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Herpes
Simplex
Viruses
INFECTIOUS DISEASE AND THERAPY
Series Editor
Burke A. Cunha
Winthrop-University Hospital
Mineola, and
State University o f New York School o f Medicine
Stony Brook, New York
1. Parasitic Infections in the Compromised Host,

edited by Peter D. Walzer and Robert M. Genta
2. Nucleic Acid and Monoclonal Antibody Probes:
Applications in Diagnostic Methodology, edited by
Bala Swaminathan and Gyan Prakash
3. Opportunistic Infections in Patients with the Acquired
Immunodeficiency Syndrome, edited by
Gifford Leoung and John Mills
4. Acyclovir Therapy for Herpesvirus Infections, edited by
David A. Baker
5. The New Generation of Quinolones, edited by
Clifford Siporin, Carl L. Heifetz, and John M. Domagala
6. Methicillin-Resistant Staphylococcus aureus: Clinical
Management and Laboratory Aspects, edited by
Mary T. Cafferkey
7. Hepatitis B Vaccines in Clinical Practice, edited by
Ronald W. Ellis
8. The New Macrolides, Azalides, and Streptogramins:
Pharmacology and Clinical Applications, edited by
Harold C. Neu, Lowell S. Young, and Stephen H. Zinner
9. Antimicrobial Therapy in the Elderly Patient, edited by
Thomas T. Yoshikawa and Dean C. Norman
10. Viral Infections of the Gastrointestinal Tract:
Second Edition, Revised and Expanded, edited by
Albert Z. Kapikian
11. Development and Clinical Uses of Haemophilus b
Conjugate Vaccines, edited by Ronald W. Ellis
and Dan M. Granoff
12. Pseudomonas aeruginosa Infections and Treatment,
edited by Aldona L. Baltch and Raymond P. Smith
13. Herpesvirus Infections, edited by Ronald Glaser
and James F. Jones
14. Chronic Fatigue Syndrome, edited by
Stephen E. Straus
15. Immunotherapy of Infections, edited by K. Noel Masihi
16. Diagnosis and Management of Bone Infections,
edited by Luis E. Jauregui
17. Drug Transport in Antimicrobial and Anticancer
Chemotherapy, edited by Nafsika H. Georgopapadakou
18. New Macrolides, Azalides, and Streptogramins in
Clinical Practice, edited by Harold C. Neu,
Lowell S. Young, Stephen H. Zinner,
and Jacques F. Acar
19. Novel Therapeutic Strategies in the Treatment of
Sepsis, edited by David C. Morrison and John L. Ryan
20. Catheter-Related Infections, edited by Harald Seifert,
Bernd Jansen, and Barry M. Farr
21. Expanding Indications for the New Macrolides,
Azalides, and Streptogramins, edited by
Stephen H. Zinner, Lowell S. Young, Jacques F. Acar,
and Harold C. Neu
22. Infectious Diseases in Critical Care Medicine,
edited by Burke A. Cunha
23. New Considerations for Macrolides, Azalides,
Streptogramins, and Ketolides, edited by
Stephen H. Zinner, Lowell S. Young, Jacques F. Acar,
and Carmen Ortiz-Neu
24. Tickborne Infectious Diseases: Diagnosis
and Management, edited by Burke A. Cunha
25. Protease Inhibitors in AIDS Therapy, edited by
Richard C. Ogden and Charles W. Flexner
26. Laboratory Diagnosis of Bacterial Infections,
edited by Nevio Cimolai
27. Chemokine Receptors and AIDS, edited by
Thomas R. O'Brien
28. Antimicrobial Pharmacodynamics in Theory
and Clinical Practice, edited by Charles H. Nightingale,
Takeo Murakawa, and Paul G. Ambrose
29. Pediatric Anaerobic Infections: Diagnosis and
Management, Third Edition, Revised and Expanded,
Itzhak Brook
30. Viral Infections and Treatment, edited by
Helga Ruebsamen-Waigmann, Karl Deres, Guy Hewlett,
and Reinhold Welker
31. Community-Aquired Respiratory Infections, edited by
Charles H. Nightingale, Paul G. Ambrose,
and Thomas M. File
32. Catheter-Related Infections: Second Edition,
Harald Seifert, Bernd Jansen and Barry Farr
33. Antibiotic Optimization: Concepts and Strategies in
Clinical Practice (PBK), edited by Robert C. Owens, Jr.,
Charles H. Nightingale and Paul G. Ambrose
34. Fungal Infections in the Immunocompromised Patient,
edited by John R. Wingard and Elias J. Anaissie
35. Sinusitis: From Microbiology To Management,
edited by Itzhak Brook
36. Herpes Simplex Viruses, edited by Marie Studahl,
Paola Cinque and Tomas Bergström
37. Antiviral Agents, Vaccines, and Immunotherapies,
Stephen K. Tyring
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First published 2006 by CRC Press
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Preface
Ubiquitous viruses among humans, herpes simplex viruses (HSV) are

believed to be several million years old. Infection by these viruses may cause
various conditions ranging from mucocutaneous disease to more severe
afflictions of the central nervous system (CNS) or life-threatening infections
in immunocompromised patients. Physicians from almost all medical spe-
cialties will encounter patients with HSV infections that may require treat
ment. Recently acquired knowledge of pathogenesis and diagnostics in
already described clinical diseases caused by HSV, as well as identification
of new syndromes or diseases where herpes simplex may or may not play
a causal role, may be unknown to a large proportion of medical personnel.
This book on HSV not only summarizes the clinical aspects of HSV
infections, but also throws light on basic molecular virology to facilitate a
better understanding of different disease manifestations. The purpose of this
volume is to compile diagnostics and management of patients with HSV
infections with basic virology relevant to an understanding of pathogenetic
mechanisms. This book will be a good aid to clinicians treating patients with
HSV infections, as well as diagnostic microbiologists who will find suitable
and updated information for their respective professions.
The first chapter of Section I deals with the evolution of the viruses
followed by an update on the natural course of primary and recurrent infec
tions as well as epidemiology. The next chapter concerns pathogenesis in
different experimental and clinical manifestations of HSV infections. The
following chapter on diagnostics demonstrates the improvement in this
field that has occurred over the last decennium, mainly due to the introduc
tion of genome detection by PCR, and to the development of new and type-
discriminating serological methods. The second, more clinical, part of the
book begins with a chapter on treatment of HSV infections utlilizing estab
lished as well as new drugs where clinical trials are ongoing. The subsequent
/il
iV Preface
chapters document the different disease manifestations of HSV infections,

including their prognosis and treatment.
In separate chapters, well-known diseases such as gingivostomatitis,
recurrent labial infection, HSV infections of the skin, acute and recurrent
genital infection, and ocular infection are described. CNS diseases such as
herpes simplex encephalitis and meningo/radiculo/myelitis, as well as dis
eases recently discovered to be caused by herpes simplex such as paresis
of the facial nerve are covered. HSV infections in immunocompromised
patients and in pregnant women including congenital and neonatal infec
tions are specially addressed in later parts of the book. Lastly, recent
advances in prevention of infection and disease by vaccination and future
outlooks in HSV research are discussed.
Marie Studahl
Paola Cinque
Tomas Bergström
Contents
Preface . . . . Hi
Contributors . . . . xi
SECTION I. UNDERSTANDING AND DIAGNOSING HERPES

SIMPLEX VIRUS
1. Evolution of Herpes Simplex Viruses 1

Rory J. Bowden and Duncan J. McGeoch
Introduction . . . . 1
Origins of the Family Herpesviridae . . . . 2
Relationships Within the Subfamily
Alphaherpesvirinae . . . . 4
Evolution of Alphaherpesvirus Genome
Structures and Gene Sets . . . . 8
Classes and Mechanisms of HSV Genomic Variation . . . . 11
Studies of HSV Variability Using Restriction
Sites and DNA Sequences . . . . 16
Studies of HSV-1 Population Relationships
and Origins . . . . 20
Prospects . . . . 25
References . . . . 26
V
vi Contents
2. Herpes Simplex Virus Vaccines and the Viral Strategies

Used to Evade Host Immunity ............................................ 35
Lauren M. Hook and Harvey M. Friedman
Vaccines for Prevention or Treatment of
Herpes Simplex Virus (HSV) . . . . 35
Novel Directions in HSV Vaccine Design . . . . 41
Conclusions . . . . 48
3. The Natural History and Epidemiology

of Herpes Simplex Viruses ................................................... 55
André J. Nahmias, Francis K. Lee, and Susanne Beckman-Nahmias
Phase I—The Coevolution (or EVO-EPI) Phase . . . . 57
Phase II—The Infrastructure Phase . . . . 59
Phase III—The Modern Phase . . . . 60
Viewing the Natural History and Epidemiology
of HSV-1 and HSV-2 in Context of the Major
Recent Changes in the World . . . . 65
Epidemiology of HSV-1 Infection . . . . 66
Epidemiology of Herpes Simplex Virus Type 2 . . . . 72
HSV Interactions with Other Sexually
Transmitted Infections, Particularly HIV . . . . 78
Challenges for Research and Public
Health Policies During Phase IV . . . . 80
4. Pathogenesis.......................................................................... 99
Tomas Bergström
HSV Structure and Replication . . . . 100
Natural Infection . . . . 103
Genetic Susceptibility of the Host . . . . 107
HSV Virulence . . . . 108
References . . . . I ll
5. Understanding and Diagnosing Herpes Simplex Virus . . . 119
Eva Thomas
Contents v/7
Detection of Viral Genomes, Whole Virus,

and Viral Antigens . . . . 121
Detection of Antiviral Antibodies . . . . 130
Laboratory Diagnosis of Specific HSV Infections . . . . 133
SECTION II. DISEASE MANIFESTATIONS OF HSV

AND TREATMENT
6. Antiviral Treatment............................................................. 153

Gerald Kleymann
Historical Aspects of Antiviral Therapy . . . . 153
Treatment of Herpes Simplex Virus (HSV)
Infections . . . . 154
Mechanism of Action . . . . 166
Toxicity of Chemotherapy . . . . 167
Resistance . . . . 169
Drug Discovery . . . . 171
7. Primary Herpes Simplex Gingivostomatitis and Recurrent

Orolabial Infection ............................................................. 177
Jacob Amir
Pathogenesis . . . . 177
Epidemiology . . . . 178
Transmission and Virus Shedding . . . . 179
Clinical Manifestations . . . . 180
Complications . . . . 181
Diagnosis . . . . 182
Therapy . . . . 183
8. Herpesvirus Infections of the Skin ...................................... 191

Karan K. Sra, Gisela Torres, and Stephen K. Tyring
Herpes Whitlow . . . . 191
Herpes Gladiatorum and HSV Folliculitis . . . . 192
Eczema Herpeticum . . . . 193
Erythema Multiforme (EM) . . . . 194
V III Contents
Localized Cutaneous HSV . . . . 195

Disseminated Cutaneous HSV . . . . 195
Post-Operative HSV-1 Reactivation . . . . 196
9. Acute and Recurrent Genital Herpes Simplex

Virus Infection ................................................................... 203
George Kinghorn
Natural History . . . . 204
Clinical Features . . . . 210
Diagnosis . . . . 215
Management . . . . 217
10. Herpes Simplex Viruses Ocular D isease............................ 239

Thomas J. Liesegang
Pathophysiology . . . . 241
Ocular Disease Manifestations . . . . 248
Diagnostics . . . . 259
Treatment . . . . 260
Concluding Comments . . . . 265
11. Herpes Simplex Encephalitis and Other Neurological Syndromes

Caused by Herpes Simplex Virus-1 ................................... 275
Marie Studahl and Birgit Sköldenberg
Clinical Disease . . . . 277
Diagnostic Strategies . . . . 282
Pathogenesis . . . . 299
Conclusion . . . . 305
Contents ix
12. Neurological Disease in Herpes Simplex Virus Type 2

(HSV-2) Infection ............................................................... 317
Elisabeth Aurelius
Aseptic Meningitis . . . . 318
Associated Mucocutaneous Lesions . . . . 321
Associated Neurological Complications . . . . 321
Recurrent Herpetic Disease . . . . 322
Primary Meningitis . . . . 326
Recurrent Meningitis . . . . 327
Meningitis with Associated Neurological
Symptoms . . . . 328
Radiculomyelopathy . . . . 328
Neuritis . . . . 329
Myelitis . . . . 329
Encephalitis, Brainstem Encephalitis . . . . 331
13. Herpes Simplex Virus and Bell’s Palsy ............................ 339

Yasushi Furuta
Anatomy of the Facial Nerve . . . . 340
General Aspects of Bell’s Palsy . . . . 341
Reactivation of HSV as a Cause of Bell’s Palsy
(Hypothesis) . . . . 341
Primary HSV Infection and Facial Paralysis . . . . 344
Animal Models of Facial Paralysis by
HSV Infection . . . . 344
Latency of HSV in the Geniculate Ganglia . . . . 346
Virus Isolation in Patients with Bell’s Palsy . . . . 346
Serological Study of Herpes Virus Infections . . . . 348
Detection of HSV Genomes in Patients
with Bell’s Palsy . . . . 350
Mechanism by which HSV Causes Facial
Paralysis . . . . 352
Conflicting Issues Against HSV Etiology
in Bell’s Palsy . . . . 354
Treatment of Bell’s Palsy . . . . 355
Summary . . . . 356
X Contents
14. Herpes Simplex Virus Infections in Immunocompromised

Patients ............................................................................... 363
Fulvio Crippa and Paola Cinque
HSV Infections in Transplanted Patients . . . . 364
HSV Infections in HIV Infected Patients . . . . 370
15. Neonatal Herpes Simplex Virus Infection.......................... 395

Kevin S. Buckley, David W. Kimberlin, and Richard J. Whitley
Acquisition of Intrauterine and Neonatal Infections . . . . 397
Clinical Manifestations . . . . 399
Virological Diagnosis . . . . 404
16. Future Outlooks ................................................................. 411

Tomas Bergstrom, Paola Cinque,, and Marie Studahl
Index . . . . 415
About the Editors . . . . 421
Contributors
Jacob Amir Department of Pediatric C, Schneider Children’s Medical

Center of Israel, Petah Tikva and Sackler School of Medicine, Tel Aviv
University, Tel Aviv, Israel
Elisabeth Aurelius Karolinska Intistute, Unit of Infectious Diseases,

Karolinska University Hospital, Stockholm, Sweden
Susanne Beckman-Nahmias Emory University, Atlanta, Georgia, U.S.A.
Tomas Bergström Department of Clinical Virology, Göteborg University,

Göteborg, Sweden
Rory J. Bowden MRC Virology Unit, Institute of Virology,

Glasgow, U.K.
Kevin S. Buckley Division of Pediatric Infectious Diseases, University of

Alabama at Birmingham, Birmingham, Alabama, U.S.A.
Paola Cinque Clinic of Infectious Diseases, Scientific Institute San

Raffaele, Milan, Italy
Fulvio Crippa Clinic of Infectious Diseases, Scientific Institute San

Raffaele, Milan, Italy
Harvey M. Friedman Infectious Disease Division, Department of

Medicine, University of Pennsylvania School of Medicine, Philadelphia,
Pennsylvania, U.S.A.
xi
xii Contributors
Yasushi Furuta Department of Otolaryngology-Head & Neck Surgery,

Hokkaido University Graduate School of Medicine, Kita-ku, Sapporo,
Japan
Lauren M. Hook Infectious Disease Division, Department of Medicine,

University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania,
U.S.A.
David W. Kimberlin Division of Pediatric Infectious Diseases, University

of Alabama at Birmingham, Birmingham, Alabama, U.S.A.
George Kinghorn Department of Genitourinary Medicine, Royal

Hallamshire Hospital, Sheffield Teaching Hospitals NHS Foundation
Trust, Sheffield, U.K.
Gerald Kleymann Department of Chemistry and Pharmacy,

Interfakultäres Institut Für Biochemie, Tübingen, Germany
Francis K. Lee Emory University, Atlanta, Georgia, U.S.A.
Thomas J. Liesegang Department of Ophthalmology, Mayo Clinic,

Jacksonville, Florida, U.S.A.
Duncan J. McGeoch MRC Virology Unit, Institute of Virology,

Glasgow, U.K.
André J. Nahmias Emory University, Atlanta, Georgia, U.S.A.
Birgit Sköldenberg Division of Medicine, Unit of Infectious Diseases,

Karolinska Institute, Karolinska University Hospital, Stockholm, Sweden
Karan K. Sra Department of Dermatology, Center for Clinical Studies,

Houston, Texas, U.S.A.
Marie Studahl Department of Infectious Diseases, Sahlgrenska University

Hospital, Göteborg University, Göteborg, Sweden
Eva Thomas Department of Pathology and Laboratory Medicine,

University of British Columbia, and Children’s and Women’s Health
Centre, Vancouver, British Columbia, Canada
Gisela Torres Department of Dermatology, Center for Clinical Studies,

Houston, Texas, U.S.A.
Contributors xiii
Stephen K. Tyring Department of Dermatology, Center for Clinical

Studies, and University of Texas Health Science Center, Houston,
Texas, U.S.A.
Richard J. Whitley Division of Pediatric Infectious Diseases, University of

Alabama at Birmingham, Birmingham, Alabama, U.S.A.
SECTION I. UNDERSTANDING AND DIAGNOSING HSV
1
Evolution of Herpes Simplex Viruses
Rory J. Bowden and Duncan J. McGeoch

, ,
MRC Virology Unit Institute o f Virology, Glasgow Scotland\ U.K.
INTRODUCTION
The evolution of the two herpes simplex virus (HSV) species, HSV-1 and
HSV-2, falls naturally into two main components. The first of these con
cerns the ancient evolutionary history of the origins and development of
HSV-1 and HSV-2 within the contexts of the family Herpesviridae, the sub
family Alphaherpesvirinae, and the Simplexvirus genus to which the HSV
species are assigned. As we shall explain, we believe that these processes
took place over a timeframe of several hundred million years (MY), with
the most recent-dated event being the divergence of the HSV-1 and HSV-
2 lineages some 9 MY ago. The second part of our treatment concerns
the evolutionary processes that have generated and are active in contempor
ary populations of HSV-1 and HSV-2, which are composed of lineages that
we estimate have arisen well within the last million years.
Investigation of both these evolutionary phases depends on availability
of herpesviral genomic sequences. For the first phase of ancient development,
the primary analytical route comprises comparison of encoded amino acid
sequences for equivalent genes from across the herpesvirus family, with pro
minent use of methods for construction of phylogenetic trees. The second
phase, of population biology of each HSV species, was initiated two decades
ago using restriction nuclease profiles for the DNAs of HSV isolates to mea
sure diversity, while its modern practice employs DNA sequences from
selected genomic regions of isolates. Availability of sufficient comparative
/
2 Bowden and McGeoch
sequence data is only now approaching a level adequate for such analyses, so
our appreciation and understanding of recent HSV evolution is still at an
early stage.
ORIGINS OF THE FAMILY HERPESVIRIDAE
Common Descent of Mammalian Avian Herpesviruses
DNA sequence analyses of the genomes of mammalian and avian herpes

viruses have shown that these viruses are all related by descent from a com
mon ancestral virus. The evidence derives from a large subset (around 40) of
the viral genes that have been found to have counterparts in every comple
tely sequenced herpesvirus genome, and representatives of which have been
seen in partly sequenced genomes (1,2). The linear arrangements in genomes
of these “core” genes are partially preserved even among highly diverged
herpesviruses. The identified set of core genes ranges from the most con
served, whose encoded amino acid sequences are unambiguously related
across the family, to the least similar, where assignment is based on scant
local similarities only. Membership thus fades into uncertainty at this lower
end rather than being definitively bounded. In addition, there are three
genes not in the core set whose distribution and genomic locations argue
that they were present in the common ancestral virus but were subsequently
lost in some lineages. We thus identify a “minimal ancestral set” of 43 genes
(1,3). The core genes mostly encode components of the icosahedral capsid,
proteins concerned with DNA replication and packaging of replicated DNA
into nascent capsids, and virion surface glycoproteins involved in entry into
and egress from cells. On the other hand, most genes specifying proteins
with control functions and most genes encoding proteins of the virion tegu
ment and surface are specific to sublineages of the viruses.
Phylogenetic relationships among mammalian and avian herpesviruses
have been investigated using gene sequence data, and detailed phylogenetic
trees have been derived (4). In addition, the limited sequence data available
for reptilian (turtle) herpesviruses indicate that they are also part of this virus
group (5,6). The summary tree shown in Figure 1 depicts major lineages and
sublineages, and their relationships. The three deepest branches of the tree
correspond to the three taxonomic subfamilies, the Alpha-, Beta-, and Gam-
maherpesvirinae, and the eight terminal branches to genera in the subfamilies
(7). In this chapter, we are concerned with HSV-1 and HSV-2, which belong
to the Simplexvirus genus or ocl group of the Alphaherpesvirinae.
Robust phylogenetic trees derived from alignments of amino acid
sequences of up to eight genes from the core set revealed within several of
the sublineages patterns of relationships among virus species that mirrored
the branching patterns of corresponding lineages of mammalian host species
during divergent evolution of the host lines (4,8,9). This observation was
Evolution of Herpes Simplex Viruses 3
Figure 1 Major lineages for mammalian and avian herpesviruses. This summary
phylogenetic tree was derived from alignment of encoded amino acid sequences
for sets of conserved gene, using a maximum likelihood method with a molecular
clock imposed. Major lineages equivalent to genera are shown, and with recent taxo
nomic additions ( Mardivirus and lltovirus). Lineages belonging to the Alpha-, Beta-,
and Gammaherpesvirinae are also labeled with the appropriate Greek letter plus a
numeral. The least certain portions of the tree adjacent to the root are shown as
dashed lines. In terminal branches, heavy lines indicate the region in each lineage
with multiple branches. Source: From Refs. 4, 7.
consistent with cospeciation of herpesviruses with their hosts being a promi

nent mechanism in the genesis of herpesvirus lineages. While this idea was
not new, the sequence-based trees available in the last few years have
provided both good evidence in its support and also a semiquantitative
foundation that has then allowed estimation of a timescale for herpesvirus
evolution by transfer of spéciation dates from mammalian paleontology. It
has to be registered that there are complexities and limitations, as with any
estimate of evolutionary time based on differences among nucleic acid or
protein sequences. The herpesvirus phylogeny also shows instances of lineages
whose origins were interpreted as having been by transfer of a founding
virus between host species. Overall, we consider that the cospeciation-based
timescale for herpesviruses is a powerful tool in interpretation of the family’s
evolution.
The least certain portions of the tree in Figure 1 are the deep branches
nearest the root (shown as dashed lines), primarily because these are most
distant from the input sequences (equivalent to the branch tips) and infer
ences concerning them are accordingly most dependent on the modelling
algorithm used. Early work (Ref. 9, using the neighbor-joining algorithm)
gave an estimate for the root locus of around 200 million years before the
present, while later more extensive and sophisticated analyses gave a figure
of 400 million years (based on Ref. 4, with more input data, modelling by
maximum-likelihood methods with gamma distribution of substitution rates

and an updated paleontological timescale). For our present purposes, three
points regarding the root of the tree are relevant. First, the root corresponds
to a very remote period in the history of life on our planet. Second, the most
recent common ancestral species (at the root) was already a recognizable
herpesvirus, with a complement of at least 43 genes and with a capsid struc
ture and DNA replication systems comparable to the modern versions.
Third, this ancestor must therefore have had a substantial previous evolu
tionary development.
Early Evolution of Herpesviruses
Considering first the early evolution of herpesviruses that gave rise to the
most recent common ancestor of present day lineages, we can conclude little
of detail or certainty. In addition to the group of herpesviruses so far dis
cussed, of mammals, birds and reptiles, there are two other sets of viruses
currently assigned to the family Herpesviridae: a group of fish and amphi
bian viruses, and a single invertebrate (oyster) virus (2). Both display the
characteristic virion architecture of herpesviruses, and indeed this is the
basis of their assignment to the family. However, very little of their gene
complements are detectably related to those of the mammalian/avian/
reptile group. Like the latter, the fish/amphibian group comprises a set of
lineages that are clearly related by gene content (although with wide overall
divergence), while the oyster virus is distinct in its gene contents from both
of the vertebrate virus groups. Our interpretation of this situation is that the
three groups are probably genuinely related in having a common origin of
their capsid genes, but that most of the remainder of the gene complements
of the present day species is in each case acquired after divergence of these
three ancient lineages. This comparison thus sketches an early stage of evo
lutionary development, of what might be termed a “pre-herpesvirus.” A
final point regarding the origins of the herpesviruses is that certain aspects
of their mechanisms for packaging DNA are similar to those of DNA bac
teriophages, suggesting some very distant connection (2).
RELATIONSHIPS W ITH IN THE SUBFAMILY

ALPHAHERPESVIRINAE
Proposed Cospeciations of Viruses and Hosts

Four major lineages are recognized in the Alphaherpesvirinae. Two of these
are populated sparsely and exclusively by avian viruses, namely Mardivirus
(a3) and Iltovirus (a4). It has been speculated that these two genera arose by
way of interspecies transfers of early alphaherpesviruses, most likely from
mammals to birds. The mammalian viruses in the subfamily lie in a
clade with the avian virus lineages as outgroups, and with two genera,
Figure 2 Lineages in the subfamily Alphaherpesvirinae. This tree expands the

alphaherpesvirus portion of the phylogenetic tree in Figure 1, with emphasis on
the HSV-proximal branches. As in Figure 1, the heavy lines indicate in terminal
branches the locations of multiple branches. Abbreviations: OW, old world; HSV,
herpesvirus; BHV-2, bovine herpesvirus 2; NW, new world; OWP, old world pri
mate; V, virus; ILTV, infectious laryngotracheitis virus.
Simplexvirus (al) and Varicellovirus (a2). The main lineages within the
Alphaherpesvirinae are expanded in Figure 2.
In the a2 group, the branching pattern for the primate, artiodactyl,
perissodactyl, and carnivore viruses is congruent with the pattern for their
host lineages, at least for the level of detail shown in Figure 2; the compar
ison is effectively at the level of order in mammalian taxonomy. The struc
ture of the a l portion of the tree is a little more complicated: this group
comprises primate viruses, except for the occurrence of one bovine herpes
virus (BHV-2) and two marsupial herpesviruses (MaHV-1 and MaHV-2).
Setting aside these anomalies, the branching pattern of old world primate
(OWP) and new world primate viruses follows the pattern for the host
lineages. Thus, the a2 clade displays characteristics of cospeciation for four
orders of placental mammals, while the a l clade evinces this characteristic
for primates only. The fact that primate virus lineages occur in both these
groups in a manner consistent with cospeciation leaves unresolved the nat
ure of the original divergence event between the a l and a2 lineages. We can
not reliably distinguish with available data the branching orders for the
lineages of BHV-2, the marsupial viruses, and the OWP viruses. We suppose
that BHV-2 and the marsupial viruses arose from two separate transfers
from OWP viruses at some distant time following separation of the OWP
and new world primate host lineages. The nearest relatives of the HSV spe
cies are in the clade labelled as OW monkey viruses in Figure 2.
Three such viruses are represented in the tree, namely simian B virus
(SBV), simian agent 8 (SA8), and herpesvirus papio 2; these have the formal
taxonomic names of Cercopithecine herpesvirus 1, 2, and 16, respectively.
These OW monkey viruses were placed in the tree shown on the basis of
only one gene, that for virion glycoprotein B. However, their relationship
to the HSV species and other species in the immediate a l locality has been
confirmed by data for several other genes (US3, US4, and US6; not shown)
and is regarded as secure.
Relationships of the HSV Species and Other OWP Viruses
Establishing the origins of the HSV species in the context of their develop
ment within the OWP virus clade turns out to be an uncertain undertaking,
for several distinct reasons. A major consideration is the paucity of data
available for a l viruses of nonhuman OWPs. As shown in Figure 2, there
is a well-defined clade of OW monkey viruses, and the HSV species lie in
a sister clade that is unambiguously separated from the OW monkey virus
clade by an extended branch. However, the HSV species are the only viruses
in this sister clade: there are no viruses of other hominoids represented (i.e.,
species of chimpanzee, gorilla, orangutan, and gibbon), which we might
expect to be associated with the HSV clade. Overall, there is surprisingly lit
tle information available on a l herpesviruses of apes. Papers from the 1980s
and earlier described HSV-like infections in captive apes, with isolation of
viruses that by the criteria of the day were very similar to HSV (10,11). Sur
veys of sera from captive and wild apes detected antibodies against HSV-like
viruses (11,12). However, we have found no recent account of isolation or
characterization of such viruses, let alone DNA sequences. This lack of
description for a class that we believe must include the closest relatives of
the HSV species is a seriously limiting factor for evaluation of the immediate
antecedents of HSV-1 and HSV-2.
The next complexity concerns the relationship between the HSV spe
cies. HSV-1 and HSV-2 are each other’s closest known relative, but their
genomes are actually quite diverged. Based on recent phylogenetic analysis
(4), our current best estimate is that the two lineages diverged 9 million years
ago. Allowing for the uncertainties of such a calculation, it remains clear
that the separation is of considerable antiquity. For comparison, the spread
of modern humans across the planet took place over the last 0.1 million
years, while divergence of the human lineage from those of other hominoids
took place between 5.5 (chimpanzee) and 14.6 (gibbon) million years
before the present (13). Thus, we estimate that splitting of the two HSV
lineages occurred long before the emergence of modern humans, and within
approximately the same timeframe as spéciation of the hominoids. We

therefore arrive at the evaluation that key to understanding the evolutionary
context of the divergence of HSV-1 and HSV-2 would be a phylogenetic tree
incorporating related viruses of hominoids—which certainly exist but for
which we have no data at all. We also wish to enter reservations regarding
use of any argument for cospeciation with host in considering the emergence
of the HSVs. The instances outlined above for which postulating occurrence
of cospeciation apparently worked well concerned high-level taxa (mamma
lian orders) and dates involving several tens of millions of years, but it is
easy to imagine that the situation might become more complicated with clo
sely related sets of species and more recent divergence dates.
The occurrence in a single host species of more than one herpesvirus of
a given genus and with quite closely related genomes, as with HSV-1 and
HSV-2, is not uncommon. Other cases are known, for example in the a2
group with artiodactyl and perissodactyl viruses and in the y2 group with
primate viruses (14-16). We imagine that such cases could arise by several
routes, such as an episode of geographic isolation of two host populations
followed by re-establishment of contact, or divergence of virus lines in
two related host species followed by transfer between these hosts, or by
evolution within one host species into a novel tissue niche. The present
day differential propensities of HSV-1 and HSV-2 are compatible with the
last of these, although it is to be noted that there is no absolute difference
in their capability for growth in different tissues. Overall, we do not perceive
any cogent reason for supporting any particular spéciation scenario. Gentry
et al. (17) proposed that changes in mating habits of human ancestors
provided conditions for divergence of HSV-1 and HSV-2, but we regard this
hypothesis as over-specific, given the various possibilities.
Our general vision of the lineage leading to HSV, in the timeframe of
the last 10 or 20 million years, is of existence in hominoid species, with
separation into separate HSV-1 and HSV-2 lineages perhaps midway in this
period, and probably with many related virus species in the evolving range of
hominoids. The fact that HSV-1 and HSV-2 appear to us as closest relatives
may be misleading, given the lack of appropriate comparisons. Viewing the
HSV species separately, we regard the following as plausible scenarios. First
and most simply, each HSV might be the human-specific member in one or
the other of two distinct groups of hominoid viruses each conforming to
cospeciational relationships. Next, they might instead have a closer relation
ship with viruses of nonhuman hominoids: the HSV species might be very
similar to ape viruses as a result of interspecies transfer. Such multiple trans
fers could generate mixing of lineages for human and nonhuman viruses;
examples of this last mode are provided by lineages of primate immunodefi
ciency viruses (on a much shorter timescale) (18) and primate hepadnaviruses
(hepatitis B viruses) (19). Lastly, there is a third class of possible descent of
the HSV lineage: the relationship of HSV to other primate viruses might be
neither cospeciational nor especially close to other hominoid viruses; for

instance, transfer from some nonhominoid host might have occurred.
EVOLUTION OF ALPHAHERPESVIRUS GENOME STRUCTURES

AND GENE SETS
Evolution of Genomic Unique and Repeat Elements
All alphaherpesvirus genomes contain long and short unique regions (UL
and Us) and a pair of large repeat elements that flank the Us sequence in
opposing orientations (Rs). Some, including HSV-1 and HSV-2, also possess
a distinct pair of large repeat elements that flank U L (RL). These large-scale
features are illustrated for HSV in Figure 3, with conventions of genome
segment naming. All of the ancestral genes, common to the three subfamilies,
lie in the UL component. An equivalent arrangement of long and short
unique sequences with flanking repeats is also found in the Cytomegalovirus
genus of the Betaherpesvirinae, but the U s, Rs, and RL components of these
betaherpesvirus genomes are considered to be unrelated to the alphaherpes
virus sequences. It thus appears that the alphaherpesvirus S region (Us
plus its flanking R s copies) emerged after divergence from the Beta- and
Gammaherpesvirinae and before the appearance of the a l to a4 lineages.
For the purposes of this analysis, we regard R L elements as comprising
some thousands of base pairs (and containing protein coding genes) and
exclude the much smaller similarly placed sequences (of several tens of base
pairs) that are found in a2 genomes. By this definition, R L elements occur
only in the a l and a3 groups. However, on the basis of their sequences
and gene contents, a l RL and a3 RL are unrelated (20). We therefore regard
as likely that the common ancestral alphaherpesvirus genome was of form
Figure 3 Gross genome structure of HSV. The linear form of the HSV genome as
found in virions is depicted. Regions of unique sequence (U L and U s) are shown as
heavy lines, and major flanking repeat elements as open boxes. The terminal copy of
R L (designated TRL) is oppositely oriented to the internal copy IRL, and similarly for
TRS and IRS. The unit consisting of TRL- U L-IR L is termed the long or L region,
and IRS- U S-T R S the short or S region. The locations are indicated by short direct
repetitions at the genome termini (a sequences) and of an oppositely oriented copy
at the junction between IRL and IRs {a'). A scale bar is shown at the foot.
U l-IR s- U s-T R s, and that R L elements subsequently developed indepen

dently in the a l and a3 lineages. Inasmuch as HSV RL contains the ICPO
gene while in a2 genomes the homologous gene is present as a single copy
in a near-terminal location within UL, the a l R L elements must have origi
nated by duplication of a region that previously constituted one extremity of
U L. In the a l group, HSV-1, HSV-2, the OW monkey viruses SBV and SA8
(21,22) and BHV-2 (23) all possess R L elements, as do MaHV-2 (24) and
HVS-1 (25), so we can conclude that RL developed at an early point in
the a l lineage. However, it has been reported that MaHV-1 (26) lacks R L
regions, and the branching pattern for the a l sublineages in Figure 2 would
then suggest that RL repeat pairs were present in the genomes of ancestors
of MaHV-1 but that one R L copy has been lost in MaHV-1.
Genes Specific to the Simplexvirus Genus
We consider that HSV-1 and HSV-2 possess equivalent sets of genes, which
encode 74 distinct proteins (27,28). This estimate excludes some proposals
for additional genes that we regard as unproven and leaves aside as unre
solved any protein coding function of the latency-associated transcript locus.
Forty-three of the HSV genes belong to the ancestral set. In principle, the
other 31 HSV genes might have arisen early in the development of the Her-
pesviridae and then been lost in other lineages, or have first appeared after
the early Alphaherpesvirinae lineage split from that leading to the Beta-
and Gammaherpesvirinae. We regard as more plausible that new acquisition
within the Alphaherpesvirinae was the predominant route and for the present
discussion we treat this as the only mode. Twenty-two of these 31 genes are
also represented in the complete sequences of avian Mardivirus genomes (20),
present before the a3 lineage diverged from those leading to a l and a2. Three
of the remaining nine HSV genes have counterparts in the a l lineage [and one
of the three also has an a A homologue (29)], so these were also acquired at an
early stage in the development of the subfamily. This leaves six HSV genes
that are considered to have arisen within the a l lineage, namely U L56,
RL1, C/55, U S 8 A , C/577, and US12. The only complete genome sequences
presently available for viruses of the a l group are of HSV-1 and HSV-2
(27,28) so that our view of when the six a 1-specific HSV genes entered the
lineage is incomplete. We know from limited sequence data that the OW
monkey viruses SA8 and SBV have homologues of five of these HSV genes,
namely U L 56 , C/55, U S 8 A , C/577, and C/572 (30,31) (A. Dolan and D.
McGeoch, unpublished data), but there is no counterpart of the sixth gene,
RL1, at the corresponding locus in the genomes in these OW monkey viruses
(A. Dolan and D. McGeoch, unpublished data). However, there is a homo
logue of HSV RL1 in the marsupial virus MaHV-1, in an apparently equiva
lent genomic location (32). Thus, given the lines of descent shown in Figure 2,
we have to propose either that the RL1 gene was gained before divergence of
the marsupial virus and OWP virus lineages and then lost in the OW monkey
virus clade or—perceived as less likely—that the gene was independently
gained in the marsupial virus and HSV lineages. Our interim conclusion is
thus that none of the HSV complement of genes is unique to HSV-1 and
HSV-2, and all may have been present in the a l lineage before the divergence
of the HSV and OW monkey virus clades.
Evolution of Genomic S Regions
A number of attributes of the alphaherpesvirus S region indicate that it has

experienced a pronounced level of evolutionary activity. Five of the six a l-
specific HSV genes are in S. Among these, US12 and also US8A may have
evolved de novo (28). HSV U s also contains five genes for virion surface gly
coproteins, three of which have apparently evolved by way of duplication and
subsequent divergence, namely US4, US6, and US7, encoding glycoproteins
G, D, and I, respectively (33). Gene US5 encodes a very small type 1 glycopro
tein, which plausibly might be a member of the same gene family that has
undergone extensive deletion. Comparisons of the locations of the boundaries
between U s and Rs across the range of alphaherpesvirus sequences show that
the boundaries are dynamic—in some genomes a given gene may be in U s
while in others it is located across the border in R s (34). This phenomenon
is considered to have its basis in partly illegitimate recombination between
genomes, with one homologous crossover between copies of Rs and one het
erologous crossover involving two loci in Us copies; an equivalent mechanism
is presumed to have been involved in genesis of a l RL elements. R s elements
typically have a higher G +C content than the unique portions of the same
genome; in the HSV species, R s exhibits the highest G +C content of the
genome, at 80% G +C over its 6.7 kbp. This feature too may be a result of
recombination processes, with a biased gene conversion mechanism acting
to drive shift in overall base composition (35,36). The RL element of HSV
has similar characteristics to Rs, with a high G +C content (72% in HSV-1,
75% in HSV-2). The Rs and RL elements are involved in generating genome
orientation isomers by recombination during normal virus replication so that
with HSV four isomers in total are produced, with each genomic region (S and
L) occurring in either orientation with respect to the other.
Comparison of Genes of HSV-1 and HSV-2

While neither HSV-1 nor HSV-2 possesses any genes absent in the other,
their coding sequences are moderately diverged. Alignment of the two sets
of coding sequences (excluding US4, which is grossly truncated in HSV-1)
requires introduction of gapping characters equivalent to 2% of the
alignment length. These aligned coding regions have an average of 0.14 non-
synonymous substitutions per nonsynonymous site, while the corresponding
figure for synonymous substitution is 0.47 (28). There are only two
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E GA L I T É
DES
HOMMES ET DES FEMMES.
A LA REYNE.
M. D C . XX II .
A LA REYNE.
adame,
Ceux qui s’adviserent de donner un Soleil pour devise au Roy vostre

Pere, avec ce mot, Il n’a point d’Occident pour moy, firent plus qu’ils ne
pensoient: parce qu’en representans sa grandeur qui voit presque tousjours ce Prince
des Astres sur quelqu’une de ses terres, sans intervale de nuict; ils rendirent la
devise hereditaire en vostre Majesté, presageans vos vertus, & de plus, la beatitude
des François sous vostre Auguste presence. C’est dis-je chez vostre Majesté,
Madame, que la lumiere des vertus n’aura point d’Occident, ny consequemment
l’heur & la felicité de nos Peuples qu’elles esclaireront. Or comme vous estes en
l’Orient de vostre aage & de vos vertus ensemble, Madame, daignez prendre courage
d’arriver en mesme point au midy de luy & d’elles, je dis de celles qui ne peuvent
meurir que par temps & culture: car il en est quelques unes des plus
recommendables, entre autres la Religion, la charité vers les pauvres, la chasteté &
l’amour conjugale, dont vous avez touché le midy dés le matin. Mais certes il faut le
courage requis à cet effort aussi grand & puissant que vostre Royauté, pour grande &
puissante qu’elle soit: les Roys estant battus de ce malheur, que la peste infernale
des flatteurs qui se glissent dans les Palais, leur rend la vertu & la clair voyance sa
guide & sa nourrice, d’un accez infiniment plus difficile qu’aux inferieurs. Je ne scay
qu’un seur moyen à vous faire esperer, d’atteindre ces deux midys en mesme instant:
c’est qu’il plaise à V. M. se jetter vivement sur les bons livres de prudence & de
mœurs: car aussi tost qu’un Prince s’est relevé l’esprit par cet exercice, les flatteurs
se trouvans les moins fins ne s’osent plus jouër à luy. Et ne peuvent communement
les Puissans & les Roys recevoir instruction opportune que des mors: parce que les
vivans estans partis en deux bandes, les foux & meschans, c’est à dire ces flateurs
dont est question, ne sçavent ny veulent bien dire pres d’eux; les sages & gens de
bien peuvent & veulent, mais ils n’osent. C’est en la vertu certes, Madame, qu’il faut
que les personnes de vostre rang cherchent la vraye hautesse & la Couronne des
Couronnes: d’autant qu’ils ont puissance & non droit de violer les loix & l’equité, &
qu’ils trouvent autant de peril & plus de honte que les autres hommes à faire ce coup.
Aussi nous apprend un grand Roy luy mesme, que toute la gloire de la fille du Roy
est par dedans. Quelle est cependant ma rusticité, tous autres abordent leurs Princes
& Roys en adorant & loüant, j’ose aborder ma Reyne en preschant? Pardonnez
neantmoins à mon zele, Madame, qui meurt d’envie d’ouyr la France crier ce mot, avec
applaudissement, La lumiere n’a point d’Occident pour moy, par tout où passera
vostre Majesté nouveau Soleil des vertus: & d’envie encore de tirer d’elle, ainsi que
j’espere de ses dignes commencemens, une des plus fortes preuves du Traicté que
j’offre à ses pieds, pour maintenir l’egalité des hommes & des femmes. Et non
seulement veu la grandeur unique qui vous est acquise par naissance & par mariage,
vous servirez de miroir au sexe & de sujet d’emulation aux hommes encore, en
l’estenduë de l’Univers, si vous vous eslevez au prix & merite que je vous propose:
mais aussi tost, Madame, que vous aurez pris resolution de vouloir luyre de ce bel &
precieux esclat, on croira que tout le mesme sexe esclaire en la splendeur de vos
rayons. Je suis de vostre Majesté
MADAME,
Tres-humble & Tres-obeissante
servante & subjecte.
Gournay.
EGALITÉ DES HOMMES ET DES
FEMMES.
a pluspart de ceux qui prennent la cause, des femmes, contre cette

orgueilleuse preferance que les hommes s’attribuent, leur rendent le
change entier: r’envoyans la preferance vers elles. Moy qui fuys toutes
extremitez, je me contente de les esgaler aux hommes: la nature
s’opposant pour ce regard autant à la superiorité qu’à l’inferiorité. Que dis-
je, il ne suffit pas à quelques gens de leur preferer le sexe masculin, s’ils ne les
confinoient encores d’un arrest irrefragable & necessaire à la quenoüille, ouy mesme
à la quenouille seule. Mais ce qui les peut consoler contre ce mespris, c’est qu’il ne
se faict que par ceux d’entre les hommes ausquels elles voudroient moins
ressembler: personnes à donner vray semblance aux reproches qu’on pourroit vosmir
sur le sexe feminin, s’ils en estoient, & qui sentent en leur cœur ne se pouvoir
recommander que par le credit de l’autre. D’autant qu’ils ont ouy trompetter par les
ruës, que les femmes manquent de dignité, manquent aussi de suffisance, voire du
temperament & des organes pour arriver à cette-cy, leur eloquence triomphe à
prescher ces maximes: & tant plus opulemment, de ce que, dignité, suffisance,
organes & temperament sont beaux mots: n’ayans pas appris d’autre part, que la
premiere qualité d’un mal habill’homme, c’est de cautionner les choses soubs la foy
populaire & par ouyr dire. Voyez tels esprits comparer ces deux sexes: la plus haute
suffisance à leur advis où les femmes puissent arriver, c’est de ressembler le
commun des hommes: autant eslongnez d’imaginer, qu’une grande femme se peust
dire grand homme, le sexe changé, que de consentir qu’un homme se peust eslever
à l’estage d’un Dieu. Gens plus braves qu’Hercules vrayement, qui ne desfit que
douze monstres en douze combats; tandis que d’une seule parolle ils desfont la
moitié du Monde. Qui croira cependant, que ceux qui se veulent eslever & fortifier de
la foiblesse d’autruy, se puissent eslever ou fortifier de leur propre force? Et le bon
est, qu’ils pensent estre quittes de leur effronterie à vilipender ce sexe, usants d’une
effronterie pareille à se loüer & se dorer eux mesmes, je dis par fois en particulier
comme en general, voire à quelque tort que ce soit: comme si la verité de leur
vanterie recevoit mesure & qualité de son impudence. Et Dieu sçait si je congnois de
ces joyeux vanteurs, & dont les vanteries sont tantost passées en proverbe, entre les
plus eschauffez au mespris des femmes. Mais quoy, s’ils prennent droict d’estre
galans & suffisans hommes, de ce qu’ils se declarent tels comme par Edict; pourquoy
n’abestiront ils les femmes par le contrepied d’un autre Edict? Et si je juge bien, soit
de la dignité, soit de la capacité des dames, je ne pretends pas à cette heure de le
prouver par raisons, puisque les opiniastres les pouroient debattre, ny par exemples,
d’autant qu’ils sont trop communs; ains seulement par l’aucthorité de Dieu mesme,
des arcsboutans de son Eglise & de ces grands hommes qui ont servy de lumiere à
l’Univers. Rangeons ces glorieux tesmoins en teste, & reservons Dieu, puis les
Saincts Peres de son Eglise, au fonds, comme le tresor.
Platon à qui nul n’a debattu le tiltre de divin, & consequemment Socrates son
interprete & Protecole en ses Escripts; (s’il n’est là mesme celuy de Socrates, son
plus divin Precepteur) leur assignent mesmes droicts, facultez & functions, en leurs
Republiques & par tout ailleurs. Les maintiennent, en outre, avoir surpassé maintefois
tous les hommes de leur Patrie: comme en effect elles ont inventé partie des plus
beaux arts, ont excellé, voire enseigné cathedralement & souverainement sur tous les
hommes en toutes sortes de perfections & vertus, dans les plus fameuses villes
antiques entre autres Alexandrie, premiere de l’Empire apres Rome. Dont il Hypathia.
est arrivé que ces deux Philosophes, miracles de Nature, ont creu donner
plus de lustre à des discours de grand poix, s’ils les prononçoient en leurs livres par
la bouche de Diotime & d’Aspasie: Diotime que ce dernier ne craint point d’appeller
sa maistresse & Preceptrice, en quelques unes des plus hautes sciences, luy
Precepteur & maistre du genre humain. Ce que Theodoret releve si volontiers en
l’Oraison de la Foy, ce me semble; qu’il paroist bien que l’opinion favorable au sexe
luy estoit fort plausible. Apres tous ces tesmoignages de Socrates, sur le faict des
dames; on void assez que s’il lache quelque mot au Sympose de Xenophon contre
leur prudence, à comparaison de celle des hommes, il les regarde selon l’ignorance &
l’inexperience où elles sont nourries, ou bien au pis aller en general, laissant lieu
frequent & spatieux aux exceptions: à quoy les deviseurs dont est question ne
s’entendent point.
Que si les dames arrivent moins souvent que les hommes, aux degrez
d’excellence, c’est merveille que le deffaut de bonne instruction, voire l’affluence de la
mauvaise expresse & professoire ne face pis, les gardant d’y pouvoir arriver du tout.
Se trouve til plus de difference des hommes à elles que d’elles à elles mesmes, selon
l’institution qu’elles ont prinse, selon qu’elles sont eslevées en ville ou village, ou
selon les Nations? Et pourquoy leur institution ou nourriture aux affaires & Lettres à
l’egal des hommes, ne rempliroit elle ce vuide, qui paroist ordinairement entre les
testes des mesmes hommes & les leurs: puis que la nourriture est de telle importance
qu’un de ses membres seulement, c’est à dire le commerce du monde, abondant aux
Françoises & aux Angloises, & manquant aux Italiennes, celles cy sont de gros en
gros de si loing surpassées par celles là? Je dis de gros en gros, car en detail les
dames d’Italie triumphent par fois: & nous en avons tiré deux Reynes à la prudence
desquelles la France a trop d’obligation. Pourquoy vrayment la nourriture ne
frapperoit elle ce coup, de remplir la distance qui se void entre les entendemens des
hommes & des femmes; veu qu’en cet exemple icy le moins surmonte le plus, par
l’assistance d’une seule de ses parcelles, je dis ce commerce & conversation: l’air
des Italiennes estant plus subtil & propre à subtilizer les esprits, comme il paroist en
ceux de leurs hommes, confrontez communement contre ceux là des François & des
Anglois? Plutarque au Traicté des vertueux faicts des femmes maintient; que la vertu
de l’homme & de la femme est mesme chose. Seneque d’autre part publie aux
Consolations; qu’il faut croire que la Nature n’a point traicté les dames ingratement,
ou restrainct & racourcy leurs vertus & leurs esprits, plus que les vertus & les esprits
des hommes: mais qu’elle les a doüées de pareille vigueur & de pareille faculté à
toute chose honeste & loüable. Voyons ce qu’en juge apres ces deux, le tiers chef du
Triumvirat de la sagesse humaine & morale en ses Essais. Il luy semble, dit il, & si ne
sçait pourquoy, qu’il se trouve rarement des femmes dignes de commander aux
hommes. N’est ce pas les mettre en particulier à l’egale contrebalance des hommes,
& confesser, que s’il ne les y met en general il craint d’avoir tort: bien qu’il peust
excuser sa restrinction, sur la pauvre & disgraciée nourriture de ce sexe. N’oubliant
pas au reste d’alleguer & relever en autre lieu de son mesme livre, cette authorité que
Platon leur depart en sa Republique: & qu’Anthistenes nioit toute difference au talent
& en la vertu des deux sexes. Quant au Philosophe Aristote, puisque remuant Ciel &
terre, il n’a point contredit en gros, que je scache, l’opinion qui favorise les dames, il
l’a confirmée: s’en rapportant, sans doubte, aux sentences de son pere & grand pere
spirituels, Socrates & Platon, comme à chose constante & fixe soubs le credit de tels
personnages: par la bouche desquels il faut advoüer que le genre humain tout entier,
& la raison mesme, ont prononcé leur arrest. Est il besoing d’alleguer infinis autres
Erasme Epist: & anciens & modernes de nom illustre, ou parmy ces derniers,
Colloq. Politia: Epist. Erasme, Politien, Agripa, ny cet honneste & pertinent Precepteur
Agripa Precel: du des courtizans: outre tant de fameux Poëtes si contrepoinctez
sexe feminin tous ensemble aux mespriseurs du sexe feminin, & si partisans
Courtizan. de ses advantages aptitude & disposition à tout office & tout
exercice louable & digne? Les dames en verité se consolent,
que ces descrieurs de leur merite ne se peuvent prouver habiles gens, si tous ces
esprits le sont: & qu’un homme fin ne dira pas, encores qu’il le creust, que le merite &
passedroit du sexe feminin tire court, pres celuy du masculin; jusques à ce que par
arrest il ait faict declarer tous ceux là buffles, affin d’infirmer leur tesmoignage si
contraire à tel decry. Et buffles faudroit il encores declarer des Peuples entiers & des
plus sublins, entre autres ceux de Smyrne en Tacitus: qui pour obtenir jadis à Rome
presseance de noblesse sur leurs voisins, allegoient estre descendus, ou de Tantalus
fils de Jupiter ou de Theseus petit fils de Neptune ou d’une Amazone, laquelle par ce
moyen ils contrepesoient à ces Dieux. Pour le regard de la loy Salique, qui prive les
femmes de la couronne, elle n’a lieu qu’en France. Et fut inventée au temps de
Pharamond, pour la seulle consideration des guerres contre l’Empire duquel nos
Peres secoüoient le joug: le sexe feminin estant vray semblablement d’un corps
moins propre aux armes, par la necessité du port & nourriture des enfans. Il faut
rémarquer encores neantmoins, que les Pairs de France ayans esté créez en
Hotman pour premiere intention comme une espece de personniers des Roys,
ainsi que leur nom le declare: les dames Pairaisses de leur chef
l’etymologie des Pairs:
du Tillet & Math. ont seance, privilege & voix deliberative par tout où les Pairs en
Histoire du Roy pour ont & de mesme estendue. Comme aussi les Lacedemoniens ce
les Dames Pairresses. brave & genereux Peuple, consultoit de toutes affaires privées &
Plut. publiques avec ses femmes. Bien a servy cependant aux
François, de trouver l’invention des Regentes, pour un equivalent des Roys; car
sans cela combien y a il que leur Estat fust par terre? Nous sçaurions bien dire
aujourd’huy par espreuve, quelle necessité les minoritez des Roys ont de cette
recepte. Les Germains ces belliqueux Peuples, dit Tacitus, qui apres plus de deux
cens ans de guerre, furent plustost triumphéz que vaincus; portoient dot à leurs
femmes, non au rebours. Ils avoient au surplus des Nations, qui n’estoient jamais
regies que par ce sexe. Et quand Ænee presente à Didon le sceptre d’Ilione, les
scoliastes disent, que cela provient, de ce que les dames filles aisnées, telle qu’estoit
cette Princesse, regnoient anciennement aux maisons Royalles. Veult-on deux plus
beaux envers à la loy Salique, si deux envers elle peut souffrir? Si ne mesprisoient
pas les femmes nos anciens Gaulois, ny les Carthaginois aussi; lors qu’estans unis
en l’armée d’Hanibal pour passer les Alpes, ils establirent les dames Gauloises
arbitres de leurs differends. Et quand les hommes desroberoient à ce sexe en
plusieurs lieux, part aux meilleurs advantages; l’inegalité des forces corporelles plus
que des spirituelles, ou du merite, peut facilement estre cause du larrecin & de sa
souffrance: forces corporelles, qui sont vertus si basses, que la beste en tient plus
par dessus l’homme, que l’homme par dessus la femme. Et si ce mesme
Historiographe Latin nous apprend, qu’où la force regne, l’equité, la probité, la
modestie mesme, sont les attributs du vainqueur; s’estonnera-on, que la suffisance &
les merites en general, soient ceux de nos hommes, privativement aux femmes.
Au surplus l’animal humain n’est homme ny femme, à le bien prendre, les sexes
estants faicts non simplement, mais secundum quid, comme parle l’Eschole: c’est à
dire pour la seule propagation. L’unique forme & difference de cet animal, ne consiste
qu’en l’ame humaine. Et s’il est permis de rire en passant, le quolibet ne sera pas
hors de saison, nous apprenant; qu’il n’est rien plus semblable au chat sur une
fenestre, que la chatte. L’homme & la femme sont tellement uns, que si l’homme est
plus que la femme, la femme est plus que l’homme. L’homme fut creé masle &
femelle, dit l’Escriture, ne comptant ces deux que pour un. Dont Jesus-Christ est
appellé fils de l’homme, bien qu’il ne le soit que de la femme. Ainsi parle apres le
grand Sainct Basile: La vertu de l’homme & de la femme est mesme chose, Homil. I.
puis que Dieu leur a decerné mesme creation & mesme honneur: masculum
& fœminam fecit eos. Or en ceux de qui la Nature est une & mesme, il faut que les
actions aussi le soient, & que l’estime & loyer en suitte soient pareils, où les œuvres
sont pareilles. Voila donc la deposition de ce puissant pilier, & venerable tesmoing de
l’Eglise. Il n’est pas mauvais de se souvenir sur ce poinct, que certains ergotistes
anciens, ont passé jusques à cette niaise arrogance, de debattre au sexe feminin
l’image de Dieu à difference de l’homme: laquelle image ils devoient, selon ce calcul
attacher à la barbe. Il failloit de plus & par consequent, desnier aux femmes l’image
de l’homme, ne pouvant luy ressembler, sans qu’elles ressemblassent à celuy auquel
il ressemble. Dieu mesme leur a departy les dons de Prophetie Olda Debora.
indifferamment avec les hommes, les ayant establies aussi pour Juges,
instructrices & conductrices de son Peuple fidelle en paix & en guerre: & qui plus est,
rendu triumphantes avec luy des hautes victoires, qu’elles ont aussi maintefois
emportées & arborées en divers lieux du Monde: mais sur quelles gens, à vostre
advis? Cyrus & Theseus: à ces deux on adjouste Hercules, lequel elles ont sinon
vaincu, du moins bien battu. Aussi fut la cheute de Pentasilée, couronnement de la
gloire d’Achilles: oyez Seneque & Ronsard parlans de luy.
L’Amazone il vainquit dernier effroy des Grecs.

Pentasilée il rua sur la poudre.
Ont elles au surplus, (ce mot par occasion) moins excellé de foy, qui comprend toutes
les vertus principales, que de suffisance & de force magnanime & guerriere?
Paterculus nous apprend, qu’aux proscriptions Romaines, la fidelité des enfans fut
nulle, des affranchis legere, des femmes tresgrande. Que si Sainct Paul, suyvant ma
route des tesmoignages saincts, leur deffend le ministere & leur commande le silence
en l’Eglise: il est evident que ce n’est point par aucun mespris: ouy bien seulement,
de crainte qu’elles n’esmeuvent les tentations, par cette montre si claire & publique
qu’il faudroit faire en ministrant & preschant, de ce qu’elles ont de grace & de beauté
plus que les hommes. Je dis que l’exemption de mespris est evidente, puisque cet
Apostre parle de Thesbé comme de sa coadjutrice en l’œuvre de nostre Seigneur,
sans toucher le grand credit de Saincte Petronille vers sainct Pierre: & puis aussi que
la Magdeleine est nommée en l’Eglise egale aux Apostres, par Apostolis. Voire que
l’Eglise & eux-mesmes ont permis une exception de ceste reigle Entre autres au
de silence pour elle, qui prescha trente ans en la Baume de Calendrier des Grecs,
Marseille au rapport de toute la Provence. Et si quelqu’un publié par Genebrard.
impugne ce tesmoignage de predications, on luy demandera que
faisoient les Sibyles, sinon prescher l’Univers par divine inspiration, sur l’evenement
futur de Jesus-Christ? Toutes les anciennes Nations concedoient la Prestrise aux
femmes, indifferemment avec les hommes. Et les Chrestiens sont au moins forcez de
consentir, qu’elles soyent capables d’appliquer le Sacrement de Baptesme: mais
quelle faculté de distribuer les autres, leur peut estre justement deniée; si celle de
distribuer cestuy-là, leur est justement accordé? De dire que la necessité des petits
enfans mourans, ait forcé les Peres anciens d’establir cet usage en despit d’eux: il est
certain qu’ils n’auroient jamais creu que la necessité les peust dispenser de mal faire,
jusques aux termes de permettre violer & diffamer l’application d’un Sacrement. Et
partant concedans ceste faculté de distribution aux femmes, on void à clair qu’ils ne
les ont interdites de distribuer les autres Sacremens, que pour maintenir tousjours
plus entiere l’auctorité des hommes; soit pour estre de leur sexe, soit afin qu’à droit
ou à tort, la paix fust plus asseurée entre les deux sexes, par la foiblesse &
Epist. ravallement de l’un. Certes sainct Jerosme escrit sagement à nostre propos;
qu’en matiere du service de Dieu, l’esprit & la doctrine doivent estre
considerez, non le sexe. Sentence qu’on doit generaliser, pour permettre aux Dames
à plus forte raison, toute action & science honneste: & cela suyvant aussi les
intentions du mesme sainct, qui de sa part honnore & auctorise bien fort leur sexe.
Davantage sainct Jean l’Aigle & le plus chery des Evangelistes, ne mesprisoit pas les
femmes non plus que sainct Pierre, sainct Paul & ces deux Peres, j’entends saint
Basile & sainct Jerosme; puis qu’il leur addresse ses Epistres Electra.
particulierement: sans parler d’infinis autres Ss: ou Peres, qui font pareille
addresse de leurs Escrits. Quand au faict de Judith je n’en daignerois faire mention
s’il estoit particulier, cela s’appelle dependant du mouvement & volonté de son
auctrice: non plus que je ne parle des autres de ce qualibre; bien qu’ils soient
immenses en quantité, comme ils sont autant heroiques en qualité de toutes sortes,
que ceux qui couronnent les plus illustres hommes. Je n’enregistre point les faicts
privez, de crainte qu’ils semblent, non advantages & dons du sexe, ains boüillons
d’une vigueur privée & specialle. Mais celuy de Judith merite place en ce lieu, parce
qu’il est bien vray, que son dessein tombant au cœur d’une jeune dame, entre tant
d’hommes lasches & faillis de cœur, à tel besoing, en si haulte & si difficile entreprise,
& pour tel fruict, que le salut d’un Peuple & d’une Cité fidelle à Dieu: semble plustost
estre une inspiration & prerogative divine vers les femmes, qu’un traict purement
voluntaire. Comme aussi le semble estre celuy de la Pucelle d’Orleans, accompagné
de mesmes circonstances environ, mais de plus ample & large utilité, s’estendant
jusques au salut d’un grand Royaume & de son Prince.
Æneid. I.
allusion. Cette illustre Amazone instruicte aux soins de Mars,
Fauche les escadrons & brave les hazars:
Vestant le dur plastron sur sa ronde mammelle,
Dont le bouton pourpré de graces estincelle:
Pour couronner son chef de gloire & de lauriers,
Vierge elle ose affronter les plus fameux guerriers.
Adjoustons que la Magdelene est la seule ame, à qui le Redempteur ait jamais
prononcé ce mot, & promis cette auguste grace: En tous lieux où se preschera
l’Evangile il sera parlé de toy. Jesus-Christ d’autrepart, declara sa tres heureuse &
tres glorieuse resurrection aux dames les premieres, affin de les rendre, dit un
venerable Pere ancien, Apostresses aux propres Apostres: cela, comme lon sçait,
avec mission expresse: Va, dit il, à cette cy mesme, & recite aux Apostres & à Pierre
ce que tu as veu. Surquoy il faut notter, qu’il manifesta sa nouvelle naissance
esgalement aux femmes qu’aux hommes, en la personne d’Anne fille de Phannel, qui
le recongneut en mesme instant, que le bon vieillard Sainct Simeon. Laquelle
naissance, d’abondant, les Sybilles nommées, ont predite seules entre les Gentils,
excellent privilege du sexe feminin. Quel honneur faict aux femmes aussi, ce songe
survenu chez Pilate; s’addressant à l’une d’elles privativement à tous les hommes, &
en telle & si haulte occasion. Et si les hommes se vantent, que Jesus-Christ soit nay
de leur sexe, on respond, qu’il le failloit par necessaire bien sceance, ne se pouvant
pas sans scandale, mesler jeune & à toutes les heures du jour & de la nuict parmy les
presses, aux fins de convertir, secourir & sauver le genre humain, s’il eust esté du
sexe des femmes: notamment en face de la malignité des Juifs. Que si quelqu’un au
reste est si fade; d’imaginer masculin ou feminin en Dieu, bien que son nom semble
sonner le masculin, ny consequemment besoin d’acception d’un sexe plustost que de
l’autre, pour honnorer l’incarnation de son fils; cettuy cy monstre à plein jour, qu’il est
aussi mauvais Philosophe que Theologien. D’ailleurs, l’advantage qu’ont les hommes
par son incarnation en leur sexe; (s’ils en peuvent tirer un advantage, veu cette
necessité remarquée) est compensé par sa conception tres precieuse au corps d’une
femme, par l’entiere perfection de cette femme, unique à porter nom de parfaicte
entre toutes les creatures purement humaines, depuis la cheute de nos premiers
parens, & par son assumption unique en suject humain aussi.
Finalement si l’Escripture a declaré le mary, chef de la femme, la plus grande
sottise que l’homme peust faire, c’est de prendre cela pour passedroict de dignité.
Car veu les exemples, aucthoritez & raisons nottées en ce discours, par où l’egalité
des graces & faveurs de Dieu vers les deux especes ou sexes est prouvée, voire leur
unité mesme, & veu que Dieu prononce: Les deux ne seront qu’un: & prononce
encores: L’homme quittera pere & mere pour suivre sa femme; il paroist que cette
declaration n’est faicte que par le besoin expres de nourrir paix en mariage. Lequel
besoin requeroit, sans doubte, qu’une des parties cédast à l’autre, & la prestance des
forces du masle ne pouvoit pas souffrir que la soubmission vînt de sa part. Et quand
bien il seroit veritable, selon que quelques uns maintiennent, que cette soubmission
fut imposée à la femme pour chastiement du peché de la pomme: cela encores est
bien esloigné de conclure à la pretendue preferance de dignité en l’homme. Si lon
croioit que l’Escripture luy commendast de ceder à l’homme, comme indigne de le
contrecarrer, voyez l’absurdité qui suivroit: la femme se treuveroit digne d’estre faicte
à l’image du Createur, de jouyr de la tressaincte Eucaristie, des mysteres de la
Redemption, du Paradis & de la vision voire possession de Dieu, non pas des
advantages et privileges de l’homme: seroit ce pas declarer l’homme plus precieux &
relevé que telles choses, & partant commettre le plus grief des blasphemes?
FIN.
L’IMPRIMEUR A RANGÉ
ces vers icy pour emplir le reste
de la feuille.
AUTHEUR INCERTAIN.
Lumine Acron dextro captus, Leonilla sinistro,

Et potis est forma vincere uterque Deos.
Blande puer, lumen quod habes concede sorori:
Sic tu cæcus Amor, sic erit illa Venus.
VERSION.
Lys & sa jeune mere aussy beaux que les Dieux:

De deux costez divers ont perdu l’un des yeux.
Lys, donne ton bon œil à ta mere Argentine;
Tu seras Cupidon, elle sera Cyprine.
AUTREMENT.
Lyse & son petit Lys aussy beaux que les Dieux,
De deux costez divers ont perdu l’un des yeux.
Si Lys donne l’autre œil à sa mere admirée;
Il est l’aveugle Amour, & Lyse Cytherée.
EX HORATIO.
Dial.
Donec gratus eram tibi,

Nec quisquam potior brachia candidæ
Cervici iuvenis dabat,
Persarum vigüi Rege beatior.
Donec non alia magis
Arsisti, neque erat Lydia post Chloen,
Lydia multi nominis,
Romana vigui clarior Ilia.
Me nunc Thrassa Chloe regit,
Dulces docta modos & Cytharæ sciens:
Pro qua non metuam mori,
Si parcent animæ fata superstiti.
Me torret face mutua
Thurini Calais filius Orinthi:
Pro quo bis patiar mori,
Si parcent puero fata superstiti.
Quid si prisca redit Venus,
Diductosque iugo cogit aheneo?
Si flava excutitur Chloe
Reiectæque patet ianua Lydiæ?
Quanquam sidere pulchrior
Ille est, tu levior cortice & improbo
Iracundior Adria;
Tecum vivere amem, tecum obeam libens.
DIALOGUE D’HORACE
ET DE LYDIE.
Tandis que mon Amour t’enflammoit constamment,

Tandis qu’un jeune amy, bravant ma jalousie,
Ne pressoit ton beau col d’un mol embrassement,
J’ay flory plus heureux qu’un Monarque d’Asie.
Devant que ton esprit brisast sa loyauté,
Devant qu’il eust chery d’une aveugle folie
Cloé plus que Lydie, illustre de beauté,
J’ay surmonté l’esclat de la Romaine Ilie.
Cloé Greque sans pair me possede à son tour
Par son luth & sa voix qui scait charmer l’oreille:
Et mourrois volontiers, victime de l’Amour,
Pour conserver mourant cette jeune merveille.
Calaïs Thurien épris de mes appas,
Par un revers gentil de ses attraits me blesse:
Et souffrirois deux fois la rigueur du trespas,
Pour sauver du tombeau cette belle jeunesse.
Quoy si l’amour premier ressuscitant son feu
Ramenoit soubs ton joug mon ame revoltée?
Quoy si mon cœur solide éternisant son vœu,
Ma Lydie est reçeue & Cloé rejettée?
Encor qu’il soit plus beau qu’un astre au front des cieux,
Toy plus leger qu’un liege & plus mutin que l’onde;
Je veux rouler mes jours aux prisons de tes yeux,
Je veux que mon cercueuil tes obseques seconde.
INCERTAIN SUR L’HORLOGE
DE SABLE.
Exiguus vitro pulvis qui dividit horas,

Et levis angustum sæpe recurrit iter,
Olim Alcipus erat: qui Marthæ ut vidit ocellos
Arsit, & est subito factus ab igne cinis.
Irrequiete cinis, miseros testabere amantes,
More tuo, nulla posse quiete frui.
VERSION
Ce peu de poudre, helas qui fîle en ces deux verres,

Courant & recourant sur ses estroictes erres,
Affin de marquer l’heure & mesurer le jour,
Estoit jadis Alcipe esclave de l’Amour.
Bruslé des yeux de Marthe il coula tout en cendre:
Et faut, cendre inquiete, en ton aspect comprendre:
Qu’un miserable esprit blessé par un bel œil
N’a jamais de repos s’il te manque au cercueil.
Au lecteur
Cette version numérisée reproduit dans son intégralité la version

originale.
La ponctuation a pu faire l’objet de quelques corrections mineures.
L’orthographe a été conservée. Seuls quelques mots ont été modifiés.
La version originale pouvant être difficile à déchiffrer, nous avons fait
suivre celle-ci d’une version modernisée dans laquelle nous avons remplacé
ſ et ß par s, ã par an ou am, ẽ par en ou em, le ĩ par in, õ par on ou om, le ũ
par um, ainsi que les abréviations de vo9 et pl9 par vous et plus.
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Textbook Herpes Simplex Viruses First Edition Tomas Bergstrom Ebook All Chapter PDF

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Herpes Simplex Viruses First Edition

Cell Biology of Herpes Viruses 1st Edition Klaus

Evolution Second edition Carl T. Bergstrom

Real-Time Rendering 4th Edition Tomas Akenine-Mo■Ller

A Planet of Viruses Third Edition Zimmer

Molecular and Cellular Biology of Viruses 1st Edition

War, Revolution, and Nation-Making in Lithuania,

Molecular Virology of Human Pathogenic Viruses 1st

Viruses as Therapeutics Methods and Protocols 1st

1. Parasitic Infections in the Compromised Host,

Reissued 2018 by CRC Press

© 2006 by Taylor & Francis

No claim to original U.S. Government works

A Library of Congress record exists under LC control number: 2005052899

ISBN 13: 978-1-138-10594-2 (hbk)

Ubiquitous viruses among humans, herpes simplex viruses (HSV) are

chapters document the different disease manifestations of HSV infections,

SECTION I. UNDERSTANDING AND DIAGNOSING HERPES

1. Evolution of Herpes Simplex Viruses 1

2. Herpes Simplex Virus Vaccines and the Viral Strategies

3. The Natural History and Epidemiology

Detection of Viral Genomes, Whole Virus,

SECTION II. DISEASE MANIFESTATIONS OF HSV

6. Antiviral Treatment............................................................. 153

7. Primary Herpes Simplex Gingivostomatitis and Recurrent

8. Herpesvirus Infections of the Skin ...................................... 191

Localized Cutaneous HSV . . . . 195

9. Acute and Recurrent Genital Herpes Simplex

10. Herpes Simplex Viruses Ocular D isease............................ 239

11. Herpes Simplex Encephalitis and Other Neurological Syndromes

12. Neurological Disease in Herpes Simplex Virus Type 2

13. Herpes Simplex Virus and Bell’s Palsy ............................ 339

14. Herpes Simplex Virus Infections in Immunocompromised

15. Neonatal Herpes Simplex Virus Infection.......................... 395

16. Future Outlooks ................................................................. 411

Jacob Amir Department of Pediatric C, Schneider Children’s Medical

Elisabeth Aurelius Karolinska Intistute, Unit of Infectious Diseases,

Susanne Beckman-Nahmias Emory University, Atlanta, Georgia, U.S.A.

Tomas Bergström Department of Clinical Virology, Göteborg University,

Rory J. Bowden MRC Virology Unit, Institute of Virology,

Kevin S. Buckley Division of Pediatric Infectious Diseases, University of

Paola Cinque Clinic of Infectious Diseases, Scientific Institute San

Fulvio Crippa Clinic of Infectious Diseases, Scientific Institute San

Harvey M. Friedman Infectious Disease Division, Department of

Yasushi Furuta Department of Otolaryngology-Head & Neck Surgery,

Lauren M. Hook Infectious Disease Division, Department of Medicine,

David W. Kimberlin Division of Pediatric Infectious Diseases, University

George Kinghorn Department of Genitourinary Medicine, Royal

Gerald Kleymann Department of Chemistry and Pharmacy,

Francis K. Lee Emory University, Atlanta, Georgia, U.S.A.

Thomas J. Liesegang Department of Ophthalmology, Mayo Clinic,

Duncan J. McGeoch MRC Virology Unit, Institute of Virology,

André J. Nahmias Emory University, Atlanta, Georgia, U.S.A.

Birgit Sköldenberg Division of Medicine, Unit of Infectious Diseases,

Karan K. Sra Department of Dermatology, Center for Clinical Studies,

Marie Studahl Department of Infectious Diseases, Sahlgrenska University

Eva Thomas Department of Pathology and Laboratory Medicine,

Gisela Torres Department of Dermatology, Center for Clinical Studies,

Stephen K. Tyring Department of Dermatology, Center for Clinical

Richard J. Whitley Division of Pediatric Infectious Diseases, University of

Rory J. Bowden and Duncan J. McGeoch

ORIGINS OF THE FAMILY HERPESVIRIDAE

Common Descent of Mammalian Avian Herpesviruses

DNA sequence analyses of the genomes of mammalian and avian herpes

consistent with cospeciation of herpesviruses with their hosts being a promi

U l-IR s- U s-T R s, and that R L elements subsequently developed indepen