Division Outlines 10.1 Cell Cycle 10.2 M Phase 10.3 Significance of Mitosis 10.4 Meiosis 10.5 Significance of Meiosis ‘n 1858, about 200 years after cells were first seen under the microscope, the German physiologist, Rudolf Virehow proposed the third dictum in cell theory, ‘omnis cellula e cellule’ (all cells come from pre-existing cells), CELL CYCLE Since then scientists have discovered that the formation of new cells includes both nuclear (karyokinesis) and cytoplasmic processes (cytokinesis). Like life eycle of individuals, cells also have a life cycle, the cell cycle. As » multicellular organism grows, this type of cell cycle is Cell Cycle and Cell | xvout 25 million cell divisions occur evey common. second in our own body. AS parts of a plant reach thei fal form, most cells stop dividing (cell cycle arres .d period of growth during whic and enter an extende' differentiate and mature. For example, leaf cell stp only a few millimeters long; thy they dividing when a leaf is continue to grow as the leaf and funetion for the rest oft leafs life. Leaf cells cannot normally be dividing again: the leat is damaged, very little regeneration is possible. I contrast, cortex cells in the stem of root also stop dividing before the organ is mature, but if damage occurs th remaining cells resume division and remain active until the(op : + difereniation (rene point or eetton pont) ‘dimes |e ete. eae (sere neeandvncanaped ee ere kent repleabon. X wie / 0 a eminent | OVA s G1 oan Mong oe Inactive Cake Om? a es molecule ee os Mey ng se $39 2, ey FS crinccge/ /oe8s % ee wait / Biren ierphase 4 FF ommome DQ Se %% £2 oa canes a % & ots structure ana ame 3 tel casmonaieerety 8 3S cyunbieod Cand mene cel cycle DML 2 Cyclin Dis broken rergttve only during Gx phase down at the beginning of Gy (check point or spindle check point | z CGoaurs near the end ofthe metaphase | | sage of miosis to determine if sister Ghromatis are correct attached tothe | spindle microtubules. eee Figure 10.1 The a layer of protective cork. ‘Some cells live for many years, even hundreds of years, but others die shortly after they mature, for example, RBCs of man lives only 120 days. Bark cells are more protective if they are dead, many flower parts die only few days after the flower first opens and gland cells often die after a brief period of secretion. Some cells never stop points a the tips of roots and shoots a are those that form wood and bark (the damage is sealed off with dividing. Cells in the growing re constantly cycling as cambium). The cell e oo? co or Cell grors and o% ogs® and roe str G2 ‘G2 check point Ensures col siza and protein reserves fre enough andi al the chromosomes ae replicated without mista, cell cycle and its control by three check points. cycle (Figure 10.1) can be divided into growth phase anda division phase. The phases of cell growth are collectively called interphase and phases of cell division are called mitosis. PHASES OF CELL CYCLE Duration of cel eycle may vary from hours to many years. typical human cell lives for 90 hours, whereas the cell cycle of Paramecium aurelia is about 6 hours. ‘The growth phase or interphase alternates with division or mitotic phase. a CHAPTER 10 Cell cycle and cell divisionya ner edt that betwee sway ass ac gost plse 68 x. Most meristernatie in interph Growth Phas During statics of ell visions cells: Were sit phrase, or inte cals the resting PMNS cells spend about 907 plant cells sp ss tn 195%, Alma Howard designate the phases OF MKD Eo ata this by measuring the iron ia rowing wot tips oF broad beans i ng pst ofthe 30 FE je apent in intemphase and about 20 pereec gynthesis. The length of a cell nding on the type of cell, the id many other eyele it resting tof their time cre Pete were the Fs sand G2 THY af radioactive (7) (licia faba). ‘They 1 eyele in nt of eas € proved that broad beans is SP spent in DNA. the cell eel cle varies tremendously, deper health, age temperature ype of plant, its factors. In 24 hour average duration of cell eyele of a human, cell division lasts for only about an hour ‘The interphase is the stage between two divisions and is the longest. Ina typical human cell, 89 out of 90 hours are spent in interphase. During this phase metabolic activities are high and no division occurs and hence also called resting phase. GI Phase GI phase or gap 1 phase or first growth period is the first stage after division. One the most important process it undergoes is the synthesis of nucleotides that are used for the synthesis ofthe next round of DNA replication. In algae, the cell cycle is brief for several hours. Short cell cycles also ‘occur in growing embryos and roots. On the other hand cell cycle may be from day to months in plants that grow slowly. The G1 phase also varies in accordance to the cell eyele. In ‘general, G1 is the longest part of the cell cycle. It can last even for years The nerve cells, however, remain permanently in GI phase. Generally this phase constitutes 25-50 percent of the total interphase. During G1 phase the daughter cells grows in size. mRNA, FRNA, and tRNA as well as proteins synthesized. wd uses Undergoes cell cycle ares it does not enter inGI 5, and G2 te 'se where it begin Phase and instead enters CO phase where it begii 2 D} UNIT it Cell: Structure and Functions sn and Gene amplification in Gy — iis stop in GI when they cease E doreduplicatio most plant cells ente, many cel in fo matt their DNA before they begin may involve just a single sf DNA replication, 1 in synthesis 2 smeleus becomes gigantic. THIS Process jg ied to occur in 80 perceny Although dividing and be § phinse arnt replicate to differentiate, T } rest cycle and the nue plication, estima s. dular cells, and other cells endor in is ndoreduplication occurs 1 maturing plant cell t often in hairs, glam ly rapid, intense metabolism, ofa mos! that must bean extremely 7 an “The normal complement of v0 copies OF each gene make messenger RNA rapidly does not seem to Such cells are also rich enough for such active cells: jn ribosome and produce large amounts of proteins, Gene amplification is similar to endoreduplication but involves only some genes that are repeatedly replicated. The amplified genes ‘are those needed for the specialized metabolism of the mature cell. For example, as a protein-rich seed develops or the oocytes of animals with yolky eggs (as amphibians) develop, the cells need large quantities of the messenger RNA that codes for the storage protein and are amplified. It would be however a waste of energy for the organisms to replicate all the other genes as well. to differentiate and mature. S Phase “Making chromatin’ is the major task of the this phase. 'S' stands for ‘synthesis of DNA'. During this phase DNA molecule replicates and the period last for 35~40 percent of the interphase. Genes that are formed of nucleotides replicate at this stage. In fact, thousands of genes are held together in 8 linear sequence in a chromosome in form of beads. The ak ey eee and are basic structural unit of the = ae Ane contain negatively charged oe ely charged histones (proteins). During the stinking pieces of DNA as well as genes are replicatedfon in protozoa Discovered ‘amitosis! (direct coll division) Description of mitosis, Used the term ‘karyokinesis! to describe ‘indirect cell division’ Created the word ‘mitosis’ Invented the terms, ‘prophase’, ‘metaphase’, and ‘anaphase’ Used the term ‘meiosis’ for reduction division Used the term, ‘telophase’. Coined the term ‘interphase’ 4749. Trembley jgai Robert Remak 4973. Schneider 4979. Schneider 479. W. Flemming 404 E. Stasburger 1890. Farmer and Moore 4894. M. Heidenhain 4913. H. Lundergardh and new histone molecules coupled with new DNA are formed. Thus, the entire chromosome is duplicated. Now the chromosome with its two chromatids are held together by centromere. Here each chromosome is a double chromosome and twice as large as it was before S phase. G2 Phase Ina few types of cell, G2 may be quite long and during much of it the cell physiology is similar to that in G1. Following S phase, the cell progresses into G2 (gap 2 or premitosis) phase during which cells prepare for cell division. This phase usually lasts only about 3 to 5 hours. The alpha and beta tubulin necessary for the spindle microtubules is synthesized and the cell is believed to produce proteins necessary for processing chromosomes and breaking down the nuclear envelop. In cultured animal tissue, if cell whose nucleus is dividing is forced to fuse with one in GI, the second cell's nucleus also begins the first steps of nuclear division. This is an evidence that during G2, the first cell produced the factors necessary to start nuclear division and that these factors are located in the cytoplasm. G2 is the second growth phase where the nucleus increases in volume, cell organelles are doubled, and spindles are formed for undergoing cell division. Gi, $ and G2 constitute the interphase portion of the cell cycle. After G2, cell division occurs. Division Phase of The Cell Cycle 1e M phase where mitosis 0c io (es in a typical human cell CY" 1 division involves two processes inesis) and (2) division of It includes th srs. This i of shot includes se duration of about 45 minut jon of 90 hours. A cell lcus (karyoki esis). the eytoplasm (ey! G0 or Differential Phase daughter cells again start (or enter) Gl .d develop. Cells that do not | phase undergo differentiation and 1, Such cells are, therefore, said to be in ese cells do not enter likely to do. After ‘After division, th phase. Here, again they grOW an proceed beyond G' ‘become a typical cel G0 phase or quiescent stage. Thus th cl yhase are $ phase as cells in GI p il differentiation the cells ultimately die, for example mature RBC, neurons and skeletal muscle. phase for long period are Cells that are arrested in G1 i cells in GO are destined said to be in GO state. Thes never to divide again, Most nerve cells in our body are in this state. What are Check Points? During the cell cycle, the cell checks to see whether it is ready to proceed with mitosis or cell division. They occur three times at G1, G2 and M phases. G1 check point. This check point makes sure that the cell is big enough and has all the ingredients to begin mitosis. G2 check point. After DNA replication in S phase the cell goes through the G2 check point where itis seen if DNA has been accurately replicated. M check point. It is the final check point that ensures that the choromosomes are attached to the mitotic spindle. CELL DIVISION The division of a cell into daughter cells is known as cell division. In fact, cell divisions lead to growth and development of every living organism. There are three kinds of cell division that occur in plants/animals. They are as follows: 1. Amitosis or direct cell division, 2. Mitosis or indirect cell division, and 3. Meisosis or reductional division. CHAPTER 10 Cell cycle and cell division (S810r (@) (2) os ) 2 a COOKED ‘Two daughter copa r nucleus biviing opus colls formed pari cot rigwe 10.2 sinos orton Arse pti into exactly equal halves te of call division yy pivision in Prokaryores sisunique and is unlike mity ity cone of the simplest mod Remak (1841), #7 Cell division in prokaryote 1c divides first followed by Amitosis cchich was first described by Robert the red blood cells of chick embryo fon, it is unlike mitosis. Here, However, the chromosom ‘on, It cytoplasmic division (Figure 40.3). The chromoso ts replicating from one poing 5) ‘Onen called direct d Icus elongates first; followed by @ constricti (circular DNA) start + DNA molecules, both remai ic (0 two. This is followed splits into two cireula membrane. The cell membr, the nacl daughter cells. joined to the cell deepens and splits the nucleus int division thus forming wo ty eytoplasmic ntess is rare but common in unicellular animals qarmvoans, carilage eels, diseased cells, old Brows brings the chromosomes to tWo ends of deeenerating cells and in cells of foetal membranes. Gell, No spindles are formed. Cytoplasmic constr the vrine amitsis, the nuclear membrane does not occurs around the middle of the cell splitting it in tn disappear (Figure 10.2) and the nucleus contents do not cells. Cell wall wall Attachment point ( DNA pie A RN ) @ % ) Bacterial ~ SY i, plasma Before replication Completion of cell division membrane OD EBT) Growth Separation of chromosome Circles a rowth of membrane between attachment points Figure 10,3 i Cell dvision in bacteria (prok, ‘aryotic cell) ) UNIT Cen: Cell : Structure and Function; isEAM PHASE Mitosis is @ duplication or equational division (Figuro joa). Ht is the most common cell di multicellular organis the number oF its cells is increasing sion that any 1 uses, s their body is growing and German cytologist, Walther Flemming, referred to the variou sof chromatin appearance as mitosen (ihreads) giving rise to the name ‘mitosis. Flemming’s work in the 1870s inspired many studies of mitosis. By the lartofthe twentieth century, biologists knew 88 Was the foundation for nuclear di sexual reproduction. Mitosis is called du because the nuclear genes in chromosomes are first copied; then one set of genes is separated from the other and each one is packed into its own nucleus, Each daughter nucleus is basically a duplicate of the original mother nucleus. Mitosis, thus, produces nuclei that are more or exact copies of the original nucleus, Mitosis, the separation of chromosomes and the formation of two genetically identical daughter nuclei. It is followed by cytokinesis, but there are many exceptions to this pattem, For example, in seed development, the nuclei of some tissues divide lication therefore, refers to many times before they become separated by cell walls, furthermore, in many species of algae, cytokinesis occurs only during sexual reproduction, Vegetative stages of these algae are permanently multinucleate. Mitosis can be studied in several types of meristematic cells, either in whole plants, in cell cultures or in wound or tumour tissues. For example, we can easily see the phenomenon in onion root tips. Thousands of genes are linked together in just few chromosomes. Each of these gene sets are duplicated in S phase by splitting of chromatid into two sister chromatids. In mitosis, one half of the doubled large chromosome (with sister chromatids) goes to each of the cell and the other half totheotherend, When that happens with each chromosome, each end of the cell automatically receives one full set of genes, The mechanism of separation of chromatids is explained in few stages of mitosis namely prophase, metaphase, anaphase and telophase (karyokinesis). This is followed by cytoplasmic division called cytokinesis. mTOR ees anx2 ‘Asomatic cell with 2n chromosomes Two daughter cells each with 2n chromosomes after mitosis Figure 10.4 A generalization of mitosis. FEE Prophase (= chromosome condensation) Prophase is the first stage of mitosis, In the early stage, the nucleus becomes spheroid and the cytoplasm gets viscous. ‘The chromatin of the nucleus shortens to from thick and elongated chromosomes. Each of the two centrosomes (with their centriole pairs) separate and migrate towards the opposite poles of the cell. Astral rays are formed around each centriole pair and together it is called aster. ‘The chromatin further condenses and the size of the chromosomes are considerably reduced. By late prophase the chromosome with two long chromatids (sister chromatids) appear as two parallel threads attached at a constriction point called the centromere. The centromere cot ists of a specific sequence of DNA that is required for the segregation of chromosomes later in mitosis. This double-thread form of the chromosome, is still considered to be one chromosome because it has just one centromere, The end of prophase is marked by the disappearance of nucleoli and the nuclear envelop. Without a nuclear boundary, chromosomes come into the cytoplasm and their behaviour appears to be somewhat uncontrolled. Chromosomes are, at their shortest, near the end of prophase and the beginning of the next metephase (Figure 10.5). sa CHAPTER 10. Cell cycle and cell divisionPLANT CELL, aa a ANIMAL CELL, ___— Contriotes — Nuclous ——__| Three important processes take place inthis phase. E Nuclootus: on Plasma membrane Cell wall synthesis. (histones) .« Duplication of centriole J 'Synthesis of proteins and enemy rich compounds for mitosis Chromosomes The chromosomes condense. Spindles are formed between = connotes nat move to = opposite poles Spindle fibres BS w 3 S Spindle fibres attach to & the chromosomes that lie & along the equator = Equator of 2 spindle w V shaped ( — = chromosome separate and x + i s move to the opposite poles LS H Reappearance = of nuclear (> } Chromosomes (with single chromatids) arrive at each membrane | Pole. New nuclear | membranes are formed and | cytoplasmic division begins Sa & Cell plate ee New cell wall Daughter cells formed have the same number of — CP, ‘chromosomes that original le es C nucleus (of the parent cell) RSP, had earlier Figure 10.5 Stages of mitosis. IT IM Cell : Structure and Functions[0A Metaphase (=chromosome alignment) ‘afer the nuclear envelope disso called spindle fibers, accumulate near Ty metaphase, Spindle fibers. parallel ib the chromosomes in which are made of microtubules, are collectively called jure spindle the spindle apparatus (spindle). ‘Th is elongated, with ity ends pointing towards, apparat opposite poles of the cell, Fibers from opposite poles ofthe spindle attach to cach chromosome on either side + to a dise-shaped structure called the of its eontrome complex of proteins that binds to the kinetochore centromere (Figure 10.6), Most fibers are not attached to kinetochore, because there are usually many more fibers than chromosomes, The unattached fibres are called ne fee to distinguish them from kinetochore- ‘As observed by light n chromosomes line up independently of one another in a circle around the circumference of the spindle apparatus. The plane of this circle called the metaphase plate (or equatorial plate) is perpendicular to the axis of the spindle (Figure 10.6). The position of chromosomes along metaphase plate is maintained by the balance of forces between two sets of kinetochore-bound fibres, one set from each of pole of the spindle. The chromosomes jeroscopy, the metaphase Spindle fibres Chromosomal fibres Kinetochore Secondary constriction Metaphase plate \died under condensed and best st he end of aphase are ology. By # microscope for their morph romosomes have all been forced into position in the cell. Disordered ‘The chromosomes are omeres and the of mm the wwe, the cl tight plane at asp hromosomes: becom ily for the splitting of their centre ¢ ordered. then 1 separation of their chrom ED Anaphase (= chromosome sep: aration) which is the shortest stage of mitosis, same centromere (i.e. Sister ite poles of the During anap! chromatids attached to the chromatids) separate and move to opposi spindle, Centromeres split before the chromatids separate and are pulled along the 's near the center of a chromosome, U shape as they axis of the spindle, When the two centromere i chromosome arms form a V or J oF are dragged through the cytoplasm. Upon separation, the word ‘chromatid’ no longer applies; each structure js considered as a separate chromosome (i.e, daughter) because it has its own centromere pindle elongates during anaphase, In most cells, the s| thereby increasing the distance between the poles. This movement further separates the two set of chromosomes. ‘As they move along the spindle, chromosomes get separated into two genetically identical nuclei Polar fibres x W > —S 8 AE Figure 10.6 Spindle fibres formed during cell division CHAPTER 10 Cell cycle and cell division (585, se ED ooo of dau e re ge at the wo poring telophase, aut ac eo ends and the spindle paras sets: ANN envelope is formed around ete oF HE NET Chromosomes an nce appear ine nese MEET readily elongate and deconden The chromosomes S| ain into aifase ehromatit cytokinesis (= division into cells oF D phase) begins + telophase) the cell chromosomes animal cells yp that © By the time the begins in le fibres develo .quator leading As early as late anaphase to prepare for cytokinesis. reach of the cell poles, cytokinesis (Figure 10.7). A ring of contract! shortens and creates constriction at the & to two daughter cells. In plants as cytokinesis continues, dictyosome vesicle laden with cell-wall precursors form phragmoplast along with microtubules. The vesicles along with phragmoplast form a single pate like vesicle called cell plate (Figure 10.8). The cell plate grows as dietyosor to the outer edge of the central vesicle. The new walls in the central vesicle grow outward along their edges. The cell plate continues to grow in this manner until the central vesicle fuses with the plasma membrane of the parent cell, thereby becoming part of the plasma me vesicles are added Vesicles forming phargmoplasts | i Cell plate Cell ~ membrane cel al : Nucleus: Golgi bodies i and endoplasmic reticulum Phragmoplast last membranes fc form form small vesicles in early tel vesicles fuse to form phragm lophase. The plasma mer amopasis atthe ells and ofr enna nnn er contents ( (poly- centre of the cell. The |, The spindle fibre transport these vesicles atthe conte, saccharides) to form a cell : cell plate, Figure 10.8 Cell wall formatioy Plant cel Il formation in plant New plasma membrane The cell plate forms the mi lamella around which cell materials deposit to form 02% thus dividing the cell intomembranes of the daughter cells. At the same time, new cell walls in the central vesicle join the wall of the HL walls is the last step parent cell. Fusion of the cytokinesis. As eytokin that surrounds the parent cell loosens and stretches, thereby enabling subsequent growth of the daughter cells by cell expansion, Shortly after cytokinesis, the daughter cells deposit additional primary cell wall materials on both sides of the newly formed cell wall and on all of the pre-existing is nears completion, the wall ta varies with cell types: ‘a dividing cell number of plasmodesmal tokinesis ‘Cytokinesis splits the cytoplasm of ini dsghier cells, In plants and few slage, evoke! ragmoplast and cell plate. The cell 1a division plane that is indie. The new cell wall cells are derived involves a phi formed from phragmoplast in he mitotic s perpendicular to th and plasma membranes of from the cell pl ughter late. measure of the amount of mitotic otic index is a oe ion of cells. It is calculated as the tivity in a popula walls a8 well, TWo features of the cytokinesis just a eee nitosis at any onetime. described characterize plants and few algae. One is the formation of a cell plate that involves a phragmoplast. In most other organisms rigid cell walls, cytokinesis occurs by constriction of the parent wall and plasma percentage me SIGNIFICANCE OF MITOSIS ‘The significance of mitosis is that it involves duplication of the genetic material and its equal distribution to each of two daughter cells. There is little or no variations. ‘The process provides the cells that are needed for the growth, An example would be a zygote developing into a functioning multicellular organism. For this to occur, there must be an increase in the number of cells from one cell to millions of cells in a human body. Mitosis supplies the cells for repair of worn or damaged tissues, e.g., in the human alimentary canal, the skin, lung, and blood cells. It maintains the chromosome number. Daughter cells have an identical set of chromosomes compared to the parent. This set of chromosomes function together as part of a tissue or an organ or an organism. Asexual reproduction produces offsprings that are genetically similar to the parents. membrane. In animal cells, which lack cell walls, cells are pinched off by the constriction of the parent plasma membrane. In some algae, however, a cell plate forms from microtubules oriented perpendicular to the spindle apparatus, not parallel as in a phragmoplast. In such algae, the cell-plate forming structure is called a phycoplast (phyco refers to algae). The second feature of cytokinesis that chraracterizes cell division in plants and some algae is the formation of plasmodesmata. As dictyosome vesicles accumulate in the growing cell plant, they trap parts of the endoplasmic reticulum, After cytokinesis, part of ER that were caught between fused vesicles connect the cytoplasm of the daughter cells. Indirect evidence shows that the Cainsicriieees Mysteries of the Mitotic Spindle How is Cell Cycle Controlled? Mitosis is dominated by the spindle apparatus. The mechanisms of spindle function have received much attention in the studies of mitosis, but the precise way that forces are formed by the spindle is still not fully understood. Two kinds of forces contribute to chromosome movement during anaphase. One force is the pull by kinetochore-bound microtubules on either side of each centromere, which moves sister chromatids in opposite directions. The other force is the push by microtubules that converge atthe poles of the spindle but are not attached to kinetochores. This pushing force eae the CHAPTER 10 Cell cycle and cell division (5871After an hour => ~— mined, one fi fon when cells are examin nds. ‘ter colchicine injec phase but Not Beyond. This is treatment (injection) the calls oF he Toc ofthe cals feaChINE TAT own the spindle formation a ana ea home meat sae fecossay Uli eon plate (equator) ct in metaphase stage and therefore, researches a sah colchicine before chromosomal studies i e 5 a often treat 7 crotubules pull while others push? - i ‘each other. How can some micro! poles of the spindle apparatus in opposite directions from sled apart Polar adie ns forse it ic longer; a sliding force pushes opposing microt ss Poverty boro at a the middle of the spindle apparatus. This suggests that opposing fibers somehow pole overlap those of the other p rat Panos slide past one another, thereby pushing their respective poles in opposite directions. This sliding microtubules hypothesis is supported by studies of isolated spindles of Stephanopyxis turris, a freshwater diatom (a type of alga). It is seen that ide during mitosis once the spindle fibres separate, the spindle movement stops. The mechanism by which microtubules is unknown, but one suggestion it that it resembles the movement of flagella. Spindle movement and flagellar movement are similar because they can both be blocked by inhibiting the hydrolysis of ATP. This means that ATP provides energy for ‘microtubule sliding and for theit movement. Unlike spindles, however, flagella contain dynein, large protein complex that converts energy from ATP into a sliding force between flagellar microtubles. Dynein has not yet been found in the mitotic spindle. Furthermore, spindle microtubules are oriented inditections opposite to their slide mates, while flagellar microtubules are oriented in the same direction, The pulling force of microtubules that are attached to a chromosome is probably caused by disassembly of microtubules. partial evidence for this hypothesis is that the alkoloid, taxol’, which supresses microtubules disassembly, inhibits chromosome movement. Further evidence is thatthe alkoloid colchicine, which inhibits assembly of microtubules, causes spindle fibers to shorten faster than normal; this accelerates the movement of chromosomes 1148 not known whether the disassembly of microtubules occures only atthe kinetochores or both at the kinetochores “An anticancerous drug extracted from Taxus baccala 88) UNIT lI Cell: Structure and Functionsand at the poles. The mode for mierotublule disassembly is also incomplete how chromosomes are se ied during mitosis, Cell Division Cycle Genes in Cell Cycle During the past few ye activate enzymes of the cell eyele, and on cell division eyele (ee) similar to those i far regulate transitions from the GI phase to the Studies of one of the yeast gen: called a mitotic function. Thi faster cell div covery was made when ede 2S was cell division affects cell size and floral development. Plant scientists clues about how ede genes work in plants, This is the first step toward the cell eyele w cell size and plant developme: ‘Types of Mitosis Various kind of mitosis occur in organisms. Some of them are given below: Intranuclear or Pre-Mitosis. In some acellular organisms (Amoeba and yeast) mitotic events occur within the nuclear envelop which remains intact. The nucleus and cytoplasm divides together. Extra-Nuclear Mitosis. In multicellular organisms, the clear membrane breaks and mitotic events occur in cytoplasm. It is also called eumitosis. Anastral Mitosis, Plants do not have centrioles and asters are not formed. Such mitosis without asters is anastral Astral or Amphiastral. In animals asters are formed and such mitosis is called astral type Endomitosis or Endoduplication. The multiplication of chromosomes (and DNA) without nuclear divisions is called endomitosis or endo- polyploidy or endoduplication. This increases the number of chromosome sets per cell. The polytene chromosomes are produced by endomitosis. Also liver cells show endoploid nuclei with 4 to 8 sets of chromosomes. ymmetrical Mitosis. The mitosis producing two equal cells is called symmetrical mitosis. symmetrical Mitosis. This leads to two unequal cells esearch on the cell eyele has f ts have also been found in plants and '§ phase or from the C jucer gene (tle 28) on in some cells oftheir flower buds, thereby causing th plants also produced flowers faster but without petals. This experiment sh jot fully explain consequently, we still canns s that ich are proteins ces of eyelins, whic sts, More recently, ede genes ave been discovered SO ced on the properti primarily in ¥6 Lof the ede genes that ha nimals, Most phase to mitosis rave revered how plant ede genes might fobacco plant containing ede underwent I to be smaller than normal. Such hat controls the timing of lc 2S in tobacco provides Jar regulation that links a peatedly 0. erred to toba cir daughter cell ows how a genes t believe that the action of ed ‘understanding the molecu! after division. Free Nuclear Division. divides several division. This conditions. Such : neytial conditions in epidermis of Ascaris Sometimes nucleus Te | times without cytoplasmic Jeads to multinucleate free nuclear division leads to sy or coenoeytic conditions in fungi. Eg MEIOSIS Meiosis is the process during which the chromosome number is reduced to half in daughter cells. Each daughter cell contains only one member of each pair of homologous chromosomes. Thus, meiosis ensures production of a haploid phase in the life cycle and fertilization ensures ‘a diploid phase (Figure 10.9). Without meiosis, sexual reproduction would not be possible. Meiosis was first worked out by Strasburger in 1888 but later on studied in more detail by Farmer and Moore in 1905 who also coined the term ‘meiosis’. Unlike mitosis, in which the chromosomes are doubled and then divided between two daughter nuclei in a single division, the doubling of the chromosomes prior to meiosis is followed by two sequential divisions that distribute the chromosomes among four nuclei. To ensure that each daughter nucleus formed by meiosis has one member of each of pair of homologous an elaborate process of chromosome pairing CHAPTER 10 Cell cycle and cell division (a ¢ eoe carats Gun + polar bodies 7 (3) Figure 10.9 Meiosis produces zeurs (Figure 10.10). As they are paired, homologous hromosomes engage in process of genetic recombination rat produces chromosomes with new combinations of aaternal and paternal alleles. The reproductive cells of various kind are called MEIOSIS I v ttf © | sperms (4) nt structures: neiocytes which undergo meiosis to produce ga Meiosis takes place in reproductive cells during the format, of gametes in animals, In protoctists, fungi in higher p meiosis lead to spore formation which later form gam, yD £) yy by mitosis. Each chromosome doubles tse thus _, Homologous Chromatids => ving two => chromosomes, separate ‘chromatids coparate SS FF ow > > xs s@ PS ee a | Prophase Prophase! Metaphase! —Telophase | Telophasell Figure 10.10 A generalized diagram showing meiosis. 5) UNIT Cel: Structure and Functions‘The following three groups can be identi oups ean be identified on the basis of where they occur, eritee de Gametic or Terminal Meiosis es all multicettular animals, otic divisions: In this group, whieh inc! many protozoa and a few lower plan are closely linked to the differentiation of the spermato70% Spermatogor reich then undergo two meiotic divisions to form four relati undifferentiated. spermatids : complete transformatioy en : transformation to become the highly specialized x atozoan, In Gao vertebrates, oogonia become mary Which then enter a gre c be a greatly extended ee and becomes filled with yolk and other materials. It is on I complete (ie. the oocyte has reached essentially the same state ia become primary spermatocytes, which oocy During this prophase, the primary oocyte after the differentiation of the oocyte as when it is fertili is fertilized) that the meiotic divisions occur. aag0 @unn ag v y Hat ganetoniyte (ooten poten ke @ Zygote emis Sper if Sperms (Baby, 2n) u ANIMAL, Female gametophyte @ 200 Vertebrate eggs are typically fertifized at a stags bofore the completion of meiosis (usually Meiosis is completed after fetiization, while th cytoplasm (Figure 10.14). at metaphase 11). he sperm enters in the ee Sporic or Intermediate Meiosis includes all higher plants, the meiotic nrclated to either gamete oF Je with the union of a ete (the ovum), In this group, whiel divisions take plac fertilization, If we begin the life eyel = (pollen cell) and a fermale gam the diploid zygote formed undergoes mitosis and develops iio a diploid sporophyte. Atsome stage in the development of sporophyte, sporogenesis (which includes meiosis) ‘occurs producing spores that germinate directly to haploid tsametophyte. The gametophyte ean bean independent Stabe or as in the ease of seed plants, a tiny structure retained ‘within the ovules. Regardless, the gametes are produced by the haploid gametophyte by mitosis. ®_.@ Transient diploid cell fa stage W male gam ) ®®® sous COO soa i (embryo sac) [ g ® ® NEW New FUNGUS FUNGUS NEW PLANT ure 10.11 Life cycle in humans, plants and fungi man and higher plants thre: , plants and fungi, In hi 4 ‘ehonn pl © cells after meiosis (in females) are CHAPTER 10 Cell cycle and cell division “591 ER 10 Cell cycl oe 2hZygotic or Initial Meiosis ' ncludes only few i ation in the: " st after the fertili meat are bp tes Liga wad afer ee sometimes several algae the few In the group, which (Chlamyviomonas, Spins meiotic divi stage of eye! complete, Moss tes fog tine 0 comets 5 «lays, unlike mitosis which fakes few" 0 hours. cet memvane wuctear | chromosome membvane METAPHASE ~ 1 oa sIS-1 a Me otorotypic division) 3 two round of division without allowing avolve a : Meth Henna atter the first division. ‘The two Avision, ‘ Leas a vejosis { (heterotyple division) and meiosis + called meiosi ae homotyple division) and each contains four phases sini to those of mitosis (Figuro 10.42) In the first meiotic division the diph Peis Meios cell is divideg : JAPHASE —1 ee Gronoromesconerse Sarg the oqo seat ae WNTERPHAEES ‘over ocours and spindle fibres ses re FOUR DAUGHTER CELLS Esch cell has hal the number of chromosomes i originally, ‘nad (n) with single chromatid TELOPHASE -i1 or MEIOSIS — 11 begins ch san cvopoone aa Qe >! RCS : N Two pausHTeR ceuLs Each cell has half the number of chromosomes (n) with two (sister) chromatids each, PROPHASE - 1, hromosomes recondense and {he spindle fibres form between {he centrioles which move apart. METAPHASE ~ 11 Spindles attach to chromosomes Figure 10.12 Dynamics of, meiosis, mes with one chromatid each ” cone ae ANAPHASE - 1 Cetcndes again Sister chromatids separate and move ‘0 opposite poles u : Stuctue and Functionsinto two haploid cells, that is, the chromosome number of daughter cell is halfofthe parent cell, Hence it is ealted the eterotypic division (meiosis 1) which includes prophase b, ctaphase 1, Anaphase I and telophase I. Prophase 1. All the events that characterize prophase of mitosis also occur in prophase 1 of meiosis. 1 is the longer than prophase of Tongest phase, evel Nucleolus uelear membrane break down, centrioles inde forms, microtubules attach ntromeres and chromosomes condense and become visible. In addition to these processes, special interactions of the chromosomes occur which are unique to prophase Tas opposed to prophase of mitosis, Because of this, prophase I is divided into five stages; namely leptotene, zygotene, pachytene, diplotene, and diakinesis. 1. Leptotene (or Leptonema = slender thread stage). The chromosomes begin to condense and appear indistinct as long uncoiled filaments or threads (Figure 10.13). The chromosomes (2n) appear in a series of beads called chromomeres (represents genes). The earlier divided centrioles move further apart. The homologous chromosomes, i.e., paternal and maternal pairs become ‘opposed to each other. These leptotene chromosomes that are irregularly arranged polarize to the centriole forming a bouquet. These groups of chomosome get attached to the nuclear membrane (bouquet stage). 2. Zygotene (mating thread or zygonema). During this, stage a remarkable pairing of chromosomes occurs. As there are two sets of chromosomes, one from the paternal side and one from the maternal side and these have gone through $ phase in the preceding interphase, each chromosome now has two chromatids; altogether there are four sets of genes of two sets of chromosomes. During zygotene, each chromosome of one set pairs with the equivalent chromosome (its homologous chromosome or homologue) of the other set (Figure 10.14). This pairing is called synapsis (or syndesis). With remarkable accuracy the two homologous chromosomes in each pair become almost perfectly aligned from end to end. A structure, the synaptonemal complex, is present between the paired homologous chromosomes. A pair of homologous +hromosome close together forming a bivalent’. Therefore, ‘Bivalent is used in reference to chromosomes (=2). Figure 10.13 Leptotene stage the number of chromosomes in a cell is twice the number of bivalents and each bive maternal chromosome. Sines chromatids, there are 4 chrot called the tetrad”. alent has one parental and one ‘e each chromosome has two matids in a homologous pair Synaptinemal Complex ‘The synaptinemal complex, which forms during synapsis protein axis along each chromosome. The begins as a homologous parts of synaptinemal complex links chromosomes, allele for allele. Each chromosome has a ribonucleoprotein lateral longitudinal elements. These Figure 10.14 Zygotene stage. ‘Tetrad is used in reference to chromatids of chromosomes (=in to form central element of synaptinemal complex (Figure 10.15). Homologous chromatin strands are 100 widely sep ‘ated which are at least 100nm apart, by the synaptinemal proteins for base-pair recognition to synaptinemal complex can join regions of chromosomes that are different, This is crucial for allowing lentical alleles at the same locus to fuse and exchange I. Chromosome exchange occurs in specific regions of remal complex that are called recombination Rodules named so because of large protein containing spheres associated with them, These nodules comprise several enzymes which ensure bringing closer the homologous segments or DNA from opposite chromatids Pairing in Zygonemal chromosomes 1. Proterminal Synapsis ‘The two homologous chromosomes start pairing from the tips (terminals) and move towards the centromere, 2. Procentric Synapsis The pairing begins from the centromere and Progress towards the terminal end of the ‘chromosomes, 3. Random Synapsis Pairing occur at many points randomly, 3. Pachytene or Pachynema (thick thread) stage, The chromosomes continue to condense and become shorter and thicker. This stage is pachytene. The synaptinemal complex seems to be involved in the process of crossing over that occurs now. In several places in each chromosome, the DNA (chromosome) of each homologous breaks (called nicking). A piece of parental homologous attaches at this point with the maternal one and equivalent piece of maternal homologous joins the paternal chromosome, The point of this interchange where the homologous. chromosome remain joined is called chiasma (plural, chiasmata), The interchange of chromatin material is called crossing over and occurs between the non-sister chromatids of the homologous chromosome (Figure 10.16), The chromatid breaks are initiated by enzyme, endonuclease and the reunion is brought about by an Cross over at chiasma Figure 10.15 Zygotene stage. enzyme called ligase. The process of rejoining is termes as annealing. Thus, we see in pachytene, mutual exchange of non sister chromatids. This brings about new combination g genes on each chromatid, which is called recombination, 4. Diplotene or diplonema. This stage follows th Pachytene. The homologous chromosomes begin to mov away (disjunction) from each other but do not separa Figure 10.16 Diplotene stage.Two chromatids of a homolog (sistor chromatids) Figure 10.17 The chromosomes (tetrad) are connected with spindle fibres in metaphase | of the chromosomes of the pair connect to the spindle poles. completely because they are held together at chiasmata (sing. chiasma) where they appear to be tangled together (Figure 10.17). All four chromatids of the paired homologous chromosomes (tetrads) are clearly visible at this stage. As. the separates begin to move along the length of chromosome from the centromere towards the end. This is termed as inalization which is seen by the end of diplotene chromosome the chiasmata ter stage. In certain animal cells like oocytes of amp! reptiles and birds the diplotene chromosome unwind and assume a large size called lampbrush chromosome. These decondensed diplotene chromosome help in synthesis of RNA. 5. Diakinesis, At this final stage, homologous chromosomes continue to separate and chiasmata are pushed to the ends of the chromosomes. Thus terminalization is completed in this stage. The homologous chromosomes become untangled and are ans, paired only at the centromeres. At this stage the chromosomes are best seen as short ind thick bivalents which tend to repel each other and Kinotochores of homolog | Telrad meiosis. Both the kinetochores of of the nucleus just inside the Jeolus and the nuclear s make migrate to the periphery nuclear membrane. The nucl membrane begins to disappear. The spindle fibre: their appearance in the cytoplasm. Prophase I. Itis the most complicated stage of meiosis. ‘The remaining stages are simple and quite similar to the stages of mitosis, Metaphase I. Spindle microtubules move the bivalents (tetrads) to the centre of the cell or metaphase plate. res of a pole join the two kinetochore The spindle of a chromosome. Similarly the two kinetochore of its homologue joins the opposite pole with its spindle fibres. Anaphase I. The homologous chromosomes separate completely from each other, moving to opposite ends of the spindle, The centromeres do not divide and each chromosome continues to consist of two chromatids. In mitosis the centromeres replicate and each chromosome divides into two chromosomes, each with just one chromatid, But in the metaphase-anaphase | transition, homologous chromosomes separate from each other and cach still has two chromatids. One set of chromosomes is pulled away from the other set and two new nuclei are CHAPTER 10 Cell cycle and cell division {595/0e cach has formed. These nuclei are now haploid, bec only one set of chromosomes; the homologue of each chromosome in one nucleus is now in the other micteus Unlike the conditions in mitosis, the two chromatids af hromosome do nat separate in meiosis 1. However, each parental and matemal homologous chromosomes after the crossover, with new combinations, segregate during. anaphase 1, This is independent of the homologous pairs, hence, corresponds to ‘Mendel’ law of independent assortment Telophase 1. As the chromosomes are still doubled 2 state, they do not need to undergo Also, because and are in a an interphase with its GI, S and G2. telophase 1 is basically the opposite of prophase II, some ‘organisms go directly from anaphase | to metaphase II, skipping telophase I and prophase II. In most organisms, however, there is at least a partial telophase I in which chromosomes start to uncoil and the nucleolus and nuclear envelope start to reappear. If the cells actually progress fully to interphase, no replication of the DNA ‘occurs-the S phase is completely missing. This interval is called interkinesis. Cytokinesis may occur, but it is not unusual for this to be absent also and for both daughter nuclei (i.e., both masses of chromatin) to stay in the original, undivided mother cell. EEX] Meiosis I! or Homotypic Division If a telophase I occurs, then prophase II is necessary to prepare the nucleus for division. Prophase II is not subdivided into stages like prophase I. Metaphase II is short and at the end of it, the centromeres divide, thereby separating each chromosome (with two chromatids) into two chromosomes, just as in metaphase of mitosis, but different from metaphase I. Anaphase II then separates each new chromosome from its replicate and in telophase I, new nuclei are formed. Each nucleus contains just ne set of chromosomes, each with a single chromatid. his follows cytokinesis leading to two more daughter lis. However, the cytoplasmic division could be in 0 ways as discussed belo Successive Cytokinesis. After meiosis | (heterotypic livision), two cells are formed by cytokinesis. Again I, ells divide in homotypic division or meiosis thereby, giving rise to 4 daughter cells. 2, Siuultancous Cytokinests. After me it ange in tetrahedron form, thus Four furry ead to form haploid cells Ty summarize, during meiosis 1, each chromogog, + undergoes synapsis wig of the paternal chromosom the homologous chromosome of the maternal 4g Crossing over results in new combinations OF genes 5 tne chromosomes. Spindle microtubules pull the pain homologous chromosomes to opposite Poles, so at eae pole there is only one set of chromosomes and not typ The new nuclei that form temporarily after meiosis |, thus haploid (n). When these nuclei undergo meiosis the two chromatids of each chromosomes are separate (as in mitosis) and the resulting nuclei each have , haploid set of chromosomes, each chromosomes wit only one chromatid. EQS) Significance of Meiosis Meiosis is necessary to ensure that the diploig chromosome number is reduced to a haploid number. The number of chromosomes must be halved during meiosis, Meiosis consists of two successive cell divisions, but only one cycle of chromosome replication. There is much variation due to each parent contributing half their genetic material (23 homologous chromosomes a5 in human) to the new organism. Meiosis separates chromosomes, halves the diploid (2n) number of chromosomes and introduces some variation into the haploid (n) cells that are produced and doubling the chromosome number at the time of fertilization is avoided. Crossing over occurs during prophase I. This is when all four chromatids are exactly aligned with each other causing non-sister chromatids to cross over, break and reassemble so that parental gene combinations are replaced by recombinants, Crossing over, independent assortment of. chromosomes and random fertilization are important Sources of genetic variation. New genetic material is thus provided for natural selection and evolution. The differences between various stages of mitosis and meiosis are enumerated in Table 4 and Table 2.gable 10.1 Differences between mitosis and meiosis. ‘Nucleus divides once. im bwalents of 4. Nucleus divides twice prophase chromosomes join to for In prophase chromosomes remain as separate units 2. In firs In fate eornosores ts into wo oper 4, In metaphase each chromosome splts to separate 3. Meet metaphase each bivalent 9 chromatcs. chromosomes. sed to opposite POeS- ‘4. inanaphase chromatids are pulled to opposite poles. «4 infest anaphase chromosomes 272 Pu | + To daughter cals ae formed, Spoor daughter eats are formed: cious 08 an other OF Foon daughter coll has te same chromosome number 2s 6. None ofthe dauger cole MO the same n0 asthe parent had. ag numberof chromosomes present * the parent cell, Le., diploid, the parent, Le., diploid. =D & &S Prophase AR Homologous pair do not ross over (or do not pair) 4 DNASYNTHESIS P Metaphase Chromatids separate felophase Two identical diploid (2n) cells: are formed Each daughter cell has the same chromosome number as 7. 4 pipLoicett > jhter cell has hi Each daughie Fie, call is haplow. in the parent cell (&) ¥I prophase ! Homologous pair cross ‘over and form pairs iit Motaphase | wy 3 Homologous chromosomes separate (®) as Telophase | Prophase Il eS Metaphase II Anaphase I! (chromatids separate) Four non-identical haploid cells are formedTable 10.2 Differences between the stages of mitosis and meiosis. Prophase Metaphase: Mitosis 1. Itis comparatively a shorter phase. 2. Chromosomes are shorter, thicker, and non-granular, 3. Each chromosome is made up of two chromatids. 4. Pairing of homologous chromosome, chiasmata formation and crossing over do not occur. 1. In metaphase, chromosomes are in the form of tetrads consisting of four chromatids. 2. The tetrads orient on the equator in such a way that their two centromeres are placed equidistant from the equator and face the poles while their arms are directed towards the equator. 3. The centromeres do not divide but the homologous chromosomes simply separate. 1. The chromosomes in anaphase are bivalent, each consisting of two chromatids. 2. The chromosomes in this stage are short and thickened. Moiosis 1. It is a longer phase comprising five sub-stages, 2. Chromosomes are thinner, longer, and granular. 3. Each chromosome appears double due to ty, chromatids. 4, Homologous chromosomes pair up. This is followeg by chiasmata formation. Crossing over occurs, In metaphase the chromosome is a dyad, consisting of two chromatids only. The orientation of dyads is just opposite to the 2 centromeres facing equator and arms directeq towards the poles. 3. The centrosome of each bivalent divides during metaphase into two and the members dyad separate, 1. The anaphase chromosomes which separate out are single. 2. The chromosomes in this Stage are longer and less thickened.