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Biology 331: Chapter 3 Chromosomal Basis of Heredity Meiosis & Mitosis: Review your Intro Biology notes If you have difficulty stop by my office Meiosis...a review: Interphase: – Chromosomes and centrosomes replicate • Much like mitosis Meiosis I: Prophase I: – Chromosomes begin to condense – Synapsis: Homolgous chromosomes come together as pairs – Tetrad: A complex of four sister chromatids (2 from each homolog chromosome) – Chiasmata: Where homologous chromatids "cross over" – The exchange of genetic material – 90% of the time for meiosis Meiosis Diagram Crossing Over Meiosis I continued Metaphase I: Anaphase I: – Paired sister chromatids of homologous chromosomes move toward opposite poles – Sister chromatids do not separate Telophase I and cytokinesis: Meiosis Diagram Meiosis II: Prophase II: Metaphase II: Anaphase II: – Centromeres of sister chromatids separate – Individual chromosomes move toward opposite poles Telophase II and cytokinesis: Each cell is now haploid Meiosis Diagram Mitosis –vs- Meiosis Some origins of genetic variation in sexually reproducing organisms: Without variation there is no Evolution! Independent assortment of chromosomes to gametes: Which of the homologous chromosomes is included in a gamete is random The chances are 50-50 for each chromosome Which of your parents chromosomes goes into a gamete 100% maternal, 100% paternal, or some combination The number of possible of possible combinations of gamete is 2n (n is the haploid number) For humans this is 823 or about 8 million combinations Random fertilization: Which sperm meets which egg? Multiply the possible gametes of each parent together Assuming 2 parents with 8 million possible gametes you get 64 trillion combinations Crossing over: Individual chromosomes in a gamete do not have to be exclusively maternal or paternal When chiasmata form portions of chromosomes can switch places In humans you get ~2-3 crossover events per chromosome pair! Number of possible combinations is nearly infinite Crossing Over Life Cycles: If you get chromosomes from both parents how do you avoid a doubling of genetic material? Meiosis: The process of creating gametes with half the normal chromosome complement Review from Intro. Bio. I & II Diploid Life Cycles Common in animals Gametes are the only haploid cells Meiosis occurs during the production of gametes Gametes do not undergo further division until fertilization Diploid zygotes undergo mitosis to form a multicellular adult Diploid Life Cycle Haploid life cycle Most fungi and some algae: Gametes fuse to form a transient diploid zygote Gametes undergo meiosis to form two haploid cells These haploid cells undergo mitosis to form a multicellular adult The adult produces gametes via mitosis Implications for genetics?? Haploid Life Cycle Alternation of Generations Plants and some algae: Sporophyte: Multicellular diploid stage – Meiosis in the sporophyte produces haploid spores – Spores give rise to multicellular adults without fusing with another cell Gametophyte: Multicellular haploid stage – The haploid gametophyte makes gametes via mitosis – Fertilization results in a diploid zygote – The zygote develops into the next sporophyte generation Alternation of generations Moss and Alternation of Generations Implications for genetics?? Some genes affect the sporophyte and others the gametophyte Height of gametophyte and color of sporophyte for example Mendel and his peas – Why did it work out? Topography of the chromosome set: Chromosome number: Centromere Position ….and chromosome size Banding patterns: Heterochromatin patterns Utilizes DNA stain (like Feulgen) Euchromatin: – Poorly staining – Less densely packed – Implications? Heterochromatin: – – – – – – Stains well Densely packed Can be constitutive or facultative Implications? Heterochromatin typically found at centromeres and telomeres The whole drosophila Y chomosome is heterochromatic G banding patterns: Chromosomes partially digested with proteolytic enzymes Stained with Giemsa reagent Produces light and dark banding regions G Banding G-light bands: Tend to be GC rich Not densely packed Replicate early mRNA label binds to these regions most G-dark bands: Tend to be AT rich Densely packed mRNA does not bind well to these areas Implications?? Three dimensional structure of a chromosome: The molecule: Each chromosome is ONE molecule of DNA E.coli contains about 1.3mm of DNA per cell H. sapiens contains about 2m of DNA per cell Implications for cells and cell division ?? The Chromosome Histone Proteins and packaging: Chromatin: The length of a chromosome is much less than that of a DNA molecule Chromosomes made up of chromatin Chromatin is DNA and protein Condensation of a chromosome Histones: Histones are the primary proteins The structure of histones is conserved across eukaryotes Histones form an octamer The solenoid: DNA wraps two times around each octamer This histone DNA spool is called a nucleosome The nucleosome "beaded necklace" assumes a coiled form called a solenoid The solenoid is stabilized by another histone protein (H1) Histones High order coiling: Solenoids are ~30nm across However, chromosomes are ~700nm across! Must have more structure Scaffold protein: Non histone The scaffold forms a spiral The nucleosomes loop on the scaffold Loops attach at SARs (scaffold attachment points) The DNA region where SARs occur are non-coding Scaffold SARs Sequence organization: How are genes arranged? What proportion of the DNA is genes? What is the nature of the non-coding regions? Organelle Genomes: Portion of the genome outside the nucleus Chloroplast DNA (cpDNA): Chloroplasts came from endosymbiotic cyanobacteria cpDNA is a circular double helix Haploid Each organelle has 20-80 copies of cpDNA Contains genes related to photosynthesis and those required for gene expression Has few noncoding sequences compared to nDNA Highly conserved (even the order is conserved) Uniparental inheritance Angiosperms maternal Some gymnosperms paternal Biparental inheritance has been found in some angiosperms Recombination is rare in the extreme Inherited as one allele Mitochondrial DNA (mtDNA) Came from purple bacteria endosymbiont mtDNA is a circular double helix Haploid Codes for many proteins used in cellular respiration 5-10 copies in each organelle mtDNA in plants, fungi and animals differ Plant mtDNA: Great variation in genome size – 300-2,400 kb in melons Structure varies – Can be circular or linear for example Recombination is frequent Sequence divergence is low however Evolution is slow Fungal mtDNA: Genome size variable – 26.7 - 115kb in one group Coding sequences include introns Modification of the "standard" DNA code – UGA = tryptophan not "stop" – CUN = threonine not leucine – AUA = methionine not isoleucine • In some cases Animal mtDNA: Very Compact – No introns and noncoding regions are limited – In the mouse 94% of the mtDNA genome is coding Genome size and gene order conserved Mutation rate is high – Sequence divergence is high – Evolutionary rate 5-10 times that of nDNA AUA = methionine not isoleucine UGA = tryptophan not "stop" Maternal Inheritance (Normally) No Recombination