Meiosis and Sexual Life Cycles • • • Living organisms are distinguished by their ability to reproduce their own kind Heredity – Is the transmission of traits from one generation to the next Variation – Shows that offspring differ somewhat in appearance from parents and siblings 1 Inheritance of Genes • • • Genes are segments of DNA, units of heredity Offspring acquire genes from parents by inheriting chromosomes Genetics is the scientific study of heredity and hereditary variation 2 Inheritance of Genes • • • Each gene in an organism’s DNA has a specific locus on a certain chromosome We inherit one set of chromosomes from our mother and one set from our father Two parents give rise to offspring that have unique combinations of genes inherited from the two parents - sexual reproduction 3 Asexual Reproduction • In asexual reproduction, one parent produces genetically identical offspring by mitosis Parent Bud Figure 13.2 0.5 mm 4 Sexual Reproduction • • 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 5 Sex Cells - Gametes • • Unlike somatic cells, sperm and egg cells are haploid cells, containing only one set of chromosomes At sexual maturity the ovaries and testes produce haploid gametes by meiosis 6 Meiosis • • • Reduces the chromosome number such that each daughter Cell has a haploid set of chromosomes Ensures that the next generation will have: – Diploid number of chromosome – Exchange of genetic information (combination of traits – that differs from that of either parent) 7 Meiosis • • • • Only diploid cells can divide by meiosis. Prior to meiosis I, DNA replication occurs. During meiosis, there will be two nuclear divisions, and the result will be four haploid nuclei. No replication of DNA occurs between meiosis I and meiosis II. 8 Sexual Life Cycles • • The three main types of sexual life cycles – Animals – Plants and Algae – Fungi and Protists Differ in the timing of meiosis and fertilization 9 Plants and Some Algae • • Exhibit an alternation of generations The life cycle includes both diploid and haploid multicellular stages Haploid multicellular organism (gametophyte) n Mitosis n Mitosis n n n Spores Gametes MEIOSIS Diploid multicellular organism (sporophyte) 2n FERTILIZATION 2n Zygote Mitosis (b) Plants and some algae 10 Most Fungi and Some Protists • • Meiosis produces haploid cells that give rise to a haploid multicellular adult organism The haploid adult carries out mitosis, producing cells that will become gametes Haploid multicellular organism Mitosis n Mitosis n n n Gametes MEIOSIS n FERTILIZATION 2n Zygote (c) Most fungi and some protists 11 Animals Key • • Meiosis occurs during gamete formation Gametes are the only haploid cells Haploid Diploid n n Gametes n MEIOSIS FERTILIZATION Zygote 2n Diploid multicellular organism 2n Mitosis (a) Animals 12 Meiosis Interphase • • Meiosis reduces the number of chromosome sets from diploid to haploid Meiosis takes place in two sets of divisions – – Meiosis I reduces the number of chromosomes from diploid to haploid Meiosis II produces four haploid daughter cells Figure 13.7 Homologous pair of chromosomes in diploid parent cell Chromosomes replicate Homologous pair of replicated chromosomes Sister chromatids Diploid cell with replicated chromosomes Meiosis I 1 Homologous chromosomes separate Haploid cells with replicated chromosomes Meiosis II 2 Sister chromatids separate Haploid cells with unreplicated chromosomes 13 Meiosis Phases • • • • Meiosis involves the same four phases seen in mitosis prophase metaphase anaphase telophase They are repeated during both meiosis I and meiosis II. The period of time between meiosis I and meiosis II is called interkinesis. No replication of DNA occurs during interkinesis because the DNA is already duplicated. 14 Prophase I • • • • • Prophase I occupies more than 90% of the time required for meiosis Chromosomes begin to condense In synapsis, the 2 members of each homologous pair of chromosomes line up side-by-side, aligned gene by gene, to form a tetrad consisting of 4 chromatids During synapsis, sometimes there is an exchange of homologous parts between non-sister chromatids. This exchange is called crossing over Each tetrad usually has one or more chiasmata, X-shaped regions where crossing over occurred Nonsister chromatids Prophase I of meiosis Tetrad Chiasma, site of crossing over 15 Metaphase I • • • At 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 PROPHASE I Sister chromatids Tetrad METAPHASE I ANAPHASE I Sister chromatids remain attached Centromere (with kinetochore) Chiasmata Metaphase plate Spindle Microtubule attached to kinetochore Homologous chromosomes (red and blue) pair and exchange segments; 2n = 6 Homologous chromosomes separate Tetrads line up Pairs of homologous chromosomes split up 16 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 PROPHASE I Sister chromatids Tetrad METAPHASE I ANAPHASE I Sister chromatids remain attached Centromere (with kinetochore) Chiasmata Metaphase plate Spindle Microtubule attached to kinetochore Homologous chromosomes (red and blue) pair and exchange segments; 2n = 6 Homologous chromosomes separate Tetrads line up Pairs of homologous chromosomes split up 17 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 18 Prophase II • • • Meiosis II is very similar to mitosis In prophase II, a spindle apparatus forms In late prophase II, chromosomes (each still composed of two chromatids) move toward the metaphase plate TELOPHASE I AND CYTOKINESIS PROPHASE II Cleavage furrow METAPHASE II ANAPHASE II Sister chromatids separate TELOPHASE II AND CYTOKINESIS Haploid daughter cells forming 19 Metaphase II • • • At metaphase II, the sister chromatids are at the metaphase plate Because of crossing over in meiosis I, the two sister chromatids of each chromosome are no longer genetically identical The kinetochores of sister chromatids attach to microtubules extending from opposite poles TELOPHASE I AND CYTOKINESIS PROPHASE II Cleavage furrow METAPHASE II ANAPHASE II Sister chromatids separate TELOPHASE II AND CYTOKINESIS Haploid daughter cells forming 20 Anaphase II • • At anaphase II, the sister chromatids separate The sister chromatids of each chromosome now move as two newly individual chromosomes toward opposite poles TELOPHASE I AND CYTOKINESIS PROPHASE II Cleavage furrow METAPHASE II ANAPHASE II Sister chromatids separate TELOPHASE II AND CYTOKINESIS Haploid daughter cells forming 21 Telophase II and Cytokinesis • • • • • In telophase II, the chromosomes arrive at opposite poles Nuclei form, and the chromosomes begin decondensing Cytokinesis separates the cytoplasm At the end of meiosis, there are four daughter cells, each with a haploid set of unreplicated chromosomes Each daughter cell is genetically distinct from the others and from the parent cell TELOPHASE I AND CYTOKINESIS PROPHASE II Cleavage furrow METAPHASE II ANAPHASE II Sister chromatids separate TELOPHASE II AND CYTOKINESIS Haploid daughter cells forming 22 Interphase and meiosis I MEIOSIS I: Separates homologous chromosomes INTERPHASE PROPHASE I METAPHASE I ANAPHASE I Sister chromatids remain attached Centromere (with kinetochore) Centrosomes (with centriole pairs) Sister chromatids Chiasmata Spindle Metaphase plate Nuclear envelope Homologous Microtubule chromosomes Tetrad attached to Chromatin separate kinetochore Pairs of homologous Chromosomes duplicate Tertads line up chromosomes split up Homologous chromosomes (red and blue) pair and exchange segments; 2n = 6 in this example 23 Telophase I, cytokinesis, and meiosis II MEIOSIS II: Separates sister chromatids TELOPHASE I AND CYTOKINESIS PROPHASE II Cleavage furrow Two haploid cells form; chromosomes are still double METAPHASE II ANAPHASE II Sister chromatids separate TELOPHASE II AND CYTOKINESIS Haploid daughter cells forming During another round of cell division, the sister chromatids finally separate; four haploid daughter cells result, containing single chromosomes 24 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 25 A Comparison of Mitosis and Meiosis • 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 of meiosis, homologous pairs move toward opposite poles of the cell. In anaphase II of meiosis, the sister chromatids separate 26 A Comparison Of Mitosis And Meiosis MITOSIS MEIOSIS Chiasma (site of crossing over) Parent cell (before chromosome replication) MEIOSIS I Prophase I Prophase Chromosome replication Duplicated chromosome (two sister chromatids) Chromosome replication Tetrad formed by synapsis of homologous chromosomes 2n = 6 Chromosomes positioned at the metaphase plate Metaphase Sister chromatids separate during anaphase Anaphase Telophase 2n Tetrads positioned at the metaphase plate Homologues separate during anaphase I; sister chromatids remain together Metaphase I Anaphase I Telophase I Haploid n=3 Daughter cells of meiosis I 2n MEIOSIS II Daughter cells of mitosis n n n n Daughter cells of meiosis II Sister chromatids separate during anaphase II 27 Comparison • • • • • • Meiosis DNA duplication followed by 2 cell divisions Sysnapsis Crossing-over One diploid cell produces 4 haploid cells Each new cell has a unique combination of genes • • • • • Mitosis Homologous chromosomes do not pair up No genetic exchange between homologous chromosomes One diploid cell produces 2 diploid cells or one haploid cell produces 2 haploid cells New cells are genetically identical to original cell (except for mutation) 28 Genetic Variation Produced In Sexual Life Cycles Contributes To Evolution • • • Mutations (changes in an organism’s DNA) are the original source of genetic diversity Mutations create different versions of genes Reshuffling of different versions of genes during sexual reproduction produces genetic variation 29 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 30 Independent Assortment • • • • 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 31 Independent Assortment of Chromosomes • Each pair of chromosomes sorts its maternal and paternal homologues into daughter cells independently of the other pairs Key Maternal set of chromosomes Paternal set of chromosomes Possibility 1 Possibility 2 Two equally probable arrangements of chromosomes at metaphase I Metaphase II Daughter cells Combination 1 Combination 2 Combination 3 Combination 4 32 Crossing Over • • • • Crossing over produces recombinant chromosomes, which combine genes 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 33 Crossing Over • Crossing over produces recombinant chromosomes that carry genes derived from two different parents Prophase I of meiosis Nonsister chromatids Tetrad Chiasma, site of crossing over Metaphase I Metaphase II Daughter cells Recombinant chromosomes 34 Random Fertilization • • • • Random fertilization adds to genetic variation because any sperm can fuse with any ovum (unfertilized egg) The fusion of gametes produces a zygote with any of about 64 trillion diploid combinations Crossing over adds even more variation Each zygote has a unique genetic identity 35 Evolutionary Significance of Genetic Variation Within Populations • • Natural selection results in accumulation of genetic variations favored by the environment Sexual reproduction contributes to the genetic variation in a population, which ultimately results from mutations 36 Sexual Reproduction - The Human Life Cycle Haploid gametes (n = 23) • • During fertilization, sperm and ovum fuse forming a diploid zygote The zygote develops into an adult organism Haploid (n) Diploid (2n) Ovum (n) Sperm Cell (n) FERTILIZATION MEIOSIS Ovary Testis Diploid zygote (2n = 46) Mitosis and development Multicellular diploid adults (2n = 46) 37 38 Preparing a Karyotype APPLICATION A karyotype is a display of condensed chromosomes arranged in pairs. Karyotyping can be used to screen for abnormal numbers of chromosomes or defective chromosomes associated with certain congenital disorders, such as Down syndrome. TECHNIQUE Karyotypes are prepared from isolated somatic cells, which are treated with a drug to stimulate mitosis and then grown in culture for several days. A slide of cells arrested in metaphase is stained and then viewed with a microscope equipped with a digital camera. A digital photograph of the chromosomes is entered into a computer, and the chromosomes are electronically rearranged into pairs according to size and shape. RESULTS This karyotype shows the chromosomes from a normal human male. The patterns of stained bands help identify specific chromosomes and parts of chromosomes. Although difficult to discern in the karyotype, each metaphase chromosome consists of two, closely attached sister chromatids (see diagram). Pair of homologous chromosomes 5 µm Centromere Sister chromatids 39