CELL CYCLE AND CELL DIVISION

CELL CYCLE:
Cell division is a very important process in all living organisms. During the division of a cell, DNA
replication and cell growth also take place. All these processes, i.e., cell division, DNA replication, and
cell growth, hence, have to take place in a coordinated way to ensure correct division and formation of
progeny cells containing intact genomes. The sequence of events by which a cell duplicates its genome,
synthesises the other constituents of the cell and eventually divides into two daughter cells is termed
cell cycle. Although cell growth (in terms of cytoplasmic increase) is a continuous process, DNA synthesis
occurs only during one specific stage in the cell cycle. The replicated chromosomes (DNA) are then
distributed to daughter nuclei by a complex series of events during cell division. These events are
themselves under genetic control.

Phases of Cell Cycle:

A typical eukaryotic cell cycle is illustrated by human cells in culture. These cells divide once in
approximately every 24 hours (Figure 10.1). However, this duration of cell cycle can vary from organism
to organism and also from cell type to cell type. Yeast for example, can progress through the cell cycle in
only about 90 minutes.

The cell cycle is divided into two basic phases: z Interphase z M Phase (Mitosis phase) The M Phase
represents the phase when the actual cell division or mitosis occurs and the interphase represents the
phase between two successive M phases. It is significant to note that in the 24 hour average duration of
cell cycle of a human cell, cell division proper lasts for only about an hour. The interphase lasts more
than 95% of the duration of cell cycle. The M Phase starts with the nuclear division, corresponding to the
separation of daughter chromosomes (karyokinesis) and usually ends with division of cytoplasm
(cytokinesis). The interphase, though called the resting phase, is the time during which the cell is
preparing for division by undergoing both cell growth and DNA replication in an orderly manner. The
interphase is divided into three further phases: z G1 phase (Gap 1) z S phase (Synthesis) z G2 phase (Gap
2) G1 phase corresponds to the interval between mitosis and initiation of DNA replication. During G1
phase the cell is metabolically active and continuously grows but does not replicate its DNA. S or
synthesis phase marks the period during which DNA synthesis or replication takes place. During this time
the amount of DNA per cell doubles. If the initial amount of DNA is denoted as 2C then it increases to
4C. However, there is no increase in the chromosome number; if the cell had diploid or 2n number of
chromosomes at G1, even after S phase the number of chromosomes remains the same, i.e., 2n. In
animal cells, during the S phase, DNA replication begins in the nucleus, and the centriole duplicates in
the cytoplasm. During the G2 phase, proteins are synthesised in preparation for mitosis while cell
growth continues.

M PHASE:
This is the most dramatic period of the cell cycle, involving a major reorganisation of virtually all
components of the cell. Since the number of chromosomes in the parent and progeny cells is the same,
it is also called as equational division. Though for convenience mitosis has been divided into four stages
of nuclear division, it is very essential to understand that cell division is a progressive process and very
clear-cut lines cannot be drawn between various stages. Mitosis is divided into the following four stages:
z Prophase

z Metaphase

z Anaphase

z Telophase
Prophase:
Prophase which is the first stage of mitosis follows the S and G2 phases of interphase. In the S and G2
phases the new DNA molecules formed are not distinct but interwined. Prophase is marked by the
initiation of condensation of chromosomal material. The chromosomal material becomes untangled
during the process of chromatin condensation (Figure 10.2 a). The centriole, which had undergone
duplication during S phase of interphase, now begins to move towards opposite poles of the cell. The
completion of prophase can thus be marked by the following characteristic events: z Chromosomal
material condenses to form compact mitotic chromosomes. Chromosomes are seen to be composed of
two chromatids attached together at the centromere. z Initiation of the assembly of mitotic spindle, the
microtubules, the proteinaceous components of the cell cytoplasm help in the process. You have studied
mitosis in onion root tip cells. It has 14 chromosomes in each cell. Can you tell how many chromosomes
will the cell have at G1 phase, after S phase, and after M phase? Also, what will be the DNA content of
the cells at G1, after S and at G2, if the content after M phase is 2C? CELL CYCLE AND CELL DIVISION 165
Cells at the end of prophase, when viewed under the microscope, do not show golgi complexes,
endoplasmic reticulum, nucleolus and the nuclear envelope.

Metaphase:
The complete disintegration of the nuclear envelope marks the start of the second phase of mitosis,
hence the chromosomes are spread through the cytoplasm of the cell. By this stage, condensation of
chromosomes is completed and they can be observed clearly under the microscope. This then, is the
stage at which morphology of chromosomes is most easily studied. At this stage, metaphase
chromosome is made up of two sister chromatids, which are held together by the centromere (Figure
10.2 b). Small disc-shaped structures at the surface of the centromeres are called kinetochores. These
structures serve as the sites of attachment of spindle fibres (formed by the spindle fibres) to the
chromosomes that are moved into position at the centre of the cell. Hence, the metaphase is
characterised by all the chromosomes coming to lie at the equator with one chromatid of each
chromosome connected by its kinetochore to spindle fibres from one pole and its sister chromatid
connected by its kinetochore to spindle fibres from the opposite pole (Figure 10.2 b). The plane of
alignment of the chromosomes at metaphase is referred to as the metaphase plate. The key features of
metaphase are: z Spindle fibres attach to kinetochores of chromosomes. z Chromosomes are moved to
spindle equator and get aligned along metaphase plate through spindle fibres to both poles.
Anaphase:
At the onset of anaphase, each chromosome arranged at the metaphase plate is split simultaneously
and the two daughter chromatids, now referred to as chromosomes of the future daughter nuclei, begin
their migration towards the two opposite poles. As each chromosome moves away from the equatorial
plate, the centromere of each chromosome is towards the pole and hence at the leading edge, with the
arms of the chromosome trailing behind (Figure 10.2 c). Thus, anaphase stage is characterised by Figure
10.2 a and b : A diagrammatic view of stages in mitosis 166 BIOLOGY the following key events:

z Centromeres split and chromatids separate.

z Chromatids move to opposite poles.
Telophase:
At the beginning of the final stage of mitosis, i.e., telophase, the chromosomes that have reached their
respective poles decondense and lose their individuality. The individual chromosomes can no longer be
seen and chromatin material tends to collect in a mass in the two poles (Figure 10.2 d). This is the stage
which shows the following key events:

z Chromosomes cluster at opposite spindle poles and their identity is lost as discrete elements.

z Nuclear envelope assembles around the chromosome clusters.

z Nucleolus, golgi complex and ER reform.

Cytokinesis:
Mitosis accomplishes not only the segregation of duplicated chromosomes into daughter nuclei
(karyokinesis), but the cell itself is divided into two daughter cells by a separate process called
cytokinesis at the end of which cell division is complete (Figure 10.2 e). In an animal cell, this is achieved
by the appearance of a furrow in the plasma membrane. The furrow gradually deepens and ultimately
joins in the centre dividing the cell cytoplasm into two. Plant cells however, are enclosed by a relatively
inextensible cell wall, thererfore they undergo cytokinesis by a different mechanism. In plant cells, wall
formation starts in the centre of the cell and grows outward to meet the existing lateral walls. The
formation of the new cell wall begins with the formation of a simple precursor, called the cell-plate that
represents the middle lamella between the walls of two adjacent cells. At the time of cytoplasmic
division, organelles like mitochondria and plastids get distributed between the two daughter cells. In
some organisms karyokinesis is not followed by cytokinesis as a result of which multinucleate condition
arises leading to the formation of syncytium

Significance of Mitosis :
Mitosis or the equational division is usually restricted to the diploid cells only. However, in some lower
plants and in some social insects haploid cells also divide by mitosis. It is very essential to understand
the significance of this division in the life of an organism. Are you aware of some examples where you
have studied about haploid and diploid insects? Mitosis results in the production of diploid daughter
cells with identical genetic complement usually. The growth of multicellular organisms is due to mitosis.
Cell growth results in disturbing the ratio between the nucleus and the cytoplasm. It therefore becomes
essential for the cell to divide to restore the nucleo-cytoplasmic ratio. A very significant contribution of
mitosis is cell repair. The cells of the upper layer of the epidermis, cells of the lining of the gut, and blood
cells are being constantly replaced. Mitotic divisions in the meristematic tissues – the apical and the
lateral cambium, result in a continuous growth of plants throughout their life.
MEIOSIS:
The production of offspring by sexual reproduction includes the fusion of two gametes, each with a
complete haploid set of chromosomes. Gametes are formed from specialised diploid cells. This
specialised kind of cell division that reduces the chromosome number by half results in the production
of haploid daughter cells. This kind of division is called meiosis. Meiosis ensures the production of
haploid phase in the life cycle of sexually reproducing organisms whereas fertilisation restores the
diploid phase. We come across meiosis during gametogenesis in plants and animals. This leads to the
formation of haploid gametes. The key features of meiosis are as follows:

z Meiosis involves two sequential cycles of nuclear and cell division called meiosis I and meiosis II but
only a single cycle of DNA replication.

z Meiosis I is initiated after the parental chromosomes have replicated to produce identical sister
chromatids at the S phase.

z Meiosis involves pairing of homologous chromosomes and recombination between them.

z Four haploid cells are formed at the end of meiosis II. Meiotic events can be grouped under the
following phases:

Meiosis I Meiosis II

Prophase I Prophase II
Metaphase I Metaphase II
Anaphase I Anaphase II
Telophase I Telophase II