Poliovirus - Knowledge

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are an association of RNA molecules consisting of a template RNA and several growing RNAs of varying length, are seen in both the replication complexes for (−)RNAs and (+)RNAs. For synthesis of each negative-strand and positive-strand RNAs, VPg protein in the poliovirus works as a primer. RNA-dependent RNA polymerase of the poliovirus adds two uracil nucleotides (UU) to VPg protein utilizing the poly(A) tail at the 3′-end of the +ssRNA genome as a pattern for synthesis of the negative-strand antigenomic RNA. To initiate this −ssRNA synthesis, the tyrosine hydroxyl of VPg is needed. But for the initiation of positive strand RNA synthesis, CRE-dependent VPg uridylylation is needed. Which means that VPg is once more utilized as a primer however this time it adds the two uridine triphosphates using a cis-acting replication element (CRE) as a template.

388:(IRES). This region consists of many secondary structures and 3 or 4 domains. Domain 3 is a self folding RNA element that contains conserved structural motifs in various stable stem loops linked by two four-way junctions. As IRES consists of many domains, these domains themselves consist of many loops that contribute to modified translation without a 5’ end cap by hijacking ribosomes. The interaction loop of domain 3 is known as GNRA tetraloop. The residues of adenosines A180 and A181 in the GUAA tetraloop form hydrogen bonds via non canonical base pairing interactions with the base pairs of the receptors C230/G242 and G231/C241, respectively. Genetic mutations in this region prevent viral protein production. The first IRES to be discovered was found in poliovirus RNA. 336:

cleaved, yielding mature viral proteins (4). The positive-sense RNA serves as template for complementary negative-strand synthesis, producing double-stranded replicative form (RF) RNA (5). Many positive strand RNA copies are produced from the single negative strand (6). The newly synthesized positive-sense RNA molecules can serve as templates for translation of more viral proteins (7) or can be enclosed in a capsid (8), which ultimately generates progeny virions. Lysis of the infected cell results in release of infectious progeny virions (9).

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region of the genome that have no coding activity, at least 3.7-kb distant from the initial location. This process can occurs without negatively influencing activity. CRE copies do not influence replication negatively. Uridylylation process of VPg that takes place at CRE needs the presence of 3CD that is an RNA binding protein. It is attached to the CRE directly and specifically. Because of its presence VPg can bind the CRE properly and primary production proceeds without problems.

578:. As of March 2020, wild PV-1 is highly localized to regions in Pakistan and Afghanistan. Certification of the eradication of indigenous transmission of wild PV-2 occurred in September 2015, after last being detected in 1999, and in October 2019 for wild PV-3 after last being detected in 2012. However, circulating vaccine-derived poliovirus (variant poliovirus, cVDPV) of all three serotypes continues to circulate and cause paralysis, having been detected in 32 countries in 2023. 658: 889: 55: 1142:

genetically more stable and less likely to regain virulence than the original Sabin2 strain, with three key genetic modifications. The vaccine derived from this strain, novel oral polio virus type 2 (nOPV2), was granted emergency licencing in 2021, and subsequently full licensure in December 2023. Genetically stabilsed vaccines targeting poliovirus types 1 and 3 are in development, with the intention that these will eventually completely replace the Sabin vaccines.

332: 399: 4650: 4636: 1022:" poliovirus (WPV), and the much more prevalent mutated oral vaccine strains, known as circulating vaccine-derived poliovirus (cVDPV) or variant poliovirus. Vaccines against each of the three wild strains of polio have given rise to strains of cVDPV, with cVDPV2 being most prominent. cVDPV caused 524 confirmed paralytic polio cases worldwide in 2023, and was detected in 32 countries. 395:. This polypeptide is then autocleaved by internal proteases into about 10 individual viral proteins. Not all cleavages occur with the same efficiency. Therefore, the amounts of proteins produced by the polypeptide cleavage vary: for example, smaller amounts of 3D are produced than those of capsid proteins, VP1–4. These individual viral proteins are: 829:. Second, because it can replicate very quickly, the virus overwhelms the host's organs before an immune response can be mounted. If detail is given at the attachment phase; poliovirus with canyons on the virion surface have virus attachment sites located in pockets at the canyon bases. The canyons are too narrow for access by 31: 1268:

were used to convert the DNA back into RNA, its natural state. Other enzymes were then used to translate the RNA into a polypeptide, producing functional viral particle. This whole painstaking process took two years. The newly minted synthetic virus was injected into PVR transgenic mice, to determine

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Five copies each of VP0, VP3, and VP1 whose N termini and VP4 form interior surface of capsid, assemble into a 'pentamer' and 12 pentamers form a procapsid. (The outer surface of capsid is consisting of VP1, VP2, VP3; C termini of VP1 and VP3 form the canyons which around each of the vertices; around

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infection. The mechanisms by which poliovirus enters the CNS are poorly understood. Three nonmutually exclusive hypotheses have been suggested to explain its entry. All theories require primary viremia. The first hypothesis predicts that virions pass directly from the blood into the central nervous

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The primary determinant of infection for any virus is its ability to enter a cell and produce additional infectious particles. The presence of CD155 is thought to define the animals and tissues that can be infected by poliovirus. CD155 is found (outside of laboratories) only on the cells of humans,

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After translation, transcription and genome replication which involve a single process, synthesis of (+) RNA) is realized. For the infecting (+)RNA to be replicated, multiple copies of (−)RNA must be transcribed and then used as templates for (+)RNA synthesis. Replicative intermediates (RIs), which

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of an animal RNA virus, poliovirus. DNA encoding the RNA genome of poliovirus was introduced into cultured mammalian cells and infectious poliovirus was produced. Creation of the infectious clone propelled understanding of poliovirus biology, and has become a standard technology used to study many

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million doses administered. In addition, the live virus can also circulate in under-vaccinated populations (circulating vaccine-derived poliovirus, cVDPV) and over time can revert to a neurovirulent form causing paralytic polio. Researchers have developed a poliovirus type 2 vaccine strain that is

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TgPVR21 mice have incorporated the human PVR at chromosome 13. These mice are less susceptible to poliovirus infection through the intracerebral route, possibly because they express decreased levels of hPVR. TgPVR21 mice have been shown to be susceptible to poliovirus infection through intranasal

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The CRE of poliovirus is identified as an unachieved base-paired stem and a final loop consisting of 61 nt. The CRE is found in enteroviruses. It is a highly preserved secondary RNA structural element and bedded in the genome's polyprotein-coding region. The complex can be translocated to the 5'

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succeeded in synthesizing poliovirus from its chemical code, producing the world's first synthetic virus. Scientists first converted poliovirus's published RNA sequence, 7741 bases long, into a DNA sequence, as DNA was easier to synthesize. Short fragments of this DNA sequence were obtained by

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The replication cycle of poliovirus is initiated by binding to the cell surface receptor CD155 (1). The virion forms a pore in the cell membrane through which viral RNA is released into the cytoplasm (2). Translation of the viral RNA occurs by an IRES-mediated mechanism (3). The polyprotein is

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Drake demonstrated that poliovirus is able to undergo multiplicity reactivation. That is, when polioviruses were irradiated with UV light and allowed to undergo multiple infections of host cells, viable progeny could be formed even at UV doses that inactivated the virus in single infections.

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response (specifically that of interferon alpha and beta) is an important factor that defines which types of cells support poliovirus replication. In mice expressing CD155 (through genetic engineering) but lacking the type I interferon receptor, poliovirus not only replicates in an expanded

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antibodies against poliovirus can prevent the spread of the virus to motor neurons of the central nervous system. Infection with one serotype of poliovirus does not provide immunity against the other serotypes; however, second attacks within the same individual are extremely rare.

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if the synthetic version was able to cause disease. The synthetic virus was able to replicate, infect, and cause paralysis or death in mice. However, the synthetic version was between 1,000 and 10,000 times weaker than the original virus, probably due to one of the added markers.

727:. The sustained viral replication causes secondary viremia and leads to the development of minor symptoms such as fever, headache, and sore throat. Paralytic poliomyelitis occurs in less than 1% of poliovirus infections. Paralytic disease occurs when the virus enters the 791:

Poliomyelitis is a disease of the central nervous system. However, CD155 is believed to be present on the surface of most or all human cells. Therefore, receptor expression does not explain why poliovirus preferentially infects certain tissues. This suggests that tissue

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4. These mice express the highest levels of the transgene and the highest sensitivity to poliovirus. TgPVR1 mice are susceptible to poliovirus through the intraspinal, intracerebral, intramuscular, and intravenous pathways, but not through the oral

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Although humans are the only known natural hosts of poliovirus, monkeys can be experimentally infected and they have long been used to study poliovirus. In 1990–91, a small animal model of poliomyelitis was developed by two laboratories. Mice were

348:(also known as the poliovirus receptor or PVR) on the cell surface. Interaction of poliovirus and CD155 facilitates an irreversible conformational change of the viral particle necessary for viral entry. Following attachment to the host 3893:

Koike S, Taya C, Aoki J, Matsuda Y, Ise I, Takeda H, et al. (1994). "Characterization of three different transgenic mouse lines that carry human poliovirus receptor gene--influence of the transgene expression on pathogenesis".

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approved the use of the TgPVR mouse as an alternative method of assessing the effectiveness of the vaccine against poliovirus type-3. In 2000, the mouse model was approved for tests of vaccines against type-1 and type-2 poliovirus.

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independent of CD155. A second hypothesis suggests that the virions are transported from peripheral tissues that have been bathed in the viremic blood, for example muscle tissue, to the spinal cord through nerve pathways via

2759: 810: 364:. Recent experimental evidence supports the latter hypothesis and suggests that poliovirus binds to CD155 and is taken up by endocytosis. Immediately after internalization of the particle, the viral RNA is released. 2905:"Recruitment of nectin-3 to cell-cell junctions through trans-heterophilic interaction with CD155, a vitronectin and poliovirus receptor that localizes to alpha(v)beta3 integrin-containing membrane microdomains" 379:

by the host cell. On entry, the virus hijacks the cell's translation machinery, causing inhibition of cellular protein synthesis in favor of virus-specific protein production. Unlike the host cell's mRNAs, the

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and death. In cases of paralytic disease, muscle pain and spasms are frequently observed prior to onset of weakness and paralysis. Paralysis typically persists from days to weeks prior to recovery.

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Nagata N, Iwasaki T, Ami Y, Sato Y, Hatano I, Harashima A, et al. (March 2004). "A poliomyelitis model through mucosal infection in transgenic mice bearing human poliovirus receptor, TgPVR21".

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Mendelsohn CL, Wimmer E, Racaniello VR (March 1989). "Cellular receptor for poliovirus: molecular cloning, nucleotide sequence, and expression of a new member of the immunoglobulin superfamily".

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A drawback of the attenuated virus used in the Sabin oral polio vaccine is its potential to cause vaccine-associated paralytic poliomyelitis (VAPP) in approximately one individual per every 2.7

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4 to 6 hours following initiation of infection in cultured mammalian cells. The mechanism of viral release from the cell is unclear, but each dying cell can release up to 10,000 polio

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Kitamura N, Semler BL, Rothberg PG, Larsen GR, Adler CJ, Dorner AJ, et al. (June 1981). "Primary structure, gene organization and polypeptide expression of poliovirus RNA".

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similar to those of humans and monkeys. This mouse model of human poliovirus infection has proven to be an invaluable tool in understanding poliovirus biology and pathogenicity.

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In TgPVR5 mice, the human transgene is located on chromosome 12. These mice exhibit the lowest levels of hPVR expression and are the least susceptible to poliovirus infection.

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the virus strains in monkey kidney epithelial cells introduces mutations in the viral IRES, and hinders (or attenuates) the ability of the virus to infect nerve tissue.

3568: 2784: 1064:. Upon infection, TgPVR mice show signs of paralysis that resemble those of poliomyelitis in humans and monkeys, and the central nervous systems of paralyzed mice are 986:. IPV, being inactivated, does not carry these risks, but does not induce contact immunity. IPV is more costly and the logistics of its delivery are more challenging. 692:. The protein has several domains of which domain D1 contains the polio virus binding site. Within this domain, 37 amino acids are responsible for binding the virus. 446:

Some of the (+) RNA molecules are used as templates for further (−) RNA synthesis, some function as mRNA, and some are destined to be the genomes of progeny virions.

5332: 3316:"Establishment of a poliovirus oral infection system in human poliovirus receptor-expressing transgenic mice that are deficient in alpha/beta interferon receptor" 1641:

Gómez Yafal A, Kaplan G, Racaniello VR, Hogle JM (November 1993). "Characterization of poliovirus conformational alteration mediated by soluble cell receptors".

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inactivation of three wild, virulent reference strains: Mahoney or Brunenders (PV-1), MEF-1/Lansing (PV-2), and Saukett/Leon (PV-3). Oral polio vaccine contains

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In the assembly of new virus particles (i.e. the packaging of progeny genome into a procapsid which can survive outside the host cell), including, respectively:

1117:(OPV) production. Previously, monitoring the safety of OPV had to be performed using monkeys, because only primates are susceptible to the virus. In 1999, the 1105:, and have proven susceptible to poliovirus through intracerebral, intramuscular, and intranasal routes. In addition, these mice are capable of developing the 2077: 970:

polio vaccine (IPV), which is injected. OPV is less expensive and easier to administer, and can spread immunity beyond the person vaccinated, creating

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3 (WPV3) is last known to have caused polio in 2012, and was declared eradicated in 2019. All wild-virus cases since that date have been due to type

6022: 6017: 1196: 3977:"Transgenic mice as an alternative to monkeys for neurovirulence testing of live oral poliovirus vaccine: validation by a WHO collaborative study" 1878:

Pelletier J, Sonenberg N (July 1988). "Internal initiation of translation of eukaryotic mRNA directed by a sequence derived from poliovirus RNA".

3650: 1134:, was tested in early clinical trials as a possible treatment for cancer. As of September 2022, a number of clinical trials are under way. 240: 228: 216: 4258: 974:. It has been the predominant vaccine used. However, under conditions of long-term vaccine virus circulation in under-vaccinated populations, 6074: 5346: 3429: 996:

transmission of wild poliovirus, with its last reported case in 2016. Of the three strains of WPV, the last recorded wild case caused by type

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templates during negative strand synthesis. Recombination in RNA viruses appears to be an adaptive mechanism for repairing genome damage.

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Attardi G, Smith J (1962). "Virus specific protein and a ribo-nucleic acid associated with ribosomes in poliovirus infected HeLa cells".

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Suzuki Y (May 2006). "Ancient positive selection on CD155 as a possible cause for susceptibility to poliovirus infection in simians".

2517:"Evidence for emergence of diverse polioviruses from C-cluster coxsackie A viruses and implications for global poliovirus eradication" 1018:

are the only two countries where the disease is still classified as endemic. Recent polio cases arise from two sources, the original "

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can reactivate the virus to produce a polio-inducing strain, while OPV can also, in rare circumstances, induce polio or persistent

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Poliovirus is one of the most well-characterized viruses, and has become a useful model system for understanding the biology of

7305: 3216:"Limited trafficking of a neurotropic virus through inefficient retrograde axonal transport and the type I interferon response" 1835:

Chen CY, Sarnow P (April 1995). "Initiation of protein synthesis by the eukaryotic translational apparatus on circular RNAs".

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Mueller S, Wimmer E, Cello J (August 2005). "Poliovirus and poliomyelitis: a tale of guts, brains, and an accidental event".

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of poliovirus RNA is extremely long—over 700 nucleotides—and highly structured. This region of the viral genome is called an

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Ren R, Racaniello VR (October 1992). "Poliovirus spreads from muscle to the central nervous system by neural pathways".

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are avoided. Codon use differs between the three genotypes, and appears to be driven by mutation rather than selection.

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Lozano G, Fernandez N, Martinez-Salas E (February 2016). "Modeling Three-Dimensional Structural Motifs of Viral IRES".

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Racaniello VR, Baltimore D (November 1981). "Cloned poliovirus complementary DNA is infectious in mammalian cells".

6666: 6334: 5338: 5267: 4741: 424:(3B), a small protein that binds viral RNA and is necessary for synthesis of viral positive and negative strand RNA 38: 4722: 4710: 3811:"Transgenic mice carrying the human poliovirus receptor: new animal models for study of poliovirus neurovirulence" 3099:"Efficient delivery of circulating poliovirus to the central nervous system independently of poliovirus receptor" 4345:"Molecular cloning of poliovirus cDNA and determination of the complete nucleotide sequence of the viral genome" 535:

The mutation rate in the virus is relatively high even for an RNA virus, with a synonymous substitution rate of

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Hogle JM, Chow M, Filman DJ (September 1985). "Three-dimensional structure of poliovirus at 2.9 A resolution".

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conditions of the stomach, allowing ingested viruses to infect the host and spread throughout the body via the

677:. Poliovirus is, however, strictly a human pathogen, and does not naturally infect any other species (although 381: 4203: 4129: 7185: 5298: 361: 5375: 707:. Virus is shed in the feces of infected individuals. In 95% of cases only a primary, transient presence of 513:

analysis of the RNA and protein sequences of poliovirus suggests that it may have evolved from a C-cluster

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CD155 molecules complexed with a poliovirus particle. Reconstructed image from cryo-electron microscopy.

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were incorporated into the synthesized DNA, so that it could be distinguished from natural poliovirus.

1220: 907:, the goal of permanent global cessation of circulation of the poliovirus and hence elimination of the 809: 6845: 6649: 5850: 5562: 5169: 4027:

Brown MC, Dobrikova EY, Dobrikov MI, Walton RW, Gemberling SL, Nair SK, et al. (November 2014).

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In 1981, Racaniello and Baltimore used recombinant DNA technology to generate the first infectious

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Geiger K, Stehling-Ariza T, Bigouette JP, Bennett SD, Burns CC, Quddus A, et al. (May 2024).

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Geiger K, Stehling-Ariza T, Bigouette JP, Bennett SD, Burns CC, Quddus A, et al. (May 2024).

5753: 5067: 4663: 3672: 3455: 2730:. (Yale studies in the history of science and medicine). New Haven, Conn: Yale University Press. 772: 4733: 7525: 7510: 7435: 7407: 7243: 7128: 5997: 5968: 5461: 5439: 5318: 5314: 5310: 5306: 5261: 5199: 5093: 5054: 4857: 3976: 1252: 1204: 993: 801:

repertoire of tissue types, but these mice are also able to be infected orally with the virus.

728: 704: 352:, entry of the viral nucleic acid was thought to occur one of two ways: via the formation of a 2256: 2096: 2013: 524:

of poliovirus probably occurred as a result of a change in cellular receptor specificity from

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Sabin AB (June 1956). "Pathogenesis of poliomyelitis; reappraisal in the light of new data".

2576:"Calibration of multiple poliovirus molecular clocks covering an extended evolutionary range" 485:(RdRP) catalyzes recombination by a copy choice mechanism in which the RdRP switches between 474: 376: 4668: 3267:"The alpha/beta interferon response controls tissue tropism and pathogenicity of poliovirus" 3265:

Ida-Hosonuma M, Iwasaki T, Yoshikawa T, Nagata N, Sato Y, Sata T, et al. (April 2005).

1979: 747:, resulting in the selective destruction of motor neurons leading to temporary or permanent 7482: 7473: 7179: 7117: 6804: 6248: 6239: 6161: 6108: 5929: 5820: 5710: 5542: 5177: 5006: 4544: 4497: 4411: 4356: 3763: 3003: 2528: 2479: 2308: 1887: 1844: 1330: 932: 777: 700: 612:

ceased to be recognized as a species, and the three serotypes were assigned to the species

305: 67: 3508: 3485: 3140:"Retrograde transport of intact poliovirus through the axon via the fast transport system" 457:

Each procapsid acquires a copy of the virus genome, with VPg still attached at the 5' end.

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A child receives oral polio vaccine during a 2002 campaign to immunize children in India.

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VP0, which is further cleaved into VP2 and VP4, VP1 and VP3, proteins of the viral capsid

4548: 4501: 4415: 4360: 3767: 3623: 3598: 3007: 2831: 2806: 2532: 2483: 2208:"Structure and function analysis of the poliovirus cis-acting replication element (CRE)" 1891: 1848: 7372: 7249: 5522: 5517: 5231: 4987: 4613: 4570: 4435: 4180: 4155: 4053: 4028: 3993: 3975:

Dragunsky E, Nomura T, Karpinski K, Furesz J, Wood DJ, Pervikov Y, et al. (2003).

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Horie H, Koike S, Kurata T, Sato-Yoshida Y, Ise I, Ota Y, et al. (February 1994).

3732: 3703:"Transgenic mice expressing a human poliovirus receptor: a new model for poliomyelitis" 3340: 3315: 3291: 3266: 3242: 3215: 2880: 2855: 2700: 2673: 2600: 2575: 2551: 2516: 2402: 2377: 2232: 2207: 2133: 1911: 1742: 1715: 1473: 1425: 1398: 1339: 1314: 1184: 1183:

In 1981, the poliovirus genome was published by two different teams of researchers: by

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Drake JW (August 1958). "Interference and multiplicity reactivation in polioviruses".

2183: 2158: 1955: 1930: 1618: 1593: 1569: 1545:"Entry and release of poliovirus as observed by electron microscopy of cultured cells" 1544: 1520: 1495: 570:

protein. Capsid proteins define cellular receptor specificity and virus antigenicity.

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Home of Picornaviruses (latest updates of species, serotypes, & proposed changes)

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are present in the same host cell. Kirkegaard and Baltimore presented evidence that

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Ohka S, Igarashi H, Nagata N, Sakai M, Koike S, Nochi T, et al. (August 2007).

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Yang WX, Terasaki T, Shiroki K, Ohka S, Aoki J, Tanabe S, et al. (March 1997).

2293:"Vaccine-derived polioviruses and the endgame strategy for global polio eradication" 2174: 1946: 1929:

Jang SK, Kräusslich HG, Nicklin MJ, Duke GM, Palmenberg AC, Wimmer E (August 1988).

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He Y, Mueller S, Chipman PR, Bator CM, Peng X, Bowman VD, et al. (April 2003).

1477: 308:. Because of its short genome and its simple composition—only a strand of RNA and a 7086: 6692: 6429: 6424: 6419: 6414: 6409: 6404: 6383: 5670: 5380: 5060: 4992: 4873: 4804: 4597: 4574: 4552: 4505: 4462: 4439: 4419: 4374: 4364: 4175: 4167: 4048: 4040: 3988: 3949: 3903: 3867: 3830: 3822: 3781: 3771: 3714: 3618: 3610: 3599:"Progress Toward Poliomyelitis Eradication - Worldwide, January 2022-December 2023" 3376: 3335: 3327: 3286: 3278: 3237: 3227: 3188: 3151: 3110: 3011: 2953: 2916: 2875: 2867: 2826: 2818: 2807:"Progress Toward Poliomyelitis Eradication - Worldwide, January 2022-December 2023" 2695: 2685: 2644: 2636: 2595: 2587: 2546: 2536: 2487: 2438: 2397: 2389: 2350: 2304: 2227: 2219: 2178: 2170: 2129: 1950: 1942: 1915: 1895: 1852: 1809: 1774: 1737: 1727: 1692: 1688: 1650: 1613: 1605: 1564: 1556: 1515: 1507: 1457: 1420: 1410: 1334: 1326: 1231: 1168: 1156: 971: 826: 767: 674: 600:

Polioviruses were formerly classified as a distinct species belonging to the genus

509:), which also use immunoglobulin-like molecules to recognize and enter host cells. 356:

in the plasma membrane through which the RNA is then "injected" into the host cell

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Koike S, Taya C, Kurata T, Abe S, Ise I, Yonekawa H, et al. (February 1991).

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3D macromolecular structures of the Poliovirus archived in the EM Data Bank(EMDB)

4675: 4641: 4281: 4156:"Engineering the Live-Attenuated Polio Vaccine to Prevent Reversion to Virulence" 3858:

Ohka S, Nomoto A (October 2001). "Recent insights into poliovirus pathogenesis".

3539: 3512: 3232: 2625:"Mononucleotide and dinucleotide frequencies, and codon usage in poliovirion RNA" 2492: 2467: 1778: 1732: 1560: 1188: 1075:

In TgPVR1 mice, the transgene encoding the human PVR was incorporated into mouse

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is the most common form encountered in nature, but all three forms are extremely

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use is not random; codons ending in adenosine are favoured, and those ending in

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Brandenburg B, Lee LY, Lakadamyali M, Rust MJ, Zhuang X, Hogle JM (July 2007).

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Yeh MT, Bujaki E, Dolan PT, Smith M, Wahid R, Konz J, et al. (May 2020).

2871: 1813: 1496:"Complexes of poliovirus serotypes with their common cellular receptor, CD155" 1315:"Poliovirus cell entry: common structural themes in viral cell entry pathways" 7504: 7339: 6990: 6451: 6088: 5805: 5780: 5776: 4825: 4820: 4768: 3056: 1256:

mail-order, and assembled. The complete viral genome was then assembled by a

1248: 1200: 1049: 959: 912: 841: 818: 780:. A third hypothesis is that the virus is imported into the CNS via infected 755:

acquired from their mothers. In rare cases, paralytic poliomyelitis leads to

582: 372: 349: 261: 183: 109: 4557: 4532: 4509: 4466: 3408:. Liverpool John Moores University, UK: John Wiley & Sons. p. 162. 2690: 2672:

Zhang J, Wang M, Liu WQ, Zhou JH, Chen HT, Ma LN, et al. (March 2011).

2640: 2541: 1856: 543:

substitutions/site/year. Base distribution within the genome is not random;

7059: 7011: 6963: 6915: 6836: 6621: 6457: 6134: 5943: 5810: 5784: 5686: 5536: 5214: 5075: 4609: 4566: 4369: 4189: 4062: 4002: 3961: 3879: 3632: 3614: 3390: 3349: 3300: 3251: 3156: 3139: 3115: 3098: 3023: 2965: 2930: 2921: 2904: 2889: 2840: 2822: 2709: 2609: 2560: 2501: 2452: 2443: 2426: 2362: 2316: 2241: 2192: 1821: 1786: 1751: 1700: 1654: 1529: 1434: 1415: 1348: 1160: 1106: 979: 837: 796:

is determined after cellular infection. Recent work has suggested that the

763: 744: 732: 712: 657: 586: 510: 121: 4517: 4474: 4431: 4388: 3915: 3844: 3795: 3776: 3728: 3200: 3165: 3124: 2658: 2411: 1964: 1907: 1864: 1662: 1627: 1594:"Neutralization of poliovirus by cell receptors expressed in insect cells" 1578: 1469: 888: 836:

Individuals who are exposed to poliovirus, either through infection or by

766:

phase of infection is thought to be an accidental diversion of the normal

7467: 7399: 6942: 6262: 6225: 5726: 5675: 5663: 5658: 5651: 5639: 5613: 5583: 5485: 5324: 5236: 5219: 5154: 4815: 3331: 3041:(1st ed.). Chichester, England: John Wiley & Sons. p. 158. 2674:"Analysis of codon usage and nucleotide composition bias in polioviruses" 2591: 2515:

Jiang P, Faase JA, Toyoda H, Paul A, Wimmer E, Gorbalenya AE (May 2007).

1261: 1216: 1057: 1011: 955: 736: 703:, meaning that one ingests the virus and viral replication occurs in the 696: 431:

proteins which comprise the protein complex needed for virus replication.

392: 320: 169: 4588:

Couzin J (July 2002). "Virology. Active poliovirus baked from scratch".

3701:

Ren RB, Costantini F, Gorgacz EJ, Lee JJ, Racaniello VR (October 1990).

2223: 1215:

The three-dimensional structure of poliovirus was determined in 1985 by

1048:

poliovirus receptor (TgPVR) mice are susceptible to poliovirus injected

331: 7281: 7213: 6810: 6616: 6443: 5576: 5360: 4044: 3907: 1076: 1045: 948: 849: 785: 740: 521: 506: 297: 45: 41: 4704: 3561: 751:. This is a very rare event in babies, who still have anti-poliovirus 7327: 7322: 7263: 7257: 6501: 6378: 6368: 6356: 6129: 5793: 5621: 5602: 5569: 5557: 5551: 5530: 5476: 5404: 5290: 5252: 5021: 5000: 4906: 4784: 4776: 4764: 4423: 3085: 3075: 1899: 1113:

The development of the TgPVR mouse has had a profound effect on oral

1019: 748: 716: 594: 547:

is less common than expected at the 5' end and higher at the 3' end.

544: 502: 371:. Thus, the genome enclosed within the viral particle can be used as 368: 357: 353: 301: 97: 7429: 3596: 3531: 2804: 2292: 2291:

Kew OM, Sutter RW, de Gourville EM, Dowdle WR, Pallansch MA (2005).

1000:

2 (WPV2) was in 1999, and WPV2 was declared eradicated in 2015. Type

566:

of poliovirus, PV-1, PV-2, and PV-3, each have a slightly different

398: 316:

it—poliovirus is widely regarded as the simplest significant virus.

7452: 5909: 5719: 5366: 5108: 4964: 1015: 975: 944: 830: 781: 752: 711:(virus in the bloodstream) occurs, and the poliovirus infection is 670: 563: 552: 461:

Fully assembled poliovirus leaves the confines of its host cell by

415: 3264: 539:

substitutions/site/year and a non-synonymous substitution rate of

7487: 5472: 4763: 1131: 989: 793: 708: 556: 497:

Poliovirus is structurally similar to other human enteroviruses (

466: 411:

whose function is to make multiple copies of the viral RNA genome

286: 54: 3592: 3590: 1640: 1163:. The structure of the virus was first elucidated in 1958 using 982:

infection in vaccinated individuals, particularly those who are

7169: 7107: 6855: 6794: 6687: 6185: 5841: 5081: 4208:

Global Polio Eradication Initiative - World Health Organization

3138:

Ohka S, Yang WX, Terada E, Iwasaki K, Nomoto A (October 1998).

1399:"Epidemics to eradication: the modern history of poliomyelitis" 1265: 1085: 920: 853: 822: 724: 567: 478: 313: 289: 282: 3974: 3470: 454:

this time, the 60 copies of VP0 are cleaved into VP4 and VP2.)

6740: 6730: 6725: 6715: 6710: 6705: 6700: 6307: 6302: 6297: 5732: 5491: 5123: 3587: 1799: 1713: 1099: 1061: 908: 652: 548: 529: 462: 345: 256: 84: 4649: 4401: 4026: 3808: 259:(also known as poliomyelitis), is a serotype of the species 7292: 7123: 6735: 6720: 6149: 5885: 4255:"Behind the picture: Rosalind Franklin and the polio model" 2290: 1542: 1095: 1071:

Three distinct types of TgPVR mice have been well studied:

4103:"UCSF Glioma Trial: PVSRIPO in Recurrent Malignant Glioma" 3569:"GPEI-Two out of three wild poliovirus strains eradicated" 3404:

Carter J, Saunders VA (2007). "Picornavirus replication".

2781:"GPEI-Two out of three wild poliovirus strains eradicated" 1447: 1094:

A fourth TgPVR mouse model - these "cPVR" mice carry hPVR

860:

antibodies) and are able to block poliovirus replication;

7353: 7349: 7332: 6478: 6329: 4958: 3214:

Lancaster KZ, Pfeiffer JK (March 2010). Gale Jr M (ed.).

1928: 1591: 1543:

Dunnebacke TH, Levinthal JD, Williams RC (October 1969).

865: 861: 857: 593:(weakened) strains of the three serotypes of poliovirus. 421: 279: 62:

A type 3 poliovirus capsid, protein side chains coloured

2465: 1175:, showing the polio virus to have icosahedral symmetry. 896: 30: 4077:"Update on Recombinant Poliovirus Cancer Immunotherapy" 3700: 3313: 3096: 2752:"Global eradication of wild poliovirus type 2 declared" 2205: 681:

and Old World monkeys can be experimentally infected).

4322:. New York: Oxford University Press. pp. 359–61. 4257:. Medical Research Council. 2019-03-14. Archived from 1592:

Kaplan G, Freistadt MS, Racaniello VR (October 1990).

581:

Specific strains of each serotype are used to prepare

3509:"Rinderpest virus has been wiped out, scientists say" 3500: 3137: 1493: 4631: 4231:"Two New Vaccines Join the Fight to Eradicate Polio" 3939: 3892: 2773: 2573: 2514: 2466:

Bernstein H, Bernstein C, Michod RE (January 2018).

2206:

Goodfellow IG, Kerrigan D, Evans DJ (January 2003).

528:(ICAM-1) (used by C-cluster Coxsackie A viruses) to 5775: 4487: 4342: 4153: 2375: 1767:

Cold Spring Harbor Symposia on Quantitative Biology

1235:

Model of poliovirus-binding CD155 (shown in purple)

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"Picornavirus". 3036: 2622: 2254: 2094: 2011: 2007: 2005: 2003: 1977: 7389: 5761: 4749: 3537: 3178: 2902: 7058: 7010: 6962: 6914: 3968: 3930: 3851: 2853: 2721: 2719: 2616: 2156: 1764: 1758: 935:, have spearheaded the campaign through the 7398: 6224: 4124: 4122: 3886: 3802: 3752:"Transgenic mice susceptible to poliovirus" 3743: 3694: 3603:MMWR. Morbidity and Mortality Weekly Report 2972: 2811:MMWR. Morbidity and Mortality Weekly Report 2376:Kirkegaard K, Baltimore D (November 1986). 2286: 2284: 2282: 2280: 2054:Viruses: Biology, Applications, and Control 2020:. John Wiley & Sons. pp. 161–169. 2000: 1707: 927:. 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Inactive polio vaccine is prepared by 397: 330: 4286:. London: Harper Collins. p. 296. 4283:Rosalind Franklin: The Dark Lady of DNA 4029:"Oncolytic polio virotherapy of cancer" 4020: 2623:Rothberg PG, Wimmer E (December 1981). 1669: 1355: 1308: 1306: 1284: 1125: 492: 7503: 7168: 7106: 6184: 5840: 4587: 4279: 2943: 2309:10.1146/annurev.micro.58.030603.123625 2263:. John Wiley & Sons. p. 166. 2123: 2103:. John Wiley & Sons. p. 164. 2051: 1385: 1331:10.1146/annurev.micro.56.012302.160757 1278: 1056:, and when injected directly into the 962:—oral polio vaccine (OPV), which uses 852:against poliovirus are present in the 7434: 7433: 7069:Respiratory syncytial virus G protein 5749: 4737: 4317: 4228: 3506: 3406:Virology: Principles and Applications 3039:Virology: Principles and Applications 2993: 2340: 2261:Virology: Principles and Applications 2101:Virology: Principles and Applications 2018:Virology: Principles and Applications 1984:Virology: Principles and Applications 1585: 1312: 943:has been achieved twice before, with 688:gene appears to have been subject to 300:long. The viral particle is about 30 7156: 7094: 7046: 6998: 6950: 6902: 6466: 6392: 6096: 5951: 5917: 4680: 4100: 2725: 1986:. John Wiley & Sons. p. 4. 1922: 1828: 1634: 1489: 1487: 1303: 1084:inoculation, and may be useful as a 326: 5034:Subacute sclerosing panencephalitis 4196: 3550:from the original on 26 August 2020 2909:The Journal of Biological Chemistry 2903:Mueller S, Wimmer E (August 2003). 2756:Global Polio Eradication Initiative 2157:Murray KE, Barton DJ (April 2003). 937:Global Polio Eradication Initiative 821:. First, it can survive the highly 296:(+ssRNA) genome that is about 7500 13: 3181:The Journal of Infectious Diseases 2854:Carstens EB, Ball LA (July 2009). 2134:10.1016/b978-0-12-800947-5.00014-4 2080:. Brown University. Archived from 1031: 941:eradication of infectious diseases 418:which cleave the viral polypeptide 367:Poliovirus is a positive-stranded 294:single-stranded positive-sense RNA 14: 7547: 6973:Mumps hemagglutinin-neuraminidase 5353:Severe acute respiratory syndrome 4879:Follicular dendritic cell sarcoma 4687:August 2001 Molecule of the Month 4627: 3683:from the original on 28 July 2020 3653:from the original on 22 July 2020 2472:Infection, Genetics and Evolution 1484: 915:effort begun in 1988, led by the 526:intercellular adhesion molecule-1 6335:Hepatitis B virus DNA polymerase 5339:Middle East respiratory syndrome 4664:ICTVdb virus classification 2006 4648: 4634: 4320:J.D. Bernal: The Sage of Science 1610:10.1128/JVI.64.10.4697-4702.1990 878:This section is an excerpt from 628:. The type species of the genus 71: 48:. Scale bar (white): 50 nm 7521:Human viruses described in 1909 6166:(non-enveloped circular ds-DNA) 4581: 4524: 4481: 4311: 4300:from the original on 2023-01-14 4273: 4247: 4222: 4147: 4094: 4069: 4009:from the original on 2021-02-24 3665: 3639: 3540:"Africa declared free of polio" 3519:from the original on 3 May 2019 3478:"Smallpox [Fact Sheet]" 3440: 3422: 3397: 3307: 3283:10.1128/JVI.79.7.4460-4469.2005 3258: 3207: 3172: 3131: 3090: 3063: 3030: 2987: 2979:Bodian D, Horstmann DH (1969). 2896: 2847: 2798: 2787:from the original on 2019-11-07 2762:from the original on 2015-09-28 2567: 2508: 2459: 2431:The Journal of General Virology 2425:Barr JN, Fearns R (June 2010). 2418: 2369: 2334: 2323:from the original on 2020-07-09 2248: 2199: 2175:10.1128/JVI.77.8.4739-4750.2003 2150: 2124:Louten J (2016). "Poliovirus". 2117: 2088: 2070: 2045: 2034:from the original on 2023-01-14 1971: 1947:10.1128/JVI.62.8.2636-2643.1988 1793: 1512:10.1128/JVI.77.8.4827-4835.2003 646: 4938:Adult T-cell leukemia/lymphoma 4924:Splenic marginal zone lymphoma 3538:Scherbel-Ball N (2020-08-25). 3363:Racaniello VR (January 2006). 2255:Carter J, Saunders VA (2007). 2095:Carter J, Saunders VA (2007). 2012:Carter J, Saunders VA (2007). 1980:"Viruses and their importance" 1978:Carter J, Saunders VA (2007). 1693:10.1016/j.virusres.2005.04.008 1285:Ryan KJ, Ray CG, eds. (2004). 921:United Nations Children's Fund 872: 1: 7186:Vesiculovirus matrix proteins 4602:10.1126/science.297.5579.174b 4204:"GPEI-OPV Oral polio vaccine" 3872:10.1016/S0966-842X(01)02200-4 3827:10.1128/JVI.68.2.681-688.1994 3016:10.1126/science.123.3209.1151 2297:Annual Review of Microbiology 2128:. Elsevier. pp. 257–71. 1319:Annual Review of Microbiology 1289:(4th ed.). McGraw Hill. 1272: 1223:using X-ray crystallography. 1109:after intranasal inoculation. 362:receptor-mediated endocytosis 278:Poliovirus is composed of an 5143:Tropical spastic paraparesis 5048:Lymphocytic choriomeningitis 3720:10.1016/0092-8674(90)90168-E 3233:10.1371/journal.ppat.1000791 2493:10.1016/j.meegid.2017.10.024 2468:"Sex in microbial pathogens" 2394:10.1016/0092-8674(86)90600-8 2355:10.1016/0042-6822(58)90073-4 1802:Journal of Molecular Biology 1779:10.1101/SQB.1962.027.001.026 1733:10.1371/journal.pbio.0050183 1561:10.1128/JVI.4.4.505-513.1969 1462:10.1016/0092-8674(89)90690-9 1287:Sherris Medical Microbiology 1210: 1150: 483:RNA-dependent RNA polymerase 409:RNA dependent RNA polymerase 386:internal ribosome entry site 7: 7223:Structure and genome of HIV 6280:(circular partially ds-DNA) 5506:Cytomegalovirus esophagitis 5426:Human parainfluenza viruses 5099:Ramsay Hunt syndrome type 2 4683:"Poliovirus and Rhinovirus" 3954:10.1016/j.virol.2003.12.008 3507:Ghosh P (14 October 2010). 3382:10.1016/j.virol.2005.09.015 1026: 778:retrograde axonal transport 699:. Infection occurs via the 10: 7552: 5856:Herpesvirus glycoprotein B 4172:10.1016/j.chom.2020.04.003 4130:"OPV - Oral Polio Vaccine" 3647:"Endemic Countries - GPEI" 3430:"WHO | Poliomyelitis" 2958:10.1016/j.gene.2005.12.016 2728:A History of Poliomyelitis 2257:"Picornavirus replication" 2097:"Picornavirus replication" 2014:"Picornavirus replication" 1221:Scripps Research Institute 1145: 877: 650: 273:three poliovirus serotypes 7516:Viruses described in 1909 7442: 7366: 7303: 7230: 7221: 7212: 7147: 7085: 7037: 6989: 6941: 6893: 6868: 6835: 6793: 6781: 6772: 6757: 6675: 6630: 6615: 6560: 6525: 6510: 6495: 6442: 6377: 6364: 6355: 6317: 6286: 6275: 6212: 6172: 6143: 6125: 6087: 6062: 6031: 6005: 5996: 5985: 5942: 5908: 5868: 5851:HHV capsid portal protein 5828: 5819: 5801: 5792: 5709: 5685: 5593: 5516: 5471: 5460: 5289: 5251: 5230: 5198: 5170:Cytomegalovirus retinitis 5153: 5107: 4986: 4975: 4948: 4775: 3482:World Health Organization 2872:10.1007/s00705-009-0400-2 1814:10.1016/j.jmb.2016.01.005 1397:De Jesus NH (July 2007). 1178: 1119:World Health Organization 958:. There are two kinds of 917:World Health Organization 848:. In immune individuals, 360:, or via virus uptake by 312:icosahedral protein coat 255:, the causative agent of 208: 203: 66: 61: 52: 37: 28: 23: 7536:Infraspecific virus taxa 5272:Infectious mononucleosis 5089:Chandipura vesiculovirus 4919:Hepatocellular carcinoma 4864:Nasopharyngeal carcinoma 4797:Hepatocellular carcinoma 4081:Tisch Brain Tumor Center 3452:Global Health Strategies 3193:10.1093/infdis/166.4.747 2126:Essential Human Virology 1371:. New York: Copernicus. 1010:As of August 2024, 731:(CNS) and replicates in 477:when at least two viral 275:, numbered 1, 2, and 3. 6846:Influenza hemagglutinin 5068:Encephalitis lethargica 4769:viral systemic diseases 4558:10.1126/science.1072266 4510:10.1126/science.6272391 4467:10.1126/science.2994218 4160:Cell Host & Microbe 4107:clinicaltrials.ucsf.edu 3673:"Poliomyelitis (polio)" 3081:Pediatric Poliomyelitis 2691:10.1186/1743-422X-8-146 2542:10.1073/pnas.0700451104 1857:10.1126/science.7536344 805:Immune system avoidance 771:system by crossing the 473:Poliovirus can undergo 427:2BC, 2B, 2C (an ATPase) 7408:Gag-onc fusion protein 5440:Human orthopneumovirus 5094:Herpesviral meningitis 5055:Arbovirus encephalitis 4370:10.1073/pnas.78.8.4887 4229:Gadye L (2023-06-14). 3860:Trends in Microbiology 3615:10.15585/mmwr.mm7319a4 3157:10.1006/viro.1998.9360 3116:10.1006/viro.1997.8450 2922:10.1074/jbc.M304166200 2823:10.15585/mmwr.mm7319a4 2629:Nucleic Acids Research 2444:10.1099/vir.0.020818-0 1655:10.1006/viro.1993.1621 1416:10.1186/1743-422X-4-70 1253:Stony Brook University 1236: 1205:Stony Brook University 901: 893: 814: 762:In many respects, the 729:central nervous system 705:gastrointestinal tract 665: 583:vaccines against polio 403: 337: 7378:murine leukemia virus 7269:Reverse transcriptase 7149:Indiana vesiculovirus 7021:Measles hemagglutinin 4898:Merkel-cell carcinoma 4727:NCBI Taxonomy Browser 4715:NCBI Taxonomy Browser 3777:10.1073/pnas.88.3.951 3484:(WHO). Archived from 2641:10.1093/nar/9.23.6221 2084:on February 22, 2007. 1369:The machinery of life 1234: 1227:Cloning and synthesis 899: 891: 812: 660: 475:genetic recombination 401: 334: 6109:Early 35 kDa protein 5543:Adenovirus infection 5007:Human polyomavirus 2 4836:Oropharyngeal cancer 4723:"Human poliovirus 3" 4711:"Human poliovirus 1" 3896:Archives of Virology 3332:10.1128/JVI.02675-06 2860:Archives of Virology 2592:10.1128/JVI.02354-07 1367:Goodsell DS (1998). 1126:Genetic modification 1107:bulbar form of polio 1044:Unlike normal mice, 933:The Gates Foundation 493:Origin and serotypes 306:icosahedral symmetry 68:Virus classification 5129:Post-polio syndrome 4765:Infectious diseases 4549:2002Sci...297.1016C 4543:(5583): 1016–1018. 4502:1981Sci...214..916R 4461:(4720): 1358–1365. 4416:1981Natur.291..547K 4361:1981PNAS...78.4887R 4101:UCSF (2017-06-01). 3815:Journal of Virology 3768:1991PNAS...88..951K 3448:"Polio Eradication" 3320:Journal of Virology 3271:Journal of Virology 3071:Acute Poliomyelitis 3008:1956Sci...123.1151S 3002:(3209): 1151–1157. 2915:(33): 31251–31260. 2580:Journal of Virology 2533:2007PNAS..104.9457J 2484:2018InfGE..57....8B 2437:(Pt 6): 1373–1387. 2224:10.1261/rna.2950603 2163:Journal of Virology 1935:Journal of Virology 1892:1988Natur.334..320P 1849:1995Sci...268..415C 1808:(5 Pt A): 767–776. 1598:Journal of Virology 1549:Journal of Virology 1500:Journal of Virology 1066:histocytochemically 964:weakened poliovirus 939:(GPEI). Successful 773:blood–brain barrier 721:reticuloendothelial 662:Electron micrograph 614:Human enterovirus C 265:, in the family of 7373:Rous sarcoma virus 5262:Epstein–Barr virus 5232:Respiratory system 4869:Burkitt's lymphoma 4674:2010-11-04 at the 4045:10.1002/cncr.28862 3908:10.1007/BF01310797 3575:on 7 November 2019 3488:on 3 February 2011 2052:Harper DR (2012). 1260:company. Nineteen 1237: 1185:Vincent Racaniello 1115:poliovirus vaccine 902: 894: 815: 757:respiratory arrest 690:positive selection 666: 404: 338: 292:. The genome is a 7498: 7497: 7436:Taxon identifiers 7427: 7426: 7423: 7422: 7419: 7418: 7415: 7414: 7362: 7361: 7208: 7207: 7204: 7203: 7200: 7199: 7196: 7195: 7143: 7142: 7139: 7138: 7081: 7080: 7077: 7076: 7033: 7032: 7029: 7028: 6985: 6984: 6981: 6980: 6937: 6936: 6933: 6932: 6889: 6888: 6864: 6863: 6753: 6752: 6749: 6748: 6611: 6610: 6491: 6490: 6487: 6486: 6438: 6437: 6351: 6350: 6347: 6346: 6343: 6342: 6271: 6270: 6258: 6257: 6208: 6207: 6121: 6120: 6117: 6116: 6083: 6082: 5981: 5980: 5977: 5976: 5938: 5937: 5904: 5903: 5864: 5863: 5743: 5742: 5705: 5704: 5456: 5455: 5387:Influenza virus A 5304:Human coronavirus 5194: 5193: 5184:Herpes of the eye 4596:(5579): 174–175. 4496:(4523): 916–919. 4410:(5816): 547–553. 4329:978-0-19-920565-3 4280:Maddox B (2003). 4166:(5): 736–751.e8. 4039:(21): 3277–3286. 3415:978-0-470-02386-0 3326:(15): 7902–7912. 3048:978-0-470-02386-0 2737:978-0-300-01324-5 2635:(23): 6221–6229. 2527:(22): 9457–9462. 2270:978-0-470-02386-0 2143:978-0-12-800947-5 2110:978-0-470-02386-0 2063:978-0-8153-4150-5 2027:978-0-470-02386-0 1993:978-0-470-02386-0 1886:(6180): 320–325. 1843:(5209): 415–417. 1604:(10): 4697–4702. 1378:978-0-387-98273-1 1313:Hogle JM (2002). 1296:978-0-8385-8529-0 1173:Rosalind Franklin 1165:X-ray diffraction 925:Rotary Foundation 923:(UNICEF) and the 905:Polio eradication 880:Polio eradication 798:type I interferon 695:Poliovirus is an 675:Old World monkeys 632:was changed from 515:Coxsackie A virus 327:Replication cycle 304:in diameter with 250: 249: 7543: 7491: 7490: 7478: 7477: 7476: 7463: 7462: 7461: 7431: 7430: 7396: 7395: 7387: 7386: 7228: 7227: 7219: 7218: 7166: 7165: 7154: 7153: 7104: 7103: 7092: 7091: 7087:Zaire ebolavirus 7056: 7055: 7044: 7043: 7008: 7007: 6996: 6995: 6960: 6959: 6948: 6947: 6912: 6911: 6900: 6899: 6791: 6790: 6779: 6778: 6770: 6769: 6693:3C-like protease 6628: 6627: 6523: 6522: 6508: 6507: 6464: 6463: 6390: 6389: 6384:Duplornaviricota 6375: 6374: 6362: 6361: 6284: 6283: 6222: 6221: 6182: 6181: 6170: 6169: 6141: 6140: 6094: 6093: 6003: 6002: 5994: 5993: 5949: 5948: 5915: 5914: 5838: 5837: 5826: 5825: 5817: 5816: 5799: 5798: 5770: 5763: 5756: 5747: 5746: 5469: 5468: 5464:digestive system 5381:Orthomyxoviridae 5347:SARS coronavirus 5333:MERS coronavirus 5249: 5248: 5061:Orthomyxoviridae 4984: 4983: 4950:Immune disorders 4874:Hodgkin lymphoma 4850:Kaposi's sarcoma 4758: 4751: 4744: 4735: 4734: 4730: 4718: 4701: 4699: 4698: 4689:. Archived from 4658: 4653: 4652: 4644: 4639: 4638: 4637: 4622: 4621: 4585: 4579: 4578: 4560: 4528: 4522: 4521: 4485: 4479: 4478: 4450: 4444: 4443: 4424:10.1038/291547a0 4399: 4393: 4392: 4382: 4372: 4355:(8): 4887–4891. 4340: 4334: 4333: 4318:Brown A (2007). 4315: 4309: 4308: 4306: 4305: 4277: 4271: 4270: 4268: 4266: 4251: 4245: 4244: 4242: 4241: 4226: 4220: 4219: 4217: 4215: 4200: 4194: 4193: 4183: 4151: 4145: 4144: 4142: 4140: 4126: 4117: 4116: 4114: 4113: 4098: 4092: 4091: 4089: 4088: 4073: 4067: 4066: 4056: 4024: 4018: 4017: 4015: 4014: 3996: 3972: 3966: 3965: 3937: 3928: 3927: 3902:(3–4): 351–363. 3890: 3884: 3883: 3855: 3849: 3848: 3838: 3806: 3800: 3799: 3789: 3779: 3747: 3741: 3740: 3722: 3698: 3692: 3691: 3689: 3688: 3669: 3663: 3662: 3660: 3658: 3643: 3637: 3636: 3626: 3594: 3585: 3584: 3582: 3580: 3571:. Archived from 3565: 3559: 3558: 3556: 3555: 3535: 3529: 3528: 3526: 3524: 3504: 3498: 3497: 3495: 3493: 3474: 3468: 3467: 3465: 3463: 3454:. Archived from 3444: 3438: 3437: 3436:on May 17, 2014. 3432:. Archived from 3426: 3420: 3419: 3401: 3395: 3394: 3384: 3360: 3354: 3353: 3343: 3311: 3305: 3304: 3294: 3277:(7): 4460–4469. 3262: 3256: 3255: 3245: 3235: 3211: 3205: 3204: 3176: 3170: 3169: 3159: 3135: 3129: 3128: 3118: 3094: 3088: 3067: 3061: 3060: 3034: 3028: 3027: 2991: 2985: 2984: 2976: 2970: 2969: 2941: 2935: 2934: 2924: 2900: 2894: 2893: 2883: 2866:(7): 1181–1188. 2851: 2845: 2844: 2834: 2802: 2796: 2795: 2793: 2792: 2777: 2771: 2770: 2768: 2767: 2748: 2742: 2741: 2726:Paul JR (1971). 2723: 2714: 2713: 2703: 2693: 2678:Virology Journal 2669: 2663: 2662: 2652: 2620: 2614: 2613: 2603: 2586:(9): 4429–4440. 2571: 2565: 2564: 2554: 2544: 2512: 2506: 2505: 2495: 2463: 2457: 2456: 2446: 2422: 2416: 2415: 2405: 2373: 2367: 2366: 2338: 2332: 2331: 2329: 2328: 2288: 2275: 2274: 2252: 2246: 2245: 2235: 2203: 2197: 2196: 2186: 2169:(8): 4739–4750. 2154: 2148: 2147: 2121: 2115: 2114: 2092: 2086: 2085: 2074: 2068: 2067: 2049: 2043: 2042: 2040: 2039: 2009: 1998: 1997: 1975: 1969: 1968: 1958: 1941:(8): 2636–2643. 1926: 1920: 1919: 1900:10.1038/334320a0 1875: 1869: 1868: 1832: 1826: 1825: 1797: 1791: 1790: 1762: 1756: 1755: 1745: 1735: 1711: 1705: 1704: 1676: 1667: 1666: 1638: 1632: 1631: 1621: 1589: 1583: 1582: 1572: 1540: 1534: 1533: 1523: 1506:(8): 4827–4835. 1491: 1482: 1481: 1445: 1439: 1438: 1428: 1418: 1403:Virology Journal 1394: 1383: 1382: 1364: 1353: 1352: 1342: 1310: 1301: 1300: 1282: 1169:Birkbeck College 1157:Karl Landsteiner 1140: 1098:, driven by a β- 1088:infection model. 1007: 1003: 999: 972:contact immunity 827:lymphatic system 768:gastrointestinal 701:fecal–oral route 642: 542: 538: 499:coxsackieviruses 375:and immediately 344:-like receptor, 76: 75: 57: 33: 21: 20: 7551: 7550: 7546: 7545: 7544: 7542: 7541: 7540: 7501: 7500: 7499: 7494: 7486: 7481: 7472: 7471: 7466: 7457: 7456: 7451: 7438: 7428: 7411: 7358: 7317:transactivators 7299: 7192: 7135: 7073: 7025: 6977: 6929: 6885: 6860: 6831: 6774:Influenza virus 6764:Negarnaviricota 6761: 6759: 6745: 6671: 6619: 6607: 6556: 6517:Kitrinoviricota 6514: 6499: 6497: 6483: 6455: 6434: 6381: 6366: 6339: 6313: 6279: 6267: 6254: 6204: 6165: 6147: 6133: 6127: 6113: 6079: 6058: 6027: 5989: 5988:(Duplodnaviria, 5987: 5986:circular ds-DNA 5973: 5934: 5900: 5860: 5809: 5803: 5788: 5774: 5744: 5739: 5701: 5681: 5589: 5563:Gastroenteritis 5521: 5518:Gastroenteritis 5512: 5500:Cytomegalovirus 5463: 5452: 5420:Paramyxoviridae 5409:Avian influenza 5285: 5280:Cytomegalovirus 5244:viral pneumonia 5242: 5239:nasopharyngitis 5238: 5235: 5226: 5190: 5164:Cytomegalovirus 5149: 5103: 4991: 4978: 4971: 4944: 4811:Cervical cancer 4771: 4762: 4721: 4709: 4696: 4694: 4676:Wayback Machine 4654: 4647: 4642:Medicine portal 4640: 4635: 4633: 4630: 4625: 4586: 4582: 4529: 4525: 4486: 4482: 4451: 4447: 4400: 4396: 4341: 4337: 4330: 4316: 4312: 4303: 4301: 4294: 4278: 4274: 4264: 4262: 4253: 4252: 4248: 4239: 4237: 4227: 4223: 4213: 4211: 4210:. 12 April 2024 4202: 4201: 4197: 4152: 4148: 4138: 4136: 4128: 4127: 4120: 4111: 4109: 4099: 4095: 4086: 4084: 4075: 4074: 4070: 4025: 4021: 4012: 4010: 3973: 3969: 3938: 3931: 3891: 3887: 3866:(10): 501–506. 3856: 3852: 3807: 3803: 3748: 3744: 3699: 3695: 3686: 3684: 3671: 3670: 3666: 3656: 3654: 3645: 3644: 3640: 3609:(19): 441–446. 3595: 3588: 3578: 3576: 3567: 3566: 3562: 3553: 3551: 3536: 3532: 3522: 3520: 3513:BBC News Online 3505: 3501: 3491: 3489: 3476: 3475: 3471: 3461: 3459: 3458:on 8 March 2016 3446: 3445: 3441: 3428: 3427: 3423: 3416: 3402: 3398: 3361: 3357: 3312: 3308: 3263: 3259: 3226:(3): e1000791. 3212: 3208: 3177: 3173: 3136: 3132: 3095: 3091: 3078: 3068: 3064: 3049: 3035: 3031: 2992: 2988: 2977: 2973: 2942: 2938: 2901: 2897: 2852: 2848: 2817:(19): 441–446. 2803: 2799: 2790: 2788: 2779: 2778: 2774: 2765: 2763: 2750: 2749: 2745: 2738: 2724: 2717: 2670: 2666: 2621: 2617: 2572: 2568: 2513: 2509: 2464: 2460: 2423: 2419: 2374: 2370: 2339: 2335: 2326: 2324: 2289: 2278: 2271: 2253: 2249: 2204: 2200: 2155: 2151: 2144: 2122: 2118: 2111: 2093: 2089: 2078:"Poliomyelitis" 2076: 2075: 2071: 2064: 2050: 2046: 2037: 2035: 2028: 2010: 2001: 1994: 1976: 1972: 1927: 1923: 1876: 1872: 1833: 1829: 1798: 1794: 1763: 1759: 1712: 1708: 1677: 1670: 1639: 1635: 1590: 1586: 1541: 1537: 1492: 1485: 1446: 1442: 1395: 1386: 1379: 1365: 1356: 1311: 1304: 1297: 1283: 1279: 1275: 1244:other viruses. 1229: 1213: 1189:David Baltimore 1181: 1153: 1148: 1138: 1128: 1054:intramuscularly 1034: 1032:Transgenic mice 1029: 1024: 1023: 1005: 1001: 997: 984:immunodeficient 883: 875: 807: 655: 649: 637: 620:) in the genus 616:(later renamed 591:live attenuated 540: 536: 495: 414:2A and 3C/3CD, 329: 199: 187: 135:Pisoniviricetes 70: 17: 12: 11: 5: 7549: 7539: 7538: 7533: 7528: 7523: 7518: 7513: 7496: 7495: 7493: 7492: 7479: 7464: 7448: 7446: 7440: 7439: 7425: 7424: 7421: 7420: 7417: 7416: 7413: 7412: 7410: 7405: 7403: 7393: 7391:Fusion protein 7384: 7383: 7382: 7381: 7380: 7375: 7364: 7363: 7360: 7359: 7357: 7356: 7347: 7342: 7337: 7336: 7335: 7330: 7325: 7312: 7310: 7301: 7300: 7298: 7297: 7296: 7295: 7290: 7278: 7277: 7276: 7274:HIV-1 protease 7271: 7266: 7254: 7253: 7252: 7239: 7237: 7225: 7216: 7210: 7209: 7206: 7205: 7202: 7201: 7198: 7197: 7194: 7193: 7191: 7190: 7189: 7188: 7180:matrix protein 7175: 7173: 7163: 7151: 7145: 7144: 7141: 7140: 7137: 7136: 7134: 7133: 7132: 7131: 7126: 7118:matrix protein 7113: 7111: 7101: 7089: 7083: 7082: 7079: 7078: 7075: 7074: 7072: 7071: 7065: 7063: 7053: 7041: 7035: 7034: 7031: 7030: 7027: 7026: 7024: 7023: 7017: 7015: 7005: 6993: 6987: 6986: 6983: 6982: 6979: 6978: 6976: 6975: 6969: 6967: 6957: 6945: 6939: 6938: 6935: 6934: 6931: 6930: 6928: 6927: 6921: 6919: 6909: 6897: 6891: 6890: 6887: 6886: 6884: 6883: 6877: 6875: 6866: 6865: 6862: 6861: 6859: 6858: 6853: 6848: 6842: 6840: 6833: 6832: 6830: 6829: 6828: 6827: 6819:viral envelope 6815: 6814: 6813: 6805:matrix protein 6800: 6798: 6788: 6776: 6767: 6760:negative-sense 6755: 6754: 6751: 6750: 6747: 6746: 6744: 6743: 6738: 6733: 6728: 6723: 6718: 6713: 6708: 6703: 6698: 6697: 6696: 6684: 6682: 6673: 6672: 6670: 6669: 6664: 6663: 6662: 6657: 6652: 6644:viral envelope 6639: 6637: 6625: 6613: 6612: 6609: 6608: 6606: 6605: 6600: 6595: 6590: 6585: 6580: 6575: 6569: 6567: 6558: 6557: 6555: 6554: 6553: 6552: 6547: 6539:viral envelope 6534: 6532: 6520: 6505: 6498:positive-sense 6493: 6492: 6489: 6488: 6485: 6484: 6482: 6481: 6475: 6473: 6461: 6440: 6439: 6436: 6435: 6433: 6432: 6427: 6422: 6417: 6412: 6407: 6401: 6399: 6387: 6372: 6359: 6353: 6352: 6349: 6348: 6345: 6344: 6341: 6340: 6338: 6337: 6332: 6326: 6324: 6315: 6314: 6312: 6311: 6310: 6300: 6295: 6293: 6281: 6273: 6272: 6269: 6268: 6266: 6265: 6259: 6256: 6255: 6253: 6252: 6242: 6237: 6231: 6229: 6219: 6210: 6209: 6206: 6205: 6203: 6202: 6197: 6191: 6189: 6179: 6167: 6145:Polyomaviridae 6138: 6123: 6122: 6119: 6118: 6115: 6114: 6112: 6111: 6105: 6103: 6091: 6085: 6084: 6081: 6080: 6078: 6077: 6071: 6069: 6060: 6059: 6057: 6056: 6051: 6046: 6040: 6038: 6029: 6028: 6026: 6025: 6020: 6014: 6012: 6000: 5991: 5990:Varidnaviria?) 5983: 5982: 5979: 5978: 5975: 5974: 5972: 5971: 5966: 5960: 5958: 5946: 5940: 5939: 5936: 5935: 5933: 5932: 5926: 5924: 5912: 5906: 5905: 5902: 5901: 5899: 5898: 5893: 5888: 5883: 5877: 5875: 5866: 5865: 5862: 5861: 5859: 5858: 5853: 5847: 5845: 5835: 5823: 5821:Herpes simplex 5814: 5796: 5790: 5789: 5777:Viral proteins 5773: 5772: 5765: 5758: 5750: 5741: 5740: 5738: 5737: 5736: 5735: 5723: 5715: 5713: 5707: 5706: 5703: 5702: 5700: 5699: 5691: 5689: 5683: 5682: 5680: 5679: 5667: 5655: 5643: 5631: 5624: 5618: 5617: 5605: 5599: 5597: 5591: 5590: 5588: 5587: 5580: 5573: 5566: 5554: 5548: 5547: 5546: 5545: 5533: 5527: 5525: 5514: 5513: 5511: 5510: 5509: 5508: 5496: 5495: 5494: 5481: 5479: 5466: 5458: 5457: 5454: 5453: 5451: 5450: 5443: 5436: 5435: 5434: 5414: 5413: 5412: 5411: 5372: 5371: 5370: 5369: 5357: 5356: 5355: 5343: 5342: 5341: 5329: 5328: 5327: 5295: 5293: 5287: 5286: 5284: 5283: 5276: 5275: 5274: 5257: 5255: 5246: 5228: 5227: 5225: 5224: 5223: 5222: 5217: 5204: 5202: 5200:Cardiovascular 5196: 5195: 5192: 5191: 5189: 5188: 5187: 5186: 5174: 5173: 5172: 5159: 5157: 5151: 5150: 5148: 5147: 5146: 5145: 5133: 5132: 5131: 5126: 5113: 5111: 5105: 5104: 5102: 5101: 5096: 5091: 5086: 5085: 5084: 5072: 5071: 5070: 5057: 5052: 5051: 5050: 5038: 5037: 5036: 5024: 5018: 5017: 5016: 5015: 5003: 4997: 4995: 4981: 4979:nervous system 4973: 4972: 4970: 4969: 4968: 4967: 4954: 4952: 4946: 4945: 4943: 4942: 4941: 4940: 4928: 4927: 4926: 4921: 4909: 4903: 4902: 4901: 4900: 4888: 4887: 4886: 4881: 4876: 4871: 4866: 4854: 4853: 4852: 4840: 4839: 4838: 4833: 4831:Vaginal cancer 4828: 4823: 4818: 4813: 4801: 4800: 4799: 4787: 4781: 4779: 4773: 4772: 4761: 4760: 4753: 4746: 4738: 4732: 4731: 4719: 4707: 4702: 4678: 4666: 4660: 4659: 4656:Viruses portal 4645: 4629: 4628:External links 4626: 4624: 4623: 4580: 4523: 4480: 4445: 4394: 4335: 4328: 4310: 4292: 4272: 4246: 4221: 4195: 4146: 4118: 4093: 4068: 4019: 3987:(4): 251–260. 3967: 3929: 3885: 3850: 3821:(2): 681–688. 3801: 3762:(3): 951–955. 3742: 3713:(2): 353–362. 3693: 3664: 3638: 3586: 3560: 3530: 3499: 3469: 3439: 3421: 3414: 3396: 3355: 3306: 3257: 3220:PLOS Pathogens 3206: 3187:(4): 747–752. 3171: 3130: 3109:(2): 421–428. 3089: 3062: 3047: 3029: 2986: 2971: 2936: 2895: 2846: 2797: 2772: 2758:. 2015-09-20. 2743: 2736: 2715: 2664: 2615: 2566: 2507: 2458: 2417: 2388:(3): 433–443. 2368: 2349:(1): 244–264. 2333: 2276: 2269: 2247: 2218:(1): 124–137. 2198: 2149: 2142: 2116: 2109: 2087: 2069: 2062: 2044: 2026: 1999: 1992: 1970: 1921: 1870: 1827: 1792: 1757: 1706: 1687:(2): 175–193. 1681:Virus Research 1668: 1649:(1): 501–505. 1633: 1584: 1555:(4): 505–513. 1535: 1483: 1456:(5): 855–865. 1440: 1384: 1377: 1354: 1302: 1295: 1276: 1274: 1271: 1258:gene synthesis 1228: 1225: 1212: 1209: 1197:Naomi Kitamura 1180: 1177: 1152: 1149: 1147: 1144: 1127: 1124: 1111: 1110: 1092: 1089: 1081: 1033: 1030: 1028: 1025: 951:in ruminants. 947:in humans and 884: 876: 874: 871: 806: 803: 651:Main article: 648: 645: 626:Picornaviridae 624:in the family 606:Picornaviridae 604:in the family 494: 491: 459: 458: 455: 436: 435: 432: 425: 419: 412: 342:immunoglobulin 328: 325: 268:Picornaviridae 248: 247: 246: 245: 233: 221: 206: 205: 201: 200: 195: 193: 189: 188: 181: 179: 175: 174: 167: 163: 162: 159:Picornaviridae 155: 151: 150: 147:Picornavirales 143: 139: 138: 131: 127: 126: 119: 115: 114: 107: 103: 102: 95: 88: 87: 82: 78: 77: 64: 63: 59: 58: 50: 49: 44:of poliovirus 35: 34: 26: 25: 15: 9: 6: 4: 3: 2: 7548: 7537: 7534: 7532: 7529: 7527: 7526:Enteroviruses 7524: 7522: 7519: 7517: 7514: 7512: 7511:Human viruses 7509: 7508: 7506: 7489: 7484: 7480: 7475: 7469: 7465: 7460: 7454: 7450: 7449: 7447: 7445: 7441: 7437: 7432: 7409: 7406: 7404: 7401: 7397: 7394: 7392: 7388: 7385: 7379: 7376: 7374: 7371: 7370: 7369: 7368: 7365: 7355: 7351: 7348: 7346: 7343: 7341: 7338: 7334: 7331: 7329: 7326: 7324: 7321: 7320: 7319: 7318: 7314: 7313: 7311: 7309: 7307: 7302: 7294: 7291: 7289: 7286: 7285: 7284: 7283: 7279: 7275: 7272: 7270: 7267: 7265: 7262: 7261: 7260: 7259: 7255: 7251: 7248: 7247: 7246: 7245: 7241: 7240: 7238: 7236: 7234: 7229: 7226: 7224: 7220: 7217: 7215: 7211: 7187: 7184: 7183: 7182: 7181: 7177: 7176: 7174: 7171: 7167: 7164: 7162: 7160: 7155: 7152: 7150: 7146: 7130: 7127: 7125: 7122: 7121: 7120: 7119: 7115: 7114: 7112: 7109: 7105: 7102: 7100: 7098: 7093: 7090: 7088: 7084: 7070: 7067: 7066: 7064: 7061: 7057: 7054: 7052: 7050: 7045: 7042: 7040: 7036: 7022: 7019: 7018: 7016: 7013: 7009: 7006: 7004: 7002: 6997: 6994: 6992: 6988: 6974: 6971: 6970: 6968: 6965: 6961: 6958: 6956: 6954: 6949: 6946: 6944: 6940: 6926: 6923: 6922: 6920: 6917: 6913: 6910: 6908: 6906: 6901: 6898: 6896: 6895:Parainfluenza 6892: 6882: 6879: 6878: 6876: 6874: 6872: 6867: 6857: 6854: 6852: 6851:Neuraminidase 6849: 6847: 6844: 6843: 6841: 6838: 6834: 6826: 6823: 6822: 6821: 6820: 6816: 6812: 6809: 6808: 6807: 6806: 6802: 6801: 6799: 6796: 6792: 6789: 6787: 6785: 6780: 6777: 6775: 6771: 6768: 6765: 6756: 6742: 6739: 6737: 6734: 6732: 6729: 6727: 6724: 6722: 6719: 6717: 6714: 6712: 6709: 6707: 6704: 6702: 6699: 6694: 6691: 6690: 6689: 6686: 6685: 6683: 6681: 6679: 6674: 6668: 6665: 6661: 6658: 6656: 6653: 6651: 6648: 6647: 6646: 6645: 6641: 6640: 6638: 6636: 6634: 6629: 6626: 6623: 6618: 6614: 6604: 6601: 6599: 6596: 6594: 6591: 6589: 6586: 6584: 6581: 6579: 6576: 6574: 6571: 6570: 6568: 6566: 6564: 6559: 6551: 6548: 6546: 6543: 6542: 6541: 6540: 6536: 6535: 6533: 6531: 6529: 6524: 6521: 6518: 6513: 6509: 6506: 6503: 6494: 6480: 6477: 6476: 6474: 6472: 6470: 6465: 6462: 6459: 6453: 6449: 6445: 6441: 6431: 6428: 6426: 6423: 6421: 6418: 6416: 6413: 6411: 6408: 6406: 6403: 6402: 6400: 6398: 6396: 6391: 6388: 6385: 6380: 6376: 6373: 6370: 6363: 6360: 6358: 6354: 6336: 6333: 6331: 6328: 6327: 6325: 6323: 6321: 6316: 6309: 6306: 6305: 6304: 6301: 6299: 6296: 6294: 6292: 6290: 6285: 6282: 6278: 6274: 6264: 6261: 6260: 6250: 6246: 6243: 6241: 6238: 6236: 6233: 6232: 6230: 6227: 6223: 6220: 6218: 6216: 6211: 6201: 6198: 6196: 6193: 6192: 6190: 6187: 6183: 6180: 6178: 6176: 6171: 6168: 6163: 6159: 6155: 6151: 6146: 6142: 6139: 6136: 6131: 6124: 6110: 6107: 6106: 6104: 6102: 6100: 6095: 6092: 6090: 6089:Baculoviridae 6086: 6076: 6073: 6072: 6070: 6068: 6066: 6061: 6055: 6052: 6050: 6047: 6045: 6042: 6041: 6039: 6037: 6035: 6030: 6024: 6021: 6019: 6016: 6015: 6013: 6011: 6009: 6004: 6001: 5999: 5995: 5992: 5984: 5970: 5967: 5965: 5962: 5961: 5959: 5957: 5955: 5950: 5947: 5945: 5941: 5931: 5928: 5927: 5925: 5923: 5921: 5916: 5913: 5911: 5907: 5897: 5894: 5892: 5889: 5887: 5884: 5882: 5879: 5878: 5876: 5874: 5872: 5867: 5857: 5854: 5852: 5849: 5848: 5846: 5843: 5839: 5836: 5834: 5832: 5827: 5824: 5822: 5818: 5815: 5812: 5807: 5806:Duplodnaviria 5802:linear ds-DNA 5800: 5797: 5795: 5791: 5786: 5782: 5778: 5771: 5766: 5764: 5759: 5757: 5752: 5751: 5748: 5734: 5731: 5730: 5729: 5728: 5724: 5722: 5721: 5717: 5716: 5714: 5712: 5708: 5698: 5697: 5693: 5692: 5690: 5688: 5684: 5677: 5673: 5672: 5668: 5665: 5661: 5660: 5656: 5653: 5649: 5648: 5644: 5641: 5637: 5636: 5632: 5630: 5629: 5625: 5623: 5620: 5619: 5615: 5611: 5610: 5606: 5604: 5601: 5600: 5598: 5596: 5592: 5586: 5585: 5581: 5579: 5578: 5574: 5572: 5571: 5567: 5564: 5560: 5559: 5555: 5553: 5550: 5549: 5544: 5541: 5540: 5539: 5538: 5534: 5532: 5529: 5528: 5526: 5524: 5519: 5515: 5507: 5504: 5503: 5502: 5501: 5497: 5493: 5490: 5489: 5488: 5487: 5483: 5482: 5480: 5478: 5474: 5470: 5467: 5465: 5459: 5449: 5448: 5444: 5442: 5441: 5437: 5433: 5432:Parainfluenza 5430: 5429: 5428: 5427: 5422: 5421: 5416: 5415: 5410: 5406: 5403: 5402: 5401: 5400: 5396: 5392: 5388: 5383: 5382: 5377: 5374: 5373: 5368: 5365: 5364: 5363: 5362: 5358: 5354: 5351: 5350: 5349: 5348: 5344: 5340: 5337: 5336: 5335: 5334: 5330: 5326: 5323: 5322: 5321: 5320: 5316: 5312: 5308: 5305: 5300: 5297: 5296: 5294: 5292: 5288: 5282: 5281: 5277: 5273: 5269: 5268:EBV infection 5266: 5265: 5264: 5263: 5259: 5258: 5256: 5254: 5250: 5247: 5245: 5240: 5233: 5229: 5221: 5218: 5216: 5213: 5212: 5211: 5210: 5206: 5205: 5203: 5201: 5197: 5185: 5182: 5181: 5180: 5179: 5175: 5171: 5168: 5167: 5166: 5165: 5161: 5160: 5158: 5156: 5152: 5144: 5141: 5140: 5139: 5138: 5134: 5130: 5127: 5125: 5124:Poliomyelitis 5122: 5121: 5120: 5119: 5115: 5114: 5112: 5110: 5106: 5100: 5097: 5095: 5092: 5090: 5087: 5083: 5080: 5079: 5078: 5077: 5073: 5069: 5066: 5065: 5063: 5062: 5058: 5056: 5053: 5049: 5046: 5045: 5044: 5043: 5039: 5035: 5032: 5031: 5030: 5029: 5025: 5023: 5020: 5019: 5014: 5011: 5010: 5009: 5008: 5004: 5002: 4999: 4998: 4996: 4994: 4989: 4985: 4982: 4980: 4974: 4966: 4963: 4962: 4961: 4960: 4956: 4955: 4953: 4951: 4947: 4939: 4936: 4935: 4934: 4933: 4929: 4925: 4922: 4920: 4917: 4916: 4915: 4914: 4910: 4908: 4905: 4904: 4899: 4896: 4895: 4894: 4893: 4889: 4885: 4882: 4880: 4877: 4875: 4872: 4870: 4867: 4865: 4862: 4861: 4860: 4859: 4855: 4851: 4848: 4847: 4846: 4845: 4841: 4837: 4834: 4832: 4829: 4827: 4826:Vulvar cancer 4824: 4822: 4821:Penile cancer 4819: 4817: 4814: 4812: 4809: 4808: 4807: 4806: 4802: 4798: 4795: 4794: 4793: 4792: 4788: 4786: 4783: 4782: 4780: 4778: 4774: 4770: 4766: 4759: 4754: 4752: 4747: 4745: 4740: 4739: 4736: 4728: 4724: 4720: 4716: 4712: 4708: 4706: 4703: 4693:on 2011-03-03 4692: 4688: 4684: 4679: 4677: 4673: 4670: 4667: 4665: 4662: 4661: 4657: 4651: 4646: 4643: 4632: 4619: 4615: 4611: 4607: 4603: 4599: 4595: 4591: 4584: 4576: 4572: 4568: 4564: 4559: 4554: 4550: 4546: 4542: 4538: 4534: 4527: 4519: 4515: 4511: 4507: 4503: 4499: 4495: 4491: 4484: 4476: 4472: 4468: 4464: 4460: 4456: 4449: 4441: 4437: 4433: 4429: 4425: 4421: 4417: 4413: 4409: 4405: 4398: 4390: 4386: 4381: 4376: 4371: 4366: 4362: 4358: 4354: 4350: 4346: 4339: 4331: 4325: 4321: 4314: 4299: 4295: 4293:0-00-655211-0 4289: 4285: 4284: 4276: 4261:on 2018-10-30 4260: 4256: 4250: 4236: 4232: 4225: 4209: 4205: 4199: 4191: 4187: 4182: 4177: 4173: 4169: 4165: 4161: 4157: 4150: 4135: 4131: 4125: 4123: 4108: 4104: 4097: 4082: 4078: 4072: 4064: 4060: 4055: 4050: 4046: 4042: 4038: 4034: 4030: 4023: 4008: 4004: 4000: 3995: 3990: 3986: 3982: 3978: 3971: 3963: 3959: 3955: 3951: 3948:(1): 87–100. 3947: 3943: 3936: 3934: 3925: 3921: 3917: 3913: 3909: 3905: 3901: 3897: 3889: 3881: 3877: 3873: 3869: 3865: 3861: 3854: 3846: 3842: 3837: 3832: 3828: 3824: 3820: 3816: 3812: 3805: 3797: 3793: 3788: 3783: 3778: 3773: 3769: 3765: 3761: 3757: 3753: 3746: 3738: 3734: 3730: 3726: 3721: 3716: 3712: 3708: 3704: 3697: 3682: 3678: 3674: 3668: 3652: 3648: 3642: 3634: 3630: 3625: 3620: 3616: 3612: 3608: 3604: 3600: 3593: 3591: 3574: 3570: 3564: 3549: 3545: 3541: 3534: 3518: 3514: 3510: 3503: 3487: 3483: 3479: 3473: 3457: 3453: 3449: 3443: 3435: 3431: 3425: 3417: 3411: 3407: 3400: 3392: 3388: 3383: 3378: 3374: 3370: 3366: 3359: 3351: 3347: 3342: 3337: 3333: 3329: 3325: 3321: 3317: 3310: 3302: 3298: 3293: 3288: 3284: 3280: 3276: 3272: 3268: 3261: 3253: 3249: 3244: 3239: 3234: 3229: 3225: 3221: 3217: 3210: 3202: 3198: 3194: 3190: 3186: 3182: 3175: 3167: 3163: 3158: 3153: 3149: 3145: 3141: 3134: 3126: 3122: 3117: 3112: 3108: 3104: 3100: 3093: 3087: 3083: 3082: 3077: 3073: 3072: 3066: 3058: 3054: 3050: 3044: 3040: 3033: 3025: 3021: 3017: 3013: 3009: 3005: 3001: 2997: 2990: 2982: 2975: 2967: 2963: 2959: 2955: 2951: 2947: 2940: 2932: 2928: 2923: 2918: 2914: 2910: 2906: 2899: 2891: 2887: 2882: 2877: 2873: 2869: 2865: 2861: 2857: 2850: 2842: 2838: 2833: 2828: 2824: 2820: 2816: 2812: 2808: 2801: 2786: 2782: 2776: 2761: 2757: 2753: 2747: 2739: 2733: 2729: 2722: 2720: 2711: 2707: 2702: 2697: 2692: 2687: 2683: 2679: 2675: 2668: 2660: 2656: 2651: 2646: 2642: 2638: 2634: 2630: 2626: 2619: 2611: 2607: 2602: 2597: 2593: 2589: 2585: 2581: 2577: 2570: 2562: 2558: 2553: 2548: 2543: 2538: 2534: 2530: 2526: 2522: 2518: 2511: 2503: 2499: 2494: 2489: 2485: 2481: 2477: 2473: 2469: 2462: 2454: 2450: 2445: 2440: 2436: 2432: 2428: 2421: 2413: 2409: 2404: 2399: 2395: 2391: 2387: 2383: 2379: 2372: 2364: 2360: 2356: 2352: 2348: 2344: 2337: 2322: 2318: 2314: 2310: 2306: 2302: 2298: 2294: 2287: 2285: 2283: 2281: 2272: 2266: 2262: 2258: 2251: 2243: 2239: 2234: 2229: 2225: 2221: 2217: 2213: 2209: 2202: 2194: 2190: 2185: 2180: 2176: 2172: 2168: 2164: 2160: 2153: 2145: 2139: 2135: 2131: 2127: 2120: 2112: 2106: 2102: 2098: 2091: 2083: 2079: 2073: 2065: 2059: 2055: 2048: 2033: 2029: 2023: 2019: 2015: 2008: 2006: 2004: 1995: 1989: 1985: 1981: 1974: 1966: 1962: 1957: 1952: 1948: 1944: 1940: 1936: 1932: 1925: 1917: 1913: 1909: 1905: 1901: 1897: 1893: 1889: 1885: 1881: 1874: 1866: 1862: 1858: 1854: 1850: 1846: 1842: 1838: 1831: 1823: 1819: 1815: 1811: 1807: 1803: 1796: 1788: 1784: 1780: 1776: 1772: 1768: 1761: 1753: 1749: 1744: 1739: 1734: 1729: 1725: 1721: 1717: 1710: 1702: 1698: 1694: 1690: 1686: 1682: 1675: 1673: 1664: 1660: 1656: 1652: 1648: 1644: 1637: 1629: 1625: 1620: 1615: 1611: 1607: 1603: 1599: 1595: 1588: 1580: 1576: 1571: 1566: 1562: 1558: 1554: 1550: 1546: 1539: 1531: 1527: 1522: 1517: 1513: 1509: 1505: 1501: 1497: 1490: 1488: 1479: 1475: 1471: 1467: 1463: 1459: 1455: 1451: 1444: 1436: 1432: 1427: 1422: 1417: 1412: 1408: 1404: 1400: 1393: 1391: 1389: 1380: 1374: 1370: 1363: 1361: 1359: 1350: 1346: 1341: 1336: 1332: 1328: 1324: 1320: 1316: 1309: 1307: 1298: 1292: 1288: 1281: 1277: 1270: 1267: 1263: 1259: 1254: 1250: 1249:Eckard Wimmer 1245: 1242: 1233: 1224: 1222: 1218: 1208: 1206: 1202: 1201:Eckard Wimmer 1198: 1194: 1190: 1186: 1176: 1174: 1170: 1167:by a team at 1166: 1162: 1158: 1143: 1135: 1133: 1123: 1120: 1116: 1108: 1104: 1101: 1097: 1093: 1090: 1087: 1082: 1078: 1074: 1073: 1072: 1069: 1067: 1063: 1059: 1055: 1051: 1050:intravenously 1047: 1042: 1040: 1021: 1017: 1013: 1009: 995: 991: 987: 985: 981: 977: 973: 969: 965: 961: 960:polio vaccine 957: 952: 950: 946: 942: 938: 934: 930: 926: 922: 918: 914: 913:public health 910: 909:poliomyelitis 906: 898: 890: 886: 881: 870: 867: 863: 859: 855: 851: 847: 843: 842:polio vaccine 839: 834: 832: 828: 824: 820: 819:immune system 811: 802: 799: 795: 789: 787: 783: 779: 774: 769: 765: 760: 758: 754: 750: 746: 742: 738: 734: 733:motor neurons 730: 726: 722: 718: 714: 710: 706: 702: 698: 693: 691: 687: 682: 680: 676: 672: 664:of poliovirus 663: 659: 654: 644: 641: 640:Enterovirus C 635: 631: 627: 623: 619: 618:Enterovirus C 615: 611: 607: 603: 598: 596: 592: 588: 584: 579: 577: 573: 569: 565: 560: 558: 554: 550: 546: 533: 531: 527: 523: 519: 516: 512: 508: 504: 500: 490: 488: 484: 480: 476: 470: 468: 464: 456: 452: 451: 450: 447: 444: 440: 433: 430: 429:, 3AB, 3A, 3B 426: 423: 420: 417: 413: 410: 406: 405: 400: 396: 394: 389: 387: 383: 378: 374: 373:messenger RNA 370: 365: 363: 359: 355: 351: 350:cell membrane 347: 343: 333: 324: 322: 317: 315: 314:encapsulating 311: 307: 303: 299: 295: 291: 288: 284: 281: 276: 274: 271:. There are 270: 269: 264: 263: 262:Enterovirus C 258: 254: 243: 242: 237: 234: 231: 230: 225: 222: 219: 218: 213: 210: 209: 207: 202: 198: 194: 191: 190: 186: 185: 184:Enterovirus C 180: 177: 176: 173: 172: 168: 165: 164: 161: 160: 156: 153: 152: 149: 148: 144: 141: 140: 137: 136: 132: 129: 128: 125: 124: 120: 117: 116: 113: 112: 111:Orthornavirae 108: 105: 104: 101: 100: 96: 93: 90: 89: 86: 83: 80: 79: 74: 69: 65: 60: 56: 51: 47: 43: 40: 36: 32: 27: 22: 19: 7443: 7315: 7304: 7280: 7256: 7242: 7231: 7178: 7157: 7116: 7095: 7060:glycoprotein 7047: 7012:glycoprotein 6999: 6964:glycoprotein 6951: 6916:glycoprotein 6903: 6869: 6837:glycoprotein 6817: 6803: 6782: 6676: 6667:Nucleocapsid 6642: 6631: 6622:Pisuviricota 6561: 6537: 6526: 6467: 6458:Pisuviricota 6447: 6393: 6318: 6287: 6213: 6173: 6135:Monodnaviria 6097: 6063: 6032: 6006: 5998:Epstein–Barr 5952: 5944:Adenoviridae 5918: 5869: 5829: 5811:Varidnaviria 5725: 5718: 5694: 5687:Pancreatitis 5669: 5657: 5645: 5633: 5626: 5607: 5582: 5575: 5568: 5556: 5535: 5498: 5484: 5445: 5438: 5424: 5418: 5385: 5379: 5359: 5345: 5331: 5302: 5278: 5260: 5215:Pericarditis 5207: 5176: 5162: 5135: 5117: 5116: 5074: 5059: 5040: 5026: 5005: 4988:Encephalitis 4957: 4930: 4911: 4890: 4856: 4842: 4803: 4789: 4726: 4714: 4695:. Retrieved 4691:the original 4686: 4681:Goodsell D. 4593: 4589: 4583: 4540: 4536: 4526: 4493: 4489: 4483: 4458: 4454: 4448: 4407: 4403: 4397: 4352: 4348: 4338: 4319: 4313: 4302:. Retrieved 4282: 4275: 4263:. Retrieved 4259:the original 4249: 4238:. Retrieved 4234: 4224: 4212:. Retrieved 4207: 4198: 4163: 4159: 4149: 4137:. Retrieved 4133: 4110:. Retrieved 4106: 4096: 4085:. Retrieved 4083:. 2022-09-01 4080: 4071: 4036: 4032: 4022: 4011:. Retrieved 3984: 3980: 3970: 3945: 3941: 3899: 3895: 3888: 3863: 3859: 3853: 3818: 3814: 3804: 3759: 3755: 3745: 3710: 3706: 3696: 3685:. Retrieved 3676: 3667: 3655:. Retrieved 3641: 3606: 3602: 3577:. Retrieved 3573:the original 3563: 3552:. Retrieved 3543: 3533: 3521:. Retrieved 3502: 3490:. Retrieved 3486:the original 3472: 3460:. 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Retrieved 2017: 1983: 1973: 1938: 1934: 1924: 1883: 1879: 1873: 1840: 1836: 1830: 1805: 1801: 1795: 1770: 1766: 1760: 1723: 1720:PLOS Biology 1719: 1709: 1684: 1680: 1646: 1642: 1636: 1601: 1597: 1587: 1552: 1548: 1538: 1503: 1499: 1453: 1449: 1443: 1406: 1402: 1368: 1322: 1318: 1286: 1280: 1251:'s group at 1246: 1238: 1214: 1182: 1161:Erwin Popper 1154: 1136: 1129: 1112: 1070: 1043: 1035: 988: 980:asymptomatic 953: 903: 885: 838:immunization 835: 816: 790: 764:neurological 761: 745:motor cortex 723:tissue, and 713:asymptomatic 694: 685: 683: 667: 647:Pathogenesis 639: 633: 629: 625: 621: 617: 613: 609: 605: 601: 599: 580: 571: 561: 534: 511:Phylogenetic 507:rhinoviruses 496: 471: 460: 448: 445: 441: 437: 428: 390: 366: 339: 318: 310:nonenveloped 277: 266: 260: 252: 251: 241:Poliovirus-3 239: 235: 229:Poliovirus-2 227: 223: 217:Poliovirus-1 215: 211: 196: 182: 170: 158: 146: 134: 123:Pisuviricota 122: 110: 98: 91: 81:(unranked): 18: 7468:Wikispecies 7400:oncoprotein 6512:Hepatitis C 6277:Hepatitis B 6263:Agnoprotein 6226:oncoprotein 6200:VP2 and VP3 5584:Coronavirus 5325:Common cold 5237:acute viral 5220:Myocarditis 5064:(probable) 4816:Anal cancer 4265:4 September 3677:www.who.int 3462:11 February 3375:(1): 9–16. 2981:Polioviruse 2303:: 587–635. 1773:: 271–292. 1726:(7): e183. 1325:: 677–702. 1217:James Hogle 1058:spinal cord 1012:Afghanistan 968:inactivated 956:vaccination 919:(WHO), the 873:Eradication 786:macrophages 737:spinal cord 735:within the 697:enterovirus 679:chimpanzees 630:Enterovirus 622:Enterovirus 608:. In 2008, 602:Enterovirus 503:echoviruses 393:polypeptide 382:5' end 321:RNA viruses 298:nucleotides 171:Enterovirus 24:Poliovirus 16:Enterovirus 7505:Categories 7474:Poliovirus 7444:Poliovirus 6825:M2 protein 6811:M1 protein 6617:SARS-CoV-2 5711:Urogenital 5577:Astrovirus 5537:Adenovirus 5361:SARS-CoV-2 5118:Poliovirus 4993:meningitis 4697:2010-01-07 4304:2020-05-15 4240:2024-08-10 4112:2024-08-10 4087:2024-08-10 4013:2022-04-01 3687:2020-08-28 3579:2 November 3554:2020-08-25 3523:14 October 3492:9 November 2791:2019-11-02 2766:2015-09-30 2327:2018-10-26 2038:2016-10-19 1273:References 1077:chromosome 1046:transgenic 1039:engineered 1008:1 (WPV1). 949:rinderpest 931:(CDC) and 900:Poliovirus 850:antibodies 844:, develop 831:antibodies 753:antibodies 741:brain stem 634:Poliovirus 610:Poliovirus 576:infectious 562:The three 522:speciation 377:translated 253:Poliovirus 204:Serotypes 197:Poliovirus 42:micrograph 7459:Q12438936 7367:Multiple 7264:Integrase 6502:Riboviria 6379:Rotavirus 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