300:
257:
1314:
126:
291:(ACE2). The evolved, high level of activity to mediate cell to cell fusion has resulted in an enhanced fusion capacity. Current prophylaxis against SARS-2 infection targets the spike (S) proteins that harbor the capacity for membrane fusion. Vaccinations are based on the blocking the viral S glycoprotein with the cell, thus stopping the fusion of the virus and its host cell membranes. The fusion mechanism is also studied as a potential target for antiviral development.
107:
1562:
32:
1550:
1586:
1574:
334:, also enter the cell through endocytosis. Entry via the endosome guarantees low pH and exposure to proteases which are needed to open the viral capsid and release the genetic material inside the host cytoplasm. Further, endosomes transport the virus through the cell and ensure that no trace of the virus is left on the surface, which could otherwise trigger immune recognition by the host.
150:. This attachment causes the two membranes to remain in mutual proximity, favoring further interactions between surface proteins. This is also the first requisite that must be satisfied before a cell can become infected. Satisfaction of this requisite makes the cell susceptible. Viruses that exhibit this behavior include many enveloped viruses such as
268:, viral receptors attach to the receptors on the surface of the cell and secondary receptors may be present to initiate the puncture of the membrane or fusion with the host cell. Following attachment, the viral envelope fuses with the host cell membrane, causing the virus to enter. Viruses that enter a cell in this manner included
311:; they “trick” the host cell to ingest the virions through the cell membrane. Cells can take in resources from the environment outside of the cell, and these mechanisms may be exploited by viruses to enter a cell in the same manner as ordinary resources. Once inside the cell, the virus leaves the host
366:
Once a virus is in a cell, it will activate formation of proteins (either by itself or using the host’s machinery) to gain full control of the host cell, if possible. Control mechanisms include the suppression of intrinsic cell defenses, suppression of cell signaling and suppression of host cellular
137:
Cell entry by enveloped viruses is more complicated. Enveloped viruses enter the cell by attaching to an attachment factor located on the surface of the host cell. They then enter by endocytosis or a direct membrane fusion event. The fusion event is when the virus membrane and the host cell membrane
138:
fuse together allowing a virus to enter. It does this by attachment – or adsorption – onto a susceptible cell; a cell which holds a receptor that the virus can bind to, akin to two pieces of a puzzle fitting together. The
83:
and introduces viral material into the cell. The major steps involved in viral entry are shown below. Despite the variation among viruses, there are several shared generalities concerning viral entry.
1013:
Outlaw, Victor K.; Bovier, Francesca T.; Mears, Megan C.; Cajimat, Maria N.; Zhu, Yun; Lin, Michelle J.; Addetia, Amin; Lieberman, Nicole A. P.; Peddu, Vikas; Xie, Xuping; Shi, Pei-Yong (2020-10-20).
1166:
Sebestyén, Magdolna G.; Budker, Vladimir G.; Budker, Tatiana; Subbotin, Vladimir M.; Zhang, Guofeng; Monahan, Sean D.; Lewis, David L.; Wong, So C.; Hagstrom, James E.; Wolff, Jon A. (July 2006).
958:"Inhibition of SARS-CoV-2 (previously 2019-nCoV) infection by a highly potent pan-coronavirus fusion inhibitor targeting its spike protein that harbors a high capacity to mediate membrane fusion"
248:(GFP), virus entry and infection can be visualized in real-time. Once a virus enters a cell, replication is not immediate and indeed takes some time (seconds to hours).
677:
Kumar, Binod; Dutta, Dipanjan; Iqbal, Jawed; Ansari, Mairaj Ahmed; Roy, Arunava; Chikoti, Leela; Pisano, Gina; Veettil, Mohanan Valiya; Chandran, Bala (October 2016).
378:
A cell is classified as susceptible to a virus if the virus is able to enter the cell. After the introduction of the viral particle, unpacking of the contents (viral
346:
into the cell, leaving the rest of the virus on the surface. This is restricted to viruses in which only the genome is required for infection of a cell (for example
350:
because they can be immediately translated) and is further restricted to viruses that actually exhibit this behavior. The best studied example includes the
736:
Veettil, Mohanan Valiya; Kumar, Binod; Ansari, Mairaj Ahmed; Dutta, Dipanjan; Iqbal, Jawed; Gjyshi, Olsi; Bottero, Virginie; Chandran, Bala (April 2016).
679:"ESCRT-I Protein Tsg101 Plays a Role in the Post-macropinocytic Trafficking and Infection of Endothelial Cells by Kaposi's Sarcoma-Associated Herpesvirus"
956:
Xia, Shuai; Liu, Meiqin; Wang, Chao; Xu, Wei; Lan, Qiaoshuai; Feng, Siliang; Qi, Feifei; Bao, Linlin; Du, Lanying; Liu, Shuwen; Qin, Chuan (2020-03-30).
193:
bilayer, a cell's natural barrier to the outside world. The process by which this barrier is breached depends upon the virus. Types of entry are:
852:
Campadelli-Fiume, Gabriella; Amasio, Michele; Avitabile, Elisa; Cerretani, Arianna; Forghieri, Cristina; Gianni, Tatiana; Menotti, Laura (2007).
1015:"Inhibition of Coronavirus Entry In Vitro and Ex Vivo by a Lipid-Conjugated Peptide Derived from the SARS-CoV-2 Spike Glycoprotein HRC Domain"
287:, either at the cell surface or in vesicles. Research efforts have focused on the spike protein's interaction with its cell-surface receptor,
220:: The host cell takes in the viral particle through the process of endocytosis, essentially engulfing the virus like it would a food particle.
738:"ESCRT-0 Component Hrs Promotes Macropinocytosis of Kaposi's Sarcoma-Associated Herpesvirus in Human Dermal Microvascular Endothelial Cells"
165:(or simply phages). Typical phages have long tails used to attach to receptors on the bacterial surface and inject their viral genome.
416:"Herpes simplex virus type 1 mediates fusion through a hemifusion intermediate by sequential activity of glycoproteins D, H, L, and B"
795:"Protective Immunity Based on the Conserved Hemagglutinin Stalk Domain and Its Prospects for Universal Influenza Vaccine Development"
358:
land on a cell, its central sheath pierces the cell membrane and the phage injects DNA from the head capsid directly into the cell.
1480:
342:
A third method is by simply attaching to the surface of the host cell via receptors on the cell with the virus injecting only its
1238:
199:
909:"Design of Potent Membrane Fusion Inhibitors against SARS-CoV-2, an Emerging Coronavirus with High Fusogenic Activity"
315:
by which it was taken up and thus gains access to the cytoplasm. Examples of viruses that enter this way include the
312:
288:
173:
Prior to entry, a virus must attach to a host cell. Attachment is achieved when specific proteins on the viral
1168:"Mechanism of plasmid delivery by hydrodynamic tail vein injection. I. Hepatocyte uptake of various molecules"
1590:
620:"KSHV Entry and Trafficking in Target Cells-Hijacking of Cell Signal Pathways, Actin and Membrane Dynamics"
324:
99:
enters the cell by attaching to the attachment factor located on a host cell. It then enters the cell by
1231:
1617:
1272:
347:
284:
245:
1578:
368:
1267:
355:
1473:
372:
210:: The cell membrane is punctured and made to further connect with the unfolding viral envelope.
1070:
Tang, Tiffany; Bidon, Miya; Jaimes, Javier A.; Whittaker, Gary R.; Daniel, Susan (June 2020).
375:. Often, these cytotoxic effects lead to the death and decline of a cell infected by a virus.
1224:
92:
How a virus enters a cell is different depending on the type of virus it is. A virus with a
1525:
1520:
1072:"Coronavirus membrane fusion mechanism offers a potential target for antiviral development"
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The most well-known example is through membrane fusion. In a number of viruses with a
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or by making a hole in the membrane of the host cell and inserting its viral genome.
1566:
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1322:
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1026:
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854:"The multipartite system that mediates entry of herpes simplex virus into the cell"
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130:
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508:
Proceedings of the
National Academy of Sciences of the United States of America
502:
Lakadamyali, Melike; Michael J. Rust; Hazen P. Babcock; Xiaowei Zhuang (2003).
383:
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93:
80:
58:
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of the target cell. A virus must now enter the cell, which is covered by a
1030:
907:
Zhu, Yuanmei; Yu, Danwei; Yan, Hongxia; Chong, Huihui; He, Yuxian (2020).
811:
1458:
1437:
1355:
924:
753:
563:"Visualization of Targeted Transduction by Engineered Lentiviral Vectors"
308:
215:
125:
119:
115:
100:
851:
106:
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578:
331:
316:
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1216:
1495:
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1345:
1167:
853:
793:
Khanna, Madhu; Sharma, Sachin; Kumar, Binod; Rajput, Roopali (2014).
636:
237:
22:
1183:
869:
31:
1277:
478:
379:
161:
These basic ideas extend to viruses that infect bacteria, known as
1126:
Helle F, Dubuisson J. "Hepatitis C virus entry into host cells."
307:
Viruses with no viral envelope enter the cell generally through
1530:
1293:
387:
343:
233:
229:
174:
146:
effectively become connected to complementary receptors on the
96:
394:) occurs as preparation of the next stage of viral infection:
1251:
1165:
76:
413:
269:
151:
1069:
1012:
279:
In SARS-CoV-2 and similar viruses, entry occurs through
792:
735:
504:"Visualizing infection of individual influenza viruses"
676:
420:
Proceedings of the
National Academy of Sciences, USA
110:
Schematic of different pathways of viral entry: (A)
407:
16:
Earliest stage of infection in the viral life cycle
618:Kumar, Binod; Chandran, Bala (November 14, 2016).
1604:
414:Subramanian RP, Geraghty RJ (20 February 2007).
337:
87:
1232:
906:
617:
479:"Virus entry into host cell ~ ViralZone page"
955:
251:
1143:, 6th edition." Blackwell Publishing, 2007.
354:; for example, when the tail fibers of the
1239:
1225:
71:is the earliest stage of infection in the
1103:
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989:
932:
828:
810:
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527:
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439:
1481:Laboratory diagnosis of viral infections
298:
294:
255:
124:
105:
1246:
1156:Lippincott Williams & Williams 2013
1605:
1220:
1573:
473:
471:
469:
1585:
13:
30:
14:
1629:
466:
236:is injected into the host cell's
181:bind to specific proteins called
79:comes into contact with the host
1584:
1572:
1561:
1560:
1548:
1312:
1152:Howley, Peter M; Knipe, David M
330:Many enveloped viruses, such as
1159:
1146:
1141:Introduction to Modern Virology
1133:
1120:
1088:10.1016/j.antiviral.2020.104792
1063:
1006:
949:
900:
561:Joo, K-I; P Wang (2008-05-15).
289:angiotensin-converting enzyme 2
260:Viral entry via membrane fusion
845:
786:
729:
670:
611:
554:
495:
1:
799:BioMed Research International
401:
1172:The Journal of Gene Medicine
696:10.1371/journal.ppat.1005960
325:foot-and-mouth disease virus
129:Membrane fusion mediated by
7:
858:Reviews in Medical Virology
361:
348:positive-strand RNA viruses
338:Entry via genetic injection
303:Viral entry via endocytosis
168:
88:Reducing cellular proximity
10:
1634:
1544:
1446:
1405:
1369:
1321:
1310:
1286:
1273:Social history of viruses
1258:
974:10.1038/s41422-020-0305-x
252:Entry via membrane fusion
246:green fluorescent protein
529:10.1073/pnas.0832269100
441:10.1073/pnas.0608374104
1474:Helper dependent virus
304:
261:
134:
122:
35:
1031:10.1128/mBio.01935-20
302:
295:Entry via endocytosis
259:
128:
109:
34:
1526:Virus quantification
1521:Virus classification
1139:N.J. Dimmock et al.
925:10.1128/JVI.00635-20
754:10.1128/JVI.02704-15
483:viralzone.expasy.org
274:herpes simplex virus
156:herpes simplex virus
1516:Virus-like particle
913:Journal of Virology
812:10.1155/2014/546274
742:Journal of Virology
520:2003PNAS..100.9280L
432:2007PNAS..104.2903S
244:Through the use of
1076:Antiviral Research
579:10.1038/gt.2008.87
305:
262:
135:
123:
36:
1600:
1599:
1491:Neurotropic virus
1336:Viral replication
1128:Cell Mol Life Sci
396:viral replication
390:via some form of
321:hepatitis C virus
225:Viral penetration
66:
65:
1625:
1618:Viral life cycle
1588:
1587:
1576:
1575:
1564:
1563:
1552:
1387:Phenotype mixing
1323:Viral life cycle
1316:
1241:
1234:
1227:
1218:
1217:
1212:
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1157:
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1010:
1004:
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936:
904:
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843:
842:
832:
814:
790:
784:
783:
773:
733:
727:
726:
716:
698:
689:(10): e1005960.
674:
668:
667:
657:
639:
637:10.3390/v8110305
615:
609:
608:
598:
558:
552:
551:
541:
531:
499:
493:
492:
490:
489:
475:
464:
463:
453:
443:
411:
283:mediated by the
207:Hemifusion state
185:proteins on the
120:macropinocytosis
73:viral life cycle
25:virus life cycle
19:
18:
1633:
1632:
1628:
1627:
1626:
1624:
1623:
1622:
1603:
1602:
1601:
1596:
1540:
1442:
1401:
1397:Viral evolution
1382:Antigenic shift
1377:Antigenic drift
1365:
1361:Lysogenic cycle
1317:
1308:
1282:
1254:
1245:
1215:
1184:10.1002/jgm.921
1164:
1160:
1154:Fields Virology
1151:
1147:
1138:
1134:
1125:
1121:
1068:
1064:
1011:
1007:
954:
950:
905:
901:
870:10.1002/rmv.546
850:
846:
791:
787:
734:
730:
675:
671:
616:
612:
573:(20): 1384–96.
559:
555:
514:(16): 9280–85.
500:
496:
487:
485:
477:
476:
467:
412:
408:
404:
364:
340:
297:
281:membrane fusion
254:
200:Membrane fusion
171:
133:fusion proteins
112:membrane fusion
90:
17:
12:
11:
5:
1631:
1621:
1620:
1615:
1598:
1597:
1595:
1594:
1582:
1570:
1558:
1545:
1542:
1541:
1539:
1538:
1533:
1528:
1523:
1518:
1513:
1508:
1503:
1498:
1493:
1488:
1486:Marine viruses
1483:
1478:
1477:
1476:
1466:
1461:
1456:
1454:Antiviral drug
1450:
1448:
1444:
1443:
1441:
1440:
1435:
1430:
1425:
1420:
1415:
1409:
1407:
1403:
1402:
1400:
1399:
1394:
1389:
1384:
1379:
1373:
1371:
1367:
1366:
1364:
1363:
1358:
1353:
1348:
1343:
1341:Viral shedding
1338:
1333:
1327:
1325:
1319:
1318:
1311:
1309:
1307:
1306:
1301:
1299:Viral envelope
1296:
1290:
1288:
1284:
1283:
1281:
1280:
1275:
1270:
1265:
1259:
1256:
1255:
1244:
1243:
1236:
1229:
1221:
1214:
1213:
1178:(7): 852–873.
1158:
1145:
1132:
1119:
1062:
1005:
968:(4): 343–355.
948:
899:
864:(5): 313–326.
844:
785:
748:(8): 3860–72.
728:
683:PLOS Pathogens
669:
610:
553:
494:
465:
426:(8): 2903–08.
405:
403:
400:
386:and the viral
363:
360:
352:bacteriophages
339:
336:
296:
293:
266:viral envelope
253:
250:
242:
241:
221:
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179:viral envelope
170:
167:
163:bacteriophages
144:viral envelope
89:
86:
64:
63:
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61:
56:
51:
46:
38:
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27:
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15:
9:
6:
4:
3:
2:
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1532:
1529:
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1509:
1507:
1506:Viral disease
1504:
1502:
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1449:
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1431:
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1426:
1424:
1421:
1419:
1418:Bacteriophage
1416:
1414:
1411:
1410:
1408:
1404:
1398:
1395:
1393:
1390:
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1383:
1380:
1378:
1375:
1374:
1372:
1368:
1362:
1359:
1357:
1354:
1352:
1351:Virus latency
1349:
1347:
1344:
1342:
1339:
1337:
1334:
1332:
1329:
1328:
1326:
1324:
1320:
1315:
1305:
1304:Viral protein
1302:
1300:
1297:
1295:
1292:
1291:
1289:
1285:
1279:
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1271:
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1077:
1073:
1066:
1058:
1054:
1049:
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1036:
1032:
1028:
1024:
1020:
1016:
1009:
1001:
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992:
987:
983:
979:
975:
971:
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963:
962:Cell Research
959:
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859:
855:
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836:
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813:
808:
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789:
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772:
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568:
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557:
549:
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525:
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498:
484:
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470:
461:
457:
452:
447:
442:
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429:
425:
421:
417:
410:
406:
399:
397:
393:
389:
385:
381:
376:
374:
370:
369:transcription
359:
357:
353:
349:
345:
335:
333:
328:
326:
322:
318:
314:
310:
301:
292:
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286:
285:spike protein
282:
277:
275:
271:
267:
258:
249:
247:
239:
235:
231:
227:
226:
222:
219:
218:
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208:
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202:
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196:
195:
194:
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187:cell membrane
184:
180:
176:
166:
164:
159:
157:
153:
149:
148:cell membrane
145:
141:
132:
131:paramyxovirus
127:
121:
117:
113:
108:
104:
102:
98:
95:
85:
82:
78:
74:
70:
60:
57:
55:
52:
50:
47:
45:
42:
41:
40:
39:
33:
29:
28:
24:
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20:
1589:
1577:
1565:
1553:
1469:Viral vector
1464:Helper virus
1428:Human virome
1413:Animal virus
1392:Reassortment
1330:
1268:Introduction
1248:Microbiology
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1130:. 2007 Oct 4
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486:. Retrieved
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392:nucleic acid
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228:: The viral
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191:phospholipid
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160:
136:
94:nonenveloped
91:
68:
67:
43:
1591:WikiProject
1459:Giant virus
1438:Plant virus
1356:Lytic cycle
1331:Viral entry
630:(11): 305.
373:translation
309:endocytosis
272:, KSHV and
216:Endocytosis
116:endocytosis
101:endocytosis
69:Viral entry
49:Replication
1607:Categories
1536:Virosphere
1511:Viral load
1501:Satellites
1287:Components
1082:: 104792.
805:: 546274.
488:2021-02-05
402:References
332:SARS-CoV-2
317:poliovirus
118:, and (C)
1496:Oncovirus
1433:Mycovirus
1423:Virophage
1346:Viroplasm
1192:1099-498X
1096:0166-3542
1039:2150-7511
982:1001-0602
878:1052-9276
821:2314-6133
762:1098-5514
705:1553-7374
646:1999-4915
587:0969-7128
567:Gene Ther
238:cytoplasm
140:receptors
75:, as the
23:Influenza
1613:Virology
1567:Category
1370:Genetics
1278:Virology
1200:16724360
1114:32272173
1057:33082259
1000:32231345
943:32376627
894:30771615
886:17573668
839:24982895
780:26819309
723:27764233
664:27854239
605:18480844
548:12883000
460:17299053
384:tegument
380:proteins
362:Outcomes
356:T2 phage
183:receptor
169:Overview
59:Shedding
1579:Commons
1406:By host
1263:History
1105:7194977
1048:7587434
991:7104723
934:7343218
830:4055638
771:4810545
714:5072609
655:5127019
624:Viruses
596:2575058
516:Bibcode
451:1815279
428:Bibcode
382:in the
313:vesicle
142:on the
54:Latency
1555:Portal
1531:Virome
1294:Capsid
1208:564796
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388:genome
344:genome
323:, and
234:genome
230:capsid
175:capsid
114:, (B)
97:capsid
1447:Other
1252:Virus
1204:S2CID
1025:(5).
890:S2CID
77:virus
44:Entry
1196:PMID
1188:ISSN
1110:PMID
1092:ISSN
1053:PMID
1035:ISSN
1019:mBio
996:PMID
978:ISSN
939:PMID
882:PMID
874:ISSN
835:PMID
817:ISSN
803:2014
776:PMID
758:ISSN
719:PMID
701:ISSN
660:PMID
642:ISSN
601:PMID
583:ISSN
544:PMID
456:PMID
371:and
154:and
81:cell
1180:doi
1100:PMC
1084:doi
1080:178
1043:PMC
1027:doi
986:PMC
970:doi
929:PMC
921:doi
866:doi
825:PMC
807:doi
766:PMC
750:doi
709:PMC
691:doi
650:PMC
632:doi
591:PMC
575:doi
534:PMC
524:doi
512:100
446:PMC
436:doi
424:104
270:HIV
232:or
204:or
177:or
152:HIV
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