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Anu Singh-Cundy • Gary Shin Discover Biology SIXTH EDITION CHAPTER 7 Cell Division © 2015 W. W. Norton & Company, Inc. CHAPTER 7 Cell Division, Part 1 OLYMPIC-CLASS ALGAL BLOOM 7.1 Why Cells Divide Many bacteria use binary fission for asexual reproduction Eukaryotes use mitosis to generate identical daughter cells Meiosis is necessary for sexual reproduction Cell divisions grow, maintain, and reproduce the human body 7.2 The Cell Cycle DNA is replicated in the S phase Most cell types in the adult body do not divide The cell cycle is tightly regulated 7.3 The Chromosomal Organization of Genetic Material A karyotype displays all the chromosomes in a nucleus Most human cells have two copies of each type of chromosome CHAPTER 7 Cell Division, Part 2 7.4 Mitosis and Cytokinesis: From One Cell to Two Identical Cells Chromosomes are compacted during early prophase Chromosomes are attached to the spindle in late prophase Chromosomes line up in the middle of the cell during metaphase Chromatids separate during anaphase New nuclei form during telophase The cytoplasm is divided during cytokinesis 7.5 Meiosis: Halving the Chromosome Set to Make Gametes Gametes contain half the chromosomes found in somatic cells Meiosis I is the reduction division Meiosis II segregates sister chromatids into separate daughter cells Meiosis and fertilization contribute to genetic variation in a population Crossing-over shuffles alleles The independent assortment of homologous pairs generates diverse gametes BIOLOGY MATTERS: PROGRAMMED CELL DEATH APPLYING WHAT WE LEARNED: THE GREAT DIVIDE Olympic-Class Algal Bloom • • Algae are photosynthetic eukaryotes that divide rapidly when nutrients are abundant. Large mats of floating algae are capable of stopping boats and entangling wildlife. The Capacity to Reproduce through Cell Division Is a Defining Feature of Life • The continuity of life depends on cell division. • It takes trillions of cell divisions to turn a fertilized egg into an adult human. • Millions of cell divisions take place in our bodies every day to replace the cells that have died. • Cell division is the generation of daughter cells from a parent cell. Why Cells Divide • • • • Cell division is necessary for reproduction in all life forms, and for growth and repair in a multicellular body. Asexual reproduction results in offspring that are genetically identical to the parent. Sexual reproduction combines genetic information from two individuals of opposite mating types to produce offspring. Offspring resulting from sexual reproduction are similar, but not identical, to the parents. Many Bacteria Use Binary Fission for Asexual Reproduction • Binary fission begins with replication of the genetic material. • Each of the two copies of DNA are segregated to opposite sides of the parent cell. • A partition develops to separate the cytoplasm, with each side containing a single copy of the DNA. • The resulting two daughter cells replace the single parent cell and are genetically identical to each other and to the original parent cell. Eukaryotes Use Mitosis to Produce Identical Daughter Cells • Mitotic division is the process in eukaryotes that generates two genetically identical daughter cells from a single parent cell. • Mitotic cell division involves the even distribution of replicated DNA into new daughter cells. • During a mitotic division, the nucleus divides in a process called mitosis, followed by a division of the cytoplasm called cytokinesis. Meiosis Is Necessary for Sexual Reproduction • Meiosis is a specialized form of cell division that is needed to make gametes (sex cells). • Meiosis in female animals results in gametes that will mature into eggs. • Meiosis in male animals produces gametes called sperm. • Meiosis reduces the genetic information passed to the daughter cells by half (n). • The fusion of information from the egg (n) and information from the sperm (n) restores a full set of genetic information (2n) in the zygote (fertilized egg). Fertilization Is the Merging of a Male and Female Gamete to Produce a Zygote • A gamete has half the amount of genetic information as a somatic cell (nonsex cell in the body). • A somatic cell contains a diploid set of genetic information (2n), whereas a gamete contains a haploid set (n). • The zygote receives one haploid set from each gamete (n + n). • The resulting embryo divides mitotically as it develops. Cell Divisions Grow, Maintain, and Reproduce the Human Body • Cell differentiation allows the daughter cells to become specialized for a specific set of functions. • All of the genetic material present in the zygote remains present in the somatic cells of the adult body, regardless of differentiation. • A small group of gamete-producing cells, called germ line cells, are reserved early in embryonic development for eventually producing gametes in the individual. • Adult stem cells are unspecialized cells that can divide to grow, regenerate, and repair tissues throughout our life span. In Humans, Meiosis Resumes (Females) or Begins (Males) at Puberty • In a female, meiosis begins while she’s still a fetus: about a million gametocytes (diploid precursor cells) begin meiosis, but then the process is put on hold until puberty! So, baby girls are born with about a million gametocytes that are arrested in an early stage of meiosis. • Typically, one arrested gametocyte resumes meiosis each month in one of the ovaries of a fertile female once she reaches puberty; one resulting daughter cell differentiates into an egg cell (ova). • In males, meiosis begins at puberty, when hormones signal millions of gametocytes, in both testes, to begin producing haploid cells that differentiate into sperm. • Male meiosis occurs daily and continues well into old age, but in women the supply of functioning gametocytes dwindles and usually disappears by age 50. The Cell Cycle Is Divided into Two Main Stages: Interphase and Cell Division The cell cycle can be completed in as little as 90 minutes in yeast cells. It takes about 24 hours in most human cells. • The cell cycle refers to the life cycle of a eukaryotic cell from its origin to its division by mitosis or meiosis. • Cell division includes: – Nuclear division, involving separation of chromosomes (mitosis) – Cytoplasmic division (cytokinesis), creating two daughter cells During Interphase, the Cell Grows and May Prepare to Divide Interphase is divided into three phases: G1, S, and G2 • During interphase, the cell: – Takes in nutrients and manufactures proteins and other substances – Grows – Conducts its specialized functions – Prepares to divide, including replicating the genetic material DNA Is Replicated in S Phase • The cell’s size and protein content increase during G1 and G2 phases, and checks are made to assess whether it is safe to progress to the next stage of the cell cycle. • The replication of DNA occurs in the S phase (“S” for synthesis of DNA). The G0 State Represents a “Safe Haven” Against Progression to Cancer • Many terminally differentiated cells enter a nondividing phase called G0. • The G0 phase can last from a few days to the lifetime of the organism. • Liver cells stay in G0 phase much of the time but divide once a year on average. • Neurons stay in G0 phase for their entire life span. • Many of the genes needed for DNA replication and cell division are absent in cells in the G0 phase, so these cells are resistant to entering and advancing through the cell cycle. The Cell Cycle Is Tightly Highly Regulated • Cell cycle checkpoints ensure that the cell cycle does not progress if conditions are not suitable for cell division. • The checkpoints are controlled by cell cycle regulatory proteins. • Internal and external signals can prompt the cell cycle regulatory proteins to advance the cell cycle. • However, cell cycle regulatory proteins can stop or pause the cell cycle if conditions are not favorable for cell division. Genetic Material Is Organized into Chromosomes, Each Chromosome Containing a Single DNA Molecule • Each DNA double helix is packaged with special proteins to form long strands of chromatin. • Chromatin is further compacted to form chromosomes. At the End of S Phase, the Replicated DNA Helices Remain Attached as Sister Chromatids • DNA is replicated during S phase, resulting in two identical double helices known as sister chromatids. • The identical sister chromatids are held together at a region called the centromere. The Karyotype Describes All the Chromosomes in a Nucleus • Every species has its own characteristic number of chromosomes. • The display of all the chromosomes in a somatic cell is called the karyotype. • Karyotypes can be seen more easily during mitosis, when chromosomes are highly compacted. • The number of chromosomes found in any given organism does not indicate the total number of genes, nor does it reflect the complexity of the organism. Somatic Cells Have Two Copies of Each Type of Chromosome • Eukaryotes carry two copies of each type of chromosome; together the two copies make up a pair of homologous chromosomes. • Humans have 46 chromosomes divided into 23 homologous pairs. • One set of chromosomes, called sex chromosomes, determines the sex of an individual among mammals. Mitosis and Cytokinesis: From One Cell to Two Identical Cells • Mitosis can be divided into four main phases: – Prophase – Metaphase – Anaphase – Telophase • The main role of mitosis is to separate sister chromatids and distribute one of each chromosome into each of the daughter cells. Chromosomes Condense and the Mitotic Spindle Forms in Early Prophase The Nuclear Envelope Disassembles and the Mitotic Spindle Captures the Replicated Chromosomes by the End of Late Prophase During Prophase, the Cell Prepares for the Segregation of Chromosomes • • • • • The chromatin becomes highly compacted, which reduces the risk of tangling and breakage at later steps. Two cytoskeletal structures called centrosomes begin to move toward opposite ends of the cell. Special proteins form long cylinders, called microtubules, that assemble into the mitotic spindle, which is anchored to a centrosome at its two ends. The nuclear envelope breaks down, enabling spindle microtubules to capture all the replicated chromosomes: the spindle microtubules attach to patches of protein (kinetochores) on each side of the centromere. The spindle will guide sister chromatids to the metaphase plate, and later to the opposite ends of the cell, as the later phases of mitosis unfold. Chromosomes Are Positioned at the Middle of the Cell during Metaphase • During metaphase, all of the replicated chromosomes are arranged in a plane that typically lies at the center of the cell. • The function of metaphase is to align the chromosomes precisely in order to facilitate equal distribution of chromosomes to each end of the cell. Chromatids Separate during Anaphase • During anaphase, sister chromatids are separated and pulled to opposite ends of the cell by the progressive shortening of the spindle microtubules. • Once separated, each chromatid is considered a new chromosome. New Nuclei Form during Telophase • During telophase, the mitotic spindle breaks down, and nuclear envelopes begin to form around the two new sets of chromosomes at each end of the cell. • Within each nucleus, the chromosomes decondense, as each new daughter cell begins its G1 phase. The Cytoplasm Is Divided during Cytokinesis • Cytokinesis is the process of dividing the parent cell’s cytoplasm to produce two daughter cells. • In animal cells, actin microfilaments form at the metaphase plate; as the actin filaments contract, they draw the plasma membrane in until the original cytoplasm is pinched into two new cells, the daughter cells. • Cytokinesis is the last step in the cell cycle. Cell Division in Plants Meiosis Is Necessary for Gamete Formation • In sexual reproduction, gametes of the opposite type (egg and sperm) fuse during fertilization to produce a zygote. • Each gamete is haploid (n): it contains half the diploid set of chromosomes (2n). • The zygote formed by the two haploid gametes contains one paternal homologue and one maternal homologue for each pair of homologous chromosomes. • Meiosis is a special type of cell division: its function is to generate haploid cells from a diploid parent cell by splitting up every homologous pair and sorting its members into different gametes. The Chromosome Set Is Halved During Meiosis • Meiosis occurs in two stages: --Meiosis I sorts each member of a homologous pair into two different daughter cells, reducing the chromosome sets from 2n to n. --Meiosis II separates sister chromatids in each cell produced by meiosis I into two different daughter cells. • • Meiosis produces four daughter cells from the division of each diploid precursor cell (gametocyte). In animals, the daughter cells differentiate into gametes (egg or sperm). Meiosis I is a Reduction Division: It Sorts the Two Members of Each Pair of Homologous Chromosome into Different Daughter Cells • Paternal and maternal partners of each homologous chromosome pair align themselves next to each other during prophase of meiosis I (prophase I). • A tetrad consists of one replicated maternal chromosome aligned parallel to its matching replicated paternal chromosomes, for a total of four chromatids. Crossing-Over in Prophase of Meiosis I Shuffles the Genes on the Homologous Pairs Before They Are Inherited by the Daughter Cells • During prophase I, DNA segments are swapped between nonsister chromatids in a process called crossing-over. • Crossing-over is a random process: the number and the location of a crossover can vary from one meiosis to another. • Meiosis I continues through prophase I and into metaphase I, anaphase I, and telophase I. Meiosis II Segregates Sister Chromatids into Separate Daughter Cells • Meiosis II cell division is almost exactly like mitosis. • The two haploid cells produced by meiosis I give rise to a total of four haploid cells. • The reduction in chromosome numbers achieved through meiosis I offsets the combining of chromosomes when gametes fuse during fertilization. Meiosis and Fertilization Contribute to Genetic Variation in a Population • Genetic variation is the raw material for evolution. • Mutations are the ultimate source of genetic variation in all types of organisms. • Different variations of a particular gene created through mutations are called alleles. • Through meiosis and fertilization, sexual reproduction produces offspring with new combinations of alleles, adding to the genetic diversity in a population. Multiple embryos inside a shark “egg case.” Just one fetal shark will survive after cannibalizing its siblings. Crossing-Over during Meiosis I Generates Gametes with Recombinant Chromosomes • Meiosis magnifies diversity by shuffling alleles between homologous pairs and then sorting these scrambled homologues randomly into gametes. • Crossing-over is the physical exchange of chromosomal segments between nonsister chromatids in paired-off paternal and maternal homologues. • Crossing-over exchanges alleles between the paternal and maternal chromatids, creating new groupings of alleles through the exchange of DNA segments in a process known as genetic recombination. The Independent Assortment of Homologous Pairs Generates Diverse Gametes • Independent assortment is the random distribution of the different homologous chromosome pairs into daughter cells during meiosis I. • Independent assortment of chromosomes produces gametes that are different from the parent and from each other. • The random events in meiosis, and the random fertilization of a certain egg by a particular sperm, contribute to the genetic uniqueness of offspring. BIOLOGY MATTERS PROGRAMMED CELL DEATH: GOING OUT IN STYLE • Some cells are killed off neatly and systematically when they are no longer needed or to help sculpt tissues during development. Characteristics of death by apoptosis (main type of programmed cell death in human body): • No spillage of cytoplasm, no inflammation • Mitochondria degrade • Proteins degraded • Fragmentation of DNA • Cell shrinks; eventually engulfed and digested inside immune cells APPLYING WHAT WE LEARNED: THE GREAT DIVIDE • Nutrients help cells divide rapidly, as seen in the bloom of “sea lettuce” that threatened to halt the 2008 Olympics. • Eutrophication refers to the overfertilization of an area, typically due to agricultural runoff. • Eutrophication can lead to large algal blooms. List of Key Terms: Chapter 7 anaphase (p. 160) asexual reproduction (p. 150) binary fission (p. 151) cancer cell (p. 156) cell cycle (p. 153) cell differentiation (p. 153) cell division (p. 153) cell plate (p. 161) centromere (p. 157) centrosome (p. 160) chromatin (p. 156) chromosome (p. 156) crossing-over (p. 166) cytokinesis (p. 152) diploid (p. 162) embryo (p. 152) fertilization (p. 152) G0 phase (p. 154) G1 phase (p. 154) G2 phase (p. 154) gamete (p. 152) genetic recombination (p. 166) germ line cell (p. 153) haploid (p. 162) homologous chromosome (p. 158) independent assortment of chromosomes (p. 167) interphase (p. 153) karyotype (p. 157) maternal homologue (p. 163) meiosis (p. 152) meiosis I (p. 163) meiosis II (p. 163) metaphase (p. 160) mitosis (p. 152) mitotic division (p. 152) mitotic spindle (p. 160) paternal homologue (p. 163) prophase (p. 160) S phase (p. 154) sex chromosome (p. 158) sexual reproduction (p. 150) sister chromatids (p. 157) somatic cell (p. 152) telophase (p. 161) tetrad (p. 164) tumor (p. 156) zygote (p. 152) Class Quiz, Part 1 Which of the following is not true about interphase? A. The cell grows larger during interphase. B. Chromosomes are replicated during interphase. C. A cell can go from G1 to S but not from S to G1. D. Chromosomes are segregated to daughter cells during interphase. Class Quiz, Part 2 The HeLa cells in the photo are undergoing mitosis in a lab dish. DNA is stained pink, and the mitotic spindle is yellow. (The multiple pink dots within the nuclei are regions called nucleoli where ribosomal RNA is made.) The cell labeled A is at which stage of mitosis? A. prophase B. metaphase C. anaphase D. telophase Class Quiz, Part 3 Which of the following is true about homologous chromosomes? A. You received both homologues in each pair of homologous chromosomes from your mother. B. The two homologues in each pair are sorted into different daughter cells at the end of meiosis I. C. The two daughter cells produced by mitosis have only one member of each homologous pair. D. Human skin cells have 46 pairs of homologous chromosomes. Class Quiz, Part 4 Meiosis ensures that A. all gamete receives the same alleles. B. chromosome number is doubled in the gametes. C. gametes have a haploid number of chromosomes. D. all paternal chromosomes end up in the same gamete. Class Quiz, Part 5 Compare mitotic cell divisions with meiosis by filling out the table below. The response to the first item is already filled in, by way of example. 1. IN HUMANS, THE CELL UNDERGOING THIS TYPE OF DIVISION IS DIPLOID. 2. A TOTAL OF FOUR DAUGHTER CELLS ARE PRODUCED WHEN ONE PARENT CELL UNDERGOES THIS TYPE OF DIVISION. 3. OUR SKIN MAKES MORE SKIN CELLS USING THIS TYPE OF CELL DIVISION. 4. THE DAUGHTER CELLS ARE GENETICALLY IDENTICAL TO THE PARENT CELL IN THIS TYPE OF DIVISION. 5. THIS TYPE OF DIVISION INVOLVES TWO NUCLEAR DIVISIONS. 6. THIS TYPE OF DIVISION INVOLVES TWO ROUNDS OF CYTOKINESIS. 7. MATERNAL AND PATERNAL HOMOLOGUES PAIR UP TO FORM TETRADS AT SOME POINT DURING THIS TYPE OF DIVISION. 8. SISTER CHROMATIDS SEPARATE FROM EACH OTHER AT SOME POINT DURING THIS TYPE OF CELL DIVISION. MITOTIC CELL DIVISION true MEIOSIS true Relevant Art from Other Chapters All art files from the book are available in JPEG and PPT formats online Cell Division in Prokaryotes Structure of a Prokaryote Asexual and Sexual Reproduction in Eukaryotes Crossing-Over Causes Offspring to Have a Genotype That Differs from the Genotype of Either Parent • Crossing-over is a reciprocal exchange of segments of nonsister chromatids in prophase I of meiosis, which produces chromosomes with new combinations of alleles. The Sorting of One Pair of Homologous Chromosomes Is Independent of the Sorting of Any Other Pair of Homologous Chromosomes • Which homologue of each pair goes to which pole is essentially random. 7.1 Concept Check, Part 1 1. In what way is binary fission similar to mitotic cell division? Give one difference. ANSWER: Both produce genetically identical daughter cells, but only prokaryotes undergo binary fission. 7.1 Concept Check, Part 2 2. What is the function of mitosis? ANSWER: Mitosis is necessary for asexual reproduction in both single-celled and multicellular eukaryotes, and for growth and repair of the multicellular body. 7.2 Concept Check, Part 1 1. When in the cell cycle does copying (replication) of DNA take place? ANSWER: In the S (synthesis) phase of interphase. 7.2 Concept Check, Part 2 2. What is the significance of the G₀ phase? ANSWER: A cell that leaves the cell cycle and stops dividing is in the G₀ phase, either temporarily or permanently. 7.3 Concept Check, Part 1 1. The house cat karyotype displays a total of 38 chromosomes. How many distinct DNA molecules are present in a cat skin cell toward the end of (a) G₀ phase; (b) G₁ phase; (c) S phase; (d) G₂ phase? ANSWER: (a) 38 (b) 38 (c) 76 (d) 76 7.3 Concept Check, Part 2 2. How is the karyotype of men different from that of women? ANSWER: All humans have 22 pairs of autosomes and a pair of sex chromosomes. Women have a pair of identical sex chromosomes called X chromosomes. Men have two dissimilar sex chromosomes: one X and one Y. 7.4 Concept Check, Part 1 1. What are sister chromatids? When during mitosis do they separate to become chromosomes in their own right? ANSWER: Sister chromatids are two identical DNA molecules, replicated from a single parental chromosome, that remain attached during mitosis until anaphase, when they separate to become two independent chromosomes, each headed to a different daughter cell. 7.4 Concept Check, Part 2 2. What is the difference between mitosis and cytokinesis? ANSWER: Mitosis is nuclear division; cytokinesis is the partitioning of the cytoplasm, and ultimately the whole parent cell, into two daughter cells. 7.5 Concept Check, Part 1 1. Meiosis I reduces the diploid chromosome set. What role does meiosis II play in gamete formation? ANSWER: Meiosis II separates the two sister chromatids of each replicated chromosome, to form gametes with 23 chromosomes each. 7.5 Concept Check, Part 2 2. How does meiosis contribute to genetic variation in a population? ANSWER: Meiosis generates genetically diverse gametes through two different mechanisms: recombination of alleles via crossing-over, and random assortment of homologous chromosomes.