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XII Reproduction in Angiosperms Reproduction is the process through which species can propagate by increase in their number. This propagation or continuity of species is essential both in time and space. Reproduction is the means of perpetuation of species. In plants new individuals may be produced from single parent or from two parents depending upon the type of reproduction. Modes of Reproduction: The reproduction in angiosperms is mainly of two types: (1) Asexual Reproduction and (2) Sexual Reproduction. 1. Asexual Reproduction: It is the process of reproduction in which the new individuals are formed without the fusion of gametes. 2. Sexual Reproduction: It is the process in which new individuals are formed by fusion of gametes. There is one more common method of reproduction observed in Angiosperms, i.e. vegetative reproduction or vegetative propagation. In this process, regeneration of new plants takes place from portions of vegetative organs of the plants e.g. root, stem, leaf. As the fusion of gametes is not involved, vegetative propagation is also included under asexual type of reproduction. Characteristic features of Asexual reproduction: (a) It is uniparental, as only single parent is involved in this type of reproduction. (b) The unit of propagule is a specialized or unspecialized part of the parent. (c) The propagule is formed from body (i.e. somatic) cell of the parent hence described as somatogenic reproduction. (d) There is no fusion of gametes. (e) The reduction division (i.e. meiosis) is absent. Therefore the cell division is of mitosis type. (f) It is quick method of reproduction. (g) A very large number of individuals can be produced by a parent. Characteristic features of Sexual reproduction: 1. In this method the offsprings are form indirectly through the process of formation and fusion of gametes. It results into formation of diploid oospore which is fusion product. 2. This type of reproduction is generally biparental. But it can also be uniparental. 3. The sexual reproduction occurs through special germinal cell, hence the name, germinal reproduction. 4. These cells develop special structures called sex organs which in turn give rise to gametes. 5. The reduction division (i.e. meiosis) occurs at some stage of life cycle. 6. The gametes which are haploid fuse to form diploid oospore. 7. The oospore in turn produces the offspring. 8. The number of offsprings are produced by sexual reproduction is smaller than the number of young ones produced through asexual reproduction. 9. It is relatively a slow process. 10. The offsprings produced through sexual reproduction show variation hence they are not just replicas. Vegetative Reproduction: The vegetative reproduction in angiosperms can be divided in two categories: (A) Natural Methods and (B) Artificial Methods. (A) Natural Methods: The vegetative reproduction occurs naturally in many flowering plants. The natural methods of plants multiplication in which somatic part of the plant body 1 XII separates and gives rise to the whole plant body. Thus propagule can be any part of the plant body such as root, stem, leaf, bud etc. which develops into new independent plant when given suitable conditions. The detached propagule may be root, stem, leaf or floral bud. 1. Tuberous roots (e.g. Sweet potato): This is the example of adventitious root which is modified for the purpose of storage root. It swells due to storage of starchy food and attains a cylindrical shape, hence the name tuberous root. The fleshy root of sweet potato develops the bud which are known as adventitious buds. Generally, the buds do not develop on roots hence these buds are called adventitious buds. These buds gives rise to new plant. 2. Stem Tuber (e.g. Potato): This is the underground stem structure participating in the function of vegetative propagation. In potato aerial branches grow downwards and come to store food. They become swollen due to stored food and become cylindrical in shape, hence the name tuber. The food –laden swelling has got a corky cover. The depressions on the tubers are known as “eyes” which are the nodes with axillary buds. The ‘eye’ with a small piece of tuber serves as the ‘propagule’ or ‘seed’ from which another potato plant can grow, when placed in soil in suitable environment. 3. Runner (e.g. Cynnodon – Harali): This is a modification of subaerial or creeping stem for vegetative propagation. Cynodon is a small grass member thriving well in damp soil. It develop special narrow green horizontal branches called ‘runner’s. These branches develop at the base of crown of leaves. It travels some distance on the ground, touch the soil and gives rise to adventitious roots at intervals and gives rise to crown of leaves upwards. Thus, a new plant is formed. Breaking of runners helps in vegetative propagation. The other examples are Centella (Bramhi) 4. Leaf (e.g. Bryophyllum – Panphuti): The leaves of this plants show crenate margin. The buds lie in the notches of margin of the leaf. When these leaves fall on to the soil, the buds in the margin develop into new plants. (B) Artificial Methods: These are several artificial methods of vegetative propagation. These are manmade special techniques in which somatic body of a plant is made to develop into new independent plant. These methods are used for propagation of desired verities according to one’s requirements. The best seasons to carry out these techniques are rainy and spring seasons. Accordingly there are various horticulture methods for vegetative propagation. They are (a) Cutting: Many plants like, sugarcane, Croton, Hibiscus rosa-sinensis, rose etc. can be easily grown by means of stem cutting. The cutting can be defined as, ‘the cut pieces of root, stem and leaves’. These are planted in nurseries in natural polarized fashion. For successful cutting it is necessary to induce rooting or development of adventitious roots. Leaf cuttings (e.g. Sansevieria – Snake plant): This plants can be propagated by using leaf cutting. For this purpose the leaves are cur in transverse manner into two or tree parts. These parts are planted in soil in vertical manner. For quick rooting the rooting hormone can be applied to the end which is in the soil. In this technique the formation of adventitious buds is of equal importance. Root cutting: The root cuttings from lemon, tamarind etc. sprout producing roots and shoots, if they are put in the soil. The other examples are Blackberry, Raspberry etc. Here ability to form adventitious roots and adventitious buds are the prerequisites. 2 XII (b) Budding: In vegetative propagation by budding methods, only one bud with a small portion of bark, with or without wood is utilized. The budding can be defined as grafting one bud ‘scion’ on the selected root ‘stock’. The use of budding is generally restricted to young plants and smaller branches of large trees. In this process two pieces of living plant tissue are connected together in such a manner, that they will unite. Subsequently they grow and develop as one plant. The two pieces are in the form of: 1. Root Stock: The root stock is the trunk or root material to which, buds or scions are grafted or budded. The ‘scion’ is a small shoot that is inserted by grafting into a root stock. 2. The second piece is called single budded scion. The root stock used for budding should have the desired characters like vigour, growth habit, resistance to soil borne pests etc. The root stock may be a rooted cutting or seedling which is properly established in soil. The advantages of Budding are: 1. It results into stronger union between scion and stock. 2. As each bud can give rise to a new plant, this potential can be fully utilized in budding. 3. The technique is simple and can be performed easily by the amateur. 4. The budding is quick and efficient method. Thus, many buds can be grafted within short period with higher percentage of success. The important methods of budding are: (1) Shield or ‘T’ budding, Inverted ‘T’ budding (2) Patch budding. Shield budding is used for propagation of many fruit trees and species of many ornamental plants. e.g. Citrus, Pear, Peach and Rose. The patch budding is used for species with thick bark e.g. Rubber tree, Mango, Walnut etc. The proper season is early and late summer. Procedure: Scion is a bud with a small piece of bark and cambium. The stock is given ‘T’ shaped cut. The bark is lifted to expose cambium. The bud is then inserted and the bark is allowed to come back to its normal position. The bud is then exposed. The joint is then treated with grafting wax and wrapped with bandage. The bud sprouts after 3-5 weeks. By this time the organic connection as been developed. Leaves and buds of stock are removed. Finally stock is cut above the bud graft. (c) Grafting: It is practiced in plants which do not root easily. This is also applicable for the plants processing a weak root system. Grafting can be defined as, “The art of joining parts of two plants of the same species (intraspecific) or between allied species (i.e. interspecific) in such a manner that brings about fusion or organic union of tissues.” The main part is stock. The scion is the portion which is to be grafted. The scion is selected from plant having desirable or superior characters. The grafting is commonly practiced in tree members like mango, apple, pear, Citrus, guava and rubber plant. Types of grafting: There are different types of grafting in practise. They are as follows: (a) bud grafting (b) approach grafting (c) tongue grafting (d) wedge grafting (e) crown grafting. These methods are based upon methods of uniting two parts. Thus, in a typical grafting experiment, the part of the graft which gives rise to upper part is scion and lower part is called stock. The graft or scion should have one to several buds. The root system of the other plant is allowed to remain intact and called stock (rootstock, understock). The shoot of the stock is often cut 10-30 cms above the base of root. The leaves and buds present over the stock are removed. In tongue (Slice or whip) grafting, oblique sloping cut or notch is given to both stock and scion. The two perfectly fit upon one another. They are tied together. The stock and scion should be of same diameter. In grafting, scion is fixed over the stock in such a manner that the cambia of the two come in contact. The union is covered with the grafting wax. It is the, tied with the help of 3 XII bandage, rubber or nail and wrapped with polythene tape. The buds of the stock are not allowed to sprout. They should be removed as soon as noticed. After few weeks (about 4 to 6) the cut surfaces unite and graft is successful. The strings or bandage and tape is then removed to avoid girdling (i.e. destruction of an area of bark around the periphery of stem). Now root system and shoot system of two different plants connect in such a way that they unite and later develop as a composite plant. Significance of Vegetative Reproduction: Merits: 1. The vegetative reproduction is useful for propagating plants with genetic sterility. e.g. Banana, Pine apple, Orange, Rose etc. 2. It brings about cultivation and perpetuation of seedless varieties of grapes. 3. It maintains useful characters in heterozygous plants. 4. It is convenient and easy method for propagation e.g. Mango Citrus. 5. It induces disease resistance e.g. fruits in Citrus. 6. Cultivation under unfavourable conditions. e.g. Avocado, Apple, Peach, Almond varieties. 7. Change in growth habit can be achieved by using specific root stock e.g. Applethe same cultivar (cultivated variety) dwarf, semidwarf and tall plants have been obtained. 8. Early flowering and fruiting can be achieved with the help of grafted variety as compared to the same varieties propagated by seeds e.g. Mango trees. 9. It helps in pollination and to overcome self compatibility in case of orchard plants. 10. It helps in production of ‘clones’ of useful cultivar. Demerits: 1. Vegetative reproduction cannot produce new variety. 2. It is expensive method as compared to seed multiplication. 3. It requires special trained staff for carrying out method like grafting, budding etc. 4. The plants raised through vegetative propagation have short life span as compared to those raised by seed cultivation. 5. The plants raised through vegetative reproduction show less anchorage and there is possibility of uprooting during storm as these plants develop adventitious roots whereas the seed cultivated varieties have tap root systems. Sexual Reproduction: It is a method of reproduction in which new individuals or offsprings are formed through the formation and fusion of haploid gametes. The fusion results in production of diploid zygote that grows to produce a new individual. In flowering plants, reproductive structures are flowers. The typical flower essentially consists of four different whorls present on knob shaped structure called receptacle. These whorls are known as floral whorls, the details of which we have studied in XI standard. 4 XII These whorls are, calyx, corolla, androecium and gynaecium. The four whorls are divided into two groups: (i) Non-essential whorls (e.g. calyx and corolla) and (ii) essential or reproductive whorls (e.g. androecium and gynaecium). These whorls are directly involved in the function of reproduction. Development of male gametophyte: Structure of Male gametophyte: The male reproductive whorl is called androecium. It consists of stamens. The number of stamens varies in different plants from one to many. The typical stamen is further differentiated into filament, connective and anthers. The filament is the stalk like part of the stamen. It raises the anthers to a proper height. The anthers are sac like lobes generally two in number. They are attached to the filament. The connective is the narrow strip of tissue which joins the anthers. Thus, in transverse section each anther shows two chambers. Thus, we get tetralocular structure known as, microsporangium. T. S. of anther shows presence of anther sac wall which encloses within it the sporogenous tissue, in which diploid microscope or pollen mother cells are differentiated. The pollen grains or microspores are formed from microspore mother cell after undergoing meiotic division. At maturity anther dehisces to liberate pollen grains. Thus, pollen grains are unicellular, uninucleate haploid in structure. It is the first cell of male gametophyte. The pollen grains is covered by two coats, the outer is called exine and inner is called intine. The exine is thick with smooth or spiny outer surface and the intine is thin and smooth. The exine shows presence of one or more germ pores which are meant for the exit of pollen tube. The microspore enlarges and undergoes a mitotic division to form a large vegetative cell and small generative cell. The structure representing the above male gametophyte is known as, pollen grain. The pollen grain is generally shed at this two celled stage. The pollen grains are then transferred to the stigma of proper flower. They are held by sticky secretion. The pollen grains germinates on the surface of stigma and finds a suitable surface. The pollen grains germinates to form pollen tube. The growth of the pollen tube is stimulated by surgery secretions of the stigma. The pollen tube is formed as cytoplasmic outgrowth called germ tube and continues to grow into pollen tube. The vegetative nucleus moves to the tip of the tube. It is followed by the generative nucleus. The pollen tube transverses through stigma and style and reaches the ovary. The generative nucleus of pollen tube divides mitotically to produce two male nuclei or male gametes. This completes the development of male gametophyte. Structure of Female Gametophyte: The gynaceium is the inner most whorl of the flower and consists of carpels. The carpel is comparable with megasporophyll which has folded along the midrib and the two margins unite. The midrib portion of carpel is known as, dorsal suture and fused margins correspond to ventral suture. The placenta bears ovules. The typical arpel consists of ovary, style and stigma. The ovary is basal, swollen part of the carpel and contains more or one ovules. The style is present at the top of the ovary. It bears stigma at its tip. The style raises the stigma to proper heights which facilitates the pollination. The ovary after pollination and fertilization develops into fruit and ovules are transformed into seeds. Structure of Ovule: The typical ovule is the megasporangium with its protective coats, the integuments, and is known as ovule. The typical ovule is attached to placenta on the inner wall of ovary. The ovule is attached to the ovary wall by a stalk called funiculus. In anatropous ovule the micropylecomes to lie close to the funiculus due to the unilateral growth of the ovule. It is also known as inverted ovule. This is the most common type of ovule in angiosperms and hence it is studied in detail. Structure of Anatropous Ovule: The typical anatropous ovule in vertical section shows following parts. (i) Nucellus: It is the main central part of the ovule made up of diploid parenchymatous tissue. 5 XII (ii) Integuments: The nucellus is surrounded by one or two wall layers called integuments. The ovule with one integument is termed as unitegmic while with two integuments is termed bitegmic. (iii) Micropyle: The integuments do not cover the ovule completely, but leave a small space, called micropyle. In the anatropous ovule, micropyle lies near the funicle. It helps in the entry of pollen tube. (iv) Funicle: It is the stalk of the ovule by which the ovule is attached with the placenta. (v) Hilium: The point of attachment of the body of the ovule to its stalk i.e. funicle is called hilum. (vi) Raphe: In anatropous ovule, the funicle continues beyond the hilum alongside the body of the ovule forming a sort of ridge, called raphe. It carries the food to the nucellus. (vii) Chalaza: The distal end of the funicle which is the junction of the integuments and nucellus is called the chalaza. (viii) Embryo sac: It is developed in the nucellus. It is also called as female gametophyte, where the embryo is formed after fertilization. Development of female gametophyte: The diploid megaspore mother cell becomes differentiated in the nucellus. It is slightly larger than the other cells of nucellus. It undergoes reduction division and produces a row of 4 haploid megaspores. Out of 4, upper three degenerate and only one megaspore becomes functional. It soon enlarges in size and forms embryo sac or female gametophyte. The nucleus in the megaspores undergoes three mitotic divisions to form 8 nuclei. These get arranged in two groups of 4 nuclei each, towards micropylar end and chalazal end. The 3 nuclei at the micropylar end form egg apparatus containing as egg cell and two synergid cells. Similarly three nuclei at calazal end form 3 antipodal cells. One nucleus from each pole (i.e. micropylar end and calazal end) get arranged in the centre to form polar nuclei. They fused to form secondary nucleus. In the straight ovule (i.e. orthotropous ovule) the hilum, chalaza and micropyle lie in a straight line. Pollination Types and Agencies: Pollination: Definition: It is the process of transfer of pollen grains from anthers of a flower to the stigma of the same or a different flower and is called pollination. The pollen grains may be transferred to the stigma of a flower present on the same plant or other plant of the same species. The pollen grains are nor motile. As such they cannot move themselves to the receptive stigma of the same or different flower. The pollination helps the transfer of pollen grains to the stigma. Thus, pollination forms as integral part of sexual reproduction in Angiosperms. Types of pollination: The pollination is of two types: (i) Self pollination and (ii) Cross pollination. 6 XII (i) Self pollination: In this type of pollination the pollen grains from the anther are transferred to the stigma of same flower or another flower present on the same plant e.g. sugarcane, Commelina. Autogamy is the process in which pollen grains are transferred to the stigma of same flower. Geitonogamy is the process in which pollen grains are transferred to stigma of different flower on the same plant. (ii) Cross pollination: In this type of pollination the pollen grains are transferred from anthers to the stigma of the other flower belonging to the same species. Xenogamy is the process in which pollen grains are transferred to the flower on separate plant of the same species. Geitonogamy is the process of cross pollination taking place in between two different flowers on the same plant. Pollinating agencies: The pollen grains are non-motile, therefore, cross pollination involves some agencies to carry the pollen grains from the anther to the stigma. Such agencies which carry the pollen grains are called pollinating agents. Types of pollinating agents: The pollinating agents are of two types: (A) Biotic and (B) Abiotic. (A) Biotic Pollinating agents: These are biotic components in the surrounding atmosphere and are therefore in the form of different living systems, like animals, insects etc. As per the nature of pollinating agents, the different terms are used e.g., 1. Zoology → flowers are pollinated by animals 2. Entomophily → flowers are pollinated by insects 3. Ornithophily → flowers are pollinated by birds 4. Chiropteriphily → flowers are pollinated by bats 5. Malacophily → flowers are pollinated by snails. (a) Ornithophily: When the flower is pollinated by bird, the phenomenon is known as ornithophily. In some part of the tropics, birds are important pollinators as compared to insects. The common birds participating in the pollination are Hummingbirds, sun-birds, and honey eaters. The ornithophilous flowers are: (1) Generally tubular (Nicotiana gluca), cup shaped [Callistemon (Bottle brush)]. (2) They are bright coloured. (3) They produce large quantities of pollen and nectar. (4) Their stamens and carpels (i.e. pistils) produce beyond the perianth lobes. (5) The red, orange, yellow colours attract-birds from long distances e.g. Red silk cotton tree, Coral tree, bottle brush etc. In Sterlitzia reginae pollination is brought about by sun-birds. The other common examples are, Bignonia, Sanchezia etc. (b) Entomophily: When the flower is pollinated by insects, the phenomenon is known as entomophily. Insects are chief pollinators and show various types of intimate relationship with flowers they visit. The insects which help in pollination are bees, flies, wasps, moths, beetles. The bees flies and beetles are diurnal as they are active in daytime whereas moths are nocturnal i.e. they are active during night hours. The bees are the main flower visitors and bring about the pollination to the tune of 80 per cent of all pollination done by insects. The insect pollinated flowers show the following characters: (1) The flowers are brightly coloured. (2) They possess fragrance and/or produce nectar. (3) The bees are fond of yellow, violet and purple coloured flowers. (4) The bees while collecting their food in the form of nectar and pollen, visit the flowers and bring about pollination. (5) Some flowers emit unpleasant smell e.g. Arum, Rafflesia etc. Some flies get attracted towards this smell and further bring about the pollination. (6) The orchid, Ophyrus speculum is pollinated by hairy wasp. This orchid flower exactly resembles female counterparts. Also these flowers possess an appearance and 7 XII odour similar to those possessed by the female wasp. Thus, males mistakes the Ophyrus flowers for their counterpart, land upon them to perform the act of pseudo-copulation. They repeat his pollen grains are spiny while the stigma is sticky. (8) Moths generally visit flowers at night. e.g. Salvia Helianthus etc. (9) The typical lever mechanism is seen in Salvia. (B) Abiotic Pollinating Agents: These are the non living agencies and hence passive in nature. The abiotic pollinating agents are of two types: (i) Wind and (ii) Water. (c) Anemophily: When the flower is pollinated by wind the phenomenon is known as ‘anemophily’. The common examples are Grasses, Maize, Jowar etc. In anemophily the air currents pick up the pollen grains from the air currents. The characters of anemophilous flowers are: (1) They produce large number of pollen grains as there is possibility of wastage of quite a large number of pollen grains due to passive nature of wind. (2) The flowers are small and incospicous. (3) The perianth lobes are reduced or absent. (4) The pollen grains are small, light, smooth and dry. (5) The male flowers generally occupy terminal position by means of which wind can easily blow away the discharged pollen. The fixation of anthers to the filament is versatile. Thus, anthers can freely oscillate in all the directions at the tip of filament, according to wind current. (6) In female flowers which are in axile position in Maize plant are present at lower position which is convenient for catching the pollen grains. (7) Also the stigma is feathery and long. The flowers are unisexual, pollen grains small in size, smooth and dry. The other examples are: coconut palm, Datepalm, Cannabis (Bhang) (d) Hydrophily: When the flower is pollinated by water, the phenomenon is known as hydrophily. The examples of water pollinated flowers are, Vallisneria, Lemna, Ceratophyllum, Zostera etc. The water pollination occurs in the plants which are totally submerged marine plants such as Zostera marina. This type of water pollination phenomenon is called hypohydrophily or hyphydrophily. In Caretophyllum, the male flower contains 30-40 stamens. The anthers at maturity get separated at the base. Then they float to the surface of water. They dehisce and liberate the pollen grains. The released pollen grains sink and germinate. The pollen grains then pollinate the female flowers. The important characteristics of hydrophilous flowers are: (1) The pollen grains in water pollinated plants get themselves adapted to help water pollination. Thus, in Zostera marina pollen grains are exceptionally long and needle like and resemble pollen tube. (2) The specific gravity of pollen grains is nearly same to that of marine water. Therefore, the pollen grains can freely float in water at any depth. (3) The stigma of such plant is very long. When pollen grain comes in contact with stigma it coils around the stigma. Vallisneria plant is submerged and dioecious plant. The flowers are borne under water. The male flowers get detached at maturity from parent plant and float on surface of water. The female flowers develop under 8 XII water. (4) At the time of pollination they are brought to the surface of water with the help of long slender stalk. (5) The female flower ultimately gets arranged in horizontal position. It creates a cup-shaped depression around it due to its weight. The male flower floating on the water surface gets into the depression formed by female flower. The anthers come in contact with exposed stigma to transfer pollen masses to it. (6) The pollen grains are usually without an axine as observed in Vallisneria, Certaphyllum and Zostera. The stalk of the female flower after pollination undergoes spiral twisting and ultimately brings the pollinated female flowers under the surface of water. When the flower is pollinated by water above its surface, the phenomenon of hydrophily is further described as epihydrophily or ephydrophily. Necessity and Significance of Pollination: (1) The act of pollination further paves its way to fertilization. This gurantees the production of seeds and fruits. (2) The products of pollination and fertilization i.e. seeds and fruits, in turn serve as a source of food for animals including human beings. (3) The cross pollination creates a chance of combination of characters of two plants. It further helps in the process of evolution. (4) In the process of hybrid seed production the role of pollination is very significant. Fertilization: Fusion of male gamete with the female gamete is called fertilization. In pollination the pollen grains reach the stigma and start their germination. The pollen is shed from the anther at 2-celled condition (i.e. small generative cell and large vegetative cell or tube cell). Then the exine ruptures and intine grows through the germ pore into a tube celled pollen tube. The pollen tube grows downward through the stigma into the style, ultimately reaches the ovule through micropyle. The generative nucleus divides into two male gametes. The tube nucleus later on disorganizes. Usually, the pollen tube enters the ovule through the micropyle. The process is known as, porogamy. The pollen tube may also enter through chalaza (chalazogamy) or through integuments (misogamy). Each male gamete consists of a nucleus surrounded by a sheath of cytoplasm. The male gametes are carried at the tip of pollen tube. The pollen tube reaches the ovule, penetrates the embryo sac. The tip of the pollen tube bursts to liberate the two gametes near the egg cell. One of the male nucleus fuses with egg and forms the diploid oospore. Thus, fusion of male gamete with the female gamete or egg is known as, fertilization. The second male gamete fuses with the secondary nucleus and forms the primary endosperm nucleus. It is noteworthy that two polar nuclei or secondary nucleus is diploid (n + n). When male gamete fuses with secondary nucleus the product – primary endosperm nucleus becomes triploid (i.e. 2n + n = 3n). Therefore, this fusion is also known as, triple fusion forming primary endosperm nucleus. Significance: (1) Double fertilization is unique feature of Angiosperms. During this process both the gametes released by pollen tube are utilized in fertilization. (2) The two male gametes fertilize: (a) haploid egg, forming diploid zygote which ultimately develops into a diploid embryo. (b) The secondary nucleus (2N), forming a triploid primary endosperm nucleus. It ultimately gives rise to an endosperm, which is a specialized nutritive tissue. 9 XII (3) Due to double fertilization, the embryo is supplemented with nutritive tissue, the endosperm. The endosperm thus ensures the growth and development of embryo. The endosperm in Angiosperms is post-fertilization product. (In gymnosperm endosperm is always haploid and formed before fertilization). Endosperm and Embryo Development: Development of endosperm: The formation of endosperm is initiated by mitotic divisions of primary endosperm nucleus (3n). Endosperm is a special nutritive tissue. The fusion of male gamete (n) and secondary nucleus (2n) results in formation primary endospermic nucleus (PEN) (3n). PEN undergoes many mitotic divisions to form triploid endosperm. It fills up the space of nucellus. The formation of endosperm occurs usually prior to the zygotic or oospore division. The endosperm is of three types: viz. (i) nuclear (ii) cellular and (iii) helobial. In the nuclear type, the nucleus undergoes repeated divisions. The nuclei do formed get arranged towards the periphery, leaving a large central vacuole. Then cytokinesis follows from periphery to the centre. Thus at maturity it shows cellular appearance. This, is common type of endosperm as seen in Maize, Sunflower, Wheat. 2. Cellular Type: Here every nuclear division is followed by cytokinesis, thus, giving cellular appearance from the beginning. e.g. Datura. 3. Helobial Type: Here the first mitotic division is followed by cytokinesis. This results in the formation of two unequal cells. The divisions taking place in both the cells are free nuclear. e.g. Asphodelus tenuifolius. The seeds in which endosperm is present are known as endospermic or albuminous seeds. In many seeds, however all the food stored up in the endosperm at its early stages is used by developing embryo. The endosperm is thus absent in seeds. Therefore, they are called nonendospermic seeds or exabluminous seed. The examples of endospermic seeds are Jowar, Castor etc. and non – endospermic seeds are Pea, Bean, Groundnut etc. 10 XII Development of Embryo: Fusion of male gamete with the egg cell results in the formation of zygote. The first division of the zygote produces a hypobasal cell (basal) towards the micropyle, and epibasal cell (terminal) towards chalaza. The basal cell divides repeatedly and ultimately produces a row of 4-8 cells and forms the suspensor. The terminal cell divides first vertically and then transversly forming 4-celled proembryo. The suspensor pushes the proembryo into the endosperm to enable developing embryo to receive nutrition. The suspensor absorbs food and feeds the developing embryo. The main aim of suspensor is to push the young embryo deeper into the embryo sac. The basal cell of suspensor enlarges and acts as an absorbing organ. It is called haustorial cell while its terminal cell called hypophysis cell, divides and redivides and gives rise to the apex of radicle. The proembryo by further divisions gives rise to whole of the embryo with all of its differentiated parts i.e. the radicle, plumule (stem apex) and two cotyledons (in dicot seed) or only one cotyledon (in monocot seed). The relative positions of micropyle, suspensor, embryo with cotyledons, endosperm can be understood by referring to (H.I.). As the endosperm and embryo mature, the integuments of ovule become hard. Thus, they provide protective covering called seed coat, i.e. testa. Finally there is formation of seed. 11