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Carbohydrate Profiles During Cotton Floral

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Carbohydrate Profiles During Cotton Floral J. Agronomy & Crop Science 192, 363—372 (2006) Ó 2006 Blackwell Verlag, Berlin ISSN 0931-2250 Crop/Stress Physiology Texas A&M Agricultural Research and Extension Center, Beaumont, TX, USA Carbohydrate Profiles During Cotton Floral Bud (Square) Development L. Tarpley, and G. F. Sassenrath AuthorsÕ addresses: Dr L. Tarpley (corresponding author; e-mail: [email protected]), Texas A&M Agricultural Research and Extension Center, 1509 Aggie Dr., Beaumont, TX 77713, USA; Dr G. F. Sassenrath, USDA/ARS Application and Production Technology Research Unit, 141 Experiment Station Rd, Stoneville, MS 38776, USA With 6 figures Received October 21, 2005; accepted March 31, 2006 Abstract because it promotes the placement of the anthers Cotton (Gossypium hirsutum) flower’s showy corolla relative to the stigma such that plentiful amounts of expands rapidly, then senesces quickly. Efficient opening the heavy sticky pollen can land on the stigma. This of corollas is important for self- and cross-pollination, and helps ensure that multiple seeds set in each carpel, indirectly lint yield. Carbohydrate relations are integral to thus providing a high potential lint yield (Poehl- bud water relations and growth, although they are not well man 1977). Furthermore, corolla size has some- understood. Soluble sugars and starch in developing floral times been related to pollinator attractiveness. buds and flowers of field-grown cotton plants were analysed. Buds contained significantly more sugar than Which pollinator attractiveness has sometimes starch. Sucrose and its hydrolysis products strongly been related to pollination efficiency, and pollin- contributed to increased sugars during corolla expansion, ation efficiency is often related to seed set. Thus the and contributed )0.211 MPa to osmotic potential. Water efficient opening of the large inflorescences could concentration was constant throughout expansion at possibly benefit cross-pollination when desired. )1 860 g kg FW, suggesting sucrose import and hydrolysis The elongation rate of the floral bud increases are coordinated with other drivers of expansion. Floral during the last week preceding anthesis (Stewart sugar content declined sharply during senescence (69 % in 1986). This acceleration is thought to reflect a shift 24 h). Although remobilization to other organs could partially explain this decline, we suggest the possibility from cell-division activity to cell expansion, but the that most sugar is broken down via respiration, resulting in manner by which cell expansion in the cotton the production of metabolic water and the significant 7 % corolla is achieved is not known. A decrease in the increase in floral water concentration during senescence. In cellular osmotic potential (increase in osmotic summary, imported sucrose and its hydrolysis products pressure) is probably involved because decreases increased rapidly during cotton floral bud growth, and in osmotic potential often indirectly contribute to supported bud expansion by decreasing the water potential cell expansion by decreasing the water potential in of bud tissue. the cell. Water then moves into the cell down its Key words: anthesis — carbohydrate composition potential gradient. The movement of water into the — cotton — floral bud — plant growth and cell increases the cellular hydrostatic (turgor) pres- development — sucrose sure, which can then act as a driving force for cell expansion by pushing out against the cell wall matrix, which is somewhat elastic in growing cells Introduction such as these. Cotton flowers are showy and ephemeral: rapid However, even if a decrease in cellular osmotic bud elongation leads to a large flower that often potential is involved, a number of questions rela- enters senescence in <1 day. Cotton is both self- ting to cotton floral bud expansion remain unan- and cross-pollinated. The efficient opening of the swered. For example, how is the osmotic potential large inflorescence in cotton fields where self- decreased? If there is substantial accumulation of pollination is being encouraged is important material, such as carbohydrate polymer that can be U.S. Copyright Clearance Centre Code Statement: 0931–2250/2006/9205–0363 $15.00/0 www.blackwell-synergy.com 364 Tarpley and Sassenrath broken down to provide an internal source of starch) from sucrose could assist in the regulation osmoticum during rapid expansion, then are there of or buffering of the degree of osmotic potential other contributors to cell expansion, such as present in the bud tissue cells. Starch accumulation relaxation of the cell wall? What processes are has been shown to occur in floral buds of a number occurring in the bud after bloom and during early of species, possibly including the cotton floral bud senescence, and are these consequential to proces- during early expansion (Zhao and Oosterhuis ses affecting cell expansion in bud elongation? 1998). Many of the species that possess starch- The water used by, or passing through, the accumulating buds have non-ephemeral flowers cotton floral bud is obtained primarily from the that open and close several times over the course of phloem because the floral bud has a higher water several days and nights (Evans and Reid 1988, van potential than that of the surrounding tissue Doorn and van Meeteren 2003). The cotton flower (Trolinder et al. 1993). Thus water movement in does not open again once it has closed, so the role the xylem would be out of the floral buds to the of any starch accumulation in relation to the cotton neighbouring leaves or meristematic tissues (Trol- floral bud expansion is not clear. Perhaps, the inder et al. 1993). The water potential of the accumulation could possibly be regulating solute phloem, in contrast, can include a large hydrostatic content in the bud and thus the rate of bud pressure initially developed by loading of sucrose expansion. The ability to synthesize and accumu- and other solutes into the phloem in source tissues late starch could also possibly act as a means of (van Bel 2003). The phloem is thus capable of buffering the rate of unloading. If active transport supplying solution, and thus water, to the floral relays are present between axially adjoined sieve bud despite the presence of the relatively high water tubes (Thompson and Holbrook 2003), then sud- potential in the floral buds. The supply of water to den changes between loading and unloading rates the floral bud primarily via the phloem has been could cause rapid changes in sucrose concentration demonstrated for flowers of a number of species in the phloem. These kinds of sudden changes (Chapotin et al. 2003). might occur in a rapidly expanding floral bud that This flow in the phloem is largely the hydrostatic is primarily dependent on imported sucrose as an pressure-driven movement of a sucrose-rich solu- osmoticum source. tion. With the possible exception of potassium, Metabolic respiration machinery is found in other solutes are typically present at a much lower floral tissues because they are rapidly developing concentration than sucrose in the phloem. Potas- and growing organs. If this machinery was present sium can be present at up to three-fourths the at high levels, then the metabolic respiration could solute potential of sucrose in the phloem of some affect the carbohydrate balance of the tissues, plants (Lang 1983). If we are to understand the which could lead to effects on the water relations osmoticum/carbohydrate dynamics of the cotton of the tissues. This machinery, however, has not floral bud, then we especially need to understand been documented to exist at high enough levels to the fates of sucrose in the growing floral bud. have a major effect on the carbohydrate balance of Conceptually the carbohydrate relationships of growing floral buds. Nonetheless, respiratory the cotton floral buds should be dominated by breakdown of sucrose or its hydrolysis products different kinds of processes at different stages of could provide some non-reversible regulation of the development. For example, during early bud osmotic potential in the growing floral buds. development, regulation of the rate of expansion Processes that increase the net effect of the is probably necessary. In contrast, during anthesis hydrostatic pressure in the floral bud cells will tend rapid expansion needs to be achieved. After to promote the rapid expansion that occurs during anthesis, the flower or plant might be able to later stages of bud development. These processes recover some of the costs of flowering, for example, would also need to be active during early bud by recycling carbohydrates for use in other parts of expansion, and include factors that lead to an the plant. increase in the hydrostatic pressure, as well as those During the early phases of bud expansion, some that decrease the needed amount of hydrostatic osmoticum is needed to help drive the expansion, pressure to achieve a certain rate of expansion. but photoassimilate that is not used as osmoticum Processes that decrease the osmotic potential in could be used to assist controlled expansion in the floral bud tissue, such as the hydrolysis of various ways. For instance, the regulated forma- starch into oligosaccharides and glucose and the tion and breakdown of carbohydrate polymers (e.g. hydrolysis of sucrose into glucose and fructose, Carbohydrates During Cotton Floral Bud Development 365 would help attract water from the phloem or other from 3 days pre-anthesis to the day of anthesis surrounding tissues. The additional water would (Trolinder et al. 1993). If an increase in soluble lead to an increase in hydrostatic pressure that sugars is involved in cotton corolla expansion, the would help drive expansion in floral bud cells increase is likely to be regulated in coordination (Salisbury and Ross 1978). Regulating the carbo- with other driving forces of expansion, such as cell hydrate hydrolytic activity could thus provide a wall extension. Cells, however, cannot enlarge way of regulating the floral bud cell expansion. indefinitely without additional deposition of cellu- However, unless this regulation of hydrolytic lar structural material.
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