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Chapter 13 Meiosis and Sexual Life Cycles Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Q: What accounts for family resemblance? Fig. 13-1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Overview: Variations on a Theme • Living organisms are distinguished by their ability to reproduce their own kind • Genetics is the scientific study of heredity and variation • Heredity is the transmission of traits from one generation to the next • Variation is demonstrated by the differences in appearance that offspring show from parents and siblings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Q: 자손은 어떻게 부모와 닮은 형질을 획득하게 되나? 13.1: Offspring acquire genes from parents by • Concept inheriting chromosomes • In a literal sense, children do not inherit particular physical traits from their parents • It is genes that are actually inherited Inheritance of Genes • Genes are the units of heredity, and are made up of segments of DNA • Genes are passed to the next generation through reproductive cells called gametes (sperm and eggs) • Each gene has a specific location called a locus (=address) on a certain chromosome • Most DNA is packaged into chromosomes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Comparison of Asexual and Sexual Reproduction • In asexual reproduction – One parent produces genetically identical offspring by mitosis A hydra reproduces by budding. The bud, a localized mass of mitotically dividing cells, develops into a small hydra, which detaches from the parent Bud Figure 13.2 (a) An individual that reproduces asexually gives rise to a clone, a group of genetically Parent identical individuals from the same parent Genetic differences in asexually reproducing organisms can be arisen by changes in the DNA called mutations (See Chapter17) 0.5 mm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In sexual reproduction – Two parents give rise to offspring that have unique combinations of genes inherited from the two parents Figure 13.2(b) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Redwoods Q: 유성생식 생활사의 특징은? • Concept 13.2: Fertilization and meiosis alternate in sexual life cycles • A life cycle is the generation-to-generation sequence of stages in the reproductive history of an organism Q: 염색체의 행동패턴은 인간 생활사와 다른 유성생식 과정과 어떤 관련성이 있나? Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Q: 인간세포에서 발견되는 염색체 세트의 차이는? • Human somatic cells (any cell other than a gamete) have 23 pairs of chromosomes • A karyotype is an ordered display of the pairs of chromosomes from a cell • The two chromosomes in each pair are called homologous chromosomes, or homologs • Chromosomes in a homologous pair are the same length and carry genes controlling the same inherited characters • The sex chromosomes (성염색체) are called X and Y • Human females have a homologous pair of X chromosomes (XX) • Human males have one X and one Y chromosome • The 22 pairs of chromosomes that do not determine sex are called autosomes (상염색체=보통염색체) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Each pair of homologous chromosomes includes one chromosome from each parent • The 46 chromosomes in a human somatic cell are two sets of 23: one from the mother and one from the father • A diploid cell (2n) has two sets of chromosomes • For humans, the diploid number is 46 (2n = 46) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A gamete (sperm or egg) contains a single set of chromosomes, and is haploid (n) • For humans, the haploid number is 23 (n = 23) • Each set of 23 consists of 22 autosomes and a single sex chromosome • In an unfertilized egg (ovum), the sex chromosome is X • In a sperm cell, the sex chromosome may be either X or Y Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 다음과 같이 2n=6인 세포를 가정했을 때 아래의 질문에 답하시오 (1) What is the haploid number of the following cell? (2) Is a “set” of chromosomes haploid or diploid? Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 인간 생활사에서 염색체의 행동 패턴은? • The human life cycle begins when a haploid sperm from the father fuses with a haploid egg from the mother • Fertilization is the union of gametes (the sperm and the egg) • The fertilized egg is called a zygote and has one set of chromosomes from each parent • The zygote produces somatic cells by mitosis and develops into an adult • At sexual maturity, the ovaries and testes produce haploid gametes • Gametes are the only types of human cells produced by meiosis, rather than mitosis • Meiosis results in one set of chromosomes in each gamete • Fertilization and meiosis alternate in sexual life cycles to maintain chromosome number Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The human life cycle Key Haploid gametes (n = 23) Haploid (n) Ovum (n) Diploid (2n) Sperm Cell (n) FERTILIZATION MEIOSIS Ovary Testis Mitosis and development Figure 13.5 Multicellular diploid adults (2n = 46) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Diploid zygote (2n = 46) 유성생식의 다양성 • The alternation of meiosis and fertilization (key events that contribute to genetic variation among offspring) is common to all organisms that reproduce sexually • The three main types of sexual life cycles differ in the timing of meiosis and fertilization Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Three types of sexual life cycles: animals • Gametes are the only haploid cells in animals • They are produces by meiosis and undergo no further cell division before fertilization • Gametes fuse to form a diploid zygote that divides by mitosis to develop into a multicellular organism Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Three types of sexual life cycles: Plants and algae • Plants and some algae exhibit an alternation of generations • This life cycle includes both a diploid and haploid multicellular stage • The diploid organism, called the sporophyte, makes haploid spores by meiosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Three types of sexual life cycles: fungi and protists • In most fungi and some protists, the only diploid stage is the singlecelled zygote; there is no multicellular diploid stage • The zygote produces haploid cells by meiosis • Each haploid cell grows by mitosis into a haploid multicellular organism • The haploid adult produces gametes by mitosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 유성생식 유형간의 차이점과 유사점 • Depending on the type of life cycle, either haploid or diploid cells can divide by mitosis • However, only diploid cells can undergo meiosis • In all three life cycles, the halving and doubling of chromosomes contributes to genetic variation in offspring Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 감수분열이란? • Concept 13.3: Meiosis reduces the number of chromosome sets from diploid to haploid • Like mitosis, meiosis is preceded by the replication of chromosomes • Meiosis takes place in two sets of cell divisions, called meiosis I and meiosis II • The two cell divisions result in four daughter cells, rather than the two daughter cells in mitosis • Each daughter cell has only half as many chromosomes as the parent cell Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Stages of Meiosis (see Next Figure 13.7) • • After chromosomes duplicate, two divisions follow – Meiosis I (reductional division): homologs pair up and separate, resulting in two haploid daughter cells with replicated chromosomes – Meiosis II (equational division) sister chromatids separate The result is four haploid daughter cells with unreplicated chromosomes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Outline of Meiosis Ref: Chapter 4 of Genetics edited by Hartwell et al. Female (2n) Male (2n) Ovary : Testis : Germ cells (2n) Germ cells (2n) Meiosis Eggs (n) Sperms (n) Gametes Fertilization Zygote (=embryo: 2n) Mitosis (100 times in somatic cells) Offspring (2n) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 제1 감수분열의 특징 • Meiosis I is preceded by interphase, when the chromosomes are duplicated to form sister chromatids • The sister chromatids are genetically identical and joined at the centromere • The single centrosome replicates, forming two centrosomes 제1 감수분열의 진행단계 • Division in meiosis I occurs in four phases – Prophase I – Metaphase I – Anaphase I – Telophase I and cytokinesis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 13.8a Prophase I Centrosome (with centriole pair) Sister chromatids Chiasmata Spindle Telophase I and Cytokinesis Anaphase I Metaphase I Sister chromatids remain attached Centromere (with kinetochore) Metaphase plate Homologous chromosomes Fragments of nuclear envelope Homologous chromosomes separate Microtubule attached to kinetochore Each pair of homologous chromosomes separates. Chromosomes line up Duplicated homologous chromosomes (red and blue) by homologous pairs. pair and exchange segments; 2n 6 in this example. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Cleavage furrow Two haploid cells form; each chromosome still consists of two sister chromatids. Prophase I • Prophase I typically occupies more than 90% of the time required for meiosis • Longest (90% of meiosis), most complex phase : takes many days, months, even years (arrested at prophase I until ovulation) • Chromosomes begin to condense • In synapsis, homologous chromosomes loosely pair up, aligned gene by gene • In crossing over, nonsister chromatids exchange DNA segments • Each pair of chromosomes forms a tetrad, a group of four chromatids • Each tetrad usually has one or more chiasmata, X-shaped regions where crossing over occurred Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Metaphase I • • • In metaphase I, tetrads line up at the metaphase plate, with one chromosome facing each pole Microtubules from one pole are attached to the kinetochore of one chromosome of each tetrad Microtubules from the other pole are attached to the kinetochore of the other chromosome Anaphase I • In anaphase I, pairs of homologous chromosomes separate • One chromosome moves toward each pole, guided by the spindle apparatus • Sister chromatids remain attached at the centromere and move as one unit toward the pole Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Telophase I and Cytokinesis • In the beginning of telophase I, each half of the cell has a haploid set of chromosomes; each chromosome still consists of two sister chromatids • Cytokinesis usually occurs simultaneously, forming two haploid daughter cells • In animal cells, a cleavage furrow forms; in plant cells, a cell plate forms • No chromosome replication occurs between the end of meiosis I and the beginning of meiosis II because the chromosomes are already replicated Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Meiosis – Prophase I (“thin” in Greek) (“Conjugation”) (“thick”) (=zipper with remarkable precision) Pachytene Figure 4.13 bivalent: each synapsed chromosome pair (=tetrad for four chromatids) Pairing b/w X and Y chromosomes: limited pairing via a small region of homology Recombination nodules: a structure appeared along the synaptonemal complex; a site of crossing-over b/w nonsister chromaitds (maternal & paternal) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Meiosis-Prophase I-continued Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mechanism of Crossing-over Crossing over during prophase produces recombined chromosomes Bivalent or tetrad Fig. 4.14 a-c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mechanism of Crossing-over Fig. 4.14 d, e Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Meiosis I – Metaphase and Anaphase Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Meiosis – Telophase I and Interkinesis Fig. 4.13e Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Meiosis – Prophase II and Metaphase II Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 제2 감수분열의 진행단계 • Division in meiosis II also occurs in four phases: – Prophase II – Metaphase II – Anaphase II – Telophase II and cytokinesis • Meiosis II is very similar to mitosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 13.8b Prophase II Metaphase II Anaphase II Telophase II and Cytokinesis During another round of cell division, the sister chromatids finally separate; four haploid daughter cells result, containing unduplicated chromosomes. Sister chromatids separate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Haploid daughter cells forming Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings A Comparison of Mitosis and Meiosis • Mitosis conserves the number of chromosome sets, producing cells that are genetically identical to the parent cell • Meiosis reduces the number of chromosomes sets from two (diploid) to one (haploid), producing cells that differ genetically from each other and from the parent cell • The mechanism for separating sister chromatids is virtually identical in meiosis II and mitosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 13.9b SUMMARY Property Mitosis Meiosis DNA replication Occurs during interphase before mitosis begins Occurs during interphase before meiosis I begins Number of divisions One, including prophase, metaphase, anaphase, and telophase Two, each including prophase, metaphase, anaphase, and telophase Synapsis of homologous chromosomes Does not occur Occurs during prophase I along with crossing over between nonsister chromatids; resulting chiasmata hold pairs together due to sister chromatid cohesion Number of daughter cells and genetic composition Two, each diploid (2n) and genetically identical to the parent cell Four, each haploid (n), containing half as many chromosomes as the parent cell; genetically different from the parent cell and from each other Role in the animal body Enables multicellular adult to arise from zygote; produces cells for growth, repair, and, in some species, asexual reproduction Produces gametes; reduces number of chromosomes by half and introduces genetic variability among the gametes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Three events are unique to meiosis, and all three occur in meiosis l – Synapsis and crossing over in prophase I: Homologous chromosomes physically connect and exchange genetic information – At the metaphase plate, there are paired homologous chromosomes (tetrads), instead of individual replicated chromosomes – At anaphase I, it is homologous chromosomes, instead of sister chromatids, that separate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Inquiry: What prevents the separation of sister chromatids at anaphase I of meiosis? • Sister chromatid cohesion allows sister chromatids of a single chromosome to stay together through meiosis I • Protein complexes called cohesins are responsible for this cohesion • In mitosis, cohesins are cleaved at the end of metaphase • In meiosis, cohesins are cleaved along the chromosome arms in anaphase I (separation of homologs) and at the centromeres in anaphase II (separation of sister chromatids) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 유성생식과정에서 발생한 유전변이의 역할은?13.4: Genetic variation produced • Concept sexual life cycles contributes to evolution in • Mutations (changes in an organism’s DNA) are the original source of genetic diversity (유전적 다양성) • Mutations create different versions of genes called alleles (대립유전자) • Reshuffling of alleles during sexual reproduction produces genetic variation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Origins of Genetic Variation Among Offspring • The behavior of chromosomes during meiosis and fertilization is responsible for most of the variation that arises in each generation • Three mechanisms contribute to genetic variation: Independent Assortment of Chromosomes Crossing Over Random Fertilization Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Independent Assortment of Chromosomes • Homologous pairs of chromosomes orient randomly at metaphase I of meiosis • In independent assortment, each pair of chromosomes sorts maternal and paternal homologues into daughter cells independently of the other pairs • The number of combinations possible when chromosomes assort independently into gametes is 2n, where n is the haploid number • For humans (n = 23), there are more than 8 million (223) possible combinations of chromosomes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Crossing Over • Crossing over produces recombinant chromosomes, which combine DNA inherited from each parent • Crossing over begins very early in prophase I, as homologous chromosomes pair up gene by gene • In crossing over, homologous portions of two nonsister chromatids trade places • Crossing over contributes to genetic variation by combining DNA from two parents into a single chromosome Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Random Fertilization • Random fertilization adds to genetic variation because any sperm can fuse with any ovum (unfertilized egg) • The fusion of two gametes (each with 8.4 million possible chromosome combinations from independent assortment) produces a zygote with any of about 70 trillion(조) diploid combinations No wonder brothers and sisters can be so different: You really are unique!!! Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Evolutionary Significance of Genetic Variation Within Populations • Natural selection results in the accumulation of genetic variations favored by the environment • Sexual reproduction contributes to the genetic variation in a population, which originates from mutations Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Privet shrubs and humans each have a diploid number of 46 chromosomes per cell. Why are the two species so dissimilar? (a) Privet chromosomes undergo only mitosis. (b) Privet chromosomes are shaped differently. (c) Human chromosomes have genes grouped together differently. (d) The two species have appreciably different genes. (e) Privets do not have sex chromosomes. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings