Introduction Chapter (Corrected)

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I.

INTRODUCTION

Vegetables are an essential component of a well-balanced diet and serve as a


supplement. India is ranked second to China in terms of full-scale vegetable
production (Anonymous, 2023). However, present production is significantly lower
than required to meet the needs of the growing population. India's annual vegetable
consumption is expected to reach 220 million metric tons by 2025 (Anonymous,
2024). Vegetables have an important role in nutrition because they provide vital
carbohydrates, proteins, nutrients, minerals, lipids,basic salts and dietary fibre. As the
majority of the Indian population is vegetarian, the importance of vegetable products
becomes increasingly relevant.

Tomato (Solanum lycopersicon L.) (2n = 24), is a member of the Solanaceae


family and one of the most widely consumed vegetables worldwide (Linnaeus, 1753).
It is also an essential crop in horticulture and holds a prominent place in the
worldwide vegetable economy. It can be grown in a variety of climatic conditions
from tropical to temperate and is also grown under protected conditions. Certainly, it
is the most popular garden crop, ranks first among processing crops and is the second
most consumed vegetable after potato. It is among the most important ‘defensive
nourishment’ due to its unique nutritional value and abundant production.

The term "tomato" in English originated from the Spanish word "tomate,"
which is derived from the "tomatl" word in the Nahuatl language of the Aztecs. It is
believed to have originated in western South America and had been cultivated in
Central American Peru-Equador region (Jenkins, 1948). The Portuguese introduced it
to India in the early 16th century, and commercial cultivation began in the 19 th
century with the advent of the British. It is now one of the most popular and
commonly cultivated vegetable crops in India and around the world. Tomato is used
in salads, ketchup, sauces, soups, pickles, purees and many other dishes. Ripe fruits
are rich in ascorbic acid, vitamins A and B, organic acids, and minerals such as
potassium and sodium.

India’s tomato production in 2023-24 is 213.20 lakh tonnes. In 2021-22, the


area of tomato cultivation was around 0.84 lakh hectares, with production of 20.33
lakh tones and productivity of 24.2 kg/hectare. Madhya Pradesh contributes 14.6%,
followed by Andhra Pradesh with 11% and Karnataka with 10.2% to total tomato
production in India (Anonymous,2021). The crop is often grown in plains during the
winter (October–April) and in hilly areas of the country during the summer (April–
October). From the time of emergence until harvest tomato plants face an enormous
number of biotic stresses, such as insect pests and diseases. Further, abiotic stresses
that tomato is extremely sensitive include high temperatures, salinity, dryness,
excessive moisture and natural contamination. There are around 200 known tomato
diseases worldwide that degrade tomato quality and lower crop yields (Atherton and
Rudich, 1986).

The tomato crop is susceptible to a number of fungal, viral, nematode, and


bacterial infections. The crop is attacked by a series of catastrophic diseases such as
tomato mosaic virus (ToMV), tomato spotted wilt virus (TSWV), tomato leaf curl
virus (TLCV), tomato bunchy top virus (TBTV), buckeye rot (Phytophthora
nicotianae var. parasitica), septoria leaf spot (Septoria lycopersici), Fusarium wilt
(Fusarium oxysporum f.sp. lycopersici), anthracnose (Colletotrichum phomoides),
powdery mildew (Leveillula taurica), bacterial wilt (Ralstonia solanacearum),
bacterial canker (Clavibacter michiganensis pv. michiganensis), bacterial leaf spot
(Xanthomonas campestris pv. vesicatoria). As the crop is attacked by a series of such
diseases one after another, the issue gets worse (Thakur and Tripathi, 2015).

Among the fungal diseases, Alternaria solani (Ellis and Martin)-induced early
blight, known as target spot disease, is one of the most devastating in the world,
causing loss in tomato-growing regions of India. The pathogen Alternaria solani
belongs to the class Deuteromycetes, order Moniliales, family Dematiaceae, genus
Alternaria. This airborne, soil-dwelling causative organism causes tomato fruit rot,
collar rot, and early blight (Datar and Mayee, 1981). Early blight in tomato caused by
the fungus Alternaria solani, is a significant disease affecting tomato crop worldwide.
It is the most serious disease, according to Mathur and Shekhawat (1986), causing up
to 50 to 80 per cent losses in tomato yields. According to Saha and Das (2012), every
1% increase in disease severity resulted in a yield loss of 0.75 to 0.77 tons/ha.

The genetics of resistance to early blight is complex, involving multiple genes


and quantitative trait loci (QTLs), with resistance often inherited in a polygenic
manner. This means that multiple genes contribute to the overall resistance of the
plant. The variability in resistance to early blight among tomato varieties is substantial
and provides a crucial genetic resource for breeding programs aimed at developing
resistant varieties. Wild relatives of tomatoes, such as Solanum pimpinellifolium and
Solanum habrochaites, have been identified as valuable sources of resistance genes
(Foolad et al., 2002). Recent advancements in genomics and molecular biology have
deepened the understanding of the genetic mechanisms underlying early blight
resistance. Studies have identified several effective QTLs associated with resistance,
such as EB-1.2, EB-5, and EB-9, in breeding lines (Anderson et al., 2021). Significant
QTLs on chromosomes 5, 6, and 9, contributing 10–35% to early blight resistance,
have also been identified in interspecific crosses involving wild relatives (Ashrafi &
Foolad, 2015). The use of genome-wide association studies (GWAS) has further
facilitated the identification of candidate genes involved in the resistance response,
showcasing successful genotypic selection in breeding programs (Chaerani et al.,
2007). These advancements emphasize the potential of integrating molecular tools
into conventional breeding strategies for improving early blight resistance.

Small RNAs, including microRNAs (miRNAs) and small interfering RNAs


(siRNAs), play a crucial role in regulating gene expression in response to pathogen
attack. These small RNAs can modulate the expression of resistance genes and other
defense-related genes, contributing to the plant’s overall resistance to early blight
(Chaerani & Voorrips, 2006). Genetic maps and molecular markers are essential tools
in the study of early blight resistance. These tools allow researchers to identify and
map QTLs associated with resistance traits. Marker-assisted selection (MAS) uses
these markers to select plants with desirable traits along with those conferring
resistance to early blight, significantly speeding up the breeding process. MAS has
revolutionized tomato breeding by enabling the precise selection of plants with
resistance to early blight (Foolad et al., 2008). By using molecular markers linked to
resistance genes, breeders can efficiently incorporate these genes into new tomato
varieties. This approach reduces the time and resources that are required to develop
resistant varieties compared to traditional breeding methods.

Early blight is extremely difficult to control once it has spread throughout the
crop (Smith and Kotcon, 2002). Although many researchers advise a combination of
conventional and chemical measures, primarily the use of fungicides and plant
extracts, to prevent the development of the disease and prevent significant yield losses
as a result of it, no practical, cost-effective method of managing the disease has yet
been developed. The primary focus of the present study is the identification of
resistance sources against Alternaria solani involving a comprehensive screening of
the tomato mini-core collection, aiming not only to identify resistant accessions but
also to map genetic determinants through GWAS. Field screening of the Tomato
Association Mapping Panel (TAMP), which represents the mini-core of tomato, is
anticipated to capture the plant factors that restrict Alternaria solani's invasion of
tomato plants, thereby enabling the mapping of gene loci across the tomato genome.
A thorough analysis of TAMP accessions for various morphological traits directly or
indirectly contributing to resistance against Alternaria solani and the utilization of
DNA markers data in GWAS will lead to deeper insights into both the fundamental
and practical aspects of resistance to this pathogen in terms of mapping genes
conferring resistance.

GWAS is a powerful method for identifying alleles associated with specific


traits in crop species by detecting the relationship between genotype and phenotype in
unrelated natural populations with LD patterns, without the need for segregated
populations (Sauvage et al., 2014). This approach offers high mapping resolution, as
demonstrated in a study, identifying SNP markers linked to Fusarium wilt resistance
in the natural germplasm of tomatoes (Kawicha et al., 2014). The genome-wide
identification of NLR (nucleotide-binding leucine-rich repeat) revealed highest
concentration of NLR genes on chromosomes 4, 5, and 11 co-localized with major
QTLs for late blight resistance in tomatoes (Bashir et al., 2022). These NLR genes are
essential in initiatig defense responses during pathogen invasion, underlining their
significance in disease tolerance and as positive regulators of resistance (Khahani et
al., 2021; Danan et al., 2011). Fine mapping of identified QTLs can further pinpoint
promising regions for disease resistance genes (Kumar and Nadarajah, 2020)
providing valuable resources for developing improved cultivars through marker-
assisted selection and genomic selection in tomato breeding programs.

The integration of genetic resources, advanced genomic tools, and molecular


markers has significantly enhanced our ability to breed tomatoes resistant to early
blight. The involvement of small RNAs in regulating resistance genes adds another
layer of complexity and potential for improving resistance. Marker-assisted selection
remains a powerful tool in the fight against early blight, ensuring the development of
robust and resilient tomato varieties. Therefore, taking into consideration the
importance of the crop and the adverse effects caused by this pathogen, the current
studies entitled ‘Genome-Wide Association Studies for Early Blight (Alternaria
solani) Disease Resistance in Tomato (Solanum lycopersicum L.)’ was carried out
with the following objectives:

1. To study the genetic variability for early blight disease resistance in tomato
association mapping panel (TAMP).

2. To determine the marker-traits association for traits governing early blight


resistance through association mapping in tomato association mapping panel.

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