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MBE Advance Access published online on February 2, 2005

Molecular Biology and Evolution, doi:10.1093/molbev/msi099
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Molecular Biology and Evolution © Society for Molecular Biology and Evolution 2005; all rights reserved.
Accepted January 25, 2005

Research Article

Consideration of RNA Secondary Structure Significantly Improves Likelihood-Based Estimates of Phylogeny: Examples from the Bilateria

Maximilian J Telford 1*, Michael J Wise 2, and Vivek Gowri-Shankar 3

1 Department of Biology, University College London, Darwin Building, Gower Street, London, WC1E 6BT UK. Tel: +44 (0)2076792554; Fax: +44 (0)2076797096
2 School of Biomedical and Chemical Sciences, University of Western Australia
3 Department of Computer Science, University of Manchester

* To whom correspondence should be addressed.
Maximilian J Telford, E-mail: m.telford{at}ucl.ac.uk


   Abstract

Sequences from ribosomal RNA genes have made a huge contribution to our current understanding of metazoan phylogeny and indeed the phylogeny of all of life. That said, some parts of this rRNA based phylogeny remain unresolved. One approach to increase the resolution of these trees would be to use more appropriate models of sequence evolution in phylogenetic analysis.

RNAs transcribed from ribosomal RNA genes have a complex secondary structure mediated by base pairing between sometimes distant regions of the rRNA molecule. The pairing between the stem nucleotides has important consequences for their evolution which differs from that of unpaired loop nucleotides. These differences in evolution should ideally be accounted for when using rRNA sequences for phylogeny estimation.

We use a novel permutation approach to demonstrate the significant superiority of models of sequence evolution that allow stem and loop regions to evolve according to separate models and, in common with previous studies, we show that 16 state models that take base pairing of stems into account are significantly better than simpler, 4-state single-nucleotide models. One of these 16 state models has been applied to the phylogeny of the Bilateria using small subunit ribosomal RNA sequences. Our optimal tree largely echoes previous results based on SSU in particular supporting the tripartite Bilaterian tree of deuterostomes, lophotrochozoans and ecdysozoans. There are also a number of differences, however, perhaps most important of which is the observation of a clade consisting of the gastrotrichs plus platyheminthes that is basal to all other lophotrochozoan taxa. Use of 16 state models also appears to reduce the Bayesian support given to certain biologically improbable groups found using standard 4 state models.

Keywords: Ribosomal RNA; phylogeny; Bilateria; Secondary structure Maximum likelihood Bayesian analysis.
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