Preface Horticultural products for human consumption are an important part of the market economy. Any waste due to pathogen attack and spoilage in the field but mostly during the postharvest phase, results in significant economic losses. Postharvest diseases are very significant under the present scenario of postharvest management, which is insufficient to meet the requirements in several countries. Such losses are more pronounced as the losses occur closer to the time of produce sale. Microbial decay is one of the main factors that determine losses compromising the quality of the fresh produce. Optimal distribution of fresh horticultural products entails prolonging their freshness and nutritional quality as long as possible after harvest. Losses caused by postharvest diseases are greater than generally realized because the value of fresh fruits and vegetables increases several-fold while passing from the field to the consumer. Owing to its significance, ‘Postharvest Plant Pathology’, as a subject is being studied either in Undergraduate or Postgraduate degree programmes at Agricultural and Horticultural Universities, for which a specific syllabus has been prescribed by the Indian Council of Agricultural Research (ICAR), New Delhi, India. At present, there is scarcity of a book for the use by the students that covers the entire syllabus prescribed by the ICAR, New Delhi for ‘Postharvest Plant Pathology’ course. This book explicates the fundamental aspects of postharvest diseases of crops and is conveniently divided into ten chapters, providing the latest information on the concept & types of postharvest diseases, economically significant postharvest pathogens & diseases of major crops, factors governing postharvest diseases, storage conditions, food safety issues, quiescence in post harvest pathogens, detailed & recent information on major mycotoxins, various approaches of postharvest disease management, integrated management strategies, biochemical & molecular aspects of postharvest diseases, apart from which, an exclusive chapter for discussing the postharvest nematode diseases and their management is also furnished. Impressive diagrams at appropriate places, convincing tables and suitable graphs / illustrations have been furnished. A bibliography providing the list of references cited has also been included. The information presented in this book, ref lecting an extensive literature search, will be useful for teachers, researchers, students in several departments including Plant Pathology, Microbiology, Food Technology, Postharvest Technology, Environmental Sciences, officials of the State Departments of Horticulture & Agriculture, personnel of Commercial Production Centers, Plant Quarantine, Certification agencies and Policy planners who are concerned with the production and supply of horticultural and agricultural produce of high quality and acceptable for human consumption.

Food waste and loss is an enormous issue worldwide. The FAO states that ‘Hunger is still one of the most urgent development challenges, yet the world is producing more than enough food’, but a vast amount of what is produced gets lost on its way from the field to the consumer. According to the FAO, about 30% of cereals, 20% of dairy products, 35% of fish and seafood, 45% of fruits and vegetables, 20% of meat, 20% of oilseed and pulses and 45% of roots and tubers are lost or wasted. Postharvest food loss is defined as measurable qualitative and quantitative food loss along the supply chain, starting at the time of harvest till its consumption or other end uses. Food losses can either be the result of a direct quantitative loss, for example, during the process of harvest or transport or arise indirectly due to losses in crop quality such as undesired sprouting or water loss. Postharvest food loss consists of many factors, which have not changed much in the last 40 years and so postharvest food loss in general has not changed much. Figure 1 illustrates the main reasons for food loss along the supply chain and is based on ‘The food pipeline’ published by Bourne in 1977. Food loss is highest during storage due to pathogens (insects, bacteria and moulds), environmental conditions (e.g. rain, humidity, heat and frost), sprouting and quality loss (rancidity, water loss and saccharification) or animals (rodents and birds). Recently published data for Switzerland have shown that about 53%-55% of the initial fresh potato production and 41%-46% of the initial processing potato production are lost mostly due to pathogen infection, saccharification, water loss of tubers and early sprouting during storage (Buchholz et al, 2018).

Fruits, nuts and vegetables play a significant role in human nutrition, especially as sources of vitamins, minerals, dietary fibre, and antioxidants. Increased consumption of a variety of fruits and vegetables on a daily basis is highly recommended because of associated health benefits, which include reduced risk of some forms of cancer, heart disease, stroke, and other chronic diseases. Both quantitative and qualitative losses occur in horticultural commodities between harvest and consumption. Qualitative losses, such as loss in edibility, nutritional quality, caloric value, and consumer acceptability of fresh produce, are much more difficult to assess than are quantitative losses. Quality standards, consumer preferences and purchasing power vary greatly across countries and cultures and these differences influence marketability and the magnitude of postharvest losses. Postharvest losses vary greatly across commodity types, with production areas and the season of production. Losses of fresh fruits and vegetables in developed countries are estimated to range from 2 percent for potatoes to 23 percent for strawberries, with an overall average of 12 percent losses between production and consumption sites. In contrast, the range of produce losses in developing countries varies widely. Losses at the retail, food-service, and consumer levels are estimated at approximately 20 percent in developed countries and about 10 percent in developing countries (Adel A. Kader and Rosa S. Rolle, 2004 ). Overall, about one third of horticultural crops produced are never consumed by humans. Reduction of postharvest losses can increase food availability to the growing world population, decrease the area needed for production, and conserve natural resources. Strategies for loss prevention include use of genotypes that have longer postharvest-life, use of integrated crop management systems and good agricultural practices that result in good keeping quality of the commodity and use of proper postharvest handling practices in order to maintain the quality and safety of fresh produce.

Postharvest loss of fruits and vegetables is defined as “that weight of wholesome edible product (exclusive of moisture content) that is normally consumed by human and that has been separated from the medium and sites of its immediate growth and production by deliberate human action with the intention of using it for human feeding but which for any reasons fails to be consumed by human.” Not only quantity and quality but even the appearance of fruits and vegetables are affected, and their market value is reduced. Most ‘skin deep’ injuries such as ‘fly speck’ in apples do not affect the edible part. Some infections are not harmful as they occur on inedible parts and can be trimmed off before consumption ego Colletotrichum spots on the outer leaves of cabbage]. Fresh fruits and vegetables are perishable and highly prone to these losses because they are composed of living tissues. These tissues must be kept alive and health throughout the process of marketing. These are composed of thousands of living cells which require care and maintenance (Alao, 2000). The pathological rots are the most serious which is followed by mechanical injury. Pathological rots in combination with mechanical damage cause serious damage to the perishables. Environmental factors such as temperature, relative humidity and oxygen balance most especially during storage are also greatly responsible for damage also environmental conditions such as temperature and humidity are responsible for rendering fruits and vegetables susceptible for pathological attacks (Yahaya and Mardiyya, 2019). Fungi and bacteria are the major pathogens, which cause postharvest diseases in crops leading to about 30 % loss. Normally diseases will be more at improper temperature and relative humidity. Apart from causing disease, pathogens are also involved in ethylene production, which hastens the deterioration of the product.

Postharvest handling is the stage of crop production immediately following harvest, including cooling, cleaning, sorting and packing. The instant a crop is removed from the ground, or separated from its parent plant, it begins to deteriorate. Postharvest activities include harvesting, handling, storage, processing, packaging, transportation and marketing. Losses of horticultural produce are a major problem in the postharvest chain. The diseases which develop on harvested parts of the plants like seeds, fruits and also on vegetables are the postharvested diseases. The amount or extent of damage depends mainly on the pathogen(s) involved, on the condition of the products and the condition of storage. Major tips to avoid postharvest losses include Tips to avoid postharvest losses include assessing maturity, to check your water quality, to check water temperature, avoid injury, keep the produce cool and proper storage. “Storage” means the phase of the postharvest system during which the products are kept in such a way as to guarantee food security other than during periods of agricultural production at the marketing level, to balance the supply and demand of agricultural products, thereby stabilizing market prices. Grain, which includes dry kitchen ingredients such as flour, rice, millet, couscous, cornmeal, and so on, can be stored in rigid sealed containers to prevent moisture contamination or insect or rodent infestation. For kitchen use, glass containers are the most traditional method. Maturity indices are the sign or indication the readiness of the commodity for harvest. It is the basis for determining harvest date. Postharvest losses occur between harvest and the moment of human consumption. They include on-farm losses, such as when grain is threshed, winnowed, and dried, as well as losses along the chain during transportation, storage, and processing.

In managing diseases of horticultural crops, pathogen quiescence has important implications in timing prophylaxis, the reduction of stresses that trigger the transition of quiescence to aggression, and managing the harvested crop to prolong quiescence to the point at which yield is no longer affected. The inception of pathogen quiescence and its maintenance on or within the host implies a dynamic equilibrium between the host, the pathogen, and their environment. Physiological and physical changes in the host, its environment, or both trigger changes in that equilibrium to permit the pathogen to resume aggression. Microorganisms lying on the host surface constitute a potential latent infection; as can be the case with Botrytis cinerea conidia that are prevented from germinating by antagonistic bacteria on leaf surfaces (William R. Jarvis, 1994). Quiescence can run the spectrum from the ungerminated spore; through symptomless, internal infections; to visible but nonexpanding lesions, such as ghost spot of tomato (Lycopersicon esculentum) to expanding lesions on banana caused by Colletotrichum musae and on avocado caused by C. gloeosporioides. A latent, dormant, or quiescent parasitic relationship is a condition in which the pathogen spends long periods during the host’s life in a quiescent stage until, under specific circumstances, it becomes active. Epidemiologists might refer to the period from spore landing until the parasitized tissue produces new spores as a latent period (Dov Prusky, 1996). The term “period of quiescence” is referred to differentiate the quiescent parasitic relationship from the latent period. Quiescence can be enforced during the various processes of fungal attack, which suggests that quiescence may or could occur during any of the processes leading from fungal germination to colonization.

Mycotoxins are naturally occurring toxins produced by certain molds and can be found in food. The molds grow on a variety of different crops and foodstuffs including cereals, nuts, spices, dried fruits, apples and coffee beans, often under warm and humid conditions. Mycotoxins can cause a variety of adverse health effects and pose a serious health threat to both humans and livestock. The adverse health effects of mycotoxins range from acute poisoning to long-term effects such as immune deficiency and cancer. A scientific expert committee jointly convened by WHO and the Food and Agriculture Organization of the United Nations (FAO), called JECFA, is the international body responsible for evaluating the health risk from natural toxins including mycotoxins. International standards and codes of practice to limit exposure to mycotoxins from certain foods are established by the Codex Alimentarius Commission based on JECFA assessments. Mycotoxins are toxic compounds that are naturally produced by certain types of molds (fungi). Molds that can produce mycotoxins grow on numerous foodstuffs such as cereals, dried fruits, nuts and spices. Mold growth can occur either before harvest or after harvest, during storage, on/in the food itself often under warm, damp and humid conditions. Most mycotoxins are chemically stable and survive food processing. Several hundred different mycotoxins have been identified, but the most commonly observed mycotoxins that present a concern to human health and livestock include aflatoxins, ochratoxin A, patulin, fumonisins, zearalenone and nivalenol/deoxynivalenol. Mycotoxins appear in the food chain as a result of mold infection of crops both before and after harvest. Exposure to mycotoxins can happen either directly by eating infected food or indirectly from animals that are fed contaminated feed, in particular from milk.

The potential for significant economic losses from decay caused by postharvest pathogens is greater than is often realized. Harvesting and handling, and the attendant costs for labor and postharvest chemical treatments, can increase the cost of fresh fruits and vegetables several fold from the field to the consumer. Therefore, any produce lost in storage is worth more economically than an equivalent amount lost in the field. Considering economics and the problem of world hunger, it is imperative to understand the importance of postharvest disease and to continually strive to reduce postharvest losses from pathogens. In recent years, many international organizations have identified the necessity of reducing postharvest food losses as an important worldwide goal. While postharvest losses vary from season to season, an average loss of 10% to 30% after harvest is an average estimate. Significantly higher losses are not uncommon in developing countries. The extent of these losses justifies increased research efforts in postharvest pathology (Carl E. Sams, 1994). Chemical treatments and irradiation have effectively reduced the number of microorganisms on various commodities. However, consumer fears associated with chemical residues and the effects of ionizing radiation on fresh produce are forcing the development of alternative protection methods. Also, many pathogens have developed resistance to several commonly used chemicals, while other decaycausing microorganisms currently cannot be controlled by chemicals. Therefore it is better to seek control strategies that will supplement or replace current disease control methods.

A wide variety of fungal and bacterial pathogens cause postharvest disease in fruits and vegetables. Some of these infect produce before harvest and then remain quiescent until conditions are more favourable for disease development after harvest. Other pathogens infect produce during and after harvest through surface injuries. In the development of strategies for postharvest disease control, it is imperative to take a step back and consider the production and postharvest handling systems in their entirety. Traditionally fungicides have played a central role in postharvest disease control. However, trends towards reduced chemical usage in horticulture are forcing the development of new strategies. Controlling or reducing disease relies on integrated crop and postharvest management, with attention to fungicide application, crop hygiene and nutrition, and the management of ripening, to optimise advantages conferred by the plant’s natural resistance factors that prevent and delay disease development. Agricultural production and food distribution systems have evolved over many years to a complex system which allows for nearly year-round supply of most fresh commodities through longterm storage and long distance shipments to consumers. The expectation of year round supplies has resulted in increased challenges for postharvest handling systems. The availability of fungicides for disease management likely played a major role in the evolution of our current system of food distribution (Mitcham, 1999). With the recent loss of registration of many postharvestfungicides and more expected under the Food Quality Protection Act (FQPA), as well as consumer pressure to reduce the amount of chemicals used on food, the challenge to maintain our current system of food distribution increases.

Postharvest food loss has been considered an unavoidable affliction since food storage first began. These losses are much more painful and costlier in terms of money and man-hours. Such losses due to plant parasitic nematodes, in particular, is severe in tropical regions especially in developing countries like India, where proper storage, transmit and marketing facilities are inadequate. In tropical agriculture, increase in production rate is generally low and, by itself, unable to keep pace with the requirements of the increasing population. Hence, such food shortage demands the need to reduce postharvest loss due to nematodes. Postharvest food losses also constitute a vast complex of physical and biological changes, which makes it very difficult to represent in a single figure. As suggested by different sources, an overall loss of at least 10 per cent occurs in the developing countries. However, the magnitude may range from negligible to cent percent in particular nematode species and crop. In addition to causing direct losses individually, phytonematodes also favour the establishment of secondary, passive micro- organisms like fungi, bacteria, viruses etc. Nematodes cause mechanical wounds, which serve as avenue for the entry of secondary microorganisms (Ravichandra, 2008). This complex association results in severe damage to planting material. Postharvest diseases due to nematodes can be managed through restricting the movement of infected planting material to uninfected areas; use of uninfected, healthy planting materials; use of treated planting materials; improvements in the cultural practices at the growing site; timely and careful harvesting; use of nematicides both at pre- and postharvest stages; employment of some physical means like heat, radiation etc; improved storage techniques and integration of these methods. By maintaining hygienic conditions during harvest, transportation, storage and packing most of the postharvest damage due to nematodes can be minimized. Of all the methods, however, regulatory measures like quarantine is more practical and effective in restricting the movement of nematode infected plant materials and contaminated soil into a state or country. Contaminated plant material should be treated (hot water or nematicides) to kill the plant parasitic nematodes before planting to prevent the further postharvest damage. These various measures are primarily aimed at either reduction or wiping out of stages of nematode from the site or protection of host from infection or creation of unfavorable host and/or environmental conditions for the nematodes.

Postharvest losses of fruits and vegetables are even higher and are estimated to be 40-50%. Postharvest fruit rotting are a major cause of those losses and are chiefly caused by fungal pathogens after fruit ripening. In a manner similar to foliar diseases that occur in the field, several factors as fungal pathogenicity, host response and environment determine the outcome of host resistance or susceptibility. However, in postharvest diseases fruit ripening is another major component that will determine fruit resistance. The molecular trigger induces the germ tube to differentiate an appressorium. It has been shown that excess nutrients over ride it and protein synthesis is not required once germination has occurred. Protein synthesis during conidium germination and appressorium formation on cellulose membranes were observed (Pathak, 1997). A specific polypeptide (95 kDa) was synthesized only when appressoria matured (a period lasting from 12-24 h after germination). This molecule is involved in the synthesis of the melanin precursor, Scytalone. Appressoria grown in the presence of cyclohexamide do not mature nor they subsequently differentiate a penetration peg. Appressorium maturation involves the formation of melanin. It has been shown that melanization of appressoria is necessary for formation of a successful penetration peg. The melanin appears to give the appressorium mechanical strength, but pigmentation is only one factor required for penetration competent appressoria. At restrictive temperatures, pigmentation of conidia of C. lagenarium mutants could be restored with dihydroxyphenylalanine without restoration of appressorial competence. Non-pathogenic mutants have been used to determine factors important for pathogenicity. Cutinase production was essential for penetration of papaya fruits. The cutinase produced during infection of papaya fruit by C. gloeosporioides has been purified and characterized as a glycoprotein of 24 kDa. Polyclonal antibodies raised against this enzyme do not cross react with cutinase from Fusarium so/ani, and lesions do not develop when papaya fruits are inoculated with spore suspensions containing the antibody preparation.