Developmental Biology

Developmental Biology in botany deals with the process of reproduction and all the series of
developmental stages required to form a new plant from reproductive structures.

Reproduction

The vital characteristics of living organisms by which new individuals are formed and so formed
individuals are similar to their parents are called reproduction. It is of two types- Asexual and sexual
reproduction.

Asexual reproduction

The reproduction in which new individuals are formed from a single parent without the involvement
of gametes and their fusion is called asexual reproduction. It is also called apomixis. The individuals
of the new generation are exactly similar to their parent and are known as clone (unit- remete).

Asexual reproduction is of following types: Agamospermy, spore formation, and vegetative

propagation. Here we discuss vegetative propagation.

Vegetative propagation

It is the type of reproduction in which new plants are produced directly from any vegetative part of
the parent plant. The part of the plant which produces new plant is called propagule. Vegetative
propagation is of following types:

Natural vegetative propagation

Artificial vegetative propagation

Types of vegetative propagation

Natural vegetative propagation

The vegetative reproduction which occurs naturally without human effort is called natural vegetative
propagation. This type of reproduction takes place in various ways-
a) By roots: E.g sweet potato,Dahlia, Asparagus e.t.c. by modified tuberous roots. The leafy shoot
arises from buds of the root are called slips. Some plants that propagate by common roots

are Dalbergia, Alnus, Rosa e.t.c.

b) By stems: Different types of stems are capable of producing new plants such as:

⇒Underground stems
1) Rhizome: Thickened, prostrate underground stem storing food is called rhizome. Example-
Banana, ginger, turmeric e.t.c.

2) Corm: Swollen, unbranched, vertically grown underground stem is called corm.

Example- Colocasia, Crocus e.t.c.
3) Bulb: Underground, condensed stem containing fleshy leaves is called bulb. Example- Onion,

Garlic e.t.c.

4) Tuber: Swollen ends of underground stem storing food is called tuber. Example-

Potato,Marsileae.t.c.

5) Sucker: The short underground stem branches that arise at the base of the aerial shoots is called a
sucker. Example- Mint,Chrysanthemum e.t.c.

⇒Sub-aeiral stems:
 Creeping and rooted stems which are neither aerial nor underground are called sub-aerial stems.

1) Runner: The narrow, green, prostrate stem which has distinct nodes and internodes is called a
runner. Eg- Cynodon, Centella e.t.c.

2) Stolen: The stem which grows laterally from the base of a stem and runs horizontally is called a
stolon. Eg- Strawberry e.t.c.

3) Offset: The stem which is found in hydrophytes and is like the runner but has only one internode
been called offset. Eg- Eichhornia, Pistiae.t.c.

⇒ Aerial stems:

Normal stem also gives rise to plants. Eg- Sugarcane, rose e.t.c.

C] By leaves: Plants likeBryophyllum, Begonia, Adiantum e.t.c. reproduce vegetatively with the
help of leaves.

d) By bulbils: The modified multicellular floral or vegetative fleshy buds are known as bulbils. Eg-
Garlic, Pineapple,Cycas, Oxalis e.t.c.

Artificial Vegetative Propagation

The vegetative reproduction which occurs by human efforts is called artificial vegetative
propagation. This method is applied for the quick production of new plants and combination of the
qualities of two varieties of plants.

a) Cutting: The portion of any plant organ such as stem, root or leaf, used for vegetative propagation
is called cutting. Stem cuttings are most commonly used for this purpose. Factors such as the
optimum length and diameter of the cutting, age of the parent plant and season are to be
considered while selecting a cutting for each species. Some of the plants propagated by stem
cuttings are sugarcane, rose, Hibiscus e.t.c.

b) Layering: Layering is a technique for plant propagation in which a portion of an aerial stem is
encouraged to grow roots while still attached to the parent plant and then removed and planted as
an independent plant. Layering is done on the one-year-old branch during a rainy season. A soft
basal branch is defoliated and small injury or cut is giving in the middle. The injured part is either
pegged down in the soil or covered by grafting clay to produce adventitious roots on it. Until the
production of roots, it is supplied with water. After the production of roots, it can be separated from
the parent plant. Layering is of different types- Soil layering (Serpentine or Mould) and air layering

(Gootee).

⇒ Serpentine layering is the layering in which a single branch is pegged in soil at many places to
produce many plants from a single branch. Mould layering is the layering in which a branch is
pegged in soil at one place and its apical bud is exposed in the air. The place which is pegged in the
soil gives rise to roots and it can be cut to plant separately.

⇒ Air layering is the type of layering which is done in the air on woody non-flexible branches. A
healthy branch is defoliated and given an injury. The injured region is then covered by grafting clay
and the clay is covered again by a rug which is tied with the help of a rope. This is called gootee. It is
then supplied with regular water until the formation of roots. Application of a root-promoting
substance to the exposed wound is sometimes beneficial. Grafting clay is the mixture of cow dung,
soil, finely cut hay and water. Air layering is also called gootee.

c) Grafting: Grafting is the technique of joining two plants in such a way that they unite and later
develop a composite plant. Grafting is done between the two closely related dicotyledonous plants
having vascular cambia. The rooted supporting portion of one plant, called stock, is joined with a
twig of another plant's shoot system, called scion.

There are numerous uses for grafting:

- To create new plants

- To create dwarf plants

- To increase hardiness

- To increase disease resistance

- To change the natural plant form

A grafted plant consists of a scion- a short piece of stem with two or more buds, and the stock-
which is the lower portion of the graft. There must be a contact with the cambium layer of the scion
and the stock for successful growth to occur.
source: [Link]

fig: Grafting

It is also essential that the graft doesn't dry out. This is prevented by wrapping the grafted area with
a wax, grafting compound or rubber/ polyethene banding.

The scion and stock must also be compatible plants. Like apple to apple, orange to orange (Same
genus).

The main reason for grafting is to asexually propagate of orchard trees, shade trees, and roses. There
are several grafting methods commonly used by propagators:
Whip and tongue grafting is commonly used to propagate fruit trees especially apple and pear. The
root of a young seedling tree is used for the rootstock. The scion is dormant twig containing three or
four buds- it is the diameter of a pencil. The stock and scion are cut at a slant angle. The whip or
tongue is made by making a single cut one-half inch deep into the sloping cut- both the scion and the
rootstock have a tongue cut. Insert the scion tongue cut in the rootstock cut. Line up the cambium
layers and bind with graft tape. Seal with wax or grafting compound to prevent moisture loss.

Cleft grafting is often used to join a smaller scion to a larger stock. It is usually done in the late
winter. The branch or tree is cut straight across. With a knife or grafting tool, a split of the slit is
made in the cut end of the limb or rootstock. Next a scion piece 3-5" long with one or two buds is cut
and the end sharpened into a thin wedge.

Budding is similar to grafting except that the scion is reduced to a single bud with a small portion of
bark or wood attached. The single bud scion is joined with the root stick to form a new plant. It is
done in the spring or fall when the bark separates easily from the wood. It is faster, easier, and more
economical than grafting. No wax is needed and cambium doesn't need to be aligned. Less scion is
needed here.

The other types of grafting are crown grafting, side grafting, wedge grafting e.t.c.

d) Approach grafting: It is also known as inarching. It is used to join together plants that are
otherwise difficult to join. The plants are grown close together and then joined so that each plant
has roots below and growth above the point of union. Both scion and stock retain their respective
parents that may or may not be removed after joining. The graft can be successfully accomplished
any time of the year.

The distinguishing feature of approach grafting is that two independently growing, self-sustaining
plants are grafted together. These self-sustaining characteristics of both plants which are to be
grafted ensure the survival of both even if the grafting attempt is, for some reason, not successful.
However, odds of being successful are greatly enhanced because of the active growing condition of
both plants involved and absence of a time limitation required for the healing of the graft union to
occur before the dependent scion does from the lack of sustenance.

Fig: Approach grafting

e) Micropropagation: It is a process of plant propagation in which a plant tissue is excised and
cultured in an artificially prepared nutrient medium under controlled laboratory conditions. The
ability of every living plant cell to produce the entire plant is called totipotency. This is being
exploited industrially to multiply plants which are difficult to propagate by conventional means.

Advantages of vegetative propagation-

1) It is easier, quicker, and less expensive method of propagation.

2) Characters of the parent plant are preserved. A superior variety can be produced by vegetative
means.

3) It is the only method of reproduction in plants which do not reproduce viable seeds such as
seedless grapes, rose, banana e.t.c.

4) It is easier to get rid of pathogens by vegetative propagation.

5) It is possible to raise a large number of selected strains by vegetative propagation.

6) By tissue culture, a large number of disease-free identical plants can be grown in a very short
period.

Disadvantages of vegetative propagation-

1) The quality of the plants cannot be improved.

2) Offspring do not show genetic variations.

3) Unwanted characters cannot be eliminated from plants.

4) Diseases in the parent plants are transmitted to the offsprings.

Sexual reproduction in angiosperm

The process of reproduction in which two parents are involved to produce new generation
by the fusion of meiotically formed male and female gametes is called sexual reproduction.

In higher plants, sexual reproduction takes two steps:

Sporogenesis: It is the process of formation of spores.

Gametogenesis: It is the process of formation of gametes.

Sporogenesis
Types of sporogenesis.
microsporogenesis and megasporogenesis
Microsporogenesis
Microsporogenesis is the process of development of microspores or pollen grains from
microspore mother cell inside the microsporangium/pollen sac of anther.

Stamen is the male sex organ of a flower which has three parts- Anther, Connective, and
Filament. The fertile part of the stamen is anther. On either side of connective, there is an
anther lobe. Each anther lobe contains two chambers or pollen sacs called microsporangia.
Therefore, there are four microsporangia per anther i:e tetrasporangiate. Such anthers are
called dithecous. In Malvaceae, anther is bisporangiate and monothecous.

Each microsporangium/pollen sac develops from a group of archesporial cells. In immature

cells of the anther, one cell gets distinct and contains abundant cytoplasm, called
archesporial cell. The archesporial cell then divides periclinally to form the outer primary
parietal cell and inner primary sporogenous cell. The parietal cell further divides to give rise
to outer layers of anther and sporogenous cell enlarges and behaves as microspore mother
cell. Microspore mother cell then divides meiotically to form microspores or pollen tetrad.
 s

Micro Gametogenesis
The process of formation of male gametophyte from functional pollen grain is called micro
gametogenesis. Pollen is haploid structure. Each pollen is globose, unnucleated, dust-like
yellowish minute structure. A pollen grain has two layered wall- an outer exine and inner
intine. At some places, exine is absent and called germ pore from where emerges a pollen
tube.
Pollen grain divides mitotically to form two unequal cells- small generative and larger
vegetative cell. At this two-celled stage, pollination takes place. The generative cell gives rise
to two male gametes while the vegetative cell forms pollen tube which is made from intine.
Fig: Micro gametogenesis

Female gametes are formed inside ovules which follows two steps: megasporogenesis and
mega gametogenesis.
Megasporogenesis
The process of development of megaspore inside megasporangium from megaspore mother
cell is called megasporogenesis.
The gynoecium is composed of the carpel. Each carpel bears a stigma, style, and an ovary.
The ovary contains ovules and arrangement of ovules inside ovary are called placentation.
Structure of an ovule(megasporangium)
The ovules are borne on a cushion like tissue called placenta. Each ovule is an ovoid
structure which is attached to the placenta by means of a stalk called funicle. The ovule
contains a mass of thin-walled parenchymatous cells called nucellus. The nucellus is covered
by one or two multicellular coats, called integuments. The integument does not cover
nucellus completely. A narrow passage called micropyle is left at the tip of the ovule. The
base of nucellus from where the integuments arise is called chalaza.

In young ovule, a hypodermal cell in nucellus becomes

large to form the archesporial cell. The archesporial
cell divides periclinally to form the outer primary
parietal cell and inner primary sporogenous cell. A
primary parietal cell divides anticlinal many times to
form the wall of the embryo sac. The primary
sporogenous cell enlarges itself to form megaspore
mother cell. Megaspore mother cell divides meiotically
to form four haploid megaspores. Out of four
megaspores, three towards micropyle degenerate and
only one towards chalaza remain functional.

Mega gametogenesis (development of female gametophyte)

The process of formation of female gamete from functional megaspore is called mega
gametogenesis. Functional megaspore undergoes three times mitosis to form embryo sac
with 8 nuclei. Initially, there are 4 nuclei at each micropyle and chalaza end but one of them
from each pole move towards centre and fuse together at the time of fertilization to from a
diploid secondary nucleus. Three cells at chalaza end are called antipodal cells which later
degenerate. Out of 3 cells of the micropylar end, one enlarges in size and is called egg cell.
Other two cells are called as synergids which secrete chemicals to guide pollen tube.

Fig: Mega gametogenesis

Pollination
The process of transfer of pollen grains from the anther to the stigma of the flower is called
pollination. Pollination occurs at the two-celled stage of pollen grains. Pollination is of two types-
Self-pollination and cross-pollination.
Types of pollination

Self-Pollination
The pollination in which pollen grains are transferred to the stigma of the same flower or different
flower of the same plant is called self-pollination. However, true self-pollination (autogamy) involves
some flower. Thus, it is of two different types- Autogamy and Geitonogamy.

Autogamy is the self-pollination in which pollen grains are transferred to the stigma of the same
flower.
Geitonogamy is the pollination in which pollen grains are transferred to the stigma of a
different flower of the same plant. It is, therefore, ecologically cross pollination but genetically self-
pollination.

True self-pollination (autogamy) is of two types- direct and indirect. Direct autogamy is the type of
autogamy in which pollination occurs due to the direct touch between the anthers and stigma by
growth or bending (e.g.-Catharanthus, Mirabilis). Indirect autogamy occurs due to factors like dew,
gravity etc.

The factors which favour self-pollination are as follows-

1) Bisexuality - The flower having both male and female reproductive organs is a bisexual flower.

2) Homogamy - The process of maturity of male and female organs of bisexual flower at the same
time is called homogamy.

3) Cleistogamy - The self-pollination that occurs in a closed flower is called cleistogamy. Eg-
Balsam, Oxalis.

4) Chasmogamy - The self-pollination that occurs in an open flower is called chasmogamy.

Eg- Mirabilis,Sunflower.

5) Bud pollination - The self-pollination that occurs in bud condition is called bud pollination. Eg- Pea,
Wheat, Rice etc.
Advantages of self-pollination:

1) It maintains pure lines.

2) It is more economical as there is less number of pollen grains required.

3) It is independent of other factors.

4) Flower do not require to make attractive parts.

5) It ensures seed production.

6) It eliminates some bad recessive characters.

Disadvantages of self-pollination:

1. New characters are not introduced.

2. Adaptability of race decreases.

3. There is no variation in the new generations.

4. Continuous self-pollination may lead to the extinction of the race.

5. It may carry unwanted characters to new individuals.

6. It has no role in evolution.

Cross-pollination

It is also called allogamy (but allogamy represents xenogamy and geitonogamy). Based on plants,
cross-pollination is of two types- Xenogamy and hybridism.

Xenogamy is the cross pollination which takes place between two different flowers of two
different plants of same species.

Hybridism is the cross pollination which takes place between two genetically different plants.

Cross pollination is always carried out by some agents. Based on the agents, it is of different types-
Anemophily, Hydrophily, and Zoophily.

Anemophily
.• Pollination by air is called anemophily.

• Plants produce an enormous amount of pollen when compared to the number of ovules available
for pollination to compensate the uncertainties of pollination.

• Flowers with well-exposed stamens are usually pollinated by anemophily.

• Large feathery stigma to trap airborne pollen grains.

• Most winds pollinated flower contains single ovule in one ovary and numerous flower packed into
an inflorescence.

• Pollen grains are light and non-sticky.

Hydrophily
• Pollination by an abiotic agent like water is called hydrophily.

• This type of pollination is very rare, about 30 genera, mostly monocot.

•Vallisneria, Hydrilla, and the common example for hydrophily.

• All aquatic plants are not Hydrophily.

• Pollen grains released into the surface of the water and carried to the stigma by the air current as
in Vallisneria.

• In sea grains, the flowers are submerged.

• Pollen grains are protected from wetting by mucilaginous covering.

Pollination by biotic agent

• Majority of flowering plants use a range if animals as pollinating agents.

• Among the animal, insects particularly bees, are the dominant biotic agents for pollination.

• Insect pollinating flowers are very large, colorful, fragrant, and rich in nectar.

• Small flowers present in a cluster to make them conspicuous.

• Flower pollinated by flies and beetles secrete foul odours.

• Nectar and pollen grains are the usual floral rewards for insects.

• In some species, floral rewards are in providing safe places to lay eggs.

• A species of moth and Yucca plant cannot complete their life cycle without each other. The moth
deposits its eggs in the locule of the ovary and the flower in turn gets pollinated by the moth.

• Many insects may consume pollen or nectar without bringing about pollination. Such floral visitors
are referred as pollen/nectar robbers,

Based on the biotic agents involved, pollination is of different types:

1. Entomophily- pollination by insects

2. Ornithophily - pollination by birds

3. Chiropterophily - pollination by bats

4. Malacophily - pollination by snails

5. Ophiophagy - pollination by snakes

6. Anthophily - pollination by human beings in plant breeding technique.

Contrivances to ensure cross-pollination:

1) Unisexuality or dicliny- The flowers having single sex organ is called the unisexual flower.

2) Dichogamy- The maturity of male and female reproductive organ at the different time is called
dichogamy. It may be protandry (anthers mature earlier than pistil), eg- Sunflower, salvia e.t.c. or
protogyny (pistil matures earlier than anthers), eg- Mirabilis, Gloriosa e.t.c.
3) Prepotency- The phenomenon in which pollen grains of the same flower germinates more rapidly
on the stigma of another flower is called prepotency. Eg- Apple, grape e.t.c.

4) Self-sterility- The phenomenon in which pollen grains of the same flower fail to germinate on the
stigma of that flower is called self-sterility. Eg- Tobacco, Potato e.t.c.

5) Heterostyly- The phenomenon having different heights of male and female reproductive organs in
a flower is called heterostyly. Eg- Primula, Jasmine e.t.c.

6) Herkogamy- The physical barrier present between male and female organs of a flower to ensure
cross-pollination is called herkogamy.

Advantages of cross-pollination:

1) The self-sterile plants can continue their race.

2) It induces variations in new generation.

3) It helps in evolution.

4) It creates better adaptability in the progeny.

5) New and more useful varieties can be produced by this method.

6) Productivity is high.

7) The defective characters are eliminated.

Disadvantages of cross-pollination:

1. It is highly expensive as a large number of pollen grains are required.

2. It is dependent on other factors.

3. Some undesirable characters may remain in the race.

4. The useful characters may be spoiled.

5. The plant has to create a large number of characters to attract biotic agents.

6. Seed production is less ensured.

Fertilization And Embryogenesis

Fertilization
The process of fusion of male and female gametes to form the zygote is called fertilization. In
angiosperms, the male gametes are carried to the egg cell by pollen tube, this phenomenon is called
siphonogamy.

The pollen grains reach the surface of stigma by pollination. Pollen grains absorbs moisture from the
surface of stigma. As a result, pollen grains swells and the intine comes out in the form of a tube
called pollen tube. Pollen tube is driven forward by dissolving the tissue of style with the help of
enzymes present at its tip. The direction of the pollen tube is controlled by the chemicals secreted
by synergids. Such movement by chemicals is known as a chemotactic movement.

The pollen tube enters the ovary by three routes -

Through chalaza

Through integuments OR

Through micropyle

The entry of pollen tube through the micropyle is the most common and is called as porogamy.

The entry of pollen tube through the chalaza is called as chalazogamy.

The entry of pollen tube through the integuments is called as mesogamy.

Chalazogamy and mesogamy are collectively called chalazogamy.

The pollen tube enters through synergids and breaks up to release its two male [Link] of
these two male gametes, one fuses with the egg cell to form a zygote. This process is
called syngamy. Another male gamete travels little far and fuses with the secondary nucleus to form
endosperm mother cell. This process is called triple fusion. Since two types of fusion i:e syngamy and
triple fusion, takes place in the embryo sac the phenomenon is called double fertilization.

Syngamy results in the formation of the zygote which later divides to give rise to an embryo.

Triple fusion results in the formation of endosperm mother cell which later divides to form
endosperm tissue. Endosperm tissue is the nutritive tissue for the growing embryo.

Double fertilization is the fusion of two male gametes brought by a pollen tube to two different cells
of the same female gametophyte in order to produce two different structures.
Fig: Fertilization

Embryogenesis

The process of formation of an embryo from a zygote is known as embryogenesis. It can be studied
under two headings:

 Dicot embryogenesis
 Monocot embryogenesis

Dicot Embryogenesis
The zygote elongates and then divides by a transverse wall into two unequal cells. The larger basal
cell is called suspensor cell and the other towards the chalaza is called terminal or embryo cell. The
suspensor cell divides transversely a few times to produce a filamentous suspensor of 6-10 cells. The
suspensor helps in pushing the embryo in the endosperm. The first cell of suspensor towards the
micropylar end becomes swollen and functions as haustorium. The haustorium helps to draw
nutrition. The last cell of the suspensor towards chalaza is called hypophysis which later gives rise to
the radicle and root cap.

The embryo cell undergoes two vertical divisions and one transverse division to form eight-celled
octant structure. Four cells towards the chalaza are known as epibasal cells and four towards
micropyle are called hypobasal cells. The epibasal cells later give rise to two cotyledons and plumule.
The hypobasal cells produce the hypocotyl.

The octant divides periclinally to produce an outer layer of protoderm. The inner cells differentiate
into procambium and ground meristem. Protoderm forms epidermis, procambium gives rise to
vascular strand and ground meristem produces cortex and pith,

Initially, the embryo is globular and undifferentiated. Early embryo with radial symmetry is called
proembryo. It later becomes heart-shaped. The growth rate of cotyledons is very high and plumule
remains at the place from where cotyledons are separated.
 Fig; dicot embryogenesis

Monocot Embryogenesis

The zygote elongates and then divides transversely to form basal and terminal cells. The basal cell
towards the micropylar end becomes large and called as suspensor cell. It may function as
haustorium. The terminal cell divides by transverse wall to form two cells. The top cell after a series
of divisions forms plumule and a single cotyledon. Cotyledon called scutellum, grows rapidly and
pushes the terminal plumule to one side. The plumule comes to lie in a depression. The middle cell,
after many divisions' forms hypocotyl and radical. It also adds a few cells to the suspensor. In some
cereals, both plumule and radicle get covered by sheaths developed from scutellum
called coleoptiles and coleorhiza respectively.

Apomixis and polyembryony

Apomixis is very common in Asteraceae and grasses.

Seeds are produced without fertilization.

Apomixis is a type of asexual reproduction which mimics the sexual reproduction.

The diploid egg cell is formed without meiosis and develops into seed without fertilization.

In Citrus and Mango, the nucellar cells start dividing, protrude into the embryo sac and develop into
an embryo.

Ovule having more than one embryo is termed as polyembryony.

Hybrid plants are developed by apomixis to maintain the genetic identity.