Debunga ialah serbuk halus hingga kasar yang mengandungi mikrogametofit benih tumbuhan, yang menghasilkan gamet jantan (sel sperma). Butiran debunga mempunyai lapisan keras yang melindngi sel sperma semasa proses pemindahan dari stamen ke pistil tumbuhan berbunga atau dari kon jantan ke kon betina tumbuhan konifer. Apabila debunga tiba di pistil serasi tumbuhan berbunga, ia bercambah dan menghasilkan tiub debunga yang memindahkan sperma kepada ovul ovari pokok tersebut. Butiran individu debunga cukup kecil untuk memerlukan pembesaran bagi melihat perincian.
Struktur debunga [sunting]
Debunga sendiri bukan merupakan gamet jantan, tetapi setiap bijian debunga menandungi sel (cuma satu sel dalam kebanyakan pohon berbunga dan beberapa dalam tumbuhan benih lain) vegetatif (tidak membiak) dan sel generatif (membiak) yang mengandungi dua nukleus: a tube nucleus (that produces the pollen tube) and a generative nucleus (that divides to form the two sperm cells). The group of cells is surrounded by a cellulose-rich cell wall called the intine, and a resistant outer wall composed largely of sporopollenin called the exine.
Pollen is produced in the microsporangium (contained in the anther of an angiosperm flower, male cone of a coniferous plant, or male cone of other seed plants). Pollen grains come in a wide variety of shapes (most often spherical), sizes, and surface markings characteristic of the species (see Electron micrograph at top right). Pollen grains of pines, firs, and spruces are winged. The smallest pollen grain, that of the Forget-me-not (Myosotis spp.), is around 6 µm (0.006 mm) in diameter. Wind-borne pollen grains can be as large as about 90-100 µm. The study of pollen is called palynology and is highly useful in paleoecology, paleontology, archeology, and forensics.
In angiosperms, during flower development the anther is composed of a mass of cells that appear undifferentiated, except for a partially differentiated dermis. As the flower develops, four groups of sporogenous cells form with in the anther, the fertile sporogenous cells are surrounded by layers of sterile cells that grow into the wall of the pollen sac, some of the cells grow into nutritive cells that supply nutrition for the microspores that form by meiotic division from the sporogenous cells. Four haploid microspores are produced from each diploid sporogenous cell called a microsporocyte, after meiotic division. After the formation of the four microspores, which are contained by callose walls, the development of the pollen grain walls begins. The callose wall is broken down by an enzyme called callase and the freed pollen grains grow in size and develop their characteristic shape and form a resistant outer wall called the exine and an inner wall called the intine. The exine is what is preserved in the fossil record.
The pollen wall protects the sperm nucleus while the pollen grain is moving from the anther to the stigma, it protects the vital genetic material from drying out and solar radiation. The pollen grain surface is covered with waxes and proteins, which are held in place by structures called sculpture elements on the surface of the grain. The outer pollen wall prevents the pollen grain from shrinking and crushing the genetic material during desiccation and it is composed of two layers. These two layers are the tectum and the foot layer, which is just above the intine. The tectum and foot layer are separated by a region called the columella, which is composed of strengthening rods. The outer wall is constructed with a resistant biopolymer called sporopollenin. The pollen tube passes through the wall by way of structures called apertures.
Pollen apertures are any modification of the wall of the pollen grain. These modifications include thinning, ridges and pores, they serve as an exit for the pollen contents and allow shrinking and swelling of the grain caused by changes in moisture content. Furrows in the pollen grain are called colpi, which along with pores, are a chief criteria for the identifying pollen classes.
Pollen grains may have furrows, the orientation of which (relative to the original tetrad of microspores) classify the pollen as colpate or sulcate. The number of furrows or pores helps classify the flowering plants, with eudicots having three colpi (tricolpate), and other groups having one sulcus.
Except in the case of some submerged aquatic plants, the mature pollen-grain has a double wall, a thin delicate wall of unaltered cellulose (the endospore or intine) and a tough outer cuticularized exospore or exine. The exine often bears spines or warts, or is variously sculptured, and the character of the markings is often of value for identifying genus, species, or even cultivar or individual. In some flowering plants, germination of the pollen grain often begins before it leaves the microsporangium, with the generative cell forming the two sperm cells.
The transfer of pollen grains to the female reproductive structure (pistil in angiosperms) is called pollination. This transfer can be mediated by the wind, in which case the plant is described as anemophilous (literally wind-loving). Anemophilous plants typically produce great quantities of very lightweight pollen grains, sometimes with air-sacs. Non-flowering seed plants (e.g. pine trees) are characteristically anemophilous. Anemophilous flowering plants generally have inconspicuous flowers. Entomophilous (literally insect-loving) plants produce pollen that is relatively heavy, sticky and protein-rich, for dispersal by insect pollinators attracted to their flowers. Many insects and some mites are specialized to feed on pollen, and are called palynivores.
In non-flowering seed plants, pollen germinates in the pollen chamber, located beneath and inside the micropyle. A pollen tube is produced, which grows into the nucellus to provide nutrients for the developing sperm cells. Sperm cells of Pinophyta and Gnetophyta are without flagella, and are carried by the pollen tube, while those of Cycadophyta and Ginkgophyta have many flagella.
When placed on the stigma of a flowering plant, under favorable circumstances, a pollen grain puts forth a pollen tube which grows down the tissue of the style to the ovary, and makes its way along the placenta, guided by projections or hairs, to the micropyle of an ovule. The nucleus of the tube cell has meanwhile passed into the tube, as does also the generative nucleus which divides (if it hasn't already) to form two sperm cells. The sperm cells are carried to their destination in the tip of the pollen-tube.
Debunga sebagai pembawa maklumat ekologi tumbuhuan [sunting]
A Russian theoretical biologist, Vigen Geodakyan (Geodakian), has suggested that the quantity of pollen reaching a pistillate flower can transmit ecological information and also regulate evolutionary plasticity in cross-pollinating plants. Plentiful pollen indicates optimum environmental conditions (for example a plant that is situated at the center of its natural range, in ideal growing conditions, with a large number of male plants nearby, and favorable weather conditions), whereas a small amount of pollen indicates extreme conditions (at the borders of its range, with a deficiency of male plants, and adverse weather conditions). Geodakian believes that the quantity of pollen reaching a pistillate flower defines the sex ratio, dispersion and sexual dimorphism of a plant population. High pollen quantity leads to a reduction of these characteristics and stabilization of a population. Small quantity leads to their increase and destabilization of a population.
Dependence of the secondary sex ratio on the amount of fertilizing pollen was confirmed on four dioecious plant species from three families — Rumex acetosa (Polygonaceae), Melandrium album (Cariophyllaceae), Cannabis sativa and Humulus japonicus (Cannabinaceae). (see summary of all these data in review article).
Dependence of offspring phenotype variety on amount of pollen was observed by Ter-Avanesyan in 1949. All three studied species of plants (cotton plant, black-eyed pea, and wheat) showed dependence in the direction forecast by the theory — fertilization with a small amount of pollen resulted in an increase in the diversity of the offspring. Ter-Avanesian writes that as a result of a limited pollination “instead of homogenous sorts we get populations”.
Debunga dalam rekod fosil [sunting]
Pollen's sporopollenin outer sheath affords it some resistance to the rigours of the fossilisation process that destroy weaker objects; it is also produced in huge quantities. As such, there is an extensive fossil record of pollen grains, often disassociated from their parent plant. The discipline of palynology is devoted to the study of pollen, which can be used both for biostratigraphy and to gain information about the abundance and variety of plants alive — which can itself yield important information about paleoclimates. Pollen is first found in the fossil record in the late Devonian period[perlu rujukan] and increases in abundance until the present day.
Demam jerami [sunting]
Lihat juga [sunting]
|Wikimedia Commons mempunyai media berkaitan: Debunga|
- European Pollen Database
- Evolution of sex
- Pollen count
- Pollen source
- Polyphenol antioxidant
- Sex ratio
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- Pleasants, Jm; Hellmich, Rl; Dively, Gp; Sears, Mk; Stanley-horn, De; Mattila, Hr; Foster, Je; Clark, P dll. (October 2001). "Corn pollen deposition on milkweeds in and near cornfields" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 98 (21): 11919–24. doi:10.1073/pnas.211287498. PMC 59743. PMID 11559840. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=11559840.
- Kenneth R. Sporne (1972). "Some Observations on the Evolution of Pollen Types in Dicotyledons". New Phytologist 71 (1): 181–185. doi:10.1111/j.1469-8137.1972.tb04826.x. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1469-8137.1972.tb04826.x.
- Walter S. Judd and Richard G. Olmstead (2004). "A survey of tricolpate (eudicot) phylogenetic relationships". American Journal of Botany 91: 1627–1644. doi:10.3732/ajb.91.10.1627. (full text)
- Geodakyan V. A. (1977). The Amount of Pollen as a Regulator of Evolutionary Plasticity of Cross-Pollinating Plants. “Doklady Biological Sciences” 234 N 1-6, 193–196.
- Соrrеns С. (1922) Geschlechtsbestimmung und Zahlenverhaltnis der Geschlechter beim Sauerampfer (Rumex acetosa). “Biol. Zbl.” 42, 465-480.
- Rychlewski J., Kazlmierez Z. (1975) Sex ratio in seeds of Rumex acetosa L. as a result of sparse or abundant pollination. “Acta Biol. Cracov” Scr. Bot., 18, 101-114.
- Correns C. (1928) Bestimmung, Vererbung und Verteilung des Geschlechter bei den hoheren Pflanzen. Handb. Vererbungswiss., 2, 1-138.
- Mulcahy D. L. (1967) Optimal sex ratio in Silene alba. “Heredity” 22 № 3, 41.
- Riede W. (1925) Beitrage zum Geschlechts- und Anpassungs-problem. “Flora” 18/19
- Kihara H., Hirayoshi J. (1932) Die Geschlechtschromosomen von Humulus japonicus. Sieb. et. Zuce. In: 8th Congr. Jap. Ass. Adv. Sci., p. 363—367 (cit.: Plant Breeding Abstr., 1934, 5, № 3, p. 248, ref. № 768).
- Geodakyan, V.A. & Geodakyan, S.V., (1985) Is there a negative feedback in sex determination? “Zurnal obschej biol.” 46 201-216 (in Russian). ).
- Ter-Avanesyan D. V. (1949). Tr. Prikl. Bot, Genet, Selekt., 28 119.
- Ter-Avanesian D. V. (1978) Significance of pollen amount for fertilization. “Bull. Torrey Bot. Club.” 105 N 1, 2–8.
Pautan luar [sunting]
- Pollen and Spore Identification Literature
- Paleobotany and Palynology at the Florida Museum of Natural History
- Palynology at the University of Arizona
- Palynology at French Institute of Pondicherry
- Palynology at the University of Sheffield
- SEM coloured and confocal microscope images of pollen
- Pollen Biology and Pollen Allergy (Radboud University Nijmegen)
- Palynology in Utrecht, the Netherlands
- The flight of the pollen cloud Outcrossing from transgenic maize and quantifying outcrossing rates
- The Evolutionary Theory of Sex: Mechanisms of Regulation of the Population Parameters
- PalDat (database comprising palynological data from a variety of plant families)
- Very large pollen accumulation in the sea
- Pollen movement dynamics Co-extra research project on coexistence and traceability of GM an non-GM supply chains