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ENCYCLOPEDIA OF VIROLOGY
FOURTH EDITION

Volume 1
 ENCYCLOPEDIA OF VIROLOGY
FOURTH EDITION

EDITORS IN CHIEF

Dennis H. Bamford
Molecular and Integrative Biosciences Research Programme
Faculty of Biological and Environmental Sciences
University of Helsinki, Helsinki, Finland

Mark Zuckerman
South London Specialist Virology Centre
King’s College Hospital NHS Foundation Trust
London, United Kingdom
and
Department of Infectious Diseases
School of Immunology and Microbial Sciences, King’s College London Medical School
London, United Kingdom

Volume 1

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EDITORS IN CHIEF

Dennis H. Bamford, PhD, is Professor Emeritus of Virology at the Faculty of Biological and
Environmental Sciences, University of Helsinki, Finland. He obtained his PhD in 1980 from
the Department of Genetics, University of Helsinki. During 1981–1982 he was an EMBO
postdoctoral fellow at the Public Health Research Institute of the City of New York, United
States, and during 1983–1992 he worked as a Senior Scientist at the Academy of Finland.
In 1993 he was appointed Professor of General Microbiology at the University of Helsinki.
He was awarded the esteemed Academy Professorship twice, in 2002–2007 and 2012–2016,
and he also served twice as the Director of the Finnish Center of Excellence (in Structural
Virology, 2000–2005, and in Virus Research, 2006–2011). Prof. Bamford has had continuous
external research funding (e.g., from several European Union, Academy of Finland, TEKES and
Jusélius Foundation funds, as well as the Human Frontier Science Program). He is an EMBO
member and has held several positions of trust in scientific and administrative organizations.
Prof. Bamford has published approx. 400 articles in international peer-reviewed journals in
virology, microbiology, biochemistry, and molecular biology (36 of them in high impact
journals). About half of the primary articles have been published with international collaborators
showing high international integration. He has also been invited to give 56 keynote and plenary
presentations in major international meetings. Prof. Bamford has supervised over 35 Master’s and
over 40 PhD theses. Seven of his graduate students or post docs have obtained a professorship and a similar number have a principal
investigator status. Prof. Bamford has studied virus evolution from a structure-centered perspective, showing that seemingly unrelated
viruses, such as bacteriophage PRD1 and human adenovirus have similar virion architecture. When the corona virion architecture was
gradually revealed, it was observed that its structural elements were close to those seen in RNA bacteriophage phi6 so that phi6 has been
actively used as surrogate for pathogenic viruses - quite a surprise!

Dr. Mark Zuckerman is Head of Virology, Consultant Medical Virologist, and Honorary Senior
Lecturer at South London Specialist Virology Centre, King’s College Hospital NHS Foundation
Trust and King’s College London Medical School, Department of Infectious Diseases, School of
Immunology and Microbial Sciences in London, United Kingdom. His interests include the
clinical interface between developing molecular diagnostic tests relevant to the local population
of patients, respiratory virus infections, herpesvirus infections in immunocompromised patients
and blood-borne virus transmission incidents in the healthcare setting. He has chaired the UK
Clinical Virology Network, Royal College of Pathologists Virology Specialty Advisory Committee
and Virology Examiners Panel and is a member of the Specialty Advisory Committee on
Transfusion Transmitted Viruses. He is a co-author on four editions of the “Mims’ Medical
Microbiology” textbook, has written chapters in a number of other textbooks and has over 100
publications in international peer-reviewed journals and is an associate editor for two journals.

v
EDITORIAL BOARD
Editors in Chief
Dennis H. Bamford
Molecular and Integrative Biosciences Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki,
Finland

Mark Zuckerman
South London Specialist Virology Centre, King’s College Hospital NHS Foundation Trust, London, United Kingdom and Department of
Infectious Diseases, School of Immunology and Microbial Sciences, King’s College London Medical School, London, United Kingdom

Section Editors
Claude M. Fauquet
St Louis, MO, United States
Michael Feiss
Department of Microbiology and Immunology, Carver College of Medicine, University of Iowa, Iowa City, IA, United States
Elizabeth E. Fry
Department of Structural Biology, Nuffield Department of Medicine, University of Oxford, Oxford, United Kingdom
Said A. Ghabrial†
Department of Plant Pathology, University of Kentucky, Lexington, KY, United States

Eric Hunter
Department of Pathology and Laboratory Medicine, Emory University School of Medicine and Emory Vaccine Center, Emory University,
Atlanta, GA, United States

Ilkka Julkunen
Institute of Biomedicine, University of Turku, Turku, Finland

Peter J. Krell
Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada

Mart Krupovic
Archaeal Virology Unit, Institut Pasteur, Paris, France

Maija Lappalainen
HUS Diagnostic Center, HUSLAB, Clinical Microbiology, University of Helsinki and Helsinki University Hospital, Helsinki, Finland

Hubert G.M. Niesters

Department of Medical Microbiology and Infection Prevention, Division of Clinical Virology, University Medical Center Groningen, Gro-
ningen, The Netherlands
Massimo Palmarini
MRC-University of Glasgow Centre for Virus Research, Glasgow, United Kingdom
David Prangishvili
Institut Pasteur, Paris, France and Ivane Javakhishvili Tbilisi State University, Tbilisi, Georgia

David I. Stuart
Department of Structural Biology, Nuffield Department of Medicine, University of Oxford, Oxford, United Kingdom and Diamond Light
Source, Didcot, United Kingdom

Nobuhiro Suzuki
Institute of Plant Stress and Resources (IPSR), Okayama University, Kurashiki, Japan

†
Deceased.

vii
SECTION EDITORS

Claude Fauquet received his PhD in biochemistry from University Louis Pasteur in Strasburg,
France in 1974. Dr. Fauquet joined the Institut de Recherche pour le Dévelopement (IRD)
and worked there as a plant virologist for 28 years, and served in Ivory Coast, West Africa for
14 years. In 1991, he founded the International Laboratory for Tropical Agricultural
Biotechnology (ILTAB) at The Scripps Research Institute, CA, United States. ILTAB was then
hosted by the Donald Danforth Plant Science Center, St. Louis, MO, from 1999 to 2012. In
2003, he co-founded the Global Cassava Partnership for the 21st Century (GCP21), which he
directed until 2019 and which goal is to improve the cassava crop worldwide.
Dr. Fauquet is an international leader in plant virology including taxonomy, epidemiol-
ogy, molecular virology, and in gene-silencing as an antiviral strategy. He was Secretary of the
International Committee on Taxonomy of Viruses (ICTV) for 18 years and the editor of
several ICTV Reports including the VIIIth ICTV Report in 2005.
He has published more than 300 research papers in reviewed journals and books. He is a
fellow of the American Association for the Advancement of Science, of the American
Phytopathological Society and a member of the St. Louis Academy of Sciences. In 2007,
Dr. Fauquet was knighted “Chevalier de l’Ordre des Palmes Académiques” by the French
Minister of High Education and Research.

Dr. Michael Feiss is Professor Emeritus in the Department of Microbiology and Immunology of
the Carver College of Medicine at the University of Iowa, IA, United States. Dr. Feiss received his
PhD in Genetics at the University of Washington followed by a postdoctoral traineeship in the
laboratory of Dr. Allan Campbell at Stanford. Dr. Feiss is a microbial geneticist who studies virus
assembly with an emphasis on how a DNA virus, bacteriophage lambda, packages viral DNA into
the empty prohead shell. The lab investigates how sites in the viral DNA orchestrate the initiation
and termination of the DNA packaging process. This work includes comprehensive examination
of the DNA recognition sites. A related interest is study of terminase, the viral DNA packaging
enzyme, including the functional domains for protein–DNA and protein–protein interactions. A
second focus has been the roles of the bacterial host’s IHF and DnaJ proteins in the lytic life cycle
of the virus. More recent work has involved a genetic dissection of the role of terminase’s
ATPase center that powers translocation of viral DNA into the prohead. This interest in the ATP
hydrolysis-driven packaging motor involves a multidisciplinary collaboration examining the
kinetics of DNA packaging during individual packaging events. Finally, recent studies have also
looked at how the packaging process has diverged among several lambda-like phages, including
phages 21, N15, and Gifsy-1.

Elizabeth E. Fry is a senior postdoctoral scientist in structural biology at the University of

Oxford, Oxford, United Kingdom, where she received her DPhil. for studies relating to the
structure determination of Foot-and-Mouth Disease Virus. Specializing in structural virology, Dr.
Fry has studied many virus/viral protein structures but her primary focus is on picornavirus
structure and function, in particular receptor interactions and virus uncoating. She is particularly
interested in rationally designing virus-like-particles as next generation vaccines to reduce the
inherent risks in handling live viruses.

ix
x Section Editors

Said A. Ghabrial† received his BSc in 1959 from Cairo University, Cairo, Egypt, and his PhD
from Louisiana State University, Baton Rouge, LA, United States, in 1965. Dr. Ghabrial did
postdoctoral research at the University of California, Davis, CA, United States, before returning
to Cairo, where he served as a plant virologist in the Ministry of Agriculture. He returned to
the United States in 1970 to do postdoctoral research at Purdue University, West Lafayette, IN.
In 1972, he joined the Plant Pathology Department at the University of Kentucky, Lexington, KY,
United States, where he rose to the rank of professor in 1986 and worked until 2013.
Dr. Ghabrial has served as an associate and senior editor of Phytopathology. He served on the
editorial boards of the Encyclopedia of Virology, 3rd edition and Encyclopedia of Plant Pathol-
ogy, and edited a thematic issue of Advances in Virus Research on “Mycoviruses”. He was a member
of the American Phytopathological Society (APS) and the American Society for Virology (ASV); in
July 2002 he was elected as a Fellow of the American Phytopathological Society. He also acted as
Chair of the ICTV Subcommittee on Fungal Viruses in 1987–1993 and 2011–2014.
His long professional career allowed him to make many scientific achievements in phy-
topathology and virology. Among them are molecular dissection of a legume-infecting RNA
virus, bean pod mottle virus (BPMV), development of BPMV-based vectors, discovery of a
transmissible debilitation disease of the phytopathogenic ascomycete, Helminthosporium vic-
toriae (Cochliobolus victoriae), establishment of a viral etiology of the H. victoriae disease, and
advancement of structural biology of diverse fungal viruses.

Eric Hunter, PhD, is Professor of Pathology and Laboratory Medicine at Emory University,
Atlanta, GA, United States. He serves as Co-Director of the Emory Center for AIDS Research
and is a Georgia Research Alliance Eminent Scholar.
Dr. Hunter’s research focus has been the molecular virology and pathogenesis of retroviruses,
including human immunodeficiency virus. He has made significant contributions to the under-
standing of the role of retroviral glycoprotein structural features during viral entry and providing
unique insights into the assembly and replication of this virus family. In recent years the emphasis of
his research has been on HIV transmission and pathogenesis, defining the extreme genetic bottleneck
and selection of viruses with unique traits during HIV heterosexual transmission. He has described
the selection of fitter viruses at the target mucosa, a gender difference in the extent of selection bias,
and a role for genital inflammation in reducing selection. His research has defined the impact of HIV
adaptation to the cellular immune response on immune recognition and control of HIV after
transmission, as well as on virus replicative fitness in vitro and in vivo. Recent work highlights the roles that virus replicative fitness and sex
of the host play in defining disease progression in a newly infected individual. His bibliography includes over 300 peer-reviewed articles,
reviews, and book chapters. He has also been the recipient of four NIH merit awards for his work on retrovirus and HIV molecular biology.
Dr. Hunter served as the Editor in Chief of the journal AIDS Research and Human Retroviruses for 10 years. He was Chair of the
AIDS Vaccine Research Subcommittee which is charged with providing advice and consultation on AIDS vaccine research to the
National Institute of Allergy and Infectious Diseases and continues to serve on editorial boards for several academic journals and
on external advisory committees for several government, academic, and commercial institutions.

Ilkka Julkunen graduated as an MD/PhD in 1984 from the Department of Virology, University of
Helsinki, Helsinki, Finland. He worked as a postdoctoral research fellow at Memorial Sloan-
Kettering Cancer Center in New York, United States, in 1986–1989, followed by positions as a
senior scientist, group leader and research professor at Finnish Institute for Health and Welfare in
1989–2013. In 2013 he became a Professor of Virology at the University of Turku, Turku, Finland.
The research interests of Dr. Julkunen have concentrated on innate and adaptive humoral
immunity in viral and microbial infections. He has studied intracellular signaling and RIG-I and
TLR-mediated activation of interferon system in human macrophages and dendritic cells and stable
cell lines in response to human and avian influenza, Sendai, Zika and coronavirus infections. In
addition, he has analyzed the downregulation of innate immunity by viral regulatory proteins from
influenza, HCV, flavi-, filo- and coronaviruses. He has expertise in vaccinology, biotechnology and
development of methods to analyze antiviral immunity, he has also been actively involved in
research training and collaborations with biotechnological industry.

†
Deceased.
 Section Editors xi

Peter Krell started his career in virology early as a summer high school student working for the
Canadian Forestry Service studying the resistance of nuclear polyhedrosis viruses (now called
baculoviruses) to environmental exposure with Dr. Fred T. Bird at the Insect Pathology Research
Institute in Sault Ste. Marie, ON, Canada. He received his BSc and MSc in biology from Carleton
University studying the iridovirus Tipula Iridescent Virus with Dr Peter Lee, in Ottawa, the Cana-
dian capital. For his PhD he headed east to Dalhousie University in Halifax, Nova Scotia on the
Atlantic coast. In addition to enjoying the salt sea air, fresh cod, lobster and mussels, he studied the
molecular biology of polydnaviruses under the guidance of Dr Don Stoltz. Heading south to Texas
A&M University in College Station, TX, United States, as a Postdoctoral Fellow he worked with Dr.
Max Summer, of baculovirus fame, and Dr. Brad Vinson continuing to study polydnaviruses, but
also became steeped in the early days of molecular baculovirology. He then accepted a faculty
position in the Department of Microbiology and Immunology at the University of Guelph in
Guelph, ON, Canada. There he switched to baculovirus research, which was more tractable, due in
part to available cell cultures and focused on viral DNA replication and functional genomics,
particularly on chitinase, cathepsin and ME53. In collaboration with Dr. Eva Nagy he studied molecular biology of different animal
viruses, notably Fowl Avian adenoviruses and their development as vaccine vectors, but also on the birnavirus infectious pancreatic
necrosis virus, the coronavirus porcine endemic diarrhea virus, fowlpox virus and the paramyxovirus Newcastle disease virus. He has been
involved extensively with virus taxonomy, being active in the International Committee on Taxonomy of Viruses (ICTV) as member of the
Polydnaviridae and Baculoviridae study groups, national representative of Canada on the ICTV, member of the Executive Committee for the
ICTV and Chair of the ICTV Invertebrate Virus Subcommittee. In terms of governance, Peter Krell was President of the Canadian Society
of Microbiology, Secretary and later President of the Society for Invertebrate Pathology, as well as being on the Editorial Boards of the
Canadian Journal of Microbiology and the ASM Journal of Virology. While at the University of Guelph, he rose through the ranks to Professor
and is currently University Professor Emeritus.

Mart Krupovic is the Head of the Archaeal Virology Unit in the Department of Microbiology at the
Institut Pasteur of Paris, France. He received his MSc in Biochemistry in 2005 from the Vilnius
University, Vilnius, Lithuania and PhD in 2010 in general microbiology from the University of
Helsinki, Helsinki, Finland. His current research focuses on the diversity, origin, and evolution of
viruses, as well as molecular mechanisms of virus–host interactions in archaea. He has published
over 170 journal articles and serves as an editor or on the editorial boards of Biology Direct, Research
in Microbiology, Scientific Reports, Virology, and Virus Evolution. He is also a member of the Executive
Committee of the International Committee on Taxonomy of Viruses (ICTV) and chairs the
Archaeal Viruses Subcommittee of the ICTV.

Maija Lappalainen, MD, PhD, Associate Professor of Clinical Microbiology, is the Head of
Clinical Microbiology in the HUS Diagnostic Center, HUSLAB, University of Helsinki and
Helsinki University Hospital, Helsinki, Finland. In her thesis during the years 1987–1992 she
studied the incidence and diagnostics of congenital toxoplasmosis. After PhD, her research
interest has been in diagnostic clinical virology, viral hepatitis, respiratory infections, viral
infections in the immunocompromised patients and viral infections during pregnancy.
xii Section Editors

Hubert G.M. Niesters (1958) studied biology and chemistry in Nijmegen, the Netherlands. After
obtaining his PhD in Utrecht (Prof. dr. M. Horzinek and Prof. dr. B. van der Zeijst, 1987) on the
molecular epidemiology of infectious bronchitis virus, he worked as a post-doctoral fellow with
Prof. dr. Jim Strauss at the California Institute of Technology (Pasadena, United States) on the
replication of Alphaviruses. He received a Niels Stensen fellowship (The Netherlands) and an E.S.
Gosney fellowship (Caltech) during this period.
After returning to the Netherlands (1989), he became a research associate in medical
microbiology at the Diagnostic Medical Center (Delft) but moved back to clinical virology as a
senior research associate in 1991 at the Erasmus University Medical Center Rotterdam (Head
Prof. dr. Ab Osterhaus). From 1993 to 2007, he was responsible for the molecular diagnostics
unit. During this period, he was involved in the discovery and characterization of several new
viruses and variants. In 2007, he became full professor and director of the Laboratory of Clinical
Virology within the Department of Medical Microbiology at the University Medical Center
Groningen and University of Groningen.
He has been actively involved in the implementation and development of new technologies like real-time amplification and
automation within clinical virology. He has been focusing on molecular diagnostics and its use and the clinical value in a
transplant setting, as well as in monitoring treatment of hepatitis viruses. Recently, his interest focuses on rapid regional epide-
miology, automation including MiddleWare solutions for molecular diagnostics, as well as the cost–benefit of rapid point-of-
impact molecular testing. Special interest is focused on raising awareness for the detection of enteroviruses (enterovirus D68) and
its relationship with acute flaccid myelitis (AFM).
Since 2017, he is the Chair of the executive board of QCMD (Quality Control of Molecular Diagnostics, Glasgow). He is an auditor
and team leader for the Dutch Council of Accreditation and Co-Editor in Chief of the Journal of Clinical Virology. He is an
(co)-author of more than 250 peer-reviewed papers, chapters and reviews including emerging viruses, such as enterovirus D68 and
hepatitis E virus (H-index 80).
For his entire work, he received in 2016 the “Ed Nowakowski Senior Memorial Clinical Virology Award” from the Pan
American Society for Clinical Virology.

Massimo Palmarini is the Director of the MRC-University of Glasgow Centre for Virus Research
and Chair of Virology at the University of Glasgow, Glasgow, United Kingdom. A veterinarian by
training, his research programs focus on the biology, evolution and pathogenesis of arboviruses
and the mechanisms of virus cross-species transmission. His work is funded by the MRC and the
Wellcome Trust. Massimo Palmarini has been elected Fellow of the Academy of Medical Sciences,
of the Royal Society of Edinburgh and of the Royal Society of Biology and he was a Wolfson-Royal
Society Research Merit Awardee. He is a Wellcome Trust Investigator.

David Prangishvili, PhD, Honorary Professor at the Institut Pasteur, Paris, France, and Professor
at Tbilisi State University, Tbilisi, Georgia, is one of the pioneers in studies on the biology of
Archaea and their viruses. His scientific career spans ex-USSR (Institute of Molecular Biology,
Moscow; 1970–1976), Georgia (Georgian National Academy of Sciences, Tbilisi; 1976–1991),
Germany (Max-Planck Institute for Biochemistry, Munich; University of Regensburg; 1991–2004)
and France (Institut Pasteur, Paris, 2004–2020). In the research groups headed by him, several
dozens of new species and eight new families of archaeal viruses have been discovered and
characterized, which display remarkable diversity of unique morphotypes and exceptional gen-
ome contents. The results of his research contribute to the knowledge on viral diversity on our
planet and change the field of prokaryotic virology, leading to the notion that viruses of
hyperthermophilic Archaea form a particular group in the viral world, distinctive from viruses of Bacteria and Eukarya, and to the
recognition of the virosphere of Archaea as one of the distinct features of this Domain of Life. David Prangishvili is a member of the
Academia Europaea, the European Academy of Microbiology, and the Georgian National Academy of Sciences.
 Section Editors xiii

David I. Stuart is MRC Professor of Structural Biology in the Nuffield Department of Med-
icine, Oxford University, Oxford, United Kingdom, Life Science Director at Diamond Light
Source and Director of Instruct-ERIC (pan-European organisation providing shared access to
infrastructure and methods for structural biology). He has diverse interests in structural
virology from picornaviruses, double-stranded RNA viruses and enveloped RNA viruses. His
drive to develop structural techniques led to the determination of the structure of Bluetongue
virus (1995) and then the first membrane containing virus, PRD1. More recently, he has been
at the fore-front of bringing Cryo-EM technology to bear on virus structure determination and
its future role in visualizing virus function in cellulo. In addition to basic science he has a
strong commitment to structural vaccinology and the development of antiviral drugs.

Dr. Nobuhiro Suzuki, PhD, received his MSc (1985) in phytopathology and PhD (1989) in
virology from Tohoku University in Sendai, Japan. Dr. Suzuki currently serves as a full Professor of
the Institute of Plant Stress and Resources, formerly Institute of Plant Sciences and Bioresouces at
Okayama University and as an Editor of Virus Research, Frontiers in Virology, Journal of General Plant
Pathology, Virology Journal, and Biology. He has also been Guest Editor to PLoS Pathogens, PNAS, and
mBio, and an Editorial Board member of Virology and Journal of Virology.
Suzuki Laboratory focuses on characterization of diverse viruses infecting phytopathogenic
fungi and exploration of their interplays. Recent achievements include the discovery of a neo-virus
lifestyle exhibited by a (+)ssRNA virus and an unrelated dsRNA virus in a plant pathogenic fungus
and of multilayer antiviral defense in fungi involving Dicer. Prior to coming to Kurashiki,
Okayama Prefecture, he was a visiting fellow of the Center for Agricultural Biotechnology at the
University of Maryland Biotechnology Institute (UMBI), College Park, MA, United States, for 4 years (1997–2001) to study molecular
biology of hypoviruses in the laboratory of Professor Donald L. Nuss. Before visiting UMBI, he served as an assistant professor and a
lecturer of the Biotechnology Institute at the Akita Prefectural College of Agriculture, Japan, for 11 years (1988–1998) where he was
engaged in a project on molecular characterization of rice dwarf phytoreovirus, a member of the family Reoviridae. He received awards
from the Japanese Phytopathological Society of Japan and Japanese Society for Virology for his outstanding achievements in plant and
fungal virology.
FOREWORD
I am delighted to write the foreword to this wonderful Fourth Edition of the Encyclopedia of Virology. The Third Edition was
published in 2008, how the world has changed in the intervening years. The release of the updated fourth edition could not be
more timely or more prescient. It is superb and a huge tribute to the authors, Elsevier the publisher, and to the brilliant editors,
Dennis Bamford and Mark Zuckerman.
SARS-CoV-2 has dominated the world since it emerged in 2019 and affected every continent and every aspect of life. A reminder,
if it were needed, of the impact of infectious diseases, the importance of virology and the vulnerability and interconnectivity of
our world.
There is no doubt that with rapidly changing ecology, urbanization, climate change, increased travel, and fragile public health
systems, epidemics and pandemics will become more frequent, more complex and harder to prevent and contain. Most of these
epidemics will be caused by viruses, those we know about and maybe able to predict and some we do not know of that will
emerge from animals, plants or the environment. Our changing climate will change the epidemiology of viruses, their vectors and
the infections they cause, hence the critical importance of this totally revised Fourth Edition of the Encyclopedia of Virology which
brings together research and an understanding of viruses in animals, plants, bacteria and fungi, the environment, and among
humans. Never has a holistic, one-health understanding been more important.
That starts with an understanding of the fundamentals of virology, a field of science that has been transformed in the years since
the Third Edition. An understanding transformed by embracing traditional fields of molecular and structural biology, genomics,
and influenced by immunology, genetics, pharmacology and increasingly by epidemiology and mathematics. Events of 2020 and
2021 also show why it is so important to integrate within traditional virology an understanding of the animal and human health
and behavior, of climate change and its impact on the ecology of viruses, plant sciences and vectors. And why we must understand
the viruses we think we know well, and those viruses less extensively studied. Research is critical to this, research that pushes the
boundaries of what we know, has the humility to seek answers to things we do not understand and shares that knowledge with the
widest possible community. That research will be most exciting at the interface between disciplines, most impactful when
dynamic, open, inclusive, global, and collaborative.
This is what the Fourth Edition of the Encyclopedia of Virology, the largest reference source of research in virology sets out to
achieve. It is a wonderful contribution to a critical field of knowledge. It contains new chapters, every chapter revised and updated
by a dedicated global community who have come together to provide what is a brilliant and inspiring reference. It is an honor to
contribute in a very small way to the timely release of the Fourth Edition of the Encyclopedia of Virology.

Jeremy Farrar

xv
PREFACE
The fourth edition of the Encyclopedia of Virology is encyclopedic, but we wanted to move away from an alphabetical list, apart from
where it was more logical, to a vision that encompassed a different structure. Articles describing novel trends as well as original
discoveries in specific subfields of virology have been distributed into a set of five volumes, namely Fundamentals of Virology, Human
and Animal Viruses, Plant Viruses, Bacterial, Archaeal, Fungal, Algal and Invertebrate Viruses, and Diagnosis, Treatment and Prevention
of Virus Infections.
We had hoped that the new edition would ‘go viral’ but it was ironic that the time to publication 12 years after the previous edition
had been made a bit longer due to a virus infection. The world encountered a devastating global pandemic, COVID-19, caused by a new
type of a coronavirus, SARS-CoV-2. Scientists in many disciplines all over the world started immediate efforts to discover solutions as to
how to mitigate and stop the spread of the pandemic. Virology moved from being a highly specialized subject to one in which everyone
became a virologist, proving just how significant the different aspects of virology are in terms of understanding the nature of viral
infection. Since the previous edition, the growth in the field of general virology has been enormous, including huge advances in basic
science, identification of novel viruses, diagnostic methods, treatment and prevention. Taking this into account, the introduction of the
articles within the Encyclopedia are very timely and crucial for providing a wealth of knowledge of the latest findings in the field of
virology to a vast range of people, whether school students, undergraduates, postgraduates, teachers, scientists, researchers, journalists
and others interested in infections and the conflict between the host and the pathogen.
Pandemic viruses have become a serious public concern in the changing world. We can ask ourselves whether we have reached
the point in which nature can no longer cope with the consequences of increased population density and human activities that are
harmful to the environment. Although several pandemics have threatened mankind before, this COVID-19 pandemic has high-
lighted the massive adverse economic consequences towards the wellbeing of society and the importance of research in virology.
We aimed to produce a Major Reference Work that differs in approach to others and binds all the virology disciplines together.
Chapters have been included on origin, evolution and emergence of viruses, environmental virology and ecology, epidemiology,
techniques for studying viruses, viral life cycles, structure, entry, genome and replication, assembly and packaging and taxonomy and
viral–host interactions. Information has been included on all known species of viruses infecting bacteria, fungi, plants, vertebrates and
invertebrates. Additional topics include antiviral classification and examples of their use in management of infection, diagnostic assays
and vaccines, as well as the economic importance of viral diseases of crops and their control.
This edition used viral classification according to the 9th Report of the International Committee on Taxonomy of Viruses published
in 2012. Updating it to the 10th Report in 2020 was affected by the pandemic and can be found online at http://ictv.global/report/.
We wish to acknowledge the hard work, interest, flexibility and patience, during such difficult times both socially and
professionally, of everybody involved in the process of writing this edition of the Encyclopedia of Virology, especially Katarzyna
Miklaszewska, Priscilla Braglia, Sam Crowe and colleagues at Elsevier. We sincerely thank all the authors and section editors for
their excellent contributions to this edition.

Book Cover Image: Viruses are obligate parasites and all cells have their own viruses increasing the total number of viruses to the
estimated astronomical number of 1031 that extends the number of stars in the universe. The viral string illustrates how pandemic
viruses surround the globe. The original picture was created by Dr. Nina Atanasova (Finnish Meteorological Institute and University of
Helsinki) and amended by Matthew Limbert at Elsevier.

Dennis H. Bamford
Mark Zuckerman

xvii
HOW TO USE THE ENCYCLOPEDIA
Structure of the Encyclopedia
All articles in the encyclopedia are arranged thematically as a series of entries within subjects/sections, apart from volume 2 where
there it was more logical to have articles arranged alphabetically.

There are three features to help you easily find the topic you are interested in: a thematic contents list, a full subject index, and
contributors.

1. Thematic contents list: The alphabetical contents list, which appears at the front of each volume, lists the entries in the order
that they appear in the encyclopedia.
2. Index: The index appears at the end of volume 5 and includes page numbers for quick reference to the information you are
looking for. The index entries differentiate between references to a whole entry, a part of an entry, and a table or figure.
3. Contributors: At the start of each volume there is a list of the authors who contributed to all volumes.

xix
LIST OF CONTRIBUTORS

Stephen T. Abedon Aleksandra Alimova

The Ohio State University, Mansfield, OH,
United States
Peter Abrahamian
Agricultural Research Service, US Department of
Agriculture, Beltsville, MD, United States
Jônatas S. Abrahão
Federal University of Minas Gerais, Belo Horizonte,
Brazil
Florence Abravanel
Toulouse University Hospital, Toulouse, France and
Toulouse University Paul Sabatier, Toulouse, France
Nicola G.A. Abrescia
Center for Cooperative Research in Biosciences, Basque
Research and Technology Alliance, Derio, Spain;
Ikerbasque, Basque Foundation for Science, Bilbao,
Spain; and Center for Biomedical Research in the Liver
and Digestive Diseases Network, Carlos III Health
Institute, Madrid, Spain
Gian Paolo Accotto
Institute for Sustainable Plant Protection, National
Research Council of Italy, Torino, Italy
Elisabeth Adderson
St. Jude Children’s Research Hospital, Memphis, TN,
United States and University of Tennessee Health
Sciences Center, Memphis, TN, United States
Mustafa Adhab
University of Baghdad, Baghdad, Iraq
Alexey A. Agranovsky
Lomonosov Moscow State University, Moscow, Russia
Nasim Ahmed
National Institute for Biotechnology and Genetic
Engineering, Faisalabad, Pakistan
Maher Al Rwahnih
University of California, Davis, CA, United States
Olufemi J. Alabi
Texas A& M AgriLife Research and Extension Center,
Weslaco, TX, United States
Aurélie A. Albertini
Institute for Integrative Biology of the Cell (I2BC),
French Alternative Energies and Atomic Energy
Commission, French National Center for Scientific
Research, Paris-Sud University, University of
Paris-Saclay, Gif-sur-Yvette, France
The City University of New York (CUNY), School of
Medicine, The City College of New York, New York,
NY, United States
Juan C. Alonso
National Biotechnology Center–Spanish National
Research Council, Madrid, Spain
Imran Amin
National Institute for Biotechnology and Genetic
Engineering, Faisalabad, Pakistan
Stephanie E. Ander
University of Colorado School of Medicine, Aurora, CO,
United States
Danielle E. Anderson
Duke-NUS Medical School, Singapore, Singapore
Ida Bagus Andika
Qingdao Agricultural University, Qingdao, China
Ana C.d.S.P. Andrade
Federal University of Minas Gerais, Belo Horizonte,
Brazil
Juana Angel
Pontifical Javeriana University, Bogota, Colombia
Vanesa Anton-Vazquez
King’s College Hospital, London, United Kingdom
Guido Antonelli
Sapienza University of Rome, Rome, Italy
Josefa Antón
University of Alicante, Alicante, Spain
Nanako Aoki
Tokyo University of Agriculture and Technology, Fuchu,
Japan
Timothy D. Appleby
King’s College Hospital, London, United Kingdom
Miguel Arenas
Department of Biochemistry, Genetics and Immunology,
University of Vigo, Vigo, Spain and CINBIO
(Biomedical Research Center), University of Vigo, Vigo,
Spain
Basil Arif
Laboratory for Molecular Virology, Great Lakes Forestry
Centre, Sault Ste Marie, ON, Canada

xxi
xxii List of Contributors
Vicente Arnau
Institute for Integrative Systems Biology (I2SysBio),
University of Valencia–Spanish National Research
Council, Valencia, Spain
Gaurav Arya
Duke University, Durham, NC, United States
Leyla Asadi
University of Alberta, Edmonton, AB, Canada
Sassan Asgari
The University of Queensland, Brisbane, QLD, Australia
Nina S. Atanasova
Finnish Meteorological Institute, Helsinki, Finland and
University of Helsinki, Helsinki, Finland
Houssam Attoui
UMR1161 Virologie, INRAE – French National
Research Institute for Agriculture, Food and
Environment, ANSES, Ecole Nationale Vétérinaire
d’Alfort, University of Paris-Est, Maisons-Alfort, France
Silvia Ayora
National Biotechnology Center–Spanish National
Research Council, Madrid, Spain
Walid Azab
Free University of Berlin, Berlin, Germany
Sasha R. Azar
The University of Texas Medical Branch, Galveston, TX,
United States
Fengwei Bai
The University of Southern Mississippi, Hattiesburg, MS,
United States
Dalan Bailey
The Pirbright Institute, Pirbright, United Kingdom
S.C. Baker
Loyola University of Chicago, Maywood, IL,
United States
Fausto Baldanti
University of Pavia, Pavia, Italy and Scientific Institute
for Research, Hospitalization and Healthcare, San
Matteo Polyclinic Foundation, Pavia, Italy
Logan Banadyga
Public Health Agency of Canada, Winnipeg, MB,
Canada
Ashley C. Banyard
Animal and Plant Health Agency, Addlestone,
United Kingdom; University of West Sussex, Falmer,
United Kingdom; and St. George's Medical School,
University of London, London, United Kingdom
Xiaoyong Bao
The University of Texas Medical Branch, Galveston, TX,
United States
Yiming Bao
Beijing Institute of Genomics, Chinese Academy of
Sciences, University of Chinese Academy of Sciences,
Beijing, China
Alan D.T. Barrett
The University of Texas Medical Branch, Galveston, TX,
United States
Diana P. Baquero
Archaeal Virology Unit, Institut Pasteur, Paris, France
and Sorbonne University, Paris, France
Moshe Bar-Joseph
Agricultural Research Organization, Volcani Center, Bet
Dagan, Israel
Rachael S. Barr
Bristol Royal Hospital for Children, Bristol,
United Kingdom
Ralf Bartenschlager
Heidelberg University, Heidelberg, Germany
David L.V. Bauer
Francis Crick Institute, London, United Kingdom
Oliver W. Bayfield
University of York, York, United Kingdom
Sally A. Baylis
Paul-Ehrlich-Institute, Langen, Germany
Philippa M. Beard
The Pirbright Institute, Pirbright, United Kingdom and The
Roslin Institute, University of Edinburgh, United Kingdom
Paul Becher
University of Veterinary Medicine, Hannover, Germany
Björn Becker
Saarland University, Saarbrücken, Germany
Karen L. Beemon
Johns Hopkins University, Baltimore, MD, United States
Martin Beer
Friedrich-Loeffler-Institute, Insel Riems, Germany
Jose Miguel Benito
Health Research Institute of the Jiménez Díaz
Foundation, Autonomous University of Madrid and Rey
Juan Carlos University Hospital, Móstoles, Spain
Mária Benkő
Institute for Veterinary Medical Research, Center for
Agricultural Research, Budapest, Hungary
 List of Contributors xxiii

Max Bergoin Maxime Boutier

National Institute of Scientific Research – Armand-
Frappier Health Research Centre, Laval, QC, Canada
Sabrina Bertin
Council for Agricultural Research and Economics,
Research Center for Plant Protection and Certification,
Rome, Italy
Shweta Bhatt
University of Copenhagen, Copenhagen, Denmark
Dennis K. Bideshi
California Baptist University, Riverside, CA,
United States and University of California, Riverside,
CA, United States
Yves Bigot
INRAE – French National Research Institute for
Agriculture, Food and Environment, Nouzilly, France
Richard J. Bingham
University of York, York, United Kingdom
Vera Bischoff
Institute for Chemistry and Biology of the Marine
Environment, Oldenburg, Germany
Kate N. Bishop
Francis Crick Institute, London, United Kingdom
Lindsay W. Black
The University of Maryland School of Medicine,
Baltimore, MD, United States
Romain Blanc-Mathieu
Institute for Chemical Research, Kyoto University, Kyoto,
Japan
Soile Blomqvist
National Institute for Health and Welfare, Helsinki,
Finland
Bryony C. Bonning
University of Florida, Gainesville, FL, United States
Lisa M. Bono
Rutgers, The State University of New Jersey,
New Brunswick, NJ, United States
Alexia Bordigoni
Aix-Marseille University, CNRS, IRD, Mediterranean
Institute of Oceanography, Marseille, France and
Aix-Marseille University, IRD257, Assistance-Publique
des Hôpitauxde Marseille, UMR Microbes, Evolution,
Phylogeny and Infections (MEPHI), IHU Méditerranée
Infection, Marseille, France
Mihnea Bostina
University of Otago, Dunedin, New Zealand
University of Liège, Liège, Belgium
P.R. Bowser
Cornell University, Ithaca, NY, United States
Daniel Bradshaw
Public Health England, London, United Kingdom
Claude Bragard
University of Louvain, Louvain-la-Neuve, Belgium
Aaron C. Brault
Centers for Disease Control and Prevention, Fort Collins,
CO, United States
Nicolas Bravo-Vasquez
St. Jude Children’s Research Hospital, Memphis, TN,
United States
Rob W. Briddon
University of Agriculture, Faisalabad, Pakistan
Thomas Briese
Columbia University, New York, NY, United States
Paul Britton
The Pirbright Institute, Pirbright, United Kingdom
Thomas J. Brouwers
Athena Institute, VU Amsterdam, Amsterdam, The
Netherlands
Kevin E. Brown
Frimley Park Hospital, Frimley, United Kingdom and
Immunisation and Countermeasures Division, Public
Health England, London, United Kingdom
Corina P.D. Brussaard
NIOZ Royal Netherlands Institute for Sea Research,
Den Burg, Texel, The Netherlands and Utrecht
University, Utrecht, The Netherlands
Harald Brüssow
Laboratory of Gene Technology, Department of
Biosystems, KU Leuven, Leuven, Belgium
Joachim J. Bugert
Bundeswehr Institute of Microbiology, Munich, Germany
Jozef J. Bujarski
Northern Illinois University, DeKalb, IL, United States
and Polish Academy of Sciences, Poznan, Poland
Laura Burga
University of Otago, Dunedin, New Zealand
Sara H. Burkhard
University Hospital of Zurich, Zurich, Switzerland
Cara C. Burns
Centers for Disease Control and Prevention, Atlanta,
GA, United States
xxiv List of Contributors

Felicity Burt Patrizia Cavadini

University of the Free State, Bloemfontein, South Africa The Lombardy and Emilia Romagna Experimental
Zootechnic Institute, Brescia, Italy
Kerry S. Burton
Leamington Spa, United Kingdom Supranee Chaiwatpongsakorn
Nationwide Children’s Hospital, Columbus, OH,
Sarah J. Butcher United States
University of Helsinki, Helsinki, Finland
Supriya Chakraborty
Mathias Büttner Jawaharlal Nehru University, New Delhi, India
Leipzig University, Leipzig, Germany
Yu-Chan Chao
Jesse Cahill Institute of Molecular Biology, Academia Sinica, Taipei,
Sandia National Labs, Albuquerque, NM, United States Taiwan
Marianna Calabretto Tyler P. Chavers
Sapienza University of Rome, Rome, Italy Centers for Disease Control and Prevention, Atlanta,
GA, United States
Thierry Candresse
The National Research Institute for Agriculture, Food Keping Chen
and the Environment, University of Bordeaux, Villenave Jiangsu University, Zhenjiang, China
d′Ornon, France Xiaorui Chen
Alan J. Cann Genomics Research Center, Academia Sinica, Taipei,
University of Leicester, Leicester, United Kingdom Taiwan
Yanping Chen
Lorenzo Capucci
Bee Research Laboratory, Agricultural Research Service,
The Lombardy and Emilia Romagna Experimental
US Department of Agriculture, Beltsville, MD,
Zootechnic Institute, Brescia, Italy
United States
Irene Carlon-Andres
Dayna Cheng
University of Oxford, Oxford, United Kingdom
National Cheng Kung University, Tainan, Taiwan
José M. Casasnovas Quentin Chesnais
National Center for Biotechnology, Spanish National University of Strasbourg, Colmar, France
Research Council (CSIC), Madrid, Spain
Sotaro Chiba
J.W. Casey Nagoya University, Nagoya, Japan
Cornell University, Ithaca, NY, United States
Wah Chiu
R.N. Casey Stanford University, Stanford, CA, United States
Cornell University, Ithaca, NY, United States
David Chmielewski
Sherwood R. Casjens Stanford University, Stanford, CA, United States
University of Utah, Salt Lake City, UT, United States
Irma E. Cisneros
Antonella Casola The University of Texas Medical Branch, Galveston, TX,
The University of Texas Medical Branch, Galveston, TX, United States
United States Lark L. Coffey
José R. Castón University of California, Davis, CA, United states
National Center for Biotechnology, Spanish National Alanna B. Cohen
Research Council, Madrid, Spain Rutgers University, New Brunswick, NJ, United States
Carlos E. Catalano Jeffrey I. Cohen
University of Colorado Anschutz Medical Campus, Laboratory of Infectious Diseases, National Institute of
Skaggs School of Pharmacy and Pharmaceutical Allergy and Infectious Diseases, National Institutes of
Sciences, Aurora, CO, United States Health, Bethesda, MD, United States
Roberto Cattaneo Seth Coleman
Mayo Clinic, Rochester, MN, United States Rice University, Houston, TX, United States
 List of Contributors xxv

Miquel Coll Amy Davis

Institute for Research in Biomedicine, Barcelona, Spain St Jude Children’s Research Hospital, Memphis, TN,
and Institute for Molecular Biology of Barcelona, United States
Barcelona, Spain
William O. Dawson
John Collinge Citrus Research and Education Center, Lake Alfred, FL,
UCL Institute of Prion Diseases, London, United States and University of Florida, Lake Alfred, FL,
United Kingdom United States
Carina Conceicao Erik De Clercq
The Pirbright Institute, Pirbright, United Kingdom Rega Institute for Medical Research, KU Leuven,
Leuven, Belgium
Gabriela N. Condezo
National Center for Biotechnology, Spanish National Raoul J. de Groot
Research Council, Madrid, Spain Utrecht University, Utrecht, The Netherlands
Michaela J. Conley Juan C. de la Torre
MRC-University of Glasgow Centre for Virus Research, The Scripps Research Institute, La Jolla, CA,
Glasgow, United Kingdom United States
Charles A Coomer Marcelo De las Heras
University of Oxford, Oxford, United Kingdom University of Zaragoza, Zaragoza, Spain
Anne K. Cordes Juliana Gabriela Silva de Lima
Hannover Medical School, Institute of Virology, Hannover, Federal University of Rio Grande do Norte, Natal, Brazil
Germany
Athos S. de Oliveira
Mauricio Cortes Jr. University of Brasília, Brasília, Brazil
Department of Chemistry, College of Arts and Sciences,
Nicole T. de Stefano
Fort Wayne, IN, United States
University of Florida, Gainesville, FL, United States
Robert H.A. Coutts
Greg Deakin
University of Hertfordshire, Hatfield, United Kingdom
NIAB-EMR, East Malling, United Kingdom
Jeff A. Cowley
Philippe Delfosse
CSIRO Livestock Industries, Brisbane, QLD, Australia
University of Luxembourg, Esch-sur-Alzette, Luxembourg
Robert W. Cross
Natacha Delrez
The University of Texas Medical Branch, Galveston, TX,
University of Liège, Liège, Belgium
United States
Tatiana A. Demina
Henryk Czosnek
Molecular and Integrative Biosciences Research Program,
The Hebrew University of Jerusalem, Rehovot, Israel
Faculty of Biological and Environmental Sciences,
Håkon Dahle University of Helsinki, Helsinki, Finland
Department of Biological Sciences, University of Bergen,
Ismail Demir
Bergen, Norway
Department of Biology, Karadeniz Technical University,
Janet M. Daly Trabzon, Turkey
University of Nottingham, Sutton Bonington,
Zihni Demirbağ
United Kingdom
Department of Biology, Karadeniz Technical University,
Subha Das Trabzon, Turkey
Okayama University, Kurashiki, Japan
X. Deng
Indranil Dasgupta Loyola University of Chicago, Maywood, IL,
University of Delhi, New Delhi, India United States
Sibnarayan Datta Cécile Desbiez
Defence Research Laboratory, Defence Research and Plant Pathology Unit, INRAE – French National
Development Organisation (DRDO), Tezpur, Assam, Research Institute for Agriculture, Food and
India Environment, Montfavet, France
xxvi List of Contributors

Christelle Desnues Lucy Dorrell

Aix-Marseille University, CNRS, IRD, Mediterranean
Institute of Oceanography, Marseille, France and
Aix-Marseille University, IRD 257, Assistance-Publique
des Hôpitaux de Marseille, UMR Microbes, Evolution,
Phylogeny and Infections (MEPHI), IHU Méditerranée
Infection, Marseille, France
Samantha J. DeWerff
University of Illinois at Urbana-Champaign, Urbana, IL,
United States
Daniele Di Carlo
Sapienza University of Rome, Rome, Italy
Arturo Diaz
La Sierra University, Riverside, CA, United States
Alfredo Diaz-Lara
University of California, Davis, CA, United States
Ralf G. Dietzgen
The University of Queensland, St. Lucia, QLD, Australia
Michele Digiaro
International Center for Advanced Mediterranean
Agronomic Studies (CIHEAM), Mediterranean
Agronomic Institute of Bari, Valenzano, Italy
Michael Dills
Montana State University, Bozeman, MT, United States
Wayne Dimech
National Serology Reference Laboratory, Fitzroy, VIC,
Australia
Savithramma P. Dinesh-Kumar
University of California, Davis, CA, United States
Linda K. Dixon
The Pirbright Institute, Pirbright, United Kingdom
Valerian V. Dolja
Oregon State University, Corvallis, OR, United States
University of Oxford, Oxford, United Kingdom
Rosemary A. Dorrington
Rhodes University, Grahamstown, South Africa
Andor Doszpoly
Hungarian Academy of Sciences, Budapest, Hungary
Andreas Dotzauer
University of Bremen, Bremen, Germany
Simon B. Drysdale
St George’s University Hospitals NHS Foundation Trust,
London, United Kingdom and St George’s, University of
London, London, United Kingdom
Robert L. Duda
University of Pittsburgh, Pittsburgh, PA, United States
Carol Duffy
University of Alabama, Tuscaloosa, AL, United States
Siobain Duffy
Rutgers, The State University of New Jersey,
New Brunswick, NJ, United States
David D. Dunigan
University of Nebraska–Lincoln, Lincoln, NE,
United States
Stéphane Duquerroy
University of Paris-Saclay, Orsay, France and Institut
Pasteur, Paris, France
Bas E. Dutilh
Utrecht University, Utrecht, The Netherlands and
Radboud University Medical Center, Nijmegen,
The Netherlands
Michael Edelstein
Faculty of Medicine, Bar Ilan University, Ramat Gan,
Israel

Aušra Domanska Herman K. Edskes

University of Helsinki, Helsinki, Finland National Institutes of Health, Bethesda, MD,
United States
Leslie L. Domier
Agricultural Research Service, US Department of Rosina Ehmann
Agriculture, Urbana, IL, United States Bundeswehr Institute of Microbiology, Munich, Germany

Pilar Domingo-Calap Toufic Elbeaino

Institute for Integrative Systems Biology (I2SysBio),
University of Valencia-CSIC, Valencia, Spain
Tatiana Domitrovic
Federal University of Rio de Janeiro, Rio de Janeiro,
Brazil
Sarah M. Doore
Michigan State University, East Lansing, MI,
United States
International Center for Advanced Mediterranean
Agronomic Studies (CIHEAM), Mediterranean
Agronomic Institute of Bari, Valenzano, Italy
Joanne B. Emerson
University of California, Davis, CA, United States
Ann Emery
University of North Carolina at Chapel Hill, Chapel
Hill, NC, United States

Christine E. Engeland
University Hospital Heidelberg and German Cancer
Research Center, Heidelberg, Germany and
Witten/Herdecke University, Witten, Germany
Luis Enjuanes
National Center for Biotechnology – Spanish National
Research Council (CNB-CSIC), Madrid, Spain
Katri Eskelin
University of Helsinki, Helsinki, Finland
Rosa Esteban
Institute of Biology and Functional Genomics,
CSIC/University of Salamanca, Salamanca, Spain
Mary K. Estes
Baylor College of Medicine, Houston, TX, United States
Cassia F. Estofolete
São José do Rio Preto School of Medicine, São José do
Rio Preto, Brazil
Alyssa B. Evans
National Institutes of Health, Hamilton, MT,
United States
Øystein Evensen
Norwegian University of Life Sciences, Oslo, Norway
Alex Evilevitch
Department of Experimental Medical Science, Lund
University, Lund, Sweden
Montserrat Fàbrega-Ferrer
Institute for Research in Biomedicine, Barcelona, Spain and
Institute for Molecular Biology of Barcelona, Barcelona,
Spain
Francesco Faggioli
Council for Agricultural Research and Economics,
Research Center for Plant Protection and Certification,
Rome, Italy
Bentley A. Fane
University of Arizona, Tucson, AZ, United States
Brian A. Federici
University of California, Riverside, CA, United States
F. Fenner
Australian National University, Canberra, ACT, Australia
Isabel Fernández de Castro
Cell Structure Laboratory, National Center for
Biotechnology – Spanish National Research Council
(CNB-CSIC), Madrid, Spain
Giovanni Ferrara
University of Alberta, Edmonton, AB, Canada
List of Contributors xxvii
Elvira Fiallo-Olivé
Institute for Mediterranean and Subtropical Horticulture
“La Mayora”–Spanish National Research Council–University
of Malaga, Algarrobo-Costa, Málaga, Spain
Andrew E. Firth
University of Cambridge, Cambridge, United Kingdom
Roland A. Fleck
King’s College London, London, United Kingdom
Ricardo Flores
Polytechnic University of Valencia, Higher Council of
Scientific Research, Valencia, Spain
Ervin Fodor
University of Oxford, Oxford, United Kingdom
Anthony R. Fooks
Animal and Plant Health Agency, Addlestone,
United Kingdom; University of Liverpool, Liverpool,
United Kingdom; and St. George's Medical School,
University of London, London, United Kingdom
Patrick Forterre
Archeal Virology Unit, Institut Pasteur, Paris, France
and French National Center for Scientific Research,
Institute of Integrative Biology of the Cell, University
of Paris-Saclay, Gif sur Yvette, France
Rennos Fragkoudis
University of Nottingham, Sutton Bonington,
United Kingdom and University of Edinburgh,
Edinburgh, United Kingdom
Manuel A. Franco
Pontifical Javeriana University, Bogota, Colombia
Giovanni Franzo
Department of Animal Medicine, Production and Health
(MAPS), Padua University, Padua, Italy
Graham L. Freimanis
The Pirbright Institute, Pirbright, United Kingdom
Juliana Freitas-Astúa
Brazilian Agricultural Research Corporation (Embrapa)
Cassava and Fruits, Cruz das Almas, Brazil
Elizabeth E. Fry
Department of Structural Biology, Nuffield Department
of Medicine, University of Oxford, Oxford,
United Kingdom
Marc Fuchs
Cornell University, Geneva, NY, United States
Tsutomu Fujimura
Institute of Biology and Functional Genomics,
CSIC/University of Salamanca, Salamanca, Spain
xxviii List of Contributors

Kuko Fuke Said A. Ghabrial†

Tokyo University of Agriculture and Technology, Fuchu,
Japan
Toshiyuki Fukuhara
Tokyo University of Agriculture and Technology, Fuchu,
Japan
To S. Fung
South China Agricultural University, Guangzhou,
China
Yahya Z.A. Gaafar
Julius Kuehn Institute – Federal Research Center for
Cultivated Plants, Braunschweig, Germany
Toni Gabaldon
Barcelona Supercomputing Center-National Center for
Supercomputing, Institute of Research in Biomedicine,
and Catalan Institution for Research and Advanced
Studies, Barcelona, Spain
Morgan Gaïa
University of Paris-Saclay, Evry, France
José Gallardo
National Center for Biotechnology, Spanish National
Research Council, Madrid, Spain
Hernan Garcia-Ruiz
University of Nebraska–Lincoln, Lincoln, NE,
United States
Juan A. García
National Center for Biotechnology-Spanish National
Research Council, Madrid, Spain
Matteo P. Garofalo
The University of Texas Medical Branch, Galveston, TX,
United States
Yves Gaudin
Institute for Integrative Biology of the Cell (I2BC),
French Alternative Energies and Atomic Energy
Commission, French National Center for Scientific
Research, Paris-Sud University, University of
Paris-Saclay, Gif-sur-Yvette, France
Andrew D.W. Geering
The University of Queensland, St. Lucia, QLD, Australia
Thomas W. Geisbert
The University of Texas Medical Branch, Galveston, TX,
United States
Andrea Gentili
Council for Agricultural Research and Economics,
Research Center for Plant Protection and Certification,
Rome, Italy
Volker Gerdts
University of Saskatchewan, Saskatoon, SK, Canada
Department of Plant Pathology, University of Kentucky,
Lexington, KY, United States
Clément Gilbert
Evolution, Genomes, Behavior and Ecology Laboratory,
CNRS University of Paris-Sud UMR 9191, IRD
UMR 247, Gif-sur-Yvette, France
Robert L. Gilbertson
University of California, Davis, CA, United States
Efstathios S. Giotis
Imperial College London, London, United Kingdom and
University of Essex, Colchester, United Kingdom
Laurent Glais
French Federation of Seed Potato Growers/Research,
Development, Promotion of Seed Potato, Paris, France
and Institute for Genetics, Environment and Plant
Protection, Agrocampus West, French National Institute
for Agriculture, Food and Environment, University of
Rennes 1, Le Rheu, France
Miroslav Glasa
Biomedical Research Center, Slovak Academy of
Sciences, Bratislava, Slovakia
Ido Golding
University of Illinois at Urbana-Champaign, Urbana, IL,
United States
Esperanza Gomez-Lucia
Complutense University of Madrid, Madrid, Spain
Zheng Gong
Beijing Institute of Genomics, Chinese Academy of
Sciences, Beijing, China
Andrea González-González
University of Florida, Gainesville, FL, United States
Michael M. Goodin
University of Kentucky, Lexington, KY, United States
Alexander E. Gorbalenya
Leiden University Medical Center, Leiden, The Netherlands
Paul Gottlieb
The City University of New York (CUNY), School of
Medicine, The City College of New York, New York,
NY, United States
M.-A. Grandbastien
INRAE – French National Research Institute for
Agriculture, Food and Environment, Versailles, France
†
Deceased.
 List of Contributors xxix

Meritxell Granell Sébastien Halary

National Center for Biotechnology, Madrid, Spain and National Museum of Natural History, UMR 7245
Institute of Chemical Research of Catalonia (ICIQ), CNRS/MNHN Molécule de Communication et
Tarragona, Spain Adaptation des Micro-organismes, Paris, France
Patrick L. Green Aron J. Hall
The Ohio State University, Columbus, OH, Centers for Disease Control and Prevention, Atlanta,
United States GA, United States

Sandra J. Greive John Hammond

University of York, York, United Kingdom Floral and Nursery Plants Research, Agricultural
Research Service, US Department of Agriculture,
Diane E. Griffin Beltsville, MD, United States
Johns Hopkins Bloomberg School of Public Health,
Baltimore, MD, United States Rosemarie W. Hammond
Agricultural Research Service, US Department of
Jonathan M. Grimes Agriculture, Beltsville, MD, United States
University of Oxford, Oxford, United Kingdom
Virginia Hargest
Nigel Grimsley St Jude Children’s Research Hospital, Memphis, TN,
Integrative Biology of Marine Organisms Laboratory, United States and University of Tennessee Health
Banuyls-sur-Mer, France and Sorbonne University, Science Center, Memphis, TN, United States
Banuyls-sur-Mer, France
Scott J. Harper
Bruno Gronenborn Washington State University, Prosser, WA, United States
Institute for Integrative Biology of the Cell, CNRS, Balázs Harrach
University of Paris-Sud, CEA, Gif sur Yvette, France Institute for Veterinary Medical Research, Center for
Julianne H. Grose Agricultural Research, Budapest, Hungary
Brigham Young University, Provo, UT, United States Masayoshi Hashimoto
Scott Grytdal The University of Tokyo, Tokyo, Japan
Centers for Disease Control and Prevention, Atlanta, Muhammad Hassan
GA, United States University of Agriculture, Faisalabad, Pakistan
Duane J. Gubler Asma Hatoum-Aslan
Duke-NUS Medical School, Singapore, Singapore University of Alabama, Tuscaloosa, AL, United States
Peixuan Guo Philippa C. Hawes
College of Pharmacy, The Ohio State University, The Pirbright Institute, Pirbright, United Kingdom
Columbus, OH, United States
Janelle A. Hayes
Tongkun Guo University of Massachusetts Medical School, Worcester,
Beijing Institute of Genomics, Chinese Academy of MA, United States
Sciences, Beijing, China Guijuan He
Anne-Lise Haenni Virginia Tech, Blacksburg, VA, United States
Institut Jacques Monod, French National Center for Klaus Hedman
Scientific Research, Paris Diderot University, Paris, University of Helsinki, Helsinki, Finland and Helsinki
France University Hospital, Helsinki, Finland
Susan L. Hafenstein Albert Heim
Pennsylvania State University, Hershey, PA, Hannover Medical School, Hanover, Germany
United States
Gary L. Hein
Ahmed Hafez University of Nebraska–Lincoln, Lincoln, NE,
Biotechvana, Valencia, Spain; Pompeu Fabra University, United States
Barcelona, Spain; and Minia University, Minya, Egypt
Manfred Heinlein
Marie Hagbom IBMP-CNRS, University of Strasbourg, Strasbourg,
Linköping University, Linköping, Sweden France
xxx List of Contributors

Mercedes Hernando-Pérez Elisabeth Huguet

National Center for Biotechnology, Spanish National
Research Council, Madrid, Spain
Carmen Hernández
Institute for Plant Molecular and Cell Biology (Spanish
National Research Council–Polytechnic University of
Valencia), Valencia, Spain
Etienne Herrbach
University of Strasbourg, Colmar, France
Stephen Higgs
Biosecurity Research Institute, Kansas State University,
Manhattan, KS, United States
Bradley I. Hillman
Rutgers University, New Brunswick, NJ, United States
Deborah M. Hinton
National Institute of Diabetes and Digestive and Kidney
Diseases, National Institutes of Health, Bethesda, MD,
United States
Judith Hirsch
Plant Pathology Unit, INRAE – French National
Research Institute for Agriculture, Food and
Environment, Montfavet, France
Jody Hobson-Peters
Australian Infectious Diseases Research Centre, School of
Chemistry and Molecular Biosciences, The University of
Queensland, St Lucia, QLD, Australia
Natalie M. Holste
University of Nebraska–Lincoln, Lincoln, NE,
United States
Jin S. Hong
Kangwon National University, Chunchon, South Korea
Margaret J. Hosie
MRC-University of Glasgow Centre for Virus Research,
Glasgow, United Kingdom
Olivia G. Howell
University of Alabama, Tuscaloosa, AL, United States
Liya Hu
Baylor College of Medicine, Houston, TX, United States
Zhaoyang Hu
Jiangsu University, Zhenjiang, China
Kuan-Ying A. Huang
Chang Gung Memorial Hospital, Taoyuan, Taiwan
Yu Huang
Peking University, Beijing, China
Natalia B. Hubbs
Hanover College, Hanover, IN, United States
Research Institute on Insect Biology, French National
Center for Scientific Research, University of Tours,
Tours, France
Roger Hull
John Innes Centre, Norwich, United Kingdom
Kiwamu Hyodo
Okayama University, Kurashiki, Japan
Eugénie Hébrard
Interactions Plantes Microorganismes Environnement,
Institut de Recherche pour le Développement, Centre de
coopération internationale en recherche agronomique
pour le développement, University of Montpellier,
Montpellier, France
Martin Hölzer
University of Jena, Jena, Germany
Tetsuro Ikegami
The University of Texas Medical Branch at Galveston,
Galveston, TX, United States
Niina Ikonen
Finnish Institute for Health and Welfare, Helsinki,
Finland
Cihan İnan
Department of Molecular Biology and Genetics,
Karadeniz Technical University, Trabzon, Turkey
İkbal Agah İnce
Department of Medical Microbiology, Acıbadem
University School of Medicine, Istanbul, Turkey
Katsuaki Inoue
Diamond Light Source, Didcot, United Kingdom
Toru Iwanami
Tokyo University of Agriculture, Tokyo, Japan
Jacques Izopet
Toulouse University Hospital, Toulouse, France and
Toulouse University Paul Sabatier, Toulouse, France
Fauziah Mohd Jaafar
UMR1161 Virologie, INRAE – French National
Research Institute for Agriculture, Food and
Environment, ANSES, Ecole Nationale Vétérinaire
d’Alfort, University of Paris-Est, Maisons-Alfort, France
Andrew O. Jackson
China Agricultural University, Beijing, China
Daral J. Jackwood
The Ohio State University/OARDC, Wooster, OH,
United States
 List of Contributors xxxi

Jean-Rock Jacques Laura Kakkola

Cellular and Molecular Epigenetics (GIGA), Liège,
Belgium and Molecular Biology (TERRA), Gembloux,
Belgium
Tiffany Jenkins
Nationwide Children’s Hospital, Columbus, OH,
United States and The Ohio State University, Columbus,
OH, United States
Jeffrey D. Jensen
Arizona State University, Tempe, AZ, United States
Daohong Jiang
Huazhong Agricultural University, Wuhan, China
Zhihao Jiang
China Agricultural University, Beijing, China
Allison R. Jilbert
The University of Adelaide, Adelaide, SA, Australia
Peng Jing
Department of Chemistry, College of Arts and Sciences,
Fort Wayne, IN, United States
Xixi Jing
Central South University, Changsha, China
Meesbah Jiwaji
Rhodes University, Grahamstown, South Africa
Kyle L. Johnson
The University of Texas at El Paso, El Paso, TX,
United States and CQuentia, Memphis, TN, United
States
Welkin E. Johnson
Boston College, Chestnut Hill, MA, United States
Ian M. Jones
University of Reading, Reading, United Kingdom and
London School of Hygiene and Tropical Medicine,
London, United Kingdom
Ramon Jordan
Agricultural Research Service, US Department of
Agriculture, Beltsville, MD, United States
Thomas Joris
Cellular and Molecular Epigenetics (GIGA), Liège,
Belgium and Molecular Biology (TERRA), Gembloux,
Belgium
Ilkka Julkunen
Institute of Biomedicine, University of Turku, Turku,
Finland
Sandra Junglen
Charité - University Medicine Berlin, Berlin, Germany
Masanori Kaido
Kyoto University, Kyoto, Japan
University of Turku, Turku, Finland
Hannimari Kallio-Kokko
University of Helsinki and Helsinki University Hospital,
Helsinki, Finland
Nassim Kamar
Toulouse University Hospital, Toulouse, France and
Toulouse University Paul Sabatier, Toulouse, France
Phyllis J. Kanki
Harvard T.H. Chan School of Public Health, Boston,
MA, United States
Peter Karayiannis
University of Nicosia, Nicosia, Cyprus
Henry M. Kariithi
Kenya Agricultural and Livestock Research Organization,
Nairobi, Kenya
Brian A. Kelch
University of Massachusetts Medical School, Worcester,
MA, United States
Karen E. Keller
Horticultural Crops Research Unit, Agricultural Research
Service, US Department of Agriculture, Corvallis, OR,
United States
Japhette E. Kembou-Ringert
University of Tel Aviv, Tel Aviv, Israel
Peter J. Kerr
University of Sydney, Sydney, NSW, Australia and
CSIRO Health and Biosecurity, Black Mountain
Laboratories, Canberra, ACT, Australia
Tiffany King
Nationwide Children’s Hospital, Columbus, OH,
United States and The Ohio State University College
of Medicine, Columbus, OH, United States
Andrea Kirmaier
Boston College, Chestnut Hill, MA, United States
Thomas Klose
Purdue University, West Lafayette, IN, United States
Barbara G. Klupp
Friedrich-Loeffler-Institute, Greifswald-Insel Riems,
Germany
David M. Knipe
Harvard Medical School, Boston, MA, United States
Nick J. Knowles
The Pirbright Institute, Pirbright, United Kingdom
Guus Koch
Wageningen Bioveterinary Research, Lelystad,
The Netherlands
xxxii List of Contributors
Renate Koenig
Julius Kühn Institute – Federal Research Center for
Cultivated Plants, Braunschweig, Germany
Susanne E. Kohalmi
The University of Western Ontario, London, ON,
Canada
Hideki Kondo
Okayama University, Kurashiki, Japan
Jennifer L. Konopka-Anstadt
Centers for Disease Control and Prevention, Atlanta,
GA, United States
Eugene V. Koonin
National Center for Biotechnology Information, National
Library of Medicine, Bethesda, MD, United States and
National Institutes of Health, Bethesda, MD, United States
Marion P.G. Koopmans
Erasmus Medical Center, Rotterdam, The Netherlands
Richard Kormelink
Wageningen University and Research, Wageningen,
The Netherlands
Ioly Kotta-Loizou
Imperial College London, London, United Kingdom
Peter J. Krell
Department of Molecular and Cellular Biology,
University of Guelph, Guelph, ON, Canada
Mart Krupovic
Archaeal Virology Unit, Institut Pasteur, Paris, France
Andreas Kuhn
University of Hohenheim, Stuttgart, Germany
Jens H. Kuhn
National Institutes of Health, Frederick, MD,
United States
Richard J. Kuhn
Purdue University, West Lafayette, IN, United States
Suvi Kuivanen
University of Helsinki, Helsinki, Finland
Ranjababu Kulasegaram
Guy’s and St Thomas’ NHS Foundation Trust, London,
United Kingdom
Raghavendran Kulasegaran-Shylini
Department of Pathogen Infection, Faculty of Infectious
and Tropical Diseases, London School of Hygiene and
Tropical Medicine, London, United Kingdom
Gaurav Kumar
University of Delhi, New Delhi, India
Manish Kumar
Jawaharlal Nehru University, New Delhi, India
Gael Kurath
US Geological Survey, Western Fisheries Research
Center, Seattle, WA, United States
Satu Kurkela
University of Helsinki and Helsinki University Hospital,
Helsinki, Finland
Wan-Chun Lai
Chang Gung Memorial Hospital, Taoyuan, Taiwan
Kevin Lamkiewicz
University of Jena, Jena, Germany
Rebecca K. Lane
University of Texas Health Science Center at San
Antonio, San Antonio, TX, United States
Andrew S. Lang
Memorial University of Newfoundland, St. John’s, NL,
Canada
Daniel Carlos Ferreira Lanza
Federal University of Rio Grande do Norte, Natal, Brazil
Maija Lappalainen
HUS Diagnostic Center, HUSLAB, Clinical
Microbiology, University of Helsinki and Helsinki
University Hospital, Helsinki, Finland
Katherine LaTourrette
University of Nebraska–Lincoln, Lincoln, NE,
United States
Chris Lauber
TWINCORE – Center for Experimental and Clinical
Infection Research, Hannover, Germany
Antonio Lavazza
The Lombardy and Emilia Romagna Experimental
Zootechnic Institute, Brescia, Italy
C. Martin Lawrence
Montana State University, Bozeman, MT, United States
Hervé Lecoq
Plant Pathology Unit, INRAE – French National
Research Institute for Agriculture, Food and
Environment, Montfavet, France
Young-Min Lee
Utah State University, Logan, UT, United States
Kristen N. LeGault
University of California, Berkeley, CA, United States
James Legg
International Institute of Tropical Agriculture, Dar es
Salaam, Tanzania
 List of Contributors xxxiii

Anne Legreve Walter Ian Lipkin

University of Louvain, Louvain-la-Neuve, Belgium Columbia University, New York, NY, United States
Petr G. Leiman Jan G. Lisby
The University of Texas Medical Branch, Galveston, TX, Copenhagen University Hospital Hvidovre, Hvidovre,
United States Denmark
Stanley M. Lemon Ding X. Liu
Lineberger Comprehensive Cancer Center, The South China Agricultural University, Guangzhou, China
University of North Carolina at Chapel Hill, NC,
Qiang Liu
United States and Department of Microbiology and
University of Saskatchewan, Saskatoon, SK, Canada
Immunology, The University of North Carolina at
Chapel Hill, NC, United States Sijun Liu
Iowa State University, Ames, IA, United States
Sebastian Leptihn
Zhejiang University-Edinburgh University Institute, Carlos Llorens
Zhejiang University, Haining, China Biotechvana, Scientific Park University of Valencia,
Valencia, Spain
Dennis J. Lewandowski
University of Florida, Lake Alfred, FL, United States L. Sue Loesch-Fries
Purdue University, West Lafayette, IN, United States
Sébastien Lhomme
Toulouse University Hospital, Toulouse, France and George P. Lomonossoff
Toulouse University Paul Sabatier, Toulouse, France John Innes Centre, Norwich, United Kingdom

Dawei Li L. Letti Lopez

China Agricultural University, Beijing, China The University of Texas at Austin, Austin, TX,
United States
Guangdi Li
Central South University, Changsha, China Alan T. Loynachan
University of Kentucky, Lexington, KY, United States
Guoqing Li
Huazhong Agricultural University, Wuhan, China Garry A. Luke
University of St. Andrews, St. Andrews, United Kingdom
Yi Li
Peking University, Beijing, China M. Luo
University of Alabama at Birmingham, Birmingham,
Zhefeng Li AL, United States
College of Pharmacy, The Ohio State University,
Columbus, OH, United States Juan J. López-Moya
Center for Research in Agricultural Genomics and
Zhenghe Li Spanish National Research Council, Barcelona, Spain
Zhejang University, Hangzhou, China
Che Ma
Jia Q. Liang Genomics Research Center, Academia Sinica, Taipei,
South China Agricultural University, Guangzhou, China Taiwan
Sebastian Liebe Stuart A. MacFarlane
Institute of Sugar Beet Research, Göttingen, Germany The James Hutton Institute, Invergowrie, United Kingdom
João Paulo Matos Santos Lima Saichetana Macherla
Federal University of Rio Grande do Norte, Natal, Brazil J. Craig Venter Institute, La Jolla, CA, United States

Bruno Lina Kensaku Maejima

HCL Department of Virology, National Reference Center The University of Tokyo, Tokyo, Japan
for Respiratory Viruses, Institute of Infectious Agents,
Fabrizio Maggi
Croix-Rousse Hospital, Lyon, France and Virpath
University of Pisa, Pisa, Italy and University of Insubria,
Laboratory, International Center of Research in
Varese, Italy
Infectiology (CIRI), INSERM U1111, CNRS—UMR
5308, École Normale Supérieure de Lyon, University Melissa S. Maginnis
Claude Bernard Lyon, Lyon University, Lyon, France The University of Maine, Orono, ME, United States
xxxiv List of Contributors

Edgar Maiss Chikara Masuta

Leibniz University Hannover, Hannover, Germany Hokkaido University, Sapporo, Japan
Kira S. Makarova Carlos P. Mata
National Center for Biotechnology Information, National University of Leeds, Leeds, United Kingdom
Library of Medicine, Bethesda, MD, United States
Jelle Matthijnssens
Ariana Manglli Rega Institute for Medical Research, KU Leuven,
Council for Agricultural Research and Economics, Leuven, Belgium
Research Center for Plant Protection and Certification,
Rome, Italy Claire P. Mattison
Centers for Disease Control and Prevention, Atlanta,
Annette Mankertz GA, United States and Cherokee Nation Assurance,
Robert Koch-Institute, Berlin, Germany Arlington, VA, United States
Pilar Manrique William McAllister
The Ohio State University, Wexner Medical Center, Rowan University School of Osteopathic Medicine,
Columbus, OH, United States Stratford, NJ, United States
Shahid Mansoor Alison A. McBride
National Institute for Biotechnology and Genetic National Institute of Allergy and Infectious Diseases,
Engineering, Faisalabad, Pakistan National Institutes of Health, Bethesda, MD,
Marco Marklewitz United States
Institute of Virology, Charité – University Medicine Michael McChesney
Berlin, Berlin, Germany University of California, Davis, CA, United States
Giovanni P. Martelli† Elaine McCulloch
University of Bari Aldo Moro, Bari, Italy Quality Control for Molecular Diagnostics (QCMD),
Darren P. Martin Glasgow, United Kingdom
University of Cape Town, Cape Town, South Africa Andrew J. McMichael
Robert R. Martin University of Oxford, Oxford, United Kingdom
Horticultural Crops Research Unit, Agricultural Research Alexander McPherson
Service, US Department of Agriculture, Corvallis, OR, University of California, Irvine, CA, United States
United States
Irene K. Meki
Manuel Martinez-Garcia French National Center for Scientific Research,
University of Alicante, Alicante, Spain Montpellier, France
Francisco Martinez-Hernandez
Ulrich Melcher
University of Alicante, Alicante, Spain
Oklaoma State University, Stillwater, OK, United States
Natalia Martín-González
Tomas A Melgarejo
Autonomous University of Madrid, Madrid, Spain
University of California, Davis, CA, United States
Joaquín Martínez Martínez
Michael J. Melzer
Bigelow Laboratory for Ocean Sciences, East Boothbay,
Department of Plant and Environmental Protection
ME, United States
Sciences, University of Hawaii, Honolulu, HI,
Manja Marz United States
University of Jena, Jena, Germany
Luiza Mendonça
Andrea Marzi University of Oxford, Oxford, United Kingdom
National Institute of Allergy and Infectious Diseases,
National Institutes of Health, Hamilton, MT, Xiang-Jin Meng
United States Virginia Polytechnic Institute and State University,
Blacksburg, VA, United States
Hema Masarapu
Sri Venkateswara University, Tirupati, India Peter P.C. Mertens
University of Nottingham, Sutton Bonington,
†
Deceased. United Kingdom
 List of Contributors xxxv

Thomas C. Mettenleiter Muhammad Mubin

Friedrich-Loeffler-Institute, Greifswald-Insel Riems, University of Agriculture, Faisalabad, Pakistan
Germany
Nicolas J. Mueller
Philip D. Minor University Hospital of Zurich, Zurich, Switzerland
St Albans, United Kingdom
Emmanuelle Muller
Ali Mirazimi The French Agricultural Research Center for
National Veterinary Institute, Uppsala, Sweden and International Development, Joint Research Units–Biology
Karolinska Hospital University, Huddinge, Sweden and Genetics of Plant-Pathogen Interactions,
Montpellier, France and Biology and Genetics of Plant-
Nischay Mishra Pathogen Interactions, University of Montpellier, The
Columbia University, New York, NY, United States French Agricultural Research Center for International
Edward S. Mocarski Development, French National Institute for Agricultural
Emory University School of Medicine, Atlanta, GA, Research, Montpellier SupAgro, Montpellier, France
United States John S. Munday
Florian Mock Massey University, Palmerston North, New Zealand
University of Jena, Jena, Germany Jacob H. Munson-McGee
Volker Moennig Montana State University, Bozeman, MT, United States
University of Veterinary Medicine, Hannover, Germany and Bigelow Laboratory for Ocean Sciences, East
Boothbay, ME, United States
Ian J. Molineux
The University of Texas at Austin, Austin, TX, Hacer Muratoğlu
United States Department of Molecular Biology and Genetics,
Karadeniz Technical University, Trabzon, Turkey
Aderito L. Monjane
Norwegian Veterinary Institute, Oslo, Finland Kenan C. Murphy
University of Massachusetts Medical School, Worcester,
Jacen S. Moore MA, United States
University of Tennessee Health Science Center,
Ugrappa Nagalakshmi
Memphis, TN, United States
University of California, Davis, CA, United States
Marc C. Morais
Keizo Nagasaki
The University of Texas Medical Branch, Galveston, TX,
Kochi University, Nankoku, Japan
United States
Nazia Nahid
Cristina Moraru
GC University, Faisalabad, Pakistan and University of
Institute for Chemistry and Biology of the Marine
Agriculture, Faisalabad, Pakistan
Environment, Oldenburg, Germany
Venugopal Nair
Hiromitsu Moriyama
The Pirbright Institute, Pirbright, United Kingdom
Tokyo University of Agriculture and Technology, Tokyo,
Japan Remziye Nalçacıoğlu
Department of Molecular Biology and Genetics,
Sergey Y. Morozov
Karadeniz Technical University, Trabzon, Turkey
Lomonosov Moscow State University, Moscow, Russia
Shigetou Namba
Thomas E. Morrison
The University of Tokyo, Tokyo, Japan
University of Colorado School of Medicine, Aurora, CO,
United States Rubab Z. Naqvi
National Institute for Biotechnology and Genetic
Léa Morvan
Engineering, Faisalabad, Pakistan
University of Liège, Liège, Belgium
Rachel Nash
Benoît Moury The Pirbright Institute, Surrey, United Kingdom
Plant Pathology Unit, INRAE – French National
Institute for Agriculture, Food and Environment, C.K. Navaratnarajah
Montfavet, France Purdue University, West Lafayette, IN, United States
xxxvi List of Contributors

Maria A. Navarrete-Muñoz Hanna M. Oksanen

Biotechvana, Madrid, Spain; Institute of Health
Research-Jiménez Díaz Foundation, Autonomous
University of Madrid; and Rey Juan Carlos University
Hospital, Móstoles, Spain
Jesús Navas-Castillo
Institute for Mediterranean and Subtropical Horticulture
“La Mayora”–Spanish National Research Council–
University of Malaga, Algarrobo-Costa, Málaga, Spain
Muhammad S. Nawaz-ul-Rehman
University of Agriculture, Faisalabad, Pakistan
Christopher L. Netherton
The Pirbright Institute, Pirbright, United Kingdom
Thu V.P. Nguyen
Baylor College of Medicine, Houston, TX, United States
Molecular and Integrative Biosciences Research Program,
Faculty of Biological and Environmental Sciences,
University of Helsinki, Helsinki, Finland
Graziele P. Oliveira
Federal University of Minas Gerais, Belo Horizonte,
Brazil
Francesco Origgi
University of Bern, Bern, Switzerland
Nikolaus Osterrieder
Free University of Berlin, Berlin, Germany
Robert A. Owens
Beltsville Agricultural Research Center, Beltsville, MD
20705, United States

Annette Niehl Emine Özsahin

Julius Kühn Institute – Federal Research Center for
Cultivated Plants, Braunschweig, Germany
Hubert G.M. Niesters
Department of Medical Microbiology and Infection
Prevention, Division of Clinical Virology, University
Medical Center Groningen, Groningen, The Netherlands
Jozef I. Nissimov
University of Waterloo, Waterloo, ON, Canada
Norman Noah
London School of Hygiene and Tropical Medicine,
London, United Kingdom
Mauricio L. Nogueira
São José do Rio Preto School of Medicine, São José do
Rio Preto, São Paulo, Brazil
Johan Nordgren
Linköping University, Linköping, Sweden
University of Guelph, Guelph, ON, Canada
Sergi Padilla-Parra
University of Oxford, Oxford, United Kingdom;
Department of Infectious Diseases, Faculty of Life
Sciences and Medicine, King’s College London, London,
United Kingdom; and Randall Division of Cell and
Molecular Biophysics, King’s College London, London,
United Kingdom
Joshua Pajak
Duke University, Durham, NC, United States
Massimo Palmarini
MRC-University of Glasgow Centre for Virus Research,
Glasgow, United Kingdom
Amanda R. Panfil
The Ohio State University, Columbus, OH,
United States

C. Micha Nübling Marcus Panning

Paul-Ehrlich-Institute, Langen, Germany Institute of Virology, Freiburg University Medical Center,
Faculty of Medicine, University of Freiburg, Freiburg,
Visa Nurmi Germany
University of Helsinki, Helsinki, Finland
Vitantonio Pantaleo
Donald L. Nuss
National Research Council, Research Unit of Bari, Bari,
University of Maryland, Rockville, MD, United States
Italy
M. Steven Oberste
Centers for Disease Control and Prevention, Atlanta, Anna Papa
GA, United States Aristotle University of Thessaloniki, Thessaloniki, Greece

Hiroyuki Ogata Nikolaos Pappas

Institute for Chemical Research, Kyoto University, Kyoto, Utrecht University, Utrecht, The Netherlands
Japan
Hanu R. Pappu
Ane Ogbe Washington State University, Pullman, WA,
University of Oxford, Oxford, United Kingdom United States

Kristin N. Parent
Michigan State University, East Lansing, MI,
United States
Colin R. Parrish
Cornell University, Ithaca, NY, United States
A. Lorena Passarelli
Kansas State University, Manhattan, KS, United States
Basavaprabhu L. Patil
ICAR–Indian Institute of Horticultural Research,
Bengaluru, India
Jade Pattyn
University of Antwerp, Antwerp, Belgium
T.A. Paul
Cornell University, Ithaca, NY, United States
Lillian Pavlik
Laboratory for Molecular Virology, Great Lakes Forestry
Centre, Sault Ste Marie, ON, Canada
Susan L. Payne
Texas A& M University, College Station, TX, United States
Michael N. Pearson
The University of Auckland, Auckland, New Zealand
Mark E. Peeples
Nationwide Children’s Hospital, Columbus, OH,
United States and The Ohio State University College of
Medicine, Columbus, OH, United States
Ben Peeters
Wageningen Bioveterinary Research, Lelystad,
The Netherlands
Joseph S.M. Peiris
The University of Hong Kong, Pok Fu Lam, Hong Kong
Malik Peiris
The University of Hong Kong, Pok Fu Lam, Hong Kong
Judit J. Pénzes
National Institute of Scientific Research – Armand-
Frappier Health Research Centre, Laval, QC, Canada
Miryam Pérez-Cañamás
Institute for Plant Molecular and Cell Biology (Spanish
National Research Council–Polytechnic University of
Valencia), Valencia, Spain
Marta Pérez-Illana
National Center for Biotechnology, Spanish National
Research Council, Madrid, Spain
Jaume Pérez-Sánchez
Institute of Aquaculture Torre de la Sal, Spanish
National Research Council, Castellon, Spain
List of Contributors xxxvii
Jean-Marie Peron
Toulouse University Hospital, Toulouse, France and
Toulouse University Paul Sabatier, Toulouse, France
Karin E. Peterson
National Institutes of Health, Hamilton, MT, United States
Karel Petrzik
Biology Center CAS, Institute of Plant Molecular
Biology, České Budějovice, Czech Republic
Mahtab Peyambari
Pennsylvania State University, State College, PA,
United States
Sujal Phadke
J. Craig Venter Institute, La Jolla, CA, United States
Hanh T. Pham
National Institute of Scientific Research – Armand-
Frappier Health Research Centre, Laval, QC, Canada
Mauro Pistello
University of Pisa, Pisa, Italy
Daniel Ponndorf
John Innes Centre, Norwich, United Kingdom
Leo L.M. Poon
The University of Hong Kong, Pok Fu Lam, Hong Kong
Welkin H. Pope
University of Pittsburgh, Pittsburgh, PA, United States
Minna M. Poranen
University of Helsinki, Helsinki, Finland
Claudine Porta
The Pirbright Institute, Pirbright, United Kingdom and
University of Oxford, Oxford, United Kingdom
Samuel S. Porter
National Institute of Allergy and Infectious Diseases,
National Institutes of Health, Bethesda, MD,
United States and University of Maryland, College Park,
MD, United States
Frank A. Post
King's College Hospital NHS Foundation Trust, London,
United Kingdom
Nils Poulicard
Interactions Plantes Microorganismes Environnement,
Institut de Recherche pour le Développement, Centre de
coopération internationale en recherche agronomique
pour le développement, University of Montpellier,
Montpellier, France
David Prangishvili
Institut Pasteur, Paris, France and Ivane Javakhishvili
Tbilisi State University, Tbilisi, Georgia
xxxviii List of Contributors

B. V. Venkataram Prasad Chris M. Rands

Baylor College of Medicine, Houston, TX, United States
Lalita Priyamvada
Centers for Disease Control and Prevention, Atlanta,
GA, United States
Simone Prospero
Swiss Federal Institute for Forest, Snow and Landscape
Research WSL, Birmensdorf, Switzerland
Elisabeth Puchhammer-Stöckl
Medical University of Vienna, Vienna, Austria
Jianming Qiu
University of Kansas Medical Center, Kansas City, KS,
United States
S.L. Quackenbush
Colorado State University, Fort Collins, CO, United States
Killian J. Quinn
King’s College Hospital, London, United Kingdom
Diego F. Quito-Avila
Department of Life Sciences, ESPOL Polytechnic
University, Guayaquil, Ecuador
Frank Rabenstein
Julius Kühn Institute, Quedlinburg, Germany
Sheli R. Radoshitzky
United States Army Medical Research Institute of
Infectious Diseases, Frederick, MD, United States
Saleem U. Rahman
National Institute for Biotechnology and Genetic
Engineering, Faisalabad, Pakistan
Mbolarinosy Rakotomalala
FOFIFA, Antananarivo, Madagascar
Norma Rallon
Institute of Health Research-Jiménez Díaz Foundation,
Autonomous University of Madrid and Rey Juan Carlos
University Hospital, Móstoles, Spain
Robert P. Rambo
Diamond Light Source, Didcot, United Kingdom
Bertha Cecilia Ramirez
The Institute for Integrative Biology of the Cell,
The French Alternative Energies and Atomic Energy
Commission, French National Center for Scientific
Research, University of Paris-Sud, University of
Paris-Saclay, Gif-sur-Yvette, France
María D. Ramos-Barbero
University of Alicante, Alicante, Spain
University of Geneva Medical School and Swiss Institute
of Bioinformatics, Geneva, Switzerland
Venigalla B. Rao
The Catholic University of America, Washington, DC,
United States
Janne J. Ravantti
University of Helsinki, Helsinki, Finland
Mandy Ravensbergen
Wageningen University and Research, Wageningen,
The Netherlands
Georget Y. Reaiche-Miller
The University of Adelaide, Adelaide, SA, Australia
D.V.R. Reddy
International Crops Research Institute for the Semi-Arid
Tropics, Hyderabad, India
Vishwanatha R.A.P. Reddy
The Pirbright Institute, Pirbright, United Kingdom
Juan Reguera
Aix-Marseille University, French National Center for
Scientific Research, Marseille, France and French
National Institute of Health and Medical Research,
Marseille, France
William K. Reisen
University of California, Davis, CA, United states
Jingshan Ren
University of Oxford, Oxford, United Kingdom
Renato O. Resende
University of Brasilia, Brasilia, Brazil
Peter A. Revill
The Peter Doherty Institute of Infection and Immunity,
Royal Melbourne Hospital, Melbourne, VIC, Australia
Félix A. Rey
Institut Pasteur, Paris, France
Lara Rheinemann
University of Utah, Salt Lake City, UT, United States
Simone G. Ribeiro
Embrapa Genetic Resources and Biotechnology, Brasília,
Brazil
Daniel Rigling
Swiss Federal Institute for Forest, Snow and Landscape
Research (WSL), Birmensdorf, Switzerland
Cristina Risco
Cell Structure Laboratory, National Center for
Biotechnology – Spanish National Research Council
(CNB-CSIC), Madrid, Spain
 List of Contributors xxxix

Efraín E. Rivera-Serrano Polly Roy

Lineberger Comprehensive Cancer Center, The University Department of Pathogen Infection, Faculty of Infectious
of North Carolina at Chapel Hill, NC, United States and Tropical Diseases, London School of Hygiene and
and Department of Microbiology and Immunology, Tropical Medicine, London, United Kingdom and
The University of North Carolina at Chapel Hill, NC, University of Reading, Reading, United Kingdom
United States
Aaron P. Roznowski
Cécile Robin The University of Texas at Austin, Austin, TX,
INRAE – French National Research Institute for United States and University of Arizona, Tucson, AZ,
Agriculture, Food and Environment, UMR BIOGECO, United States
Cestas, France
Luisa Rubino
Rodrigo A.L. Rodrigues Institute for Sustainable Plant Protection, National
Federal University of Minas Gerais, Belo Horizonte, Research Council, Bari, Italy
Brazil
Olli Ruuskanen
Elina Roine
Turku University Hospital, Turku, Finland
University of Helsinki, Helsinki, Finland
Maria R. Rojas Eugene V. Ryabov
University of California, Davis, CA, United States USDA, Agricultural Research Service, Beltsville, MD,
United States
Marilyn J. Roossinck
Pennsylvania State University, State College, PA, Martin D. Ryan
United States University of St. Andrews, St. Andrews, United Kingdom
Vera I.D. Ros Ki H. Ryu
Wageningen University and Research, Wageningen, Seoul Women’s University, Seoul, South Korea
The Netherlands
Hanns-Joachim Rziha
Cristina Rosa Eberhard Karls University of Tübingen, Tübingen,
Pennsylvania State University, University Park, PA, Germany
United States
Sead Sabanadzovic
Hanna Rose
Mississippi State University, Starkville, MS,
Leibniz University Hannover, Hannover, Germany
United States
David A. Rosenbaum
University of Florida, Gainesville, FL, United States Roghaiyeh Safari
Cellular and Molecular Epigenetics (GIGA), Liège,
Shannan L. Rossi Belgium and Molecular Biology (TERRA), Gembloux,
The University of Texas Medical Branch, Galveston, TX, Belgium
United States
Azeez Sait Sahul Hameed
Michael G. Rossmann† C. Abdul Hakeem College, Melvisharam, India
Purdue University, West Lafayette, IN, United States
Nicole Samies
L. Roux University of Alabama at Birmingham, Birmingham,
University of Geneva Medical School, Geneva, Switzerland AL, United States
Simon Roux Carmen San Martín
US Department of Energy Joint Genome Institute, National Center for Biotechnology, Spanish National
Walnut Creek, CA, United States Research Council, Madrid, Spain
J. Rovnak
Ruth-Anne Sandaa
Colorado State University, Fort Collins, CO,
Department of Biological Sciences, University of Bergen,
United States
Bergen, Norway
David J. Rowlands
University of Leeds, Leeds, United Kingdom Hélène Sanfaçon
Agriculture and Agri-Food Canada, Summerland, BC,
†
Deceased. Canada
xl List of Contributors

Rafael Sanjuán Declan C. Schroeder

Institute for Integrative Systems Biology (I2SysBio), University of Reading, Reading, United Kingdom and
University of Valencia-CSIC, Valencia, Spain University of Minnesota, St. Paul, MN, United States
Neeraja Sankaran Stacey Schultz-Cherry
Utrecht University, Utrecht, The Netherlands St. Jude Children’s Research Hospital, Memphis, TN,
Fernando Santos United States
University of Alicante, Alicante, Spain Thomas F. Schulz
Cecilia Sarmiento Hannover Medical School, Institute of Virology,
Tallinn University of Technology, Tallinn, Estonia Hannover, Germany and German Center for Infection
Research, Hannover-Braunschweig Site, Braunschweig,
Takahide Sasaya Germany
National Agriculture and Food Research Organization,
Fukuyama, Japan Catherine A. Scougall
The University of Adelaide, Adelaide, SA, Australia
Preethi Sathanantham
Virginia Tech, Blacksburg, VA, United States Kimberley D. Seed
University of California, Berkeley, CA, United States
Panayampalli S. Satheshkumar
Centers for Disease Control and Prevention, Atlanta, Joaquim Segalés
GA, United States Departament of Animal Health and Anatomy, Faculty of
Veterinary Medicine, Autonomous University of
Yukiyo Sato
Barcelona, Barcelona, Spain; Animal Health Research
Okayama University, Kurashiki, Japan
Center (CReSA) – Institute of Agrifood Research and
Andreas Sauerbrei Technology (IRTA), Campus UAB, Barcelona, Spain;
Jena University Hospital, Jena, Germany and OIE Collaborating Center for the Research and
Control of Emerging and Re-emerging Swine Diseases in
Eugene I. Savenkov
Europe (IRTA-CReSA), Barcelona, Spain
Swedish University of Agricultural Sciences, Uppsala,
Sweden and Linnean Center for Plant Biology, Uppsala, Mateo Seoane-Blanco
Sweden National Center for Biotechnology, Madrid, Spain
Carita Savolainen-Kopra Madhumati Sevvana
National Institute for Health and Welfare, Helsinki, Purdue University, West Lafayette, IN, United States
Finland
Kazım Sezen
Kay Scheets Department of Biology, Karadeniz Technical University,
Oklahoma State University, Stillwater, OK, Trabzon, Turkey
United States
Arvind Sharma
Uffe V. Schenider Institut Pasteur, Paris, France
Copenhagen University Hospital Hvidovre, Hvidovre,
Denmark Sumit Sharma
Linköping University, Linköping, Sweden
Richard H. Scheuermann
J. Craig Venter Institute, La Jolla, CA, United States; James M. Sharp
University of California, San Diego, CA, United States; University of Zaragoza, Zaragoza, Spain and Edinburgh,
La Jolla Institute for Immunology, La Jolla, CA, United Kingdom
United States; and Global Virus Network, Baltimore,
MD, United States Qunxin She
Shandong University, Qingdao, China
Manfred J. Schmitt
Saarland University, Saarbrücken, Germany Keith E. Shearwin
The University of Adelaide, Adelaide, SA, Australia
James E. Schoelz
University of Missouri, Columbia, MO, United States Hanako Shimura
Hokkaido University, Sapporo, Japan
Jason R. Schrad
Michigan State University, East Lansing, MI, Reina S. Sikkema
United States Erasmus Medical Center, Rotterdam, The Netherlands
 List of Contributors xli

Aaron Simkovich Thomas E. Spencer

Agriculture and Agri-Food Canada, London, ON, University of Missouri, Columbia, MO, United States
Canada and The University of Western Ontario,
Pothur Sreenivasulu
London, ON, Canada
Sri Venkateswara University, Tirupati, India
Peter Simmonds
Ashley L. St. John
University of Oxford, Oxford, United Kingdom
Duke-NUS Medical School, Singapore, Singapore
Tarja Sironen
David K. Stammers
University of Helsinki, Helsinki, Finland
University of Oxford, Oxford, United Kingdom
Susanna Sissonen
John Stanley
Finnish Institute for Health and Welfare, Helsinki,
John Innes Centre, Colney, United Kingdom
Finland
Glyn Stanway
Michael A. Skinner University of Essex, Colchester, United Kingdom
Imperial College London, London, United Kingdom
Thilo Stehle
Douglas E. Smith University of Tuebingen, Tuebingen, Germany and
University of California, San Diego, La Jolla, CA, Vanderbilt University School of Medicine, Nashville, TN,
United States United States
Melvyn Smith Gregory W. Stevenson
Viapath Analytics, Specialist Virology Centre, King’s Iowa State University, Ames, IA, United States
College NHS Foundation Trust, London, United
Kingdom Lucy Rae Stewart
Agricultural Research Service, US Department of
Thomas J. Smith Agriculture, Wooster, OH, United States
The University of Texas Medical Branch, Galveston, TX,
United States C.C.M.M. Stijger
Wageningen University and Research Center, Bleiswijk,
Teemu Smura The Netherlands
Helsinki University Hospital and University of Helsinki,
Helsinki, Finland Peter G. Stockley
University of Leeds, Leeds, United Kingdom
Eric J. Snijder
Leiden University Medical Center, Leiden, David Stone
The Netherlands Weymouth Laboratory, Centre for Environment, Fisheries
and Aquaculture Science, Weymouth, United Kingdom
Gisela Soboll Hussey
Michigan State University, East Lansing, MI, Ashley E Strother
United States The University of Texas Medical Branch, Galveston, TX,
United States
Maria Söderlund-Venermo
University of Helsinki, Helsinki, Finland Sundharraman Subramanian
Michigan State University, East Lansing, MI,
Merike Sõmera United States
Tallinn University of Technology, Tallinn, Estonia
William C. Summers
Eun G. Song Yale University, New Haven, CT, United States
Seoul Women’s University, Seoul, South Korea
Liying Sun
Milan J. Sonneveld Northwest A& F University, Yangling, China
Erasmus University Medical Center, Rotterdam,
Wesley I. Sundquist
The Netherlands
University of Utah, Salt Lake City, UT, United States
Beatriz Soriano
Petri Susi
Biotechvana, Scientific Park University of Valencia and
University of Turku, Turku, Finland
Institute for Integrative Systems Biology (I2SysBio),
University of Valencia–Spanish National Research Curtis A. Suttle
Council, Valencia, Spain University of British Columbia, Vancouver, BC, Canada
xlii List of Contributors

Nobuhiro Suzuki Nicholas M.I. Taylor

Institute of Plant Stress and Resources (IPSR), Okayama University of Copenhagen, Copenhagen, Denmark
University, Kurashiki, Japan
Xu Tengzhi
Lennart Svensson University of California, Davis, CA, United States
Linköping University, Linköping, Sweden and Karolinska
Raquel Tenorio
Institute, Stockholm, Sweden
Cell Structure Laboratory, National Center for
Ronald Swanstrom Biotechnology – Spanish National Research Council
University of North Carolina at Chapel Hill, Chapel (CNB-CSIC), Madrid, Spain
Hill, NC, United States Robert B. Tesh
Daniele M. Swetnam The University of Texas Medical Branch, Galveston, TX,
University of California, Davis, CA, United states United States

Moriah L. Szpara Vaskar Thapa

Pennsylvania State University, University Park, PA, Pennsylvania State University, State College, PA,
United States United States

Keisuke Tabata John E. Thomas

Heidelberg University, Heidelberg, Germany The University of Queensland, Brisbane, QLD, Australia

Anna Taglienti Julie A. Thomas

Council for Agricultural Research and Economics, Rochester Institute of Technology, Rochester, NY,
Research Center for Plant Protection and Certification, United States
Rome, Italy Lynn C. Thomason
Frederick National Laboratory for Cancer Research,
Naoki Takeshita
Frederick, MD, United States
Tokyo University of Agriculture and Technology, Fuchu,
Japan Elizabeth Ashley Thompson
The University of Southern Mississippi, Hattiesburg, MS,
Kana Takeshita Urayama
United States
Tokyo University of Agriculture and Technology, Fuchu,
Japan Jeremy R. Thompson
Cornell University, Ithaca, NY, United States
Michael E. Taliansky
The James Hutton Institute, Dundee, United Kingdom Antonio Tiberini
Council for Agricultural Research and Economics,
Pan Tao Research Center for Plant Protection and Certification,
The Catholic University of America, Washington, DC, Rome, Italy
United States
Peter Tijssen
Jacqueline E. Tate National Institute of Scientific Research – Armand-
Centers for Disease Control and Prevention, Atlanta, Frappier Health Research Centre, Microbiology and
GA, United States Immunology, Laval, QC, Canada
Satyanarayana Tatineni Yuji Tomaru
Agricultural Research Service, US Department of Japan Fisheries Research and Education Agency,
Agriculture, Lincoln, NE, United States and University Kanagawa, Japan
of Nebraska–Lincoln, Lincoln, NE, United States
Laura Tomassoli
Sisko Tauriainen Council for Agricultural Research and Economics,
University of Turku, Turku, Finland Research Center for Plant Protection and Certification,
Rome, Italy
Norbert Tautz
University of Luebeck, Luebeck, Germany Ruben Torres
National Biotechnology Center–Spanish National
Paulo Tavares
Research Council, Madrid, Spain
Institute for Integrative Biology of the Cell, CEA, CNRS,
University of Paris-Sud, University of Paris-Saclay, Jia Q. Truong
Gif-sur-Yvette, France The University of Adelaide, Adelaide, SA, Australia
 List of Contributors xliii

Erkki Truve† James L. Van Etten

Tallinn University of Technology, Tallinn, Estonia University of Nebraska–Lincoln, Lincoln, NE,
Chih-Hsuan Tsai United States
Institute of Molecular Biology, Academia Sinica, Taipei, Suzanne van Meer
Taiwan University Medical Center Utrecht, Utrecht,
Roman Tuma The Netherlands
University of Leeds, Leeds, United Kingdom and Monique M. van Oers
University of South Bohemia, České Budějovice, Wageningen University and Research, Wageningen,
Czech Republic The Netherlands
Topi Turunen Mark J. van Raaij
Infectious Disease Unit, Espoo, Finland and Finnish National Center for Biotechnology, Madrid, Spain
Institute for Health and Welfare, Helsinki, Finland
Marc H.V. Van Regenmortel
Reidun Twarock University of Strasbourg, Strasbourg, France
University of York, York, United Kingdom
Piet A. van Rijn
Ioannis E. Tzanetakis Wageningen Bioveterinary Research, Lelystad, The
University of Arkansas, Fayetteville, United States Netherlands and North-West University, Potchefstroom,
South Africa
Antti Vaheri
University of Helsinki, Helsinki, Finland Alain Vanderplasschen
University of Liège, Liège, Belgium
Eeva J. Vainio
Natural Resources Institute Finland (Luke), Helsinki, Dana L. Vanlandingham
Finland College of Veterinary Medicine, Kansas State University,
Manhattan, KS, United States
Anna M. Vaira
Institute for Sustainable Plant Protection, National Olli Vapalahti
Research Council of Italy, Torino, Italy Helsinki University Hospital and University of Helsinki,
Helsinki, Finland
Steven M. Valles
Center for Medical, Agricultural and Veterinary Mark Varrelmann
Entomology, Agricultural Research Service, US Institute of Sugar Beet Research, Göttingen, Germany
Department of Agriculture, Gainesville, FL,
Nikos Vasilakis
United States
The University of Texas Medical Branch, Galveston, TX,
Adrián Valli United States
National Center for Biotechnology-Spanish National Michael Veit
Research Council, Madrid, Spain Free University of Berlin, Berlin, Germany
Rodrigo A. Valverde Česlovas Venclovas
Louisiana State University Agricultural Center, Baton Vilnius University, Vilnius, Lithuania
Rouge, United States
H. Josef Vetten
Pierre Van Damme Julius Kühn Institute, Braunschweig, Germany
University of Antwerp, Antwerp, Belgium
Marli Vlok
Rene A.A. van der Vlugt University of British Columbia, Vancouver, BC, Canada
Wageningen University and Research Center,
Wageningen, The Netherlands Anne-Nathalie Volkoff
Diversity, Genomes and Insects-Microorganisms
Bernard A.M. Van der Zeijst Interactions, National Institute of Agricultural Research,
Leiden University Medical Center, Leiden, University of Montpellier, Montpellier, France
The Netherlands
Ian E.H. Voorhees
Koenraad Van Doorslaer Cornell University, Ithaca, NY, United States
University of Arizona, Tucson, AZ, United States
Alex Vorsters
†
Deceased. University of Antwerp, Antwerp, Belgium
xliv List of Contributors

Jonathan D.F. Wadsworth Kerstin Wernike

UCL Institute of Prion Diseases, London, United Kingdom Friedrich-Loeffler-Institute, Insel Riems, Germany
Peter J. Walker Rachel J. Whitaker
The University of Queensland, St. Lucia, QLD, Australia University of Illinois at Urbana-Champaign, Urbana, IL,
United States
Paul Wallace
Quality Control for Molecular Diagnostics (QCMD), K. Andrew White
Glasgow, United Kingdom York University, Toronto, ON, Canada
Aiming Wang Anna E. Whitfield
Agriculture and Agri-Food Canada, London, ON, North Carolina State University, Raleigh, NC,
Canada United States
Jen-Ren Wang Richard Whitley
National Cheng Kung University, Tainan, Taiwan University of Alabama at Birmingham, Birmingham,
Lin-Fa Wang AL, United States
Duke-NUS Medical School, Singapore, Singapore Reed B. Wickner
Nan Wang National Institute of Diabetes and Digestive and Kidney
Institute of Biophysics, Chinese Academy of Sciences, Diseases, National Institutes of Health, Bethesda, MD,
Beijing, China United States

Xiangxi Wang Luc Willems

Institute of Biophysics, Chinese Academy of Sciences,
Beijing, China
Xiaofeng Wang
Virginia Tech, Blacksburg, VA, United States
Katherine N. Ward
University College London, London, United Kingdom
Matti Waris
University of Turku, Turku, Finland
Ranjit Warrier
Purdue University, West Lafayette, IN, United States
Daniel Watterson
Australian Infectious Diseases Research Centre, School of
Chemistry and Molecular Biosciences, The University of
Queensland, St Lucia, QLD, Australia
Marta L. Wayne
University of Florida, Gainesville, FL, United States
Friedemann Weber
FB 10 – Institute for Virology, Justus Liebig University
Giessen, Giessen, Germany
Cellular and Molecular Epigenetics (GIGA), Liège,
Belgium and Molecular Biology (TERRA), Gembloux,
Belgium
Brian J. Willett
MRC-University of Glasgow Centre for Virus Research,
Glasgow, United Kindom
Alexis Williams
The University of Texas Medical Branch, Galveston, TX,
United States
Stephen A. Winchester
Frimley Park Hospital, Frimley, United Kingdom and
Immunisation and Countermeasures Division, Public
Health England, London, United Kingdom
Clayton W. Winkler
National Institutes of Health, Hamilton, MT,
United States
Stephan Winter
Leibniz Institute – DSMZ – German Collection of
Microorganisms and Cell Cultures GmbH, Braunschweig,
Germany

Sung-Chan Wei William M. Wintermantel

Institute of Molecular Biology, Academia Sinica, Taipei, Agricultural Research Service, US Department of
Taiwan Agriculture, Salinas, CA, United States

Robin A. Weiss Jennifer Wirth

University College London, London, United Kingdom Montana State University, Bozeman, MT, United States
Tao Weitao Yuri I. Wolf
Southwest Baptist University, Bolivar, MO, National Center for Biotechnology Information, National
United States Library of Medicine, Bethesda, MD, United States
Another random document with
no related content on Scribd:
 Un exemple de la complexité de la moindre sensation : celle du
chaud, du froid. Nous croyons qu’elle est enregistrée par le
thermomètre, par l’unique thermomètre. C’est inexact. Il y a des
jours où le thermomètre est très bas et où nous avons chaud et
réciproquement. Pour recueillir toutes les coordonnées, tous les
éléments de notre impression, il faudrait en même temps employer
le baromètre qui mesure la pression, l’hygromètre qui mesure
l’humidité, l’anémomètre qui mesure la force du vent, le radiomètre
qui mesure celle du soleil. Peut-être d’autres, qui mesureraient la
tension électrique…
Rien n’est simple.

Nous appelons souvent mensonge une des faces de la vérité,

une de celles qui ne nous apparaissent pas.
Pliés à concevoir l’unité du vrai, nous accusons de fausseté tout
ce qui n’est point à cette couleur unique du vrai.

La vérité est si diverse qu’on peut en peindre deux aspects

opposés, également exacts et sincères. Cela dépend du point de
vue où l’on s’est placé.
Il est plus juste encore de représenter la vérité comme une
matière plastique, analogue à la glaise, que chacun pétrit à son
empreinte.
Les avocats exploitent précisément cette plasticité des faits. Ils
s’efforcent de leur donner une figure favorable à la cause qu’ils
défendent. Ce n’est pas de l’artifice. C’est de l’art.

L’idée d’unité fait qu’on s’étonne de voir un même être se

présenter sous des aspects différents. On l’accuse de manquer de
sincérité. Mais nous ne nous étonnons pourtant pas qu’un même
arbre soit, l’hiver et l’été, nu ou couvert de feuilles. L’apparence
change. L’armature reste.
 Habituons-nous à la diversité d’un même être, avant qu’il soit
diminué à nos yeux du fait d’être divers. Tel homme aime une femme
et en parle légèrement. Pourtant, il est sincère, tour à tour, dans la
tendresse et l’ironie.
Un ami ne nous trahit pas parce qu’il médit un tantinet de nous.
Ce sont des états différents. Voilà tout.
Vous protestez ? Vous vous cabrez ? Mais vous-même, vous
savez bien que vous avez été dans ces états différents, sans cesser
d’être vous-même. Examinez-vous. N’avez-vous pas été tour à tour
tendre et dur, cruel et bon, sensible et sec ?
Cette diversité de l’être apparaît dans les témoignages en justice.
L’un dit d’un accusé : il était doux. Un autre : il était violent. Un
troisième : il était franc. Un autre encore : il était fourbe. Témoins à
charge, témoins à décharge trouvent dans sa vie des traits opposés
et qui peuvent être tous vrais. Car nous sommes divers.
Cela déroute notre coutume de penser selon l’unité, de juger les
hommes comme s’ils étaient des statues, d’une seule matière, tout
en bronze, tout en marbre. Et nous ne sommes qu’une mosaïque.

Au point de vue de la diversité de l’être, nous pouvons constater

sur nous-même, dans le détail de la vie, des manifestations d’un
instinct et d’un contre-instinct correspondant.
Exemple : nous sommes sociables, puisque la solitude nous est
mortelle, ainsi que le prouve le régime cellulaire. Et nous souhaitons
farouchement d’être seul en wagon. De même, l’homme a
certainement le goût et le besoin d’une compagne. Par là, il est
monogame. Et en même temps il aspire à la polygamie. Ces
contradictions sont innombrables. En sa diversité, la créature
apparaît contradictoire.
Chacun de nous est un livre dont les feuillets ne se répètent pas.
Nous-même, nous n’en savons pas déchiffrer toutes les pages. Et
nous ne savons même pas d’où vient le souffle qui les fait tourner.
 Dans l’amas des usages, il en est de bons et de mauvais. Eux
aussi sont divers. Mais notre esprit, plié à l’unité, accepte tout. La
crainte et la paresse aidant, il refuse de procéder à l’inventaire, de
conserver les uns et de détruire les autres.
On ne déplorera jamais assez cet esprit d’unité. Tout est
composé, tout est complexe.

Cette idée de diversité doit s’appliquer aussi à la multitude, aux

êtres considérés les uns par rapport aux autres. Il y a longtemps
qu’on a remarqué que les empreintes digitales diffèrent d’un individu
à l’autre. Il faut transporter cette notion dans le domaine moral. Tous
les individus sont aussi différents les uns des autres que leur
empreinte digitale.
La première conséquence, c’est qu’il ne faut pas juger un autre
d’après soi. En lui prêtant nos propres mobiles, nous risquerions fort
de nous tromper. D’autant que lui-même, tombant dans le même
travers, nous prête les siens. C’est la source d’innombrables
malentendus.
Une autre conséquence de cette diversité, c’est de faire
apparaître le défaut de la Loi, telle que nous l’ont léguée les
civilisations antiques. Pénétrée de l’esprit d’unité, elle entend
s’appliquer à tous les hommes, les juger comme s’ils étaient
identiques. Les soumettant au même gabarit, elle opprime trop les
uns, épargne trop les autres.
L’idée de diversité et l’idée de souplesse, dominées elles-mêmes
par la notion du déterminisme, présideront sans doute à la refonte
des Codes.

La vie est précaire.

Deux conceptions de la vie sont en présence. La première est

fondée sur la certitude de vivre très vieux. Celui qui l’adopte dira par
exemple : « Dans dix ans, je ferai cela », quitte à s’évaporer dans dix
jours. Elle a pour principal inconvénient de sacrifier les plus belles
années de la vie à la préparation de la vieillesse, de choisir une
situation médiocre, où l’on rogne sur le superflu, sur le nécessaire,
pendant sa jeunesse et sa maturité, pour avoir droit à cette retraite à
laquelle on ne parviendra peut-être pas…
La seconde conception consiste à prendre conscience de la
précarité de l’avenir, de la fragilité de la vie, de l’imprévu du
lendemain. C’est à elle que va ma préférence. Je la crois plus
humaine. La première ressemble trop à la sérénité animale.
On m’a souvent opposé que le sens de la précarité de l’avenir
décourageait l’effort. C’est une erreur. N’ayant pas la certitude béate
d’une longue vie, on n’en est que plus pressé d’aboutir, de réaliser
sa tâche, de créer l’œuvre qu’on veut laisser après soi. C’est un
stimulant, non pas un anesthésiant.
Et, au point de vue spécial de la retraite, de l’épargne à réserver
pour la vieillesse, cette idée de la fragilité de la vie n’entraîne pas
une frivole insouciance de l’avenir, un gaspillage au jour le jour. Non.
Elle restreint simplement la part énorme que l’on sacrifie à cette
retraite dans la société actuelle, part démesurée, puisqu’on voit des
milliers de gens choisir le métier de fonctionnaire, être malheureux
toute leur vie, pour cette fameuse retraite dont ils ne doutent pas de
jouir, et dont beaucoup ne jouiront pas.

J’ai entendu des gens qui, se plaignant de leurs occupations,

soupiraient : « On oublie de vivre… » Et c’est vrai.
Ne conçoit-on pas une existence plus souple, plus intelligente, où
l’on ait le temps de vivre ? Par exemple, ne pourrait-on pas avoir la
sagesse de se retirer plus tôt, en pleine force, au risque de n’avoir
pas autant d’aisance ? A quoi bon tant de richesse, si on meurt
avant d’en profiter ?
 Des commerçants, des industriels, des grands cultivateurs,
hésitent souvent à saisir l’occasion unique d’un voyage qui
émaillerait leur vie d’un rare souvenir. Ils obéissent plus à la routine
qu’au zèle. Qu’ils tombent gravement malades, ou qu’ils se cassent
une jambe, ils feront ainsi la preuve qu’ils ont pu abandonner pour
un temps leurs affaires sans qu’elles en souffrent sérieusement.

Pour réagir contre cette foi instinctive dans la sécurité de l’avenir,

pour laisser la place du hasard, il existe un moyen qui, si puéril qu’il
paraisse, n’en est pas moins efficace. A propos d’un voyage
prochain, on dit, on écrit, avec une lourde certitude : « Je partirai tel
jour… j’arriverai tel jour… je prendrai le train de telle heure… »
Écrivez ou dites plutôt : « Je compte partir… je me propose
d’arriver… j’ai l’intention de prendre tel train… » On est contraint de
penser les mots qu’on prononce. Par ce simple artifice, on prend le
sens de la précarité de la vie, on fait la part du destin, on s’incline
légèrement devant le dieu inconnu.

Notre vie sera courte ou longue, nous ne le savons pas d’avance.

L’important, c’est qu’elle soit toujours pleine et brillante.
Nous soufflons chacun notre bulle de savon. Nous ne savons pas
quand elle éclatera. Peut-être sera-t-elle encore petite à ce moment-
là. Peut-être sera-t-elle devenue grande. Mais l’important, c’est
qu’elle soit emplie d’un souffle sain, et qu’elle s’irise…

L’inutile tristesse.

Que d’exemples on pourrait donner du fâcheux penchant qui

nous incline à ne voir que le mauvais côté de la vie ! J’en ai déjà cité.
Vieux restes sans doute des terreurs ancestrales, du temps où
l’homme désarmé tremblait devant les monstres et peuplait la nature
de fantômes et de divinités féroces.
 Ainsi le paysan se plaint sans cesse du mauvais temps. Il
constate bien plus rarement le beau temps. D’une façon plus
générale, on constate plus volontiers le mal que le bien. Les
journaux sont le plus frappant exemple de cette tendance. Ils sont
tristes. Ils n’enregistrent que le crime et l’accident. Ils ne donnent
pas un reflet exact de la vie. Cela réagit certainement sur la
mentalité du pays. Car nous sommes des imitateurs-nés.
Pourquoi, dans ces quotidiens, ne pas donner aux événements
heureux, gais, curieux, une place analogue à celle qu’ils tiennent
dans la vie à côté des événements dramatiques ? Faut-il vraiment du
sang pour exciter l’intérêt ? Plus tard, en feuilletant nos journaux, on
s’étonnera de voir que neuf portraits sur dix représentaient des
assassins.

Pourquoi, lorsqu’on a commencé de bâtir des gares, des

casernes, des hôpitaux, des lycées, a-t-on fait « triste ? » Pourquoi
n’avoir pas fait riant ? Il n’en eût guère coûté davantage. Je sais
deux gares charmantes. L’une, où le poste d’aiguillage est pavoisé
de roses. L’autre, où la vigne entoure les piliers de la marquise et fait
à son fronton une frise admirable. L’exemple est à suivre.
Les seuls établissements publics qui soient gracieux, n’étaient
pas destinés à leur emploi actuel : les ministères, installés dans des
palais désaffectés. Et encore, bien vite, l’esprit administratif a su les
enlaidir, à grand renfort de banquettes, de cloisons, de tapis verts,
taches d’encre et garçons de bureau.

Ouvrez un dictionnaire de synonymes. Les mots tristes ont

beaucoup plus d’équivalents que les mots gais.
Autre preuve du triste penchant qu’on pourrait redresser : La
plupart des présages annoncent un événement fâcheux : le sel
répandu, les couverts croisés, treize à table, la glace brisée, etc. Il
n’y en a qu’un qui soit optimiste. « Araignée du soir, espoir ». Encore
semble-t-il avoir été forgé pour faire pendant à : « Araignée du matin,
chagrin ».
En somme, il y a peu de présages de bonheur.

Nous avons un mot qui signifie : dire de quelqu’un du mal

injustifié. C’est médire.
Mais nous n’avons pas de mot qui signifie : dire de quelqu’un du
bien injustifié.

Une preuve encore que nous inclinons vers la tristesse : la

plupart de nos rêves sont tristes. Ce sont des « cauchemars ». Notre
esprit, libre, sans brides, lancé dans le champ infini des
imaginations, se repaît de malheurs.

Nous sommes encore si peu accoutumés au bonheur que nous

ne voulons pas y croire. La nouvelle d’une grande joie nous laisse
d’abord incrédules : « Non ? Ce n’est pas possible ? Ce n’est pas
vrai… » Le premier cri d’allégresse, c’est un cri de doute.

L’harmonie dans la vie.

Nous ne savons pas mettre d’harmonie, d’équilibre, entre nos

dépenses — ou nos épargnes — et leurs effets. Nous ne réglons
pas nos efforts sur leurs résultats. Nous obéissons encore à des
instincts que nous ne soumettons pas à la raison.
Ainsi, pour économiser le prix d’une voiture, une bourgeoise en
tenue de gala risquera, un soir de pluie, de perdre sa robe, ou même
sa vie, si la bronchite s’ensuit.
Et l’exemple du télégramme ? On s’évertue à parler petit nègre,
on compromet la clarté du texte — au risque d’être incompris, de
perdre tout le bénéfice d’une affaire, quelquefois des milliers de
francs — pour économiser un mot, un sou !

Dans une maison où l’argent coule à flot, et presque sans

contrôle, on fera la toute petite économie d’un blanchissage et l’on
assiéra son convive devant une nappe maculée d’une tache
douteuse et gênante…

On voit une femme de condition moyenne passer des heures à

ravauder une paire de chaussettes. Elle passe, à cette besogne
ingrate, un temps démesuré. Car elle essaie de restaurer, de
ressusciter cette ruine qui n’est plus que reprises. Et elle ne réfléchit
pas au prix minime de cet article.
Je crois donc qu’il faudrait mettre les jeunes filles en garde contre
cette tendance, leur apprendre l’usage harmonieux de leur temps et
de leurs occupations.

On peut citer encore, comme exemple de dépense démesurée

d’énergie, celui de l’homme qui — sans raison capitale — veut
attraper le tramway… Il court, s’accroche, se cramponne, flotte en
drapeau au flanc de la voiture. Enfin il se hisse. Il a risqué de se faire
tuer ou tout au moins de se faire broyer les jambes. Et tout cela pour
gagner deux minutes ! N’y a-t-il pas disproportion entre le risque et le
résultat ?… Ah ! si, en des occasions capitales de sa vie, qui le
laissent au contraire indolent et mou, il développait cette sauvage
ardeur !

Le « plan du réel ».

Nous ne manquons pas d’éprouver une déception chaque fois

que nous apprenons une défaillance de notre prochain. A la faveur
d’un scandale, ouvre-t-on l’armoire secrète d’une famille, y
découvre-t-on fatalement quelque linge sale ? Nous révèle-t-on dans
notre entourage quelque faiblesse, quelque turpitude ? Nous en
ressentons chaque fois une surprise déçue. Quelque chose en nous
se décroche, automatiquement. Nous n’apprenons la vie qu’à coups
d’étonnements attristés. Pourquoi ?
Surtout parce que notre éducation nous a peint un monde parfait.
Les livres qui composent la bibliothèque enfantine ne connaissent
pas les fléchissements soudains de la créature. Les personnages
n’en sont point humains. Ce sont de pures marionnettes qui s’agitent
au-dessus de la vie. Puis, au foyer, les parents — ces modèles
immédiats — apparaissent impeccables, dans une sorte de
surhumanité. Le bandeau du respect cache aux petits les fautes des
grands. Bref, nous ne tentons rien pour acheminer prudemment
l’enfant vers l’humaine vérité. Qu’arrive-t-il ? Qu’il place les êtres
dans une atmosphère idéale, sur un plan trop élevé. Pour les
ramener au plan du réel, il doit les faire descendre. Chaque fois,
c’est la chute d’un ange.
Enfin, nous portons peut-être en nous l’image d’un monde
supérieur. Elle illustrerait notre espoir qu’il se réalise. Elle serait une
anticipation. Le sentiment du divin, qui s’agite au fond des âmes,
n’est peut-être lui-même que l’aspiration vers ces temps meilleurs, la
prescience de l’homme futur. Dieu, c’est une prévision…

Ah ! pourquoi a-t-on cultivé en nous cette notion d’une humanité

dès maintenant parfaite ? Pourquoi, aux yeux de l’enfant, a-t-on
placé les individus sur un plan supérieur au réel ?
Acquérir de l’expérience, c’est perdre ensuite ces illusions dont
on prit soin de nous nourrir. Nous employons notre existence à
dépouiller, de désenchantement en désenchantement, cette
ingénuité tenace. Ce doit être une des causes de la tristesse de la
vie.
Si on nous avait peu à peu dévoilé la faiblesse et la misère
pitoyables de la créature, nous ne sentirions pas cette continuelle
« chute des feuilles » dans notre cœur. Nous n’éprouverions pas, par
exemple, un inutile déchirement à découvrir que nos parents eurent
des travers et des défaillances, comme les autres. Cela nous
paraîtrait naturel.
Erreur néfaste, de nous avoir fait vivre d’abord dans un monde si
haut, qu’ensuite la réalité nous apparaisse toujours basse. En
célébrant éperdûment dans nos poèmes et dans nos romans
« l’haleine parfumée » de toutes les amantes, nous avons mis dans
chaque baiser une déception.

« Mais, en montrant aux enfants la vie telle qu’elle est, vous allez
en faire d’affreux petits sceptiques, des pessimistes féroces et
recroquevillés ! » Telle est l’objection qui se dresse contre cette vue.
Elle ne me paraît pas fondée. L’éducation au foyer permet le tact, la
mesure, la prudence, l’art patient des préparations. Par elle,
l’adolescent ne sera-t-il pas mieux initié aux réalités que par le choc
brutal de ces réalités mêmes ? Et puis, à tout prendre, si l’existence
doit lui réserver des étonnements, ne doit-on pas préférer, à la triste
surprise des désillusions, la surprise heureuse de découvrir ce qu’il y
a — malgré tout — de joli, d’élégant dans la vie ?

Il y a peut-être de l’égoïsme aussi chez les parents qui veulent

garder à leurs enfants toute la fleur de l’ignorance pour s’en donner
l’agréable spectacle. Ainsi, je ne suis pas bien sûr que « Noël » soit
une si charmante fiction. Est-ce bien pour la joie des enfants que
nous la conservons ? N’est-ce pas pour le plaisir que nous donne la
vue de leur fraîche surprise ? Qui donc a mis en balance l’allégresse
de l’enfant le soir de Noël, avec sa secrète déception le jour où il
apprend qu’il n’y a pas de Noël ?

Il n’y a guère de collaborateurs, même de ceux dont nous

associons, dont nous soudons les noms, comme Erckmann-
Chatrian, qui ne se soient, à un moment de leur vie, brouillés et
combattus. Chaque fois que nous apprenons une de ces haines
intimes, c’est en nous une déception, un décrochement d’illusion.
Pourquoi ? Parce que nous portons en nous, par hérédité, par
éducation, peut-être par instinct du futur, cette notion des affections
fraternelles, parfaites, immuables. La vie se charge de la détruire. Si
nous n’avions pas cette idée préconçue, et cultivée en nous, toute
union de ce genre serait à nos yeux l’heureuse surprise.

S’il était bien entendu, bien admis, que la discrétion humaine est
toujours relative, qu’elle n’est jamais absolument étanche,
imperméable, on n’aurait pas de déception chaque fois qu’on
apprend une indiscrétion.
Et, de plus, on ne confierait à personne ce qu’on ne veut
absolument pas divulguer.

Physiquement, on n’est pas toujours en beauté ; quand on bâille,

on est laid. Eh bien, de même, on ne peut pas toujours être beau
moralement. On a ses moments d’abandon, ses bâillements.
Prenons-en bravement notre parti. Et ne soyons pas déçu de ces
fléchissements de notre prochain.

Quand nous passons à la réalisation de nos projets, il se produit

toujours une sorte de retrait analogue à celui du métal quand il se
refroidit dans son moule. Nous pourrions, sans verser dans le
pessimisme ni le scepticisme, nous éviter ces désillusions. Nous
pourrions prévoir ce retrait, comme on prévoit celui du métal.
 CHAPITRE IV
VUES MORALES

Les mains propres. — Le déterminisme. — La loi d’équilibre. — Le contrôle. — La

bravoure. — La mort. — La religion.

Les mains propres.

De longs siècles nous ont donné le goût de nous laver. Ce goût

est devenu presque un instinct, une nécessité. Nous souffrons de ne
pas nous sentir nets, physiquement, et non seulement sur les parties
visibles de nous-mêmes, mais sur notre corps tout entier. Ce goût de
la propreté se répand. Il pénètre dans des classes de plus en plus
modestes et de plus en plus nombreuses. Il descend la pyramide
sociale. Car on peut s’imaginer la société comme une pyramide : au
sommet, le tout petit groupe des privilégiés ; puis des couches qui
vont en s’élargissant au fur et à mesure qu’on s’éloigne plus de la
richesse et du pouvoir. Eh bien, si la propreté est devenue un
instinct, comment affirmer que la probité n’en deviendra pas un
également, par un travail analogue ? Nous n’étions pas plus
contraints d’être propres que nous ne le sommes d’être honnêtes.
Pour qui établit une continuité absolue entre l’être physique et l’être
moral, les deux besoins se confondent.
On éprouve la même satisfaction et on retire le même avantage à
être propre physiquement et à être propre moralement. Les taches
gênent autant sur la conscience et sur les mains.
D’ailleurs, toutes les notions nouvelles sur la solidarité, sur
l’interdépendance des êtres, sur la mutuelle répercussion de leurs
actes, tendent à démontrer qu’on a encore plus d’intérêt à être probe
qu’à être propre.

L’argot, qui élabore la langue de demain, avait prévu cette fusion

du physique et du moral, cette étroite analogie de la probité et de la
propreté. Pour une mauvaise action, un acte vil, il dit : une saleté,
une crasse.

Quand on n’établit pas de différence entre ce qu’on appelle le

physique et ce qu’on appelle le moral, entre le corps et l’âme, quand
on les bloque en un tout, quand on les confond aussi étroitement
que le cierge et sa flamme, cette vue des « mains propres » apparaît
toute naturelle. Il semble normal que l’individu, par l’effet continu
d’une lente culture, après avoir éprouvé le besoin d’être propre
physiquement, éprouve le besoin d’être propre moralement.

Et peut-être même une nouvelle étape sera-t-elle franchie.

Quand la propreté s’affine, elle devient l’élégance. Après qu’on s’est
lavé, on aspire à se parer. Ce sont les phases successives de la
toilette. Peut-être la propreté morale aura-t-elle aussi son luxe, son
élégance, se parera-t-elle de ces vertus nobles et charmantes qui
sont la grâce de la créature.

Le déterminisme.

Être déterministe c’est, surtout, croire que nos actes, que nos
paroles, sont déterminés par des influences qui s’exercent sur nous,
des réactions qui se développent en nous, mais dont nous ne
sommes pas maîtres, pas plus que nous ne sommes maîtres des
phénomènes de notre vie physique. Toutes ces forces se combinent,
se composent, aboutissent à une résultante, qui est notre acte ou
notre parole. Notre conscience enregistre cette délibération, mais ne
la dirige pas. Si j’analyse le plus simple de mes gestes, saisir un
objet, marcher vers un but, je vois que j’obéis à un ensemble de
sollicitations que je n’ai pas provoquées spontanément.
Au moment où nous croyons prendre librement une résolution,
toutes ces voix intérieures ont déjà délibéré, conclu à cette décision.
Nous n’avons que l’illusion de la volonté. La sphère mentale est un
véritable parlement, où les instincts se groupent, délibèrent,
expriment enfin, par un vote décisif, la volonté de l’organisme tout
entier, comme le vote parlementaire exprime la volonté nationale.
Même dans notre langue usuelle, les mots trahissent ce travail
intérieur : on pèse le pour et le contre, on balance, on se résout, on
se détermine.

C’est une erreur de croire que la formule : « Ce n’est pas sa

faute » ne puisse pas servir de règle de vie, ne puisse pas être la
base d’une morale. Penser fortement que ce n’est pas plus la faute
d’un homme s’il est irascible que s’il est phtisique, regarder les vices,
les travers, enfin toutes les tares de l’esprit, du même œil que les
tares du corps, les accueillir avec la même pitié, comment ne sent-
on pas ce qu’une telle conception entraîne fatalement de
mansuétude, de patience, et comme elle est vraiment large et
charitable ?

Si nos instincts nous dirigent, c’est à nous à améliorer les bons, à

réduire les mauvais. Il y a là tout un entraînement, toute une
orthopédie à pratiquer. De même que nous fortifions un buste
malingre par l’exercice, nous pouvons développer un instinct faible
par l’éducation. Quelle vue féconde ! Traiter les maladies morales
comme les maladies physiques, cela signifie aussi les soigner,
remonter à leurs causes — ce qu’on n’a jamais fait — essayer de les
guérir. Mais qu’on y fasse bien attention. Cela signifie aussi se
défendre contre elles. Où l’on dit coupable, il faut dire nuisible. C’est
la même sauvegarde, avec plus d’indulgence et sans l’idée sombre
de vindicte. Évidemment l’heure n’est pas venue où une telle
maxime : « Ce n’est pas sa faute », inspirera les lois. Mais dès
maintenant, pour qui en est fermement convaincu, saturé, elle est,
en toutes circonstances, devant soi et devant les autres, à tout
instant, le guide sûr et juste…

Les deux grands mots profonds des enfants sont : « Ce n’est pas
ma faute » et : « Je ne l’ai pas fait exprès ». Ils ont bien raison. C’est
tout le fond du déterminisme. Mais s’ensuit-il qu’il n’y ait pas de suite
à donner à leurs petits délits ? Et que cela les absolve et leur
permette de recommencer ? Cent mille fois non ! Voilà l’erreur des
gens qui prétendent que nous sommes libres de faire le bien et le
mal et qui accusent les doctrines déterministes d’être dissolvantes.
Pas du tout. Quand un enfant attrape la rougeole, ce n’est pas sa
faute. Il ne l’a pas fait exprès. On le soigne, pourtant. Eh bien, de
même quand il a commis une faute. On s’efforce d’en découvrir les
causes, d’en montrer les inconvénients, d’en éviter le retour.

On peut agir sur ses sentiments dans la mesure où l’on peut agir
sur son organisme. Il semble qu’on puisse soigner une crise morale
comme on soigne une maladie, en puisant dans le désir de guérir
l’énergie nécessaire à la cure. Les remèdes sont analogues dans les
deux cas. Les révulsifs, les dérivatifs, deviennent, dans la maladie
morale, les distractions qui détournent l’esprit de sa hantise et le
portent vers d’autres objets. Il y a des anesthésiants qui endorment
la sensibilité douloureuse, par exemple une saine fatigue, un labeur
acharné. Il y a l’opération, qui coupe court, comme l’absence, le long
voyage, le silence. Il y a l’homéopathie, qui combat le mal par le mal,
l’amour par l’amour… Il y a enfin la recherche des causes, qui
démontre parfois l’origine toute fortuite, la nature fragile de la crise et
qui permet de l’atteindre à sa source.

Entre la théorie du déterminisme et celle qui nous suppose libres

de choisir entre le bien et le mal, il n’y a au fond qu’une question de
mots. Se croire de la volonté, ou savoir qu’on n’a que l’illusion de la
volonté, cela revient au même. Ce sont deux façons différentes de
voir la nature humaine.
Mais le déterminisme est plus indulgent. Et il est aussi plus
fécond, parce qu’il indique les moyens directs de se soigner, de
s’amender, de s’améliorer.
Il faut s’élever violemment contre la confusion du déterminisme et
du fatalisme. Le fataliste s’abandonne à sa destinée. Tandis que le
déterministe envisage les moyens de se mettre en garde contre lui-
même, de trouver des remèdes à ses défaillances.
Si l’on conçoit chaque décision sous la forme simpliste d’une
pesée, où des poids jetés dans les deux plateaux de la balance
finissent par la faire osciller et se fixer dans un certain sens, on sent
bien qu’on pourra faire intervenir de nouveaux poids pour la faire
pencher dans le meilleur sens. Se sent-on tenté par une mauvaise
action, sollicité par un vice ? On appelle à son secours des aides
nouvelles. On se représente les dangers, les inconvénients que l’on
va risquer.

Si l’on a admis que le travail de nos décisions ressemble à celui

d’une opération où l’on met des poids dans les deux plateaux de la
balance, il y aura intérêt à se livrer consciemment à cette besogne
de détermination, à porter en deux colonnes les raisons pour et les
raisons contre. On pèsera mieux.

Il faut considérer les peines du code comme des contre-poids. La

crainte qu’elles inspirent est un poids que l’on jette dans la balance
de nos décisions.

A la lecture des faits-divers, on a conscience qu’un grand nombre

de drames auraient pu être évités si, au préalable, on avait soigné
mentalement le meurtrier. Car souvent on voit qu’il était en
puissance de crime. Mais on ne soigne pas le cerveau, parce que
c’est l’âme, l’intangible, le différent du corps.

Je crois que les hommes ne se disent pas d’avance : je vais être

noir, je vais être fatal. Ils se laissent entraîner. Ils glissent au mal et
au crime.

Pour se déterminer, il faut chercher la solution de moindre

inconvénient. On ne peut pas toujours éviter de faire de la peine. Il
faut faire la moindre peine.

Je suis très persuadé que les mots exercent une influence

mécanique sur la pensée. J’entends par là que l’on contraint sa
pensée à couler dans le moule des mots qu’on prononce. Elle prend
leur empreinte. On pense comme on parle. Ainsi, en déterminisme,
le simple artifice qui consiste à dire : « C’est de son fait », au lieu de :
« C’est de sa faute », modifie la mentalité. L’emploi même de la
formule efface de l’esprit l’idée de blâme, de responsabilité au sens
religieux, d’expiation, la remplace par une constatation apaisée,
sereine, où déjà se lève l’indulgence.

La loi d’équilibre.

La loi d’équilibre qui semble régir l’univers et suspendre les

astres dans l’espace, cette même loi doit gouverner le détail de notre
vie. Elle veut qu’à toute action corresponde une réaction égale. On
ne dira plus : tout se paye. Mais peut-être dira-t-on : tout se
compense. Il faut seulement que cette loi ait le temps de jouer.
Elle ne connaît pas d’exception. Quand elle semble en défaut,
nous pouvons nous en prendre à nos sens ou à notre savoir bornés.
L’exemple est classique de Leverrier découvrant par le calcul
l’existence de Neptune. La loi d’équilibre exigeait, pour être
respectée, que cette planète fût.
Ainsi, tout se balance, le positif et le négatif, le beau et le laid, le
bon et le mauvais. Tous les contraires se neutralisent. Toutes les
forces opposées s’annihilent dans un équilibre final. Et la notion, en
somme, est rassurante, de cette harmonie faite d’antagonisme, de
ce néant fécond.

La loi d’équilibre nous ouvre des vues consolantes. Car elle veut
que le bien et le mal se compensent. Ainsi, dans le monde
innombrable des plantes, il doit y en avoir autant de bienfaisantes
que de malfaisantes. Je crois qu’on n’a pas arraché aux simples
tous leurs secrets. Les hommes ont découvert des poisons végétaux
qui, en quelques secondes, amènent l’organisme de la santé à la
mort. La loi d’équilibre permet de prévoir qu’on découvrira, en
opposition avec ces toxiques, des toniques qui amèneront
instantanément à la santé un organisme au seuil de la mort. Non pas
des contre-poisons, mais de puissants révulsifs qui ressusciteront
l’individu près de succomber à un choc, un traumatisme, une
asphyxie.
De même, dans une autre direction, on peut prévoir que
l’équilibre se rétablira entre les villes et les campagnes. Avant qu’un
pendule ne prenne la verticale, il oscille à droite, puis à gauche. Ce
qui manque à une vie humaine, c’est le temps de voir le pendule
prendre l’équilibre. On n’assiste qu’à une oscillation.

Tout se compense, à la condition que cet équilibre ait le temps de

se réaliser. C’est bien en effet un équilibre qui s’établit entre le bon
et le mauvais, au cours d’une existence.
Supposez un joueur de roulette qui jouerait un nombre de coups
infini… Les mathématiques prouvent que ses gains et ses pertes se
seraient équilibrés.
 Je crois que le mal et le bien se compensent dans une existence,
car le total d’une vie, comme le résultat d’une longue opération de
jeu, doit tendre vers zéro, vers l’équilibre qui régit tout.
Seulement, nous avons dans l’esprit l’idée d’expiation qui vient
de la terreur inspirée aux premiers hommes par le mystérieux
univers qui les entourait. Nous voulons que le mal qu’on endure
expie le mal qu’on a fait.

Il doit y avoir en nous un sens du juste et de l’injuste. Quelque

chose nous avertit, par exemple, qu’il y a de l’injustice dans le cas
du fermier, obligé de travailler pour verser de l’argent à un fils de
famille qui n’a fait qu’hériter, dans le cas de l’ouvrier qui trime avec
les mêmes gestes pendant cinquante ans, pour enrichir un patron
qui se donne tous les plaisirs de l’oisiveté.
Mais nous devons cultiver, développer ce sens de la justice.
C’est un des aspects de cette loi d’équilibre qui régit les mondes.
Rien d’étonnant à ce que ce sens ne soit pas renforcé en nous. Car
c’est la vérité scientifique, en découvrant par exemple la loi de la
gravitation, qui a dévoilé cette notion de l’équilibre universel.
Découverte relativement récente, notion encore neuve. C’est à nous
à la dégager, à la faire resplendir en nous.

Le Contrôle.

Il est utile de se rendre compte du rôle en chacun de nous de

l’inconscient.
C’est une zone du cerveau que l’état de nos connaissances ne
nous permet pas encore de circonscrire, où s’élaborent nos pensées
avant de parvenir aux régions où nous en prenons conscience.
Antre obscur, où se cuisinent, où s’agitent nos résolutions avant
d’émerger au jour.
Il est bon d’imaginer l’existence de cette zone où certaines de
nos pensées naissent, flottent, s’ébauchent. Peut-être est-ce la
partie du cerveau la plus ancienne, celle qui ressemble le plus à
celle des autres animaux, celle de l’instinct, tandis que les régions,
où s’exerce notre contrôle, sont des conquêtes récentes, le propre
de l’homme, le signe du progrès et du perfectionnement.
Parfois, bien que tout ce travail ait lieu à notre insu, cette
ébauche de notre décision est très poussée. Elle jaillit, elle émerge
toute armée, toute prête, dans la lumière de notre conscience. A
peine avons-nous le temps de l’examiner. Déjà elle agit, déjà elle
commande nos gestes. C’est l’impulsion, avec tous ses charmes et
tous ses dangers.
Donc, pensons à tout ce travail caché, secret pour nous-mêmes,
toute cette préparation qui parfois va nous surprendre.
Quand on a le sentiment de cet inconscient, on s’explique bien
des décisions subites, qui se sont élaborées dans l’antre, dans
l’ombre de nous-mêmes…

A l’inverse, il arrive fréquemment que des paroles, des images,

bref le butin de nos sens, tombent dans notre inconscient sans que
nous les percevions au passage, sans que nous ayons eu le temps
de les enregistrer, un peu comme un visiteur se glisse dans
l’immeuble sans être vu du concierge.

Grâce aux journaux, nous nous soucions de toutes sortes de faits

que nous eussions ignorés sans cette lecture. A propos
d’événements qui ne nous touchent pourtant pas directement, nous
nous créons des petites joies, des petits chagrins, des triomphes et
des défaites. Nous savons ce qui se passe dans l’univers et dans
l’esprit des autres. Nous participons plus que nos ancêtres à la vie
générale. L’information nous arrive en ondes rapides. Et notre
conscience du monde s’en trouve prolongée.
Ainsi elle s’étend en surface. De même qu’elle doit,
physiologiquement, gagner en profondeur. Au fur et à mesure que
des êtres se civilisent, le domaine de la conscience doit conquérir