Discovery and evolution of novel hemerythrin genes in annelid worms.
Despite extensive study on hemoglobins and hemocyanins, little is known about hemerythrin (Hr) evolutionary history. Four subgroups of Hrs have been documented, including: circulating Hr (cHr), myohemerythrin (myoHr), ovohemerythrin (ovoHr), and neurohemerythrin (nHr). Annelids have the greatest diversity of oxygen carrying proteins among animals and are the only phylum in which all Hr subgroups have been documented. To examine Hr diversity in annelids and to further understand evolution of Hrs, we employed approaches to survey annelid transcriptomes in silico.



Sequences of 214 putative Hr genes were identified from 44 annelid species in 40 different families and Bayesian inference revealed two major clades with strong statistical support. Notably, the topology of the Hr gene tree did not mirror the phylogeny of Annelida as presently understood, and we found evidence of extensive Hr gene duplication and loss in annelids. Gene tree topology supported monophyly of cHrs and a myoHr clade that included nHrs sequences, indicating these designations are functional rather than evolutionary.



The presence of several cHrs in early branching taxa suggests that a variety of Hrs were present in the common ancestor of extant annelids. Although our analysis was limited to expressed-coding regions, our findings demonstrate a greater diversity of Hrs among annelids than previously reported.
Departamento de Genética, Laboratório de Biologia Evolutiva Teórica e Aplicada, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, Brazil.|Department of Biological Sciences, Molette Biology Laboratory for Environmental and Climate Change Studies, Auburn University, Auburn, AL, 36849, USA.
Elisa M
EM
Costa-Paiva
Department of Biological Sciences, Molette Biology Laboratory for Environmental and Climate Change Studies, Auburn University, Auburn, AL, 36849, USA.|Warm Springs Fish Technology Center, U.S. Fish and Wildlife Service, 5308 Spring ST, Warm Springs, GA, 31830, USA.
Nathan V
NV
Whelan
Department of Biological Sciences, Molette Biology Laboratory for Environmental and Climate Change Studies, Auburn University, Auburn, AL, 36849, USA.
Damien S
DS
Waits
Department of Biological Sciences, Molette Biology Laboratory for Environmental and Climate Change Studies, Auburn University, Auburn, AL, 36849, USA.
Scott R
SR
Santos
Departamento de Genética, Laboratório de Biologia Evolutiva Teórica e Aplicada, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, Brazil.
Carlos G
CG
Schrago
Department of Biological Sciences, Molette Biology Laboratory for Environmental and Climate Change Studies, Auburn University, Auburn, AL, 36849, USA. ken@auburn.edu.
Kenneth M
KM
Halanych
D000818:Animals; D000845:Annelida / Q000145:classification / Q000235:genetics*; D001483:Base Sequence; D001499:Bayes Theorem; D019143:Evolution, Molecular; D006422:Hemerythrin / Q000737:chemistry / Q000235:genetics*; D010802:Phylogeny; D016415:Sequence Alignment
D006422:Hemerythrin
Annelida; Blood pigments; Respiratory proteins; Transcriptome