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Journal of Virology, December 2004, p. 13613-13626, Vol. 78, No. 24
0022-538X/04/$08.00+0     DOI: 10.1128/JVI.78.24.13613-13626.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.

Mucosal Human Papillomaviruses Encode Four Different E5 Proteins Whose Chemistry and Phylogeny Correlate with Malignant or Benign Growth

Ignacio G. Bravo* and Ángel Alonso

Deutsches Krebsforschungszentrum, Heidelberg, Germany

Received 7 May 2004/ Accepted 28 July 2004

We performed a phylogenetic study of the E2-L2 region of human mucosal papillomaviruses (PVs) and of the proteins therein encoded. Hitherto, proteins codified in this region were known as E5 proteins. We show that many of these proteins could be spurious translations, according to phylogenetic and chemical coherence criteria between similar protein sequences. We show that there are four separate families of E5 proteins, with different characteristics of phylogeny, chemistry, and rate of evolution. For the sake of clarity, we propose a change in the present nomenclature. E5{alpha} is present in groups A5, A6, A7, A9, and A11, PVs highly associated with malignant carcinomas of the cervix and penis. E5ß is present in groups A2, A3, A4, and A12, i.e., viruses associated with certain warts. E5{gamma} is present in group A10, and E5{delta} is encoded in groups A1, A8, and A10, which are associated with benign transformations. The phylogenetic relationships between mucosal human PVs are the same when considering the oncoproteins E6 and E7 and the E5 proteins and differ from the phylogeny estimated for the structural proteins L1 and L2. Besides, the protein divergence rate is higher in early proteins than in late proteins, increasing in the order L1 < L2 < E6 {approx} E7 < E5. Moreover, the same proteins have diverged more rapidly in viruses associated with malignant transformations than in viruses associated with benign transformations. The E5 proteins display, therefore, evolutionary characteristics similar to those of the E6 and E7 oncoproteins. This could reflect a differential involvement of the E5 types in the transformation processes.

* Corresponding author. Mailing address: Deutsches Krebsforschungszentrum, Im Neuenheimer Feld-242, 69120 Heidelberg, Germany. Phone: 49 6221 424943. Fax: 49 6221 424932. E-mail: i.bravo{at}dkfz.de.

Journal of Virology, December 2004, p. 13613-13626, Vol. 78, No. 24
0022-538X/04/$08.00+0     DOI: 10.1128/JVI.78.24.13613-13626.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.



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