Help Line Mob
9357388588
Notes
SVOENB Sec
Instructional Video Sec
MCQs


Cell cycle and cell division

Are you aware that all organisms, even the largest, start their life from a single cell? You may wonder how a single cell then goes on to form such large organisms. Growth and reproduction are characteristics of cells, indeed of all living organisms. All cells reproduce by dividing into two, with each parental cell giving rise to two daughter cells each time they divide. These newly formed daughter cells can themselves grow and divide, giving rise to a new cell population that is formed by the growth and division of a single parental cell and its progeny. In other words, such cycles of growth and division allow a single cell to form a structure consisting of millions of cells.

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). 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:
-Interphase
-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:

-G1 phase (Gap 1)
-S phase (Synthesis)
-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.


How do plants and animals continue to grow all their lives? Do all cells in a plant divide all the time? Do you think all cells continue to divide in all plants and animals? Can you tell the name and the location of tissues having cells that divide all their life in higher plants? Do animals have similar meristemat c tissues?

Some cells in the adult animals do not appear to exhibit division (e.g., heart cells) and many other cells divide only occasionally, as needed to replace cells that have been lost because of injury or cell death. These cells that do not divide further exit G1 phase to enter an inactive stage called quiescent stage (G0) of the cell cycle. Cells in this stage remain metabolically active but no longer proliferate unless called on to do so depending on the requirement of the organism.
In animals, mitotic cell division is only seen in the diploid somatic cells. However, there are few exceptions to this where haploid cells divide by mitosis, for example, male honey bees. Against this, the plants can show mitotic divisions in both haploid and diploid cells. From your recollection of examples of alternation of generations in plants (Chapter 3) identify plant species and stages at which mitosis is seen in haploid cells.
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 (karyokinesis), it is very essential to understand that cell division is a progressive process and very clear-cut lines cannot be drawn between various stages.

Karyokinesis involves following four stages:


-Prophase
-Metaphase
-Anaphase
-Telophase


Prophase

Prophase which is the first stage of karyokinesis 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 intertwined. 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 centrosome, 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:
-Chromosomal material condenses to form compact mitotic chromosomes. Chromosomes are seen to be composed of two chromatids attached together at the centromere.
- Centrosome which had undergone duplication during interphase, begins to move towards opposite poles of the cell. Each centrosome radiates out microtubules called asters. The two asters together with spindle fibres forms mitotic apparatus.

You have studied mitosis in onion root tip cells. It has 16 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?

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:
-Spindle fibres attach to kinetochores of chromosomes.
-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 daughter 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 remains directed 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 the following key events:
-Centromeres split and chromatids separate.
-Chromatids move to opposite poles.

Telophase

At the beginning of the final stage of karyokinesis, 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 each set of chromatin material tends to collect at each of the two poles (Figure 10.2 d). This is the stage which shows the following key events:
-Chromosomes cluster at opposite spindle poles and their identity is lost as discrete elements.
-Nuclear envelope develops around the chromosome clusters at each pole forming two daughter nuclei.
-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 the separation of cytoplasm called cytokinesis at the end of which cell division gets completed (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 (e.g., liquid endosperm in coconut).


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 usually results in the production of diploid daughter cells with identical genetic complement. 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: -Meiosis involves two sequential cycles of nuclear and cell division called meiosis I and meiosis II but only a single cycle of DNA replication.
-Meiosis I is initiated after the parental chromosomes have replicated to produce identical sister chromatids at the S phase.
- Meiosis involves pairing of homologous chromosomes and recombination between non-sister chromatids of homologous chromosomes.
- Four haploid cells are formed at the end of meiosis II.
Meiotic events can be grouped under the following phases: Meiosis I - Prophase I, Metaphase I, Anaphase I, Telophase I,
Meiosis II- Prophase II, Metaphase II, Anaphase II, Telophase II,

Meiosis I

Prophase I: Prophase of the first meiotic division is typically longer and more complex when compared to prophase of mitosis. It has been further subdivided into the following five phases based on chromosomal behaviour, i.e.,
Leptotene, Zygotene, Pachytene, Diplotene and Diakinesis.
During leptotene stage the chromosomes become gradually visible under the light microscope. The compaction of chromosomes continues throughout leptotene. This is followed by the second stage of prophase I called zygotene. During this stage chromosomes start pairing together and this process of association is called synapsis. Such paired chromosomes are called homologous chromosomes. Electron micrographs of this stage indicate that chromosome synapsis is accompanied by the formation of complex structure called synaptonemal complex. The complex formed by a pair of synapsed homologous chromosomes is called a bivalent or a tetrad. However, these are more clearly visible at the next stage. The first two stages of prophase I are relatively short-lived compared to the next stage that is pachytene. During this stage, the four chromatids of each bivalent chromosomes becomes distinct and clearly appears as tetrads. This stage is characterised by the appearance of recombination nodules, the sites at which crossing over occurs between non-sister chromatids of the homologous chromosomes. Crossing over is the exchange of genetic material between two homologous chromosomes. Crossing over is also an enzyme-mediated process and the enzyme involved is called recombinase. Crossing over leads to recombination of genetic material on the two chromosomes. Recombination between homologous chromosomes is completed by the end of pachytene, leaving the chromosomes linked at the sites of crossing over. The beginning of diplotene is recognised by the dissolution of the synaptonemal complex and the tendency of the recombined homologous chromosomes of the bivalents to separate from each other except at the sites of crossovers.

These X-shaped structures, are called chiasmata. In oocytes of some vertebrates, diplotene can last for months or years.

The final stage of meiotic prophase I is diakinesis.
This is marked by terminalisation of chiasmata. During this phase the chromosomes are fully condensed and the meiotic spindle is assembled to prepare the homologous chromosomes for separation. By the end of diakinesis, the nucleolus disappears and the nuclear envelope also breaks down. Diakinesis represents transition to metaphase.

Metaphase I:
-The bivalent chromosomes align on the equatorial plate
(Figure 10.3). -The microtubules from the opposite poles of the spindle attach to the kinetochore of homologous chromosomes.

Anaphase I:

The homologous chromosomes separate, while sister chromatids remain associated at their centromeres (Figure 10.3). Telophase I: The nuclear membrane and nucleolus reappear, cytokinesis follows and this is called as dyad of cells (Figure 10.3). Although in many cases the chromosomes do undergo some dispersion, they do not reach the extremely extended state of the interphase nucleus. The stage between the two meiotic divisions is called interkinesis and is generally short lived. There is no replication of DNA during interkinesis. Interkinesis is followed by prophase II, a much simpler prophase than prophase I.


Meiosis II

Prophase II: Meiosis II is initiated immediately after cytokinesis, usually before the chromosomes have fully elongated. In contrast to meiosis I, meiosis II resembles a normal mitosis. The nuclear membrane disappears by the end of prophase II (Figure 10.4). The chromosomes again become compact.
Metaphase II: At this stage the chromosomes align at the equator and the microtubules from opposite poles of the spindle get attached to the kinetochores (Figure 10.4) of sister chromatids.
Anaphase II: It begins with the simultaneous splitting of the centromere of each chromosome (which was holding the sister chromatids together), allowing them to move toward opposite poles of the cell (Figure 10.4) by shortening of microtubules attached to kinetochores.
Telophase II: Meiosis ends with telophase II, in which the two groups of chromosomes once again get enclosed by a nuclear envelope; cytokinesis follows resulting in the formation of tetrad of cells i.e., four haploid daughter cells (Figure 10.4).

Significance of meiosis

Meiosis is the mechanism by which conservation of specific chromosome number of each species is achieved across generations in sexually reproducing organisms, even though the process, per se, paradoxically, results in reduction of chromosome number by half. It also increases the genetic variability in the population of organisms from one generation to the next. Variations are very important for the process of evolution.

Exercises
1. What is the average cell cycle span for a mammalian cell?
2. Distinguish cytokinesis from karyokinesis.
3. Describe the events taking place during interphase.
4. What is Go (quiescent phase) of cell cycle?
5. Why is mitosis called equational division?
6. Name the stage of cell cycle at which one of the following events occur:
(i) Chromosomes are moved to spindle equator.
(ii) Centromere splits and chromatids separate.
(iii) Pairing between homologous chromosomes takes place.
(iv) Crossing over between homologous chromosomes takes place.
7. Describe the following:
(a) synapsis (b) bivalent (c) chiasmata
Draw a diagram to illustrate your answer.
8. How does cytokinesis in plant cells differ from that in animal cells?
9. Find examples where the four daughter cells from meiosis are equal in size and where they are found unequal in size.
10. Distinguish anaphase of mitosis from anaphase I of meiosis.
11. List the main differences between mitosis and meiosis.
12. What is the significance of meiosis?
13. Discuss with your teacher about
(i) haploid insects and lower plants where cell-division occurs, and
(ii) some haploid cells in higher plants where cell-division does not occur.
14. Can there be mitosis without DNA replication in ‘S’ phase?
15. Can there be DNA replication without cell division?
16. Analyse the events during every stage of cell cycle and notice how the following two parameters change
(i) number of chromosomes (N) per cell
(ii) amount of DNA content (C) per cell


Go back -top
MCQs
SV021110.0012020 Cell Cycle

021110.001. A cell cycle is :
(A) the time from the formation of a cell until its death. (B) the series of events that takes place from the formation of a cell until it divides again. (C) the sequence of events that assures each daughter cell of a set of chromosomes identical with that of its parent cell (mitosis) (D) the growth of a cell until it is large enough to divide again.

B

021110.002. A diploid somatic cell :
(A) cannot undergo division again (B) can undergo mitosis but not meiosis (C) can undergo mitosis or meiosis (D) can undergo meiosis but not mitosis.

B

021110.003. A cell is in metaphase if :
(A) its chromosomes are visible as distinct thread-like structures (B) the nuclear membrane is not visible (C) the chromosomes are lined up on the equator of the spindle (D)the chromosomes are separated into two distinct groups attached to the spindle (E) the chromosomes are found in two compact groups in two small patches of cytoplasm that are in the process of separating into two distinct cells.

C

021110.004. A cell inprophase of mitosis can be distinguished froma Cell in prophase I of meiosis by :
(A)the presence of only half as many chromosomes in the meiotic cell (B) the formation of tetrads in the meiotic cell (C) the presence of twice as many chromosomes in the meiotic cell. (D)None of the above.

B

021110.005. The function of mitotic cell division in the life history of an organism is :
(A) reproduction of identical individuals if the organism is unicellular (B) growth of an individual if the organism is multicellular (C) repair of injured tissue (D) All of the above.

D

021110.006. Substances that interfere with microtubule function interfere with cell division because:
(A)microtubules must be distributed equally to the new cells (B) microtubules are involved in the precise separation of the chromosomes which ensures that into each daughter cell (C) without microtubules, cytokinesis cannot take place and syncytium is formed (D) microtubules are essential for the disappearance of the nuclear membrane, and without them the chromosomes have to stay too close together within the nuclear membrane to be able to separate into two new nuclei.

B

021110.007. The chief significance of mitosis is that it :
(A)ensures genetic homogeneity of the cells (B) reduces the chromosome number to half (C) causes transformation of DNA to RNA (D) increases the water contents of the cell.

A

021110.008. Movement of chromosomes during anaphase is :
(A) dependent on association of spindle fibres with kinetochore. (B) independent of spindle fibres (C) due to cytoplasmic streaming (D) due to excess of ATP formed in mitochondria.

A

021110.009. Cell would normally proceed to mitosis without interruption :
(A)once it has entered the S-period (B) once it has entered the Grperiod (C) at any time during cell division activity (D)None of the above.

A

021110.010. In a cell number of chromosomes after first meiosis is 44. The number of chromosomes in its daughter cells after completion of meiosis is :
(A) 44 (B) 22 (C) 11 (D) 66.

A

021110.011. Mechanism ensuring continuity in mitosis is :
(A) crossing over (B) halfing the chromosome number between the two daughter cells (C) segregation of paternal and maternal chromosome (D) formation of two daughter cells with identical DNA.

D

021110.012. Which is the most active stage of cell cycle in terms of gene action ?
(A) Metaphase (B) Anaphase (C) Telophase (D) S-phase of interphase.

D

021110.013. The cause of chromosome laggards (laggards) during meiosis is:
(A) dicentric chromosome (B) inversion (C) acentric chromosome (D) duplication of a gene.

C

021110.014. An angiospermic plant has 12 chromosomes in its pollen grains. How many chromosomes will it have in its endosperm ?
(A)48 (B)36 (C) 24 (D) 12.

B

021110.015. The synaptonemal complex is a structure visible from:
(A)diplotene through metaphase I (B) zygotene through pachytene (C) pachytene through diplotene (D)leptotene through diplotene.

B

021110.016. The best stage of mitosis for studying chromosome morphology is : (A) metaphase (B) anaphase (C) interphase (D) telophase

A

021110.017. How many meiosis shall be required to produce 100 grains of wheat?
(A) 150 (B) 400 (C) 100 (D) 125.

D

021110.018. In an adult human being, growth and cell division stop in: (A) skin cells (B) bone marrow (C) testes and ovary (D) bone cells.

D

021110.019. Phragmoplast is the :
(A) cell plate formed during cytokinesis of plant cells (B) cell membrane formed during cytokinesis of animal cells (C) plastid which helps in cytokinesis (0) plastid capable of self-duplication

A

021110.020. Which pair of the following are known for chromosome theory of heredity? (A) Morgan and Bridges (B) Boveri and Beadle (C) Sutton and Boveri (D) Morgan and Boveri

C

021110.021. During cell division one of the cytoplasmic organelles plays an important role, it is :
(A) mitochondria (B) centriole (C) golgi complex (D) is not precise one. B
021110.022. Each longitudinal half of a chromosome of late prophase is known as: (A) chromatin (B) chromatin fibre (C) chromatid (D) half chromatid.
C
021110.023. As a cell proceeds towards mitosis, it can be halted by depriving the cell of oxygen or by poisoning its oxidative enzyme with carbon monoxide. What does this indicate? (A) Mitosis is a type of cellular respiration (B) Mitosis is an enzyme controlled event (C) Mitosis involves an expenditure of energy (D) Mitosis yields energy that the cell uses to make ATP molecules
C
021110.024. During cell division one of the cytoplasmic organelles plays an important role, it is :
(A) mitochondria (B) centriole (C) golgi complex (D) is not precise one.
B
021110.025. Each longitudinal half of a chromosome of late prophase is known as:
(A) chromatin (B) chromatin fibre (C) chromatid (D) half chromatid.
C
021110.026. As a cell proceeds towards mitosis, it can be halted by depriving the cell of oxygen or by poisoning its oxidative enzyme with carbon monoxide. What does this indicate? (A) Mitosis is a type of cellular respiration (B) Mitosis is an enzyme controlled event (C) Mitosis involves an expenditure of energy (D) Mitosis yields energy that the cell uses to make ATP molecules
C
021110.027. Each longitudinal half of a chromosome of late prophase is known as:
(A) chromatin (B) chromatin fibre (C) chromatid (D) half chromatid.
C
021110.028. The spindle fibre during cell division arises from:
(A) centromere (B) nucleus (C) aster (D) mitochondria.
C
021110.029. Mitosis consists of :
(A) a series of nuclear division (B) cleavage of cytoplasm (C) karyokinesis followed by cytokinesis (D) formation of daughter nuclei
C
021110.030. During metaphase :
(A) chromosomes occupy equatorial zone (B) chromosomes are not yet ready for division (C) chromosomes orient themselves into any place on the mitotic spindle (D) chromosomes assemble at one of the poles of mitotic spindle.
A
021110.031. The points where the crossing over takes place between chromatids of two homologous chromosomes of a pair are: (A) centromere (B) chiasmata (C) chromomere (D) chromocentre
B
021110.032. Which of the following is capable of self-reproduction in cell division ?
(A) protein molecule (B) Fat molecule (C) Carbohydrate molecule (D) DNA molecule.
D
021110.033. Mitosis and meiosis differ because during meiosis: (A) homologous chromosomes pair and exchange parts (B) chromosome number is halved (C) four nuclei formed are not similar (D) All the above.
D
021110.034. During cell division nuclear membrane reappears in : (A) interphase (B) prophase (C) telophase (D) S-phase.
C
021110.035. In which stage of mitosis, all the chromosomes are arranged on equatorial plate ? (A) Prophase (B) Metaphase (C) Anaphase (D) Telophase.
B
021110.036. During prophase the chromosomes are : (A) larger and coiled (B) much contracted, thickened and coiled (C) much contracted, thickened and uncoiled (D) larger and straight.
B
021110.037. Cell division or mitosis is a normal process in living cells but sudden and abnormal mitosis in an organ results in : (A) new organ (B) new organism (C) zygote (D) cancer.
D
021110.038. Meiosis is a type of cell division in which: (A) the chromosome number is reduced to half (B) the chromosome number remains the same (C) the chromosome number is doubled (D) the chromosome number is reduced to one fourth.
A 021110.039. In which one of the following stages chromosomes appear as long thin threads in :
(A) leptotene (B) zygotene (C) pachytene (D) diplotene. A
021110.040. During Meiosis 1, in which stage synapsis takes place?
(A)Pachytene (B) Zygotene (C) Diplotene (D) Leptotene.
B
021110.041. One of the following types of cell division results in reduction of chromosomes :
(A) mitosis (B) binary fission (C) meiosis (D) amitosis.
C
021110.042. Chiasmata are formed by exchange between non-sister chromatids of homologous chromosome was first indicated by :
(A) Altman (B) Sax (C) Janssen (D) T.H. Morgan.
C
021110.043. How many times the mitotic division must occur in the cell of root tip to form 256 cells ?
(A) 128 (B) 255 (C) 64 (D) 8.
D
021110.044. When a diploid cell undergoes meiosis :
(A) Homologous chromosomes segregate (B) All dominant genes segregate into daughter Cells (C) Homologous chromosomes pair but do not separate (D) All linkage groups get distributed
A
021110.045. Amitosis is a usual process of cell division in :
(A) eukaryotic cells (B) prokaryotic cells (C) spore mother cells (D) meristematic cells.
B


1110 Notes

Cell Division


Cell Cycle (Howard and Pelc, 1953)

Cell cycle is a series of programmed cyclic changes by which the cell duplicates its contents and divides into two. The sequence of events is genetically controlled.
Time interval between two successive divisions is called generation time. Cell cycle has two parts-long non dividing I-phase and short dividing M-phase.
cell cycle is defined as the period between successive divisions of a cell. Interphase (I-phase).

It is complex of changes that occurs in a newly formed cell before it is able to divide. Interphase is also called intermitosis, preparatory phase or energy phase.
Previously it was known as resting phase.
The nondividing state of mature cell/nucleus is also called interphase.
It lasts throughout the life of the cell.
For example, nerve cells do not divide after birth and, therefore, last throughout the life of the individual.
Inter mitotic interphase is the period of intense biosynthetic activity wherein the cell doubles its size and duplicates its chromosome complement along with the increase in number of various cell organelles.

Interphase also accomplishes three important processes which are preparatory to cell division:
(a) Replication of DNA and synthesis of nuclear proteins such as histones.
(b) Duplication of centriole in animal cells. Daughter centriole develops at right angle to parent centriole.
(c) Synthesis of energy rich components for providing energy.

Interphase has three stages.
(i) G1-Phase (First Growth or Gap Phase, Pre-Synthetic Phase, Post-Mitotic Phase). It is the longest phase of cell cycle. Cell and its nucleus (to a lesser extent) grow in size.
There is synthesis of biochemicals like RNAs, proteins, enzymes for DNA synthesis, amino acids for histone formation, nucleotides and ATP. Cell organelles also increase in number.
Duration of G1 phase is longer in cells dividing infrequently. It is shorter in frequently dividing cells. In G1 phase a cell has three options.
The G0 phase is a period in the cell cycle in which cells exist in a quiescent state. G0 phase is viewed as either an extended G1 phase, where the cell is neither dividing nor preparing to divide, or a distinct quiescent stage that occurs outside of the cell cycle.
(a) Continue cycle and enter S-phase.
(b) Stop cell cycle and enter Go phase for undergoing differentiation.
(c) Get arrested in G1 phase from where it may enter cell cycle or pass into G0 phase.
(Internal Checkpoints During the Cell Cycle: The cell cycle is controlled at three checkpoints.
The integrity of the DNA is assessed at the G1 checkpoint. Proper chromosome duplication is assessed at the G2 checkpoint.
Attachment of each kinetochore to a spindle fiber is assessed at the M checkpoint. Cyclins are a family of proteins that control the progression of cells through the cell cycle by activating cyclin-dependent kinase (Cdk) enzymes.
Through phosphorylation, Cdks signal the cell that it is ready to pass into the next stage of the cell cycle.
As their name suggests, Cyclin-Dependent Protein Kinases are dependent on cyclins, another class of regulatory proteins.
Cyclins bind to Cdks, activating the Cdks to phosphorylate other molecules. A cyclin-dependent kinase complex (CDKC, cyclin-CDK) is a protein complex formed by the association of an inactive catalytic subunit of a protein kinase, cyclin-dependent kinase (CDK), with a regulatory subunit, cyclin. Once cyclin-dependent kinases bind to cyclin, the formed complex is in an activated state.)
The deciding factor is storage of energy rich compounds and availability of mitogens. The stage where this decision is made can be called check point.
Once this check point is crossed, the cell reaches a state called ante phase (Bullough, 1952) whereby it will divide even under unfavourable conditions. There is abundant storage of ATP energy.

(ii) S-Phase (Synthetic Phase). DNA of each chromosome replicates followed by synthesis of histones. As a result each chromosome undergoes replication producing two chromatids.
S-phase is also known as invisible stage of M-phase. Synthesis of other biochemicals also continues. Centrioles replicate towards the end of S-phase or beginning of G2-phase.

(iii) G2-Phase (Second Growth or Gap Phase, Post-Synthetic Phase, Pre-mitotic Phase).
There is again intense synthesis of proteins and RNAs. Cell organelles increase in number. Protein tubulin is formed.
G0-Phase. It is phase of cell differentiation when cell cycle is stopped about the middle of G1 Phase due to activation of certain genes.
The genes allow the cell to grow to a particular size, assume particular shape and come to perform certain specific functions.
Various phases of cell cycle are controlled by proteins cyclins and kinases that take part in phosphorylation and dephosphorylation.

M-Phase. It is the phase of cell division. Cell division consists of nuclear division or karyokinesis and protoplast division or cytokinesis.
Cell division or M-phase is of three types - amitosis, mitosis and meiosis.

AMITOSIS (Direct Division; Robert Remak, 1855)
It was discovered by Remak (1841, 1855) and studied by Flemming (1882).
In amitosis, nucleus elongates, constricts in the middle and divides directly into two daughter nuclei. Chromatin does not condense to form chromosomes.
Spindle is not produced. Nuclear division is followed by cytokinesis or division of cytoplasm. It occurs through cleavage or constriction, e.g., cartilage cells, degenerate cells, meganucleus of Paramecium, cells of foetal membranes and endosperm.
Some workers include procaryotic cell division under amitosis. As amitosis does not distribute chromatin equitably, it leads to structural and functional irregularities.

Mitosis

It is also called somatic division because it occurs during formation of somatic or body cells.
Mitosis is studied in plants in the regions of meristems, e.g., stem tip, root tip (Onion 2n = 16).
In animals it is studied in bone marrow, skin, base of nails, etc.
Mitosis is equational division in which a parent cell divides into two identical daughter cells, each of which contains the same number and kind of chromosomes as are present in the parent cell.
It is centric in animal cells and acentric (without participation of centrosome) in plants.
Mitosis occurs in two steps, karyokinesis and cytokinesis.

Karyokinesis. It is the stage of nuclear division (indirect nuclear division) which is continuous but is divided into four stages for the sake of convenience –
prophase,
metaphase,
anaphase and
telophase.

1. Prophase.
It is the longest phase of karyokinesis. In early prophase or spireme stage,
the chromatin fibres condense through spiralisation to form elongated chromosomes.
There is increased viscosity and refractivity of cytoplasm.
Animal cells become nearly rounded. The ends of chromosomes are not distinguishable.
The nucleus appears as a ball of wool.
Centrosome has already divided. The daughter centrosomes begin to move away from each other.
In mid prophase, chromosomes shorten and become distinct with each having two chromatids attached to narrow point called centromere.
The centrosomes develop astral rays and migrate farther. In late prophase, the centrosomes reach the poles, form asters and begin to develop spindle fibres.
Nucleolus degenerates and nuclear envelope starts breaking.
In plant cells, centrosome are absent. Spindle fibres develop without them. Polar ends are negatively charged.
They are organised in both plants and animals with the help of Ca2+ containing protein calmodulin.

Prometaphase. It is considered to be second phase by certain cytologists. Metaphase is then called third stage of karyokinesis.
During this stage the nuclear envelope disappears so that nucleoplasm comes in contact with cytoplasm.
A differentiation between the two disappears. A fluid area occurs in the centre of the cell. Chromosomes can move freely in this area.

Spindle apparatus or mitotic apparatus gets organiscd. It is spindle-shaped colourless fibrous body which can be observcd with the help of pol arising microscope.
A spindle fibre consists of 4 - 20 microtubules formed of protein tubulin.


Spindle fibres converge at the two ends or poles.
Spindle has the maximum diameter in the middle. It is called equator. In dividing animal cells, the spindle bears asters at the two poles. Such a spindle is called amphiaster or centric.
In plant cells spindle is acentric or anastral. Fibres of the spindle are of two main types,
continuous (from pole to pole) and discontinuous (radiating from one pole but not reaching the other).
The term eumitosis or extranuclear mitosis is used for organisation of spindle in which nuclear envelope degenerates.
In many protists, fungi and algae, the nuclear envelope does not degenerate during mitosis. Instead, spindle is formed inside the nucleus. It is called intranuclear mitosis or premitosis.

2. Metaphase.
Chromosomes are the shortest and the thickest in metaphase.
Each chromosome gets attached to two discontinuous spindle fibres or chromosome fibres, one from each pole, in the region of its kinetochores (a complex of proteins associated with the centromere of a chromosome during cell division, to which the microtubules of the spindle attach.). The latter have plus ends for this purpose.
Chromosome fibres contract and bring the chromosomes over the equator. The phenomenon is called congression. Centromeres of all the chromosomes are present over the equator. Therefore, chromosomes form a sort of apparent plate called equatorial or metaphasic plate. Limbs of chromosomes project in different directions. Metaphase is the best time to see chromosomes.

3. Anaphase.

It is the phase of shortest duration.
APC (anaphase promoting complex) develops. It degenerates proteins binding the two chromatids in the region of centromere. As a result, the centromere of each chromosome divides.
This converts the two chromatids into daughter chromosomes each being attached to the spindle pole of its side by independent chromosomal fibre.
The chromosomes move towards the spindle poles with the centromeres projecting towards the poles and the limbs trailing behind.
There is corresponding' shortening of chromosome fibres. Anaphasic chromosomes appear differently depending upon the position of their centromeres - V -shaped (metacentric), L-shaped (submetacentric), J-shaped (acrocentric) and I-shaped (telocentric). The two pole-ward 'moving chromosomes of each type remain attached to each other by interzonal fibres. Ultimately, two groups of chromosomes come to lie at the spindle poles.

4. Telophase.
It is the stage of reconstitution of nuclei.
Spindle fibres converge at the two ends or poles.
Spindle has the maximum diameter in the middle. It is called equator. In dividing animal cells, the spindle bears asters at the two poles. Such a spindle is called amphiaster or centric.
In plant cells spindle is acentric or anastral. Fibres of the spindle are of two main types,
continuous (from pole to pole) and discontinuous (radiating from one pole but not reaching the other).
The term eumitosis or extranuclear mitosis is used for organisation of spindle in which nuclear envelope degenerates.
In many protists, fungi and algae, the nuclear envelope does not degenerate during mitosis. Instead, spindle is formed inside the nucleus. It is called intranuclear mitosis or premitosis.
Chromosomes uncoil, elongate, lose their stainability and form chromatin fibres.
Nucleolar organisers form nucleoli in one or more pairs of chromosomes.

Nucleoplasm and nuclear envelope appear so that two daughter nuclei are formed.

Dinomitosis.
It is type of nuclear division found in dinoflgellates in which the nuclear envelope persists. Microtubular spindle is not formed.
Chromosomes move while attached to inner membrane of nuclear envelope.
Karyochorisis. Intranuclear spindle is formed with spindle pole bodies (SPBs) developing at the two ends.

Cytokinesis
Cytokinesis is the division of cytoplasm of a cell undergoing karyokinesis to form two daughter cells.
I t begins towards the middle of anaphase and is completed simultaneously with the completion of telophase.
Non occurrence of cytokinesis produces multinucleate coenocyte or syncytium. Cytokinesis also brings about nearly equitable distribution of various cell organelles like mitochondria, plastids, lysosomes, Golgi apparatus, endoplasmic reticulum, cytoplasmic matrix, etc. Mitochondria and plastids are known to multiply through fission.
Details of multiplication of other organelles are not -yet known.


Cytokinesis is of two types, cleavage and cell plate.

Cleavage Cytokinesis.
A mid body of dense vesicular and fibrous material appears in the equatorial region.
Microfilaments appear in the peripheral region.
Their activity constricts the cell membraY1e. As a result a centripetal furrow or cleavage develops in the middle.
The furrow deepens and divides the parent protoplast into two uninucleate protoplasts or cells. Cleavage is the usual method of cytokinesis in animals.
It also occurs in some lower plants where wall material is deposited in the furrow between the two daughter protoplasts.
Cell Plate Cytokinesis.
It occurs in plants. Middle part of spindle persists.
It gets interdigited with microtubules to form a complex structure called phragmoplast. Vesicles having pectic compounds and other wall materials appear in the middle of phragmoplast. They fuse and form a film or cell plate with membrane on either side.
This divides the parent binucleate cell into two daughter uninucleate cells.
Cell plate grows centrifugally and functions as middle lamella. Primary wall is deposited on its either side by the two daughter protoplasts.

Importance.

(i) Growth. Mitosis is essential for formation of new cells required for growth of all multicellular organisms. A human neonate has 6 X 1012 cells all derived from a single celled zygote. The adult human body has 1014 cells.
(ii) Repair. Old and worn out cells are regularly replaced. The new cells are formed through mitosis. It is estimated that daily 5 X 109 cells are being lost and replaced in the upper layer of epidermis, lining of gut, RBCs and WBCs.
(iii) Maintenance of Surface Volume Ratio.
(iv) Maintenace of Nucleocytoplasmic Ratio.
(v) Reproduction. Mitosis is a method of multiplication in unicellular organisms
(vi) Opportunity for Differentiation. Multicellular condition provides opportunity for differentiation and division of labour
(vii) Maintenance of Chromosome Number. It maintains similar number and type. of chromosomes in all the cells. The latter is important in healing, asexual reproduction, replacement of worn out cells and regeneration.
(viii) Somatic Mutations. It maintans somatic mutations by vegetative reproduction, e.g., Navel Orange.

00
Meiosis
1. It occurs only once in the life cycle. The cells undergoing meiosis are called meiocytes (e.g., oocytes, spermatocytes, microsporocytes, megasporocytes).
2. Meiosis is a double division in which a diploid cell divides twice to form four haploid cells. Interphase is single so that DNA or chromosome replication occurs once. It changes single stranded or monad chromosome into double stranded chromosome.
3. Meiosis can be
(a) gametic (occurring at the time of gamete formation, e.g., animals),
(b) zygotic (occurring at the time of zygote or zygospore germination, e.g., Ulothrix, Spirogyra, Chlamydomonas) and
(c) sporic (at the time of microspore and megaspore formation, e.g., most plants comprising bryophytes, pteridophytes, gymnosperms and angiosperms).
Meiosis is studied in anthers of unopencd flowersl buds and testis of Grasshopper.
Interphase
It is a stage prior to karyokinesis.
Interphase has three phases -G1, S and G2. Chromosomes replicate during S-phase except for small parts (0.3% of total which occurs during zygotene).
Size of nucleus increases to about 3 times. Centrosome replicates in G2 phase which is short.
M-phase has two steps or divisions, meiosis I and
meiosis II.
The essential processes that occur in meiosis are:
(i) Two successive divisions without replication of DNA in the period of interkinesis.
(ii) Crossing over and chiasma formation.
(iii) Segregation of sister chromatids.

Meiosis I

It is the actual reduction division which is also called heterotypic division because it brings about change from diploid to haploid state.
In meiosis I, the two chromatids of a chromosome often become different due to crossing over.

Prophase I. It is long, complex and divisible into five stages –
leptotene,
zygotene,
pachytene,
diplotene and
diakinesis.

(i) Leptotene (Leptonema).
Chromatin fibres condense and form chromosomes.
The chromosomes often show chromomeres.
They are attached to nuclear envelope by both of their ends through a specialised structure called attachment plate.
They may also develop basket-like arrangement called bouquet stage (diverging from a common point lying near centrosome).
Chromosome number is diploid where there are two chromosomes of each type called homologous chromosomes.
Their chromatids are not clear because of the development of nucleoprotein core between them.

(ii) Zygotene (Zygonema).
Homologous chromosomes join laterally in the process of synapsis (Montgomery, 1901) syndesis to form bivalents.
Number of bivalents is half the number of individual chromosomes.
Bivalents are actually tetrads but the individual chromatids of the two chromosomes are not clear due to the presence of nucleoprotein core between them.
Depending upon the area of initiation, synapsis can be procentric (starting from centromeres),
proterminal (pairing beginning from telomeric regions and proceeding inwards) and intermediate (= random pairing starts at several points).
Pairing proceeds from the starting regions towards other parts in a zipper like manner.
It brings alleles of the two homologous chromosomes exactly opposite each other.
The two chromosomes of a bivalent are held together by nucleoprotein core.
The whole structure is called synaptinemal or synaptonemal complex (Moses, 1956).

(iii) Pachytene (= Pachynema).
Bivalents may remain in pachytene stage for days.
Dense areas or recombination nodules (Zickler, 1977), having multienzymes, appear here and there over the bivalents.
There is breakage and re-union of chromatid segments.
At places it results in exchange of segments between nonsister chromatids of a bivalent.
The phenomenon is called crossing over. It brings about chromosome recombinations.
However, individual chromatids are not clear except towards the end of pachytene when synaptinemal complex begins to dissolve.

(iv) Diplotene (Diplonema). At most places synaptinemal complex dissolves.
Chromatids become clear and the bivalents are now called tetrads.
At places homologous chromosomes remain attached to each other. The points of attachment between nonsister chromatids of two homologous chromosomes are called chiasmata.
They are sites of previous crossing over where synaptinemal cdmplex persists. Chiasmata were first seen by Johannsen (1909).
In animal oocytes, diplotene stage is often prolonged. In oocytes of many fishes, amphibians, reptiles and birds,
the diplotene chromosomes decondense, elongate and form lampbrush chromosomes. They take part in rapid synthesis of various RNAs.

(v) Diakinesis. Chiasmata shift towards the chromosome ends (terminalisation). RNA synthesis stops. Nucleolus degenerates.
Nuclear envelope breaks at places. A spindle begins to develop, with (in animals) or without centrioles.

Metaphase I.

A bipolar fibrous spindle appears in the area of nucleus. It has asters at the two poles in animal cells (amphiaster) while the same are absent in plant cells (anastral).
The chromosome pairs get attached to spindle poles by discontinuous fibres. In a pair, each chromosome is attached to only one spindle pole of its side. Congression brings the bivalents/tetrads over the equator of the spindle.
The chromosome limbs lie over the equator while the centromeres are projected outwardly towards the poles.
On the equator, the chromosome bivalents form an apparent double whorl or double metaphasic plate.

Anaphase I.
Chiasmata disappear completely and the homologous chromosomes separate. The process is called disjunction.
The separated chromosomes (univalents) show divergent chromatids and are called dyads.
They move towards the spindle poles and ultimately form two groups of haploid chromosomes.

Telophase I.
Chromosomes elongate. Nucleoplasm and nuclear envelope appear over each chromosome group forming nuclei. Nucleolus is rarely formed.
Interkinesis.
It is brief intrameiotic interphase, which is found in some cases in order to synthesise deficient biochemicals. DNA synthesis does not occur.

Significance.
Meiosis I gives stimulus for formation of gametes or spores.
It reduces the chromosome number to half, performs random separation of paternal and maternal chromosomes, shows gene recombinations due to crossing over and produces occasional aberration due to non¬disjunction.


Meiosis II

It is homotypic or equational division which is meant for '"maintaining the haploid number, converting dyad chromosome state into monad state and separating the two chromatids of a chromosome which have become different due to crossing over.
DNA replication is absent.
Prophase II. The chromatin fibres shorten to form chromosomes.
Nucleolus and nuclear envelope break down. Spindle is formed in the area of each nucleus.
Both telophase I and prophase II are omitted in some organisms where anaphase I directly leads to metaphase II, e.g., Trillium.
Metaphase II.
Chromosomes come to lie at the equator of the spindle forming a single metaphasic plate.
The centromere of each chromosome gets attached by both its surfaces to the spindle poles of their sides by distinct chromosome fibres.
Anaphase II.
Centromere of each chromosome divides into two.
This separates the two chromatids of a chromoosome into two independent daughter chromosomes.
Each daughter chromosome is attached to spindle pole of its side by a chromosome fibre.
Chromosomes move towards the spindle poles forming two groups. Since there were two spindles, a total of four groups are formed.
Telophase II.
The four groups of chromosomes organise themselves into four haploid nuclei. Chromosomes de condense and elongated to return to interphase condition.
Nuclear envelope is formed from remains of old nuclear envelope and endoplasmic reticulum.
Nucleolus develops from NOR of certain chromosomes. It is caused by formation of rRNA and its association of ribosomal proteins.
Cytokinesis.
Cytokinesis may occur after each division (successive type) or simultaneously at the end of meiosis. It is generally through cleavage.
In case of plants, wall material is deposited in the furrows. Cytokinesis gives rise to four haploid cells.

Importance.
(i) Variations. Meiosis produces a lot of variations due to
(a) independent assortment of chromosomes
(b) crossing over (c) irregular disjunction
(d) gene mutations during replication and nicking for crossing over
(ii) Polyploidy. Failure of chromosomes to separate during anaphase I leads to polyploidy.
(iii) Maintenance of chromosome number generation after generation.
(iv) Sexual Reproduction. It produces gamete forming structures and is, therefore, essential for sexual reproduction.
Spindle fibres converge at the two ends or poles.
Spindle has the maximum diameter in the middle. It is called equator. In dividing animal cells, the spindle bears asters at the two poles. Such a spindle is called amphiaster or centric.
In plant cells spindle is acentric or anastral. Fibres of the spindle are of two main types,
continuous (from pole to pole) and discontinuous (radiating from one pole but not reaching the other).
The term eumitosis or extranuclear mitosis is used for organisation of spindle in which nuclear envelope degenerates.
In many protists, fungi and algae, the nuclear envelope does not degenerate during mitosis. Instead, spindle is formed inside the nucleus. It is called intranuclear mitosis or premitosis.

00
5091 509