Sexual reproduction in flowering plants

PRATIVA SHUKLA
ZOOLOGY [ M.PHILL]
 A Fascinating Organ of Angiosperms

 Morphologically flower is a modified shoot meant for sexual reproduction.

Flower

Floral whorls

Calyx Corolla Androecium Gynoecium

(sepals) (Petals) (Stamen) (Carpel)

Reproductive whorls
Flower showing various parts
Pre-Fertilization : structure and event

 All flowering plants (angiosperm) show sexual reproduction.

 Flowers are the sites of sexual reproduction.

 Several hormonal and structural changes result in differentiation and development of the
floral primordium.

 Inflorescences bear the floral buds and then flowers.

 Pre-Fertilization : structure and event
structure of a flower

 A typical flower has 2 parts :

Androecium

Gynoecium

 Androecium : it is the male reproductive part of the flower.

 It consists of a whorl of stamens. Their number and length are variable in different species.
 Parts of a stamen : Filament

Anther
 Pre-Fertilization : structure and event
structure of a flower

 Filament is a long and slender stalk.

 Its proximal end is attached to the thalamus or the petal of flower.
 Terminal and typically bilobed. Each lobe has 2 thecae (dithecous)
 Often a longitudinal groove runs lengthwise separating the theca.
 Pre-Fertilization : structure and event
Transverse section of Anther

 The anther is a tetragonal structure consisting of four microsporangia located at the corners (2 in each lobe).
 The microsporangia develop to pollen sacs. They extend longitudinally all through the length of an anther and are
packed with pollen grains.
 Structure of Microsporangium

 A typical microsporangium is near circular in outline.

 It is surrounded by 4 wall layers :
 Epidermis
 Endothecium
 Middle layers
 Tapetum
 The outer 3 layers give protection and help in dehiscence of anther to release the pollen.
 The tapetum(innermost layer) nourishes the developing pollen grains.
 Cells of the tapetum contain dense cytoplasm and generally have more the one nucleus.
 In young anther,each microsporangium has sporogenous tissue at centre. It consists of compactly arranged
homogenous diploid cells (sporogenous cells)
 Microsporogensis

 As the anther develops, each sporogenous cell (microspore or pollen mother cell) undergoes meiotic divisions to
form microspore tetrads (microspores arranged in a cluster of four cells).
 The formation of microspores from a pollen mother cell (PMC) through meiosis is called microsporogenesis.
 As the anther mature and dehydrate, the microspores dissociate from each other and develop into pollen grains.
 Each microsporangium contains thousands of pollen grains. They are released with the dehiscence of anther.
 Pollen grain (male gametophyte)

 Generally spherical.
 25-50 mm in diameter.
 Cytoplasm is surrounded by a plasma membrane.
 A pollen grain has a two-layered wall : exine and intine
 Exine : the hard outer layer made up of sporopollenin (highly resistant organic material). It can withstand high
temperature and strong acids and alkali. Enzymes cannot degrade sporopollenin.
 Exine has apertures called germ pores where sporopollenin is absent.
 Pollen grains are preserved as fossils due to the presence of sporopollenin
 Exine exhibits patterns and designs.
 Intine : the inner wall
 It is a thin and continuous layer made up of cellulose and pectin.
 A matured pollen grain contains 2 cells :
 Vegetative cell : it is bigger, has abundant food reserve and a large irregularly shaped nucleus.
 Generative cell : it is small and floats in the cytoplasm of the vegetative cell. It is spindle shaped with dense
cytoplasm and a nucleus.
 Over 60% angiosperms shed their pollen grains at 2- celled stage.
 In others, generative cell divides mitotically to give 2 male gametes. Thus pollen grains are shed at 3- celled stage.
 The shed pollen grains have to land on the stigma before they lose viability.
 The viability period of pollen grains is variable. It depends on temperature and humidity.
 Viability of pollen grains of some cereals (rice, wheat etc) is 30 min. some members of Rosaceae and solanceae have
viability for months.
Economic importance of pollen grain

 These are rich nutrients. Pollen tablets are used as food supplements. Pollen tablets and syrups
increase performance of athletes and race horses.

 They are stored for years in liquid nitrogen (-196 degree C). They are used as pollen banks in
crop breeding programmes.

 Pollen grains of some plants (e.g. Parthenium or carrot grass) are allergic for some people. It
leads to chronic respiratory disorders- asthma, bronchitis, etc
 Gynoecium :

 Female reproductive part.

 It may have a single pistil (monocarpellary) or more than one pistil (multicarpellary)
 In multicarpellary, the pistils may be fused together (syncarpous) or free (apocarpous)
 Each pistil has three parts :
 Stigma : Landing platform for pollen grains.
 Style : Elongated slender part beneath the stigma.
 Ovary : Basal bulged part.
 Inside the ovary is the ovarian cavity (locule) in which placenta is located.
 Arising from the placenta are the ovules (megasporangia).
 The number of ovules in an ovary may be one (wheat, paddy, mango etc) to many (papaya, water melon, orchids
etc)
 Structure of Megasporangium (ovule)

 Ovule is attached to the placenta by a stalk (funicle).

 Junction between the body of ovule and funicle is called hilum.
 Each ovule has 1 or 2 protective envelops (integuments) except at the tip where a small opening (micropyle) is present.
 Opposite the micropylar end is the chalaza (basal part).
 Enclosed within the integuments, there is a mass of cells called nucellus. Its cells contain reserve food materials.
 Inside the nucellus is the embryo sac (female gametophyte).
 An ovule generally has a single embryo sac formed from a megaspore.
Structure of Megasporangium (ovule)

 It is the formation of megaspores from megaspore mother cell (MMC)

 Ovules generally differentiate a single MMC in the micropylar region of the nucellus.
 It is a large cell containing dense cytoplasm and a prominent nucleus.
 MMC undergoes meiosis to produce 4 megaspores.
Structure of Megasporangium (ovule)

 It is the formation of megaspores from megaspore mother cell (MMC)

 Ovules generally differentiate a single MMC in the micropylar region of the nucellus.
 It is a large cell containing dense cytoplasm and a prominent nucleus.
 MMC undergoes meiosis to produce 4 megaspores.
 Formation of female gametophyte
(embryo sac)

 In majority of flowering plants, one megaspore is functional while the other three degenerate.
 The functional megaspore develops into the female gametophyte.
 The embryo sac formation from a single megaspore is called monsporic development.
 The nucleus of functional megaspore divides mitotically to form 2 nuclei. They moves to opposite poles,
forming 2-nucleate embryo sac.
 The nuclei again divide two times forming 4-nucleate and 8-nucleate stages of the embryo sac.
 These divisions are free nuclear, i.e. nuclear divisions are not followed immediately by cell wall formation.
 After the 8-nucleate stage, cell walls are laid down leading to the organization of the typical female gametophyte. Large
central cell.
 6 of the 8 nuclei are surrounded by cell walls and organized into cells. Remaining 2 nuclei (polar nuclei) are situated
below the egg apparatus in the large central cell.
 Distribution of cells in the embryo sac

 A typical mature embryo sac is 8-nucleate and 7- celled.

 3 cells are grouped at the micropylar end and form egg apparatus. It consists of 2 synergids and one egg cell.
 Synergids have special cellular thickenings at the micropylar tip called filiform apparatus. It helps to guide the pollen
tubes into the synergid.
 3 cells at the chalazal end are called the antipodals.
 The large central cell has two polar nuclei.
 Pollination

 It is the transfer of pollen grains from the anther to the stigma of a pistil.
 TYPES OF POLLINATION : a) Autogamy (self-pollination)
 b) Geitonogamy
 c) Xenogamy
 Based on the source of pollen, pollination is 3 types :
a) Autogamy (self – pollination) :
 It is the transfer of pollen grain from the anther to stigma of the same flower.
 In flowers with exposed anthers $ stigma, complete autogamy is rare.
 Autogamy in such flowers requires synchrony in pollen release and stigma receptivity. Also, anther $ stigma should be
close to each other.
 Plants like viola (common pansy), Oxalis $ Commelina produce 2 types of flowers :
 Chasmogamous flowers
 Cleistogamous flowers
Viola
oxalis
Chasmogamous cleistogamous
 Autogamy (self-pollination)

 CHASMOGAMOUS FLOWERS : They are similar to flowers of other species with exposed anthers and stigma
 CLEISTOGAMOUS FLOWERS : They do not open at all.
 Anther and stigma lie close to each other.
 They are autogamous.
 When anther dehisce in the flower buds, pollen grains come in contact with stigma for pollination.
 Cleistogamous flowers produce assured seed –set even in the absence of pollinators.
 Cleistogamy leads to inbreeding depression.
Geitonogamy

 It is the transfer of pollen grains from the anther to the stigma of another flower of the
same plants.
 It is functionally cross-pollination involving a pollinating agent. But it is genetically
similar to autogamy since the pollen grains come from the same plant.
 Xenogamy

 It is the transfer of pollen grains from anther to the stigma of a different plant.
 It brings genetically different pollen grains to the stigma.
Agent of pollination
Abiotic agents : Wind

 Pollination by wind is called anemophily.

 More common abiotic agent.
 Wind pollinated flowers often have a single ovule in each ovary and numerous flowers
packed into an inflorescence.
 E.g – Corncob – the tassels are the stigma and style which wave in the wind to trap
pollen grains.
 Wind-pollination is quite common in grasses.
 Abiotic agents : Wind

 WAYS FOR EFFECTIVE POLLINATION :

 Flowers produce large amount of pollen.
 Pollen grains are light and non- sticky.
 They often possess well-exposed stamens (for easy dispersion of pollens into wind currents).
 Large, feathery stigma to trap air-borne pollen grains.
 Abiotic agents : Water

 Pollination by water is called hydrophily.

 It is quite rare. It is limited to about 30 genera, mostly monocotyledons.
 E.g. Vallisneria $ Hydrilla (fresh water), Zostera (marine sea-grasses) etc.
 But in lower plants, water is a regular mode of transport for the male gametes. Distribution of some bryophytes $
pteridophytes is limited because they need water for the transport of male gametes and fertilization.
 In Vallisneria, the female flower reaches the surface of water by the long stalk and the male flowers or pollen grains are
released on to the surface of water. They are carried by water currents and reach the female flowers.
 In sea grasses, female flowers remain submerged in water. Pollen grains are long and ribbon like. They are carried
inside the water and reach the stigma.
 Abiotic agents : Water

 The pollen grains of most of the water pollinated species have a mucilaginous covering to protect from wetting.
 Not all aquatic plants use hydrophily.
 In most of aquatic plants (water hyacinth,water lily etc), the emerge above the level of water for entomophily or
anemophily.
 Biotic agents : Animals

 Used by majority of flowering plants.

 E.g. Bees, butterflies, flies, beetles, wasps, ants, moths, birds, bats, primates, reptiles etc.
 Pollination by insects (Entomophily), particularly bees is more common.
 Often flowers of animal pollinated plants are specifically adapted for a particular species of animal.
 FEATURES OF INSECT – POLLINATED FLOWERS
 Large, colourful, fragrant and rich in nectar. Nectar $ pollen grains are the floral rewards for pollination.
 Small flowers from inflorescence to make them visible.
 The flowers pollinated by flies and beetles secrete foul odours to attract these animals.
 The pollen grains are generally sticky.
 Biotic agents : Animals

 When the animal comes in contact with the anthers and the stigma, its body gets pollen grains.
 When its comes in contact with the stigma, it results in pollination.
 Some plants provide safe places as floral reward to lay eggs. E.g. Amorphophallus (it has the tallest flower of 6 feet).
 A moth species and the plant Yucca cannot complete their life cycles without each other. The moth deposits its eggs in the
locule of ovary. The flower gets pollinated by moth. The larvae come out of the egg as seeds start developing.
 Many insects consume pollen or nectar without bringing about pollination. They are called pollen/nectar robbers.
 Outbreeding Device

 Hermaphrodite flowers can undergo self-pollination (autogamy).

 Continued self-pollination results in inbreeding depression.
 To avoid self-pollination and encourage cross-pollination, there are some devices in plants.
 a) Avoiding synchronization : Here, the pollen is released before the stigma becomes receptive or before the release of
pollen.
 b) Arrangement of anther $ stigma at different positions.
 c) Self-incompatibility : It is genetic mechanism to prevent self-pollen (from the same flower or other flowers of the
same plant) from fertilization by inhibiting pollen germination or pollen tube growth in the pistil.
 d) Production of unisexual flowers : if male $ female flowers are present on the same plant (i.e. monoecious, e.g, castor $
maize), it prevents autogamy but not geitonogamy.
 In dioecious plants (e.g. papaya), male and female flowers are present on different plant (dioecy). This prevents both
autogamy and geitonogamy.
 Pollen – pistil Interaction

 It is the process in which pistil recognizes compatible or incompatible pollen through the chemical components produced
by them.
 If the pollen is compatible (right type), the pistil accepts it and promotes post-pollination events.
 Pollen grains germinates on stigma to produce a pollen tube through one of the germ pores. The contents of pollen grain
move into the pollen tube. Pollen tubes grows through the tissue of stigma and style and reaches the ovary.
 Pollen – pistil Interaction

 If the pollen is incompatible (wrong type), the pistil rejects pollen by preventing pollen germination or the pollen tube
growth.
 In plants which shed pollen grains at 2- celled condition (a vegetative cell $ a generative cell), the generative cell divides
into two male gametes during pollen tube growth.in plants which shed pollen in the 3-celled condition, pollen tubes
carry 2 male gametes from the beginning.
 Pollen tubes reaches the ovary, then enters the ovule through micropyle and then enters one of the synergids through the
filiform apparatus. The filiform apparatus present at the micropylar part of the synergids guides the entry of pollen tube.
 Artificial hybridisation

 It is a crop improvement programme in which desired pollen grains are used for pollination.
 This is achieved by following techniques :
 Emasculation
 Bagging
 EMASCULATION : Removal of anthers from the bisexual flowers bud of female parent before the anther dehisces.
 BAGGING : Here, emasculated flowers are covered with a suitable bag ( made up of butter paper) to prevent
contamination of its stigma with unwanted pollen.
 When the stigma attains receptivity, mature pollen grains collected from anthers of male parent are dusted on the stigma.
Then the flowers are rebagged and allowed to develop the fruits.
 For unsexual flowers, there is no need for emasculation. Female flower buds are bagged before the flowers open. When
the stigma becomes receptive, pollination is carried out using the desired pollen and the flower rebagged.
 Double Fertilization

 After entering one of the synergids, the pollen tube releases the 2 male gametes into the cytoplasm of the synergid. One
male gamete moves towards the egg cell and fuses with its nucleus (syngamy) to form zygote(diploid).
 The other male gametes moves towards the two polar nuclei located in the central cell and fuses with them to produce a
triploid primary endosperm nucleus (PEN). As it involves fusion of 3 haploid nuclei, it is called triple fusion.
 Since 2 types of fusion (syngamy $ triple fusion) take place in an embryo sac, it is called double fertilization.
 It is an event unique to flowering plants.
 The central cell after triple fusion becomes primary endosperm cell(PEC) and develops into the endosperm. Zygote
develops into an embryo.
 Post – Fertilization : structures $ events

Post-fertilization event

 Endosperm
endo
Embryo development Ovule into seed Ovary into fruit
development
Endosperm development

 Primary endosperm cell divides repeatedly to form a triploid endosperm tissue.

 Endosperm cells are filled with reserve food materials. They are used for nutrition of the
developing embryo.
 In common endosperm development, PEN undergoes successive nuclear divisions to
give free nuclei. This stage is called free-nuclear endosperms.
 Number of free nuclei varies greatly.
 Endosperm becomes cellular due to cell wall formation.
 Tender coconut water is a free-nuclear endosperm (made up of thousands of nuclei) and
the surrounding white kernel is the cellular endosperm.
 Endosperm development

 Embryo develops at micropylar end of the embryo sac where the zygote is situated.
 Most zygotes divide only after the formation of some endosperm. This gives nutrition to developing embryo.
 In monocots $ dicots, seeds differ greatly. But embryogeny (early embryonic developments) is similar.
 Dicotyledonous Embryo

 It has an embryonal axis and 2 cotyledons.

 The portion of embryonal axis above the level of cotyledons is the epicotyl, which terminates with the plumule (stem
tip).
 The cylindrical portion below the level of cotyledons is hypocotyl that terminates with the radicle (root tip).
 The root tip is covered with a root cap.
 Monocotyledonous Embryo

 They possess only one cotyledon.

 Cotyledon of the grass family is called scutellum.
 It is situated lateral to the embryonal axis. At its lower end, the embryonal axis has the radicle and root cap enclosed in
coleorrhiza (an undifferentiated sheath).
 Portion of embryonal axis above the level of attachment of scutellum is the epicotyl. It has a shoot apex and a few leaf
primordial enclosed in coleoptile ( a hollow foliar structure).
 Seed from ovule

 Seed is the fertized ovule formed inside fruits.

 It is the final product of sexual reproduction.
 It consists of seed coats, cotyledons $ embryo axis.
 The cotyledons are simple, generally thick and swollen due to storage food.
 Mature seeds are 2 types :
 a) Non-albuminous seeds : They have no residual endosperm as it is completely consumed during embryo development.
E.g. pea, groundnut, beans.
 b)Albuminous seeds : They retain some endosperm. E.g. wheat, maize, barley, castor, coconut, sunflower.
 Occasionally, in some seeds (black pepper, beet etc) remnants of nucellus are also persistent. It is called perisperm.

 Integuments of ovules harden as tough protective seed coats. It has a small pore (micropyle) trhough which oxygen $
water enter into the seed during germination.

 As the seed matures, it becomes dry by reducing water content (10-15 % moisture by mass). The metabolic activity of the
embryo slows down. It may enter a state of inactivity (dormancy).

 Under favourable conditions (moisture, oxygen $ suitable temperature), they germinate.

Structure of some seed
Advantage of seeds

 1. Since pollination and fertilization are independent of water, seed formation is more dependable.

 2. Better adaptive strategies for dispersal to new haitats. It helps the species to colonize in other
areas.

 3. They have food reserves. So young seedlings are nourished until they are capable of
photosynthesis.

 4. The hard seed coat protects the young embryo.

 5. Being products of sexual reproduction, they generate new genetic combinations and variations.
Viability of seeds after dispersal

 In a few species, the seeds lose viability within a few months. Seeds of many species live
for several years.

 Some seeds can remain alive for hundreds of years. The oldest is that of a lupine
(Lupinus arcticus) excavated from Arctic Tundra. The seed geminated and flowered after
an estimated record of 10,000 years of dormancy.

 2000 years old viable seed is of the date palm (phoenix dactylifera) discovered during the
archeological excavation at king Herod’s palace near the Dead Sea.
 Fruit from ovary

 The ovary develops into a fruit.

 Trasformation of ovules into seeds and ovary into fruit proceeds simuitaneously.
 The wall of ovary develops into pericarp (wall of fruit)
 The fruits may be fleshy (e.g. guava, orange, mango, etc.) or dry (e.g. groundnut,mustard etc).
 Fruits are 2 types :
 True fruits : in this, fruit develops only from the ovary. Other floral parts degenerate and fall off.
 False fruit : in this, the thalamus also contributes to fruit formation. E.g. apple, strawberry, cashew etc.
 In some species,fruits develop without fertilization. Such fruits are called parthenocarpic fruits. E.g. Banana.
 Parthenocarpy can be induced through the application of growth hormones. Such fruits are seedless.
 Apomixis and polyembryony

 Development of apomictic seeds :

 In some species, diploid egg cell is formed without reduction division and develops into the embryo without
fertilization is called Apomixis.
 Occurrence of more than one embryo in a seed is called polyembryonay.

Multiple embryos

 single seed
 Importance of apomixes in hybrid seed
industry

 If the seeds from hybrids are sown, the plants in the progeny will segregate and lose hybrid characters.
 Production of hybrid seeds is costly. So hybrid seeds are also expensive.
 If the hybrids are made into apomicts, there is no segregation of characters in hybrid progeny. So farmers can
keep on using hybrid seeds to raise new crop.