Journal of Molecular Evolution

, Volume 72, Issue 3, pp 253–264 | Cite as

Polymorphisms in the Extracellular Region of Dopamine Receptor D4 Within and Among Avian Orders

  • Hideaki Abe
  • Shin’ichi Ito
  • Miho Inoue-MurayamaEmail author


Polymorphisms in the dopamine receptor D4 gene (DRD4) have been widely investigated to assess their correlation with variations in animal behavior. We precisely examined polymorphisms in the extracellular region of DRD4 in 75 avian species belonging to 16 orders and detected high degrees of polymorphism at inter- and intraordinal levels. The existence of a variable number of proline repeats (2 to 12 times) in the extracellular region was a common feature in all Neognathae, and a strong codon bias at synonymous sites was found among Passeriformes, Galliformes, and other non-passerine Neoaves. Furthermore, significantly higher values of the pairwise disparity index were detected in Passeriformes, suggesting either a substantial difference in the evolutionary processes or a higher level of mutation rate in the passerine clade. The differences in both codon bias and other genetic parameters among avian taxa would be explained by different levels of selective pressure on the extracellular region of DRD4. Our study suggested that different conformations determined in a sequence-dependent manner at the extracellular region could be one of the key factors affecting the efficiency and accuracy of DRD4 expression. Our findings further imply a possibility that behavioral diversity, which would be important during the processes of adaptive radiation, may be enhanced by the selection acting on indels or single-nucleotide substitutions in the extracellular region of DRD4.


Dopamine receptor D4 (DRD4Polymorphism Codon bias Extracellular region Avian Neoaves Proline repeats Selective pressure 



For blood and tissue samples, the authors thank the following institutions: Kobe Kacho-en (Hyogo prefecture, Japan), Fuji Kacho-en (Shizuoka prefecture, Japan), Kakegawa Kacho-en (Shizuoka prefecture, Japan), Matsue Vogel Park (Shimane prefecture, Japan), Fukui Nature Conservation Center (Fukui prefecture, Japan), Research Center for Wildlife Management, Gifu University (Gifu prefecture, Japan), Tennoji Zoo (Osaka prefecture, Japan), and Yokohama Zoological Gardens (Kanagawa prefecture, Japan). We are grateful to the following persons for great support in the laboratory work (in alphabetical order): Kazumi Kato, Emi Kawaguchi, and Boniface B. Kayang at Kyoto University. Thanks also to Shuichi Matsumura at Gifu University for useful comments on earlier versions of this paper. This study was supported financially by the Ministry of Education, Culture, Sports, Science and Technology (MEXT) with a Grant-in-aid for Science Research (#21310150 to MI-M) and Global Center of Excellence Program “Formation of a Strategic Base for Biodiversity and Evolutionary Research: from Genome to Ecosystem.”


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Copyright information

© Springer Science+Business Media, LLC 2011

Authors and Affiliations

  • Hideaki Abe
    • 1
  • Shin’ichi Ito
    • 2
  • Miho Inoue-Murayama
    • 1
    Email author
  1. 1.Wildlife Research Center of Kyoto UniversityKyotoJapan
  2. 2.Faculty of Applied Biological SciencesGifu UniversityGifuJapan

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