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Hari Har Ram
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Hari Har Ram
Hari Har Ram: Former Professor Department of Plant Breeding and GeneticsG.B. Pant University of Agriculture and Technology Pantnagar, Udham Singh Nagar-263 153 Uttarakhand, India

Former Head Department of Vegetable ScienceG.B. Pant University of Agriculture and Technology Pantnagar, Udham Singh Nagar-263 153 Uttarakhand, India

Former Vice President Krishidhan Vegetable Seeds, PuneFormer FAO Consultant (Myanmar, Libya, DPR Korea)

The present book entitled “Commercial Plant Breeding-1 Vegetable Crops” contains 32  chapters  bifurcated  into  two  parts,  part  I  “Commercial  Plant  Breeding  and Regulatory Affairs” having initial 14 chapters deals with commercial plant breeding, the  vegetable  seed  business,  vegetable  seed  supply  chain  management,  all  India coordinated  research  project  (vegetable  crops),  world  vegetable  centre  (AVRDC), and seed and IPR related acts. The part II “Commercial Breeding of Vegetable Crops” having remaining 18 chapters is devoted to commercial breeding of individual major vegetable crops relevant to seed industry.

The  book  is  intended  for  undergraduate  and  post  graduate  students  of  state agricultural  universities  and  agricultural  colleges  for  the  courses  on  vegetable breeding  in  general  and  commercial  vegetable  breeding  in  particular  and  the professionals across ICAR Institutes, SAUs and Seed Industry. First of its kind, the present book aims to bring academic and vegetable seed business worlds both on a common platform

Author Speak

0 Start Pages

Preface The present book entitled “Commercial Plant Breeding-I-Vegetable Crops” was conceived after going through the syllabus of B. Sc. Agriculture (Hon.) as per recommendation of Indian Council of Agricultural Research - V Dean Committee for Colleges of Agriculture throughout India and also having discussion with Mr. Sumit Jain, New India Publishing Agency, New Delhi. As per this syllabus, Commercial Plant Breeding is listed under elective courses and the course content includes plant breeding approaches and seed production procedures with emphasis on hybrid seed production and related activities as practiced by seed companies across major field and vegetable crops. Although, the proposed course ‘Commercial Plant Breeding’ includes both field and vegetable crops in the same course, it was considered more prudent to prepare two books, one dealing with vegetable crops and the other dealing with field crops, separately as nationally and internationally these two groups of the crops are dealt with separately at all levels. Even, private seed companies and ICAR institutes and state agricultural universities deal with these two groups of crops separately for teaching/research purposes. Therefore, it was agreed with the publisher that there will be two books; the first dealing with vegetable crops and the second dealing with field crops and that is how Commercial Plant Breeding-I-Vegetable Crops has been prepared. Initial few chapters are devoted to introducing the students with vegetable seed business, and seed companies’ activities which play key roles in taking forward the R&D products, namely, open-pollinated seeds and hybrid seeds to the end users i.e., the farmers across the country as per suitability of the product in a particular geographic region/location. The related activities crucial to core commercial plant breeding and product delivery are IPR regulations and seed related acts, multi-location testing, commercial launch of the product, parental seed purity, multiplication of parental seed/foundation seed, commercial seed production, direct import of commercial seed, product development, grow-out tests, assurance of seed quality, and managing entire supply chain so that the seed is delivered to the farmers nearest to his location. After these chapters, the book coverage shifts to the breeding of individual major vegetable crops which make business sense to the private seed companies. In these crop-wise chapters on breeding of individual vegetable crops, the focus is on product design and requirements, commercial breeding goals, commercial plant breeding approaches (line breeding, product development), emasculation and pollination, and hybrid seed production techniques with emphasis on how these are handled at production plots on commercial scale, and of course with needful exposition of theoretical concepts behind commercial breeding activities so that a commercial plant breeder appreciates the logic behind what he is doing in the field/lab. I have included an adequate literature review covering landmark publications from ancient times to the present.

1 Commercial Plant Breeding

Defining-Commercial Plant Breeding The term commercial plant breeding originates by putting prefix ‘commercial’ before ‘plant breeding’ and is used as an adjective to plant breeding which surely is a noun. Naturally, commercial plant breeding remains largely plant breeding with slight twist that commercial plant breeding aims at delivering commercial products to be delivered to the end users that is the farmers as per the need of the consumers/markets. Thus, the farmers plant those products (open-pollinated cultivars and hybrids) which are easily accepted by the consumers and which are easy to be handled by the traders as the products reach to the traders and then from traders to the retailers and ultimately to the consumers. Along with consumers’ acceptance, these kinds of products must meet certain level of yield, maturity and resistance/tolerance to diseases and pests. Accordingly, these are designed and bred by the commercial seed companies which operate as per demand of the markets. Private seed companies operate on a business model and generate their own revenue to keep the companies running and therefore these seed companies are bound to design and breed commercially viable products. Thus, all plant breeding activities which occur at the research farm and laboratory of private seed companies aimed at designing and delivering commercially viable products are called as commercial plant breeding activities. While designing such project and running the plant breeding activities at the research farm and laboratories of the private seed companies, the emphasis has to be on ‘how to do’ and ‘how quickly to do’ rather than ‘why to do’. ‘Why to do’ issue is not totally ignored and this gets due consideration while formulating the research project based on extensive deliberations between various departments, namely, R&D, Sales and Marketing, Product Development, and Seed Production. The term commercial plant breeding is generally not in use in public sector research institutes and the agricultural universities and certainly not in class room teachings and discussions while teaching courses on plant breeding and related courses under the domain of plant breeding. There the focus is on theory in contrast to applications which form the backbone of commercial plant breeding as practiced by the commercial organizations such as private sector seed companies which have to earn revenue by selling the products to meet the operational expenditure of running the companies and also to earn profit to run the activities of the company on long term basis in a sustainable manner and to invest in R&D. Therefore, a working definition of commercial plant breeding could be as follows.

1 - 64 (64 Pages)
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2 Vegetable Seed Business in India

The general recommendation for intake of fruits and vegetables is at least 400 g per person as recommended by World Health Organization (WHO), five servings of 80 g per day or about 146 kg per person per annum. For vegetables, the recommended intake requirement is at least 300 g per day per individual. This 300 g requirement must include 50 g leafy vegetables, 50 g roots and tubers and 200 g other vegetables (Kapur, 2016). Current annual vegetable production including potato, based on recent three years (2015-16, 2016-17, 2017-18) average is around 177 million tons from 10 million ha giving a productivity of 17.7 ton/ha. With this average and stagnating production and assuming Indian population at 133 CR (1.33 billion) and 25% postharvest losses, per capita per day availability of vegetables comes to 273 g leaving a gap of 27 g between the recommended dose and the availability of vegetables per head per day in India. By 2050, the projected population of India is 1.5 billion and to meet the requirement of this population, vegetable production will have be 220 million tons, of course making allowance of 25% post-harvest losses. Considering all kind of constraints on the available land area of vegetables, the vegetable area will be around 10 million ha and consequently, major gain of production will have to come from productivity increase from current level of about 18 tons/ha to 22 tons/ha and this seems to be a gigantic task as during last several years, the productivity of vegetables in India has been between 17 and 18 tons/ha, an indication that it has reached to a plateau. This situation calls for fall massive interventions on technological fronts in terms of innovative products with focus on better hybrids with higher yields, and resistance to biotic and abiotic stresses deploying precision breeding technologies using more of marker assisted breeding, genomics and gene editing tools and delivering high quality seeds to the farmers with assurance of high performance. This chapter looks into the present seed scenario and the emerging trends with respect to new seeds and new technologies and examines the need to facilitate rules, regulations and IPR environment for more investment in vegetable research and development and quality seed production and distribution. Improved seed with shift towards hybrid seed will be the key driver to increase vegetable production and productivity in India.

65 - 77 (13 Pages)
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3 Multi-Location Testing and Product Development

Genotype × environment interaction underlies the efforts of crop ecologists, agronomists, and plant breeders to define ecological regions, mega-environments, and ecotypes and to specify the adaptation and yield stability of individual cultivars. Multi-location testing or multiple year testing gives rise to an important term, genotype x environment interaction. “Interaction” is a statistical term signifying differential response of genotypes over the environments. The environments could be in terms of more numbers of diverse locations or years. Multi-location trials (MLTs) are also called as multiple environment trials (METs) and these have been adequately reviewed by Yan and Hunt (1998) and Kang et al. (2004). The following sections of this chapter derive extensive information on MLT/MET from these two sources. Introduction The genotype-environment-interaction (GEI) is a complex phenomenon that undermines the repeatability of experimental results and consequently reduces the selection efficiency. Agricultural researchers have long been concerned of the various implications of GEI in plant breeding programmes. Understanding of the implications of GEI structure/nature is important in crop breeding programmes as a significant GEI can seriously impair selection of superior genotypes in new crop introduction and cultivar/hybrid development programmes. Crop breeders of field and vegetable crops, etc. are in better position to decide whether crop cultivars should be developed for all the environments of interest or if the breeders should develop specific cultivars for specific target/mega environments. Variation among genotypes in phenotypic sensitivity to the environment may require the development of locally adapted varieties. Genotype-by-environment is widely observed and reported issue. Large numbers of publications both on GEI and stability parameters are pointers to the facts that it is a complex and intriguing issue and these are expected to be more relevant in the light of the fact that greater emphasis is going to be placed on sustainability of agricultural systems.

78 - 86 (9 Pages)
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4 Grow Out Test

Genetic purity (GP) is a crucial seed quality factor. Several steps and precautions are involved in achieving the desired levels of genetic purity in the seed. Right from the choice of genetically pure seed for seed production up to final packing of the seed produced, at any stage, genetic purity can get affected. Field inspection and postharvest checks help in verifying conformity to seed quality standards especially genetic purity. Wherever seed characteristics are clear, examination of samples in the quality assurance laboratory for other distinguishable varieties (ODV) gives an idea of the genetic purity status. Among the several methods available for GP assessment, Grow Out Test is one. In this context, some basic principles and procedures and guidelines relating to GOT are presented with more of information in tabular forms. In brief, the aspects covered include objectives, principles, kinds of GOT, genetic purity standards, procedure, main factors for observations, evaluation, and recording and reporting. Table 4.1 gives the submitted sample size for GOT; and Table 4.2 shows the minimum number of days required for evaluation from sowing and Table 4.3 deals with maximum weight of submitted sample and minimum weight of working sample for GPM, GOT and ODV.

87 - 97 (11 Pages)
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5 Seed Supply Chain Management

Supply chain management or inventory management is an important and integral component of commercial breeding activities practiced by the private seed companies whose over all research efforts in terms of developing and commercially releasing high performing cultivars and hybrids get defeated unless the product reaches to the end users that is the farmers. Seed supply chain management is the lifeline of any seed business especially when the seed business is under private sector seed companies whose delivery is very strong. Supply chain management refers to the process of ordering, storing, and using a company’s inventory. These include the management of raw materials, components, and finished products, as well as warehousing and processing such items and ultimately delivering virtually to the door steps of the farmers through a strong network of dealers/distributors/retailers. For companies with complex supply chains and manufacturing processes, balancing the risks of inventory gluts and shortages is especially difficult and challenging. The process of supply chain management is a continuously evolving process, ready to deal with the unforeseen situations like the situation created by COVID-19 pandemic forcing countries after countries going for lockdown and bringing the entire business operations including material and men transport to a grinding halt. However, even under these extremely challenging situations, governments fully know that agricultural activities cannot be fully stopped to avoid resurgence of hunger and malnutrition. However, despite the window allowed by the government for running the supply of essential goods including agricultural inputs like seed, agrochemicals and farm machines, manufacturing the agro-inputs including seeds and their distribution to the endpoints has become extremely difficult. A few companies have devised new processes to run seed supply using completely new and innovative approaches, for example, In Bayer’s first-ever e-commerce collaboration in India, the company has tied up with Agro-Star, a Pune-based e-commerce firm in the agricultural segment. Under the partnership, farmers can order Bayer’s seeds and crop protection products for their entire crop lifecycle and receive agronomic advisory through Agro-Stars digital agri-tech platform. Home delivery of agri-inputs can currently be availed by farmers in North, West and Central India, with plans to extend the scope to other geographies in the future. In the current COVID-19 lockdown situation, the partnership has enabled last mile delivery of products for multiple crops directly to farmer’s doorsteps.

98 - 112 (15 Pages)
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6 Seed Import and Export

The International movement of seed takes place for (i) germplasm for R&D purposes, (ii) experimental lines and hybrids for screening purposes, (iii) basic seed for multiplication purposes, and (iv) commercial varieties and hybrids for marketing (Ranganathan, 2019). Vegetable seed import and export for research purposes is handled by National Bureau of Plant Genetic Resources (NBPGR), New Delhi where the importing research organization in India first gets an import permit from NBPGR following due procedures as available on NBPGR website ( Once the importing research organization, like ICAR institutes, state agricultural universities, private seed companies, etc get the import permit which specifies the list of material to be imported and the seed source, the import permit is sent to the research seed supplying agency who in turn attaches a phytosanitary certificate along with the copy of the import permit to the consignment and sends the same to NBPGR. NBPGR retains part of seed and releases the seed to the indenter after necessary quarantine test and charging quarantine fee. The importing organization is expected to give a brief report about the material to the NBPGR after its evaluation at its farm/lab. The bulk seed import (vegetable seed for commercial purposes) is handled by Directorate of Plant Protection, Quarantine and Storage, Faridabad, Haryana ( with no condition of testing by government system. This exemption of testing is not available for field crop seed. It needs to be clarified that the bulk vegetable seed import has been simplified as per New Seed Policy-1988 which has put the vegetable seeds among others under OGL (open general license) and this provision facilitates the vegetable seed in bulk and the private seed companies have taken advantage of this provision. The Indian private seed companies are importing large quantities of seed of cauliflower, cabbage, broccoli, radish, carrot, beet root and even vegetable peas. The bulk vegetable seed import for commercial purposes is done as per following procedure.

113 - 135 (23 Pages)
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7 Release and notification of Vegetable Varieties in india

Varietal release and notification comes under the purview of the Seeds Act which is intended to provide for regulating the quality of certain notified seeds for sale and for marketing and for the matters connected therewith. Released and notified varieties are offered for seed certification by the seed certification agencies which ensure that the varieties under certification meet certain prescribed field and seed standards failing which they stand to be rejected during the process of seed certification. This ensures seed quality and its availability to the farmers. Purpose of Release The purpose of Release of cultivars is to introduce the newly evolved varieties to the public and the farmers in particular for general cultivation in the region for which the variety is released and notified ensuring availability of seed of varieties according to their suitability and adaptation and avoiding mismatch. In other words, release of a cultivar is in the nature of a recommendation to the farmers for its adoption. Therefore, notification of a variety is linked to the release of a variety, though the release process itself does not have statutory cover.

136 - 151 (16 Pages)
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8 The Seed Act 1966

The Seed Act 1966 was passed by the Indian Parliament on 29th December 1966 and it came into force from 2nd October, 1969. In addition, Seed Regulation Order was also passed in December 1983. The sole objective of Seed Act and Order has been to ensure that the Indian farmers get good quality seeds of important crops. Important provisions and guidelines for the farmers are given below: The Act is applicable for the seeds notified by the Central Government from time to time. Seed quality is regulated at two stages viz. seed production and seed certification. For this purpose, seed testing laboratories and seed certification agencies are to be established in each state. Maharashtra has been the first state to implement this act. Now all the states have implemented this act.

152 - 163 (12 Pages)
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9 New Seed Policy 1988

New Seed Policy Ministry of Agriculture, vide Letter No.11-71/88-SD-1 dated September 16,1988, has brought out ‘New Policy on Seed Development’, with the aim of providing to the farmers the best planting materials available in the world so as to increase productivity and thereby increasing farm income and export earnings. The Policy covers the import of seeds of coarse cereals/pulses/oil seeds; vegetable and flower seeds; bulbs/tubers of flowers; cuttings/ saplings/bud/wood etc., of flowers; and seeds & planting material of fruits. The following are the guidelines for the import of above categories of seed/planting material under the Seed Policy Import of Seeds of Coarse Cereals/Pulses/Oil Seeds The import of seeds of coarse cereals/pulses/oil seeds is permitted for a period not exceeding two years by companies that have technical/financial collaboration with companies abroad, provided that the latter agree to supply parental lines/nucleus or breeder seed technology to the Indian firm within two years of first shipment of commercial consignment. The bulk import is permitted by the importer based on the recommendation issued by the Department of Agriculture & Cooperation (DAC) after considering the trial/evaluation report of the varieties/hybrids, which he intends to import. PPA or any other officer notified for the said purpose issues the import permit based on the recommendation of DAC. However the recommendation of DAC is not required for import of trial material. The quantities of the seed permitted for import for initial trials by the importer, for ICAR multi-location trials & agronomic trials or combined trials (vide Letter No. 11-71/88/S.D.I dated 30th January 1989 issued by the Ministry of Agriculture) as per Para 4.3.5 of Seed Policy shall be in accordance with Schedule-XII and also the quantities of seeds of parental lines required to be sampled and submitted in sealed condition for accession to gene bank of National Bureau of Plant Genetic Resources (NBPGR), New Delhi (vide Letter No. 11-102/88-S.D.I dated 5th February 1991) as per Para 4.3.8 of Seed Policy. The imported consignment shall be subjected to detailed testing for a period of 30-35 days and the same purpose the consignment will be held under detention in AAI warehouse or any customs bonded warehouse at the cost of importer.

164 - 170 (7 Pages)
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10 Protection of Plant Varieties and farmers' Rights Act, 2001

Protection of Plant Varieties and Farmers’ Rights Act-2001 (PPVFRA-2001) was enacted by government of India in 2001 to create a sui generis system of providing a legal framework as an IPR mechanism to protect the varieties of crop plants in India along with grant of breeders’ and farmers’ rights in the interest of all the stakeholders and also to comply with WTO guidelines to which India is a legal signatory. The salient features of this act are as follows. The Protection of Plant Varieties and Farmers’ Rights Act-2001, Act no. 53 of 2001, (30th october, 2001) This act is in place to provide for the establishment of an effective system for protection of plant varieties, the rights of farmers and plant breeders and to encourage the development of new varieties of plants. It recognizes and protects the rights of the farmers in respect of their contribution made at any time in conserving; improving and making available plant genetic resources for the development of new plant varieties; it encourages accelerated agricultural development in the country; it protects plant breeders’ rights to stimulate investment for research and development, both in the public and private sector, for the development of new plant varieties; it facilitates the growth of the seed industry in the country which will ensure the availability of high quality seeds and planting material to the farmers. To give effect to the aforesaid objectives, it is necessary to undertake measures for the protection of the rights of farmers and plant breeders; as India, has ratified the Agreement on Trade Related Aspects of Intellectual Property Rights should, inter alia, make provision for giving effect to sub-paragraph (b) of paragraph 3 of article 27 in Part II of the said Agreement relating to protection of plant varieties. Accordingly, this act is in place.

171 - 203 (33 Pages)
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11 Biological diversity Act 2002

The increasing concerns about dwindling biological resources globally led to the Convention on Biological Diversity in 1992. This convention for the first time, recognized sovereign rights of States over their biological resources and emphasized that access to genetic resources should be only for environmentally sound purposes and should be subject to national legislations. The access has to be on mutually agreed terms which inter-alia would include recognition of associated Traditional Knowledge of indigenous communities and equitable benefit sharing arrangements. Taking cognizance of the provisions of the CBD, and to deal with extensive pressure on our biological resources, India has enacted an umbrella legislation called Biological Diversity Act-2002 (No. 18 of 2003), and also notified the Biological Diversity Rules-2004 for guidance and compliance by various stakeholders, including the Union and State Governments, Non-state sectors, and individuals. The Act aims at conservation of biological resources as well as facilitating access to them in a sustainable manner and through a just process. The salient features of this act are as follows (NBA, 2004).

204 - 222 (19 Pages)
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12 Dus Testing and Registration of Plant Varieties

In order to provide for the establishment of an effective system for protection of plant varieties, the rights of farmers and plant breeders and to encourage the development of new varieties of plants it has been considered necessary to recognize and protect the rights of the farmers in respect of their contribution made at any time in conserving, improving and making available plant genetic resources for the development of the new plant varieties. Moreover to accelerate agricultural development, it is necessary to protect plants breeders’ rights to stimulate investment for research and development for the development of new plant varieties. Such protection is likely to facilitate the growth of the seed industry which will ensure the availability of high quality seeds and planting material to the farmers. India having ratified the Agreement on Trade Related Aspects of the Intellectual Property Rights has to make provision for giving effect to Agreement. To give effect to the aforesaid objectives the Protection of Plant Varieties and Farmers’ Rights Act, 2001 has been enacted in India

223 - 244 (22 Pages)
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13 All india Coordinated Research Project (Vegetable Crops)

All India Coordinated Research Project (AICRP) on vegetable crops (VC) was started during the IV five-year plan in 1970-71, to provide a national grid for multi-location testing of the vegetable technologies developed by various research institutes and state agricultural universities. The headquarter of the project was at the Division of Vegetable Crops, Indian Agricultural Research Institute (IARI), New Delhi and the first Project Coordinator (Vegetable Crops) joined the project in July 1971. During 1986, level of the project was elevated to the Project Directorate. Further, in a significant development during 1992, headquarter of the directorate was shifted from IARI, New Delhi to the present day Indian Institute of Vegetable Research (IIVR) at Varanasi. In a significant development, in eleventh five-year plan the National Seed Project (Vegetables) was merged with AICRP (VC). Presently the AICRP Vegetable Crops is a network of 36 regular and 18 voluntary centres. The network is headed by the Project Coordinator and it’s headquarter functions from IIVR (Indian Institute of Vegetable Research), Varanasi, main campus. The AICRP centers are located in different agro-climatic zones of the country. The centers in the network include ICAR institutes, central institutes, state agricultural and traditional universities and a few other public and private research organizations.

245 - 254 (10 Pages)
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14 World Vegetable Centre (AVRdC)

The World Vegetable Center (previously called as Asian Vegetable Research and Development Centre, AVRDC) with headquarter at Shanhua, Taiwan conducts research, builds networks, and carries out training and promotion activities to raise awareness of the role of vegetables for improved health and global poverty alleviation. Vegetables can alleviate poverty by creating new jobs and new sources of income for farmers and landless laborers, improve health by providing essential micronutrients lacking in diets, enhance learning and working capacities of adults and children through improved diets and health, and improve the sustainability of food production practices by diversifying cropping systems. The Center’s research and development work focuses on breeding improved vegetable lines, developing and promoting safe production practices, reducing postharvest losses, and improving the nutritional value of vegetables ( The centre’s mission is Research and Development to realize the potential of vegetables for healthier lives and more resilient livelihoods

255 - 262 (8 Pages)
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15 Tomato

Tomato (Lycopersicum esculentum Miller, 2n = 2x = 24, also known as Solanum lycopersicum, Family Solanaceae) has been the leading vegetable crop globally. Tomato genome is composed of approximately 950 Mb of DNA, more than 75% of which is heterochromatin and largely devoid of genes. The global production of tomato in 2017 has been 182 million tons from 5 million ha area giving a productivity of 36.4 tons/ha. Top 10 countries in tomato production in descending order are China, India, Turkey, USA, Egypt, Iran, Italy, Spain, Mexico and Brazil. Tomato is planted almost throughout the year in different parts of India in an area of 8 lakh ha (0. 8 million ha) with production of 20.8 million tons giving a productivity of 26 ton/ha. Tomato area in India is almost 10% of total vegetable area. The states accounting for major tomato production in India are Andhra Pradesh, Orissa, Telangana, Madhya Pradesh, Karnataka, Maharashtra, West Bengal, Chhattisgarh, Bihar, Gujarat, and Himachal Pradesh. Out of total area, about 78% is planted with F1 hybrid seed while remaining 22% of the area still goes to open-pollinated varieties. The average productivity of tomato close to 26 tons/ha is still very low compared with the global productivity of 36.4 tons/ha. However, with the introduction of high performing and disease resistant hybrids, the productivity and production are continuously improving. Most of hybrid tomato seed (95 tons) is sold by the private seed companies. There are more than 375 tomato hybrids in the commercial chain in India produced and marketed by different companies. About 50 hybrids out of these 375 hybrids are being grown on more than 80% of the area under tomato in India. Leading companies in tomato hybrid seed segment are Seminis (Bayer), Syngenta, Rasi HyVeg, and Namdhari (NHB, 2015; Kapur, 2016).

263 - 296 (34 Pages)
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16 Chilli

Chilli (Capsicum annuum, 2n = 2x = 24, family Solanaceae) also known as hot-pepper, classified as often-cross-pollinated crop is consumed both as fresh green and dry spice type. It is a rich source of vitamin C and pungency to the consumers and world food industries. The global area and production statistics often run into different figures due to inclusion of some times only fresh green chilies, sometimes red-dry chillies and sometimes inclusion of sweet peppers also in this broad category. The global production of fresh and dry chillies in 2014 has been reported to be 32.32 million tons where China alone accounted for 16.12 million tons followed by India with over 5 million tons of production. Other major countries growing chilli are Mexico, Indonesia, Spain, Turkey, USA, Korea, Bangladesh, Pakistan, Nigeria, Thailand and Ethiopia. More than 65% of global green and dry chillies production is in Asia, while Europe contributes 12% and Africa around 10%. China is the leading producer of green and dry chillies, Area under chillies in China is less than that in India, but productivity is almost double than that in India.

297 - 324 (28 Pages)
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17 Eggplant

Solanum melongena L. (2n = 2x = 24, family Solanaceae, often cross-pollinated) is known as eggplant in the United States and aubergine in France and England. It is one of the few cultivated solanaceous species originating from the Old World. It is known as brinjal in its home country, India, where it was domesticated long ago and where the greatest diversity exists. Widely grown in temperate and tropical Asian countries, eggplant has been also a common crop in the Middle East and around the Mediterranean basin and is now cultivated worldwide. Although an Old World taxon, S. melongena unexpectedly shares strong genetic similarities with New World Solanum species, namely, tomato and potato. These syntenic features have been discovered and will undoubtedly assist in an unlocking of the genes controlling eggplant key horticultural traits (Daunay, 2008). As per FAOSTAT website-2018 quoted by Ministry of Agriculture and Farmers Welfare, Government of India, Horticulture Statistics Division (2018), the world production of eggplant was 51.29 million tons from 1.79 million ha with a productivity of 28.65 tons/ha during 2016. Top ten countries in eggplant production during this period were China, India, Egypt, Turkey, Spain, Iran, Indonesia, Italy, Japan, and Philippines. The bulk of eggplant production came from top ranking China (0.78 million ha area, 32.03 million tons production and 41.06 tons/ha productivity) followed by India (0.66 million ha area, 12.60 million tons production and 19.09 tons/ha productivity) occupying the second rank. Productivity-wise, Spain is top rank holder with productivity close to 65 tons. Leading brinjal producing states in India are West Bengal, Odisha, Gujarat, Bihar, Andhra Pradesh, Madhya Pradesh, Maharashtra, Chhattisgarh, Karnataka, Haryana, Jharkhand and Assam.

325 - 347 (23 Pages)
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18 Okra

Okra or lady’s finger (Abelmoschus esculentus L. Moench, 2n = 2x = 130, family, Malvaceae) is classified as fruit vegetable like tomato, and eggplant and is grown in the tropical and sub-tropical regions of world. Okra growing major countries are India (occupying the first rank), Turkey, Iran, Iraq, Egypt, Saudi Arabia, Cameroon, Mexico, Lebanon, Israel, Jordan, Greece, Yugoslavia, Bangladesh, Afghanistan, Pakistan, Myanmar, Japan, Malaysia, Brazil, Nigeria, Ethiopia, Sudan, Ghana, Benin, Cyprus, and southern USA. Though, virtually it is not grown in Europe and north America, yet lot of people in these countries have started liking this common vegetable because of good amount of vitamin A and folic acid, besides, carbohydrates, phosphorous, magnesium, and potassium. In India, okra is extensively grown throughout the country but the states with substantial acreage under okra are Gujarat, Maharashtra, Andhra Pradesh, Telangana, Uttar Pradesh, Madhya Pradesh, West Bengal, Assam, Rajasthan, Tamil Nadu, Karnataka, Haryana and Punjab, both in rainy and spring-summer season. The global total area and production under okra is reported to be close to 1.15 million ha and 7.9 million tons giving a productivity of 6.8 tons/ha. India with approximately 0.5 million ha (5 lakh ha) area and production and productivity of an average of 6 million tons and 12 tons/ha, respectively accounts for close to 75% of share of world production of okra and hence has been accorded a prominent place in the list of crops under R&D by the private seed companies in India. In India, okra has huge export potential as fresh vegetable to the neighbouring countries in the Middle East, like, Bahrain, Qatar, Kuwait, Saudi Arabia, Muscat, Tehran, and Abu Dhabi and SouthEast Asia, particularly Singapore, Mauritius, Malaysia, Sri Lanka and Bangladesh. Therefore, any company with its own R&D in India treats okra a worthwhile crop to keep it under its portfolio and naturally okra becomes an important crop for commercial breeding. Unless a crop makes business sense on account of substantial acreage, production and seed sale volume, there will be no point in launching a full-fledged commercial breeding programme on that particular crop. Thus, okra qualifies a good candidate for commercial breeding in India.

348 - 373 (26 Pages)
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19 Cabbage

Cabbage (Brassica oleracea, ssp. capitata, 2n = 2x = 18, family Brassicaceae = Cruciferae, cross-pollinated due to sporophytic incompatibility) comes under generic name cole crops. It is a rich source of calcium, and vitamins A and C. It is generally consumed as cooked vegetable, boiled or stewed, raw salad, soup and sauerkraut which is cabbage cut into fine slices that undergo lactic acid fermentation in a brine made of its own juice with salt. Sauerkraut was traditionally made at home in Germany and other European countries, but nowadays it is mainly an industrial product. According to Indian Horticulture Database-2014 (NHB, 2015), global production of cabbage spread over more than 125 countries has been 70.64 million tons from 2.42 million ha area with a productivity of 29.2 tons/ha. Of this total global cabbage production, top ranking China (32.80 million tons from 0.98 million ha) and second ranking India (9.04 million tons from 0.40 million ha) jointly contributed about 59% of the production. Other countries in the descending order of the production were Russian Federation, Japan, South Korea, Ukraine, Indonesia, Poland, Romania and USA. In terms of productivity (tons/ha), South Korea (71.2 t/ha) and Japan (67.6 t/ha) were far ahead than the rest of the countries which were in the range of 20.1 to 42.9 tons/ha. During 2017-18, also Indian cabbage production was 9.04 million tons from 0.40 million ha (4 lakh ha) area giving a productivity of 22.6 t/ha. Thus in India, cabbage area seems to be stabilized at 4 lakh ha. The top ten cabbage producing states in India are West Bengal, Bihar, Odisha, Assam, MP, Gujarat, Chhattisgarh, Jharkhand, Karnataka and UP (Horticulture Statistics at a Glance-2018, MAFW, 2018).

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20 Cauliflower

Cauliflower (Brassica oleracea var. botrytis, 2n = 2x = 18, Family Brassicaceae, cross-pollinated, sporophytic self-incompatibility) is traditional European crop that has become widespread in Asia. As per Horticulture Statistics at a Glance-2018, published by Ministry of Agriculture and Farmers Welfare, Government of India (MAFW, 2018), during 2016, global production of cauliflower and broccoli was 25.23 million tons from 1.34 million ha with a productivity of 18.82 tons/ha. Bulk of area and production (>70%) was accounted for by China, I rank (0.52 million ha area, 10.26 million tons production, 19.73 tons/ha productivity) and India, II rank (0.43 million ha area, 8.20 million tons production, 19.0 tons/ha productivity). Other countries in the descending order were USA, Spain, Mexico, Italy, France, Poland, Pakistan and Egypt. Within India, as per 2017-18 figures, top 10 cauliflower growing states are West Bengal (74, 000 ha), Bihar (62, 000 ha), MP (46, 000 ha), Orissa (41, 000 ha), Haryana (40, 000 ha), Gujarat (25, 000 ha), Chhattisgarh (24, 000 ha), Jharkhand (22, 000 ha), UP (18, 000 ha) and Punjab (18, 000).

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21 Bottle gourd

Bottle gourd (Lagenaria siceraria, 2n = 2x = 22, family Cucurbitaceae, monoecious, cross-pollinated but genetically behaving like self-pollinated crop), is grown primarily in India on large scale for consumption as cooked vegetables. Bottle gourd is considered to be one of the earliest species of plants to be domesticated by humans. Bottle gourds are regularly grown and consumed in various parts of the world, particularly Asia. Fruits of bottle gourds are harvested young and used as a vegetable. The mature, dried fruits have been used throughout the world for making storage jars and utensils and hence the name bottle gourd. Shoots and leaves are also cooked and eaten. Lately, bottle gourd juice is also becoming popular in India as a health drink. Other countries growing bottle gourd on limited scale are Sri Lanka, Indonesia, Malaysia, Thailand, Philippines, China, Hong Kong, Tropical Africa, Colombia and Brazil. After seed scooping, the hard rind is used as containers also. Seeds are used for oil extraction and oil is used as cooking oil. During 2017-18, India produced 27 lakh tons (2.7 million tons) of bottle ground from 1.60 lakh ha (0.16 million ha) and this translated into a productivity of 17 tons/ ha (Horticulture Statistics at a Glance-2018, MAFW, 2018). Top six states producing bottle gourd on significant scale are Bihar, UP, Haryana, MP, Odisha, and Pujab, production being in the range of 135,000 to 650,000 tons.

403 - 413 (11 Pages)
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22 Bitter gourd

Bitter gourd (Momordica charantia L., 2n = 2x = 22, family Cucurbitaceae, monoecious, cross-pollinated but genetically behaving like self-pollinated crop), also known as balsam pear, bitter melon, bitter cucumber, and African cucumber is rich source of vitamin C and iron and credited with medicinal use in management of diabetes. Fruits are cooked with other vegetables, stuffed, stir-fried, or added in small quantities to beans and soups to provide a slightly bitter flavor. The bitterness of most cucurbits is mainly due to cucurbitacins. The bitterness of bitter gourd is due to the cucurbitacin-like alkaloid momordicine and triterpene glycosides (momordicoside K and L) as reviewed by Behera et al. (2010). Bitter gourd is grown on approximately 340,000 hectares across Asia, with India and China accounting for more than half the production. Estimated area of bitter gourd in India is about 80,000 ha with production reaching to 10,00,000 resulting into a productivity of 12 tons/ha. States accounting for major area of bitter gourd are Chhattisgarh, Telangana, Andhra Pradesh, Orissa, Madhya Pradesh, Uttar Pradesh, Bihar, Tamil Nadu, Haryana and Assam.

414 - 427 (14 Pages)
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23 Sponge Gourd

Smooth gourd or sponge gourd (2n =2x = 26, Luffa cylindrica Roem syn. aegyptiaca, family Cucurbitaceae, monoecious, cross-pollinated but genetically behaving like self-pollinated crop) is not a global crop and is a vegetable crop very much indigenous to India and the adjoining countries. As per estimates, the acreage of this crop in India is about 50,000 ha spread over almost all the states but on a limited scale. However, UP, Bihar and West Bengal are the main states growing sponge gourd. Origin and Domestication It is agreed that sponge gourd may have originated from Asian sub-tropical areas, particularly India and India itself should be the centre of domestication as well because global presence of this crop is negligible (Kalloo, 1993). NBPGR, New Delhi publications unanimously mention Indo-Chinese and Hindustani gene centres as primary centres/domestication/secondary centres for Luffa species and in their publications, Luffa aegyptiaca is more commonly used than Luffa cylindrica for sponge gourd (Pandey et al., 2019).

428 - 433 (6 Pages)
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24 Ridge Gourd

Ridge gourd also known as ribbed gourd or angled gourd or silky gourd or angled loofah or vegetable gourd (Luffa acutangula (Roxb. L., 2n = 2x, family Cucurbitaceae, mostly monoecious and cross-pollinated but behaving like self-pollinated crop in term of no inbreeding depression) is not a crop of global importance; rather it is grown and consumed primarily in India and the adjoining countries which used to be part of greater India. In India, estimated area is 50,000 ha spread over almost entire country but the leading states are UP, Bihar, West Bengal, MP, Chhattisgarh, and Maharashtra, Karnataka, Tamil Nadu and north east hill region. Origin and Domestication Ridge gourd has similar origin as that of sponge gourd (refer to page 428).

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25 Cucumber

Cucumber (Cucumis sativus var. sativus L., 2x = 2n = 14, normally monoecious, cross-pollinated) is a member of the economically important family Cucurbitaceae which includes squash/pumpkin (Cucurbita ssp.), watermelon [Citrullus lanatus (Thunb.) Matsum. & Nakai], melon (Cucumis melo L.) and gourds. After tomato (Solanum lycopersicum L.) and watermelon, cucumber and melon are cultivated more widely than any other vegetable species globally accounting for about 2.5 million ha and producing 43 million tons giving a productivity of over 17 tons/ha under field and greenhouse culture. Production of cucumber is the second largest of all cucurbits, where China, Iran, Russia, Turkey and the United States represent about 75% of the world production. During 2017-18, India produced 1.25 million tons of cucumber from 0.08 million ha (80, 000) area resulting into a productivity of close to 16 tons/ha (MAFW, 2018). Top ten cucumber producing states in India in the same period are Haryana (17000 ha), MP (9500 ha), Karnataka (8300 ha), Assam (6900 ha), Maharashtra (4500 ha), Andhra Pradesh (4400 ha), Bihar (3900 ha), Punjab (3600 ha), Kerala (3400 ha) and UP (3300 ha).

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26 Pumpkin

Pumpkin (Cucurbita moschata, 2n = 2x = 40, family Cucurbitaceae, monoecious, cross-pollinated, behaving like self-pollinated crop genetically) is grown globally throughout in the tropical, sub-tropical and temperate regions of both the New and Old World. The term pumpkin is normally applied to the edible fruit of any species of Cucurbita when ripe as a table vegetable or in pie (Whitaker and Robinson, 1986). Squash and pumpkin are the words quite often used interchangeably. However, the term squash or summer squash is commonly used for Cucurbita pepo which is eaten as immature fruit also. The term pumpkin is used generally for all kinds of five cultivated species under the genus Cucurbita but is quite commonly used for C. moschata. Besides C. pepo (summer squash) and C. moschata (winter or baking squash), the other three species under the genus Cucurbita are C. mixta (squash), C. maxima (winter or baking squash), and C. ficifolia (squash, fig-leaf gourd, Malabar gourd). While compiling statistics FAO clubs squash and gourd together with pumpkin. In 2017, world production of pumpkins (including squash and gourds) was 27 million tons, with China (7.8 million tons from 0.4 million ha) and India (5.0 million tons from 0.5 million ha) accounting for 47% of the total production. Russia, Ukraine, and the United States each produced about one million ton. As per APEDA (The Agricultural and Processed Food Products Export Development Authority), India produced 1.44 million tons of pumpkin which translates into 72000 ha area assuming a productivity of 20 tons/ha during 2015-16. The major states growing pumpkin in India are Orissa, UP, MP and Chhattisgarh. It is clarified that in context of India, pumpkin means C. moschata only although IIHR released a variety of C. maxima under the name Arka Suryamukhi in 1987 (Ram, 2012). However, now it is nowhere to be seen. Fruits of C. pepo (summer squash or zucchini) now can be seen in limited scale in superstores and limited scale cultivation is certainly happening under protected cultivation of this crop in India. Under this chapter, only C. moschata is being dealt with under the term pumpkin. Pumpkin is now becoming popular in India due to its high productivity, nutritive value, good storability, long period of availability and better transportability. It is rich in vitamins particularly precursor of vitamin A and minerals.

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27 Watermelon

Watermelon (Citrullus lanatus (Thunb.) Matsam. & Nakai, 2n = 2x = 22, family Cucurbitaceae, cross-pollinated crop) is grown commercially in areas with long frost-free warm periods. Three-fourths of the world production is grown in Asia, with China the leading country in production. Watermelon is a nutrient-rich food full of vitamins, minerals and antioxidants. It also contains niacin, thiamin, vitamin B-6, zinc, magnesium and many more nutrients. There are many health benefits to watermelon as it is low in calories so it promotes phyto-nutrients and anti-oxidants essential for the optimization of health. At global level, China tops in watermelon production (79 million tons) with other countries falling far behind, namely, Turkey (3.9 million tons), Iran (3.8 million tons), and India (2.5 million tons). During 2017-18, India produced 2.5 million tons of watermelon from 0.1 million ha (1 lakh ha) acreage and thus Indian watermelon productivity is 25 tons/ha. Top five states having area more than 10,000 ha in India are West Bengal (17,000 ha), UP (14,000 ha), Odisha (12,000 ha), AP (11,000 ha) and Karnataka (10,000 ha).

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28 Muskmelon

Melon or muskmelon (Cucumis melo L., 2n = 2x = 24, family Cucurbitaceae, mostly andromonoecious, cross-pollinated as well as self-pollinated) has latest global production close to 30 million tons where top ten producing countries are China (16.0 million tons), Turkey (1.8 million tons), Iran (1.6 million tons), Egypt (1.1 million tons), India (1.0 million tons), Kazakhstan (0.9 million tons), USA (0.8 million tons), Spain (0.7 million tons), Italy (0.6 million tons), Guatemala (0.6 million tons). India produced 1.2 million tons melons from 54000 ha acreage during 2017-18 giving a productivity of 22 tons/ha (MAFW, 2018). US melons encompass the netted, salmon-flesh cantaloupe; the smooth-skinned green fleshed ‘Honey Dew’, the wrinkled-skinned, white-fleshed ‘Golden Beauty’ and several other dessert melons. Other forms with different fruit characters (white-fleshed, green-fleshed, having striped thin skin and juicy) are commonly found in India and in Orient as indigenous land-race cultivars. These are getting displaced by the cultivar types as cantaloupe, Madhuras and Galia due to their long shelf life. Melon is mainly cultivated for the consumption of the fruits which can be harvested immature; in this case, the fruit is not sweet and can be eaten raw, cooked or pickled. In most cases, the fruit is harvested at maturity with high sugar content (mainly sucrose). Fruit is mainly eaten raw; marginal uses are cubes canned in syrup, in “fruits confits”, candies, ice-creams, biscuits and also in cosmetics. Locally, seeds can also be consumed (Pitrat, 2008).

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29 Onion

Onion (Onion cepa, 2n = 2x 16, highly cross-pollinated, family Amaryllidaceae), is a valuable vegetable crop for the people all over the world. Tomato, onion and potato put together have been given a nick name as TOP seeing their combined importance globally. As per Ministry of Agriculture and Farmers Welfare-Horticulture Statistics Division, Government of India (MAFW, 2018), the global acreage, production and productivity of onion during 2016 were 4.96 million ha, 93.17 million tons and 18.80 tons/ha. Top 10 onion producing countries in the world are China, India, Egypt, USA, Iran, Turkey, Russian Federation, Pakistan, Brazil and Argentina. During 2016, China occupied I rank with area, production and productivity figures of 1.09 million ha, 23.91 million tons and 22.00 tons/ha. India was closely at II rank with corresponding figures of 1.20 million ha, 19.20 million tons and 16.18 tons/ha. Productivity-wise, USA is at the top rank (56.40 tons/ha), followed by Iran (37.95 tons/ha) and Egypt (36.71 tons/ha). The Netherlands is the world’s largest onion exporter with a total of around 800,000 tons, which is ca. 80% of its total annual production (Shigyo and Kik, 2008). Within India, which produced 19.20 million tons of onion from 1.20 million ha (12 lakh ha), the major onion producing states during 2017-18 have been as given

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30 Radish

Radish (Raphanus sativus, 2n = 2x = 18, family Brassicaceae or Cruciferae, cross-pollinated crop) is grown over 7 million ha where China, Japan, India, South Korea, and Europe are the main producers. As per Horticulture Statistics at a Glance-2018 (MAFW, 2018), India produced 30 lakh tons of radish from 2 lakh ha acreage with a productivity of 15 tons/ha during 2017-18. Ten top ranking states in India with acreage within the range of 6000 to 33000 ha are West Bengal, Haryana, Bihar, Assam, Punjab, Chhattisgarh, Orissa, Madhya Pradesh, Uttar Pradesh and Karnataka. Today, radishes are grown throughout the world. Different local people prefer to use various parts of the radish plants including roots, leaves, sprouts, seed pods and oil from seeds as their food according to their own custom. Radishes are low in calories and high in vitamin C, folate, and potassium. Radishes contain sulfurous compounds, such as sulforaphane, which have anti-cancer properties, and are expectorant.

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31 Carrot

Carrot (Daucus carota, 2n = 2x = 18, family Apiaceae or Umbelliferae, cross-pollinated) is among the top ten most economically important vegetable crops globally in terms of area, production and market value (Simon et al, 2008). Almost half of the more than 42 million tons of the global production -by far- is harvested in China, followed by Uzbekistan, Russia and the United States of America. Ukraine, Poland, United Kingdom, Germany, France and Japan complete the top 10.This production comes from approximately an area of 1.5 million ha giving a productivity of 28 tons/ha. India produced 1.6 million tons of carrot during 2017-18 from 96000 ha area resulting into a productivity of 16.66 tons/ha. In India, top 7 states in carrot production are Haryana, Bihar, Punjab, MP, UP, Karnataka and Assam. Because of breeding of cultivars adapted to both summer and winter seasons, carrot is available round the year. The conventional orange colour varieties of Europe are rich in beta carotene, a precursor of vitamin A and contain appreciable quantity of thiamine and riboflavin. The red carrots are rich in lycopene, an important pigment for healthy eyes and general body growth, besides having anti-cancerous properties. Black carrots are rich in anthocyanins. Further, carrots have medicinal properties. Carrot has two distinct groups with regard to flowering and seed production. These are tropical and temperate types. Tropical or oriental cultivars are annual and adapted to high temperature conditions and these do not require low temperature exposure for flowering. They flower and set seeds freely in Indian plains after winter months with the advent of long day conditions in spring. The temperate/European types behave like biennial crops and require 6-8 weeks of vernalization at mean temperature of 4.8 to 10° C after they have passed the full root development phase for production of seed stalks (Sureja and Kalia, 2012).

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32 Vegetable Pea

Vegetable pea also known as garden pea (Pisum sativum, 2n = 2x = 14, family Fabaceae, formley Leguminosae, sub-family Papilionaceae), is rich in vitamins and sugars. Its green immature grains are consumed as fresh vegetables. Picked peas, are canned, frozen and dehydrated or freeze dried for super stores markets. Those associated with the immature pod are the snow, sugar or sugar snap types where pods are consumed. The total pea production world-wise has steadily increased owing to increasing green pea volume and top five countries are China (1.4 million ha), India (0.50 million ha), United States (0.08 million ha), United Kingdom (0.04 million ha) and Algeria (0.04 million ha). For field pea, the major producing countries are Russia, China, Canada, Europe, Australia and United States. India produced 5.42 million tons of garden pea from 0.54 million ha area during 2017-18 giving a productivity of 10 tons/ha. The major garden pea producing states in India in descending order are UP, MP, Punjab, Himachal Pradesh, West Bengal and Assam. UP alone accounts for 40% of the total garden pea area. Within UP, major districts contributing to the vegetable pea are Jalaun, Lalitput, Jhansi, Mahoba, Kasganj, Etah, Sultanpur, Azamgarh, Hamirpur, and Amethi (MAFW, 2018).

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33 End Pages

References Abhary, M., Pati, B. L., Fauquet, C. M. 2007: Molecular biodiversity, taxonomy, and nomenclature of Tomato yellow leaf curl-like viruses. In: H. Czosnek (ed.). Tomato Yellow Leaf Curl Virus Disease: Management, Molecular Biology, Breeding for Resistance, 85—118. Springer, Dordrecht, The Netherlands. Acquaah, G. 2007. Principles of Genetics and Breeding. Blackwell Publishing, USA, UK and Australia. pp 569. Adamus, A., Kielkowska, A., Michalik, B. 2004. Carrot haploid production through induced parthenogenesis. In: Genetic variation for plant breeding. J. Vollmann, H. Grausgruber, and P. Ruckenbauer (eds.), BOKU, Vienna. pp. 451-454. Adluri P, Baldoldiya G. M, Nath P. (2017) Screening of Bhut Jolokia (Capsicum chinense Jacq.) germplasm of North East India against chili leaf curl virus. Int J Pure Appl Biosci 5:1189–1196. Agrama, H.A., Scott, J. W. 2006. Quantitative trait loci for tomato yellow leaf curl virus and tomato mottle virus resistance in tomato. J. Am. Soc. Hortic. Sci. 131:267–272. Agrawal, R. L. 1998. Fundamentals of plant breeding and hybrid seed production. Publishers Inc., Enfield, New Hampshire, USA. Allard, R. W. 1960. Principles of Plant Breeding. John Wiley and Sons, New York, London and Toppan Company, Tokyo. pp 485. Allard, R. W. 1999. Principles of Plant Breeding, 2nd ed., Wiley, New York. Ambrose, M. 2008. Garden Pea. In In J. Prohens and F. Nuez (eds.). Handbook of Plant Breeding-Vegetables II, Springer. pp 3-26. Ambrose, M. J. 2004. A novel allele at the afila (Af) locus and new alleles at the tendril-less (Tl) locus. Pisum Gen. 36, 1-2. Anbinder, H., Reuveni, M., Azari, R., Paran, H., Nahon, S., Shlomo, H., Chen, L., Lapidot, M., Levin, I. 2009. Molecular desection of tomato leaf curl virus resistance in tomato line derived from Solanum peruvianum.Theor. Appl. Genet. DOI 10.1007/s00122-009-1060-z.

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