Plant embryogenesis
Encyclopedia
Plant embryogenesis is the process that produces a plant embryo from a fertilised 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 root
s.
occurs naturally as a result of sexual fertilization and the formation of the zygotic embryo
s. The embryo along with other cells from the motherplant 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.
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
) 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, cotyledon
s, 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, 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 cotyledon
s 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.
In a dicot embryo, the hypophysis, which is the uppermost cell of the suspensor
, differentiates to form part of the root cap. Plant cells can also be induced to form embryos in plant tissue culture
; these embryos are called somatic embryos, which are used to generate new plants from single 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 gymnosperm
s 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 bud
s that form on stem
s. 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.
s; 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.
Zygote
A zygote , or zygocyte, is the initial cell formed when two gamete cells are joined by means of sexual reproduction. In multicellular organisms, it is the earliest developmental stage of the embryo...
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
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
and root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
s.
Seeds
EmbryogenesisEmbryogenesis
Embryogenesis is the process by which the embryo is formed and develops, until it develops into a fetus.Embryogenesis starts with the fertilization of the ovum by sperm. The fertilized ovum is referred to as a zygote...
occurs naturally as a result of sexual fertilization and the formation of the zygotic embryo
Zygote
A zygote , or zygocyte, is the initial cell formed when two gamete cells are joined by means of sexual reproduction. In multicellular organisms, it is the earliest developmental stage of the embryo...
s. The embryo along with other cells from the motherplant develops into the seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
or the next generation, which, after germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...
, grows into a new plant.
Embryogenesis may be divided up into two phases, the first involves morphogenetic events
Morphogenesis
Morphogenesis , is the biological process that causes an organism to develop its shape...
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
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
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
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...
and seedling growth. Embryogenesis involves cell growth and division, cell differentiation and programmed cellular death
Apoptosis
Apoptosis is the process of programmed cell death that may occur in multicellular organisms. Biochemical events lead to characteristic cell changes and death. These changes include blebbing, cell shrinkage, nuclear fragmentation, chromatin condensation, and chromosomal DNA fragmentation...
. The zygotic embryo is formed following double fertilisation
Fertilisation
Fertilisation is the fusion of gametes to produce a new organism. In animals, the process involves the fusion of an ovum with a sperm, which eventually leads to the development of an embryo...
of the ovule, giving rise to two distinct structures: the plant embryo and the endosperm
Endosperm
Endosperm is the tissue produced inside the seeds of most flowering plants around the time of fertilization. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This makes endosperm an important source of nutrition in human diet...
which together go on to develop into a seed. Seeds may also develop without fertilization, which is referred to as apomixis
Apomixis
In botany, apomixis was defined by Winkler as replacement of the normal sexual reproduction by asexual reproduction, without fertilization. This definition notably does not mention meiosis...
. Plant cells can also be induced to form embryos in plant tissue culture
Plant tissue culture
Plant tissue culture is a collection of techniques used to maintain or grow plant cells, tissues or organs under sterile conditions on a nutrient culture medium of known composition. Plant tissue culture is widely used to produce clones of a plant in a method known as micropropagation...
; such embryos are called somatic embryos.
Following fertilization, the zygote undergoes an asymmetrical cell division
Cell division
Cell division is the process by which a parent cell divides into two or more daughter cells . Cell division is usually a small segment of a larger cell cycle. This type of cell division in eukaryotes is known as mitosis, and leaves the daughter cell capable of dividing again. The corresponding sort...
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
Octant
An octant is one of eight divisions.-Octant in the plane :Traditionally wind direction is given as one of the 8 octants because that is more accurate than merely giving one of the 4 quadrants, and the wind vane typically does not have enough accuracy to bother with more precise indication.-Octant...
) 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, cotyledon
Cotyledon
A cotyledon , is a significant part of the embryo within the seed of a plant. Upon germination, the cotyledon may become the embryonic first leaves of a seedling. The number of cotyledons present is one characteristic used by botanists to classify the flowering plants...
s, hypocotyl
Hypocotyl
The hypocotyl is the stem of a germinating seedling, found below the cotyledons and above the radicle .-Dicots:...
, 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, 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 cotyledon
Cotyledon
A cotyledon , is a significant part of the embryo within the seed of a plant. Upon germination, the cotyledon may become the embryonic first leaves of a seedling. The number of cotyledons present is one characteristic used by botanists to classify the flowering plants...
s and shoot meristem
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
, while the lower layer produces the hypocotyl
Hypocotyl
The hypocotyl is the stem of a germinating seedling, found below the cotyledons and above the radicle .-Dicots:...
and root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
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.
In a dicot embryo, the hypophysis, which is the uppermost cell of the suspensor
Suspensor
-Fungi:Suspensors are filamentous structural formations employed by Fungi in holding a zygospore between two strains of hyphae.-Plants:Suspensors are also found in plant zygotes in angiosperms in connecting the endosperm to an embryo. During embryo development in angiosperm seeds, normal...
, differentiates to form part of the root cap. Plant cells can also be induced to form embryos in plant tissue culture
Plant tissue culture
Plant tissue culture is a collection of techniques used to maintain or grow plant cells, tissues or organs under sterile conditions on a nutrient culture medium of known composition. Plant tissue culture is widely used to produce clones of a plant in a method known as micropropagation...
; these embryos are called somatic embryos, which are used to generate new plants from single cells.
Plant growth and buds
Embryonic tissue is made up of actively growing cellsCell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....
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
Sporophyte
All land plants, and some algae, have life cycles in which a haploid gametophyte generation alternates with a diploid sporophyte, the generation of a plant or algae that has a double set of chromosomes. A multicellular sporophyte generation or phase is present in the life cycle of all land plants...
and gametophyte
Gametophyte
A gametophyte is the haploid, multicellular phase of plants and algae that undergo alternation of generations, with each of its cells containing only a single set of chromosomes....
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 gymnosperm
Gymnosperm
The gymnosperms are a group of seed-bearing plants that includes conifers, cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek word gymnospermos , meaning "naked seeds", after the unenclosed condition of their seeds...
s 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
Apomixis
In botany, apomixis was defined by Winkler as replacement of the normal sexual reproduction by asexual reproduction, without fertilization. This definition notably does not mention meiosis...
) and becomes a plant embryo.
This embryonic condition also occurs in the bud
Bud
In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of the stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be specialized to develop flowers or short shoots, or may have...
s that form on stem
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
s. 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 embryogenesis
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 protoplastProtoplast
Protoplast, from the ancient Greek πρῶτον + verb πλάθω or πλάττω , initially referred to the first organized body of a species.Protoplast has several biological definitions:...
s; development of synthetic seed technology. Cells derived from competent source tissue are cultured to form an undifferentiated mass of cells called a callus
Callus (cell biology)
Plant callus is a mass of undifferentiated cells derived from plant tissue for use in biological research and biotechnology. In plant biology, callus cells are those cells that cover a plant wound. To induce callus formation, plant tissues are surface sterilized and then plated onto in vitro...
. 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.