Apomixis Apomixis is the production of an embryo that bypasses the usual process of meiosis and fertilization. The genotype of the embryo and resulting plant will be the same as the seed parent. This is clonal seed production ! Apomixis Apomixis occurs in over 300 species from at least 35 plant families. It is most common in the Asteraceae Graminaceae Rosaceae Rutaceae Goat’s beard (Tragopogon) Apomixis Only a few economically important food crops exhibit apomixis. These include Citrus, Mango (Mangifera), and Mangosteen (Garcinia). Several grass species and cultivars are apomicts including Kentucky blue grass. Orange (Citrus) Apomixis Types of apomixis: Gametophytic Diplospory Apospory Sporphytic Adventive embryony Apple (Malus) Apomixis Gametophytic apomixis Embryo derived from a nonreduced gametophyte cell. Diplospory Apospory Dandelion (Taraxacum) Apomixis Normal sexual development from a reduced gamete. Apomixis Diplospory Megaspore mother cell does not finish or initiate meiosis. Egg sac organizes normally but cells are diploid. Apomixis Diplospory Embryo initiates development without being fertilized. Endosperm develops either with or without fertilization. In some cases (Poa), fertilization is required for endosperm formation. The egg does not get fertilized because it is already growing. Common in grasses and Asteraceae. Eastern gamagrass (Tripsacum dactyloides) Apomixis Apospory The megaspore mother cell undergoes normal meiosis, but the resulting cells degrade before forming an embryo sac. Apomixis Apospory Additional cell(s) in the nucellus become densely cytoplasmic and take on the role of apomictic initials. These cells bypass meiosis to form their own embryo sac within the same ovule. Apomixis Apospory Endosperm must be fertilized (pseudogamy). Aposporous apomicts are found in the Rosaceae, Asteraceae and in some grasses. Garden rose Apomixis Adventive embryony The megaspore mother cell undergoes normal meiosis and forms a normal sexual embryo sac. These cells are fertilized by male sperm cells as in normal embryogenesis. Normal Megaspore Mother Cell Nucellus Adventive embryony Meiosis Egg Sac Apomixis Adventive embryony However, cells in the nucellus become embryogenic and form multiple embryos in the micropylar region of the ovule. Adventive embryony Normal Megaspore Mother Cell Nucellus Meiosis Egg Sac Apomixis Adventive embryony Cells in the nucellus become embryogenic. These develop into somatic embryos. Apomixis Adventive embryony These asexual embryos do not produce their own embryo sac but grow into the embryo sac of the sexual embryo. Endosperm must be fertilized. Common in tropical fruit trees like citrus and mango. Apomixis Adventive embryony Multiple seedlings (polyembryony) from a single seed in Citrus. One of the seedlings will be sexual and the others will be clones. Multiple embryos Single seed Apomixis Adventive embryony Single seed Pachira contains up to 5 seedlings per seed. The large seedling is sexual and the smaller are nucellar derived. Multiple embryos Apomixis Somatic embryogenesis Somatic embryogenesis is also the formation of embryos from vegetative tissue that does not involve gamete fusion. Seed Development Plant Hormones and Seed Development Auxin Cytokinin Gibberellin Ethylene Abscisic acid Seed Development Plant Hormones and Seed Development Growth and differentiation of the embryo. Accumulation of food reserves. Storage for use during germination. Growth and development of fruit. Seed Development Plant Hormones and Seed Development Auxin Abundant in developing seeds. Relatively high levels throughout seed development before declining during maturation drying. Required for normal embryogenesis. High during cell division activity including the endosperm. Seed Development Plant Hormones and Seed Development Auxin This is critical for orienting the eight nuclei into their proper positions. Chalazal Low Auxin gradient Early in embryo sac development an auxin gradient is established from the micropylar to chalazal regions of the embryo sac. High Micropylar Seed Development Plant Hormones and Seed Development Auxin The gradient is from the apical to basal portions of the embryo and is required for the appropriate bipolar (shoot and root) symmetry of the embryo. Auxin gradient An auxin gradient is also important for normal embryo formation. High Low Seed Development Plant Hormones and Seed Development Auxin Auxin from seeds can be the signal for fruit to develop. Seed Development Plant Hormones and Seed Development Cytokinins Abundant in developing seeds. The highest concentration of cytokinins is found during the cell division stages of embryo and endosperm development. Key role is in cell division. Seed Development Plant Hormones and Seed Development Cytokinins The major role for cytokinins during seed development is the control of endosperm development. Cytokinin determines the number of cells in the endosperm and the timing for cellularization. Endosperm Seed Development Plant Hormones and Seed Development Gibberellins Abundant in developing seeds. Various forms high in Stages I and II. GA mutants form normal embryos but seed size is reduced. Seed Development Plant Hormones and Seed Development Gibberellins That gibberellins are important in the development of cotyledons as storage reserve organs. Cotyledons Endosperm Oak (Quercus) Seed Development Parthenocarpy Parthenocarpy is fruit production without seed formation. Both auxins and gibberellins can be important to induce parthenocarpy. Seed Development Parthenocarpy Vegetative parthenocarpy does not require pollination. Examples - fig and pear. Stimulative parthenocarpy requires pollination. Example – grapes. Seed Development Plant Hormones and Seed Development Abscisic Acid Abundant in developing seeds. ABA high in Stage II. Facilitates storage reserve accumulation. Prevents precocious germination. Seed Development Plant Hormones and Seed Development Abscisic Acid ABA induces expression of genes for storage proteins. Redrawn from Finkelstein and Crouch, 1987 Seed Development Plant Hormones and Seed Development Abscisic Acid Seed Development Precocious germination (Vivipary) Seeds by-pass maturation drying and there is premature germination while seeds are still on the mother plant. Tomato Seed Development Precocious germination (Vivipary) Premature germination is generally undesirable. Genetic mutation in response to ABA. Rhipsalis Seed Development Precocious germination (Vivipary) For a few species, vivipary is the normal type of seedling development. Mangrove species are important plants that provide stabilization and habitat along the edge of waterways. Mangrove (Rhizophora) Seed Development Precocious germination (Vivipary) In mangrove, the seed germinates while still on the plant. Black Mangrove (Avicennia gerinans) Mangrove (Rhizophora mangle) Seed Development Precocious germination (Vivipary) Seed Development Plant Hormones and Seed Development Ethylene Ethylene is produced during seed development. Ethylene mutants produce normal seeds. Seed Development Plant Hormones and Seed Development Ethylene Appears to be important in de-greening prior to maturation drying. Seed Development Plant Hormones and Seed Development There is an interesting interaction between ethylene, gibberellin and ABA in controlling programmed cell death in endosperm in cereals. Endosperm Corn Seed Development Plant Hormones and Seed Development There must be a mechanism in place that programs endosperm cells to die, while adjacent aleurone and embryo cells continue the maturation process. Aleurone Endosperm Wheat
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