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General Introduction 01hfdst.02-055 09-07-2002 12:31 Pagina 9 Chapter 1 General introduction Flower bulbs are a major export product of the Netherlands. In 2001, the Netherlands produced an estimated number of 10 billion flower bulbs, which is 65% of the total world production.Tulip, narcissus and lilies are the most important crops. More than half of the flower bulbs is used for the forcing market for cut flowers and pot plants and the rest for the dry sales market for parks and home gardens. More than 75% of the total Dutch production is exported to the United States, Japan and to countries within the European Union. (Sources: Internationaal Bloembollen Centrum, Productschap Tuinbouw) 9 01hfdst.02-055 09-07-2002 12:31 Pagina 10 Chapter 1 Flower bulbs Periodicity of flower bulbs Under natural circumstances, flower bulbs are exposed to seasonal changes in temperature, rainfall and light (Le Nard and De Hertogh, 1993a).The growth cycle of the bulbs is adapted to that periodicity. Depending on the season of flowering, growth activity of the bulbs takes place during spring or autumn. Spring flowering bulbs have a rest period during the summer and resume growth during autumn (Le Nard and De Hertogh, 1993a). They require a warm-cold-warm temperature sequence for growth and development.Typical examples are the tulip, freesia, narcissus and hyacinth. Summer flowering bulbs rest in winter and resume growth in spring. Typical examples of summer flowering bulbs are lily, allium and gladiolus. The time of flowering is not related to the time of flower initiation. Flower initiation can take place during different periods of the year and at different stages of bulb development. Hartsema (1961) distinguished seven periods of floral initiation, which are depicted in Figure 1.1. planting floweringharvesting storage replanting flowering flower initiation 1 2 3 5/6 4 Figure 1.1 Scheme of possible flower initiation/formation periods in flower bulbs according to Hartsema (1961). 1. Flowers are formed more than a year before flowering (e.g. certain amaryllis and nerine species) 2. Flowers are formed almost a year before flowering, just after the previous flowering period (e.g. narcissus) 3. Flower formation occurs after harvest and during storage (e.g. tulip, crocus and hyacinth) 4. Flower formation starts during or towards the end of the storage period, but has to be completed after planting (e.g. lily, dahlia) 5. Flowers are formed after replanting at low temperatures (e.g. iris) 6. Flowers are formed after replanting in spring (e.g. anemone, freesia and gladiolus) 7. Flower and leaf formation alternate and are not dependent on temperature periodicity (e.g. Hippeastrum (amaryllis)) (not shown) 10 01hfdst.02-055 09-07-2002 12:31 Pagina 11 General introduction Forcing From the previous, it may be concluded that most flower bulbs have a life cycle of one year or longer, either flowering in spring or in summer. Nevertheless year-round production of cut flowers has become common practice. For the control of flowering, needed for the year-round production of flowers, bulbs are subjected to specific temperature regimes to affect their growth and development. The use of artificial growth conditions for the flowering of bulbs, to simulate those required by nature, is called “forcing” (Le Nard and De Hertogh, 1993a). Forcing of flower bulbs starts at harvest during the rest period.The applied temperatures during the phase of enlargement affect the physiological state of the bulb at harvest. However, the temperatures in this period can hardly be controlled since it usually occurs outside on the field. Elevated temperatures applied directly after harvesting are used to control the time and rate of flower formation (Le Nard and De Hertogh, 1993a). Subsequently, low temperatures are applied to differentiate the flower bud and initiate rooting.The applied low temperatures are usually necessary to break the rest period or the so-called dormancy of flower bulbs, which is defined as the state of a healthy bulb, characterized by little or no external growth of the shoot and roots (Borochov et al., 1997; Le Nard and De Hertogh, 1993a).After the requirements to break dormancy are fulfilled, bulbs are planted and transferred to a rooting room (to promote root development) or directly to the greenhouse. Depending on the chosen temperature regime flowering can be promoted, retarded or prevented. Disorders in flower bulb development Detailed knowledge of bulb periodicity is important for the control of flowering. When forcing conditions are not properly chosen, physiological disorders in bulb development may occur.The use of incorrect temperature regimes and light conditions during storage and planting often result in leaf aberrations and bud abortion (De Hertogh and Le Nard, 1993b). Furthermore, exposure to ethylene during storage or planting can induce disorders like gummosis, necrosis, flower abscission and abortion (Kamerbeek and De Munk, 1976). Flower bud abortion is a large problem in the forcing of flower bulbs and thus an important subject of this study. De Hertogh and Le Nard (1993b) defined flower abortion as:“the failure of a bulb to produce a marketable flower after the floral organs have been formed”. Bud abortion can emerge after planting in multiple forms, ranging from the absence of a flower (‘blindness’) to the bleaching and withering of the bud. Unfortunately, the effects of the applied forcing conditions and thus the induction of physiological disorders are usually not visible externally. Sometimes, the presence of the disorders can be determined by cutting the 11 01hfdst.02-055 09-07-2002 12:31 Pagina 12 Chapter 1 bulb shortly before planting, but in practice the effects become apparent at flowering.When more knowledge is gained about the processes underlying physiological aberrations like bud abortion, their development might become detectable at an early stage during storage.This would avoid the further costs of storage and planting and it potentially increases the chances to minimize the damage. Knowledge of the physiology of flower bulbs and the availability of a tool to detect distorted developments, will also enable the optimization of forcing conditions of new ornamental flower bulb genera. Finally, with the development of a sensitive detector for aberrations, quality can be better guaranteed. The aim of this study was therefore to gain a better insight in the processes underlying physiological disorders in flower bulbs induced by storage conditions, and to search for tools to assess their internal quality. The present study of the physiological disorders in flower bulbs is focused on the effect of storage temperatures on the two most important crops in bulb breeding: tulip and lily.Tulip and lily are also typical examples of spring and summer flowering bulbs, respectively. Tulip Morphology of the tulip The morphology of a tulip bulb is depicted in Figure 1.2. A tulip bulb usually consists of two to six fleshy scales, enclosing the shoot.The scales are enlarged Figure 1.2 Longitudinal section of a tulip bulb cv. Apeldoorn. The tunic has been removed. 1. site of root primordia 2. basal plate 3. daughter bulb 4. scale 5. first internode of the stem 6. upper internode of the 9 stem (‘neck’) 7. pistil 8. stamen 9. leaves. 8 7 6 4 5 3 2 1 12 01hfdst.02-055 09-07-2002 12:31 Pagina 13 General introduction leaves and serve as the primary storage organs.The outer dry and brown scale is called the tunic and protects the bulbs from infections and mechanical damage.The basal plate is the short fleshy stem axis of the bulb (De Hertogh and Le Nard, 1993a) from which the shoot and roots develop.The basal plate comprises a network of vascular bundles connecting the scales and roots to the shoot. Daughter bulbs are developing in the axils of the scales. Growth of the daughter bulbs occurs mainly during planting.They require a minimal size to be able to flower after planting; otherwise they produce a leaf (Le Nard and De Hertogh, 1993b). Forcing of tulips For the year-round production of tulip bulbs and cut flowers, different forcing programs are nowadays used in the Netherlands.The bulbs can be left uncooled after harvest (in June/July) and planted outdoors or in the greenhouse for flowering around April, according to the natural periodicity. To promote flowering, bulbs are dry-stored 1-2 weeks at moderate (20 °C) or high (30-35 °C) temperatures directly after harvest. Subsequently, the bulbs are transferred to low temperatures (2-9 °C) and dry-stored, stored on trays or on water for a certain period. Hereafter the bulbs are transferred to a rooting room or directly to the greenhouse (17-20 °C). This way, the first flowers can be obtained in November.When the storage temperatures remain higher or when the transfer from high to low temperatures is delayed, flowering is also delayed. To r etard flowering to summer or fall, bulbs are frozen stored (so-called ice-tulips). To obtain these ice-tulips, bulbs are stored until November at moderate temperatures (17-20 °C). Subsequently, the bulbs are either planted in trays for three to six weeks at 9 or 5 °C, to enable rooting, or packed in soil after the cold period. Hereafter, bulbs are frozen at -1.5 to -2 °C. After the summer period the trays are placed in the greenhouse or outside and flowers are obtained in autumn. However, this procedure has not yielded good flower quality, so far. Physiology of the tulip After harvest and during subsequent storage of the bulbs, flower bud and root differentiation are the primary processes occurring. In June/July, a few weeks after harvesting, the flower bud has reached flower stage G (Cremer et al., 1974).This means that all floral organs have been formed, i.e.
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