Layers from outside to inside in angiosperm ovule:
C — Integuments (outermost protective layer → becomes seed coat)
A — Parietal cell layer
B — Middle layer
D — Nucellus (innermost — contains embryo sac with female gamete)
Sequence: C → A → B → D
The ovule is the structure inside the ovary that develops into a seed after fertilisation. Structure from outside to inside: Funiculus: stalk attaching ovule to placenta of ovary wall. Hilum: point of attachment. Integuments: protective layers. Most angiosperms = bitegmic (two integuments: outer and inner). Some = unitegmic (one integument — Asteraceae, Solanaceae). Micropyle: opening at the tip where pollen tube enters. Chalaza: base of ovule, opposite micropyle. Nucellus: parenchymatous nutritive tissue enclosed by integuments. Homologous to megasporangium. Embryo sac: female gametophyte develops within nucellus. Parietal cell layer and middle layer: nucellar tissue layers around the embryo sac. After fertilisation: integuments → seed coat (outer integument → testa, inner integument → tegmen). Nucellus → perisperm (in some seeds). Embryo sac → embryo + endosperm.
Megasporogenesis: formation of haploid megaspores from diploid megaspore mother cell (MMC) in the nucellus. Process: A single cell in the nucellus differentiates as the MMC (megaspore mother cell). MMC undergoes meiosis → 4 megaspores (megaspore tetrad). In most angiosperms (Polygonum type): 3 of 4 megaspores degenerate. Only 1 functional megaspore (chalazal megaspore — nearest the chalaza) survives. This functional megaspore → female gametophyte (embryo sac) by mitosis. Female gametophyte development (megagametogenesis): functional megaspore → 3 mitotic divisions without cell wall formation → 8-nucleate stage → cell organisation → 7-cell, 8-nucleus embryo sac. Stages of mitosis: 1 nucleus → 2 → 4 → 8. Then cytokinesis: 3 antipodal cells at chalazal end, 2 synergids + 1 egg cell at micropylar end, 1 central cell with 2 polar nuclei.
Standard (Polygonum-type) embryo sac has 7 cells and 8 nuclei. Egg apparatus (micropylar end): 1 egg cell (n) — female gamete. 2 synergids (n) — guide pollen tube, filiform apparatus at micropylar end for pollen tube reception. Antipodal cells (chalazal end): 3 cells (n) — degenerate after fertilisation, nutritive role. Central cell: 1 large cell (2n — contains 2 polar nuclei n+n). After double fertilisation: egg + sperm 1 → zygote (2n) → embryo. Polar nuclei + sperm 2 → primary endosperm nucleus (3n) → endosperm. Synergids degenerate after pollen tube penetration. Antipodals degenerate. Total nuclei count: 1 (egg) + 2 (synergids) + 3 (antipodals) + 2 (polar nuclei) = 8 nuclei.
Double fertilisation is unique to angiosperms (discovered by Nawaschin, 1898). Two sperm cells from one pollen grain: Syngamy: sperm 1 (n) + egg (n) → zygote (2n) → embryo. Triple fusion: sperm 2 (n) + 2 polar nuclei (n+n=2n) → primary endosperm nucleus (3n) → endosperm. Both events occur simultaneously. Significance: ensures endosperm forms only when fertilisation occurs → resources invested only in seeds that will develop. Triploid endosperm provides nutrition for developing embryo. Pollen tube entry: through micropyle (porogamy — most common), or through chalaza (chalazogamy), or through integuments. Pollen tube enters synergid → releases 2 sperm → one goes to egg, one to central cell.
After double fertilisation: zygote (2n) undergoes mitotic divisions → embryo development. Primary endosperm nucleus (3n) → endosperm development. Integuments → seed coat. Embryo development stages: Proembryo → globular embryo → heart embryo → torpedo embryo → mature embryo. In dicots (Capsella bursa-pastoris studied): Globular: spherical mass, no differentiation. Heart: two cotyledon primordia visible. Torpedo: elongated, cotyledons and axis differentiated. Mature embryo: radicle, plumule, cotyledons, hypocotyl, epicotyl. Endosperm: Cellular type: cell walls form after each nuclear division (Gossypium/cotton). Nuclear type: free nuclear divisions first, then cell walls form (Coconut milk = liquid nuclear endosperm). Helobial type: intermediate (in monocots). Seeds: albuminous (endosperm persists) or non-albuminous (endosperm absorbed by embryo, stored in cotyledons).
Placentation = arrangement of ovules on the placenta inside the ovary. Marginal: placenta along ventral suture of monocarpellary ovary. Peas and beans (Fabaceae). Axile: placenta at central axis of multilocular ovary. Tomato, lemon, orange (Solanaceae, Rutaceae). Parietal: placenta on ovary wall (wall) in unilocular ovary. Mustard, cucurbits (Brassicaceae, Cucurbitaceae). Free central: ovules on central column with no septa. Primrose, Dianthus (Primulaceae, Caryophyllaceae). Basal: single ovule at base of ovary. Sunflower, marigold (Asteraceae). Superficial: ovules attached to inner surface all around (Water lily, Nymphaea). Each type corresponds to different ovary structure and is used in plant classification.
After fertilisation: ovary → fruit (pericarp = wall of fruit). Ovules → seeds. True fruit: develops from ovary alone. False fruit (pseudocarp): thalamus or calyx also contributes. Apple, pear: thalamus (receptacle) forms fleshy edible part — ovary forms the core. Types of true fruits: Drupe: fleshy with stony endocarp (mango, peach, cherry, coconut). Berry: fleshy throughout, many seeds (tomato, banana, grapes). Capsule: dry, dehiscent, multiple seeds (poppy, cotton). Legume/pod: Fabaceae family (pea, bean). Achene: dry, single seed, thin pericarp (sunflower). Nut: dry, single seed, hard pericarp (walnut, chestnut). Caryopsis: grain (seed coat fused with pericarp — wheat, rice, maize). Parthenocarpic fruits: develop without fertilisation → seedless (banana, some grapes, cucumber varieties).
Polyembryony: more than one embryo in a seed. Causes: fertilisation of both synergids (rare), adventive embryony (sporophytic tissue of nucellus/integuments → extra embryos), cleavage polyembryony (zygote or proembryo splits). Examples: Citrus (orange, lemon): regular nucellar polyembryony — nucellar cells develop into embryos alongside the zygotic embryo. Onion (Allium), groundnut (Arachis) also show polyembryony. In Citrus: nucellar embryos are genetically identical to mother plant (same as vegetative propagation). Apomixis: seed formation without fertilisation. Types: Diplospory: unreduced egg cell (2n) develops into embryo. Apospory: somatic cells of nucellus form 2n embryo sac. Adventive embryony: nucellar/integument cells directly → embryo (Citrus). Importance: apomictic seeds produce plants genetically identical to mother → maintain hybrid vigour indefinitely → commercially valuable in hybrid crops.