42:
207:
2093:
594:. For non-enveloped viruses, the capsid itself may be involved in interaction with receptors on the host cell, leading to penetration of the host cell membrane and internalization of the capsid. Delivery of the genome occurs by subsequent uncoating or disassembly of the capsid and release of the genome into the cytoplasm, or by ejection of the genome through a specialized portal structure directly into the host cell nucleus.
520:
196:
2341:
2329:
466:
2365:
2353:
564:
interior of the helix bind three nucleotides of the RNA genome. Influenza A viruses differ by comprising multiple ribonucleoproteins, the viral NP protein organizes the RNA into a helical structure. The size is also different; the tobacco mosaic virus has a 16.33 protein subunits per helical turn, while the influenza A virus has a 28 amino acid tail loop.
31:
627:
539:-fold axial symmetry. The helical transformation are classified into two categories: one-dimensional and two-dimensional helical systems. Creating an entire helical structure relies on a set of translational and rotational matrices which are coded in the protein data bank. Helical symmetry is given by the formula
614:
between replicator communities since these communities could not survive if the number of gene parasites increased, with certain genes being responsible for the formation of these structures and those that favored the survival of self-replicating communities. The displacement of these ancestral genes
150:
mechanism of the cell. In some viruses, including those with helical capsids and especially those with RNA genomes, the capsid proteins co-assemble with their genomes. In other viruses, especially more complex viruses with double-stranded DNA genomes, the capsid proteins assemble into empty precursor
602:
It has been suggested that many viral capsid proteins have evolved on multiple occasions from functionally diverse cellular proteins. The recruitment of cellular proteins appears to have occurred at different stages of evolution so that some cellular proteins were captured and refunctionalized prior
413:
and bacteriophage φ6 have capsids built of 120 copies of capsid protein, corresponding to a T = 2 capsid, or arguably a T = 1 capsid with a dimer in the asymmetric unit. Similarly, many small viruses have a pseudo T = 3 (or P = 3) capsid, which is organized according to a T = 3 lattice, but with
563:
is the pitch of the helix. The structure is said to be open due to the characteristic that any volume can be enclosed by varying the length of the helix. The most understood helical virus is the tobacco mosaic virus. The virus is a single molecule of (+) strand RNA. Each coat protein on the
603:
to the divergence of cellular organisms into the three contemporary domains of life, whereas others were hijacked relatively recently. As a result, some capsid proteins are widespread in viruses infecting distantly related organisms (e.g., capsid proteins with the
41:
476:
An elongated icosahedron is a common shape for the heads of bacteriophages. Such a structure is composed of a cylinder with a cap at either end. The cylinder is composed of 10 elongated triangular faces. The Q number (or
218:
consists of 20 triangular faces delimited by 12 fivefold vertexes and consists of 60 asymmetric units. Thus, an icosahedral virus is made of 60N protein subunits. The number and arrangement of
481:), which can be any positive integer, specifies the number of triangles, composed of asymmetric subunits, that make up the 10 triangles of the cylinder. The caps are classified by the T (or T
365:
112:, have developed more complicated structures due to constraints of elasticity and electrostatics. The icosahedral shape, which has 20 equilateral triangular faces, approximates a
391:
162:
Structural analyses of major capsid protein (MCP) architectures have been used to categorise viruses into lineages. For example, the bacteriophage PRD1, the algal virus
292:
266:
500:
chaperone protein GroES and able to substitute for it in the assembly of bacteriophage T4 virions during infection. Like GroES, gp31 forms a stable complex with
45:
Illustration of geometric model changing between two possible capsids. A similar change of size has been observed as the result of a single amino-acid mutation
1564:
Yamada S, Matsuzawa T, Yamada K, Yoshioka S, Ono S, Hishinuma T (December 1986). "Modified inversion recovery method for nuclear magnetic resonance imaging".
586:
The virus must assemble a stable, protective protein shell to protect the genome from lethal chemical and physical agents. These include extremes of
2259:
405:
have pentamers instead of hexamers in hexavalent positions on a quasi T = 7 lattice. Members of the double-stranded RNA virus lineage, including
1666:"Biochemical and structural evidence in support of a coherent model for the formation of the double-helical influenza A virus ribonucleoprotein"
2017:
135:. The envelope is acquired by the capsid from an intracellular membrane in the virus' host; examples include the inner nuclear membrane, the
1923:
1904:
1648:
1531:
1336:
921:
672:"A Selection for Assembly Reveals That a Single Amino Acid Mutant of the Bacteriophage MS2 Coat Protein Forms a Smaller Virus-like Particle"
234:, an icosahedral structure can be regarded as being constructed from pentamers and hexamers. The structures can be indexed by two integers
120:, taking the space of a cylinder but not being a cylinder itself. The capsid faces may consist of one or more proteins. For example, the
1779:
1083:"Three RNA cells for ribosomal lineages and three DNA viruses to replicate their genomes: a hypothesis for the origin of cellular domain"
1430:
Damodaran KV, Reddy VS, Johnson JE, Brooks CL (December 2002). "A general method to quantify quasi-equivalence in icosahedral viruses".
645:
2369:
1549:
Marusich EI, Kurochkina LP, Mesyanzhinov VV. Chaperones in bacteriophage T4 assembly. Biochemistry (Mosc). 1998;63(4):399-406
1830:"Chasing the Origin of Viruses: Capsid-Forming Genes as a Life-Saving Preadaptation within a Community of Early Replicators"
937:
610:
A computational model (2015) has shown that capsids may have originated before viruses and that they served as a means of
73:. The observable 3-dimensional morphological subunits, which may or may not correspond to individual proteins, are called
70:
1038:
Krupovic M, Bamford DH (December 2008). "Virus evolution: how far does the double beta-barrel viral lineage extend?".
1918:. University of California Press. Chapter 6. The Geodesic Polyhedra of R. Buckminster Fuller and Related Polyhedra.
2396:
2010:
1892:
496:
that has a prolate head structure. The bacteriophage encoded gp31 protein appears to be functionally homologous to
100:
Capsids are broadly classified according to their structure. The majority of the viruses have capsids with either
1268:
2051:
146:
Once the virus has infected a cell and begins replicating itself, new capsid subunits are synthesized using the
1267:
Carrillo-Tripp M, Shepherd CM, Borelli IA, Venkataraman S, Lander G, Natarajan P, et al. (January 2009).
1899:. Dover Publications. pp. 142–144, Figures 4-49, 50, 51: Custers of 12 spheres, 42 spheres, 92 spheres.
1142:"Structure of an archaeal virus capsid protein reveals a common ancestry to eukaryotic and bacterial viruses"
2046:
450:
312:
733:
Lidmar J, Mirny L, Nelson DR (November 2003). "Virus shapes and buckling transitions in spherical shells".
607:), whereas others are restricted to a particular group of viruses (e.g., capsid proteins of alphaviruses).
378:-number is representative of the size and complexity of the capsids. Geometric examples for many values of
2252:
434:
121:
2003:
1986:
454:
442:
670:
Asensio MA, Morella NM, Jakobson CM, Hartman EC, Glasgow JE, Sankaran B, et al. (September 2016).
2304:
2299:
1841:
1729:
1478:
1365:
1094:
866:
807:
752:
683:
493:
147:
2357:
2294:
2279:
2041:
1566:
The
Science Reports of the Research Institutes, Tohoku University. Ser. C, Medicine. Tohoku Daigaku
640:
611:
528:
504:
446:
2328:
1810:
1621:
1063:
776:
742:
715:
231:
222:
in an icosahedral capsid can be classified using the "quasi-equivalence principle" proposed by
2391:
2269:
2114:
1964:
1919:
1900:
1869:
1802:
1757:
1695:
1644:
1613:
1573:
1527:
1504:
1447:
1393:
1332:
1298:
1249:
1214:
1173:
1122:
1055:
1020:
975:
917:
894:
835:
768:
707:
699:
438:
271:
245:
2345:
2165:
2101:
1954:
1946:
1859:
1849:
1794:
1747:
1737:
1685:
1677:
1605:
1494:
1486:
1439:
1383:
1373:
1288:
1280:
1241:
1204:
1163:
1153:
1112:
1102:
1047:
1010:
1002:
884:
874:
825:
815:
760:
691:
402:
58:
2175:
2160:
2155:
2139:
2092:
1990:
963:
604:
176:
have been placed in the same lineage, whereas tailed, double-stranded DNA bacteriophages (
136:
117:
35:
1412:
1933:
Almansour I, Alhagri M, Alfares R, Alshehri M, Bakhashwain R, Maarouf A (January 2019).
1845:
1733:
1482:
1369:
1232:
Caspar DL, Klug A (1962). "Physical principles in the construction of regular viruses".
1098:
870:
811:
756:
687:
519:
195:
2333:
2264:
2232:
2119:
2077:
1959:
1934:
1864:
1829:
1752:
1717:
1690:
1665:
1609:
1499:
1466:
1388:
1353:
1293:
1269:"VIPERdb2: an enhanced and web API enabled relational database for structural virology"
1168:
1141:
1117:
1082:
1015:
1006:
990:
889:
854:
830:
795:
632:
615:
between cellular organisms could favor the appearance of new viruses during evolution.
132:
1443:
206:
155:
that include a specialized portal structure at one vertex. Through this portal, viral
2385:
2284:
2196:
2129:
2082:
1814:
968:
470:
223:
140:
109:
1625:
1067:
991:"Involvement of the portal at an early step in herpes simplex virus capsid assembly"
719:
298:
steps from the edge of a pentamer, turning 60 degrees counterclockwise, then taking
2247:
2242:
2206:
2191:
2170:
2026:
780:
426:
398:
200:
178:
173:
1854:
1378:
1245:
695:
2237:
2216:
2134:
2109:
1354:"Periodic table of virus capsids: implications for natural selection and design"
941:
422:
215:
164:
105:
1722:
Proceedings of the
National Academy of Sciences of the United States of America
1596:
Aldrich RA (February 1987). "Children in cities--Seattle's KidsPlace program".
1193:"Membrane proteins modulate the bilayer curvature in the bacterial virus Bam35"
1146:
Proceedings of the
National Academy of Sciences of the United States of America
1140:
Khayat R, Tang L, Larson ET, Lawrence CM, Young M, Johnson JE (December 2005).
1087:
Proceedings of the
National Academy of Sciences of the United States of America
859:
Proceedings of the
National Academy of Sciences of the United States of America
800:
Proceedings of the
National Academy of Sciences of the United States of America
764:
671:
2314:
2289:
1950:
1798:
1490:
1209:
1192:
622:
227:
2274:
2211:
2201:
2124:
1742:
1664:
Ye Q, Guu TS, Mata DA, Kuo RL, Smith B, Krug RM, Tao YJ (26 December 2012).
1158:
1107:
879:
820:
650:
410:
219:
169:
74:
1968:
1873:
1806:
1761:
1699:
1508:
1451:
1397:
1302:
1253:
1218:
1177:
1126:
1059:
1024:
898:
839:
772:
711:
465:
1681:
1617:
1577:
2056:
1780:"Origin of viruses: primordial replicators recruiting capsids from hosts"
1284:
855:"Platonic and Archimedean geometries in multicomponent elastic membranes"
747:
406:
62:
1897:
The
Geometrical Foundation of Natural Structure: A Source Book of Design
1051:
1306:
131:, meaning that the capsid is coated with a lipid membrane known as the
66:
17:
1995:
703:
626:
414:
distinct polypeptides occupying the three quasi-equivalent positions
30:
2309:
1191:
Laurinmäki PA, Huiskonen JT, Bamford DH, Butcher SJ (December 2005).
591:
113:
90:
1718:"Multiple origins of viral capsid proteins from cellular ancestors"
2030:
518:
501:
464:
205:
194:
101:
54:
40:
29:
370:
In this scheme, icosahedral capsids contain 12 pentamers plus 10(
214:
The icosahedral structure is extremely common among viruses. The
124:
virus capsid has faces consisting of three proteins named VP1–3.
527:
Many rod-shaped and filamentous plant viruses have capsids with
1999:
156:
1983:
417:
T-numbers can be represented in different ways, for example
302:
steps to get to the next pentamer. The triangulation number
587:
1939:
Database: The
Journal of Biological Databases and Curation
853:
Vernizzi G, Sknepnek R, Olvera de la Cruz M (March 2011).
796:"Faceting ionic shells into icosahedra via electrostatics"
507:
that is absolutely necessary for the folding and assembly
555:
is the number of structural units per turn of the helix,
531:. The helical structure can be described as a set of
397:
Many exceptions to this rule exist: For example, the
315:
274:
248:
2225:
2184:
2148:
2100:
2065:
1935:"IRAM: virus capsid database and analysis resource"
1234:
Cold Spring Harbor
Symposia on Quantitative Biology
511:of the bacteriophage T4 major capsid protein gp23.
116:, while the helical shape resembles the shape of a
1591:
1589:
1587:
967:
359:
286:
260:
1641:Principles of Virology, Vol. 1: Molecular Biology
1559:
1557:
1555:
794:Vernizzi G, Olvera de la Cruz M (November 2007).
523:3D model of a helical capsid structure of a virus
392:List of geodesic polyhedra and Goldberg polyhedra
1984:IRAM-Virus Capsid Database and Analysis Resource
1773:
1771:
1711:
1709:
1545:
1543:
962:Alberts B, Bray D, Lewis J, Raff M, Roberts K,
590:or temperature and proteolytic and nucleolytic
77:. The proteins making up the capsid are called
429:and, depending on the type of quasi-symmetry,
182:) and herpesvirus belong to a second lineage.
2011:
1778:Krupovic M, Dolja VV, Koonin EV (July 2019).
989:Newcomb WW, Homa FL, Brown JC (August 2005).
469:The prolate structure of a typical head on a
421: = 1 can only be represented as an
8:
1828:Jalasvuori M, Mattila S, Hoikkala V (2015).
1526:. Boston: Academic Press. pp. 167–174.
1331:. Boston: Academic Press. pp. 115–123.
294:; the structure can be thought of as taking
2018:
2004:
1996:
1467:"The structure of elongated viral capsids"
1958:
1863:
1853:
1751:
1741:
1689:
1498:
1387:
1377:
1292:
1208:
1167:
1157:
1116:
1106:
1014:
888:
878:
829:
819:
746:
351:
326:
314:
273:
247:
2260:Laboratory diagnosis of viral infections
65:(repeating) structural subunits made of
916:. New York: Garland. pp. 161–162.
662:
360:{\displaystyle T=h^{2}+h\cdot k+k^{2}}
1524:Desk Encyclopedia of General Virology
1329:Desk Encyclopedia of General Virology
433: = 3 can be presented as a
165:Paramecium bursaria Chlorella virus-1
7:
2352:
1716:Krupovic M, Koonin EV (March 2017).
1352:Mannige RV, Brooks CL (March 2010).
572:The functions of the capsid are to:
535:1-D molecular helices related by an
2364:
1639:Racaniello VR, Enquist LW (2008).
1610:10.1111/j.1442-200x.1987.tb00013.x
1419:. University of Wisconsin-Madison.
1007:10.1128/JVI.79.16.10540-10546.2005
108:structure. Some viruses, such as
25:
938:"Virus Structure (web-books.com)"
914:Introduction to Protein Structure
159:is translocated into the capsid.
2363:
2351:
2340:
2339:
2327:
2091:
1465:Luque A, Reguera D (June 2010).
1417:Institute for Molecular Virology
625:
1643:. Washington, D.C.: ASM Press.
559:is the axial rise per unit and
139:membrane, and the cell's outer
374: − 1) hexamers. The
306:for the capsid is defined as:
1:
1444:10.1016/S0022-2836(02)01138-5
1327:Johnson JE, Speir JA (2009).
970:Molecular Biology of the Cell
646:Goldberg–Coxeter construction
445:and their respective duals a
1916:Polyhedra: A Visual Approach
1855:10.1371/journal.pone.0126094
1787:Nature Reviews. Microbiology
1432:Journal of Molecular Biology
1379:10.1371/journal.pone.0009423
1246:10.1101/sqb.1962.027.001.005
1040:Nature Reviews. Microbiology
696:10.1021/acs.nanolett.6b02948
1914:Pugh A (1 September 1976).
1279:(Database issue): D436-42.
912:Branden C, Tooze J (1991).
2413:
765:10.1103/PhysRevE.68.051910
651:Fullerene#Other buckyballs
53:is the protein shell of a
2323:
2089:
2052:Social history of viruses
2037:
1799:10.1038/s41579-019-0205-6
1598:Acta Paediatrica Japonica
1491:10.1016/j.bpj.2010.02.051
1210:10.1016/j.str.2005.08.020
1081:Forterre P (March 2006).
199:Icosahedral capsid of an
61:. It consists of several
974:(4th ed.). p.
89:). The capsid and inner
27:Protein shell of a virus
1951:10.1093/database/baz079
1743:10.1073/pnas.1621061114
1159:10.1073/pnas.0506383102
1108:10.1073/pnas.0510333103
880:10.1073/pnas.1012872108
821:10.1073/pnas.0703431104
582:interact with the host.
579:deliver the genome, and
451:rhombic triacontahedron
287:{\displaystyle k\geq 0}
261:{\displaystyle h\geq 1}
2253:Helper dependent virus
1273:Nucleic Acids Research
524:
473:
435:truncated dodecahedron
361:
288:
262:
211:
210:Virus capsid T-numbers
203:
122:foot-and-mouth disease
46:
38:
1682:10.1128/mBio.00467-12
522:
468:
455:pentakis dodecahedron
443:truncated icosahedron
362:
289:
263:
209:
198:
44:
33:
2305:Virus quantification
2300:Virus classification
598:Origin and evolution
313:
272:
246:
148:protein biosynthesis
2295:Virus-like particle
1846:2015PLoSO..1026094J
1734:2017PNAS..114E2401K
1728:(12): E2401–E2410.
1483:2010BpJ....98.2993L
1471:Biophysical Journal
1370:2010PLoSO...5.9423M
1099:2006PNAS..103.3669F
1052:10.1038/nrmicro2033
995:Journal of Virology
871:2011PNAS..108.4292V
812:2007PNAS..10418382V
757:2003PhRvE..68e1910L
688:2016NanoL..16.5944A
641:Geodesic polyhedron
612:horizontal transfer
576:protect the genome,
447:triakis icosahedron
83:viral coat proteins
1989:2019-10-23 at the
1522:Casjens S (2009).
1285:10.1093/nar/gkn840
741:(5 Pt 1): 051910.
525:
474:
357:
284:
258:
232:Goldberg polyhedra
212:
204:
172:and the mammalian
47:
39:
2397:Protein complexes
2379:
2378:
2270:Neurotropic virus
2115:Viral replication
1925:978-0-520-02926-2
1906:978-0-486-23729-9
1650:978-1-55581-479-3
1533:978-0-12-375146-1
1477:(12): 2993–3003.
1338:978-0-12-375146-1
923:978-0-8153-0270-4
735:Physical Review E
439:icosidodecahedron
127:Some viruses are
16:(Redirected from
2404:
2367:
2366:
2355:
2354:
2343:
2342:
2331:
2166:Phenotype mixing
2102:Viral life cycle
2095:
2020:
2013:
2006:
1997:
1972:
1962:
1929:
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1818:
1784:
1775:
1766:
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1745:
1713:
1704:
1703:
1693:
1676:(1): e00467–12.
1661:
1655:
1654:
1636:
1630:
1629:
1593:
1582:
1581:
1561:
1550:
1547:
1538:
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1401:
1391:
1381:
1349:
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1342:
1324:
1318:
1317:
1315:
1314:
1305:. Archived from
1296:
1264:
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1229:
1223:
1222:
1212:
1188:
1182:
1181:
1171:
1161:
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1110:
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1035:
1029:
1028:
1018:
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973:
959:
953:
952:
950:
949:
940:. Archived from
934:
928:
927:
909:
903:
902:
892:
882:
850:
844:
843:
833:
823:
791:
785:
784:
750:
748:cond-mat/0306741
730:
724:
723:
667:
635:
630:
629:
529:helical symmetry
494:bacteriophage T4
492:is the host for
403:papillomaviruses
390:can be found at
366:
364:
363:
358:
356:
355:
331:
330:
293:
291:
290:
285:
267:
265:
264:
259:
59:genetic material
57:, enclosing its
21:
2412:
2411:
2407:
2406:
2405:
2403:
2402:
2401:
2382:
2381:
2380:
2375:
2319:
2221:
2180:
2176:Viral evolution
2161:Antigenic shift
2156:Antigenic drift
2144:
2140:Lysogenic cycle
2096:
2087:
2061:
2033:
2024:
1991:Wayback Machine
1980:
1975:
1932:
1926:
1913:
1907:
1895:(1 June 1979).
1891:
1887:
1885:Further reading
1882:
1881:
1840:(5): e0126094.
1827:
1826:
1822:
1782:
1777:
1776:
1769:
1715:
1714:
1707:
1663:
1662:
1658:
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1638:
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1633:
1595:
1594:
1585:
1563:
1562:
1553:
1548:
1541:
1534:
1521:
1520:
1516:
1464:
1463:
1459:
1429:
1428:
1424:
1410:
1409:
1405:
1351:
1350:
1346:
1339:
1326:
1325:
1321:
1312:
1310:
1266:
1265:
1261:
1231:
1230:
1226:
1203:(12): 1819–28.
1190:
1189:
1185:
1152:(52): 18944–9.
1139:
1138:
1134:
1093:(10): 3669–74.
1080:
1079:
1075:
1037:
1036:
1032:
1001:(16): 10540–6.
988:
987:
983:
961:
960:
956:
947:
945:
936:
935:
931:
924:
911:
910:
906:
852:
851:
847:
806:(47): 18382–6.
793:
792:
788:
732:
731:
727:
669:
668:
664:
659:
631:
624:
621:
605:jelly-roll fold
600:
570:
517:
484:
480:
463:
347:
322:
311:
310:
270:
269:
244:
243:
193:
188:
186:Specific shapes
79:capsid proteins
36:cytomegalovirus
34:Schematic of a
28:
23:
22:
15:
12:
11:
5:
2410:
2408:
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2370:WikiProject
2238:Giant virus
2217:Plant virus
2135:Lytic cycle
2110:Viral entry
423:icosahedron
230:. Like the
216:icosahedron
191:Icosahedral
106:icosahedral
2386:Categories
2315:Virosphere
2290:Viral load
2280:Satellites
2066:Components
1893:Williams R
1313:2011-03-18
948:2007-07-10
657:References
505:chaperonin
228:Aaron Klug
220:capsomeres
201:adenovirus
174:adenovirus
168:(PBCV-1),
153:procapsids
75:capsomeres
63:oligomeric
2275:Oncovirus
2212:Mycovirus
2202:Virophage
2125:Viroplasm
1815:169035711
1411:Sgro JY.
1197:Structure
964:Watson JD
568:Functions
485:) number.
411:rotavirus
339:⋅
279:≥
253:≥
170:mimivirus
129:enveloped
71:protomers
2392:Virology
2346:Category
2149:Genetics
2057:Virology
1987:Archived
1969:31318422
1874:25955384
1834:PLOS ONE
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619:See also
551:, where
407:reovirus
141:membrane
2358:Commons
2185:By host
2042:History
1960:6637973
1865:4425637
1842:Bibcode
1753:5373398
1730:Bibcode
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1618:3144854
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1366:Bibcode
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831:2141786
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781:6023873
753:Bibcode
704:1532201
684:Bibcode
592:enzymes
515:Helical
509:in vivo
498:E. coli
490:E. coli
461:Prolate
453:, or a
441:, or a
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69:called
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