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C ell cycle and cell division CELL CYCLE AND CELL DIVISION Chapter Outline: • • • • • • • • Prerequisites Learning objectives Cell Cycle M Phase Significance of Mitosis Meiosis Significance of Meiosis Summary www.sciencetuts.com C ell cycle and cell division Prerequisites Hi children, this is an interesting and familiar topic which we would like to know, the following facts as prerequisites: 1. Growth is a fundamental feature of living organisms. 2. In unicellular organisms growth includes only cell enlargement. 3. In multicellular organisms growth includes cell division and cell elongation. 4. The life of multicellular organisms begins with a single celled zygote. 5. The zygote divides repeatedly and forms thousands of cells which aggregate to form different plant organs. Some of the cells retain the capacity to divide. Such cells constitute a meristem. Therefore in a plant, cell division is confined only to meristematic cells. 6. Cell division is the most important method of formation of new cells. New cells always arise from pre existing cells. Learning objectives Before discussing this present chapter, we would like to know the concept of cell cycle and cell division that how it plays an important role in our living system. This is a wonderful creation among the nature that a single cell forms a structure, consisting of millions of cells. Let us gain the knowledge over this concept as following learning objectives. 1. To understand and explain the concept of cell cycle as (a) interphase and (b) M-phase. 2. To understand the process of cell division and its significance. 3. To comprehend the process of interphase which is further proceeds as G1 phase,S- phase and M-phase. 4. To recognize the process of mitosis in which nucleus divides first and is followed by the division of cytoplasm. 5. To recognize and recall the different phases of karyokinesis in mitosis are (1) prophase (2) metaphase (3) Anaphase and (4) Telophase. 6. To identify and explain the process of Meiosis (or) reduction division. 7. To recognize and recall the following terminology in meiosis-l and meiosis-ll as Cytokinesis, Karyokinesis, Prophase-l and ll, Metaphase-l and ll, Anaphase-l and ll, Telophase-l and ll, Leptotene, Zygotene, Pachytene, Diplotene, Diakinesis, Synaptonemal complex, Bivalent, Chiasmata etc. 8. To know the importance of crossing over. 9. To explain the difference between mitotic & meiotic cell division. 2 www.sciencetuts.com C ell cycle and cell division Cell Division is of 2 types 1) Direct cell division(Amitosis) 2) Indirect cell division Direct cell divisions: Budding Furrowing Indirect cell division: Mitosis Meiosis DIRECT CELL DIVISION OR AMITOSIS In direct cell division the cell divides directly by furrowing or budding into 2 to many cells. Amitosis is a type of cell division which involves no spindle formation. The nucleus divides directly into two. The division of nucleus is followed by cytoplasmic division. This type of division is common in Bacteria, Amoeba & Yeast. They contain naked circular DNA molecule. The nucleus elongates and becomes dumbbell shaped. As the constriction deepens the nucleus is divided into two daughter nuclei. The nuclei may or may not be equal. Amoeba Amoeba INDIRECT CELL DIVISION In indirect cell division the cell divides systematically to produce 2 - 4 daughter cells. Spindle apparatus is formed. Chromosomes appear. The daughter cells are equal in all respects. The indirect cell division is of 2 types. They are Mitosis and Meiosis. Mitosis Indirect Cell Division 3 www.sciencetuts.com C ell cycle and cell division Cell Cycle The sequence of events by which a cell duplicates its genome, synthesizes the other constituents of the cell and eventually divides into two daughter cells is termed as cell cycle. Although cell growth is a continuous process, DNA synthesis occurs only during one specific stage in the cell cycle. The replicated chromosomes are then distributed to daughter nuclei by a complex series of events during cell division. These events are under genetic control. PHASES OF CELL CYCLE The duration of cell cycle varies from organism to organism and also from cell to cell type. The cell cycle is divided into 2 phases. A) Interphase B) M - Phase (Mitotic phase) INTERPHASE Phases Of Cell Cycle The stage of cell cycle in which the nucleus is not in a state of division is called Interphase. The Cell spends most of its life cycle in Interphase. Interphase is the period of interval between two successive cell divisions. Though it is called as resting phase, it is not a period of rest but a period of great metabolic activity. It is the time during which the cell is preparing for division by under growing cell growth and DNA replication in an orderly manner. Interphase Interphase is divided into 3 further phases or sub stages. 1) G1 Phase (Gap 1) 2) S phase (Synthesis) 3) G2 Phase (Gap 2) 1) G1 Phase: G1 phase corresponds to the interval between Mitosis and initiation of DNA replication. During G1 Phase the cell is metabolically active & continuously grows but does not replicate its DNA. The events that occur in G1 phase include. a) Increase in size of the cell. b) Synthesis of RNA and proteins. This process takes 5 hours. 2) S-Phase: This phase marks the period during which DNA replication takes place. During this time the amount of DNA/cell doubles, but there is no increase in chromosome number. This process takes 7 hours. 4 Phase S-Phase www.sciencetuts.com C ell cycle and cell division 3) G2 – Phase: During G2 phase, the synthesis of proteins & RNA continues. Various cell organelles are newly synthesized. Energy sources (ATP) required for spindle formation and movement of chromosomes is synthesized. It denotes gap between S-phase and M-phase. This process takes 3 hours. G2 – Phase Some cells in the adult animals do not appear to exhibit division (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 (Go) 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. M- Phase M-phase represents the phase when actually cell division occurs. It takes one hour. (Mitos means thread) Mitosis was first observed by Strass burger (1870) in plant cells & by Fleming (1882) in animal cells. Mitosis results in the formation of two identical daughter cells. Therefore mitosis may be considered as equational division. The chromosomal number of daughter cells is equal to the chromosomal number of parent cells. The daughter cells resemble the parent cell in all respects except the size. Mitotic cell division occurs in vegetative or somatic cells. Hence it is called somatic cell division. Mitosis occurs actively in apical meristems of stem or root. It helps in the growth of the organisms by increasing their size, shape and volume, hence it is called growth division. In Mitosis the nucleus divides first & is followed by division of cytoplasm. Nuclear division is called Karyokinesis and cytoplasm division is called Cytokinesis. Strass burger Fleming apical meristems KARYOKINESIS (Karyon means Nucleus; Kinesis means division) karyokinesis is the division of nucleus. It can be divided into 4 phases. They are 1) Prophase 2) Metaphase 3) Anaphase and 4) Telophase. PROPHASE (pro means first) Prophase is the first stage of Mitosis in the early prophase, the chromosomes are long slender and spread extensively in the nucleoplasm. The chromosomes gradually become short, thick and stout of condensation of chromatin. In the Mid prophase, each chromosome splits longitudinally into two parts called chromatids, but they remain united at the centromere. 5 Prophase www.sciencetuts.com C ell cycle and cell division In the late prophase, the Nuclear membrane decreases in thickness and gradually dissolves. Nucleolus also decreases in size and finally disappears. The chromosomes appear to be randomly scattered in the cytoplasm. At this stage, initiation of mitotic spindle takes place. The microtubules and the proteinaceous components of the cell cytoplasm, helps in this process. Late prophase METAPHASE (Meta means between) By this stage the complete disintegration of nuclear envelops & condensation of chromosomes is complete. At this stage the morphology of the chromosomes can be easily studied. The important changes in this phase are a) Formation of bipolar spindle fibres which are attached to the Kinetochores of chromosomes. Metaphase b) Orientation of chromosomes on the equatorial region and forming an equatorial plate or Metaphasic plate. In the Early Metaphase – a bipolar spindle apparatus is formed by the fusion of microtubules & chromosomes move towards the equator. All the centromeres along with Kinetochores are oriented along the axis of equator & their chromosomal arms are placed towards the poles. The spindle fibres start from poles extend towards equator and attaches to the kinetochore region of chromosomes and form into an apparatus known as spindle apparatus. The spindles are composed of microtubules and chemically consist of tubulin proteins. The plane of alignment of the chromosomes at metaphase is referred to as the Metaphase plate or equatorial plate. These kinds of spindle fibres are commonly observed. a) CONTINUOUS SPINDLE FIBRES: They originate at one pole and extend to the opposite pole through the equator without touching any chromosome. b) CHROMOSOMAL SPINDLE FIBRES: They originate at both the poles and extend in opposite directions towards the equator and attaches to the kinetochore region of chromosomes at the equator. c) INTERZONAL SPINDLE FIBRES: They are found suspended nearer the equator on its either side. ANAPHASE (Ana means back) The Anaphase is characterized by the following events 1) Separation of two chromatids. 2) Migration of chromatids (daughter chromosomes) to opposite poles. The centromere divides, spindle fibres begin to contract as a result, the two chromatids are separated from each other in each chromosome. The separated chromatids, along with centromere can be called daughter chromosomes. Each daughter chromosome moves away from equatorial plate the movement of chromosomes towards their respective poles takes place by pull & push mechanism of spindle fibres. Based on the position of centromere, the chromosomes exhibit V, L, I and J shapes during their movement towards opposite poles. 6 www.sciencetuts.com C ell cycle and cell division Anaphase V, L, I and J TELOPHASE (Telo means end) It is reverse of Prophase. The sets of daughter chromosomes at the poles, Organize into daughter nuclei. The chromosomes uncoil & all uncoiled chromosomes together appear as a network called chromatin reticulum. Nuclear envelope and nucleoli are again formed. Thus 2 daughter nuclei which are identical are formed. They contain the same number of chromosomes as in their mother nucleus. Telophase CYTOKINESIS Mitosis accomplishes not only the movement of duplicated chromosomes into daughter nuclei, but the cell itself divides into two daughter cells by a process called cytokinesis & its leads to the completion of cell division. In animal cells cytoplasmic division takes place by furrow in the plasma membrane. In higher plants, cytokinesis takes place by cell plate formation method. During this process, the spindle fragments gather at the equator region & form a barrel shaped structure called Phragmoplast. The vacuoles of Golgi complex enter the phragmoplast & release pectins into it. Thus the phragmoplast is changed into a liquid form of plate like structure known as cell plate. The growth of cell plate takes place centrifugally & gets connected to the parent wall. It gradually undergoes physical & chemical changes & finally forms into cement like layer known as middle lamellum thus dividing the cytoplasm into 2 parts. Cytokinesis Cell plate The cell organelles are equally distributed to both cells on either side of the middle lamellum. Primary cell wall materials like cellulose & hemicellulose are deposited on either side of middle lamellum & finally new primary cell walls are formed on either side of the middle lamellum. After the formation of two primary cell walls the parent cell is divided into two daughter cells. The division of cytoplasm by the formation of cell plate is called cell plate method. 7 www.sciencetuts.com C ell cycle and cell division 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) Cell wall Coconut DURATION OF MITOSIS: The mitotic division may be completed in 2-3 hours. Eg: 1) Root tip cells of onion – 100 min 2) Endosperm cells of pea – 180 min Temperature also effects Mitotic division In staminal hairs of Rhoeo – the nuclear division takes 30 min at 450c 75 min at 250c 135 min at 100c The time required for different phases also varies. Prophase is the longest phase. Telophase – 2nd place Anaphase – 3rd place Metaphase – Shortest phase. In cells of stigmatic hairs of some grasses The time required to complete the process At 1900c - prophase 36 – 40 min Metaphase 7 – 10 min Anaphase 15 – 20 min Telophase 20 – 35 min Significance of Mitosis 1. Growth and development in organism is caused by mitosis & it restores the surface or volume ratio of the cell. 2. The daughter cells formed by mitosis are identical with mother cell. Hence it is important in conserving the genetic integrity of the organism. 3. In unicellular organisms, mitosis helps in reproduction. vegetative and asexual propagation takes place through mitosis. 4. It maintains constant number of chromosomes in all the cells of the body. 5. Mitosis helps in wound healing. 6. Replacement of the lost parts – cells of the upper layer of epidermis, cells of the lining of the gut & blood cells. 8 www.sciencetuts.com C ell cycle and cell division MEIOSIS OR REDUCTION DIVISION (Meioum means to diminish) The type of cell division, in which the chromosomes number is reduced exactly to half is called Meiosis. The term Meiosis was coined by J.B. Farmer (1905). It occurs in reproductive cells of sexually reproducing organisms. The cells in which meiosis occurs are called meiocytes. J.B. Farmer In Meiosis the chromosomes present in meiocyte is reduced to half in Meiosis- I. & these reduced chromosomes divided by mitotically in Meiosis II. In this the mother cell divides into four daughter cells and these cells are un identical when compared to their parent cell. Every organism has definite number of chromosomes in diploid condition (2n). All organisms begin their life from zygotic stage. The zygote is formed by the fusion of gametes. Thus in diploid organisms every cell has 2 sets of chromosomes each acquired from the male and female gametes. Chromosomes with similar characters, uniform size & types belonging to parental sources are called homologous chromosomes. In the life cycle of diploid organisms, the chromosome number is reduced to half at one stage or the other. If not, at every fertilization the chromosome number would be doubled leading to abnormal number & causes unimaginable genetic disorders. Since the number of chromosomes is reduced to half, it is called reduction division. Zygotic stage Sperm Egg The key features of meiosis are 1) Meiosis involves two sequential cycles of nuclear cell division called Meiosis I and Meiosis II but only a single cycle of DNA replication occurs. 2) Meiosis- I is initiated after the parental chromosomes have replicated to produce identical sister chromatids at the S- phase of interphase. 3) Meiosis involves pairing of homologous chromosomes & recombination between them. 4) Four haploid cells are formed at the end of Meiosis – II. MEIOSIS- is divided in two : 1) Meiosis I 2) Meiosis II MEIOSIS I – further divides into 1) Karyokinesis – l 2) Cytokinesis MEOSIS II - further divides into 1) Karyokinesis – II 2) Cytokinesis 9 www.sciencetuts.com C ell cycle and cell division Karyokinesis – l: Occurs in four stages • Prophase – I • Metaphase – I • Anaphase – I • Telophase – I Prophase – I: Again has 5 stages • Leptotene • Zygotene • Pachytene • Diplotene • Diakinesis KARYOKINESIS – II: Has 4 stages • Prophase –II • Metaphase –II • Anaphase –II • Telophase – II MEIOSIS-I / HETEROTYPIC DIVISION / REPRODUCTIONAL DIVISION Karyokinesis – I: It is completed in 4 phases. In this the chromosome number is reduced to half. Cytokinesis may take place immediately or it may be delayed till the completion of telophase II. PROPHASE – l: Prophase of the first meiotic division is longer & complex. Based on the chromosomal behaviour, it has been sub divided into 5 phases. a. LEPTOTENE (or) LEPTONEMA: The nucleus & nucleolus enlarges in size. The chromosomes are long and slender. b. ZYGOTENE (or) ZYGONEMA: It is characterized by pairing of homologous chromosomes and this process of association is called synapsis. The pairs of homologous chromosomes are called bivalents. The pairing is bought about in a zipper like fashion due to protein called synaptonemal complex and may start at centromere or chromosome ends or at any other position. Based on the specific locations from which synapsis occur, it is further classified into three types. Leptotene(or) Leptonema Zygotene (or) Zygonema 1. Proterminal Synapsis: Pairing starts at the polarized ends and progresses gradually towards the other extremity. 2. Procentic Synapsis: Pairing starts at the centromere & proceeds towards the ends of the chromosomes. 3. Random or Intermediate Synapsis: Pairing of chromosomes occurs simultaneously at various places (randomly) along the length of the chromosomes. Zygotene is also characterized by the enlargement of nucleolus and initiation of spindle formation. C) Pachytene or Pachynema or Genetic Recombination stage: It is a long lasting stage (weeks or years) the chromosomes divides into two chromatids thus in each bivalent, four chromotids are seen. 10 www.sciencetuts.com C ell cycle and cell division They are called Pachytene tetrads. In a bivalent, the chromatids of the same chromosome are called sister chromotids and those of two different chromosomes as non sister chromatids. The non sister chromotids exchange their parts mutually at one, two or many places. Pachytene Such points where chromatids physically contact each other are called Chiasmata. During the formation of chiasmata, the chromotid arms first break due to the action of enzyme called “endonuclease” The broken chromatid arms mutually exchange with each other and get united by the action of an enzyme called Ligase.The formation of chiasmata lead to exchange of genetic material and result in the recombination of genetic characters. This phenomenon is known as ‘Crossing Over ‘. It is responsible for the origin of new species and thus leads to evolution. D) DIPLOTENE (or) DIPLONEMA: The beginning of diplotene can be recognized by the dissolution of the synaptonemal complex by the repulsion activity between homologous chromosomes, As a result, the bivalents of homologous chromosomes are separated from each other except at the sites of chiasmata regions. At this stage the chiasmata are clearly visible as ‘X’ shaped links between non sisters chromatids of homologous chromosomes. Diplotene In oocytes of some vertebrates diplotene can last for months or years. E) DIAKINESIS: This stage is marked by the ‘TERMINALISATION OF CHIASMATA‘ This displacement of chiasmata towards the terminal position is called Terminalisation. The bivalents become very thick & short and migrate to the periphery of the nucleus. The nucleolus begins to disappear, the nuclear membrane disrupts and the chromosomes are released into the cytoplasm. Diakinesis represents transition to metaphase. Diakinesis Metaphase – I: It is characterized by the appearance of bipolar spindle apparatus. The bivalent chromosomes move to the equator of the cell and arrange themselves with their centromeres directed towards opposite poles & their arms towards the equator. The movement of bivalent chromosomes towards the equator is called chromosomal congression. The homologous chromosomes are fused by the chiasmata at the terminal ends. ANAPHASE – I: It is characterized by the movement of each homologous towards the respective poles without the division of the centromere. This phenomenon is called Seggregation or disjunction of chromosomes. As a result, the diploid number of chromosomes is reduced to half. 11 Metaphase: I Anaphase –I www.sciencetuts.com C ell cycle and cell division TELOPHASE – I: The nuclear membrane and nucleolus reappear, cytokinesis follows and this is called diad of cells. The stage between two meiotic divisions is called Interkinesis and is generally short lived. Interkinesis is followed by prophase II. MEIOSIS II / HOMOETYPIC DIVISION / SECOND MEIOTIC DIVISION:- Telophase I Meiosis II is equivalent to Mitosis. It is completed into 2 stages. They are Karyokinesis - II & cytokinesis. Prophase ll: Is similar to mitotic prophase. The nuclear membrane, nucleolus appears and chromosomes are organized. Each chromosome has two chromatids which remain attached at centromere. All the chromosomes will be free in the cytoplasm. Prophase ll METAPHASE II: It is similar to mitotic metaphase but dissimilar to it in the formation of 2 spindle apparatus perpendicular to the one formed at the metaphase I. Metaphase II ANAPHASE II: It begins with simultaneous splitting of centromere of each chromosome allowing them to move toward opposite poles of the cell. Anaphase II TELOPHASE II: The daughter chromosomes arrived at the poles become thin and transform into chromatin. Nuclear membrane and nucleolus are reorganized. Thus at the end of Telophase –II, four daughter nuclei are formed. CYTOKINESIS: The process is similar as Mitotic division. Telophase II SIGNIFICANCE OF MEIOSIS 1) It helps in the maintenance of a constant chromosome number from one generation to the next. 2) Due to crossing over, genetic recombinations are caused which help in the origin of new species and lead to evolution. 12 www.sciencetuts.com C ell cycle and cell division MITOSIS: 1) 2) 3) 4) 5) 6) 7) 8) 9) It occurs in haploid and diploid cells. It occurs in somatic cells. Nucleus divides once. Daughter cells are identical. Two daughter cells are formed. Prophase is simple. Pairing of chromosomes does not occur. Both chiasmata & crossing over are absent. The chromosome numbers of daughter nuclei are unchanged. MEIOSIS: 1) It occurs only is diploid cells. 2) It occurs in reproductive cells. 3) Nucleus divides twice. 4) 5) 6) 7) 8) 9) Daughter cells are un identical. Four daughter cells are formed. It is complicated & shows five sub stages. Homologous chromosomes pair to form bivalents. Both chiasmata & crossing over occur between non sister chromatids. Chromosome number of daughter nuclei is reduced to half. Summary 1. According to cell theory. Cells arise from pre existing cells. The process by which this occurs is called cell division. 2. Any sexually reproducing organism starts its life cycle from a single-celled zygote. 3. Cell division does not stop with the formation of the mature organism but continues throughout its life cycle. 4. The stage through which a cell passes from one division to the next is called the cell cycle. 5. Cell cycle is divided into two phases called (i) interphase -a period of preparation for cell division, and (ii) Mitosis (M- phase) - the actual period of cell division. 6. Interphase is further subdivided into G1, S and G2. 7. G1 phase is the period when the cell grows and carries out normal metabolism. Most of the organelle duplication also occurs during this phase. 8. S phase marks the phase of DNA replication and chromosome duplication. 9. G2 phase is the period of cytoplasmic growth. 10. Mitosis is also divided into four stages namely prophase, metaphase, anaphase and telophase. 11. Chromosome condensation occurs during prophase. Simultaneously, the centrioles move to the opposite poles. The nuclear envelope and the nucleolus disappear and the spindle fibres start appearing. 12. Metaphase is marked by the alignment of chromosomes at the equatorial plate. 13. During anaphase the centromeres divide and the chromatids start moving towards the opposite poles. 13 www.sciencetuts.com C ell cycle and cell division 14. Once the chromatids reach the two poles, the chromosomal elongation starts, nucleolus and the nuclear membrane reappear. This stage is called the telophase. 15. Nuclear division is then followed by the cytoplasmic division and is called cytotokinesis. 16. Mitosis thus, is the equational division in which chromosome number of the parent is conserved in the daughter cell. 17. In contrast to Mitosis, meiosis occurs in the diploid cells, which are destined to form gametes. It is called the reduction division since it reduces the chromosome number by half while making the gametes. 18. In sexual reproduction when the two gametes fuse the chromosome number is restored to the value in the parent. 19. Meiosis is divided into two phases namely Meiosis-l and Meiosis II. In the first Meiotic division the homologous chromosomes pair to form bivalents and undergo crossing over. 20. Meiosis -l has along prophase, which is divided further into five phases. These are leptotene, zygotene, pachytene, diplotene and diakinesis. 21. During Metaphase- I the bivalents arrange on the equatorial plate. This is followed by anaphase-I. 22. Anaphase - l: In whish homologous chromosomes move to the opposite poles with both their chromatids. Each pole receives half the chromosome number of the parent cell. 23. In Telophase I, the nuclear membrane and nucleolus reappear. 24. Meiosis- II is similar to Mitosis. 25. During anaphase- II the sister chromatids separate. Thus at the end of meiosis four haploid cells are formed. 14 www.sciencetuts.com