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W1602789585 abstract "From its inception on 1 March 1913, East Malling has focused on the research needs of the fruit-growing industry. Captain R. Wellington, the first Director, defined its mission as ‘the study of problems met with in the actual culture of fruit trees and bushes’. Although the scope of research at East Malling expanded into other crops, and perennial plants more generally, the consistent focus has remained on the science underpinning the production of perennial fruit crops. It is noteworthy, too, that from the beginning East Malling had very close links to the growers of fruit (indeed their financial contributions were essential in the purchase of the land to establish the ‘station’), and those links have persisted through a variety of mechanisms including the East Malling Research (EMR) Association, projects funded via the Horticultural Development Company (which receives a levy from growers) and projects directly funded via individual companies and consortia of companies. This underlying rationale of research allied to the practical needs of growers is reflected today in the current mission and vision of EMR ‘to conduct high-quality strategic and applied research in horticultural and environmental sciences, and deliver knowledge, products and services that benefit public and private customers’ with the aim of being ‘recognised as the pre-eminent research institute in the UK, with a significant international reputation, for strategic and applied research, development and innovation in horticulture with particular emphasis on perennial and clonally propagated crops’. Throughout our 100 years, scientists at East Malling have published their findings in the Annals of Applied Biology, with the first article appearing in 1924 on ‘The mulberry “blight” in Britain’ (Wormald, 1924). Articles on the pests and diseases of fruit crops (and other crops such as hops) have been a particular feature of East Malling's contributions to the journal, with occasional contributions on topics as diverse as use of rootstocks, post-harvest storage, apple replant disease and rooting of cuttings. In this editorial, we briefly review some of the outstanding contributions that research at East Malling has made over the last 100 years to the fruit industry, examine current research themes and then turn our eye to the future and the research that we believe needs to be done to secure nutritious fruits for an expanding, and economically richer, global population. Rootstocks have been at the heart of research at East Malling throughout its 100-year history. An early article was on crown gall disease of rootstocks (Wormald & Grubb, 1924). The causative agent, Agrobacterium tumefaciens, in its disarmed form, is one of the most important tools in the molecular biologist's repertoire for plant transformation and the study of gene function (Bevan et al., 1983). Some 60 years after this report, David James' group at East Malling was the first to successfully transform and regenerate the apple variety Greensleeves (bred at East Malling during the 1960s) using the now familiar binary vector system (James et al., 1989). East Malling is most widely known for its classification and subsequent development of rootstocks for tree fruits during the first 40 years of its existence. Much of this research is summarised in an AAB article by Preston (1956) in which he refers to the early work of Ronald Hatton, a regular contributor to the Annals in its early years (Amos et al., 1930; Hatton et al., 1937). Hatton devised the nomenclature of the first Malling rootstock clones which remains in use today. As a global enterprise, it was quickly realised that the original set of Malling clones would not suffice in areas of the world where pests such as Eriosoma lanigerum (woolly apple aphid), or factors such extreme heat or cold, were common. Preston details some of the pioneering work East Malling undertook with the John Innes Research Station, and the renowned geneticist M.B. Crane, to breed rootstocks with resistance to woolly apple aphid. Besides disease tolerance, another key reason for the use of rootstocks is to control the vigour of the mature tree. The genes underlying the rootstock effects on scion vigour remain undiscovered, although key microscopic observations were made and summarised by Beryl Beakbane in a article that highlighted the morphological variation between vigorous and dwarfing rootstocks (Beakbane, 1956). Differences in the bark/wood ratio, the relative proportions of storage to conduction and strengthening tissues in stems and roots and the size of xylem vessels (i.e. secondary structure) were all correlated with dwarfing, with the most dwarfing rootstocks having more living tissue and less strengthening tissue per unit of root. In later research, morphological variation in leaf structure was also found to be correlated with dwarfing potential (Beakbane, 1967). Similarly detailed work was undertaken to understand the basis of disease resistance in rootstocks. One of the major problems of apple rootstocks is the oomycete pathogen Phytophthora cactorum, which causes a disease called collar rot (Sewell & Wilson, 1973a,b; Sewell et al., 1974). These studies, part of a larger set of articles on Phytophthora infections on apple, highlight two important phenomena, that to this day are poorly understood: the first is how a rootstock influences the resistance of susceptibility of its grafted scion to collar rot, and the second is how seasonal fluctuations in the abundance of different Phytophthora pathogens correlate with the periods of increased host susceptibility. East Malling has made significant contributions to the understanding and control of pests (both insects and mites), and diseases of perennial crops including bacterial, fungal, oomycete and viral agents. Much of the early work was published in the Annals; some highlights are reviewed here. Early pest research at East Malling focused on chemical control (e.g. Gimmingham et al., 1926; Eaton & Davies, 1950), but an appreciation of the role of insects as vectors of viruses to crops was also developing (Prentice & Harris, 1946; Posnette & Ellenberger, 1963). During the 1960s, research on pests in apple orchards focused on red spider mite, Panonychus ulmi, which causes leaf bronzing and fruit russeting (Avery & Briggs, 1968; Briggs & Avery, 1968). The control of this mite (Cranham, 1982a,b) and indeed of other pests such as the damson-hop aphid, Phorodon humili (Muir, 1979) and woolly apple aphid, E. lanigerum (Lyth & Watkins, 1981), was exacerbated by resistance to chemical insecticides—a problem which is still a challenge, although better managed, in modern crop protection programmes (Cross & Berrie, 1996). In addition, chemical control can often disrupt key natural enemies in the crop (Easterbrook, 1997a,b); exploiting pest predators and parasitoids, by either habitat augmentation (Easterbrook & Tooley, 1999) or introductions (Easterbrook et al., 2001; Fitzgerald et al., 2007, 2008) could mitigate the need for some pesticide applications. Studying the biology of pest species (Cranham, 1972, 1973) and the parameters for reproduction and population build-up, coupled with the development of techniques such as electrophoresis (Murray & Solomon, 1978; Fitzgerald et al., 1986), monoclonal antibodies or molecular markers, has permitted an understanding of ‘who eats who’ and ultimately resulted in a reduced need for many broad spectrum chemical control measures. Utilising natural enemies is a free service which can keep pests in check (Campbell, 1978; Easterbrook, 1997a,b; Campbell & Lilley, 1998; Campbell & Cone, 1999). For example, Typholodromus pyri is the main predator of P. ulmi, but is generally sensitive to insecticides sprayed to control the pest (Kapetanakis & Cranham, 1983); the cessation of such pesticides at key times in the predator's lifecycle has reduced damage by the pest. An example of the evolution of control strategies developed at East Malling for a pest is that for blackcurrant gall mite (Cecidophyopsis ribis), which vectors blackcurrant reversion virus (Thresh, 1966) and causes sterility in the bushes and galling of the buds known as big bud disorder (Gordon et al., 1994). Early control used products such as organochlorine and pyrethroid insecticides (Thresh, 1968; Dicker et al., 1972). By fully investigating the emergence of the young mites in relation to meteorological records Cross & Ridout (2001) were able to devise a prediction model of mite emergence. In later research (Cross & Harris, 2006), sulphur was shown to be the most effective acaricide for control of the gall mite; two timed, early season sprays gave superior control to pyrethroids. The gall mite forecasting model is now used regularly by growers and is the basis of management of this important pest in UK blackcurrant crops. As broad spectrum insecticides have decreased in use (Barber et al., 2003), more pests have appeared and methods to prevent, detect and control them have been needed; e.g. Lygus rugulipennis in strawberry (Easterbrook, 1997a,b), vine weevil in container grown plants (Cross et al., 1995) and summer fruit tortix, Adoxophyes orana, in orchards (Cross, 1997). These methods developed hand-in-hand with better targeting of spray applications (Cross & Berrie, 1995) and an ability to reduce the dose applied per hectare (Walklate et al., 2003; Walklate & Cross, 2012), bringing environmental benefits and reduced exposure to spray operators. Early virus work at EMR led to the characterisation and naming of the major viruses that infect strawberry (Prentice, 1948, 1949, 1952; Prentice & Woollcombe, 1951) and other host plants (Posnette, 1947; Posnette & Ellenberger, 1957; Cropley, 1961; Legg, 1964), along with the identification of their vectors (Posnette, 1952). The use of serological tests, known as enzyme-linked immunosorbent assays (ELISAs) for the detection of plant viruses were pioneered at East Malling together with other groups (Clark & Adams, 1977) alleviating the need for time-consuming grafting experiments. These tests are still used today. The development of micro-propagation techniques of strawberry and other clonally propagated crops enabled the production of virus-free plant material and has largely eradicated viruses in commercial crops. Crosse et al. (1960) reported the appearance of Erwinia amylovora, the causative agent of fireblight in pear and apple, for the first time in the UK. Work continued in the 1970s under the direction of Eve Billing and resulted in the development of a model predicting the risk of infection based on weather data (Billing, 1976) named the Billing's integrated system (BIS; Lecomte et al., 1998). Soil-borne diseases are particularly important in perennial crops where annual crop rotations are not possible. Two soil-borne diseases have been studied in particular: Verticillium, which infects strawberry and hop among other hosts, and Phytophthora species, which have a wide host range including apple, strawberry and raspberry. Work on Verticillium albo-atrum, which infects hops, identified strains differing in virulence. These were described as either nonlethal ‘fluctuating’ wilt (M strain) or the lethal ‘progressive’ wilt (V1 strain; Isaac & Keyworth, 1948). This was followed by the emergence of resistance-breaking strains in the 1960s known as ‘super-virulent’ wilt (V2 and V3 strains), suggesting polygenic resistance is in play (Sewell & Wilson, 1980) in this host–pathogen interaction. The hop-growing industry has now reduced in size in the UK, due in part to the difficulties with V. albo-atrum; more recently research has been focused on V. dahliae, which infects strawberry. Apple scab (Venturia inaequalis) is the major apple disease in the UK and consequently has featured in publications throughout the last 100 years. Fundamental research on the biology and epidemiology of V. inaequalis determined the infection conditions (Moore, 1964) and lifecycle (Jeger & Butt, 1983) of this fungus. This informed management practices for control of the disease, including growing season sprays (Burchill & Hutton, 1965) and the use of post-harvest urea treatments to promote the decomposition of leaves, thus destroying the overwintering substrate for the fungus and disrupting the lifecycle (Burchill, 1968). Studies of the epidemiology of V. inaequalis (Jeger et al., 1982) led to the development of mathematical models of the infection process and prediction of epidemics, which together with other disease and pest risk models, was developed into a grower friendly computer-based forecasting system known as ADEM (Xu & Butt, 1997). More recent work on V. inaequalis has monitored the emergence of fungicide resistance (Gao et al., 2009; Xu et al., 2010) and virulence characteristics of V. inaequalis populations (Jeger, 1981; Barbara et al., 2008; Xu et al., 2013). Research leading to the development of modern controlled atmosphere (CA) storage can be traced back to the pioneering experiments carried out by Kidd and West, utilising the facilities in the Ditton Laboratory that became incorporated into the EMR Station in 1969. Early experiments on storage soon led to recommended storage regimes for commercial apple varieties (Kidd & West, 1927, 1936). Subsequent research analysed the factors that influence the storage potential of apples, namely harvest maturity and the mineral (calcium) concentrations in fruits (Stow et al., 1985). Studies showed that relatively high concentrations of carbon dioxide (8%) can be used to maintain firmness of apples without adversely influencing the eating quality (Stow, 1986). Such elevated concentrations of carbon dioxide were also able to reduce the incidence of scald and core browning (Stow, 1986). Experiments with different apple varieties showed that it was possible to reduce oxygen levels to as low as 0.75% to aid the storage of the variety Cox's Orange Pippin in seasons when flesh firmness is low at harvest (Stow, 1989). Cox's Orange Pippin is an important UK apple variety that is renowned for its flavour and aroma, but orchard and seasonal climatic differences are larger sources of variation in firmness and ester content of apples after storage than the maturity of the fruit at the time of harvest (Knee et al., 1990). The focus on Cox's Orange Pippin continued with the analysis of storage regimes aimed at the removal of ethylene, the ripening-promoting gaseous plant hormone. Different methods can be used to remove ethylene, all of which were shown to maintain fruit firmness (Stow & Genge, 1990). The Jim Mount Building replaced the Ditton Laboratory in 1992, and the years of research into storage of various crops and cultivars led to the production of best practice guides for the production of apples and pears that included information on harvest dates and storage regimes. A problem relatively recently resolved was that of Diffuse Browning Disorder (DBD) seen in Cox's Orange Pippin. Initial investigations found that DBD occurred in fruit that had been picked at an optimum stage of maturity for storage and also had favourable mineral composition (Johnson, 2009). Susceptibility to DBD was related to orchard site, and modification of storage conditions did not prevent DBD. After careful analysis of orchard practices triazole sprays were linked to DBD formation (Johnson, 2009). It turned out that the triazole fungicides myclobutanil and penconazole caused more DBD than the triazole plant growth regulator paclobutrazol. However, when both were used together this caused significantly more DBD. DBD was absent in fruits from orchards where triazoles were not used (Johnson, 2009), and thus a solution to the problem was found. Replant disease of apple and cherry (sometimes called “soil sickness”) was first observed systematically at East Malling in 1958. It is a disease syndrome of pathogens and parasites which results in poor establishment of young trees when planted on a site from which mature trees of specific species have been removed. A variety of organisms have been implicated as causal agents including soil bacteria, parasitic nematodes, fungi and oomycetes acting alone or in combination (Mazzola & Manici, 2012). Savory (1967) described five main characteristics of the phenomenon: (a) it is specific to a degree so that apples planted after apple or quince are affected but apples after plums or cherries are not; (b) root growth is inhibited; (c) there are no leaf symptoms, but shoot growth ceases earlier than with a healthy tree; (d) there are initially direct effects on shoot growth, but relative growth rates are similar thereafter and (e) it persists in the soil for a long time. Early experiments at East Malling demonstrated that nematodes alone were not the cause and neither were toxins released from decomposing roots of the previous crops (Savory, 1969). Control of the diseases, or its alleviation, was achieved by partial sterilisation of soil with chloropicrin, but this chemical must be carefully handled (Pitcher et al., 1966; Sewell et al., 1992). The severity of the disease is affected by many factors including soil pH, nitrogen and phosphorus status (Sewell et al., 1992). While various approaches have been used to characterise the aetiology of the disease, there are still different views as to its composition. However, fungal and oomycete species of Phytophthora, Pythium and Rhizoctonia appear to be common (Mazzola, 1998; Mazzola & Manici, 2012). Recent research has aimed to understand the genes and pathways underlying complex trait variation in perennial plants, arthropod pests and plant pathogens. Plants with improved resilience to biotic and abiotic stresses are key to sustainable future food production and also increasingly required in the natural environment as invasive pests and pathogens establish themselves in new environmental niches. Many modern cultivars lack resistance to the most common pests and pathogens that attack them, and industry is dependent on a diminishing repertoire of control products. One such example is the need to deploy soil fumigants such as chloropicrin to guard against infection from the vascular wilt V. dahliae in the cultivated strawberry Fragaria × ananassa. The scheduled withdrawal of this control measure leaves the industry in a precarious position. Recent work to identify resistance to Verticillium in strawberry used QTL mapping to identify multiple loci controlling resistance to Verticillium in the field. Genetic markers (short DNA sequences) in genetic linkage with resistance genes have been identified and screened on the wider germplasm to validate their transferability (Šurbanovski et al., personal communication). Similar techniques have also been applied to apple and raspberry, in the latter case to identify genes controlling resistance to the large raspberry aphid (Sargent et al., 2007). Rapid and inexpensive DNA sequencing technologies have also contributed to the availability of whole genome sequences for a number of important crop species such as apple and peach (Velasco et al., 2010; Verde et al., 2013). Furthermore the model perennial, Fragaria vesca, which has a rapid lifecycle, has a complete genome sequence and a wealth of molecular tools available (Shulaev et al., 2011) which can be utilised to study traits such as fruit development, flowering and disease resistance (Eikemo et al., 2010; Koskela et al., 2012). These genetic and genomic resources will play an important part in understanding the evolution of different facets of perenniality and how gene regulation varies over evolutionary time to modify plant architecture (Vilanova et al., 2008), as well as how natural selection has shaped the species we harness for food production. Recent work in this area has highlighted how population genetics can provide an insight into the evolution of domestic species such as apple, and how potential hybridisation between two geographically separated apple species, Malus sylvestris and Malus sieversii, may have occurred during the spread of apples from East to West over the past 8000 years (Harrison & Harrison, 2011). Crop protection is a continually evolving and changing subject due, in part, to the introduction of new pests and diseases, climate change and changes in pesticide approvals and plant growing systems (e.g. polytunnel use). Current research at EMR works to time and target plant protection products more effectively (Fountain et al., 2012) and, most importantly, to integrate them into modern growing systems as part of Integrated Pest and Disease Management (IPDM) programmes (Cross & Berrie, 2006). Insect pheromone discovery and development (in collaboration with chemical ecologists at the Natural Resources Institute, University of Greenwich) has resulted in the identification of eight midge pheromones including the chrysanthemum gall midge Rhopalomyia longicauda (Liu et al., 2009) and the raspberry cane midge, Resseliella theobaldi (Hall et al., 2009, 2012) which are now used by growers to time applications of pesticides more accurately. Two major breakthroughs include the discovery and exploitation of the aggregation pheromone of the strawberry blossom weevil, Anthonomus rubi (Cross et al., 2006) and the identification of mirid species sex pheromones. The latter was complicated, as the pheromone consists of three compounds, one of which elicited an alarm response if applied in too high a dose (Innocenzi et al., 2004; Fountain et al., 2011a). Current research focuses on the exploitation of plant volatiles which may be repellent to pest organisms, for example cis-jasmone (Birkett et al., 2000; Pope et al., 2007), or attractive to females of the pest. Biological control agents (BCAs) play an important role in sustainable pest and pathogen management programmes, reducing the occurrence of pesticide residues and the potential emergence of pesticide resistance in target organisms. Recent research exploiting indigenous microbial antagonists from orchards has resulted in the discovery of two BCAs (Rungjindamai et al., 2013) which have inhibitory activity against Monilinia laxa, the most important disease on cherry and plum in the UK. Research has also focused on optimising the timing and combinations of commercially available BCAs and increasing the understanding of the environment in which the BCAs are introduced (i.e. interactions with other microorganisms on the plant surface) to improve their efficacy and consistency against disease targets. One technique being used at EMR is to model biocontrol dynamics. There appears to be a lack of synergy in combining two biocontrol agents to manage plant diseases (Xu & Jeger, 2013), a conclusion supported by experimental work (Xu et al., 2011). Controlling orchard pests by augmentation of the surrounding habitat (hedgerow mixes) (Nagy et al., 2010; Fountain et al., 2013) and encouraging earwigs into the trees with refuges and reduced insecticide inputs (Markó et al., 2009) promotes a free service provided by these naturally occurring predators. Resurgence in the interest of pollinating insects (Dicks et al., 2012) has been sparked by dramatic declines in bee populations worldwide. EMR is working towards identifying the most important species for each crop (Fountain et al., 2011b) so that management plans can be targeted to preserve the valuable habitat used by these essential insects. A key component for the control of pests and diseases is the application of knowledge of the epidemiology, ecology, biology and lifecycle to the implementation of management strategies to disrupt them. One such component has been the development of disease forecasting models for strawberry botrytis and mildew using in-crop climate data, which predict the optimum time for the application of protectant or eradicant treatments (Xu et al., 2000; Dodgson et al., 2009). This reduces the number of treatment applications required during the growing season compared to routine fungicide spray programmes (Saville et al., 2013). The application of fewer fungicides during the growing season is an important target for reducing residues in produce, and this was the rationale behind the zero residue management system developed for apple production. The system is based on the use of conventional pesticides up to petal fall and after harvest, but during fruit development in summer only biocontrol agents or sulphur are used. Disease control during the dormant season will minimise inoculum carryover into the new season (Berrie & Cross, 2012). Manipulating canopy structure has also been shown to be effective at producing an unfavourable microclimate for fungal development; for example, reducing raspberry cane density reduces Botrytis cinerea infection (O'Neill et al., 2009). Sustainable intensification, including greater efficiency in the use of resources, is needed to meet the world's increasing demands for food and fibre. Recent research at East Malling has tackled challenges associated with climate change, food security, food chain quality and resource use efficiency. Greater efficiencies in water and nutrient use have been achieved through an improved understanding of responses of horticultural crops to environmental stresses such as soil drying (Cameron et al., 2008; Sharp et al., 2009) and soil flooding (Atkinson et al., 2008; Else et al., 2009). Investigations into the natural variation in drought tolerance in the cultivated strawberry and some of its progenitors (Grant et al., 2012) are also helping to inform longer term efforts to develop new crops with improved water use efficiency and drought resistance. In addition to improving resource use efficiency, manipulating the supply of essential resources such as nutrients and water can also modify the production of plant secondary metabolites involved in plant pest defence. The antimalarial drug artemisinin is commercially extracted from the medicinal plant Artemisia annua, but its leaf concentration can be significantly increased by manipulating boron nutrition (Davies et al., 2011). Use of deficit irrigation to manipulate plant signalling has also been shown to have positive effects on concentrations of key phytonutrients (Dodds et al., 2007), and tolerance to powdery mildew (Xu et al., 2013), and this knowledge is being used to develop beneficial stresses that improve organoleptic quality and extend shelf-life of fresh produce. An improved understanding of the sites of biosynthesis of important phytonutrients such as l-ascorbic acid (Atkinson et al., 2013) is also improving the understanding of how pre-harvest factors can be manipulated to improve post-harvest quality. Coordination of metabolic and physiological activity between plant organs is key to the control of growth and development, especially in clonal plants such as strawberry. The acquisition of essential resources by young ramets is facilitated by the rapid development of transport tissue functionality (Atkinson & Else, 2012) that enables young unrooted ramets to preferentially acquire water and mineral ions compared to older ramets. Sustainable intensification is seen as the main route for meeting the world's increasing demands for food and fibre, so the demands for greater efficiency in the use of resources and less waste are mounting (Gregory et al., 2013). Building on our existing knowledge of crop plant physiology, biochemistry and molecular genetics, our ongoing research is utilising new genomic resources, advances in next generation sequencing, transcriptomics and metabolomics in collaborative projects with universities and industry to optimise both pre- and post-harvest factors to improve food security and minimise waste through the supply chain. Similarly, EMR entomologists and pathologists are undertaking research that will assist growers to tackle new invasive pests and diseases, which are increasing in incidence due to the increased movement of fruit and plant material around the globe. This, in conjunction with new planting systems, artificial substrates, denser plantings, the use of polythene to speed up growth and, of course, a changing and more variable climate leave will provide many challenges. EMR enters its second century with renewed confidence that its research can still contribute to, and resolve ‘problems met with in the actual culture of’ perennial and clonally propagated crops." @default.
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W1602789585 date "2013-06-11" @default.
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W1602789585 title "One hundred years of research at East Malling: science into practice for perennial fruit crops" @default.
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