
(1991) [55] adaptation of an easier classification of Naudin’s
taxonomy, categorizing C. melo into a single wild variety, Somaclonal Variations
C. melo var. agrestis, along with six cultivated types: The genetic variability in melons facilitates the integration
cantalupensis, inodorus, conomon, dudaim, flexuosus, as associated with horticultural and resistance characteristics
well as momordica. Kirkbride (1993) [44] further divided C. from wild into horticultural varieties using traditional
melo classified into two subspecies according to ovary breeding methods. However, interspecific crosses between
pubescence: C. melo subsp. Agrestis and C. melo sub sp. C. melo and Cucumis metuliferus attempted by Norton and
Melo. Granberry (1980) [93] did not yield reproducible results.
Protoplast fusion via somatic hybridization offers an
Breeding objective alternative method to hybridize cucurbit species that are
When developing superior melon varieties and hybrids, key otherwise cross-incompatible. For example, a somatic
traits to focus on include highly production, uniform fruit hybrid between C. melo and Cucumis myriocarpus has been
shape and size, early maturity, a durable netted skin with a achieving by mesophyll protoplast fusion. Another Using
few otherwise negligible presence of seed cavity, a mild protoplast fusion, a somatic hybrid between melon and
musky flavour, and attractive out-side colour. As a dessert pumpkin was developed, confirmed by isozyme patterns and
fruit, important quality parameters are TSS, consistent flesh the number and shape of chromosomes. The taste profile of
thickness, texture, and colour. The ideal sugar content the hybrid resembled that of muskmelon, indicating that its
should range from 11% to 13%, with a minimum of 10%. hybrid characteristics decreased as it reached maturity. This
Additionally, varieties as well as hybrids must be disease- method is useful for transferring desirable genes from wild,
resistant such as powdery mildew, downy mildew, as well cross-incompatible species into cultivated muskmelons.
as mosaic virus, and insect pests like the red pumpkin beetle
and fruit fly. Genetic engineering
Convetional breeding has acted a crucial function in
Approaches of advanced breeding technology in enhancing muskmelons in terms of production, quality, and
muskmelon resistance to diseases as well as pests. This approach was
Genetic Variation and Gene Map significantly facilitated the creation of new varieties.
The majority morphological traits are heavily affected by However, it faces challenges such as sexual incompatibility
ecological conditions, which reduce their reliability of using in crosses between different species and genera, as well as
phenotype to evaluate genetic diversity. In contrast, time-consuming processes. The construction of low-caliber
molecular markers based on DNA sequence variations are interspecific hybrids traits. In recent years, technologies
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including as marker-assisted breeding and genetic Mapping molecular linkages and identifying genes of
engineering have greatly advanced genetic enhancement interest
Along with varietal development, overcoming the In muskmelon, the genes responsible to respect disease
challenges associated with conventional breeding methods resistance as well as horticulture characteristics have been
The development of transgenic approaches have been mapped. The process of identifying marker(s) closed linked
effectively utilized in numerous plant, including to a specific gene of interest is referred to as gene tagging,
muskmelons (Bezirganoglu et al., 2014) [10]. Muskmelons which precedes gene mapping. Genetic map involves
are significantly threatened by fungal diseases, viral determining the precise location of a marker on a
infections, and insect pests, resulting in substantial losses in chromosome. The identification of markers associated with
production. (Kwon et al., 2001) [46]. Fungal disorders monogenic traits relies on understanding the Mundelein
affecting melons, such as downy mildew, powdery mildew segregation of the character and involves screening a limited
on leaves, and soil-borne pathogens like Rhizoctonia solani point of samples with a relatively large number of markers.
causing damping-off, root and stem rot, along with The many segregate analysis types is a potent application
Fusarium oxysporum leading to wilting, pose significant used to identify markers that are linked to specific genes
threats to production (Bezirganoglu et al., 2014) [10]. Viruses (Michelmore et al., 1991) [53]. This method, individuals from
present a substantial challenge, causing extensive damage to a segregating population are grouped based on their
muskmelon crops. Over In the previous thirty years, 2 phenotypic expression, and these pools are subsequently
primary methods have been employed to cultivate virus- 753atalyse to obtain an adequate number of markers for
resistant muskmelon varieties, including coat-protein both dominant and recessive monogenic traits. Research
mediated resistance. efforts have primarily aimed at identifying markers
Researchers have developed transgenic muskmelon associated with disease resistance and quality traits in
melon. Only one of the most destructive diseases worldwide
genotypes by introducing overexpressed coat protein genes
affecting melons is Fusarium oxysporum f. sp. Melonis.
from viruses like WMV, PRSV, and ZYMV. This
Wechter et al. (1995) [85] the Fom-2 gene, which confers
technique, known as coat-protein mediated protection or
resistance to Fusarium wilt caused through Fusarium
resistance, has proven effective in opposition to several
oxysporum f. sp. Melonis, was tagged using RAPD
viruses (Yoshioka et al., 1992; Fang and Grumet, 1993;
molecular markers in the MR-1 × AY population. A
Gonsalves et al., 1994) [88, 22, 30]. Plants expressing the
flanking marker, 596, was identified at 2 cM and later
amino-truncated core portion or antisense CP-ZYMV
converted into a Sequence Characterized Amplified Region
exhibited limited protection, while those with the full-length (SCAR) marker (Wechter et al., 1998; Wang et al., 2000) [86,
CP-ZYMV gene demonstrated high resistance (Fang and 84]
. Two additional flanking markers, AAC/CAT1 and
Grumet, 1993) [22]. Similarly, observed ribozyme-mediated ACC/CAT1, were found at 1.7 and 3.3 cM, respectively, for
resistance in muskmelon against CMV, while Huttner et al. Fom-2 in the same population (Wang et al., 2000) [84]. The
(2001) [37] reported resistance against 753atalyse yellow Fom-2 gene was mapped using RAPD markers, with
mosaic virus” and “watermelon mosaic virus” using the flanking markers E07-1.3 and G17-1.0 identified from the
same method. This resistance mechanism relies on Vedrantais × PI 161375 cross.
ribozymes, small RNA molecules that specifically cleave Brotman et al. (2002) [12] The Fusarium wilt resistance gene
viral RNA. Bordas et al. (1997) [11] demonstrated enhanced Fom-1 (effective against races 0 and 2) was mapped, with
salt tolerance in muskmelon by introducing the transgene flanking markers NBS 47-3 and K-1180 identified, spaced
HAL 1 from Saccharomyces cerevisiae. Overexpression of 2.6 and 7.0 cM apart, respectively. Several molecular
HAL 1 helped maintain high potassium levels, low sodium markers have been discovered for various virus resistance
content, and an increased potassium/sodium ratio (Serrano, genes in melon.
1996) [73]. The Zym-1 gene, conferring resistance to Zucchini yellow
Melon fruit is high-rich in vitamins and folic acid, which mosaic virus, was mapped in muskmelon using the SSR
serve as potent antioxidants and essential compounds in marker CMAG36 (Danin-Poleg et al., 2002) [16].
human metabolic reactions. The ripening process in Additionally, the Vat gene, providing resistance against
muskmelon includes a series of changes in texture, color, muskmelon Aphis gossypii, has mapped with RFLP flanking
aroma, firmness, and flavour (Lelievre et al., 1997; Jiang markers NSB 2 and AC 39 identified at 31 and 6.4 cM,
and Fu, 2000) [48, 38]. Ethylene is recognized as a crucial respectively. Apart from disease resistance, several
ripening factor in climacteric fruits (Giovannoni, 2001). monogenic agronomic traits have also been mapped in
Two essential enzymes in the ethylene biosynthesis pathway melon. Danin-Poleg et al. (2002) [16] mapped the pH locus,
are ACC synthase (ACS), which converts S-adenosyl which determines flesh acidity in melon, using an SSR
methionine into 1-aminocyclopropane-1-carboxylate (ACC), marker. They identified the flanking marker CMAT141 at
and ACC oxidase (ACO), which converts ACC into 1.7 cM and another marker, 269-0.9, at 8.7 cM on the
ethylene (Fluhr and Mattoo, 1996; Johnson and Ecker, opposite position. Perin et al. (1998) [64] recognition the
1998) [24, 39]. marker EIU5 at 6 cM for the p gene, which confers the
To improve fruit quality, reduce ethylene production, and development of five carpels in the flower.
delay ripening in muskmelon, antisense technology Silberstein et al. (2003) [78] mapped the gene a, responsible
targeting the ACS and ACO genes was employed. (Ayub et for a monoecious morphological with stamen less flowers in
al., 1996; Guis et al., 1997; Silva et al., 2004) [3-4, 32, 79]. the homozygous recessive state, contrasting with
Transgenic methods have also been used to genetically hermaphrodite flowers. They identified a flanking RFLP
enhance other important traits, such as fruit development, marker, CS-DH 21, at 7 cM. Other significant horticultural
sucrose content, aroma, and female flower development traits, such as non-yellowing and extended shelf life
(Zhang et al., 2014; Kocaman et al., 2016) [89, 45].
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(Touyama et al., 2000) [83], and the ms-3 gene were also resistance, appropriately anticipating disease phenotypes in
mapped in melon. the population. Molecular-assisted selection has been
Mapping of QTLs successfully demonstrated for phenotypic testing of Fom-2
Many agronomic traits are controlled by quantitative trait gene-linked markers (Wang et al., 2000; Burger et al., 2003)
[84, 13]
loci (QTLs). Analyzing flanking molecular markers for . Comparative studies between the muskmelon genetic
quantitative trait loci helps simplify the complicated code and Arabidopsis thaliana have reported local gene
inheritance of polygenic traits into Mendelian-like factors, collinearity, suggesting that A. thaliana ESTs could be
aiding in selection. Recombinant inbred lines (RILs), near- adjusted regarding melon genetic research. Many identified
isogenic lines (NILs), or any segregating population of wide genes responsible for disease resistance and horticultural
cross are ideal for QTL mapping due to their near- traits have been mapped and are utilized in MAS, including
homozygosity and minimal environmental effects, which an emphasis on discovering novel genes and its respect to
increase the accuracy of QTL detection. The genes fruit quality and disease resistance.”
underlying QTL can be cloned, and identified markers can
be used in molecular breeding programs. Quantitative trait Important traits for melon breeding
loci -controlled horticultural traits can be introduced into In melon breeding, the primary focus is on traits that affect
desirable parents through molecular marker-assisted yield/output and standard, encompassing disease resistance,
selection, similar to Mendelian inherited traits. Perin et al. fruit attributes such as quality, sugar content, aroma,
(2002) [64] mapped QTLs for fruit length (fl) and fruit shape nutrition, and fruit colour. Consequently, numerous studies
(fs), identifying eight QTLs for fruit shape (fs1.1, fs2.1, have aimed to determine the Genetic sequences responsible
fs8.2), and concluded that fruit shape is generally for these characteristics. This genetic information is crucial
determined during early ovary development as quantitative for selecting target genes for introducing desired
trait loci for fruit and ovary shape are cosegregated. They characteristics into melons through genome editing. This
also mapped quantitative trait loci for fruit ethylene section provides an overview of the genetic elements or
production and detected four QTLs (eth1.1, eth2.1, eth3.1, genetic markers associated with key characteristics in
eth111.1), tightly linked to the ethylene receptor gene ERS1. muskmelon breeding.
Monforte et al. (2004) mapped QTLs for earliness, total
soluble solids, fruit weight, and fruit shape from the PI Virus Resistance
161375 × Pinyonet Piel de Sapo population, identifying nine Cucumber virus (CMV)
QTLs for earliness, eight for fruit shape, six for fruit weight, The Cucumber mosaic virus triggers distinctive mosaic
and five for TSS. Singh et al. (2015) [80] reported five symptoms on muskmelon leaves, causing plant stunting and
quantitative trait loci for fruit traits, including fruit length, mottling or mosaic patterns on fruits, which can lead to
weight, number of fruits per plant, and ascorbic acid content reduced yields. Known for its ability to infect a wide range
in an intraspecific mapping population of melon. Eduardo et of hosts, this virus has been studied extensively through
al. (2004) [17] developed three NILs from PI 161375 × major QTL mapping, identifying the cmv1 gene as
Pinyonet Piel de Sapo to verify the effect of alleles at fs1.1, conferring complete recessive resistance against CMV
fs9.1, and fs11.1 on fruit shape. NILs carrying alleles at subgroup II strains (Essafi et al. 2009) [21]. For resistance to
fs1.1 and fs9.1 (PI 161375) produced more elongated fruits subgroup I strains, Two or more additional quantitative trait
than those with a Pinyonet Piel de Sapo background, with loci (cmvqw3.1 and cmvqw10.1) are required (Guiu-
fs1.1 alleles promoting longitudinal growth and fs9.1 alleles Aragonés et al. 2014) [33]. The cmv1 gene contains the gene
causing transversal growth. Quantitative trait loci in favour responsible for vascular protein sorting 41 (CmVPS41).
of disease resistance in melon have also been identified. Polymorphisms in protein subunit (amino acid) residues
Lately, Perchepied et al. (2005) [63] mapped the Fusarium (L348R or G85E) have been linked to a resistance
wilt resistance gene Fom-1.2 (Fusarium wilt race 1.2), phenotype (Giner et al. 2017, Pascual et al. 2019) [28, 61].
detecting nine QTLs in five linkage groups include Isabelle These mutations alter the intracellular distribution of
(resistant) and Vedrantais (susceptible) parents. CmVPS41, restricting viral movement within bundle sheath
cells and providing resistance to CMV (Pascual et al. 2019,
Genetic Mapping and Molecular Selection Real et al. 2023) [61, 69].
Several disease resistance genes were cloned in muskmelon.
Downy mildew resistance is controlled through few Zucchini mosaic virus (ZYMV)
dominant complementary genes (At1 and At2), which were Zucchini mosaic virus causes vein clearing and yellowing,
cloned by Balaas et al. (1992) [6]. Joobeur et al. (2004) [40] blisters, and enations on leaves, as well as Severe dwarfing
cloned the Fom-2 gene, providing resistance to Fusarium (stunting) in plants. On fruits, it results in mosaic or necrotic
wilt in muskmelon. The Vat gene inhibits muskmelon aphid cracks, marbling, tissue firmness. Pathotype F distinct
colonization as well as prevents the spread of distinct CMV samples can cause melon plants to wilt with the Fn gene
and potyviruses. Pauquet et al. (2004) [62] isolated the Vat (Risser et al. 1981) [70] Here’s an alternative phrasing:
gene, which is 6 kb in length with three introns and encodes “Instead of inducing mosaic symptoms in melons
a protein of 1473 amino acids, and verified its function in 754atalysed754 the Fn+ allele. The genetic locus known as
susceptible melon varieties. Resistance genes, once cloned, Fn, associated with limp necrosis, are found in many
can be incorporated into susceptible lines via transformation muskmelon accessions. Researchers have identified 3
or traditional breeding facilitated by marker-assisted prospective candidates’ resistance loci: Zym-1, Zym-2, and
selection. The effectiveness of MAS can be enhanced by Zym-3. A comprehensive examination of the Zym-1 locus
converting markers to SCARs or sequence-tagged locations. uncovered genes conferring resistance gene- NBL-1
Wang et al. (2000) [84] used the Fom-2 marker to screen 45 (MELO3C015354), NBL-2 (MELO3C015353), and NBL-3,
melon genetic profiles in regards to Fusarium wilt characteristic of the coiled-coil nucleotide-binding site
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found in the leucine-rich repeat family (CC-NB-LRR, identified to provide resistance to these pathogens.
abbreviated as NBL). Furthermore, Two NAC family
transcription factors (MELO3C015355 and Powdery mildew
MELO3C015357) are situated in the neighbouring region Muskmelon powdery mildew is generally attributed to
(Adler-Berke et al. 2021) [1]. Podosphaera xanthii (Px) and Golovinomyces
cichoracearum (Gc) (Li et al. 2017) [49]. G. cichoracearum
Resistance to other diseases is commonly observed in temperate and cold field
Powdery mildew, Fusarium head blight, gummy stem blight environments, whereas P. xanthii is predominantly found in
and downy mildew are significant challenges in muskmelon subtropical and tropical regions, as well as in greenhouse
cultivation, fungal-induced, oomycetes, and thread-like crops. P. xanthii is increasingly becoming the primary
fungi. These pathogens thrive in hot and humid pathogen in most countries (Bardin et al. 1999) [7]. More
environments, making them prevalent in greenhouses where than 28 physiological races of P. xanthii have been
most melons are grown. Consequently, these diseases identified through the reactions of isolates to melon
reduce muskmelon yield and cause revenue declines. differential lines (McCreight 2006) [52]. As new races
Governing these diseases is a crucial aspect of melon emerge, previously resistant melon varieties are becoming
breeding. Unlike viral diseases, fungal, oomycete, and mold susceptible. Therefore, it is crucial to develop varieties with
infections can be somewhat managed with fungicides. multiple race-specific resistance genes as well as field
However, due to the ecological and health hazards linked to resistance genes. So for, several genes and QTLs linked to
fungicide use, incorporating resistance genes into melons is powdery mildew resistance have been mapped in different
highly beneficial. Implementing resistance to these melon populations originating from diverse genetic
pathogens is crucial for successful melon breeding. backgrounds (Haonan et al., 2020) [34]. The identified QTLs
Extensive genetic resources have been studied, and have been listed by Lopez-Martín et al. (2022) [51].
numerous efforts have been made to identify resistance
genes. This section highlights the loci, genes, and markers
Table 1: Identified powdery mildew resistance loci
resistance. Consequently, MELO3C010403 is considered and ‘VED’ revealed 166 Quantitative Trait Loci, with single
The prime candidate gene for GSB resistance. prominent cluster located at chromosome 8, coinciding to
the ripening-related Quantitative Trait Loci ETHQV8.1.
Table 2: Identified gummy stem blight resistance loci Quantitative Trait Loci associated with esters, lipid-derived
Loci Hereditary type Plant introduction accession volatiles, as well as carotenoid metabolites have been
Gsb-1 Dominance PI140471 specifically detected. Target genes were suggested in favour
Gsb-2 Dominance PI157082 of the biosynthesis of compounds such as 3-(Methylthio)
Gsb-3 Dominance PI511890 propanoic acid ethyl ester. Additionally, Near-Isogenic
Gsb-4 Dominance PI482398 Lines involving overlapping introgressive segments through
Gsb-5 Recessive PI420145 such Korean accession ‘SC’ demonstrated an involvement
Gsb-6 Dominance PI482399 of the climacteric maturity Quantitative Trait Loci
(Nonaka, S. and Ezura, 2024) [57] ETHQB3.5 in VOC Metabolic synthesis. Specifically,
compounds such as 3-methylbutyl acetate, benzyl acetate, 2-
Fruit trait methylbutan-1-ol, 2-methylpropyl acetate, and 1-
Melon, a crucial crop both economically and agriculturally, methylsulfanylbutan-1-one showed changes (Zhao et al.
exhibits significant diversity in traits such as pericarp and 2023) [91].
fruit flesh colour, nutritional content, sugar levels, aroma,
and ripening characteristics. These traits play pivotal roles Shelf-life
in fruit quality and are paramount in breeding programs. Improving the shelf-life of fruit crops is vital due to its
Numerous studies have focused on understanding and direct impact on reducing edibles decline and residue.
improving these traits, including fruit flesh colour, sugar Approximately 14 per cent of global food production occurs
content, aroma, and ripening. Shahwar et al. (2023) [74] prior to loss reaching retail, with An extra 17 per cent is lost
reviewed several of these studies and identified numerous at the retail and consumer stages, amounting to USD 400
QTLs associated with these traits. Below, we present the billion annually (English et al., 2019; Forbes et al., 2021) [18,
genes identified in relation to fruit traits. 25]
. This issue is particularly critical for fruits and
vegetables, which have shorter Comparative shelf life with
Fruit colour cereals and non-perishable items items (English et al., 2019)
Pair of genes, CmOr and CmFBN1, play crucial roles in [18]
. Enhancing plant genetics to extend shelf-life not only
determining the fresh orange color of melon fruits. Melons enhances food sustainability but also reduces energy-
exhibit three primary fruit colours of orange, green, and intensive processes like temperature and humidity control
white. The orange color arises from the buildup of β- required for maintaining fruit quality during export, thereby
carotenoid, a precursor to vitamin A, regulated by the CmOr lowering costs (Lamberty and Kreyenschmidt, 2022) [47].
gene in its synthesis. Carotenoid accumulation is further Ethylene gas, a gaseous phytohormone critical for regulating
managed through the carotenoid sequestration protein fruit shelf-lives, is synthesized from methionine, first
FIBRILLIN1 (CmFBN1). A comparative proteomic analysis converting to S-adenosyl-L-methionine (SAM) by SAM
between a high β-carotene melon variety and its isogenic synthase. SAM serves as the primary methyl donor in
line with a defective CmOr gene revealed impaired plants, involved in methylation processes of lipids, proteins,
chromoplast formation. This analysis also highlighted and nucleic acids. SAM is further converted by amino
differential expression of CmFBN1. CmFBN1 enhances cyclopropane carboxylate (ACC) synthase into 5-
carotenoid accumulation triggered by CmOr by promoting methylthioadenosine, which cycles back to methionine and
the proliferation of plastid globules, facilitating carotenoid then to ACC, the precursor of ethylene. Ethylene production
sequestration within chromoplasts. Thus, both CmOr and is 756atalysed when ACC is oxidized by ACC oxidase
CmFBN1 are critical in ensuring the vibrant orange colour (ACO). ACO, a Primary enzyme in ethylene regulation,
characteristic of certain melon varieties. often exists in multiple homologs with conserved sequences
across plant species. In melons, the gene CmACO1 from the
Sugar content melon genome, highlighted in Melonet-DB, shows
Several genes related to sugar metabolism in melons have predominant expression among collected fruits.
been identified, Such factors include sucrose phosphate Investigations were demonstrated such inhibiting an ACO
synthase (CmSPS1), acidic invertase (CmAI), and sucrose gene via genetic modifications like antisense as well as
transporters (CmTST1-3). These genes play critical roles in RNA interference prolongs the keeping quality of
regulating sugar content, which is a significant trait climacteric melons through 5-14 days (Ayub et al., 1996) [3-
affecting fruit quality and sweetness. Quantitative trait loci 4]
. Additionally, mutations induced by ethyl
analysis conducted Together with varieties ‘PS’ and ‘SC’ methanesulfonate therapy and TILLING collection, such as
identified Three significant loci associated with sugar a G194D mutation, have also proven effective in prolonging
content on chromosomes 4, 5, and 7 (Argyris et al. 2017) [2]. melon shelf-life (Dahmani Mardas et al., 2010). These
These loci provide valuable insights into the genetic basis of findings underscore CmACO1 as a pivotal gene influencing
sugar accumulation in melons, contributing to efforts in melon fruit shelf-life.
breeding for enhanced fruit sweetness and quality.
Enhancements in genetic editing for agricultural
Aroma breeding
Eighty-two VOCs (Volatile Organic Compounds) The In recent studies implementing a CRISPR/Cas9 structure
composition of both melon rind and flesh was 756atalyse inside melons, between 40% and 100% of transgenic lines
utilizing gas chromatography-mass spectrometry (GC-MS). exhibited mutations in the target gene, indicating a high
An RIL group derived from the hybridization between ‘PS’ success rate. Typically, a CRISPR/Cas9 structure is
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International Journal of Advanced Biochemistry Research https://www.biochemjournal.com
introduced within the confines of crop genomes using sugar accumulation in melon (Cucumis melo L.).
Agrobacterium-mediated gene transfer technology. Frontiers in Plant Science. 2017;8:1679.
However, transformation protocols for melons often result 3. Ayub R, Guis M, Amor MB, Gillot L, Roustan JP,
in low frequencies and can induce tetraploidy in tissue Latché A, et al. Expression of ACC oxidase antisense
culture. To address these challenges, alternative approaches gene inhibits ripening of cantaloupe melon fruits.
such as the iPB method have been proposed, which allows Nature Biotechnology. 1996;14(7):862-866.
gene editing Void of certain be necessary with regard to 4. Ayub R, Guis M, Amor MB, Gillot L, Roustan JP,
plant tissue culture by employing systems like selection and Latche A, et al. Expression of ACC oxidase antisense
re-differentiation. These methods aim to improve efficiency gene inhibits ripening of cantaloupe melon fruits.
and avoid the complications associated with traditional Nature Biotechnology. 1996;14(7):862-866.
tissue culture techniques in melon transformation. 5. Bailey LH, Bailey EZ. Hortus Third. New York:
In muskmelon breeding, A large number of desirable Macmillan Pub. Co.; 1976.
characteristics like disease resilience, fruit characteristics, as 6. Balass M, Cohen Y, Bar-Joseph M. Identification of a
well as sensual expression to be frequently influenced by constitutive 45 kD soluble protein associated with
single nucleotide polymorphisms (SNPs) preferably than resistance to downy mildew in muskmelon (Cucumis
major hereditary conditions. Thus, genome editing melo L.) line PI 124111F. Physiological Molecular
technologies this may induce specific SNPs are considered Plant Pathology. 1992;41:387-396.
many experiments than easy gene disruption methods like 7. Bardin, Carlier, Nicot. Genetic differentiation in the
CRISPR/Cas9. Base editors and target-AID enzymes have French population of Erysiphe cichoracearum, a causal
been developed to enable precise single nucleotide agent of powdery mildew of cucurbits. Plant Pathology.
substitutions (e.g., C to T, G to A, A to G, T to C) in crops 1999;48(4):531-540.
like Oryza sativa and Solanum lycopersicum (Bastet et al., 8. Bastet A, Zafirov D, Giovinazzo N, Guyon-Debast A,
2019; Hunziker et al., 2020; Shimatani et al., 2017) [8, 36, 76]. Nogué F, Robaglia C, et al. Mimicking natural
A targeted AID technique has also been successfully applied polymorphism in eIF 4E by CRISPR-Cas9 base editing
during muskmelon, facilitating nucleotide and protein is associated with resistance to potyviruses. Plant
changes in genes such as CmelF4E for resistance against Biotechnology Journal. 2019;17(9):1736-1750.
MNSV (Shirazi Parsa et al., 2023) [77]. Additionally, 9. Baudracco-Arnas S, Pitrat M. A genetic map of melon
primery editing has emerged as a technology that allows (Cucumis melo L.) with RFLP, RAPD, isozyme, disease
targeted DNA nucleotide replacement through linking resistance and morphological markers. Theoretical
reverse transcriptase to a Cas9 variant (nCas9), Applied Genetics. 1996;93:57-64.
demonstrating potential applications in crops like Oryza 10. Bezirganoglu I, Hwang S, Fang TJ, Shaw J. Transgenic
sativa as well as Triticum aestivum. lines of melon (Cucumis melo L. var. makuwa cv.
Currently, one limitation of CRISPR/Cas9 and related 'Silver Light') expressing antifungal protein and
technologies is their dependence on a specific PAM chitinase genes exhibit enhanced resistance to fungal
sequence (NGG) near the target site. However, ongoing pathogens. Plant Cell Tissue Organ Culture.
research is advancing the development of Cas proteins 2014;112:227-237.
capable of recognizing diverse PAM sequences, which 11. Bordas M, Montesinos C, Dabauza M, Salvador A,
would greatly expand the spectrum of sequences that can be Roig LA, Serrano R, et al. Transfer of the yeast salt
targeted in genome editing applications. This progress tolerance gene HAL1 to Cucumis melo L. cultivars and
promises to significantly enhance the versatility and in vitro evaluation of salt tolerance. Transgenic
precision of genome modification tools for enhancing crops
Research. 1997;6:41-50.
efforts. Genome editing technology is progressing quickly
12. Brotman Y, Silberstein L, Kovalski I, Perin C,
and becoming increasingly user-friendly for breeding novel
varieties. Technically, it holds immense potential to Dogimont C, Pitrat M, Klingler J, et al. Resistance gene
expedite variety development. However, several obstacles homologs in melon are linked to genetic loci conferring
must be addressed for widespread commercial adoption, disease and pest resistance. Theoretical and Applied
including patent-related challenges. To facilitate its use in Genetics. 2002;104:1055-1063.
breeding and commercializing new varieties, efforts to 13. Burger Y, Katzire N, Tzuri G, Portnoy V, Saar U,
minimize patent fees and streamline technology transfer, Shriber S, et al. Variation in the response of melon
packaging, and consulting services are crucial. Moreover, genotypes to Fusarium oxysporum f. sp. Melonis race 1
enhancing public awareness and understanding of genome determined by inoculation test and molecular markers.
editing technology is essential to its broader acceptance and Plant Pathology. 2003;52:204-211.
application. 14. Chen K, Wang Y, Zhang R, Zhang H, Gao C.
CRISPR/Cas genome editing and precision plant
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