2/17/16 BIOLOGY 207 - Dr.Locke Lecture#11 - Chromosomes and their behavior in meiosis Required readings and problems: Reading: Open Genetics, Chapter 2 Problems: Chapter 2 Optional Griffiths (2008) 9th Ed. Readings: 32-57 (N.B. 47-49), 75-76, 103-105 Problems: 9th Ed. Ch. 3: 1-22 (not 18) Campbell (2008) 8th Ed. Readings: Concept 14.1 Concepts: How do chromosomes behave in meiosis?? 1. A characteristic chromosome set (karyotype) is specific for a species. 2. The process of meiosis occurs in diploid cells, called meiocytes, and produces four haploid nuclei (the meiotic products). 3. The synapsis of structurally (and genetically) similar chromosomes (homologues) leads to the orderly reduction in chromosome numbers in the first division of meiosis. 4. Random segregation of chromosomes and chromosome crossing-over are two features that result in genetic diversity. Biol207 Dr. Locke section Lecture#11 Fall'11 page 1 2/17/16 Cytogenetics: - cytology to study heredity (in eukaryotes) - use normal genome to identify the standard set of chromosomes in a species - use normal chromosome organization to identify abnormal The normal set is based on several principles: 1) The chromosomes from a cell are usually observed at the metaphase stage of mitosis - this is a standard point in the cell cycle to see well differentiated, separate chromosomes 2) Since all the cells of an organism have the same genetic material (same quality, quantity and organization), then the chromosomes of all body cells should be same. -extrapolate from one cell --> all cells (not always true) 3) Because different individuals of a species have their DNA organized in a similar manner the organization of their chromosomes will reflect this and be similar within the species -extrapolate: one individual --> all individuals in species Biol207 Dr. Locke section Lecture#11 Fall'11 page 2 2/17/16 Standard Metaphase chromosomes Morphological differences among chromosomes make it possible to distinguish chromosomes. Identify different chromosomes based on: 1) Size - overall length of the chromosome - varies among those in the set 2) Centromere position (primary constriction) - location of the spindle attachment telocentric - centromere is at the end, telomere acrocentic - centromere is off centre metacentric - centromere is in the middle 3) Secondary constrictions - chromosome thickness varies along the length - Nucleolus Organizer Regions (NOR) - contain the genes for rRNA - one or a few sites per genome (not necessarily one on each chromosome) - also other sites of 2° constrictions (not NOR) 4) Banding patterns - there are special staining procedures that reveal differences in staining (or bands) along each chromosome. -Each chromosome has a specific, reproducible, unique pattern that distinguishes it from the others. Biol207 Dr. Locke section Lecture#11 Fall'11 page 3 2/17/16 Result of staining and cytology: a Karyotype Each chromosome can be identified and assembled together to obtain a karyotype. Karyotype - refers to a set of photographs organizing the metaphase chromosomes based on number, size, and morphology for a cell, individual, or species Idiogram - a diagrammatic representation of the karyotype - standard diagram of the chromosomes Karyotype of a Diploid organism The karyotype would consist of "pairs of chromosomes" - one from father - one from mother - each "set" is a genome or "N" value's worth of chromosomes Example: eg. humans 46 chromosomes (23 pairs) 1N = 23 from mother - maternal in origin - N in the eggs 1N = 23 from father - paternal in origin - N in the sperm 2N = 46 chromosomes Each species has its own chromosome set - Each species' chromosome set can be defined based on number, size, and structure - There is limited variation for normal and extreme differences are usually mutants Example: - human chromosome #15 are the same, however, close observation shows that slight differences do exist among the chromosomes in the population Biol207 Dr. Locke section Lecture#11 Fall'11 page 4 2/17/16 Mitosis: 1 cell -> 2 cells – stability! Cell cycle: G1, S, G2, Mitosis --> two daughter cells - chromosome replication followed by mitosis maintains the chromosome complement at each cell generation Diploid cell Fig - replication and mitosis Haploid cell Fig - process is exactly the same Note: - chromosome number and organization are maintained through each cell generation. - daughter cells are genetically equivalent to the parent. Meiosis - 1 cell --> 4 cells (nuclei) – diversity! - germline cells (not somatic) called Meiocyte - meiocyte will produce gametes- See Fig - in males -> sperm - in females -> eggs - Diploid - made from 2 chromosome sets - 1 maternal + 1 paternal --> normal S-phase - undergo chromosome replication -->each chromosome has 2 sister chromatids Then the cell enters meiosis - process has 2 cell divisions Meiosis I (MI) and Meiosis II (MII) - both divisions follow the same basic steps (prophase, metaphase, anaphase, telophase) - the events in each are different and different from mitosis Biol207 Dr. Locke section Lecture#11 Fall'11 page 5 2/17/16 Meiosis I Prophase I - complex and lengthy - 5 separate cytological stages along a dynamic continuum 1) Leptotene - The interphase nucleus begins to condense into visible long, thin threads (chromosomes) 2) Zygotene - homologous chromosomes begin to pair up or synapse - pair up at localized points, then "zipper" up along the whole chromosome - each homolog of a pair join together through an elaborate structure called the synaptonemal complex. 3) Pachytene - chromosomes are fully synapsed as a bivalent (bi = two chromosomes) - the 2 chromosomes have 4 chromatids ****** crossing over takes place ******* - # of bivalents = # of chromosomes in one set = n 4) Diplotene - pairing loosens and chromosomes start to separate such that the chromatids become visually apparent - chiasmata appear and are the visible result of a crossover event that had occurred earlier (during pachytene) - normally see one (or more) chiasmata per chromosome pair - chiasmata contribute to proper chromosome segregation during meiosis 5) Diakinesis – continuation/extension of Diplotene - further chromosome contraction - form compact chromosomes which are easier to move to daughter cells than larger chromosomes Biol207 Dr. Locke section Lecture#11 Fall'11 page 6 2/17/16 Metaphase I - Nuclear membrane and nucleoli disappear and the chromosomes moved to the equatorial plane by the spindle apparatus (filaments of microtubules) that attach to the centromere of each chromosome - chromosomes align such that homologous chromosomes face opposite poles - Note: both orientations are possible - for any given chromosome pair the orientation happens at random Anaphase I - homologous chromosome centromeres begin to move towards opposite poles Telophase I - chromosomes reach the poles and may decondense - not ture in all organisms, some proceed directly to MII Result of MI = reduction division - homologous chromosome to each pole - each resulting nucleus (cell) now has only one set of chromosomes (1n) - one homolog from each pair of homologous chromosomes - each chromosome still has two chromatids Interphase - transient - interkinesis - no DNA replication Biol207 Dr. Locke section Lecture#11 Fall'11 page 7 2/17/16 Next: Meiosis II - equational division occurs in each of the two MI products Prophase II - haploid chromosome number - chromosomes condense and shorten Metaphase II - chromosomes move to equatorial plane - each chromosome has 2 chromatids Anaphase II - centromeres split and sister chromatids move to opposite poles Note: chromatids now become chromosomes Telophase II - four product nuclei produced - 4 X 1N products of meiosis from one diploid (2N) meiocyte The meiotic products which are haploid, such as pollen, sperm, etc., Gamete Will go on and unite with a complementary gamete (in the process of Syngamy) to form a zygote, which is 2N again (diploid). Biol207 Dr. Locke section Lecture#11 Fall'11 page 8 2/17/16 Meiosis Notes: - See animation on the WWW - each chromosome pair orients independently in MI, the reductional division - each chromatid orients independently in MII, the equational division - get the independent assortment of chromosomes and of the alleles on those chromosomes - meiosis leads to diversity among the gametes (not the same as the meiocyte) and therefore of the progeny. There is a second feature, in addition to independent assortment, that leads to genetic diversity: Crossing over - Genetic Recombination During prophase I there is a physical exchange that can occur between non-sister chromatids. This exchanges the alleles present on homologous chromosomes. This process mixes up the combination of alleles further (). Follow the chromatids through MI - crossover results in different meiotic products being produced - chromosomes are different in that they contain parts of both original parental chromosomes. - part maternal - part paternal Biol207 Dr. Locke section Lecture#11 Fall'11 page 9 2/17/16 Biol207 Dr. Locke section Lecture#11 Fall'11 page 10