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Basic Virology
Third
Edition
Edward K. Wagner
Department of Molecular Biology and Biochemistry
University of California Irvine
Mart inezJ. Hewlett
Department of Molecular and Cellular Biology
University of Arizona
David C. Bloom
Department of Molecular Genetics and Microbiology
University of Florida
David Camerini
Department of Molecular Biology and Biochemistry
University of California Irvine
Blackwell
Publishing
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Brief Contents
Preface xxi
Acknowledgments xx ix
P A R T I V I R O L O G Y A N D V I R A L D IS E A S E 1
Chapter 1 Intro duct ion Th e Impact of Viruses on O ur View of Life 3
Chapter 2 An Ou tline of Virus Replication and Viral Pathogenesis 15
Chapter 3 Virus Disease in Population s and Individual Animals 27
Chapter 4 Patterns of Some Viral Diseases of H um ans 41 *
PART I I BASIC PROPE RTIES OF VIRUSES A N D V IR U S -
CELL INT ER AC TIO N 63
Chapter 5 Virus Struc ture and Classification 65
Chapter 6 The Beginning and End of the Virus Replication Cycle 79
Chapter 7 Ho st Im m une Response to Viral Infection - Th e Na ture of the Vertebrate
Imm une Response 97
Chapter 8 Strategies to Protect Against and Co mb at Viral Infection 119
PART I I I W O R K IN G W IT H V IRUS 145
Chapter 9 Visualization and Enu me ration of Virus Particles 147
Chapter 10 Replicating and Me asuring Biological Activity of Viruses 155
Chapter 11 Physical and Chemical M anipu lation of the Structural Com pon ents of
Viruses 173
Chapter 12 Characterization of Viral Products Expressed in the Infected Cell 193
Chapter 13 Viruses Use Cellular Processes to Express their Genet ic
Information 213
P AR T IV R E P L I C A T I O N P A T T E R N S O F S P E C I F I C
VIRUSES 243
Chapter 14 Replication of Positive-sense RN A Viruses 245
Chapter 15 Replication Strategies of RNA Viruses Requ iring RNA -directed m RNA
Transcriptio n as the First Step in Viral Gene Expression 273
Chapter 16 Replication Strategies of Small and Medium -sized D N A Viruses 303
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Contents
Preface xxi
Preface to the second edition xxii
Preface to the third editio n xxii
Text organization xxiii
Specific features of this text designed to aid instructors and
students in pursuin g topics in greater depth xxvi
Dept h of coverage xxvi
Sources for further study xxvi
The Intern et xxvii
Chap ter outlines xxvii
Case studies xxvii
Review mate rial xxvii
Glossary xxvii
Acknowledgments xxix
PART I
CHAPTER 1
VIRO LOG Y AN D V IRAL DISEASE 1
Introduc tion - The Impact of Viruses on Ou r View of Life 3
Th e science of virology 3
The effect of virus infections on the host organism and populations
viral pathogenesis, virulence, and epidemiology 4
Th e interaction between viruses and their hosts 6
Th e history of virology 7
Examples of the impact of viral disease on hum an history 8
Examples of the evolutionary impact of the virus-host
interaction 9
Th e origin of viruses 9
Viruses have a constructive as well as destructive impact on
society 12
Viruses are not the smallest self-replicating path ogen s 13
Que stions for Cha pter 1 14
CHAPTER 2
An Outline of Virus Replication and Viral Pathogenesis
Virus replication 15
Stages of virus replication in the cell 17
Pathogenesis of viral infection 19
Stages of virus-induced pathology 19
15
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VI I I
CONTENTS
CHAPTER 3
CHAPTER 4
Initial stages of infection - entry of the virus into the host 20
Th e incuba tion period and spread of virus throug h the host 21
Multiplication of virus to high levels
occurrence of disease
symptoms 23
The later stages of infection
the imm une response 24
The later stages of infection
virus spread to the next
individual 24
The later stages of infection
fate of the host 24
Questions for Chapter 2 25
Virus Disease in Popu lations and Individual Animals 27
Th e nature of virus reservoirs 27
Some viruses with huma n reservoirs 28
Some viruses with verteb rate reservoirs 30
Viruses in popula tions 30
Viral epidemiology in small and large popula tions 30
Factors affecting the control of viral disease in populations 33
Anim al models to study viral pathogenesis 34
A mouse model for studying poxvirus infection and spread 35
Rabies: where is the virus durin g its long incubation period? 37
Herpes simplex virus latency 37
Mu rine models 39
Rabbit models 40
Guinea pig models 40
Questions for Chapter 3 40
Patterns o f Some Viral Diseases of Humans 41
The dynamics of hum an-viru s interactions 42
The stable association of viruses with their natural host places
specific constraints on the nature of viral disease and mode of
persistence 42
Classification of human disease-causing viruses according to
virushost dynam ics 44
Viral diseases leading to persistence of the virus in the host are
generally associated with viruses having long associations with
hum an populations 44
Viral diseases associated with acute, severe infection are
suggestive of zoonoses 48
Patte rns of specific viral diseases of hu m ans 49
Acu te infections followed by virus clearing 49
Colds and respiratory infections 49
Influenza 49
Variola 49
Infection of an accidental target tissue leading to permane nt
dam age despite efficient clearing 50
Persistent viral infections 50
Papilloma and polyomavirus infections 50
Herpesvirus infections and latency 52
Ot her complications arising from persistent infections 52
Viral and subviral diseases with long incuba tion periods 53
Rabies 53
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CONTENTS IX
PART II
CHAPTER 5
CHAPTER 6
CHAPTER 7
HIV-AIDS 53
Prion diseases 54
Some viral infections targetin g specific organ systems 54
Viral infections of nerve tissue 54
Examples of viral encep halitis with grave progno sis 55
Rabies 55
Herpes encephalitis 55
Viral encepha litis with favorable prognosis for recovery 56
Viral infections of the liver (viral hepatitis ) 56
Hepatitis A 57
Hepatitis B 57
Hepatitis C 57
Hepatitis D 57
Hepatitis E 58
Que stions for Cha pter 4 58
Problems for Part I 59
Add itional Reading for Part I 61
BASIC PROPERTIES OF VIRUSES A N D
V I R U S C E L L I N T E R A C T I O N
63
Virus Structure and Classification 65
The features of virus 65
Viral genomes 69
Viral capsids 69
Viral envelopes 72
Classification schemes 72
Th e Baltim ore scheme of virus classification 75
Disease-based classification schemes for viruses 75
Th e virosphere 77
Que stions for Chap ter 5 78
The Beginning and End o f the Virus Replication Cycle 79
Ou tline of the virus replication cycle 79
Viral entry 80
Animal virus entry into cells the role of the cellular receptor 80
Mech anisms of entry of nonenve loped viruses 83
En try of enveloped viruses 84
Entry of virus into plant cells 85
Injection of bacteriophage DNA into scherichia coli 87
Nonspecific methods of introd ucing viral genomes into cells 89
Late events in viral infection: capsid assembly and virion release 89
Assembly of helical capsids 89
Assembly of icosahedral capsids 92
Generation of the virion envelope and egress of the enveloped
virion 93
Question s for Chap ter 6 96
Host Immune Response to Viral Infection - The Nature of the
Verteb rate Imm une Response 97
The innate imm une response - early defense against pathogens 98
Toll-like receptors 99
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CONTENTS
Defensins 99
Th e adaptive imm un e response and the lymp hatic system 100
Tw o pathways of helper T response - the fork in the road 101
The immunological structure of protein 102
Role of the antigen-presenting cell in initiation of the immune
response 104
Clonal selection of im mu ne reactive lympho cytes 107
Immu ne memory 108
Co mp lem ent-m ediate d cell lysis 108
Con trol and dysfunction of imm unity 108
Specific viral responses to host immu nity 109
Passive evasion of immunity antigenic drift 110
Passive evasion of immunity internal sanctuaries for
infectious virus 110
Passive evasion of immunity imm une tolerance 110
Active evasion of imm unity - imm unosuppression 111
Active evasion of imm unity - blockage of M H C antigen
presentation 111
Con sequences of imm un e suppression to virus infections 112
Measurem ent of the imm une reaction 112
Measurem ent of cell-mediated (T-cell) imm unity 112
Measurement of antiviral antibody 112
Enzym e-linked imm uno sorb ent assays (ELISAs) 113
Neu tralization tests 114
Inhibition of hemagg lutination 114
Complement fixation 115
Questions for Chap ter 7 ^117
C H A P T E R 8 Strategies to Protect Against and Com bat Viral Infection 119
Vaccination
induction of immunity to prevent virus
infection 120
An tiviral vaccines 120
Smallpox and the history of vaccination 120
Ho w a vaccine is produced 122
Live-virus vaccines 122
Killed-virus vaccines 123
Reco mb inant virus vaccines 124
Capsid and subu nit vaccines 124
DN A vaccines 125
Edible vaccines 125
Problem s with vaccine prod uctio n and use 125
Eu kary otic cell-based defenses against virus replication 126
Interferon 126
Ind uctio n of interferon 127
Th e antiviral state 128
Me asurem ent of interferon activity 128
O th er cellular defenses against viral infection 130
Small RN A-based defenses 130
Enzym atic modification of viral genom es 131
Antiviral drugs 131
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X I
PART
III
CHAPTER9
CHAPTER
1
Targeting antiviral drugsto specific featuresof the virus
replication cycle
131
Acyclovir
and the
herpesviruses
132
Blocking influenza virus entryandvirus maturation 132
Chemotherapeutic approachesfor HIV 134
Multiple drug therapies toreduceoreliminate mutationto
drug resistance
134
Other approaches
135
Bacterial antiviral systems restriction endonucleases 135
QuestionsforChapter 8 136
Problems
for
Part
II 139
Additional Reading
for
Part
II 143
WORKING WITH VIRUS 145
VisualizationandEnumerationofVirus Particles 147
Using
the
electron microscope
to
study
and
count viruses
147
Counting (enumeration)
of
virions with
the
electron
microscope
149
Atomic force microscopy
a
rapid
and
sensitive meth od
for
visualization
of
viruses
and
infected cells, potentially
in
real
time
151
Indirect methods for coun ting virus particles 152
Questions forChapter 9 154
Replicatingand Measuring Biological ActivityofViruses 155
Cell culture techniques
156
Maintenanceof bacterial cells 156
Plant cell cultures 156
Culture
of
animal
and
human cells
157
Maintenance
of
cells
in
culture
157
Types
of
cells
157
Lossof contact inhibition ofgrowth andimmortalizationof
primary cells 159
The outcome
of
virus infection
in
cells
160
Fate
of
the virus
160
Fate
of
the cell following virus infection
162
Cell-mediated maintenanceof the intra-and intercellular
environment 162
Virus-mediated cytopathology changes
in the
physical
appearance
of
cells
163
Virus-mediated cytopathology changesin thebiochemical
propertiesofcells 163
Measurement
of
the biological activity
of
viruses
164
Quantitative measure
of
infectious centers
164
Plaque assays
164
Generation of transformed cell foci 165
Use of virus titerstoquantitatively control infection conditions 166
Examples
of
plaque assays
167
Statistical analysis
of
infection
168
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CHAPTER 11
CHAPTER 12
Dilution endpo int methods 169
The relation between dilution endpoint and infectious units of
virus 169
Ques tions for Chap ter 10 170
Physical and Chemical Manipulation of the Structural
Com ponents of Viruses 173
Viral structural proteins 173
Isolation of structural proteins of the virus 174
Size fractionation of viral structu ral prote ins 176
Det erm ining the stoichiometry of capsid proteins 177
The poliovirus capsid - a virion with equimolar capsid
proteins 178
Analysis of viral capsids that do not contain equimolar numbers
of proteins 179
Characterizing viral genomes 179
Sequen ce analysis of viral genom es 180
Mea suring the size of viral genomes 184
Direct measure of DNA genome lengths in the electron
microscope 185
Rate zonal sedimentation and gel electrophoresis for measuring
viral geno me size 185
The polymerase chain reaction detection and characterization of
extremely small quantities of viral genomes or transcripts 187
Real time PCR for precise quantitative measures of viral
DNA 189
PCR detection of RNA 190
PC R as an epidemiological tool 190
Que stions for Chap ter 11 191
Characterization of Viral Products Expressed in the Infected
Cell 193
Cha racteri zation of viral prote ins in the infected cell 193
Pulse labeling of viral proteins at different times following
infection 194
Use of imm une reagents for study of viral proteins 195
Work ing with antibodies 196
Dete ction of viral proteins using immuno fluorescence 198
Related methods for detecting antibodies bound to
antigens 201
Detecting and characterizing viral nucleic acids in infected
cells 205
Dete cting the synthesis of viral genomes 205
Characterization of viral mR NA expressed durin g infection 205
In situ hybridization 207
Further characterization of specific viral mRNA
molecules 209
Use of microarray technology for getting a complete picture of the
events occurring in the infected cell 210
Questions for Chap ter 12 212
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XIII
CHAPTER 13
PART IV
CHAPTER 14
Viruses Use Cellular Processes to Express Their Genetic
Information 213
Prokaryotic DNA replication is an accurate enzymatic model for
the process generally 21 5
Th e replication of eukaryotic DN A 216
Th e replication of viral DN A 21 7
Th e effect of virus infection on host D N A replication 217
Expression of mR NA 217
Prokaryotic transcription 219
Prokaryotic RNA polymerase 219
The prokaryotic promoter and initiation of
transcription 220
Con trol of prokaryotic initiation of transcription 220
Term ination of prokaryotic transcription 221
Eukaryotic transcription 221
Th e promo ter and initiation of transcription 221
Con trol of initiation of eukaryotic transcription 223
Processing of precursor mR NA 224
Visualization and location of splices in eukaryotic
transcripts 226
Posttranscriptional regulation of eukaryotic mRNA
function 231
Virus-induced changes in transcription and posttranscriptional
processing 232
Th e mechanism of protein synthesis 232
Euk aryotic translation 233
Prokaryotic translation 234
Virus-ind uced changes in translation 236
Questions for Chap ter 13 236
Problems for Part III 239
Additional Reading for Part III 241
R E P L I C A T I O N
VIRUSES 243
PATTERNS OF SPECIF IC
245
eplication of Positive-sense RNA Viruses
RNA viruses general considerations 246
A general picture of RNA -directed RNA replication 246
Replication of positive-sense RNA viruses whose genomes are
translated as the first step in gene expression 248
Positive-sense RNA viruses encoding a single large open reading
frame 249
Picornavirus replication 249
The poliovirus genetic map and expression of poliovirus
proteins 249
Th e poliovirus replication cycle 252
Picornavirus cytopatholog y and disease 254
Flavivirus replicatio n 25 6
Positive-sense RNA viruses encoding more than one translational
reading frame 257
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Two viral mRNAs are produced in different amounts during
togavirus infections 258
Th e viral genome 258
Th e virus replication cycle 258
Togavirus cytopatholog y and disease 262
A somewhat more complex scenario of multiple translational
reading frames and subgenomic mRNA expression: coronavirus
replication 263
Coron avirus replication 264
Cyto patho logy and disease caused by coronaviruses 266
Replication of plant viruses with RNA genomes 267
Viruses with one genome segment 268
Viruses with two genom e segments 268
Viruses with three genom e segments 269
Replication of bacteriophage with RNA genomes 269
Regulated translation of bacteriophage mR NA 269
Questions for Chap ter 14 272
C H A PT
R 1 5 Replication Strategies o f RNA Viruses Requiring RNA-directed
mRNA Transcription as the First Step in Viral Gene
Expression 273
Replication of negative-sense RNA viruses with a monopartite
genome 275
Replication of vesicular stomatitis virus - a model for
Mononegavirales 275
Vesicular stomatitis virus virion and genom e 275
Generation, capping, and polyadenylation of mR NA 276
Gen eration of new negative-sense virion RNA 278
Mech anism of host shutoff by vesicular stomatitis virus 279
Cyto patho logy and diseases caused by rhabdoviruses 280
Paramyxoviruses 280
Pathogenesis of paramyxoviruses 280
Filoviruses and their pathogenesis 28 2
Bornaviruses 282
Influenza viruses - negative-sense RN A viruses with a
multipartite genome 283
Involvem ent of the nucleus in flu virus replication 284
Generation of new flu nucleocapsids and maturation of the
virus 285
Influenza A epidemics 285
Other negative-sense RNA viruses with multipartite
genomes 288
Bunyaviruses 288
Virus structure and replication 288
Pathogenesis 291
Arenaviruses 291
Virus gene expression 292
Pathogenesis 292
Viruses with double-stranded RNA genomes 292
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CHAPTER 16
CHAPTER 17
Reovirus structu re 292
Reovirus replication cycle 294
Pathogenesis 295
Subviral pathog ens 29 5
Hep atitis delta virus 296
Viroids 297
Prions 298
Que stions for Ch apter' 15 301
Replication Strategies of Small and Medium-Sized DNA
Viruses 303
DNA viruses express genetic information and replicate their
genomes in similar, yet distinct, ways 304
Papovavirus replication 30 5
Replication of SV40 virus - the model polyomavirus 305
Th e SV40 genome and genetic map 309
Productive infection by SV40 310
Abortive infection of cells nonpermissive for SV40
replication 312
Replication of papillomaviruses 31 4
The HPV -16 genome 316
Virus replication and cytopatholog y 316
Replication of adenoviruses 319
Physical properties of adenovirus 319
Capsid structure 319
Th e adenovirus genome 319
Th e adenovirus replication cycle 319
Early events 31 9
Adenovirus DN A replication 321
Late gene expression 321
VA transcription and cytopathology 321
Transfo rmation of nonperm issive cells by adenovirus 323
Replication of some single-stranded D N A viruses 32 3
Replication of parvoviruses 323
Dependovirus DNA integrates in a specific site in the host cell
genome 324
Parvoviruses have potentially exploitable therapeutic
applications 325
DN A viruses infecting vascular plants 325
Geminiviruses 325
Single-stranded DNA bacteriophage OX174 packages its genes
very comp actly 326
Questions for Chapter 16 328
Replication of Some Nuclear-Replicating Eukaryotic DNA Viruses
wi th Large Genomes 331
Herpesvirus replication and latency 332
Th e herpesviruses as a group 332
Gen etic complexity of herpesviruses 33 3
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Co mm on features of herpesvirus replication in the host 33 3
Th e replication of the prototypical alpha-herpesvirus -
HSV 334
The HSV virion 334
Th e viral genome 334
HS V productive infection 338
HS V latency and LAT 347
HS V transcription during latency and reactivation 348
Ho w do LA T and other specific HS V genes function - may be
to accom mo date reactivation? 350
EBV latent infection of lymphocytes, a different set of problems
and answers 351
Pathology of herpesvirus infections 354
Herpesviruses as infectious co-carcinogens 354
Baculovirus, an insect virus with important practical uses in
molecular biology 355
Virion structure 355
Viral gene expression and genom e replication 356
Pathogenesis 356
Imp ortance of baculoviruses in biotechnolo gy 356
Questions for Chapter 17 357
C H A P T E R 1 8 Replication of Cytoplasmic DNA Viruses and Large
Bacteriophages 359
Poxviruses DNA viruses that replicate in the cytoplasm of
eukaryotic cells 360
The pox virion is complex and contains virus-coded transcription
enzymes 360
T he poxvirus replication cycle 361
Early gene expression 363
Geno me replication 363
Interm ediate and late stages of replication 363
Pathogenesis and history of poxvirus infections 36 4
Is smallpox virus a poten tial biological terror weapon? 364
Replication of large DN A-con taining bacteriophages 365
Co mp onen ts of large DN A-co ntaining phage virions 365
Replication of phage T7 367
The genome 367
Phage-controlled transcription 367
Th e practical value of T 7 367
T4 bacterioph age: the basic mod el for all DN A viruses 367
The T4 genome 368
Regulated gene expression dur ing T 4 replication 369
Capsid ma turatio n and release 370
Replication of phage
X:
a simple model for latency and
reactivation 370
The phage Xgenome 372
Phage
X
gene expression imm ediately after infection 37 2
Biochemistry of the decision between lytic and lysogenic
infection in
E. coli
375
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CHAPTER 19
CHAPTER 20
CHAPTER 21
A group of algal viruses shares features of its genome structure with
poxviruses and bacteriophages 376
Que stions for Cha pter 18 377
Retroviruses: Conv erting RNA to DNA 381
Retrovirus families and their strategies of replication 382
Th e molecular biology of retrovirus 383
Retrovirus structural proteins 383
Th e retrovirus genom e 384
Gene tic maps of representative retroviruses 386
Replication of retroviruses: an outline of the replication
process 386
Initiation of infection 386
Capsid assembly and ma turation 389
Action of reverse transcriptase and RNase-H in synthesis of
cDNA 389
Virus gene expression, assembly, and ma turatio n 391
Transcription and translation of viral mRN A 391
Capsid assembly and morphog enesis 391
Mechan isms of retrovirus transformation 392
Transformation through the action of
viral oncogene
a
subverted cellular growth control gene 392
Oncornavirus alteration of normal cellular transcriptional control
of growth regulation 393
Oncornavirus transformation by growth stimulation of
neighboring cells 395
Cellular genetic elements related to retroviruses 395
Retrotransposons 396
Th e relationship between transposable elements and viruses 397
Qu estions for Cha pter 19 397
Human Immunodeficiency Virus Type 1 (HIV-1) and Related
Lentiviruses 399
HI V- 1 and related lentiviruses 399
The origin of HIV -1 and AIDS 399
HI V- 1 and lentiviral replication 400
Destruction of the imm une system by HIV-1 406
Questions for Chapter 20 411
Hepadnaviruses: Variations on the Retrovirus Theme 413
Th e virion and the viral genom e 413
Th e viral replication cycle 41 5
The pathogenesis of hepatitis B virus 415
A plant hepadna virus : cauliflower mosaic virus 416
Geno me structure 416
Viral gene expression and genom e replication 416
Th e evolutionary origin of hepadnaviruses 417
Questions for Chapter 21 419
Problems for Part IV 421
Add itional Reading for Part IV 429
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CONTENTS
PART V
CHAPTER 22
CHAPTER 23
CHAPTER 24
V I R U S E S : N E W A P P R O A C H E S A N D N E W
PROBLEMS 433
The Mo lecular Genetics of Viruses 435
M utatio ns in genes and resulting changes to proteins 437
Analysis of mu tation s 43 8
Complementation 438
Recombination 439
Isolation of mutan ts 440
Selection 440
HS V thymidine kinase - a portable selectable marker 440
Screening
441
441 tool kit for molecular virologists
Viral genomes 441
Locating sites of restriction endonuclease cleavage on the viral
genome - restriction mapp ing 442
Clon ing vectors 44 3
Cloning of fragments of viral genomes using bacterial
plasmids 444
Cloning using phage X 449
Clonin g single-stranded DN A with bacteriophage M 13 451
DN A animal virus vectors 451
RN A virus expression systems 45 3
Defective virus particles 45 4
Directed mutagenesis of viral genes
Site-directed mutagenesis 45 6
Generation of recombinant viruses
Bacterial artificial chromosomes
Questions for Chapter 22 461
454
456
458
Molecu lar Pathogenesis 463
An introd uctio n to the study of viral pathogenesis 46 3
Animal models 464
Choo sing a model: natural host vs. surrogate models 464
Dev elopm ent of new models: transgenic animals 464
Hyb rid models: the SC ID-h u mouse 464
Considerations regarding the hum ane use of animals 465
Meth ods for the study of pathogenesis 466
Assays of virulenc e 46 6
Analysis of viral spread within the host 46 7
Resolving the infection to the level of single cells 47 0
Characterization of the host response 470
Imm unolog ical assays 470
Use of transgenic mice to dissect critical components of the host
im mu ne response that mo dulate the viral infection 471
Question for Chap ter 23 471
Viral Bioinformatics and Beyond
Bioinformatics 473
Bioinformatics and virology 47 3
Biological databases 47 4
473
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CHAPTER 25
APPENDIX
Primary databases 47 4
Secondary databases 475
Com posite databases 475
Oth er databases 476
Biological applications 47 6
Similarity searching tools 476
Protein functional analysis 478
Sequ ence analysis ,478
Structural modeling 478
Structural analysis 47 8
Systems biology and viruses 479
Viral internet resources 481
Questions for Chap ter 24 484
Viruses and the Future - Problems and Promises 485
Clouds on the horizon emerging disease 48 5
Sources and causes of emergen t virus disease 488
Th e threat of bioterrorism 489
What are the prospects of using medical technology to eliminate
specific viral and othe r infectious diseases? 49 0
Silver linings viruses as therape utic agents 49 0
Viruses for gene delivery 49 1
Using viruses to destroy other viruses 49 3
Viruses and nanotechnology 49 3
Th e place of viruses in the biosphere 49 4
W hy study virology? 494
Questions for Chap ter 25 495
Problems for Part V 49 7
Additional Reading for Part V 499
Resource Center 501
Books of historical and basic value 501
Books on virology 501
Mo lecular biology and biochem istry texts 503
Detailed sources 50 3
Sources for experimental protocols 503
Th e Internet 504
Virology sites 504
Important websites for organizations and facilities of
interest 505
Technical Glossary 507
Index 533