Развитие зародыша растений после оплодотворения

Process that occurs after the fertilization of an ovule to produce a fully developed plant embryo
Plant embryonic development, also plant embryogenesis, is a process that occurs after the fertilization of an ovule to produce a fully developed plant embryo. This is a pertinent stage in the plant life cycle that is followed by dormancy and germination. The zygote produced after fertilization must undergo various cellular divisions and differentiations to become a mature embryo. However, both plants and animals including humans, pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.

Embryogenesis occurs naturally as a result of single, or double fertilization, of the ovule, giving rise to two distinct structures: the plant embryo and the endosperm which go on to develop into a seed. The zygote goes through various cellular differentiations and divisions in order to produce a mature embryo. These morphogenic events 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 following morphogenic events are only particular to eudicots, and not monocots.

Following fertilization, the zygote and endosperm are present within the ovule, as seen in stage I of the illustration on this page. Then the zygote undergoes an asymmetric transverse cell division that gives rise to two cells a small apical cell resting above a large basal cell. These two cells are very different, and give rise to different structures, establishing polarity in the embryo. apical cellThe small apical cell is on the top and contains most of the cytoplasm, the aqueous substance found within cells, from the original zygote. It gives rise to the hypocotyl, shoot apical meristem, and cotyledons. Stage II, in the illustration above, indicates what the embryo looks like during the eight cell stage. According to Laux et al., there are four distinct domains during the eight cell stage. The first two domains contribute to the embryo proper. The apical embryo domain, gives rise to the shoot apical meristem and cotyledons. The second domain, the central embryo domain, gives rise to the hypocotyl, root apical meristem, and parts of the cotyledons. The third domain, the basal embryo domain, contains the hypophysis. The hypophysis will later give rise to the radicle and the root cap. The last domain, the suspensor, is the region at the very bottom, which connects the embryo to the endosperm for nutritional purposes.

Additional cell divisions occur, which leads to the sixteen cell stage. The four domains are still present, but they are more defined with the presence of more cells. The important aspect of this stage is the introduction of the protoderm, which is meristematic tissue that will give rise to the epidermis. However, in the torpedo stage of development, parts of the suspensor complex must be terminated. The appearance of a mature embryo is seen in Stage VI, in the illustration above.

The end of embryogenesis is defined by an arrested development phase, or stop in growth. This phase usually coincides with a necessary component of growth called dormancy. Dormancy is a period in which a seed cannot germinate, even under optimal environmental conditions, until a specific requirement is met. Breaking dormancy, or finding the specific requirement of the seed, can be rather difficult. For example, a seed coat can be extremely thick. According to Evert and Eichhorn, very thick seed coats must undergo a process called scarification, in order to deteriorate the coating. It also plays an important role in the establishment polarity with the plant embryo. Research has shown that the hypocotyl from both gymnosperms and angiosperms show auxin transport to the root end of the embryo. They hypothesized that the embryonic pattern is regulated by the auxin transport mechanism and the polar positioning of cells within the ovule. The importance of auxin was shown, in their research, when carrot embryos, at different stages, were subjected to auxin transport inhibitors. The inhibitors that these carrots were subjected to made them unable to progress to later stages of embryogenesis. During the globular stage of embryogenesis, the embryos continued spherical expansion. In addition, oblong embryos continued axial growth, without the introduction of cotyledons. During the heart embryo stage of development, there were additional growth axes on hypocotyls. Further auxin transport inhibition research, conducted on Brassica juncea, shows that after germination, the cotyledons were fused and not two separate structures.

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 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.

The process of androgenesis allows a mature plant embryo to form from a reduced, or immature, pollen grain. Androgenesis usually occurs under stressful conditions. 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.