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Viruses 2009, 1, 57-71; doi:10.3390/v1010057 OPEN ACCESS viruses ISSN 1999-4915 /journal/viruses Review More and More Coronaviruses: Human Coronavirus HKU1 Patrick C. Y. Woo 1,2,3,†,*, Susanna K. P. Lau 1,2,3,†,*, Cyril C. Y. Yip 3, Yi Huang 3 and Kwok-Yung Yuen 1,2,3 1 State Key Laboratory of Emerging Infectious Diseases, The University of Hong Kong, Hong Kong, China 2 Research Centre of Infection and Immunology, The University of Hong Kong, Hong Kong, China 3 Department of Microbiology, The University of Hong Kong, Hong Kong, China † These authors contributed equally to the manuscript * Authors to whom correspondence should be addressed; E-mails: pcywoo@hkucc.hku.hk (P.C. Y.W.); skplau@hkucc.hku.hk (S.K.P.L.); Tel. +852 Fax: +852 Received: 7 May 2009; in revised form: 6 June 2009 / Accepted: 11 June 2009 / Published: 11 June 2009 Abstract: After human coronaviruses OC43, 229E and NL63, human coronavirus HKU1 (HCoV-HKU1) is the fourth human coronavirus discovered. HCoV-HKU1 is a group 2a coronavirus that is still not cultivable. The G + C contents of HCoV-HKU1 genomes are 32%, the lowest among all known coronaviruses with complete genome sequences available. Among all coronaviruses, HCoV-HKU1 shows the most extreme codon usage bias, attributed most importantly to severe cytosine deamination. All HCoV-HKU1 genomes contain unique tandem copies of a 30-base acidic tandem repeat of unknown function at the N-terminus of nsp3 inside th