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Plant embryogenesis is the process that produces a plant embryo from a fertilized ovule by asymmetric cell division and the differentiation of undifferentiated cells into tissues and organs. It occurs during seed development, when the single-celled zygote undergoes a programmed pattern of cell division resulting in a mature embryo. A similar process continues during the plant's life within the meristems of the stems and roots.
Embryogenesis occurs naturally as a result of sexual fertilization and the formation of the zygotic embryos. The embryo along with other cells from the mother plant develops into the seed or the next generation, which, after germination, grows into a new plant.
Embryogenesis may be divided up into two phases, the first involves morphogenetic events which form the basic cellular pattern for the development of the shoot-root body and the primary tissue layers; it also programs the regions of meristematic tissue formation. The second phase, or postembryonic development, involves the maturation of cells, which involves cell growth and the storage of macromolecules (such as oils, starches and proteins) required as a 'food and energy supply' during germination and seedling growth. Embryogenesis involves cell growth and division, cell differentiation and programmed cellular death. The zygotic embryo is formed following double fertilisation of the ovule, giving rise to two distinct structures: the plant embryo and the endosperm which together go on to develop into a seed. Seeds may also develop without fertilization, which is referred to as apomixis. Plant cells can also be induced to form embryos in plant tissue culture; such embryos are called somatic embryos and can be used to generate new plants from single cells.
Following fertilization, the zygote undergoes an asymmetrical cell division that gives rise to a small apical cell, which becomes the embryo and a large basal cell (called the suspensor), which functions to provide nutrients from the endosperm to the growing embryo. From the eight cell stage (octant stage) onwards, the zygotic embryo shows clear embryo patterning, which forms the main axis of polarity, and the linear formation of future structures. These structures include the shoot meristem, cotyledons, hypocotyl, and the root and root meristem: they arise from specific groups of cells as the young embryo divides and their formation has been shown to be position-dependent.
In the globular stage[clarification needed], the embryo develops radial patterning through a series of cell divisions, with the outer layer of cells differentiating into the 'protoderm.' The globular embryo can be thought of as two layers of inner cells with distinct developmental fates; the apical layer will go on to produce cotyledons and shoot meristem, while the lower layer produces the hypocotyl and root meristem. Bilateral symmetry is apparent from the heart stage; provascular cells will also differentiate at this stage. In the subsequent torpedo and cotyledonary stages of embryogenesis, the embryo completes its growth by elongating and enlarging.[clarification needed]
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Plant growth and buds
Embryonic tissue is made up of actively growing cells and the term is normally used to describe the early formation of tissue in the first stages of growth. It can refer to different stages of the sporophyte and gametophyte plant; including the growth of embryos in seedlings, and to meristematic tissues, which are in a persistently embryonic state, to the growth of new buds on stems.
In both gymnosperms and angiosperms, the young plant contained in the seed, begins as a developing egg-cell formed after fertilization (sometimes without fertilization in a process called apomixis) and becomes a plant embryo. This embryonic condition also occurs in the buds that form on stems. The buds have tissue that has differentiated but not grown into complete structures. They can be in a resting state, lying dormant over winter or when conditions are dry, and then commence growth when conditions become suitable. Before they start growing into stem, leaves, or flowers, the buds are said to be in an embryonic state.
Somatic embryos are formed from plant cells that are not normally involved in the development of embryos, i.e. ordinary plant tissue. No endosperm or seed coat is formed around a somatic embryo. Applications of this process include: clonal propagation of genetically uniform plant material; elimination of viruses; provision of source tissue for genetic transformation; generation of whole plants from single cells called protoplasts; development of synthetic seed technology. Cells derived from competent source tissue are cultured to form an undifferentiated mass of cells called a callus. Plant growth regulators in the tissue culture medium can be manipulated to induce callus formation and subsequently changed to induce embryos to form from the callus. The ratio of different plant growth regulators required to induce callus or embryo formation varies with the type of plant. Asymmetrical cell division also seems to be important in the development of somatic embryos, and while failure to form the suspensor cell is lethal to zygotic embryos, it is not lethal for somatic embryos.
Notes and references
- Suárez, María F., and Peter V. Bozhkov. 2008. Plant embryogenesis. Totowa, NJ: Humana Press. p 3.
- Current topics in developmental biology; v. 67. 2005. [S.l.]: Elsevier Academic Press. p 135.
- ZMBP Embryo Patterning
- Pandey, Brahma Prakash. 2005. Textbook of botany angiosperms: taxonomy, anatomy, embryology (including tissue culture) and economic botany. New Delhi: S. Chand & Company. p 410.
- McManus, Michael T., and Bruce E. Veit. 2002. Meristematic tissues in plant growth and development. Sheffield: Sheffield Academic Press.
- Singh, Gurcharan. 2004. Plant systematics: an integrated approach. Enfield, NH: Science Publishers. p 61.
- http://www.accessexcellence.org/LC/ST/st2bgplant.html Plant Tissue Culture
- http://www.umanitoba.ca/afs/plant_science/COURSES/39_768/l15/l15.1.html A site on the developing plant embryo from a molecular genetics perspective