Plant Life Cycle and Alternation of Generations

Alternation of Generations Definition:

Alternation of generations is a type of life cycle in which subsequent generations of
plants alternate between diploid and haploid organisms.

What is Alternation of Generations?

Alternation of generations is common in plants, algae, and fungi. This can be
compared to the sexual reproduction in animals where both haploid and diploid cells
are found in every generation.

Plants alternate between the diploid sporophyte and haploid gametophyte, and
between asexual and sexual reproduction. Therefore, the life cycle of plants is known
as alternation of generations. The ability of the plants to reproduce sexually and
asexually helps them to adapt to different environments.

The alternation of generations depends upon the type of the plant. In Bryophytes, the
dominant generation is haploid and the gametophyte comprises the main plant. In
tracheophytes, the dominant generation is diploid and the sporophyte comprises the
main plant.

The plants’ life cycle in one of the two generations is dominant over the other. The
plants in the dominant generation grow larger and live longer. The plants in the non-
dominant generations are small and hardly visible. On the contrary, the dominant
generations are seen in the form of ferns, trees or other plants.

The dominant generation in vascular plants is the sporophyte, while in the non-
vascular plants is the gametophyte.

Describe the Alternation of Generations in plants –

Life cycle:
The alternation of generations include the following stages: 

The diploid sporophyte has a structure called sporangium. 

The sporangium undergoes meiosis and forms haploid spores. 
The spore develops into a gametophyte which is haploid in nature. 
The gametophyte has the reproductive organs which undergo mitosis to form haploid
gametes. 
The gametes fertilize to form a haploid zygote which matures into a mature
sporophyte. This cycle keeps repeating.

Stages of Alternation of Generations:

Following are the two stages of alternation of generations:
Sporophyte Generation:
Two haploid gametes fuse together to form a diploid zygote. This results in a
sporophyte.
The sporophyte is formed by multiple rounds of mitosis and is a multicellular
organism. On reaching maturity, the sporophyte develops reproductive organs known
as sporangia. This is one key point in the alternation of generations.
These sporangia are used to create haploid spores. These spores are released and
carried away by air and water and when the conditions are favourable they develop
into a gametophyte.

Gametophyte Generation:
This is the next generation in the alternation of generations. The spore is newly
formed and has half the DNA as the parent organism. This spore undergoes mitosis
multiple times to form a gametophyte.
The gametophyte generation creates gametes. These gametes are produced by
gametangia. These gametes are then transferred between plants or spread into the
environment.
When a gamete encounters a gamete of the opposite sex, it fuses with it to form a
zygote which eventually becomes a sporophyte.
This is the simplest version of alternation of generations. This is widely found in ferns.

Life Cycle Events in a Flowering Plant:

A flowering plant undergoes the following events during its life cycle:

Germination: A plant undergoes germination and begins to grow from seed. The
roots are formed below the soil while the leaves, roots, and stem appear above the soil.

Pollination: Pollens are carried by wind or insects to another flower. This is called
pollination.

Fertilization: The pollen travels to the ovary of the flower where the fusion of the
male and gametes take place. This is called fertilization.

Dispersal: The seeds are scattered by the wind and animals. Some of these seeds
emerge into a new plant. Thus we see how a plant life cycle begins with a seed. The
seed sprouts to form a seedling. The seedling gets converted into a new plant which
forms new seeds and the cycle continues.
 Gametogenesis in Plants: Sporogenesis and Gametogenesis

 In higher plants, the sexual reproduction completes in 2 steps:

 Sporogenesis
 Gametogenesis
 In angiosperm, flowers are the designated structures of sexual reproduction.
 A flower is termed to be complete if it possesses all its four parts: calyx,
corolla, androecium, and gynoecium.
 Androecium and gynoecium refer to the male and female organs of
reproduction respectively whereas calyx and corolla aids in reproduction.
 Gametogenesis is defined as the biological process in which the formation
of gametes take place.
 The maturation of gametes take place in the reproductive cells of androecium
and gynoecium.

Male gamete formation:

The formation of male gametes take place inside anther which consists of two steps:
1. Microsporogenesis
2. Micro gametogenesis

Microsporogenesis:
 The process of formation of microspores or pollen grains from microspore
mother cell inside the microsporangium/pollen sac of anther is termed as
microsporogenesis
 The formation of pollen grains take place in anther which is responsible for
formation of male gametes.
 A typical anther is tetrasporangiate in nature. Connective, a column of sterile
tissue connects the bilobed anther. Each lobe consists of two microsporangia
divided by strip of sterile tissue.
 Externally these lobes can be splitted by longitudinal grooves.
 At maturity, due to breakdown of partition wall, two sporangia in a lobe fuse.
 The young anther consists of homogeneous cells that are bound by a well-
defined epidermis.
 In course of development, the anther becomes four lobed, and in each lobe
some hypodermal cells become more noticeable than the others because of their
larger size and discrete nuclei.
 The cells are known as archesporium (or archesporial initials).
 The archesporial cells divide in a plane parallel to the outer wall of the anther
lobe (periclinal division), cutting off parietal cells towards the epidermis and
primary sporogenous cells towards the interior of the anther.
 The cells of the parietal layer go through a series of periclinal and anti-clinal
division to form 2-5 concentric layers of the anther wall.
 The primary sporogenous cells, either directly or after a few mitoses, plays role
as microspore mother cells.
T.S. of anther discloses the following structures:
1. Anther wall:
 The fully-fledged anther wall consists of an epidermis accompanied by a layer
of endothecium, 2 or 3 middle layers and a single layered tapetum.
2. Epidermis:
 It is the outermost layer of anther.
 These cells divide frequently by anticlinal division so as to cope up with the
rapidly enlarging internal tissue.
 Their role is to protect the anther.
3. Endothecium:
 Normally, the endothecium is single layered and emerges from the parietal cells.
 The cells adjacent to endothecium aids in the dehiscence of anther during the
course of maturation.
4. Middle layer:
 It is 2-3 layered and emerges from parietal cells.
 The cells next to endothecium help in the dehiscence of anther and others act as
storage centers for starch and other food materials essential for development of
pollen.
5. Tapetum:
 This is the innermost layer of anther wall and arises from parietal cells. It
completely surrounds the sporogenous tissue.
 It is made up of single layer of cells, each cell contains dense cytoplasm and
obvious nucleus.
 It is physiologically vital layer as all the food materials to the sporogenous
tissue must pass across it.
6. Sporogenous tissue:
 The sporogenous cells may either directly play role as microspore mother cells
(or pollen mother cells) or they may go through a few mitoses before entering
meiosis.
 In a meiotic division, each microspore mother cell yields four haploid
microspores. Microspore tetrads represent the group of four microspor
 All the pollen mother cells in an anther locule are interlinked by plasmodesmata
and with tapetal cells.
 When these cells go through meiosis the linkage between the tapetal cells and
pollen mother cells are ruptured.
 The deposition of callose (glucans) makes the walls of pollen mother cells thick.
 The plasmodesmatal connections between the pollen mother cells are replaced
by huge cytoplasmic channels.
 The channels allow passage for the transportation of cytoplasmic contents from
one cell to the other is known as cytomixis.
 Because of this, a close synchrony is maintained during meiosis in the large
number of pollen mother cells in anther locule.
  At the end of the first meiosis, the cytoplasmic channels are halted.
 Cytokinesis takes place either after first meiosis or after second meiosis
yielding tetrad.
 Mostly all the four spores within a tetrad are totally segregated from one
another and form the spores.
 Within the callose wall the microspores start to produce their individual wall.
 Then the breakdown of the common wall allows the microspores or pollen
grains to set free in the anther locule.
 Each pollen grain bears haploid nucleus in the pollen cytoplasm surrounded
externally by the plasmalemma and pollen wall.
 Pollen wall consists of outer uneven exine and inner even intine wall.

Micro-gametogenesis:

 The process of development of male gametes in the microspores or pollen

grains is termed as Micro gametogenesis.
 Microspores symbolizes the initiation of the male gametophytic generation.
 During gametogenesis, the pollen nucleus divides mitotically to yield the
unequal cells:
– The larger one is vegetative cell or tube cell which forms the pollen tube.
-The smaller one, situated towards the wall is generative cell which again
divides by mitosis to form two sperms (male gametes).
 After germination of pollen grain, the generative cell may divide inside the
pollen grain or in the pollen tube.
 At the same time when these alterations are going on, the microspore or pollen
grain is also synthesizing its wall.
– At one or more loci, the pollen wall is very thin.
– These regions are called germpores.
 Before pollen mitosis starts, 2 discrete changes in the pollen protoplast are
observed:
 The nucleus is replaced from the center towards one side of the cell, which
gives sign for the position of generative cell.
  The cytoplasm between the nucleus and the wall, on the side where vegetative
cell is to be halted become highly vacuolated.
 Initially, the cytoplasm of the vegetative cell and that of generative cell is
disconnected by two plasma membranes.
 The wall of generative cell grows inwards between the plasma lemma of the
generative cell and the intine, till the two ends of the wall connects and merge,
then the cell is finally removed off and come to lie freely in the cytoplasm of
the vegetative cell.
 Afterwards, the wall of generative cell soon vanishes, and the cytoplasm of
generative cell remains enclosed in two plasma membranes.
 The generative cell divides by a mitosis division.
 If this division takes place when they are in anther, the pollens are shed at 3
celled stage and if it takes place after pollen germination, they are shed at 2
celled stage.
 The exine ruptures and the intine forms a pollen tube.
 The vegetative nucleus enters the pollen tube. Now it is known as tube nucleus.
 The generative tube nucleus enters the tube and divides to form two generative
nuclei which finally form the two male gametes.
 The tube nucleus disorganizes.
 One of the gamete fuses with the female gamete during fertilization.

Female gamete formation:

The formation of female gametes take place inside the ovules which consists of
following steps:
1. Mega sporogenesis
2. Mega gametogenesis
Mega sporogenesis:
 The process of formation of megaspore, within the megasporangium or ovule is
called as mega sporogenesis.
 Megaspores represents the female gametophyte.
 The ovule emerges as a small mass of homogenous tissue on the placenta in the
ovary.
 Integuments arise near to the base of this tissue which forms the nucellus in
mature ovule.
– Although the integuments commence later, they grow rapidly than the
nucellus and soon covers it almost completely, omitting the region of the
micropyle.
 The archesporium modifies immediately below the nucellar epidermis.
-The archesporial cell directly plays role as the megaspore mother cell so that
the sporagenous cell is hypodermal and the nucellar tissue around it stays single
layered. -Such ovules are called tenuinucellate.
 In some others, the hypodermal archesporial cell divides obliquely, cutting an
outer parietal cell and an inner sporogenous cell.
-The parietal cell may either remain undivided or go through a few periclinal
and anticlinal divisions so that the sporogenous cell becomes embedded in the
 nucellar tissue.
– All such ovules where the sporogenous cell becomes sub-hypodermal, either
due to the formation of parietal cells or due to the divisions in the nucellar
epidermis, or both are called crassinucellate.
 The archaesporium turns more obvious than its surrounding cells due to its
larger nucleus, denser cytoplasm and larger size.
 It either directly functions as the megaspore mother cell or it divides
periclinally cutting an outer primary parietal cells and an inner sporogenous cell.
 The latter functions as the megaspore mother cell.
 Megaspore mother cell undergoes meiosis to yield four haploid megaspores.
 Only one of the four megaspores function and form the female
gametophyte, the remaining upper three megaspores degenerate.
 Normally, the chalazal megaspore of tetrad is functional.
 The megasporangium together with the integuments, is called ovule.
 It is affixed to the placenta, on the inner wall of the ovary, by a stalk called
funiculus.
 An ovule ready for fertilization constitutes of nucellar tissue covered almost
completely by one or two integuments, leaving a small opening at the apical
end.
 This opening termed as micropyle is the main passage for the entry of the
pollen tube in the ovule.
 The basal region of the ovule where funiculus is affixed is called chalaza.
 In the nucellus, the female gametophyte is present, also called embryo sac.

Mega gametogenesis:

 The process of development of female gametes in the ovule or megaspore is

known as the mega gametogenesis.
 The megaspore (n) marks the initiation of the megagametophyte generation.
 The nucleus of the megaspore divides and grows into the female gametophyte
or the embryo sac.
  The first division of the functional megaspore nucleus yields the two nuclei,
one situated at the micropylar end and the other at the chalazal end inside
the, megaspore.
 Both the nuclei divide and thus two nuclei lie at each pole.
 The third division yields 8 nuclei, 4 located at the micropylar end and the
other four at the chalazal end.
 Out of the 4 nuclei at each pole, one moves to the centre and forms
secondary nucleus after fusion.
 The three nuclei at the micropylar end forms the egg apparatus, and those
at the chalazal end forms the three antipodal cells.
 In the egg apparatus, larger cell is called the egg cell and the linked
comparatively smaller cells are known as synergids.
 In this way, embryo sac is formed.
 At the same time, the tissue of nucellus elongates and forms funicle.
 The entire structure is now termed as ovule or megasporangium.

Describe embryo sac as 7 celled and 8 nucleated with suitable diagram:

Female Gametophyte i.e. embryo sac develops from megaspore mother cell(MMC)
inside the ovule through the process megasporogenesis and megagametogenesis. The
diploid megaspore mother cell divides by meiosis to produce 4 haploid megaspores
and this process is called megasporogenesis.
Out of 4 megaspores, only one survives and the other 3 degenerate. This functional
megaspore then divides by mitosis to produce two daughter nuclei which move to
opposite pole of the ovule. These two nuclei further undergo 2 more mitotic division
divisions to produce 4 nuclei at each pole. Three nuclei at the micropyler pole
constitute the egg apparatus which contains one egg cell and two synergids. Three
nuclei at the chalazal end constitute the antipodal cells.
One nuclei leaves from each pole and moves towards centre and form a central cell
which is diploid. These nuclei are called polar nuclei. Thus, a mature embryo sac is 7
celled but 8 nucleated.
 Double Fertilization

Double Fertilization: In angiosperms, one sperm fertilizes the egg to form the 2n
zygote, while the other sperm fuses with two polar nuclei to form the 3n endosperm.
This is called a double fertilization.

After pollen is deposited on the stigma, it must germinate and grow through the style
to reach the ovule.
The microspores, or the pollen, contain two cells:
i. The pollen tube cell and
ii. The generative cell.

The pollen tube cell grows into a pollen tube through which the generative cell travels.

The germination of the pollen tube requires water, oxygen, and certain chemical
signals. As it travels through the style to reach the embryo sac, the pollen tube’s
growth is supported by the tissues of the style.

During this process, if the generative cell has not already split into two cells, it now
divides to form two sperm cells.

The pollen tube is guided by the chemicals secreted by the synergids present in the
embryo sac; it enters the ovule sac through the micropyle.

Of the two sperm cells, one sperm fertilizes the egg cell, forming a diploid zygote; the
other sperm fuses with the two polar nuclei, forming a triploid cell that develops into
the endosperm.

Together, these two fertilization events in angiosperms are known as double

fertilization. After fertilization is complete, no other sperm can enter.
The fertilized ovule forms the seed, whereas the tissues of the ovary become the fruit,
usually enveloping the seed.

Early Embryonic Development

Embryo development: Shown are the stages of embryo development in the ovule of
a shepherd’s purse (Capsella bursa). After fertilization, the zygote divides to form an
upper terminal cell and a lower basal cell. (a) In the first stage of development, the
terminal cell divides, forming a globular pro-embryo. The basal cell also divides,
giving rise to the suspensor. (b) In the second stage, the developing embryo has a
heart shape due to the presence of cotyledons. (c) In the third stage, the growing
embryo is crowded and begins to bend. (d) Eventually, it completely fills the seed.

After fertilization, embryonic development begins.

The zygote divides to form two cells:

i. The upper cell (terminal cell) and
ii. The lower cell (basal cell).

The division of the basal cell gives rise to the suspensor, which eventually makes
connection with the maternal tissue. The suspensor provides a route for nutrition to be
transported from the mother plant to the growing embryo.

The terminal cell also divides, giving rise to a globular-shaped proembryo.

In dicots (eudicots), the developing embryo has a heart shape due to the presence of
the two rudimentary cotyledons. In non-endospermic dicots, such as Capsella bursa,
the endosperm develops initially, but is then digested. In this case, the food reserves
are moved into the two cotyledons.
As the embryo and cotyledons enlarge, they become crowded inside the developing
seed and are forced to bend. Ultimately, the embryo and cotyledons fill the seed, at
which point, the seed is ready for dispersal.

Embryonic development is suspended after some time; growth resumes only when the
seed germinates. The developing seedling will rely on the food reserves stored in the
cotyledons until the first set of leaves begin photosynthesis.

Key Points
 Double fertilization involves two sperm cells; one fertilizes the egg cell to
form the zygote, while the other fuses with the two polar nuclei that form the
endosperm.
 After fertilization, the fertilized ovule forms the seed while the tissues of the
ovary become the fruit.
 In the first stage of embryonic development, the zygote divides to form two
cells; one will develop into a suspensor, while the other gives rise to a
proembryo.
 In the second stage of embryonic development (in eudicots), the developing
embryo has a heart shape due to the presence of cotyledons.
 As the embryo grows, it begins to bend as it fills the seed; at this point, the
seed is ready for dispersal.

Key Terms
 double fertilization: a complex fertilization mechanism that has evolved in
flowering plants; involves the joining of a female gametophyte with two male
gametes (sperm)
 suspensor: found in plant zygotes in angiosperms; connects the endosperm to
the embryo and provides a route for nutrition from the mother plant to the
growing embryo
 proembryo: a cluster of cells in the ovule of a fertilized flowering plant that
has not yet formed into an embryo

Cryptochrome

A large number of plants responses to blue light have been known for long time.The
blue light specific responses of higher plants include phototropism, stomatal
movement, inhibition of hypocotyl elongation, pigment biosynthesis and gene
activation.The exact identity of the blue light photoreceptor is not yet known and
hence the name cryptochrome has been given which implies a “hidden pigment”

Phytochrome

Phytochrome is a blue protein pigment responsible for the perception of light in

photo-physiological processes. It is possibly the only photoreceptor in photoperiodism
and the flowering process.

Phytochrome structure:
1. Phytochrome is a soluble chromoprotein with a molecular mass of 250 kDa.
2. It occurs as a dimer made up of two subunits, each of 125 kDa.
3. Each subunit consists of two components ― light absorbing pigment molecule,
chromophore and a polypeptide chain, apoprotein.
4. The apoprotein monomer has a molecular mass of 125 kDa.
5. Apoprotein and chromophore together make up the holochrome.
6. The chromophore is a linear tetrapyrrole similar to phycocyanin termed as
phytochromobilin and it is a ring attached to the protein through a thioether-linkage to
a cysteine residue.
7. The principal difference between the Pr chromophore and the Pfr chromophore
appears to be a cis-trans isomerization of the methane-bridge between rings C and D.

Biosynthesis:

1. Phytochrome apoprotein alone cannot absorb red or far-red light.

2. Light can be absorbed only when polypeptide is covalently linked with
phytochromobilin to form the holoprotein.
3. Phytochromobilin is synthesized within plastids.
4. After synthesis, it leaks out of the plastid into the cytosol.
5. Assembly of apoprotein with chromophore is autocatalytic, that is, it occurs
spontaneously when purified apoprotein is mixed with chromophore in test tube, for
which no cofactors are necessary.
Phytochrome Controlled Responses:

☻The regulatory effects of light on plant growth and development are visualized
most prominently at two stages in the life cycle of the plant ― firstly, at the stage of
seed germination and seedling development, and secondly, at the stage of transition
from the vegetative to the flowering phase.
☻For the sake of convenience, these diverse photomorphogenetic responses can be
classified into two types, fast responses and slow responses.

Type I: Fast Responses:

☻The type I responses include those processes in which the quantum energy
absorbed by the plant in transduced to another form of energy.
☻Examples of this type of essentially energy-transducing responses include leaf
movement of Mimosa and chloroplast movement in Mougeotia.
☻These phenomena are relatively rapid, occurring on a time scale of second and
minutes.

Type II: Slow Responses:

☻The rates and activation of certain aspects of growth and development are switched
on or modulated under the influence of the quality of light (red or far-red).
☻Examples of type II responses include stem elongation, seed germination, leaf
expansion, flower initiation and pigment biosynthesis.
☻Type II responses are relatively slow responses occurring on a time scale of hours
and days