DuplicationsBar eye: caused by duplicationDuplications: source of evolutionary novelty?Structural ChangesInversionsPowerPoint PresentationSlide 7Inversions Suppress RecombinationInversions and evolutionSlide 10Slide 11Slide 12Robertsonian Translocation: Familial Down SyndromeSlide 14Position EffectChromosome Evolution:Evolutionary change in chromosome numberSlide 18SyntenySlide 20Polyploidy_________Triploid TetraploidMany plants are polyploidsSlide 25Some autopolyploidsAllopolyploidy Resutls from hybridization between different speciesAllopolyploidsSpeciation by PolyploidizationSlide 30Artificially-Produced New Species!RhaphanobrassicaPolyploidy in AnimalsWhy less common in animals?DuplicationsBar eye: caused by duplicationDuplications: source of evolutionary novelty?•______________•______________•______________•______________Duplication is a source of new genes over evolutionary time: e.g., gene families like globins and MHC genesStructural Changes•Deletions (deficiencies)•Duplications•Inversions•TranslocationsInversionsInversions Suppress RecombinationOrganisms carrying an inversion tend to undergo little crossing over in the inversion region in both inverted and non-inverted chromosomes.If there is crossing over, half the chromatids involved in crossing over will produce non-viable gametes.Can be a cause of ________Inversions and evolutionIf loci inside an inversion affect a single trait (or suite of related traits), this means they'll be inherited together and allele combinations won’t be broken up by recombination.A suite of tightly linked loci that affect a single trait is collectively known as a SUPERGENE.•mimicry coloration in some species of butterflies•snail shell color pattern (some species)Structural Changes•Deletions (deficiencies)•Duplications•Inversions•TranslocationsRobertsonian Translocation: Familial Down SyndromePosition Effect•In some translocation mutations, the new position of the gene can affect the rate of its transcription and translation.•For example, if a highly-transcribed gene is translocated to a region close to tightly coiled, inactive heterochromatin, it can sometimes be partially engulfed by that heterochromatin. This will result in a failure of the gene to be expressed in the cells where the heterochromatin coils over the translocated gene.•This can be seen in "position effect variegation" in our old pal Drosophila:Chromosome Evolution: 2n nHuman 46 23Chimpanzee4824Horse 64 32Cat 38 19Chicken7839Geometrid moth224112Drosophila mel84Ant (Myrecia pilosula)21Tomato 24 12Horsetail216108Pink bread mold 14 7Evolutionary change in chromosome numberCentromere fusion: two acrocentric chromosomes become fused at or near their terminal centromeres, creating a single, large chromosome. Result: Often evidence of this in closely related species. Compare the NF (fundamental number): the number of chromosome arms visible at metaphase. Or compare sequence data.Centromere fission. In this case, the chromosome splits at the centromere, creating two chromosomes from one. Result:QuickTime™ and aTIFF (LZW) decompressorare needed to see this picture.SyntenyPolyploidyAutopolyploidyAllopolyploidy_________Triploid TetraploidP: 2n 2nG: n 2nF1: P: 2n 2nG:F1:Many plants are polyploids•> ______ of flowering plants are of polyploid origin.•In many genera, different species will have different ploidy levels (multiples of a base number) representing a series of polyploids. In the genus Chrysanthemum, different species have chromosome numbers of 2n = 18, 36, 54, 72, 90, and 198 - all multiples of a base chromosome number of 9.Some autopolyploids•Winesap apples•Commercial bananas•Seedless watermelons•Cultivated tiger lilyAllopolyploidyResutls from hybridization between different speciesBread wheat, oats cotton, tobacco, strawberries, coffeeXenopus frogsAllopolyploidsBread wheat______________________________________________Xenopus frogsSpeciation by PolyploidizationSome species are sufficiently closely related that their genes, when combined in a hybrid individual, provide the necessary information for a viable organism--but not for that organism to undergo normal meiosis.Homo sapiens 2n - 46Pan troglodytes 2n - 44Hybrid between the two would be 2n = 45.(BUT THIS HAS NOT HAPPENED, DESPITE WHAT THE WEEKLY WORLD NEWS WILL TELL YOU.)Speciation by PolyploidizationSome species are sufficiently closely related that their genes, when combined in a hybrid individual, provide the necessary information for a viable organism--but not for that organism to undergo normal meiosis.Homo sapiens 2n - 46Pan troglodytes 2n - 44Hybrid between the two would be 2n = 45.(BUT THIS HAS NOT HAPPENED, DESPITE WHAT THE WEEKLY WORLD NEWS WILL TELL YOU.)Artificially-Produced New Species!•Brassica oleracea (_______) x Raphanus sativa (______)•Both species 2n = 18; n = 9, chromosomes not homologous.•_______ X ______ --> hybrid has 18 chromosomes, but meiosis is abnormal.•Nondisjunction in a meristematic cell --> 36 chromosomes.•The cell is said to be allopolyploid (allotetraploid).•Normal meiosis!•These plants are self-fertile, so can produce offspring, even if there is only one such individual. •New “genus/species" was named Raphanobrassica.•Unfortunately, the hybrid combines the root of a cabbage with the flower head of a radish, rather than the reverse!RhaphanobrassicaPolyploidy in AnimalsAnimals can also produce allopolyploids (e.g. horse x donkey --> mule; lion x tiger--> liger or tigon). In mammals, these crosses tend to be sterile, but many fish and reptile species are thought to have arisen as polyploids (e,g, ________________________).Why less common in animals?Many animals have chromosomally determined sex, and polyploidy interferes with this.Most animals have several isolating mechanisms (geographic, temporal, behavioral etc.) that tend to prevent natural interbreeding between species.Many plants have meristematic tissue throughout their lives and are self-fertile: In plants, about 5 out of every 1000 gametes produced is