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Genome evolution. There are both proximate and ultimate explanations in molecular biology Mutation continually generates variation in genome content and structure Raw material for natural selection Potential for non-adaptive evolution Function is too blunt a concept for genome evolution.
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Genome evolution • There are both proximate and ultimate explanations in molecular biology • Mutation continually generates variation in genome content and structure • Raw material for natural selection • Potential for non-adaptive evolution • Function is too blunt a concept for genome evolution
The genome as phenotype • Evolutionary biologists study both pattern and process, or mechanism • What unexpected patterns do we see • Within genomes? • By comparisons among them? • What can we infer about proximal and ultimate mechanisms by • Catching evolution red-handed? • Testing genome evolutionary models?
Three open questions • How rapidly do the expression profiles of duplicated genes diverge? • What is the extent of polymorphism in gene order within species? • What is responsible for physical clusters of co-expressed genes?
Three open questions • How rapidly do the expression profiles of duplicated genes diverge? • What is the extent of polymorphism in gene order within species? • What is responsible for physical clusters of co-expressed genes?
Arabidopsis MPSS • With Blake Meyers and Barry Kesner • Massively parallel signature sequencing • Bead based expression monitoring • Estimates for nearly all genes • Estimates are good even at low expression • Sample eight tissues from Arabidopsis thaliana • Couple with genome-wide phylogenetic analysis of duplicated genes
Measuring expression distance Tissue 2 Tissue 1 Tissue 3
Measuring duplication age Rice Athal1 Athal2 T1 T2
Divergence between duplicates Expression distance Age of duplication
Does the pattern differ for • Tandem duplicates? • Transposed duplicates? • Polyploidy remnants?
Three open questions • How rapidly do the expression profiles of duplicated genes diverge? • What is the extent of polymorphism in gene order within species? • What is responsible for physical clusters of co-expressed genes?
Gene order polymorphism • With Jason Lieb and Jennifer Kriss • Using high-throughput methods to compare gene order in two yeast strains • One strain known to lack 7 ORFs present in the reference genome • The key ingredients • Comparative genomic hybridization to a whole-genome chip • Whole-genome genotyping in multiple haploid recombinants from a cross between the two strains
Detecting gene transposition • Genes that are in different chromosomal positions in the two strains will appear as deletions and amplifications in the recombinant progeny
Three open questions • How rapidly do the expression profiles of duplicated genes diverge? • What is the extent of polymorphism in gene order within species? • What is responsible for physical clusters of co-expressed genes?
Natural selection and clusters of co-expressed genes • With Jianhua Hu • Neighboring genes with similar expression profiles are more common than expected in C. elegans • Two classes of explanation • Adaptive: more efficient, less error-prone • Maladaptive: due to recent transpositions
Selection and recombination • In regions of low recombination • deleterious mutations can hitch-hike to high frequency along with favorable ones • favorable mutations are kept at low frequency by linkage to deleterious ones • The effectiveness of natural selection is directly related to recombination rate • Are clusters found in regions of high recombination (adaptive) or low (maladaptive)?
Measuring the co-expression of neighboring genes • Measure the distance in expression space from central gene to each neighbor • If neighboring genes are co-expressed, the average of these distances will be smaller than for non-neighboring genes d1 d2 d3 d4 X
Expression distance and recombination rate Spearman = -0.07 p = 0.0005
Opportunities abound! • The field of genome evolution is a vast and open playing field • Gads of genome data • Some seriously cool experimental techniques • A golden age in bioinformatics and molecular evolution