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Physiology of Flower Bud Growth and Opening

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Physiology of Flower Bud Growth and Opening Proc. Indian Acad. Sci. (Plant Sci.), Vol. 93, No. 3, July 1984, pp. 253-274 Printed in India. Physiology of flower bud growth and opening H Y MOHAN RAM and I V RAMANUJA RAO Department of Botany, University of Delhi, Delhi 110007, India Al~traet. Flower growth and opening are commonplaceevents, but physiologicallyintrieate and inadequately explained. In .this review, we have brou8ht together and evaluated informaª on this subject to for attention on the dynamicfacets of flowerdevelopment. In particular, the physiolo8icalbasis of flowerbud dormancy,nature of cleistogamy,mechanism of flowerbud growth and turgor maintenanceand role of stamens in corolla growth have been examined. The regulation of flower movementsand openingby temperature and light, and circadian rhythmsin floweropening have beendiscuss~ alongwith a considerationof the role of the petal epidermis in light pereeption. It is emphasized that studies on flower physiologyneed to be intensified in view of the lacunae in our basic knowledgeas wellas to providea sound basis for improvingyields of both agricultural and hortieultural crops. Keywords. a-amylase; blossom showers; coroUa;flower bud dormaney;flower bud growth; flower opening; flower movements; flower physiology;invertase; stamens; coffee;gladiolus; Kalanchoe; Lamiwn; Pharbitiy, Ruellia; Turnera. 1. Introduction Angiosperms constitute the dominant and most ubiquitous group of vascular plants and occupy a wide range of ecological habitats. The presence of a flower that bears ovules concealed in a hollow ovary makes them unique. De¡ what a flower is and how it carne to be established in the long course of evolutionary history have engaged the interest of generations of botanists. Lack of fossil evidence and the deep gaps between what is presently considered as the most primitive flower and the most highly evolved anthostrobilus of fossil ancestors are serious handicaps in traeing the origin of the flower. Several widely divergent viewpoints have been expressed in interpreting the fundamdntal nature of the flower, which has a varied structure and organization. Although angiosperms have breeding systems other than sexual reproduction, the main functions of the flower are: formation and release of microspores, reception of pollen, guidance of male gametophytes, gametic union, formation of seeds and associated dispersal structures and auxillary functions for the attraction or exclusion of animal vectors (Pijl 1978). Flowers display various adaptive mechanisms to speed up evolutionary responses to meet the selective pressures by promoting genetic recombi- nation and gene migration. On account of their economic, aesthetic and scientific value, flowers have captivated the attention of professional and amateur botanists alike. Naturalists have also been drawn to the study of flowers because of the close interrelationship between the latter and mammals, birds and insects. For a student of developmental botany the transition from the vegetative to the reproductive phase is a remarkable event. Despite the impressive amount of literature on flower initiation, our present knowledge about flower development, involving specific changes in gene function for the differentiation of floral organs in a close 253 254 H Y Mohan Ram and 1 V Ramanuja Rao succession on the floral axis is quite incomplete. The understanding of the inherent genetic mechanisms and their modification by the environmental and physiological factors are challenging questions for the plant scientist. A glimpse into the work of earlier scientists such as Pfeffer (1897), Goldsmith and Hafenrichter (1932), Ball (1933, 1936) and Bª (1929) on flowers displays originality of approach, a thoroughness in establishing facts anda penetration in understanding complex phenomena about flower structure, growth, movements and functions. The optical and scanning electron microscopic facilities, analytical tools, photographic appliances and the tissue culture technique available to the modern plant scientist ate so refined that dynamic phenomena like flower growth and development can be studied not only in depth but also in a multidisciplinary perspective. On account of limitation of space, only a few sr aspr of flower growth ate dealt with. Floral morphogenesis as understood through the application of in vitro culture (Johri and Ganapathy 1967; Mohan Ram and Jaiswal 1974; Konar and Kitchlue 1982) and experimental modification of flower sex have been covered elsewhere (Heslop- Har¡ 1972; Mohan Raro 1980; Durand and Durand 1984). As corolla generally constitutes the most conspicuous part of the flower, corolla expansion and divergence are considered synonymous with flower growth. In the account that follows, material from classical works that have remained remarkably novel are drawn to illustrate poignant points and advances made in the recent years are discussed at appropriate places. 2. Genetic and environmental regulation of bud dormancy, corolla growth and opening 2.1 Bud dormancy 2.la Low temperature induction ofgrowth: In most plants flower buds, once initiated, follow a continuous course ofdevelopment leading to anthesis and fruit formation. In some perennial plants especially the temperate deciduous trees, flower buds undergo a pr ofdormancy. Bud dormancy is broken by exposure to low temperatures. Under natural conditions chilling (1 -10 ~C) in winter fulfils this requirement. Ir the winters ate mild and do not extend over a sufficient length of time, the buds rail to biossom and set fruit. In sour cherry, bud growth is arrested after the sepal and petal primordia have been formed, anthers areat the stage of sporogenous cells and ovule primordia have not yet been differentiated (Felker et al 1983). In sour cherry, flower bud chilling is accompanied by meiosis, first in the anthers and then in the ovule. Importantly, starch accumulation takes place with the progression of chilling. Ultrastructural studies indicate that du¡ dormancy, carbohydrate meta- bolism plays a more crucial role than nuclear events (Felker et al 1983). However, in Rhododendron flower buds, starch accumulation occurs before the rest pe¡ and its utilization is postponed until the last weeks of cell expansion prior to anthesis (Schneider 1972). Temperature is an important factor affecting flower initiation and development in bulbous plants (Hartsema 19£ Low temperature promotes the translocation of gibberellin-like substances from the bulb scales of iris to the shoot apex (Rodrigu› Pereira 1964). Whereas the concentration ofgibberellins in the bulb scales does not vary Flower bud growth and opening 255 much, a larger bulb would contain more total gibberellins than a smaller one. Work on narcissus, tulip and hyacinth (see Rees 1972) has shown that flower initiation and differentiation takes place in warm temperatures, but flower maturation and anthesis are rapid only when the plants become exposed to low temperatures during of after flower differentiation. Although the precise role of gibberellins is not known, it has been suggested that in iris they are involved in flower initiation and flower stem development and in tulip in flower extension. In narcissus flower primordia may be laid down one year before anthesis. Thus at the time of planting, a bulb contains already formed flower buds. In Zephyranthes, flower initiation alternates with leaf formation throughout the growing period. Thus tiny immature buds and large buds approaching the point of anthesis occur in a dormant state in the bulb. In Z. rosca flower growth and opening are promoted when the temperature is lowered in the presence of water (Kerling 1949). It is also important that the period oflow temperature exposure exceeds 10 hr. Kerling (1949) was of the opinion that low temperature caused auxin release which promoted pedicel growth. Holdsworth (1961) working with Z. citrina, Z. tubispatha and with two species of Pancratium showed that increased soil moisture after rains could be the factor trigge¡ gregarious flowering. The situation is thus reminiscent of that in coffee. 2.1b Role of moisture in breaking bud dormancy in coffee: In certain coffee growing arcas, flower buds undergo a period of rest prior to meiosis after attaining a certain stage of development. Bud opening occurs only after rer a rainfall--appropriately termed "blossom shower" (Went 1917; van der Meulen 1939; Mathew and Chokkanna 1961). In Chickmagalur arca in Karnataka, flower buds ate initiated in October and remain dormant until April the following year. Failure of blossom showers can cause a se¡ set back in coffee production. Leliveld (1939) had noted that meiosis in C. canephora was dependent on the occurrence of rain. Mes (1957a) showed that flower bud dormancy could be broken and meiosis triggered by submerging the bud-bearing branches of C. arabica in water of supplying water by other mcans. Water treatment of twigs bearing buds has now become a standard method for studying meiosis in Coffea spp (Sreenivasan 1983). According to Alvim (1958) the moisture effect is also dependent on temperature, higher temperatures being more productive. Whereas Mes (1957b) believed that dormancy was caused by water stress in the young flower buds, Alvim (1960) observed that water stress was not the cause of dormancy but a condition necessary for overcoming it. It was only after experiencing a period of water stress that coffee plants become receptive to water, resulting in resumption of flower bud growth. Continuous growth under saturated conditions does not promote flower bud
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