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JUNE 2012 • VOLUME 86 • NO. 12

TABLE OF CONTENTS

SPOTLIGHT Articles of Significant Interest Selected from This Issue by the 6385 Editors STRUCTURE AND ASSEMBLY Epitope Insertion at the N-Terminal Molecular Switch of the Daniel Luque, José M. González, Josué 6470–6480 , Leads to Gómez-Blanco, Roberto Marabini 3؍Rabbit Hemorrhagic Disease T Javier Chichón, Ignacio Mena, Iván 4؍Larger T Angulo, José L. Carrascosa, Nuria Verdaguer, Benes L. Trus, Juan Bárcena, and José R. Castón Structure of Virus Polymerase in Complex with Ralph T. Mosley, Thomas E. Edwards, 6503–6511 Primer-Template RNA Eisuke Murakami, Angela M. Lam, Rena L. Grice, Jinfa Du, Michael J. Sofia, Philip A. Furman, and Michael J. Otto Analysis of Viral and Cellular Factors Influencing Herpesvirus- Katharina S. Grimm, Barbara G. Klupp, 6512–6521 Induced Nuclear Envelope Breakdown Harald Granzow, Frederik M. Müller, Walter Fuchs, and Thomas C. Mettenleiter Structural Basis for RNA Binding and Homo- Andy Ka-Leung Ng, Mandy Ka-Han 6758–6767 Formation by Influenza B Virus Nucleoprotein Lam, Hongmin Zhang, Jinhuan Liu, Shannon Wing-Ngor Au, Paul Kay- Sheung Chan, Jiahuai Wang, and Pang- Chui Shaw Capsid Structure and Its Stability at the Late Stages of Helen E. White, Michael B. Sherman, 6768–6777 SPP1 Assembly Sandrine Brasilès, Eric Jacquet, Philippa Seavers, Paulo Tavares, and Elena V. Orlova Structural Insight into the Unique Properties of Adeno- Michael A. DiMattia, Hyun-Joo Nam, 6947–6958 Associated Virus Serotype 9 Kim Van Vliet, Matthew Mitchell, Antonette Bennett, Brittney L. Gurda, Robert McKenna, Norman H. Olson, Robert S. Sinkovits, Mark Potter, Barry J. Byrne, George Aslanidi, Sergei Zolotukhin, Nicholas Muzyczka, Timothy S. Baker, and Mavis Agbandje-McKenna REPLICATION AND REGULATION OF VIRAL GENE EXPRESSION Extensive Mutagenesis of the Conserved Box E Motif in Duck Yong-Xiang Wang, Cheng Luo, Dan 6394–6407 P Protein Reveals Multiple Functions in Zhao, Jürgen Beck, and Michael Nassal Replication and a Common Structure with the Primer Grip in HIV-1 Determinants in the Maturation of Rubella Virus P200 Replicase Jason D. Matthews, Wen-Pin Tzeng, 6457–6469 Polyprotein Precursor and Teryl K. Frey A Conserved N-Terminal Domain Mediates Required DNA David J. Taggart, Jonathan K. Mitchell, 6575–6585 Replication Activities and Phosphorylation of the and Paul D. Friesen Transcriptional Activator IE1 of Autographa californica Multicapsid Nucleopolyhedrovirus

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Replication-Uncoupled Histone Deposition during Adenovirus Tetsuro Komatsu and Kyosuke Nagata 6701–6711 DNA Replication Identification of a Divalent Metal Cation Binding Site in Herpes Kevin F. Bryant, Zhipeng Yan, David H. 6825–6834 Simplex Virus 1 (HSV-1) ICP8 Required for HSV Replication Dreyfus, and David M. Knipe The N Terminus and C Terminus of Virus 1 Lauren M. Wagner, Jonathan T. Lester, 6862–6874 ICP4 Cooperate To Activate Viral Gene Expression Frances L. Sivrich, and Neal A. DeLuca

Deletion of UL21 Causes a Delay in the Early Stages of the Ekaette F. Mbong, Lucille Woodley, 7003–7007 1 Replication Cycle Elizabeth Frost, Joel D. Baines, and Carol Duffy GENETIC DIVERSITY AND EVOLUTION Characterization of a Novel Bipartite Double-Stranded RNA Mingde Wu, Fengyin Jin, Jing Zhang, 6605–6619 Mycovirus Conferring Hypovirulence in the Phytopathogenic Long Yang, Daohong Jiang, and Fungus Botrytis porri Guoqing Li Genomic Sequences of Australian Bluetongue Virus Prototype David B. Boyle, Dieter M. Bulach, 6724–6731 Serotypes Reveal Global Relationships and Possible Routes of Rachel Amos-Ritchie, Mathew M. Entry into Australia Adams, Peter J. Walker, and Richard Weir Genetic Analysis of Cytomegalovirus in Malignant Gliomas Bornali Bhattacharjee, Nicholas 6815–6824 Renzette, and Timothy F. Kowalik Human Papillomavirus Type 16 Genetic Variants: Phylogeny Iris Cornet, Tarik Gheit, Silvia 6855–6861 and Classification Based on E6 and LCR Franceschi, Jerome Vignat, Robert D. Burk, Bakary S. Sylla, Massimo Tommasino, Gary M. Clifford, and the IARC HPV Variant Study Group VIRUS- INTERACTIONS Residues within the C-Terminal Arm of the Herpes Simplex Sarah A. Connolly and Richard 6386–6393 Virus 1 B Ectodomain Contribute to Its Refolding Longnecker during the Fusion Step of Virus Entry Vicriviroc Resistance Decay and Relative Replicative Fitness in Athe M. N. Tsibris, Zixin Hu, Roger 6416–6426 HIV-1 Clinical Isolates under Sequential Drug Selection Paredes, Kay E. Leopold, Opass Pressures Putcharoen, Allison L. Schure, Natalie Mazur, Eoin Coakley, Zhaohui Su, Roy M. Gulick, and Daniel R. Kuritzkes

Oct4؉ Stem/Progenitor Swine Lung Epithelial Cells Are Targets Mahesh Khatri, Sagar M. Goyal, and 6427–6433 for Influenza Virus Replication Yehia M. Saif A3-O-Sulfated Heparan Sulfate Binding Peptide Preferentially Mohamed M. Ali, Ghadah A. Karasneh, 6434–6443 Targets Herpes Simplex Virus 2-Infected Cells Min Jung Jarding, Vaibhav Tiwari, and Deepak Shukla Glycoprotein B of Herpes Simplex Virus 2 Has More than One Martin I. Muggeridge 6444–6456 Intracellular Conformation and Is Altered by Low pH Role for TBC1D20 and Rab1 in Replication via Inbar Nevo-Yassaf, Yakey Yaffe, Meital 6491–6502 Interaction with Droplet-Bound Nonstructural Protein 5A Asher, Orly Ravid, Sharon Eizenberg, Yoav I. Henis, Yaakov Nahmias, Koret Hirschberg, and Ella H. Sklan TP-RT Domain Interactions of Duck Hepatitis B Virus Reverse Rajeev K. Boregowda, Christina Adams, 6522–6536 Transcriptase in cis and in trans during Protein-Primed and Jianming Hu Initiation of DNA Synthesis In Vitro Simultaneous Treatment of Human Bronchial Epithelial Cells Miyuki Kawase, Kazuya Shirato, Lia 6537–6545 with Serine and Cysteine Protease Inhibitors Prevents Severe van der Hoek, Fumihiro Taguchi, and Acute Respiratory Syndrome Coronavirus Entry Shutoku Matsuyama

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Cysteines in the Stalk of the Nipah Virus G Glycoprotein Are Dianna Maar, Brooke Harmon, David 6632–6642 Located in a Distinct Subdomain Critical for Fusion Activation Chu, Belinda Schulz, Hector C. Aguilar, Benhur Lee, and Oscar A. Negrete Integrin ␤1 Mediates Virus Entry through Activation Roza Izmailyan, Jye-Chian Hsao, Che- 6677–6687 of PI3K/Akt Signaling Sheng Chung, Chein-Hung Chen, Paul Wei-Che Hsu, Chung-Lin Liao, and Wen Chang Feline Is Characterized by a Short N-Terminal Region Michele Celestino, Arianna Calistri, 6688–6700 and Is Counteracted by the Feline Immunodeficiency Virus Claudia Del Vecchio, Cristiano Salata, Envelope Glycoprotein Flavia Chiuppesi, Mauro Pistello, Alessandra Borsetti, Giorgio Palù, and Cristina Parolin Murine Cytomegalovirus Targets Factor ATF4 To Zhikang Qian, Baoqin Xuan, Travis J. 6712–6723 Exploit the Unfolded-Protein Response Chapa, Nathaniel Gualberto, and Dong Yu The Ubiquitin-Specific Protease USP7 Modulates the Wiebke Jäger, Susann Santag, 6745–6757 Replication of Kaposi’s Sarcoma-Associated Herpesvirus Latent Magdalena Weidner-Glunde, Eva Episomal DNA Gellermann, Semra Kati, Marcel Pietrek, Abel Viejo-Borbolla, and Thomas F. Schulz Classical Swine Fever Virus p7 Protein Is a Involved Douglas P. Gladue, Lauren G. Holinka, 6778–6791 in Virulence in Swine Eneko Largo, Ignacio Fernandez Sainz, Consuelo Carrillo, Vivian O’Donnell, Ryan Baker-Branstetter, Zhiqiang Lu, Xavier Ambroggio, Guillermo R. Risatti, Jose L. Nieva, and Manuel V. Borca

Temperature-Dependent Survival of Turnip Crinkle Virus- Xiuchun Zhang, Xiaofeng Zhang, 6847–6854 Infected Arabidopsis Plants Relies on an RNA Silencing-Based Jasleen Singh, Dawei Li, and Feng Qu Defense That Requires DCL2, AGO2, and HEN1 The US16 Gene of Human Cytomegalovirus Is Required for Matteo Bronzini, Anna Luganini, 6875–6888 Efficient Viral Infection of Endothelial and Epithelial Cells Valentina Dell’Oste, Marco De Andrea, Santo Landolfo, and Giorgio Gribaudo Proteomic Profiling of EBNA1- Protein Interactions in Natasha Malik-Soni and Lori Frappier 6999–7002 Latent and Lytic Epstein-Barr Virus Infections CELLULAR RESPONSE TO INFECTION Herpes Simplex Virus gH/gL and gB Bind Toll- Valerio Leoni, Tatiana Gianni, Stefano 6555–6562 Like Receptor 2, and Soluble gH/gL Is Sufficient To Activate Salvioli, and Gabriella NF-␬B Campadelli-Fiume Pellino-1 Selectively Regulates Epithelial Cell Responses to Julie A. Bennett, Lynne R. Prince, Lisa 6595–6604 Rhinovirus C. Parker, Clare A. Stokes, Harold G. de Bruin, Maarten van den Berge, Irene H. Heijink, Moira K. Whyte, and Ian Sabroe Cdk1 Inhibition Induces Mutually Inhibitory Apoptosis and Xudong Li, Shijia Chen, and Ren Sun 6668–6676 Reactivation of Kaposi’s Sarcoma-Associated Herpesvirus

Initial HIV-1 -Specific CD8؉ T Cells in Acute HIV-1 Stephanie A. Freel, Ralph A. Picking, 6835–6846 Infection Inhibit Transmitted/Founder Virus Replication Guido Ferrari, Haitao Ding, Christina Ochsenbauer, John C. Kappes, Jennifer L. Kirchherr, Kelly A. Soderberg, Kent J. Weinhold, Coleen K. Cunningham, Thomas N. Denny, John A. Crump, Myron S. Cohen, Andrew J. McMichael, Barton F. Haynes, and Georgia D. Tomaras

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Coordinate Regulation of DNA Damage and Type I Interferon Wadzanai P. Mboko, Bryan C. Mounce, 6899–6912 Responses Imposes an Antiviral State That Attenuates Mouse Brittani M. Wood, Joseph M. Kulinski, Gammaherpesvirus Type 68 Replication in Primary John A. Corbett, and Vera L. Macrophages Tarakanova Frequent and Strong Antibody-Mediated Natural Killer Cell Christina F. Thobakgale, Lena Fadda, 6986–6993 Activation in Response to HIV-1 in Individuals with Kimberly Lane, Ildiko Toth, Florencia Chronic HIV-1 Infection Pereyra, Suzane Bazner, Thumbi Ndung’u, Bruce D. Walker, Eric S. Rosenberg, Galit Alter, Mary Carrington, Todd M. Allen, and Marcus Altfeld TRANSFORMATION AND ONCOGENESIS Epstein-Barr Virus Latent Membrane Protein 2A Promotes Yu-Yan Lan, Jenn-Ren Hsiao, Kung- 6656–6667 Invasion of Nasopharyngeal Carcinoma Cells through ERK/Fra- Chao Chang, Jeffrey Shu-Ming Chang, 1-Mediated Induction of Matrix 9 Chaio-Wei Chen, Hsiao-Ching Lai, Shih-Yi Wu, Tzu-Hao Yeh, Fang-Hsin Chang, Wei-Hung Lin, Ih-Jen Su, and Yao Chang

The Epstein-Barr Virus (EBV)-Induced Tumor Suppressor Eleonora Forte, Raul E. Salinas, 6889–6898 MicroRNA MiR-34a Is Growth Promoting in EBV-Infected B Christina Chang, Ting Zhou, Sarah D. Cells Linnstaedt, Eva Gottwein, Cassandra Jacobs, Dereje Jima, Qi-Jing Li, Sandeep S. Dave, and Micah A. Luftig GENE DELIVERY Human Galectin 3 Binding Protein Interacts with Recombinant Jerome Denard, Cyriaque Beley, Robert 6620–6631 Adeno-Associated Virus Type 6 Kotin, René Lai-Kuen, Stéphane Blot, Hervé Leh, Aravind Asokan, R. Jude Samulski, Philippe Moullier, Thomas Voit, Luis Garcia, and Fedor Svinartchouk VACCINES AND ANTIVIRAL AGENTS A Lentiviral Vector-Based, Herpes Simplex Virus 1 (HSV-1) Flavia Chiuppesi, Laura Vannucci, 6563–6574 Glycoprotein B Vaccine Affords Cross-Protection against HSV-1 Anna De Luca, Michele Lai, Barbara and HSV-2 Genital Infections Matteoli, Giulia Freer, Roberto Manservigi, Luca Ceccherini-Nelli, Fabrizio Maggi, Mauro Bendinelli, and Mauro Pistello Distinct Effects of Two HIV-1 Capsid Assembly Inhibitor Christopher T. Lemke, Steve Titolo, 6643–6655 Families That Bind the Same Site within the N-Terminal Uta von Schwedler, Nathalie Goudreau, Domain of the Viral CA Protein Jean-François Mercier, Elizabeth Wardrop, Anne-Marie Faucher, René Coulombe, Soma S. R. Banik, Lee Fader, Alexandre Gagnon, Stephen H. Kawai, Jean Rancourt, Martin Tremblay, Christiane Yoakim, Bruno Simoneau, Jacques Archambault, Wesley I. Sundquist, and Stephen W. Mason Aptamer That Binds to the gD Protein of Herpes Simplex Virus Subash C. B. Gopinath, Kyoko Hayashi, 6732–6744 1 and Efficiently Inhibits Viral Entry and Penmetcha K. R. Kumar A Triclade DNA Vaccine Designed on the Basis of a Fan Zhou, Guiqin Wang, Philippe 6970–6978 Comprehensive Serologic Study Elicits Neutralizing Antibody Buchy, Zhipeng Cai, Honglin Chen, Responses against All Clades and Subclades of Highly Zhiwei Chen, Genhong Cheng, Xiu- Pathogenic Avian Influenza H5N1 Feng Wan, Vincent Deubel, and Paul Zhou

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PATHOGENESIS AND IMMUNITY Endothelial Cells Elicit Immune-Enhancing Responses to Nadine A. Dalrymple and Erich R. 6408–6415 Dengue Virus Infection Mackow Association of Major Histocompatibility Complex Class I Takushi Nomura, Hiroyuki Yamamoto, 6481–6490 Haplotypes with Disease Progression after Simian Teiichiro Shiino, Naofumi Takahashi, Immunodeficiency Virus Challenge in Burmese Rhesus Taku Nakane, Nami Iwamoto, Hiroshi Macaques Ishii, Tetsuo Tsukamoto, Miki Kawada, Saori Matsuoka, Akiko Takeda, Kazutaka Terahara, Yasuko Tsunetsugu-Yokota, Naoko Iwata- Yoshikawa, Hideki Hasegawa, Tetsutaro Sata, Taeko K. Naruse, Akinori Kimura, and Tetsuro Matano Effects of Manipulating Apoptosis on Infection of Hua Wang, Taryn Gort, Daniel L. 6546–6554 Aedes aegypti Mosquitoes Boyle, and Rollie J. Clem

CD4؉ CD25؉ Regulatory T Cells Impair HIV-1-Specific CD4 T Douglas S. Kwon, Mathieu Angin, 6586–6594 Cell Responses by Upregulating Interleukin-10 Production in Tomoyuki Hongo, Kenneth M. Law, Monocytes Jessica Johnson, Filippos Porichis, Meghan G. Hart, David F. Pavlik, Daniel P. Tighe, Daniel G. Kavanagh, Hendrik Streeck, Marylyn M. Addo, and Daniel E. Kaufmann

Multifunctional CD4 Cells Expressing Gamma Interferon and Deborah M. Brown, Sarah Lee, Maria 6792–6803 Perforin Mediate Protection against Lethal Influenza Virus de la Luz Garcia-Hernandez, and Susan Infection L. Swain Pathogenicity and Transmission in Pigs of the Novel A(H3N2)v Pravina Kitikoon, Amy L. Vincent, 6804–6814 Influenza Virus Isolated from Humans and Characterization of Phillip C. Gauger, Sarah N. Schlink, Swine H3N2 Viruses Isolated in 2010-2011 Darrell O. Bayles, Marie R. Gramer, Daniel Darnell, Richard J. Webby, Kelly M. Lager, Sabrina L. Swenson, and Alexander Klimov Uncommon Pathways of Immune Escape Attenuate HIV-1 Mark A. Brockman, Denis R. Chopera, 6913–6923 Replication Capacity Alex Olvera, Chanson J. Brumme, Jennifer Sela, Tristan J. Markle, Eric Martin, Jonathan M. Carlson, Anh Q. Le, Rachel McGovern, Peter K. Cheung, Anthony D. Kelleher, Heiko Jessen, Martin Markowitz, Eric Rosenberg, Nicole Frahm, Jorge Sanchez, Simon Mallal, Mina John, P. Richard Harrigan, David Heckerman, Christian Brander, Bruce D. Walker, and Zabrina L. Brumme A Single at the Hemagglutinin Cleavage Site Yi Zhang, Yipeng Sun, Honglei Sun, 6924–6931 Contributes to the Pathogenicity and Neurovirulence of H5N1 Juan Pu, Yuhai Bi, Yi Shi, Xishan Lu, Influenza Virus in Mice Jing Li, Qingyu Zhu, George F. Gao, Hanchun Yang, and Jinhua Liu

Mice Deficient in STAT1 but Not STAT2 or IRF9 Develop a Markus J. Hofer, Wen Li, Peter 6932–6946 ,Lethal CD4؉ T-Cell-Mediated Disease following Infection with Manders, Rachael Terry, Sue Ling Lim Lymphocytic Choriomeningitis Virus Nicholas J. C. King, and Iain L. Campbell

Elite Controllers with Low to Absent Effector CD8؉ T Cell Zaza M. Ndhlovu, Jacqueline 6959–6969 Responses Maintain Highly Functional, Broadly Directed Proudfoot, Kevin Cesa, Donna Marie Central Memory Responses Alvino, Ashley McMullen, Seanna Vine, Eleni Stampouloglou, Alicja Piechocka- Trocha, Bruce D. Walker, and Florencia Pereyra

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A Novel Variant Marking HLA-DP Expression Levels Predicts Rasmi Thomas, Chloe L. Thio, Richard 6979–6985 Recovery from Hepatitis B Virus Infection Apps, Ying Qi, Xiaojiang Gao, Darlene Marti, Judy L. Stein, Kelly A. Soderberg, M. Anthony Moody, James J. Goedert, Gregory D. Kirk, W. Keith Hoots, Steven Wolinsky, and Mary Carrington A Reverse Genetics Approach To Study Feline Infectious Gergely Tekes, Danica Spies, Barbara 6994–6998 Peritonitis Bank-Wolf, Volker Thiel, and Heinz- Jürgen Thiel GENOME ANNOUNCEMENTS Complete Genome Sequence of a Novel Porcine Enterovirus Wen Zhang, Shixing Yang, Quan Shen, 7008–7009 Strain in China Liping Ren, Tongling Shan, Jianzhong Wei, Li Cui, and Xiuguo Hua Complete Genome Sequence of Porcine Kobuvirus Strain Yuchen Lin, Shaobo Xiao, Yuqing 7010 WUH1 Zeng, Tao Song, Songlin Zeng, Huanchun Chen, and Liurong Fang Genome Sequence of a Reassortant Strain of Bluetongue Virus Sushila Maan, Narender S. Maan, Marc 7011–7012 Serotype 23 from Western India Guimera, Kyriaki Nomikou, Karam Pal Singh, Gillian Pullinger, Manjunatha N. Belaganahalli, and Peter P. C. Mertens Complete Genome Sequence of a Novel Marine Siphovirus, pVp- Ji Hyung Kim, Jin Woo Jun, Casiano H. 7013–7014 1, Infecting Vibrio parahaemolyticus Choresca, Sang Phil Shin, Jee Eun Han, and Se Chang Park Complete Genome Sequence of a New-Genotype Porcine Quan Shen, Wen Zhang, Shixing Yang, 7015–7016 Norovirus Isolated from Piglets with Diarrhea Li Cui, and Xiuguo Hua A Novel Porcine Circovirus Type 2a Strain, 10JS-2, with Eleven- Lin Cai, Xue Han, Dongmei Hu, 7017 Nucleotide Insertions in the Origin of Genome Replication Xiaoxia Li, Baoyue Wang, Jianqiang Ni, Zhi Zhou, Xiuling Yu, Xinyan Zhai, and Kegong Tian Complete Genome Sequence of the First Canine Circovirus Amit Kapoor, Edward J. Dubovi, Jose 7018 Angel Henriquez-Rivera, and W. Ian Lipkin Genome Sequence of the First Human Adenovirus Type 14 Qiwei Zhang, Donald Seto, Suhui Zhao, 7019–7020 Isolated in China Li Zhu, Wei Zhao, and Chengsong Wan Complete Genome Sequence of a Dengue Virus Serotype 4 Hui Zhao, Xue-Dong Yu, Xia-Yi 7021–7022 Strain Isolated in Guangdong, China Zhang, Tao Jiang, Wen-Xin Hong, Man Yu, Feng-Yu Hu, Shun-Ya Zhu, E-De Qin, Yong-Qiang Deng, Cheng-Feng Qin, and Fu-Chun Zhang Complete Genome Sequence of Avian Bornavirus Genotype 1 Negin Mirhosseini, Patricia L. Gray, Ian 7023 from a Macaw with Proventricular Dilatation Disease Tizard, and Susan Payne Organization of the Ambisense Genome of the Helicoverpa Mohamed El-Far, Jozsef Szelei, Qian 7024 armigera Densovirus Yu, Gilles Fédière, Max Bergoin, and Peter Tijssen

Cover photograph (Copyright © 2012, American Society for Microbiology. All Rights Reserved.): Schematic of the delivery of exosome- associated cargo molecules (e.g., microRNAs, mRNAs, polypeptides, and ) into a recipient host cell. Exosomes are secreted mi- crovesicles playing important roles in intercellular communication. Bioinformatics analysis demonstrates that different cellular microRNAs are found in exosomes from cells infected with different strains of influenza viruses. Dynamic intercellular communication between infected and noninfected bystander cells could involve virus strain-specific exosome-associated microRNAs. See related article in June 2012, vol. 86, no. 11, p. 6109.)