Cell Cycle and Cell Division
Cell Cycle and Cell Division
CELL CYCLE
Cell cycle ( Howard and Pelc, 1953) is genetically controlled series of changes that occur
in a newly formed cell by which it supplicates ins contents,1undergoes, growth and
division to form two daughter .it consists of states or periods, a long nondividing growth
I-phase and a short dividing M-phase. Both have substages. I-phase represents
interphase.
The regular sequence of G1, S, G2 ( interphase) and M phase ( mitotic phase) is called
the cell cycle.
Interphase is called „resting stage”, but it is in fact a period of great activity. Three
important process, which are preparatory to cell division, take place during interphase.
Thus it is also known as preparatory phase. These processes are
- Replication of DNA along with the synthesis of nuclear proteins such as the
histones
- Synthesis of embryo rich compounds, which provide energy for mitosis, and
synthesis of proteins at the end of interphase states or periods, a long non -dividing
growth I-phase and a short dividing M-phase. Both have substages. I-phase represents
interphase.
The regular sequence of G1, S, G2 ( interphase) and M phase ( mitotic phase) is called
the cell cycle.
Interphase is called „resting stage”, but it is in fact a period of great activity. Three
important process, which are preparatory to cell division, take place during interphase.
Thus it is also known as preparatory phase. These processes are
- Replication of DNA along with the synthesis of nuclear proteins such as the
histones
- Synthesis of embryo rich compounds, which provide energy for mitosis, and
synthesis of proteins at the end of interphase
Interphase of dividing cell has been classified into three subphases -G1- Phase, S-phase
and G2-phase
G1-phase
G1 phase is also known as first growth phase or post mitotic gap phase. It is the longest
phase of cell division. In this phase different types of RNA (mRNA, tRNA, rRNA) and
proteins are synthesized.
All cell organelles ( ER, mitochondria, Golgi complex, ribosomes, plasmid in plant cell)
multiply. The duration of G1 Phase varies from cell to cell. It is shorter in frequently
dividing cells. G1 phase cell has three options.
The deciding factor for above option are availability of mitogen and energy rich
compounds. This point is called check point .
S-phase
S-phase is known as synthetic phase. In this stage replication of DNA takes place by the
synthesis of histones. As a result each chromosome under goes replication producing two
chromatids. Each chromosome carries a duplicate set of genes. A haploid cell (n)
becomes diploid (2n) and a diploid cell (2n), thus becomes tetraploid (4n) at the end of S-
phase. Repairing of damaged DNA also takes place.
G2-phase
G2 is also called second growth phase or pre-mitotic gap phase. In this phase synthesis of
DNA stops and synthesis of RNAs and proteins continues. All cell organelles multiply
and spindle formation takes place. It lasts for 2-5 hours in most cells. Some proteins
formed in this phase cause condensation of chromosomes to initiate mitosis
G0-phase
The phase in which cells fail to divide further ( do not undergo S-phase after G1-phase)
and undergo differentiation is known as G0 phase or quiescent stage. It occurs due to
non-availability of mitogen and energy rich compounds. The cells remain metabolically
active, grow in size and differentiate for particular function after attaining a particular
shape.
However some cells remain in undifferentiated state as reserve cells. They may proceed
with cell division when required e.g. fibroblasts; it helps in healing of wounds and grow
and divide again.
M-phase
The process of cell division is found to be essentially the same in all living organism and
the events are chiefly centered in the nucleus. Three type of cell divisions have been
distinguished:
Mitosis and meiosis are the two major types of cell division. The basic stage in both the
types of divisions are almost identical.
Amitosis is a direct division characterized by the splitting of nucleus followed by that of
cytoplasm.
Mitosis is a somatic cell division which takes place in vegetative cells. It maintains the
chromosome number.
Meiosis is a reduction division, occurring in the reproductive cells. The chromosome
number are reduced to half.
AMITOSIS
Occurrence
Drawbacks
As Amitosis does not distribute chromatin equitably, it results in structural and functional
abnormalities in the cell.
MITOSIS
Mitosis is a type of cell division in which chromosomes of parent cells are duplicated ( by
replication of DNA) and equally distributed ( quantitatively and qualitatively) into two
daughter nuclei. Term mitosis is derived from Greek word “Mitos” means thread or fibril
Mitosis was first observed by Strassburger in plant cells (1870) and Boveri and
Flemming in animal cell ( 1879 ). The term was coined by Flemming in 1882. It is also
known as equational division due to equal distribution of chromosomes in daughter
nuclei. It is often known as somatic cell division due to occurrence in somatic cells.
In plants all meristematic regions are the sites of mitosis e.g. root ape, shoot apex,
intercalary meristem, lateral meristem, leave, flowers, fruits, embryo, seeds etc.
In animals embryo, skin, bone marrow etc are the sites of mitosis
Mitosis is completed in two steps karyokinesis and cytokinesis. Karyokinesis
Mitosis starts with the nuclear division of parent cell known as karyokinesis ( Gk, karyon
-nucleus, kinesis – movement) . The four phases of karyokinesis are prophase,
metaphase, anaphase and telophase
Prophase
Prophase ( Gk Pro-first, phase –stage ) is often divided into three substages – early
prophase, mid prophase and late prophase. It is the first stage of mitosis proper. It is the
longest phase of Karyokinesis
Early prophase
In this sub-stage nucleus and cell become spheroid and nucleus appears as boll of wool.
Chromatin fibre condense to form elongated chromosome and this increases viscosity and
refractivity of cytoplasm
In animal cells duplicated centrioles. ( S phase of interphase) start to move towards
opposite poles of the cell. Each centriole radiates out fine microtubular fibrils called
astral rays. In animal cells and cells of lower plants, fibrils appears like spokes of a wheel
around each centriole to form an aster.
Late prophase
In this substage spindle fibres start appearing around the nucleus. The size of
chromosomes is much reduced as compared to early prophase . Spindle poles are formed
without asters in plant cells and with asters in animal cells.
Nucleolus and other cell organelles ( like mitochondria, Golgi complex, ER, vacuoles
etc) disappear. The presence of the spindle is essential for mitosis. If cells are treated with
colchicines, which inhibits spindle formation, anaphasic movement of the two groups of
chromosomes to the poles does not take place.
Metaphase
Anaphase
In anaphase ( Gk ana – up, phase – stage) chromosomes are arranged on the equatorial
plate for a short period. The centromeres of chromosomes starts to divide into two,
forming daughter chromosomes with centromere in each. Daughter chromosomes are
repulsive so, migrate towards opposite poles. Spindle fibres attached to the centromeres
shorten and pull the chromosomes to the poles. The velocity of anaphasic movement does
not depend on the size of the chromosomes. In anaphasic movement of chromosomes, the
centromeres lead the path while the limbs trails behind. So anaphasic chromes, the
centromeres lead the path while the limbs trail behind. So anaphasic chromosomes appear
as V-, L-, J- and I- shaped.
At the end of anaphase two groups of chromosomes are formed, one at each pole. The
number and types of chromosomes at each pole is the same as in the parent nucleus.
Telophase
Mitosis ends with division of cytoplasm known as cytokinesis. It is derived from greek
word “cytos” means hollow or cell, “kinesis: means movement. It starts towards the
middle of anaphase and is completed with the telophase. It is different in animal and
plants. If nuclear divison takes place without cytoplasmic division, a syncytium is formed
animal cytokinesis.
The furrow forms and deepens centripetally and finally cleaves the parent cell into two
daughter cells. This method of cytokinesis is known as cleavage method. All cell
organelles ( mitochondria, Golgi complex, lysosomes., ER, ribosomes etc) are also
distributed between two daugheter cells.
Significance of mitosis
Growth and development - A single cell zygote grows into full blown baby ( 6×1022
cells) by repeated mitosis. Plants are able to grow throughout their life due to mitotioc
division in their apical and lateral meristems. Increases in tissue mass, results from
increase in cell number called hyperplasmic. Hnece, mitosis is essential for growth and
development of a multicellular organism.
Maintenance of cell size: An overgrown somatic cell is induced to divide so that mitosis
helps in maintaining a proper surface volume ratio. It has also a high nucleocytoplasmic
ratio which is brought back to efficient level through divisions. These ratios are important
for proper functioning of cell distribution of all the chromosomes. This helps in proper
co-ordination among daughter cells.
Healing and regeneration – For healing of wounds new cells are produced by Mitosis
some organism are able to regenerate missing part of body also whole organism though
mitosis
Reparing – the mechanism for replacing old or worn out cells is called repairing. In
human body roughly 5 ×109 cells are lost from skin surface, lining of alimentary canal,
blood cell etc. these are replaced by new cells formed through mitosis.
Evidence of basic relationship – The mechanism of mitosis are similar in the majority of
organism, showing basic similarity and relationship among them
MEIOSIS
Meiosis is a process of reductional division in which the number of chromosomes per cell
is cut in half. In animals, meiosis always results in the formation of gametes, while in
other organism it can give rise to spores. The word “meiosis” comes from greek world
meioun, means “to make small”, since it results in a reduction of the chromosome
number.
The term meiosis was coined by Farmer and Moore (in 1905). The division was first
studied by Van Benedin (1887), Strassburger (1888), Sutton (1900) and Winiwater
(1900). Meiosis I &II were differentiated by Gregoire. In 1911 the American geneticist
Thomas Hunt morgan ( 1866 – 1945) observed cross-over in Drosophila melanogaster
meiosis and provided the first genetic evidence that genes are transmitted on
chromosomes
Meiosis is essential for sexual reproduction and therefore occurs in all eukaryotes (
including single –celled organisms) that reproduce sexually. Meiosis does not occur in
archaea or bacteria, which reproduce via asexual process such as binary fission.
In lower plants, and in many protists, meiosis results in formation of haploid cells that
can divide vegetatively without undergoing fertilization, referred to as spores.
The cells of a particular species have a constant number of chromosomes. In sexually
reproducing organisms male and female gametes fuse together to form the zygote. If the
gamete has the same number of chromosomes number remains constant from generation
to generation. This is because of meiotic division which reduces the chromosome number
to half.
Types of meiosis
The cells in which meiosis takes place are called meiocytes. In animals, meiocytes are of
two types, spermatocytes and oocytes. In higher plants, meiocytes are differentiated into
microsporocytes and megasporocytes. Depending upon the stage when meiosis occurs,
the latter is of three types - gametic, zygotic and sporic meiosis.
Gametic meiosis
Meiosis is most of the animal take place during the formation of gametes
(gametogenesis). It is termed as genetic meiosis. When two gametes fuse in fertilization,
a diploid zygote is formed. Gametic meiosis results in diplontic life cycle
Zygotic meiosis
In some lower plants meiosis takes place in the zygote and the resulting organism are
haploid. It is called zygotic meiosis. Organism having zygotic meiosis have haplontic life
cycle.
Sporic meiosis
Meiosis I
In meiosis I, the homologous pairs in a diploid cell separate, producing two haploid cells,
so it is also referred to as a reduction division. Like mitosis, it is studied under for stages
– prophase, metaphase, anaphase and telophase.
Prophase I
Prophase I is more complicated and prolonged stage . prophase I is divided into five sub-
phases: Leptotene, zygotene, pachytene, diplotenme and diakinesis.
Leptotene
Leptotene also known as leptonema is a first stage of prophase I during which individual
chromosomes begin to condense into long strands within the nucleus which are loosly
interwoven. However the two sister chromatids are still so tightly bound that they are in
distinguishable from one another.
Leptotene chromosomes may be irregularly arranged or may be polarized towards the
centrioles forming a „bouquet‟. Electron microscope studies have shown that bouquet
formation results when a group of chromosomes is attached close together on the nuclear
membrane . There are two sets of chromosomes in a diploid cell undergoing meiosis, one
set contributed by the male parent and other by the female parent. These are always two
similar chromosomes, having the same size, form and structure. They are called
homologous chromosomes. One of them is paternal chromosome and the other maternal
chromosome.
Zygotene
During zygotene or zygonema the chromosomes become shorter anad thicker. The
homologous chromosomes come to lie side by side in pairs . ( G. zygon = yolk; tene =
thread). This pairing of homologus chromosomes is known as synapsis or syndesis. A
pair of homologous chromosomes lying together is called a bivalent. The chromatids are
still not visible. A fibrillar, somewhat ladder-like , organelle, called synaptonemal
complex, develops between the synapsed homologous chromosomes. It is thought to
stabilize the paired condition of chromosomes till crossing over is completed.
Pairing of two homologous chromosomes begins when their corresponding ends come
together on the nuclear matrix. Pairing may occur in one of the following three ways-
(i) Proterminal pairing : It starts at the ends and proceeds towards the middle
(ii) Procentric pairing: it begins at the centromeres and progresses towards the ends.
(iii) Random ( intermediate) pairing: It commences at many point towards the ends.
The synaptonemal complex is attached at both ends through its lateral element to the
inner surface of the nuclear membrane. The central elements is not attached directly .
Also arising from the lateral elemnent is another series of smaller loops. These loops fuse
in the middle line to make up the central element.
Pachytene
Diplotene
During diplotene or diplonema the synaptic forces keeping the homologous chromosomes
together come to an end. The homologous chromosomes start separating ( G. diplos =
double; tene = thread). This is called disjunction. It makes chromatids more distinct and
the tetrads very clear. Separation of homologous chromosomes does not take place at the
points called chiasmata (singular, chiasma). The chiasmata make the sites where crossing
over occurred during pachytene ( Gr . chiasma = crosspiece). They help in holding
homologous chromosomes together.
The number and position of chiasmata varies with the length of the chromosomes and
with the species. Chiasmata are found in the meiosis of almost all eukaryotic organism.
However, achiasmatic meiosis ( meiosis without chiasma) has been reported in some
organisms, e.g. males of higher dipteral ( includeing Drosophila), Panorpa ( scorpion fly),
many mantids and roaches, some grasshoppers and scorpions. A chiasma formed at the
ends of chromosomes is called termical chiasma. Chiasmata formed along the lengths of
chromosomes are called interstitial chiasmata.
During diplotene the chiasmata begins to be displaced along the length of the
chromosome. The terminal chiasma slips off the ends of the chromosomes, and its
position is taken up by an interstitial chiasma, which is now called the terminal chiasma.
This process is called terminalization. As diplotene progresses the number of interstitial
chiasmata becomes lesser in number. The terminalization may be due to electrostatic
force or despiralization of chromosomes.
When terminalization is completed the homologous remain in contact through the
terminal chiasma. The degree of terminalization is expressed by the terminalization
coefficient (T).
The synaptonemal complexes mostly disappear during diplotene. In certain regions short
segments may persist. The most common regions where the complexes persists are, near
the ends of the bivalents where the lateral elements are attached to the nuclear membrane,
and at the sites of chiasmata formation. With the disappearance of the synaptonemal
complexes the axial filaments become unpaired.
In diplotene, the chromosomes may unfold to nearly normal form and start transcription
of mRNA and rRNA to build up food reverves in the cytoplasm. This process is most
profound in the primary ooctytes of amphibians, reptiles and brids. In some species, the
chromosomes enlarge greatly, assuming lampbrush form.
Diakinesis
Diakinesis is not sharply differentiated from diplotene. The chromosomes become more
conctracted. The bivalents are more evenly distributed in the nucleus and migrate towards
the periphery. RNA synthesis stops. Nucleolus degenerates. A spindle begins to develop,
with or without centrioles.
Prometaphase I
The nuclear membrane disappears in prometaphase I and the chromosomes reach their
maximum contraction. Spindle formation begins.
Metaphase I
The chromosomes now become arranged on a equator of the cell, The spindle is formed.
Spindle fibres becomes attached to the centromeres of the two homologous
chromosomes. The two centromeres of each bivalent lie on opposite side of the equatorial
plate.
The attachment of tetrads to the spindle fibres in metaphase I is different from that of
mitotic metaphase chromosomes. Each homologous chromosome has two kinetochore,
one for each of its two chromatids. Both the kinetochores of a homologous chromosome
connect to the same spindle pole. The two kinetochores of its homologous join the
opposite spindle pole. In metaphase I of meiosis there are bivalents, each bivalent
consisting of two centromeres.
Anaphase I
During anaphase I, from each tetrad two chromatids of a chromosome move as a unit (
dyad) to one pole of a spindle, and the remaining two chromatids of its homologue
migrate to the opposite pole.
Thus, the homologous chromosomes of each pair, rather than the chromatids of a
chromosome, are separated. As a result, half of the chromosomes, which appear in early
prophase, go to each pole. It is here in the anaphase I that the real reduction in the poles is
still double and consists of two chromatids. This is in contrast to the single-stranded
chromosomes of mitotic anaphase
The paternal and maternal chromosomes of each homologous pair segregate during
anaphase I independently of the other chromosomes. Anaphase I is cytological event that
corresponds to Mendel‟s law of independent assortment. Although the paternal and
maternal chromosomes of a homologous pair have the genes for the same traits, either
chromosome of a pair may carry different alleles of same genes. Therefore, independent
assortment of homologous chromosomes in anaphase I introduces genetic variability.
Telophase I
During telophase I the chromosomes at each pole of the spindle uncoil and elongate, but
remain straight and often do not assume interphase form. The satellite chromosome
develop forms around the chromosomes and nucleoli. The spindle and the astral rays
gradually disappear.
The cytoplasm divides at its middle by cleavage ( constriction) in an animal cell and by
cell plate formation in plant cell. This produces two daughter cell, each has received only
one chromosome from each homologous pair. Thus it has half the number of
chromosomes, but double the amount of nuclear DNA as each chromosome is still
double.
The daughter cells formed by meiosis I are called secondary spermatocytes or secondary
oocytes in male and female animals.
Cell enter a period of rest as interkinesis or interphase II. No DNA replication occurs
during this stage. Protein and RNA synthesis may occur. It is important for bringing true
haploidy.
Meiosis II
The second meiotic division is essentially similar to mitosis. In this division, the two
chromatids of each chromosome separate from each other and go to separate daughter
cells. With the result, the number of chromosomes remains the same as produced by
meiosis I, Meiosis II is, therefore, known as homotypic division. If however, differs from
mitosis in that DNA does not duplicate, while centromere do so. It has 4 phases –
Prophase II, metaphase II, anaphase II, and telophase II.
Prophase II
Metaphase II
In metaphase II the chromosomes become oriented on the equatorial plate and have the
relationship to the spindle as in mitosis.
Anaphase II
Anaphase II, where the centromeres are cleaved, allows the kinetochores to pull the sister
chromatids apart. The sister chromatids by convention are now called sister
chromosomes, and they are pulled towards opposite poles.
Telophase II
In telophase II the group of chromosomes at each pole of the spindle gets enclosed by a
nuclear envelope. Nucleoli are laid sown, Astral rays and spindles are lost.
Cytokinesis
Cytoplasm divides at its middle by furrowing in an animal cell and by cell plate
formation in a plant cell. This produces two daughter cells. The latter have half the
number of chromosomes and half the amount of nuclear DNA. These cells are mature
gametes in animals and spores in plants.
Cytokinesis may occur after each nuclear division. In such cases, it is said to be of
successive type. First the diploid parent cell divides by heterotypic division into two
haploid cells, which then produce four haploid cells by homotypic division. The four
daughter cells may form a linear or isobilateral tetrad. Often cytokinesis is delayed until
both the nuclear divisions are completed, so that four cells are simultaneously formed,
each with a haploid nucleus. The cytoplasmic division in such cases is said to be of
simultaneous type
Significance of meiosis
Formation of gametes – Meiosis forms gametes that are essential for sexual reproduction.
Genetic information – It switches on the genetic information for the development of
gametes or gametophytes and switches off the sporophytic information
Meiosis facilitates stable sexual reproduction – Without the halving of ploidy, or
chromosome count, fertilization would result in zygotes that have twice the number of
chromosomes than the zygotes from the previous generation. Successive generations
would have an exponential increase in chromosome count, resulting in an unwildy
genome that would cripple the reproductive fitness of the species. Most importantly,
however, meiosis produces genetic variety in gametes that propagates to offspring.
Recombination and independent assortment allows for a greater diversity of genotype in
the population. As a system of creating diversity, meiosis allows a species to maintain
stability under environment changes.
Crossing over- It introduces new combination of traits or variations.
Mutations – Chromosomal and genomatic mutations can take place by irregularities of
meiotic divisions. Some of these mutations are useful to the organism and are perpetuated
by natural selection.
Evidence of basic relationship of organisms – Details of meiosis is essentially similar in
the majority of organisms showing their basic similarity and relationship.