title: The serine/threonine kinase 33 is present and expressed in palaeognath birds but has become a unitary pseudogene in neognaths about 100 million years ago.
abstract: BACKGROUND
  Serine/threonine kinase 33 (STK33) has been shown to be conserved across all major vertebrate classes including reptiles, mammals, amphibians and fish, suggesting its importance within vertebrates. It has been shown to phosphorylate vimentin and might play a role in spermatogenesis and organ ontogenesis. In this study we analyzed the genomic locus and expression of stk33 in the class Aves, using a combination of large scale next generation sequencing data analysis and traditional PCR.
  
  
  RESULTS
  Within the subclass Palaeognathae we analyzed the white-throated tinamou (Tinamus guttatus), the African ostrich (Struthio camelus) and the emu (Dromaius novaehollandiae). For the African ostrich we were able to generate a 62,778 bp long genomic contig and an mRNA sequence that encodes a protein showing highly significant similarity to STK33 proteins from other vertebrates. The emu has been shown to encode and transcribe a functional STK33 as well. For the white-throated tinamou we were able to identify 13 exons by sequence comparison encoding a protein similar to STK33 as well. In contrast, in all 28 neognath birds analyzed, we could not find evidence for the existence of a functional copy of stk33 or its expression. In the genomes of these 28 bird species, we found only remnants of the stk33 locus carrying several large genomic deletions, leading to the loss of multiple exons. The remaining exons have acquired various indels and premature stop codons.
  
  
  CONCLUSIONS
  We were able to elucidate and describe the genomic structure and the transcription of a functional stk33 gene within the subclass Palaeognathae, but we could only find degenerate remnants of stk33 in all neognath birds analyzed. This led us to the conclusion that stk33 became a unitary pseudogene in the evolutionary history of the class Aves at the paleognath-neognath branch point during the late cretaceous period about 100 million years ago. We hypothesize that the pseudogenization of stk33 might have become fixed in neognaths due to either genetic redundancy or a non-orthologous gene displacement and present potential candidate genes for such an incident.
authors.affiliation: Institute for Molecular Genetics, Johannes Gutenberg University Mainz, Johann-Joachim-Becherweg 32, 55128, Mainz, Germany. lautweit@uni-mainz.de.
authors.forename: Tobias
authors.initials: T
authors.lastname: Lautwein
authors.affiliation: Institute for Molecular Genetics, Johannes Gutenberg University Mainz, Johann-Joachim-Becherweg 32, 55128, Mainz, Germany. Steffen.Lerch@unimedizin-mainz.de.|Departement of Neurology, University Medical Center, Johannes Gutenberg-University Mainz, Langenbeckstr.1, 55131, Mainz, Germany. Steffen.Lerch@unimedizin-mainz.de.
authors.forename: Steffen
authors.initials: S
authors.lastname: Lerch
authors.affiliation: Institute for Molecular Genetics, Johannes Gutenberg University Mainz, Johann-Joachim-Becherweg 32, 55128, Mainz, Germany. Daniel.Schaefer@med.uni-duesseldorf.de.|Departement of Pediatric Oncology, Hematology and Clinical Immunology, University Children's Hospital, Medical Faculty, Heinrich Heine University, Moorenstr. 5, 40225, Düsseldorf, Germany. Daniel.Schaefer@med.uni-duesseldorf.de.
authors.forename: Daniel
authors.initials: D
authors.lastname: Schäfer
authors.affiliation: Institute for Molecular Genetics, Johannes Gutenberg University Mainz, Johann-Joachim-Becherweg 32, 55128, Mainz, Germany. eschmidt@uni-mainz.de.
authors.forename: Erwin R
authors.initials: ER
authors.lastname: Schmidt
mesh_terms: D000818:Animals; D001717:Birds / Q000235:genetics*; D019143:Evolution, Molecular; D005786:Gene Expression Regulation; D016678:Genome; D017346:Protein Serine-Threonine Kinases / Q000235:genetics*; D014714:Vertebrates
chemical_list: D017346:Protein Serine-Threonine Kinases