X chromosome evolution in Drosophila
Although the X chromosome is usually similar to the autosomes in size, gene density and cytogenetic appearance, theoretical models predict that its hemizygosity in males may cause unusual patterns of evolution. The sequencing of several genomes has indeed revealed differences between the X chromosome and the autosomes in the rates of gene divergence, patterns of gene expression and rates of gene movement between chromosomes. In this thesis, I have attempted to investigate some of these patterns and their possible causes. The first two chapters consist of theoretical and empirical work intended to analyse the rates of evolution of coding sequences of X-linked and autosomal loci, with particular emphasis on faster-X evolution, the theory that more effective selection on the X can lead to higher rates of adaptive evolution on this chromosome. By analyzing X-linked and autosomal coding sequence in several species of Drosophila, we found some evidence for more effective selection on the X, particularly evident in the higher levels of codon usage bias detected at X-linked loci. We argue that this could be due to higher levels of recombination on the X chromosome increasing its effective population size (NeX) relative to the autosomal effective population size (NeA). To further investigate this hypothesis, we have modeled the effect of increased NeX/NeA on rates of evolution and confirmed that this can contribute to faster-X evolution. The last two chapters deal with the evolution of sex-biased genes and the possible causes for their differential accumulation on the X. We used EST data to create expression profiles for D. melanogaster male-, female- and unbiased genes. Our results suggest that the expression levels of sex-biased genes are incompatible with the accepted iii model of sex-biased gene evolution. We also show that the deficit of testis-expressed genes that is observed in Drosophila seems to be stronger for highly expressed genes. In fact, for very lowly expressed genes, we observe a small excess of testis-expressed genes on the X. We attempt to discuss this pattern in view of what is currently known about the evolution of sex-biased gene expression.