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Outline: Mitosis and the Cell Cycle 1. Why we need cell division 2. Stages of the cell cycle 3. How bacteria are different 4. Regulating the cell cycle – how does a cell know when to go to the next stage? 5. Cancer – what happens when the cell cycle is unregulated Fig. 12-2 Cell division is important for: 100 µm (a) Reproduction 20 µm 200 µm (b) Growth and development (c) Tissue renewal Cell division results in genetically identical daughter cells • Most cell division (mitosis) results in daughter cells with identical genetic information, DNA • A special type of division (meiosis) produces nonidentical daughter cells (gametes, or sperm and egg cells). These cells have half as much DNA as cells made by mitosis. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Overview: Cell Division • 2 ways to divide the cell’s DNA: mitosis and meiosis Mitosis Meiosis Makes new somatic (body) cells Makes reproductive cells (eggs and sperm) Purpose: Growth, development, repair Purpose: Reproduction New daughter cells have the same number of chromosomes as the parent cell New daughter cells have half as many chromosomes as the parent cell “My toes are made by mitosis, my ova are made by meiosis.” Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Organization of the Genetic Material • All the DNA in a cell constitutes the cell’s genome • A genome can consist of a single DNA molecule (common in prokaryotic cells) or a number of DNA molecules (common in eukaryotic cells) • DNA molecules in a cell are packaged into chromosomes, which are DNA molecules wrapped around proteins called histones. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-3 20 µm • Every eukaryotic species has a characteristic number of chromosomes in each cell nucleus • Somatic cells (nonreproductive cells) have two sets of chromosomes (1 from mom, 1 from dad) • Gametes (reproductive cells: sperm and eggs) have half as many chromosomes as somatic cells • Eukaryotic chromosomes are made condensed chromatin, a material made of DNA and protein Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Human Gamete Cell Chromosomes Human Somatic Cell Chromosomes Each chromosome is part of a pair of 2 homologous chromosomes Each chromosome is by itself – doesn’t have a homologous chromosome Distribution of Chromosomes During Eukaryotic Cell Division • In preparation for cell division, DNA is replicated and the chromosomes condense • Each duplicated chromosome has two identical sister chromatids, which separate during cell division • The centromere is the narrow “waist” of the duplicated chromosome, where the two chromatids are most closely attached Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-UN3 Fig. 12-4 0.5 µm Chromosomes Chromosome arm DNA molecules Chromosome duplication (including DNA synthesis) Centromere Sister chromatids Separation of sister chromatids Centromere Sister chromatids • Eukaryotic cell division consists of: – Mitosis, the division of the nucleus – Cytokinesis, the division of the cytoplasm • Gametes are produced by a variation of cell division called meiosis • Meiosis yields nonidentical daughter cells that have only one set of chromosomes, half as many as the parent cell Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Outline: Mitosis and the Cell Cycle 1. Why we need cell division 2. Stages of the cell cycle 3. How bacteria are different 4. Regulating the cell cycle – how does a cell know when to go to the next stage? 5. Cancer – what happens when the cell cycle is unregulated Phases of the Cell Cycle • The cell cycle consists of – Mitotic (M) phase (mitosis and cytokinesis) • Mitosis has 5 subphases – Interphase (cell growth and copying of chromosomes in preparation for cell division) • Interphase has 3 subphases Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Interphase (about 90% of the cell cycle) can be divided into subphases: – G1 phase (“first gap”) – S phase (“synthesis”) – G2 phase (“second gap”) • The cell grows during all three phases, but chromosomes are duplicated only during the S phase Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-5 G1 S (DNA synthesis) G2 G1 + S + G2 = Interphase M phase = Mitosis • Mitosis is conventionally divided into five phases: – Prophase – Prometaphase – Metaphase – Anaphase – Telophase • Cytokinesis is well underway by late telophase BioFlix: Mitosis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-UN1 G1 S Cytokinesis Mitosis G2 MITOTIC (M) PHASE Prophase Telophase and Cytokinesis Prometaphase Anaphase Metaphase Fig. 12-6a G2 of Interphase Prophase Prometaphase Fig. 12-6b G2 of Interphase Chromatin Centrosomes (with centriole (duplicated) pairs) Prophase Early mitotic Aster spindle Nucleolus Nuclear Plasma envelope membrane Prometaphase Centromere Chromosome, consisting of two sister chromatids Fragments of nuclear envelope Kinetochore Nonkinetochore microtubules Kinetochore microtubule The Mitotic Spindle: A Closer Look • The mitotic spindle is an apparatus of microtubules that controls chromosome movement during mitosis • During prophase, assembly of spindle microtubules begins in the centrosome, the microtubule organizing center • The centrosome replicates, forming two centrosomes that migrate to opposite ends of the cell, as spindle microtubules grow out from them Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • An aster (a radial array of short microtubules) extends from each centrosome • The spindle includes the centrosomes, the spindle microtubules, and the asters Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • During prometaphase, some spindle microtubules attach to the kinetochores (parts of the centromere that attach to the spindle fibers) of chromosomes and begin to move the chromosomes • At metaphase, the chromosomes are all lined up at the metaphase plate, the midway point between the spindle’s two poles Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-7 Aster Centrosome Sister chromatids Microtubules Chromosomes Metaphase plate Kinetochores Centrosome 1 µm Overlapping nonkinetochore microtubules Kinetochore microtubules 0.5 µm Fig. 12-UN4 Fig. 12-6c Metaphase Anaphase Telophase and Cytokinesis Fig. 12-6d Metaphase Anaphase Metaphase plate Spindle Centrosome at one spindle pole Telophase and Cytokinesis Cleavage furrow Daughter chromosomes Nuclear envelope forming Nucleolus forming • In anaphase, sister chromatids separate and move along the kinetochore microtubules toward opposite ends of the cell • The microtubules shorten by depolymerizing at their kinetochore ends Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Nonkinetochore microtubules from opposite poles overlap and push against each other, elongating the cell • In telophase, genetically identical daughter nuclei form at opposite ends of the cell Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Cytokinesis: A Closer Look • In animal cells, cytokinesis occurs by a process known as cleavage, forming a cleavage furrow • In plant cells, a cell plate forms during cytokinesis Animation: Cytokinesis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-9a 100 µm Cleavage furrow Contractile ring of microfilaments Daughter cells (a) Cleavage of an animal cell (SEM) Fig. 12-9b Vesicles forming cell plate Wall of parent cell Cell plate 1 µm New cell wall Daughter cells (b) Cell plate formation in a plant cell (TEM) Fig. 12-10a Nucleus Nucleolus 1 Prophase Chromatin condensing Fig. 12-10b Chromosomes 2 Prometaphase Fig. 12-10c 3 Metaphase Fig. 12-10d 4 Anaphase Fig. 12-10e Cell plate 5 Telophase 10 µm Fig. 12-UN5 Outline: Mitosis and the Cell Cycle 1. Why we need cell division 2. Stages of the cell cycle 3. How bacteria are different 4. Regulating the cell cycle – how does a cell know when to go to the next stage? 5. Cancer – what happens when the cell cycle is unregulated Binary Fission • Prokaryotes (bacteria and archaea) reproduce by a type of cell division called binary fission • In binary fission, the chromosome replicates (beginning at the origin of replication), and the two daughter chromosomes actively move apart Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-11-1 Cell wall Origin of replication E. coli cell Two copies of origin Plasma membrane Bacterial chromosome Fig. 12-11-2 Cell wall Origin of replication E. coli cell Two copies of origin Origin Plasma membrane Bacterial chromosome Origin Fig. 12-11-3 Cell wall Origin of replication E. coli cell Two copies of origin Origin Plasma membrane Bacterial chromosome Origin Fig. 12-11-4 Cell wall Origin of replication E. coli cell Two copies of origin Origin Plasma membrane Bacterial chromosome Origin The Evolution of Mitosis • Since prokaryotes evolved before eukaryotes, mitosis probably evolved from binary fission • Certain protists exhibit types of cell division that seem intermediate between binary fission and mitosis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-12 Bacterial chromosome (a) Bacteria Chromosomes Microtubules Intact nuclear envelope (b) Dinoflagellates Kinetochore microtubule Intact nuclear envelope (c) Diatoms and yeasts Kinetochore microtubule Fragments of nuclear envelope (d) Most eukaryotes Outline: Mitosis and the Cell Cycle 1. Why we need cell division 2. Stages of the cell cycle 3. How bacteria are different 4. Regulating the cell cycle – how does a cell know when to go to the next stage? 5. Cancer – what happens when the cell cycle is unregulated The eukaryotic cell cycle is regulated by a molecular control system • The frequency of cell division varies with the type of cell – some cells stop dividing altogether • These cell cycle differences result from regulation due to signaling molecules Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Evidence for Cytoplasmic Signals • The cell cycle appears to be driven by specific chemical signals present in the cytoplasm • We know this because if you put cytoplasm from one cell into a different cell, it can move that cell ahead in the cell cycle Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-13 EXPERIMENT Experiment 1 S G1 Experiment 2 M G1 RESULTS S S When a cell in the S phase was fused with a cell in G1, the G1 nucleus immediately entered the S phase—DNA was synthesized. M M When a cell in the M phase was fused with a cell in G1, the G1 nucleus immediately began mitosis—a spindle formed and chromatin condensed, even though the chromosome had not been duplicated. The Cell Cycle Control System • The events of the cell cycle are directed by a cell cycle control system, which is similar to a clock • The cell cycle control system is regulated by internal and external controls • The clock has specific checkpoints where the cell cycle stops until a go-ahead signal (one of those molecules in the cytoplasm) is received Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-14 G1 checkpoint Control system G1 M G2 M checkpoint G2 checkpoint S • For many cells, the G1 checkpoint seems to be the most important one • If a cell receives a go-ahead signal at the G1 checkpoint, it will usually complete the S, G2, and M phases and divide • If the cell does not receive the go-ahead signal, it will exit the cycle, switching into a nondividing state called the G0 phase Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-15 G0 G1 checkpoint G1 (a) Cell receives a go-ahead signal G1 (b) Cell does not receive a go-ahead signal The Cell Cycle Clock: Cyclins and Cyclin-Dependent Kinases • Two types of regulatory proteins are involved in cell cycle control: cyclins and cyclin-dependent kinases (Cdks) • The concentration of cyclins varies throughout the cell cycle • Cdks are dependent on cyclins, which activate them • When cyclin and Cdk bind together, they form the active molecule MPF that triggers a cell’s passage past the G2 checkpoint into the M phase Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-16 RESULTS 5 30 4 20 3 2 10 1 0 100 200 300 Time (min) 400 0 500 Fig. 12-17b Degraded cyclin G2 Cdk checkpoint Cyclin is degraded MPF Cyclin (b) Molecular mechanisms that help regulate the cell cycle Cyclin accumulation Cdk Stop and Go Signs: Internal and External Signals at the Checkpoints • Cyclins and cdks are proteins that can be made by ribosomes with instructions from DNA • You have genes in your DNA that give instructions for how to make these proteins • Gene expression is regulated – each gene can be turned on (causing that protein to be made) or off at different times • Internal and external signals lead to the genes for cyclins and cdks being turned on or off. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Examples of signals • An example of an internal signal (comes from inside the cell) is that kinetochores not attached to spindle microtubules could send a molecular signal that delays anaphase • Some external signals (come from outside the cell) are growth factors, proteins released by certain cells that stimulate other cells to divide • For example, platelet-derived growth factor (PDGF) stimulates the division of human fibroblast cells in culture Fig. 12-18 Scalpels Petri plate Without PDGF cells fail to divide With PDGF cells proliferate Cultured fibroblasts 10 µm Outline: Mitosis and the Cell Cycle 1. Why we need cell division 2. Stages of the cell cycle 3. How bacteria are different 4. Regulating the cell cycle – how does a cell know when to go to the next stage? 5. Cancer – what happens when the cell cycle is unregulated • Another example of external signals is densitydependent inhibition, in which crowded cells stop dividing • Most animal cells also exhibit anchorage dependence, in which they must be attached to a stable surface in order to divide • Normal cells make a protein called p53 to stop the cell cycle if there is a problem. In extreme cases, p53 can cause a damaged cell to die (apoptosis) • Cancer cells exhibit neither density-dependent inhibition nor anchorage dependence, and mutations in the gene for p53 are the most common mutations leading to cancer Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-19 Anchorage dependence Density-dependent inhibition Density-dependent inhibition 25 µm 25 µm (a) Normal mammalian cells (b) Cancer cells Loss of Cell Cycle Controls in Cancer Cells • Cancer cells do not respond normally to the body’s control mechanisms; they grow and divide out of control • Cancer cells may not need growth factors to grow and divide: – They may make their own growth factor – They may convey a growth factor’s signal without the presence of the growth factor – They may have an abnormal cell cycle control system Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • A normal cell is converted to a cancerous cell by a process called transformation – Happens due to mutations that change the DNA, making the cell ignore regulatory checkpoints and measures. Mutations in p53 are the most common. • Cancer cells form tumors, masses of abnormal cells within otherwise normal tissue • The lump is called a benign tumor if the cancer cells remain in the same place • Malignant tumors invade surrounding tissues and can metastasize, exporting cancer cells to other parts of the body, where they may form secondary tumors Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 12-20 Lymph vessel Tumor Blood vessel Cancer cell Metastatic tumor Glandular tissue 1 A tumor grows from a single cancer cell. 2 Cancer cells invade neighboring tissue. 3 Cancer cells spread to other parts of the body. 4 Cancer cells may survive and establish a new tumor in another part of the body.