Somatic embryogenesis

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Somatic embryos are mainly produced in vitro and for laboratory purposes, using either solid or liquid nutrient media which contain plant growth regulators (PGRs). The main PGRs used are auxins but can contain cytokinin in a smaller amount.[1] Somatic embryogenesis is a process where a plant or embryo is derived from a single somatic cell or group of somatic cells. This is in contrast to zygotic embryogenesis, where ,in diploid species, two haploid cells combine to form one diploid cell. Somatic embryos are produced when somatic cells are restructured through a series of morphological and biochemical changes in the embryogenic pathway.[2] Development of somatic embryos is not that different from zygotic embryos.
[3]

Three examples of somatic embryogenesis from nature are ovules in Paeonia and on the leaves

of Aspleniumand Kalanchoe. Shoots and roots are monopolar while somatic embryos are bipolar, allowing them to form a whole plant without culturing on multiple media types. Somatic embryogenesis has served as a model to understand the physiological and biochemical events that occur plant developmental processes as well as a component to biotechnological advancement.[4] The first documentation of somatic embryogenesis was by Steward et al in 1958 and Reinert in 1959 with carrot cell suspension cultures.[5][6]

Switchgrass somatic embryos

Contents
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1 Direct and indirect embryogenesis 2 Plant regeneration via somatic embryogenesis 3 Factors influencing somatic embryogenesis 4 Uses of somatic embryogenesis 5 Problems associated with somatic embryogenesis 6 Tracking and Fate Maps

7 Angiosperms 8 Gymnosperms 9 See Also 10 References 11 External links

[edit]Direct

and indirect embryogenesis

Somatic embryogenesis has been described to occur in two ways: directly or indirectly[7] Direct embryogenesis occurs when embryos are started directly from explant tissue creating an identical clone while indirectly occurs from unorganized tissue (callus).

[edit]Plant

regeneration via somatic embryogenesis

Plant regeneration via somatic embryogenesis occurs in five steps: initiation of embryogenic cultures, proliferation of embryogenic cultures, prematuration of somatic embryos, maturation of somatic embryos and plant development on nonspecific media. Initiation and proliferation occur on a medium rich in auxin, which induces differentiation of localized meristematic cells. The auxin typically used is 2,4-D. Once transferred to a medium with low or no auxin, these cells can then develop into mature embryos. Germination of the somatic embryo can only occur when it is mature enough to have functional root and shoot apices.[8]

[edit]Factors

influencing somatic embryogenesis

Factors and mechanisms controlling cell differentiation in somatic embryos are relatively ambiguous. Certain compounds excreted by plant tissue cultures and found in culture media have been shown necessary to coordinate cell division and morphological changes.[9] These compounds have been identified by Chung et al.[10] as various polysaccharides, amino acids, growth regulators, vitamins, low molecular weight compounds and polypeptides. Several signaling molecules known to influence or control the formation of somatic embryos have been found and include extracellular proteins, arabinogalactan proteins and lipochitooligosaccharides. Temperature and lighting can also affect the maturation of the somatic embryo.

[edit]Uses

of somatic embryogenesis

Plant transformations Mass propagation[11]

[edit]Problems

associated with somatic embryogenesis

High chance of mutations Difficult method

Loss of regenerative ability High percentage of albino shoots during regeneration Not possible with all plant species and must be optimized for each species and its use

[edit]Tracking

and Fate Maps

Understanding the formation of a somatic embryo through establishment of morphological and molecular markers is important for construction of a fate map. The fate map is the foundation in which to build further research and experimentation. Two methods exist to construct a fate map: synchronous celldivision and time-lapse tracking. The latter typically works more consistently because of cell-cyclealtering chemicals and centrifuging involved in synchronous cell-division.[2]

[edit]Angiosperms
Embryo development in angiosperms is divided into several steps. The zygote is divided asymmetrically forming a small apical cell and large basal cell. The organizational pattern is formed in the globular stage and the embryo then transitions to the cotyledonary stage.[12]Embryo development differs in monocots and dicots. Dicots pass through the globular, heart-shaped, and torpedo stages while monocots pass through globular, scuetellar, and coleoptilar stages.[13] Many culture systems induce and maintain somatic embryogenesis by continuous exposure to 2,4dichlorophenoxyacetic acid. Abscisic acid has been reported to induce somatic embryogenesis in seedlings. After callus formation, culturing on a low auxin or hormone free media will promote somatic embryo growth and root formation. In monocots, embryogenic capability is usually restricted to tissues with embryogenic or meristematic origin. Somatic cells of monocots differentiate quickly and then lose mitotic and morphogenic capability. Differences of auxin sensitivity in embryogenic callus growth between different genotypes of the same species show how variable auxin responses can be.[14] Carrot Daucus carota was the first and most understood species with regard to developmental pathways and molecular mechanisms.[2] Time-lapse tracking by Toonen et al (1994) showed that morphology of competent cells can vary based on shape and cytoplasm density. Five types of cells were identified from embryonic suspension: spherical cytoplasm-rich, spherical vacuolated, oval vacuolated, elongated vacuolated, and irregular shaped cells. Each type of cell multiplied with certain geometric symmetry. They developed into symmetrical, asymmetrical, and aberrantly-shaped cell clusters that eventually formed embryos at different frequencies.[15]This indicates that organized growth polarity do not always exist in somatic embryogenesis.[2]

[edit]Gymnosperms
Embryo development in gymnosperms occurs in three phases. Proembryogeny includes all stages prior to suspensor elongation. Early embryogeny includes all stages after suspensor elongation but before root meristem development. Late embryogeny includes development of root and shoot meristems.[12] Time-lapse tracking in Norway Spruce Picea abies revealed that neither single cytoplasmic-rich cells nor vacuolated

cells developed into embryos. Proembryogenic masses (PEMs), an intermediate between unorganized cells and an embryo composed of cytoplasmic-rich cells next to a vacuolated cell, are stimulated with auxinand cytokinin. Gradual removal of auxin and cytokinin and introduction of ABA[disambiguation needed
]

will allow an embryo to form.[2]Using somatic embryogenesis has been considered for mass production of

vegetatively propagated pine clones and cryopreservation of germplasm. However, the use of this technology for reforestation and breeding of pine trees is in its infancy