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Expansion of tandem gene clusters Gene duplication by unequal crossing-over Divergence of coding and regulatory sequences Evolution of HOX clusters The homeobox is a highly conserved DNA-binding domain HOX4 homeodomain Drosophila Amphioxus Mouse Human Chick Frog Fugu Zebrafish Evolution of HOX clusters in vertebrates - Vertebrates have multiple HOX clusters - Paralogous HOX genes may Mouse Fugu have partly redundant functions - Some genes and clusters become specialized for distinct functions - Different lineages lose some genes and acquire new functions for others Zebrafish Using gene replacement to test HOX gene function Using gene replacement to test HOX gene function Functionally equivalent HOX paralogs HOX gene labial required for tritocerebral commissure Genetic rescue strategy lab-Gal4 lab-; lab-Gal4 / UAS - lab red = HRP; green = Lab Other HOX genes can partially substitute for lab Other HOX genes can partially substitute for lab Distinct roles of Ubx and abd-A in the midgut dpp expression in the fly midgut is activated by Ubx, but repressed by abd-A Activation and repression are mediated by the same binding sites within a dpp enhancer Onychophoran Ubx causes homeotic transformations in Drosophila Onychophoran Ubx can regulate some Drosophila target genes dSRF dpp Onychophoran Ubx cannot fully substitute for Drosophila Ubx Chimeric Ubx proteins Mapping the functional specificity of Onychophoran Ubx Functional specificity of Ubx maps outside the homeodomain Conservation and change in Ubx proteins Mapping the functional specificity of Ubx proteins Functional comparison of Drosophila and Artemia Ubx Evolution of a new repression domain Non-HOX functions of derived HOX genes Maternal gradient of bicoid establishes AnteriorPosterior axis in the Drosophila embryo bicoid gradient formation in Drosophila zen functions in Dorso-Ventral patterning sna sog zen zen is a derived HOX gene HOX-like expression of zen homologue in Cupiennius Evolution of zen and bcd from an ancestral HOX gene Genetic control of segmentation in Drosophila ftz deletion does not affect segmentation in Tribolium Antennae HOX-like expression of the Chelicerate ftz homologue Archegozetes longisetosus Effects of ectopic ftz expression in Drosophila Dm 36% 4% Sg en expression A homeodomain-deficient ftz allele ftz expression in Schistocerca Homeotic antenna to leg transformations Dm-ftz Tc-ftz Tc-ftz Sg-ftz Dll-Gal4 Tribolium ftz retains homeotic potential wt Dm-ftz Tc-ftz Dm-Scr wt Dm-ftz Tc-ftz Dm-Antp Evolution of ftz function Comparison of ftz proteins between Drosophila and Schistocerca Expression and function of ftz-F1 ftz-F1 ftz ftz-F1 mutant en ftz-F1 encodes a nuclear hormone receptor Physical interaction between ftz and ftz-F1 Transcriptional activation by ftz / ftz-F1 complex In yeast In Drosophila embryos Novel functions of divergent HOX genes Loss and gain of protein-protein interaction motifs