Abstract
Genome size varies widely across organisms, with no apparent tie to organismal complexity. While genome size is inherited, there is no established evolutionary model for this trait. Hypotheses have been postulated for the observed variation in genome sizes across species, most notably the effective population size hypothesis, the mutational equilibrium hypothesis, and the adaptive hypothesis. While much data has been collected on genome size, the above hypotheses have largely ignored impacts from phylogenetic relationships. In order to test these competing hypotheses, genome sizes of 87 Sophophora species were analyzed in a comparative phylogenetic approach using Pagel’s parameters of evolution, Blomberg’s K, Abouheif’s Cmean and Moran’s I. In addition to testing the mode and rate of genome size evolution in Sophophora species, the effect of number of taxa on detection of phylogenetic signal was analyzed for each of these comparative phylogenetic methods. Sophophora genome size was found to be dependent on the phylogeny, indicating that evolutionary time was important for predicting the variation among species. Genome size was found to evolve gradually on branches of the tree, with a rapid burst of change early in the phylogeny. These results suggest that Sophophora genome size has experienced gradual changes, which support the largely theoretical mutational equilibrium hypothesis. While some methods (Abouheif’s Cmean and Moran’s I) were found to be affected by increasing taxa numbers, more commonly used methods (λ and Blomberg’s K) were found to have increasing reliability with increasing taxa number, with significantly more support with fifteen or more taxa. Our results suggest that these comparative phylogenetic methods, with adequate taxon sampling, can be a powerful way to uncover the enigma that is genome size variation through incorporation of phylogenetic relationships.
Highlights
When considering trait evolution, sequence of the genotype is traditionally inspected for evidence of selection or drift, through methods such as DN/DS ratios
The relationships in this phylogeny are supported by other large Drosophila phylogenies in the literature [12, 36,37,38]. These results suggest that the constructed phylogeny is representative of the true relationships found in Sophophora, and should have reliable branch lengths
We look at measures of phylogenetic signal (Pagel’s λ, Blomberg’s K, Abouheif’s Cmean, and Moran’s I) and at measures of mode and tempo of evolution (Pagel’s δ and κ) in order to test three hypotheses of genome size evolution: low effective population size hypothesis, mutational equilibrium hypothesis, and an adaptive hypothesis
Summary
Sequence of the genotype is traditionally inspected for evidence of selection or drift, through methods such as DN/DS ratios. These tests, are not applied to genome size. The wide variation in genome size is not generally attributed to coding DNA sequences, but rather to repetitive and nongenic DNA [1, 3, 4] For these reasons, we are analyzing the variation in DNA content in a comparative phylogenetic context with the goal of establishing an evolutionary model for genome size from proposed hypotheses
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