title: A genomic timescale for the origin of eukaryotes.
abstract: BACKGROUND
  Genomic sequence analyses have shown that horizontal gene transfer occurred during the origin of eukaryotes as a consequence of symbiosis. However, details of the timing and number of symbiotic events are unclear. A timescale for the early evolution of eukaryotes would help to better understand the relationship between these biological events and changes in Earth's environment, such as the rise in oxygen. We used refined methods of sequence alignment, site selection, and time estimation to address these questions with protein sequences from complete genomes of prokaryotes and eukaryotes.
  
  
  RESULTS
  Eukaryotes were found to evolve faster than prokaryotes, with those eukaryotes derived from eubacteria evolving faster than those derived from archaebacteria. We found an early time of divergence (approximately 4 billion years ago, Ga) for archaebacteria and the archaebacterial genes in eukaryotes. Our analyses support at least two horizontal gene transfer events in the origin of eukaryotes, at 2.7 Ga and 1.8 Ga. Time estimates for the origin of cyanobacteria (2.6 Ga) and the divergence of an early-branching eukaryote that lacks mitochondria (Giardia) (2.2 Ga) fall between those two events.
  
  
  CONCLUSIONS
  We find support for two symbiotic events in the origin of eukaryotes: one premitochondrial and a later mitochondrial event. The appearance of cyanobacteria immediately prior to the earliest undisputed evidence for the presence of oxygen (2.4-2.2 Ga) suggests that the innovation of oxygenic photosynthesis had a relatively rapid impact on the environment as it set the stage for further evolution of the eukaryotic cell.
authors.affiliation: Astrobiology Research Center and Department of Biology, 208 Mueller Laboratory, The Pennsylvania State University, University Park, Pennsylvania 16802, USA. sbh1@psu.edu
authors.forename: S B
authors.initials: SB
authors.lastname: Hedges
authors.forename: H
authors.initials: H
authors.lastname: Chen
authors.forename: S
authors.initials: S
authors.lastname: Kumar
authors.forename: D Y
authors.initials: DY
authors.lastname: Wang
authors.forename: A S
authors.initials: AS
authors.lastname: Thompson
authors.forename: H
authors.initials: H
authors.lastname: Watanabe
mesh_terms: D000818:Animals; D001105:Archaea / Q000235:genetics; D005051:Eubacterium / Q000235:genetics; D005057:Eukaryotic Cells; D019143:Evolution, Molecular; D014644:Genetic Variation / Q000235:genetics; D016678:Genome; D020745:Genome, Archaeal; D016680:Genome, Bacterial; D018503:Genome, Protozoan; D005872:Giardia / Q000235:genetics; D008957:Models, Genetic; D010802:Phylogeny