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Samenvatting van alle hoorcolleges van week 1 t/m 8 van de cursus Virologie (BMW31405)
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  • Course
  • Virologie (BMW31405)
  • Institution
  • Universiteit Utrecht (UU)
  • Book
  • Principles of Virology

Deze gedetailleerde samenvatting behandelt alles wat aan bod is gekomen tijdens de hoorcolleges gedurende week 1 t/m 8 van de cursus Virologie aan de Universiteit Utrecht (BMW31405). Daarnaast zijn de corresponderende hoofdstukken uit het boek Principles of Virology (4th edition Flint) ook in de sa...

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  • January 8, 2023
  • 297
  • 2021/2022
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  • virion
  • morfologie
  • classificatie virussen
  • baltimore classificatie
  • virale levenscyclus
  • entry
  • replicatie
  • genexpressie
  • assemblage
  • pathogenese
  • uitscheiding
  • transmissie
  • hematogene verspreiding
  • neurale versp

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  • Biomedische Wetenschappen
  • Virologie (BMW31405)
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Samenvatting virologie
Week 1 t/m 8



Biomedische Wetenschappen
2021-2022




Door Nicole (studente BMW)

,Inhoudsopgave
Algemene introductie virologie ............................................................................................................... 8
Geschiedenis ....................................................................................................................................... 8
Ivanosfki........................................................................................................................................... 8
Beijerinck ......................................................................................................................................... 8
Loeffler en Frosch ............................................................................................................................ 8
Walter Reed en Jesse William Lazear .............................................................................................. 9
Definitie virus ...................................................................................................................................... 9
Virion ................................................................................................................................................. 10
Karakteriseren virionen ................................................................................................................. 11
Morfologie ..................................................................................................................................... 12
Classificatie van virussen ................................................................................................................... 14
Baltimore classificatie.................................................................................................................... 14
Virussen en de cel.................................................................................................................................. 15
Virale levenscyclus............................................................................................................................. 15
Entry .............................................................................................................................................. 16
Virale replicatie.............................................................................................................................. 18
Virale genexpressie ....................................................................................................................... 22
Virale assemblage.......................................................................................................................... 22
Pathogenese .......................................................................................................................................... 24
Uitscheiding ....................................................................................................................................... 24
Respiratoire secretie ..................................................................................................................... 24
Transmissie ........................................................................................................................................ 24
Virale verspreiding............................................................................................................................. 26
Hematogene verspreiding ............................................................................................................. 28
Neurale verspreiding ..................................................................................................................... 30
Tropisme ............................................................................................................................................ 31
Virale virulentie ................................................................................................................................. 34
Meten virulentie ............................................................................................................................ 34
Factoren virulentie ........................................................................................................................ 34
Afweermechanismen van de gastheer .............................................................................................. 36
Adaptieve afweer (specifiek) ......................................................................................................... 36
Patronen van infectie ........................................................................................................................ 39
Herpes simplex virus ..................................................................................................................... 40
Humaan immunodeficiëntievirus (HIV) ......................................................................................... 40
Mechanismen van ziekte ................................................................................................................... 40

1

, Complicaties .................................................................................................................................. 40
Structuur van virionen ........................................................................................................................... 41
Functies ............................................................................................................................................. 41
Bescherming van het genoom ....................................................................................................... 42
Aflevering van het genoom ........................................................................................................... 42
Overige functies............................................................................................................................. 42
Terminologie ..................................................................................................................................... 42
Analysemethoden ............................................................................................................................. 43
Elektronenmicroscopie.................................................................................................................. 43
Cryo-elektronenmicroscopie ......................................................................................................... 43
Tomografie .................................................................................................................................... 45
X-ray kristallografie ....................................................................................................................... 45
Eiwitmantel ....................................................................................................................................... 46
Helicale symmetrie ........................................................................................................................ 46
Icosahedrale symmetrie ................................................................................................................ 48
Membraan omvattende virussen ...................................................................................................... 53
Aanhechting: receptoren en receptorbindende eiwitten ..................................................................... 55
Entry .................................................................................................................................................. 55
Aanhechting ...................................................................................................................................... 56
Virale receptoren identificeren ..................................................................................................... 57
Virus-receptor interacties ................................................................................................................. 60
Naakte virussen ............................................................................................................................. 60
Membraan omvattende virussen .................................................................................................. 61
Penetratie en ontmanteling .................................................................................................................. 63
Penetratie .......................................................................................................................................... 63
Membraan omvattende virussen .................................................................................................. 64
Naakte virussen ............................................................................................................................. 71
Virale replicatie ................................................................................................................................. 75
Replicatie in de kern ...................................................................................................................... 75
Virale RNA synthese .............................................................................................................................. 77
Positief (+) strengige RNA virussen ................................................................................................... 77
Negatief (–) strengige RNA virussen.................................................................................................. 77
RNA-afhankelijke RNA synthese ........................................................................................................ 78
RNA-afhankelijke RNA polymerase ............................................................................................... 79
Supplementaire eiwitten (accessoire eiwitten) ............................................................................ 82
Template........................................................................................................................................ 84

2

, Primer ............................................................................................................................................ 86
Replicatie positief (+) strengige RNA virussen................................................................................... 88
Poliovirus ....................................................................................................................................... 88
Sindbis virus ................................................................................................................................... 90
Replicatie negatief (–) strengige RNA virussen ................................................................................. 94
Influenza A virus ............................................................................................................................ 94
Rhabdovirussen ............................................................................................................................. 99
DNA virussen: Transcriptie mechanismen .......................................................................................... 102
Promotor ......................................................................................................................................... 102
RNA polymerasen ............................................................................................................................ 103
Meten transcriptie........................................................................................................................... 103
Run-off assay ............................................................................................................................... 103
Luciferase assay ........................................................................................................................... 104
Transcriptieproces ........................................................................................................................... 105
Variatie in promotoren ................................................................................................................ 106
Stimulatie van transcriptie .......................................................................................................... 110
Enhancers .................................................................................................................................... 112
DNA virussen: Replicatie mechanismen .............................................................................................. 115
Simian virus 40 (SV40) ..................................................................................................................... 116
Large T antigen (LT) ..................................................................................................................... 117
Replicatie ..................................................................................................................................... 118
Self-priming ..................................................................................................................................... 121
Replicatie experimenten ................................................................................................................. 124
Processing van virale (pre-)mRNA’s .................................................................................................... 126
Capping ............................................................................................................................................ 126
Cap snatching .............................................................................................................................. 130
Poly-adenylering .............................................................................................................................. 131
Splicing............................................................................................................................................. 133
Alternatieve splicing .................................................................................................................... 136
Overige post-transcriptionele modificaties..................................................................................... 139
Methylering ................................................................................................................................. 139
2’-O-methyltransferasen ............................................................................................................. 141
RNA-editing ................................................................................................................................. 142
RNA turnover ............................................................................................................................... 145
mRNA transport naar het cytoplasma......................................................................................... 147
Transcriptional host shut-off door virussen ................................................................................ 149

3

, Non-coding RNA’s........................................................................................................................ 150
Regulatie van translatie ....................................................................................................................... 156
Ribosomen ....................................................................................................................................... 156
tRNA’s .......................................................................................................................................... 157
Aminoacyl-tRNA synthetasen ...................................................................................................... 158
Polyribosomen............................................................................................................................. 159
Translatie ......................................................................................................................................... 160
Initiatie van translatie.................................................................................................................. 161
Elongatie ...................................................................................................................................... 163
Terminatie ................................................................................................................................... 164
Diversiteit van translatie strategieën .......................................................................................... 165
Regulatie van translatie ................................................................................................................... 169
Geïntegreerde stress respons (eIF2) ........................................................................................... 170
Virus-geïnduceerde host shut-off via eIF4F ................................................................................ 173
Gevolgen host shut-off ................................................................................................................ 176
Intracellulair transport van virion componenten ................................................................................ 179
Transport naar de kern (na entry) ................................................................................................... 180
Transport naar het plasmamembraan (na replicatie) ..................................................................... 182
Membraan eiwitten..................................................................................................................... 182
Niet-membraan eiwitten ............................................................................................................. 192
Transport naar intracellulaire membranen ..................................................................................... 193
Transport van virale genoom .......................................................................................................... 195
Assemblage, exit en maturatie ............................................................................................................ 197
Bestuderen assemblage .................................................................................................................. 197
1. Vorming capside .......................................................................................................................... 198
Influenza A virus .......................................................................................................................... 200
HIV ............................................................................................................................................... 201
Herpes simplex virus 1 (HSV-1) ................................................................................................... 202
2. Verpakken van het genoom ........................................................................................................ 203
Herpes simplex virus 1 (HSV-1) ................................................................................................... 204
HIV ............................................................................................................................................... 205
Influenza A virus .......................................................................................................................... 205
Incorporatie van andere virion componenten ............................................................................ 206
3. Budding........................................................................................................................................ 206
4. Afgifte .......................................................................................................................................... 207
Naakte virussen ........................................................................................................................... 207

4

, Membraan omvattende virussen ................................................................................................ 208
5. Maturatie..................................................................................................................................... 214
Type verspreiding ............................................................................................................................ 215
Evolutie en genetische diversiteit van RNA en DNA virussen ............................................................. 216
Diversiteit ........................................................................................................................................ 216
Mutaties ...................................................................................................................................... 216
Selectie ........................................................................................................................................ 217
Recombinatie/reassortment ....................................................................................................... 219
Bronnen virale diversiteit ............................................................................................................ 220
Virus evolutie............................................................................................................................... 221
Opkomende virussen........................................................................................................................... 222
HIV-1 ................................................................................................................................................ 223
SARS-CoV-1 ...................................................................................................................................... 223
MERS-CoV ........................................................................................................................................ 223
SARS-CoV-2 ...................................................................................................................................... 223
Vaccins ................................................................................................................................................. 224
Geschiedenis ................................................................................................................................... 224
Vaccinatie op grote schaal............................................................................................................... 225
Uitroeiing ..................................................................................................................................... 225
Basis van vaccinaties ....................................................................................................................... 226
Immuungeheugen ....................................................................................................................... 226
Veiligheid, effectiviteit en praktisch ............................................................................................ 227
Soorten vaccins ............................................................................................................................... 229
Geattenueerde vaccins (replicatie-competent) .......................................................................... 230
Geïnactiveerde vaccins ................................................................................................................ 231
Subunit vaccins ............................................................................................................................ 233
Nucleïnezuur-gebaseerde vaccins ............................................................................................... 234
Antigeniciteit verbeteren ................................................................................................................ 235
Antivirale middelen ............................................................................................................................. 236
Geschiedenis ................................................................................................................................... 237
Aangrijpingspunten ......................................................................................................................... 238
Screeningmethoden ........................................................................................................................ 239
Mechanisme-gebaseerde screening............................................................................................ 239
Cel-gebaseerde screening ........................................................................................................... 240
Structuur-gebaseerde drug design (in silico)............................................................................... 241
Screening van antivirale activiteit in vitro ....................................................................................... 242

5

, Dosering bepalen......................................................................................................................... 243
Voorbeelden van antivirale middelen ............................................................................................. 244
Voorkomen van aanhechting ...................................................................................................... 244
Voorkomen van penetratie en ontmanteling.............................................................................. 245
Voorkomen van reverse transcriptie van retrovirussen (RNA naar DNA) ................................... 245
Voorkomen van assemblage, maturatie en afgifte ..................................................................... 247
Highly active antiretroviral therapy (HAART) .................................................................................. 248
Virussen en het afweersysteem .......................................................................................................... 249
Cel-autonome intrinsieke antivirale afweer.................................................................................... 250
Apoptose ..................................................................................................................................... 251
Aangeboren en adaptieve immuunrespons .................................................................................... 253
Aangeboren immuunrespons ...................................................................................................... 253
Adaptieve immuunrespons ......................................................................................................... 253
Aangeboren immuunrespons .......................................................................................................... 254
IFN-α/β ........................................................................................................................................ 254
Herkenning van virussen ............................................................................................................. 255
IFN gentranscriptie ...................................................................................................................... 262
IFN respons .................................................................................................................................. 264
Adaptieve immuunrespons ............................................................................................................. 269
Antilichamen ............................................................................................................................... 270
Virale suppressie ......................................................................................................................... 272
Overige virale evasie mechanismen ................................................................................................ 273
DNA virussen vs. RNA virussen .................................................................................................... 273
HIV: ontdekking, transmissie en pathogenese .................................................................................... 274
HIV epidemie ................................................................................................................................... 274
Ontdekking van HIV ......................................................................................................................... 274
Oorsprong van HIV .......................................................................................................................... 274
Transmissie ...................................................................................................................................... 275
Verloop van infectie ........................................................................................................................ 276
1. Acute fase ................................................................................................................................ 277
2. Asymptomatische/chronische fase ......................................................................................... 277
3. Ontwikkeling van AIDS............................................................................................................. 277
Entry van HIV ................................................................................................................................... 278
Pathogenese .................................................................................................................................... 281
Immuundysfunctie ...................................................................................................................... 281
Centraal zenuwstelsel.................................................................................................................. 283

6

, Kanker.......................................................................................................................................... 284
HIV levenscyclus .................................................................................................................................. 285
Genoomorganisatie ......................................................................................................................... 285
Virale replicatiecyclus ...................................................................................................................... 290
Entry ............................................................................................................................................ 290
Replicatie ..................................................................................................................................... 291
Budding en maturatie.................................................................................................................. 294
Antivirale middelen ......................................................................................................................... 295
Limitaties ..................................................................................................................................... 296
Preventiemethoden..................................................................................................................... 296




7

,Algemene introductie virologie
Geschiedenis
Er zijn aanwijzingen dat virussen al vanaf 1500 jaar voor Christus bestaan, maar pas sinds iets meer
dan een eeuw zijn virussen voor het eerst ontdekt. Deze ontdekking zorgde ook voor het ontwikkelen
van behandelingen voor virussen, waardoor het aantal sterfgevallen door virussen de laatste
decennia sterk afgenomen is. De founding fathers van de virologie zijn Dmitri Iosifovich Ivanofski
(1864-1920) en Martinus Willem Beijerinck (1851-1931).

Ivanosfki
De Latijnse vertaling van virus is gif (toxine). Deze vertaling is ontstaan naar aanleiding van laesies in
een tabaksblad, waarvan men zich afvroeg of deze ontstaan zijn door gif of infectieuze agentia.
Hiertoe werd gebruik gemaakt van porseleinen filtersystemen (Chamberland filter), waarbij de
poriën zo klein zijn dat bacteriën tegengehouden worden. Hierdoor zou het filtraat steriel moeten
zijn. Dit filtraat werd vervolgens gebruikt om een nieuw tabaksblad te inoculeren. Hierdoor
ontstonden opnieuw laesies, waaruit bleek dat de laesies dus veroorzaakt worden door iets kleiner
dan bacteriën. Het tabaksmozaïekvirus is dus geen bacteriële infectie.

Beijerinck
Beijerinck herhaalde het experiment van Ivanofski, meerdere keren met het extract van het nieuw
geïnfecteerde tabaksblad. Dit wordt ook wel een seriële passage genoemd, waarbij het extract als
het ware steeds verdund wordt. Hieruit volgt dus dat het geen gif zou zijn, omdat dit uitgedund zou
moeten raken. Het extract bevat dus iets dat in staat is om zich te repliceren en kleiner is dan
bacteriën. Beijerinck concludeerde vervolgens dat het een vloeibare levensvorm (contagium vivum
fluidum) was.

Loeffler en Frosch
Loeffler en Frosch waren de eerste ontdekkers van een animaal virus, namelijk het mond- en
klauwzeervirus. Hierbij maakten zij gebruik van filters met nog kleinere poriën, waardoor zelfs de
kleinste virussen niet kunnen passeren.




Figuur 1 Schematische weergave van de werking van Chamberland en Berkefeld filters.




8

, Walter Reed en Jesse William Lazear
Walter Reed ontdekte het eerste humane virus, namelijk het gele koorts virus. Hiervoor werden
humane vrijwilligers gebruikt, waarbij het virus geïnfecteerd werd met behulp van de
eerdergenoemde methoden. Jesse William Lazear ontdekte later dat het virus overgedragen wordt
door muggen. Deze virussen worden ook wel Arthropod-borne (ARBO) virussen genoemd. Later
werd ontdekt dat dit type virussen onder controle kan worden gebracht door reductie van de
muggenpopulatie (vector).

Na de ontwikkeling van de elektronenmicroscoop konden virussen bestudeerd worden. Hieruit kon
men afleiden dat virussen een a-cellulaire structuur hebben. Daarnaast komen virussen voor in alle
organismen. Er is een enorme verscheidenheid aan virussen en een grote variatie aan
replicatiemechanismen.

Definitie virus
Een virus voldoet aan een aantal criteria, waaronder:
• Kleine infectieuze agentia met een a-cellulaire organisatie.
➢ Virussen zijn geen organismen.
• Reproduceren enkel in levende cellen.
➢ Virussen worden geproduceerd in levende cellen.
➢ Virussen zijn dus obligaat intracellulaire parasieten.
• Bestaan uit nucleïnezuren voor het genoom (DNA of RNA), dat beschermd wordt door een
eiwitmantel (naakte virussen).
➢ Sommige virussen hebben hieromheen nog een membraan dat uit lipiden bestaat, dit is
de membraan enveloppe (membraan omvattende virussen). Hierin bevinden zich virus-
gecodeerde eiwitten, die doorgaans uitsteken, waardoor ze spikes genoemd worden.
• Worden overgedragen van de ene gastheer naar de andere gastheer via een virion.




Figuur 2 Schematische weergave van de structuur van een virus.




9

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