Piscine orthoreovirus: Difference between revisions
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Piscine orthoreovirus (PRV) is an Orthoreovirus species that infects fish species exclusively , and has been found present at higher concentration in fish with various diseases.<ref name=" |
Piscine orthoreovirus (PRV) is an Orthoreovirus species that infects fish species exclusively , and has been found present at higher concentration in fish with various diseases.<ref name="Palacios_2010">{{cite journal | vauthors = Palacios G, Lovoll M, Tengs T, Hornig M, Hutchison S, Hui J, Kongtorp RT, Savji N, Bussetti AV, Solovyov A, Kristoffersen AB, Celone C, Street C, Trifonov V, Hirschberg DL, Rabadan R, Egholm M, Rimstad E, Lipkin WI | display-authors = 6 | title = Heart and skeletal muscle inflammation of farmed salmon is associated with infection with a novel reovirus | journal = PloS One | volume = 5 | issue = 7 | pages = e11487 | date = July 2010 | pmid = 20634888 | pmc = 2901333 | doi = 10.1371/journal.pone.0011487 }}</ref> These diseases include heart and skeletal muscle inflammation (HSMI), jaundice syndrome, proliferative darkening syndrome and erythrocytic body inclusion syndrome.<ref name="Palacios_2010" /><ref name="Takano_2016">{{cite journal | vauthors = Takano T, Nawata A, Sakai T, Matsuyama T, Ito T, Kurita J, Terashima S, Yasuike M, Nakamura Y, Fujiwara A, Kumagai A, Nakayasu C | display-authors = 6 | title = Full-Genome Sequencing and Confirmation of the Causative Agent of Erythrocytic Inclusion Body Syndrome in Coho Salmon Identifies a New Type of Piscine Orthoreovirus | journal = PloS One | volume = 11 | issue = 10 | pages = e0165424 | date = 2016-10-27 | pmid = 27788206 | pmc = 5082797 | doi = 10.1371/journal.pone.0165424 }}</ref><ref name="Garseth_2013">{{cite journal | vauthors = Garseth ÅH, Ekrem T, Biering E | title = Phylogenetic evidence of long distance dispersal and transmission of piscine reovirus (PRV) between farmed and wild Atlantic salmon | journal = PloS One | volume = 8 | issue = 12 | pages = e82202 | date = 2013-12-11 | pmid = 24349221 | doi = 10.1371/journal.pone.0082202 }}</ref><ref name = "Cartagena_2020">{{Cite journal| vauthors = Cartagena J, Jiménez C, Spencer E |date= November 2020 |title=Detection of Piscine orthoreoviruses (PRV-1b AND PRV-3a) in farmed Coho salmon with jaundice syndrome from Chile |journal=Aquaculture|volume=528|pages=735480|doi=10.1016/j.aquaculture.2020.735480|issn=0044-8486}}</ref> PRV is thought to mainly affect [[Aquaculture|aquacultured]] and [[Mariculture|maricultured]] fish stocks, and recent research has been focused around the susceptibility of wild stock. However, whether PRV is virulent, specifically with respect to HSMI, it has not been unanimously agreed upon.<ref>{{cite journal | vauthors = Zhang Y, Polinski MP, Morrison PR, Brauner CJ, Farrell AP, Garver KA | title = High-Load Reovirus Infections Do Not Imply Physiological Impairment in Salmon | language = English | journal = Frontiers in Physiology | volume = 10 | pages = 114 | date = 2019 | pmid = 30930782 | doi = 10.3389/fphys.2019.00114 }}</ref> PRV has been in the public eye mostly due to a potential linkage to farmed Atlantic Salmon exhibiting HSMI.<ref name="Palacios_2010" /> Public concern has been raised regarding the possibility of open ocean net farms transmitting PRV to wild populations being a factor in their recent decline.<ref>{{Cite journal| vauthors = Noakes DJ, Beamish RJ, Kent ML |date= March 2000 |title=On the decline of Pacific salmon and speculative links to salmon farming in British Columbia |journal=Aquaculture|volume=183|issue=3-4|pages=363–386|doi=10.1016/s0044-8486(99)00294-x|issn=0044-8486}}</ref> PRV is not confirmed to be pathogenic in wild salmon stocks. |
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== Classification == |
== Classification == |
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Phylogenetic analysis of each segment of the PRV genome initially placed PRV in the [[Reoviridae]] family, subfamily [[Spinareovirinae]], based on sequence and structural similarity to known [[Reoviruses]].<ref name=" |
Phylogenetic analysis of each segment of the PRV genome initially placed PRV in the [[Reoviridae]] family, subfamily [[Spinareovirinae]], based on sequence and structural similarity to known [[Reoviruses]].<ref name="Palacios_2010" /> Phylogenetic sequence analysis indicated PRV was equally related to the genuses [[Orthoreovirus|Orthoreoviridae]] and [[Aquareovirus|Aquareoviridae]] upon initial discovery.<ref name="Palacios_2010" /> As a result PRV was first believed to have evolved separately from a common ancestor related to both [[Orthoreovirus|Orthoreoviridae]] and [[Aquareovirus|Aquareoviridae]] in the subfamily [[Spinareovirinae]]. |
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Currently, PRV is officially classified as an Orthoreovirus.<ref>{{Cite web|title=ICTV|url=https://talk.ictvonline.org/taxonomy/p/taxonomy-history?taxnode_id=201904974|access-date=2021-03-16|website=talk.ictvonline.org}}</ref> Placement into the genus Orthoreoviridae and not Aquareoviridae has been argued for the following reasons: Higher sequence fidelity within homologous sequences<ref name=" |
Currently, PRV is officially classified as an Orthoreovirus.<ref>{{Cite web|title=ICTV|url=https://talk.ictvonline.org/taxonomy/p/taxonomy-history?taxnode_id=201904974|access-date=2021-03-16|website=talk.ictvonline.org}}</ref> Placement into the genus Orthoreoviridae and not Aquareoviridae has been argued for the following reasons: Higher sequence fidelity within homologous sequences<ref name="Kibenge_2013" />, the same segment numbers, lack of a fusogenic-associated small transmembrane protein (FAST)<ref name=":4" />, presence of a fiber viral attachment protein<ref name="Markussen_2013" />, lack of syncytia formation and GC nucleoside percentage. <ref name="Markussen_2013" /> |
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Opposing arguments to this placement include fact that the only other non-fusogenic orthoreovirus species is mammalian Orthoreovirus (MRV), and that while aquareoviridae have many known fish viruses orthoreoviridae doesn't have any other known.<ref name=":11">{{ |
Opposing arguments to this placement include fact that the only other non-fusogenic orthoreovirus species is mammalian Orthoreovirus (MRV), and that while aquareoviridae have many known fish viruses orthoreoviridae doesn't have any other known.<ref name=":11">{{cite journal | vauthors = Attoui H, Fang Q, Jaafar FM, Cantaloube JF, Biagini P, de Micco P, de Lamballerie X | title = Common evolutionary origin of aquareoviruses and orthoreoviruses revealed by genome characterization of Golden shiner reovirus, Grass carp reovirus, Striped bass reovirus and golden ide reovirus (genus Aquareovirus, family Reoviridae) | journal = The Journal of General Virology | volume = 83 | issue = Pt 8 | pages = 1941–1951 | date = August 2002 | pmid = 12124458 | doi = 10.1099/0022-1317-83-8-1941 }}</ref><ref>{{cite journal | vauthors = Duncan R | title = Extensive sequence divergence and phylogenetic relationships between the fusogenic and nonfusogenic orthoreoviruses: a species proposal | journal = Virology | volume = 260 | issue = 2 | pages = 316–28 | date = August 1999 | pmid = 10417266 | doi = 10.1006/viro.1999.9832 }}</ref> Other arguments include the fact that the S1 and S4 sequences in PRV have no known homologues to either genus,<ref name="Palacios_2010" /> and that almost all orthoreoviridae and aquareoviridae are bicistronic for their viral attachment protein, while PRV is monocistronic.<ref name=":8" /><ref name="Markussen_2013" /> PRV exhibits 5' terminal sequences on its segments that don't align to either genus.<ref name="Palacios_2010" /> There have been a few species of aquareoviridae identified as non-fusogenic, including GCRV104 and GCRVGD108.<ref>{{cite journal | vauthors = Nibert ML, Duncan R | title = Bioinformatics of recent aqua- and orthoreovirus isolates from fish: evolutionary gain or loss of FAST and fiber proteins and taxonomic implications | journal = PloS One | volume = 8 | issue = 7 | pages = e68607 | date = 2013-07-04 | pmid = 23861926 | doi = 10.1371/journal.pone.0068607 }}</ref> The outer clamp protein of PRV is found on the bicistronic segment S1, the first of any known orthoreovirus or aquareovirus to encode this protein on a polycistronic segment.<ref name=":4" /> |
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== Genome and Structure == |
== Genome and Structure == |
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=== Genome === |
=== Genome === |
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Piscine orthoreovirus has a segmented [[Baltimore classification|dsRNA]] genome made up of 10 individual linear segments, cumulatively measuring around 23,600bp.<ref name=" |
Piscine orthoreovirus has a segmented [[Baltimore classification|dsRNA]] genome made up of 10 individual linear segments, cumulatively measuring around 23,600bp.<ref name="Palacios_2010" /><ref name="Markussen_2013">{{cite journal | vauthors = Markussen T, Dahle MK, Tengs T, Løvoll M, Finstad ØW, Wiik-Nielsen CR, Grove S, Lauksund S, Robertsen B, Rimstad E | display-authors = 6 | title = Sequence analysis of the genome of piscine orthoreovirus (PRV) associated with heart and skeletal muscle inflammation (HSMI) in Atlantic salmon (Salmo salar) | journal = PloS One | volume = 8 | issue = 7 | pages = e70075 | date = 2013-07-29 | pmid = 23922911 | pmc = 3726481 | doi = 10.1371/journal.pone.0070075 }}</ref> It has a [[GC-content|GC content]] of 47%. Each segment has conserved terminal sequences. The 3' end sequence (UCAUC-3') is the same as Orthoreoviridae and Aquareoviridae. The 5' end sequence (5'-GAUAAA/U) shows complete uniqueness within PRV. <ref name="Palacios_2010" /><ref name=":11" /> These segments are referred to as L1-3 (Long), M1-3(Medium) and S1-4 (Short) based on length and comparison with homologous segments in orthoreoviridae and aquareoviridae. L1 is the longest at 3916bp and S4 the shortest at 1040bp.<ref name="Palacios_2010" /><ref name="Markussen_2013" /><ref name="Kibenge_2013">{{cite journal | vauthors = Kibenge MJ, Iwamoto T, Wang Y, Morton A, Godoy MG, Kibenge FS | title = Whole-genome analysis of piscine reovirus (PRV) shows PRV represents a new genus in family Reoviridae and its genome segment S1 sequences group it into two separate sub-genotypes | journal = Virology Journal | volume = 10 | issue = 1 | pages = 230 | date = July 2013 | pmid = 23844948 | pmc = 3711887 | doi = 10.1186/1743-422X-10-230 }}</ref> |
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11 proteins are confirmed to be encoded.<ref name=" |
11 proteins are confirmed to be encoded.<ref name="Markussen_2013" /> S1 exhibits bicistronicity with 2 overlapping open reading frames, while the remaining proteins are either confirmed as monocistronic or have been thought to be bicistronic with no further evidence. <ref name="Kibenge_2013" /><ref name=":4">{{cite journal | vauthors = Key T, Read J, Nibert ML, Duncan R | title = Piscine reovirus encodes a cytotoxic, non-fusogenic, integral membrane protein and previously unrecognized virion outer-capsid proteins | journal = The Journal of General Virology | volume = 94 | issue = Pt 5 | pages = 1039–1050 | date = May 2013 | pmid = 23343626 | doi = 10.1099/vir.0.048637-0 }}</ref><ref name="Markussen_2013" /> The proteins that each segment encodes for are as follows, using a standardized naming system across reovirus genera:<ref name="Markussen_2013" /><ref name=":8">{{cite journal | vauthors = Yan X, Parent KN, Goodman RP, Tang J, Shou J, Nibert ML, Duncan R, Baker TS | display-authors = 6 | title = Virion structure of baboon reovirus, a fusogenic orthoreovirus that lacks an adhesion fiber | journal = Journal of Virology | volume = 85 | issue = 15 | pages = 7483–95 | date = August 2011 | pmid = 21593159 | doi = 10.1128/JVI.00729-11 }}</ref> |
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* L1-λ3 - Shell Protein (Inner capsid protein) |
* L1-λ3 - Shell Protein (Inner capsid protein) |
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* L2-λ2 - Turret protein <ref name=" |
* L2-λ2 - Turret protein <ref name="Markussen_2013" /> |
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* L3-λ1 - [[RNA-dependent RNA polymerase|RdRP]] |
* L3-λ1 - [[RNA-dependent RNA polymerase|RdRP]] |
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* M1-μ2 - [[Nucleoside-triphosphatase|NTPase]] |
* M1-μ2 - [[Nucleoside-triphosphatase|NTPase]] |
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* M2-μ1 - Outer capsid protein |
* M2-μ1 - Outer capsid protein |
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* M3-μNS - unknown function, <ref>{{ |
* M3-μNS - unknown function, <ref>{{cite journal | vauthors = Haatveit HM, Nyman IB, Markussen T, Wessel Ø, Dahle MK, Rimstad E | title = The non-structural protein μNS of piscine orthoreovirus (PRV) forms viral factory-like structures | journal = Veterinary Research | volume = 47 | issue = 1 | pages = 5 | date = January 2016 | pmid = 26743679 | pmc = 4705589 | doi = 10.1186/s13567-015-0302-0 }}</ref> |
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* S1-σ3 - Outer Clamp Protein (Capsid protein) |
* S1-σ3 - Outer Clamp Protein (Capsid protein) |
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* p13 - Cytotoxic Nonstructural Protein<ref name=":3">{{ |
* p13 - Cytotoxic Nonstructural Protein<ref name=":3">{{cite journal | vauthors = Kuehn R, Stoeckle BC, Young M, Popp L, Taeubert JE, Pfaffl MW, Geist J | title = Identification of a piscine reovirus-related pathogen in proliferative darkening syndrome (PDS) infected brown trout (Salmo trutta fario) using a next-generation technology detection pipeline | journal = PloS One | volume = 13 | issue = 10 | pages = e0206164 | date = 2018-10-22 | pmid = 30346982 | pmc = 6197672 | doi = 10.1371/journal.pone.0206164 }}</ref> |
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* S2-σ2 - Core Clamp Protein |
* S2-σ2 - Core Clamp Protein |
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* S3-σNS - unknown function |
* S3-σNS - unknown function |
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* S4-σ1 - Viral Attachment Protein<ref name=" |
* S4-σ1 - Viral Attachment Protein<ref name="Markussen_2013" /> |
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=== Subdivisions === |
=== Subdivisions === |
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[[File:PRV M2 Sequence compariston.png|thumb|Dendogram of M2 segment nucleotide sequences for different PRV genotypes and subgenotypes.|405x405px]]Piscine reovirus has been grouped into multiple different genotypes based on sequence diversity. Although multiple ways of subdividing PRV have been proposed,<ref name=" |
[[File:PRV M2 Sequence compariston.png|thumb|Dendogram of M2 segment nucleotide sequences for different PRV genotypes and subgenotypes.|405x405px]]Piscine reovirus has been grouped into multiple different genotypes based on sequence diversity. Although multiple ways of subdividing PRV have been proposed,<ref name="Garseth_2013" /> the system most often used in the literature subdivides it into 2 genotypes, I and II, which further divide into Ia and Ib, and IIa and IIb, respectively.<ref name="Godoy_2021">{{cite journal | vauthors = Godoy M, Medina DA, Suarez R, Valenzuela S, Romero J, Kibenge M, Wang Y, Kibenge F | display-authors = 6 | title = Extensive Phylogenetic Analysis of Piscine Orthoreovirus Genomic Sequences Shows the Robustness of Subgenotype Classification | journal = Pathogens | volume = 10 | issue = 1 | pages = 41 | date = January 2021 | pmid = 33430212 | doi = 10.3390/pathogens10010041 }}</ref><ref name="Takano_2016" /><ref name=":10">{{cite journal | vauthors = Godoy MG, Kibenge MJ, Wang Y, Suarez R, Leiva C, Vallejos F, Kibenge FS | title = First description of clinical presentation of piscine orthoreovirus (PRV) infections in salmonid aquaculture in Chile and identification of a second genotype (Genotype II) of PRV | journal = Virology Journal | volume = 13 | issue = 1 | pages = 98 | date = June 2016 | pmid = 27296722 | pmc = 4906990 | doi = 10.1186/s12985-016-0554-y }}</ref><ref name="Kibenge_2019">{{cite journal | vauthors = Kibenge FS | title = Emerging viruses in aquaculture | journal = Current Opinion in Virology | volume = 34 | pages = 97–103 | date = February 2019 | pmid = 30711892 | doi = 10.1016/j.coviro.2018.12.008 | series = Emerging viruses: intraspecies transmission • Viral Immunology }}</ref> These divisions and subdivisions are based on sequence diversity within segment S1.<ref name=":10" /><ref name="Kibenge_2013" /> |
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==== Genotype I (PRV-1) <ref name=" |
==== Genotype I (PRV-1) <ref name="Kibenge_2013" /> ==== |
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====== Ia ====== |
====== Ia ====== |
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This sub genotype of PRV is found primarily in farmed Atlantic salmon in Norway, Chile and Canada. It has been associated with populations exhibiting HSMI. It has been found in farmed [[Chinook salmon]] in Canada exhibiting Jaundice syndrome, as well as farmed [[Rainbow trout]] and Coho showing HSMI like symptoms in Chile and Canada.<ref>{{Cite journal| |
This sub genotype of PRV is found primarily in farmed Atlantic salmon in Norway, Chile and Canada. It has been associated with populations exhibiting HSMI. It has been found in farmed [[Chinook salmon]] in Canada exhibiting Jaundice syndrome, as well as farmed [[Rainbow trout]] and Coho showing HSMI like symptoms in Chile and Canada.<ref>{{Cite journal| vauthors = Di Cicco E, Ferguson HW, Kaukinen KH, Schulze AD, Li S, Tabata A, Günther OP, Mordecai G, Suttle CA, Miller KM | display-authors = 6 |date=2018-10-01|title=The same strain of Piscine orthoreovirus (PRV-1) is involved in the development of different, but related, diseases in Atlantic and Pacific Salmon in British Columbia |journal=FACETS|volume=3|issue=1|pages=599–641|doi=10.1139/facets-2018-0008|issn=2371-1671}}</ref> |
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====== Ib ====== |
====== Ib ====== |
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This sub genotype of PRV is found primarily in farmed Atlantic and Coho salmon in Norway and Chile, additionally being found in farmed Rainbow trout in Chile.<ref |
This sub genotype of PRV is found primarily in farmed Atlantic and Coho salmon in Norway and Chile, additionally being found in farmed Rainbow trout in Chile.<ref name = "Cartagena_2020" /> It has been present in many salmon with HSMI. It has also been found in Coho populations exhibiting Jaundice syndrome. |
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==== Genotype 2 <ref name=":10" /> ==== |
==== Genotype 2 <ref name=":10" /> ==== |
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Genotype 2 was first discovered when comparing the S1 sequences among farmed Atlantic salmon in Chile, and further reiterated with both M2 and whole genome analysis. |
Genotype 2 was first discovered when comparing the S1 sequences among farmed Atlantic salmon in Chile, and further reiterated with both M2 and whole genome analysis.<ref name="Polinski_2020">{{cite journal | vauthors = Polinski MP, Vendramin N, Cuenca A, Garver KA | title = Piscine orthoreovirus: Biology and distribution in farmed and wild fish | journal = Journal of Fish Diseases | volume = 43 | issue = 11 | pages = 1331–1352 | date = November 2020 | pmid = 32935367 | doi = 10.1111/jfd.13228 }}</ref><ref name="Godoy_2021"/> Despite being within one genotype, the sub genotypes IIa and IIb exhibit much higher inter-sequence diversity than do Ia and Ib. |
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====== IIa (PRV-3) ====== |
====== IIa (PRV-3) ====== |
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PRV-3 has been associated with pathological heart lesions in rainbow trout.<ref>{{ |
PRV-3 has been associated with pathological heart lesions in rainbow trout.<ref>{{cite journal | vauthors = Vendramin N, Kannimuthu D, Olsen AB, Cuenca A, Teige LH, Wessel Ø, Iburg TM, Dahle MK, Rimstad E, Olesen NJ | display-authors = 6 | title = Piscine orthoreovirus subtype 3 (PRV-3) causes heart inflammation in rainbow trout (Oncorhynchus mykiss) | journal = Veterinary Research | volume = 50 | issue = 1 | pages = 14 | date = February 2019 | pmid = 30777130 | doi = 10.1186/s13567-019-0632-4 }}</ref><ref name="Godoy_2021" /> It has also been found in farmed Coho Salmon, [[Brown trout]], and Rainbow trout exhibiting HSMI across Northern Europe and Chile.<ref name="Kibenge_2019" /><ref name="Kibenge_2019" /> It has also been found in wild brown trout with proliferative darkening syndrome in Central Europe.<ref name=":3" /> Comparison with PRV-2 shows an 80.1% and 90.5% similarity for nucleotides and amino acids, respectively.<ref name=":12">{{cite journal | vauthors = Dhamotharan K, Vendramin N, Markussen T, Wessel Ø, Cuenca A, Nyman IB, Olsen AB, Tengs T, Krudtaa Dahle M, Rimstad E | display-authors = 6 | title = Molecular and Antigenic Characterization of Piscine orthoreovirus (PRV) from Rainbow Trout (Oncorhynchus mykiss) | journal = Viruses | volume = 10 | issue = 4 | date = April 2018 | pmid = 29614838 | pmc = 5923464 | doi = 10.3390/v10040170 }}</ref> Comparison with PRV-1 showed a 72.9% similarity in nucleotide sequence and 80.0% amino acid sequence similarity. |
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====== IIb (PRV-2) ====== |
====== IIb (PRV-2) ====== |
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PVR-2 has only been found in farmed Coho salmon in Japan exhibiting Erythrocytic Inclusion Body Syndrome (EIBS).<ref name=" |
PVR-2 has only been found in farmed Coho salmon in Japan exhibiting Erythrocytic Inclusion Body Syndrome (EIBS).<ref name="Takano_2016" /> Only one genome of PRV-2 has been sequenced thus far, which has reduced confidence in phylogenetic placements.<ref name=":4" /> Comparison with PRV-1 showed a 73.4% and 80.3% similarity in nucleotide sequence and amino acid sequence, respectively.<ref name=":12" /> |
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=== Structure === |
=== Structure === |
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[[File:PRV virion proteins.png|thumb|440x440px|Electrogram image of PRV with immunogold labelled proteins in black arrows. Left: Viral attachment protein (σ1) on outer capsid. Right: Outer clamp protein (σ3) on outer capsid.]] |
[[File:PRV virion proteins.png|thumb|440x440px|Electrogram image of PRV with immunogold labelled proteins in black arrows. Left: Viral attachment protein (σ1) on outer capsid. Right: Outer clamp protein (σ3) on outer capsid.]] |
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As a member of genus orthoreovirus, the virion form of PRV is a non-enveloped icosahedral capsid with two layers, the outer and core. The diameter of the total virion encompassing the outer layer is approximately 70nm, with the inner core layer measuring around 39nm in diameter.<ref>{{ |
As a member of genus orthoreovirus, the virion form of PRV is a non-enveloped icosahedral capsid with two layers, the outer and core. The diameter of the total virion encompassing the outer layer is approximately 70nm, with the inner core layer measuring around 39nm in diameter.<ref>{{cite journal | vauthors = Wessel Ø, Braaen S, Alarcon M, Haatveit H, Roos N, Markussen T, Tengs T, Dahle MK, Rimstad E | display-authors = 6 | title = Infection with purified Piscine orthoreovirus demonstrates a causal relationship with heart and skeletal muscle inflammation in Atlantic salmon | journal = PloS One | volume = 12 | issue = 8 | pages = e0183781 | date = 2017-08-25 | pmid = 28841684 | doi = 10.1371/journal.pone.0183781 }}</ref> |
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| ⚫ | The inner layer consists of the shell protein, λ3, as well as the inner clamp protein σ2, which is thought to play a role in the structural stabilization of the icosahedron.<ref>{{cite journal | vauthors = Zhou ZH, Zhang H, Jakana J, Lu XY, Zhang JQ | title = Cytoplasmic polyhedrosis virus structure at 8 A by electron cryomicroscopy: structural basis of capsid stability and mRNA processing regulation | journal = Structure | volume = 11 | issue = 6 | pages = 651–63 | date = June 2003 | pmid = 12791254 | doi = 10.1016/s0969-2126(03)00091-1 }}</ref> The outer layer is thought to play a stabilizing role for the inner capsid and is made up of μ1(Outer Capsid protein) as well as σ3 (Outer Clamp Protein). Additionally, the outer layer has fibre proteins (σ1) that mediate viral attachment and entry into the host. |
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| ⚫ | PRV is a turreted reovirus, exhibiting a turret protein (λ2) on the five-fold axes of its inner capsid icosahedron.<ref name="Markussen_2013" /><ref>{{cite journal | vauthors = Miyazaki N, Uehara-Ichiki T, Xing L, Bergman L, Higashiura A, Nakagawa A, Omura T, Cheng RH | display-authors = 6 | title = Structural evolution of reoviridae revealed by oryzavirus in acquiring the second capsid shell | journal = Journal of Virology | volume = 82 | issue = 22 | pages = 11344–53 | date = November 2008 | pmid = 18787002 | pmc = 2573255 | doi = 10.1128/JVI.02375-07 }}</ref> The homologous proteins for L2 in MRV and ARV have both guanyltransferase and methyltransferase activity. Although not entirely conserved, the active regions of (λ2) exhibit fidelity to MRV and ARV sequences, suggesting λ2 is the turret protein and that it plays a similar role in the 5' capping of transcribed viral mRNA. |
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| ⚫ | The inner layer consists of the shell protein, λ3, as well as the inner clamp protein σ2, which is thought to play a role in the structural stabilization of the icosahedron<ref>{{ |
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| ⚫ | PRV is a turreted reovirus, exhibiting a turret protein (λ2) on the five-fold axes of its inner capsid icosahedron.<ref name=" |
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== Discovery == |
== Discovery == |
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PRV was first identified in 2010 via high throughput DNA sequencing and bioinformatics approaches which determined the virus was present in maricultured Atlantic salmon affected by Heart and Skeletal Muscle Inflammation (HSMI), and while control of the spread of PRV was advised, a causal relationship between PRV and HSMI was not demonstrated at that time.<ref name=" |
PRV was first identified in 2010 via high throughput DNA sequencing and bioinformatics approaches which determined the virus was present in maricultured Atlantic salmon affected by Heart and Skeletal Muscle Inflammation (HSMI), and while control of the spread of PRV was advised, a causal relationship between PRV and HSMI was not demonstrated at that time.<ref name="Palacios_2010" /> |
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In 2012, several wild fish species off the coast of Norway tested positive for PRV indicating that PRV is present in wild populations, however the majority of sample tests yielded negative results indicating low prevalence of PRV off the Norwegian coast.<ref>{{ |
In 2012, several wild fish species off the coast of Norway tested positive for PRV indicating that PRV is present in wild populations, however the majority of sample tests yielded negative results indicating low prevalence of PRV off the Norwegian coast.<ref>{{cite journal | vauthors = Wiik-Nielsen CR, Løvoll M, Sandlund N, Faller R, Wiik-Nielsen J, Bang Jensen B | title = First detection of piscine reovirus (PRV) in marine fish species | journal = Diseases of Aquatic Organisms | volume = 97 | issue = 3 | pages = 255–8 | date = January 2012 | pmid = 22422096 | doi = 10.3354/dao02425 }}</ref> |
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There has been speculation that PRV 1a was introduced to the West Coast of North America from a North Atlantic source. PRV 1a was introduced to Chile from North Pacific and North Atlantic sources<ref name=" |
There has been speculation that PRV 1a was introduced to the West Coast of North America from a North Atlantic source. PRV 1a was introduced to Chile from North Pacific and North Atlantic sources<ref name="Polinski_2020" /> |
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== Distribution == |
== Distribution == |
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[[File:Cultus Lake cutthroat trout.jpg|thumb|330x330px|Cutthroat trout that tested positive for PRV in Cultus Lake, British Columbia, Canada.]] |
[[File:Cultus Lake cutthroat trout.jpg|thumb|330x330px|Cutthroat trout that tested positive for PRV in Cultus Lake, British Columbia, Canada.]] |
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PRV has been found to have a nearly worldwide distribution with various studies detecting the presence of PRV in fisheries in the Atlantic off the coast of the UK, Ireland, and Norway, as well as in both the Atlantic and Pacific Coasts off of North and South America. PRV has been detected in pacific farmed and wild fish species as far north as Alaska, and as far south as Chile.<ref name=" |
PRV has been found to have a nearly worldwide distribution with various studies detecting the presence of PRV in fisheries in the Atlantic off the coast of the UK, Ireland, and Norway, as well as in both the Atlantic and Pacific Coasts off of North and South America. PRV has been detected in pacific farmed and wild fish species as far north as Alaska, and as far south as Chile.<ref name="Kibenge_2013" /><ref name="Polinski_2020" /><ref>{{cite journal | vauthors = Siah A, Morrison DB, Fringuelli E, Savage P, Richmond Z, Johns R, Purcell MK, Johnson SC, Saksida SM | display-authors = 6 | title = Piscine Reovirus: Genomic and Molecular Phylogenetic Analysis from Farmed and Wild Salmonids Collected on the Canada/US Pacific Coast | journal = PloS One | volume = 10 | issue = 11 | pages = e0141475 | date = 2015-11-04 | pmid = 26536673 | pmc = 4633109 | doi = 10.1371/journal.pone.0141475 }}</ref> |
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PRV-1 is less common in wild salmon than it is in farmed salmon, and while wild Sea Trout tend to exhibit PRV-3 infection, the same can not be said for wild Atlantic Salmon populations.<ref name=" |
PRV-1 is less common in wild salmon than it is in farmed salmon, and while wild Sea Trout tend to exhibit PRV-3 infection, the same can not be said for wild Atlantic Salmon populations.<ref name="Polinski_2020" /> |
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== Interaction with Host == |
== Interaction with Host == |
||
=== Infection Dynamics === |
=== Infection Dynamics === |
||
The kinetics of PRV infection have been identified into three distinct phases:<ref |
The kinetics of PRV infection have been identified into three distinct phases:<ref name="Polinski_2020" /><ref>{{cite journal | vauthors = Polinski MP, Marty GD, Snyman HN, Garver KA | title = Piscine orthoreovirus demonstrates high infectivity but low virulence in Atlantic salmon of Pacific Canada | journal = Scientific Reports | volume = 9 | issue = 1 | pages = 3297 | date = March 2019 | pmid = 30867461 | doi = 10.1038/s41598-019-40025-7 }}</ref> |
||
# early entry and systemic dissemination |
# early entry and systemic dissemination |
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| Line 100: | Line 98: | ||
=== Disease Prevention === |
=== Disease Prevention === |
||
HSMI is one of the most significant diseases in Atlantic salmon aquaculture in Norway. In efforts to mitigate this disease, two studies have been done, using vaccines that target PRV-1. One study uses a formalin-killed virus preparation strategy, and the other study uses DNA that expresses non-structural proteins. While both vaccines seem to have shown moderate protection against HSMI, neither of them could prevent the infection of PRV-1. There has also been attempts to create a strain of Atlantic Salmon that was ‘HSMI-resistant’ and while these fish seem to be somewhat resistant to HSMI, they are not resistant against PRV. Another form of treatment was to use specific feed formulations that would reduce the effects of HSMI and while this was successful, the PRV infection still remained.<ref name=" |
HSMI is one of the most significant diseases in Atlantic salmon aquaculture in Norway. In efforts to mitigate this disease, two studies have been done, using vaccines that target PRV-1. One study uses a formalin-killed virus preparation strategy, and the other study uses DNA that expresses non-structural proteins. While both vaccines seem to have shown moderate protection against HSMI, neither of them could prevent the infection of PRV-1. There has also been attempts to create a strain of Atlantic Salmon that was ‘HSMI-resistant’ and while these fish seem to be somewhat resistant to HSMI, they are not resistant against PRV. Another form of treatment was to use specific feed formulations that would reduce the effects of HSMI and while this was successful, the PRV infection still remained.<ref name="Polinski_2020" /> |
||
Currently, there are no strategies that seem to be able to eliminate PRV once it has been introduced to a population of naïve fish. The best form of prevention so far has been egg-surface disinfection in farmed salmon to limit the spread of PRV and while it is possible to remove PRV from a confined aquaculture system with intense molecular testing and disinfection, the salmon may become infected with PRV once exposed to saltwater environments.<ref name=" |
Currently, there are no strategies that seem to be able to eliminate PRV once it has been introduced to a population of naïve fish. The best form of prevention so far has been egg-surface disinfection in farmed salmon to limit the spread of PRV and while it is possible to remove PRV from a confined aquaculture system with intense molecular testing and disinfection, the salmon may become infected with PRV once exposed to saltwater environments.<ref name="Polinski_2020" /> |
||
== Prevalence in Aquaculture == |
== Prevalence in Aquaculture == |
||
PRV has been found to be most prevalent in Europe, and the Americas with relatively few reports in Asia. The prevalence of PRV varies greatly depending on the region and host species. PRV surveillance occurs most significantly in North America and Norway<ref name=" |
PRV has been found to be most prevalent in Europe, and the Americas with relatively few reports in Asia. The prevalence of PRV varies greatly depending on the region and host species. PRV surveillance occurs most significantly in North America and Norway<ref name="Polinski_2020" /> |
||
'''Europe:''' |
'''Europe:''' |
||
PRV-1 has been present in Norwegian farmed Atlantic Salmon since roughly 1988; within the rest of Europe, PRV is suspected to be common, but reports in some countries remain limited. Iceland has reported finding high PRV-1 prevalence in its Atlantic salmon. In 2018, PRV-3 was reported at Brown Trout and Rainbow trout farms in Denmark and Germany.<ref name=" |
PRV-1 has been present in Norwegian farmed Atlantic Salmon since roughly 1988; within the rest of Europe, PRV is suspected to be common, but reports in some countries remain limited. Iceland has reported finding high PRV-1 prevalence in its Atlantic salmon. In 2018, PRV-3 was reported at Brown Trout and Rainbow trout farms in Denmark and Germany.<ref name="Polinski_2020" /> |
||
'''North America:''' |
'''North America:''' |
||
PRV seems to exhibit high infectivity in North America, with a single infected fish often resulting in infection of a farms entire fish stock. PRV-1 tends to be initially detected following the return of farmed salmon from seawater. It’s likely that PRV-1 has been present in North America for decades, or possibly longer.<ref name=" |
PRV seems to exhibit high infectivity in North America, with a single infected fish often resulting in infection of a farms entire fish stock. PRV-1 tends to be initially detected following the return of farmed salmon from seawater. It’s likely that PRV-1 has been present in North America for decades, or possibly longer.<ref name="Polinski_2020" /> |
||
'''South America:''' |
'''South America:''' |
||
In Chile, farmed Atlantic salmon show high rates of PRV-1 infection, while wild fish populations show low rates of infection. Wild Coho Salmon do however show high rates of PRV-3 infection. Rainbow trout show high rates of infection with PRV-1.<ref name=" |
In Chile, farmed Atlantic salmon show high rates of PRV-1 infection, while wild fish populations show low rates of infection. Wild Coho Salmon do however show high rates of PRV-3 infection. Rainbow trout show high rates of infection with PRV-1.<ref name="Polinski_2020" /> |
||
'''Asia:''' |
'''Asia:''' |
||
Within Asia, PRV-2 has only been found in Japan and it has been linked to Erythrocytic Inclusion Body Syndrome (EIBS) in farmed Coho Salmon.<ref name=" |
Within Asia, PRV-2 has only been found in Japan and it has been linked to Erythrocytic Inclusion Body Syndrome (EIBS) in farmed Coho Salmon.<ref name="Polinski_2020" /> |
||
==== Host range: ==== |
==== Host range: ==== |
||
Revision as of 06:37, 10 April 2021
| Piscine orthoreovirus | |
|---|---|
| |
| Analysis of PRV-1 using Transmission Electron Microscopy (Scale bar 100nm). | |
Virus classification 
| |
| (unranked): | Virus |
| Realm: | Riboviria |
| Kingdom: | Orthornavirae |
| Phylum: | Duplornaviricota |
| Class: | Resentoviricetes |
| Order: | Reovirales |
| Family: | Sedoreoviridae |
| Subfamily: | Spinareovirinae |
| Genus: | Orthoreovirus |
| Species | |
|
Piscine orthoreovirus | |
Piscine orthoreovirus (PRV) is an Orthoreovirus species that infects fish species exclusively , and has been found present at higher concentration in fish with various diseases.[1] These diseases include heart and skeletal muscle inflammation (HSMI), jaundice syndrome, proliferative darkening syndrome and erythrocytic body inclusion syndrome.[1][2][3][4] PRV is thought to mainly affect aquacultured and maricultured fish stocks, and recent research has been focused around the susceptibility of wild stock. However, whether PRV is virulent, specifically with respect to HSMI, it has not been unanimously agreed upon.[5] PRV has been in the public eye mostly due to a potential linkage to farmed Atlantic Salmon exhibiting HSMI.[1] Public concern has been raised regarding the possibility of open ocean net farms transmitting PRV to wild populations being a factor in their recent decline.[6] PRV is not confirmed to be pathogenic in wild salmon stocks.
Classification
Phylogenetic analysis of each segment of the PRV genome initially placed PRV in the Reoviridae family, subfamily Spinareovirinae, based on sequence and structural similarity to known Reoviruses.[1] Phylogenetic sequence analysis indicated PRV was equally related to the genuses Orthoreoviridae and Aquareoviridae upon initial discovery.[1] As a result PRV was first believed to have evolved separately from a common ancestor related to both Orthoreoviridae and Aquareoviridae in the subfamily Spinareovirinae.
Currently, PRV is officially classified as an Orthoreovirus.[7] Placement into the genus Orthoreoviridae and not Aquareoviridae has been argued for the following reasons: Higher sequence fidelity within homologous sequences[8], the same segment numbers, lack of a fusogenic-associated small transmembrane protein (FAST)[9], presence of a fiber viral attachment protein[10], lack of syncytia formation and GC nucleoside percentage. [10] Opposing arguments to this placement include fact that the only other non-fusogenic orthoreovirus species is mammalian Orthoreovirus (MRV), and that while aquareoviridae have many known fish viruses orthoreoviridae doesn't have any other known.[11][12] Other arguments include the fact that the S1 and S4 sequences in PRV have no known homologues to either genus,[1] and that almost all orthoreoviridae and aquareoviridae are bicistronic for their viral attachment protein, while PRV is monocistronic.[13][10] PRV exhibits 5' terminal sequences on its segments that don't align to either genus.[1] There have been a few species of aquareoviridae identified as non-fusogenic, including GCRV104 and GCRVGD108.[14] The outer clamp protein of PRV is found on the bicistronic segment S1, the first of any known orthoreovirus or aquareovirus to encode this protein on a polycistronic segment.[9]
Genome and Structure
Genome
Piscine orthoreovirus has a segmented dsRNA genome made up of 10 individual linear segments, cumulatively measuring around 23,600bp.[1][10] It has a GC content of 47%. Each segment has conserved terminal sequences. The 3' end sequence (UCAUC-3') is the same as Orthoreoviridae and Aquareoviridae. The 5' end sequence (5'-GAUAAA/U) shows complete uniqueness within PRV. [1][11] These segments are referred to as L1-3 (Long), M1-3(Medium) and S1-4 (Short) based on length and comparison with homologous segments in orthoreoviridae and aquareoviridae. L1 is the longest at 3916bp and S4 the shortest at 1040bp.[1][10][8]
11 proteins are confirmed to be encoded.[10] S1 exhibits bicistronicity with 2 overlapping open reading frames, while the remaining proteins are either confirmed as monocistronic or have been thought to be bicistronic with no further evidence. [8][9][10] The proteins that each segment encodes for are as follows, using a standardized naming system across reovirus genera:[10][13]
- L1-λ3 - Shell Protein (Inner capsid protein)
- L2-λ2 - Turret protein [10]
- L3-λ1 - RdRP
- M1-μ2 - NTPase
- M2-μ1 - Outer capsid protein
- M3-μNS - unknown function, [15]
- S1-σ3 - Outer Clamp Protein (Capsid protein)
- p13 - Cytotoxic Nonstructural Protein[16]
- S2-σ2 - Core Clamp Protein
- S3-σNS - unknown function
- S4-σ1 - Viral Attachment Protein[10]
Subdivisions
Piscine reovirus has been grouped into multiple different genotypes based on sequence diversity. Although multiple ways of subdividing PRV have been proposed,[3] the system most often used in the literature subdivides it into 2 genotypes, I and II, which further divide into Ia and Ib, and IIa and IIb, respectively.[17][2][18][19] These divisions and subdivisions are based on sequence diversity within segment S1.[18][8]
Genotype I (PRV-1) [8]
Ia
This sub genotype of PRV is found primarily in farmed Atlantic salmon in Norway, Chile and Canada. It has been associated with populations exhibiting HSMI. It has been found in farmed Chinook salmon in Canada exhibiting Jaundice syndrome, as well as farmed Rainbow trout and Coho showing HSMI like symptoms in Chile and Canada.[20]
Ib
This sub genotype of PRV is found primarily in farmed Atlantic and Coho salmon in Norway and Chile, additionally being found in farmed Rainbow trout in Chile.[4] It has been present in many salmon with HSMI. It has also been found in Coho populations exhibiting Jaundice syndrome.
Genotype 2 [18]
Genotype 2 was first discovered when comparing the S1 sequences among farmed Atlantic salmon in Chile, and further reiterated with both M2 and whole genome analysis.[21][17] Despite being within one genotype, the sub genotypes IIa and IIb exhibit much higher inter-sequence diversity than do Ia and Ib.
IIa (PRV-3)
PRV-3 has been associated with pathological heart lesions in rainbow trout.[22][17] It has also been found in farmed Coho Salmon, Brown trout, and Rainbow trout exhibiting HSMI across Northern Europe and Chile.[19][19] It has also been found in wild brown trout with proliferative darkening syndrome in Central Europe.[16] Comparison with PRV-2 shows an 80.1% and 90.5% similarity for nucleotides and amino acids, respectively.[23] Comparison with PRV-1 showed a 72.9% similarity in nucleotide sequence and 80.0% amino acid sequence similarity.
IIb (PRV-2)
PVR-2 has only been found in farmed Coho salmon in Japan exhibiting Erythrocytic Inclusion Body Syndrome (EIBS).[2] Only one genome of PRV-2 has been sequenced thus far, which has reduced confidence in phylogenetic placements.[9] Comparison with PRV-1 showed a 73.4% and 80.3% similarity in nucleotide sequence and amino acid sequence, respectively.[23]
Structure
As a member of genus orthoreovirus, the virion form of PRV is a non-enveloped icosahedral capsid with two layers, the outer and core. The diameter of the total virion encompassing the outer layer is approximately 70nm, with the inner core layer measuring around 39nm in diameter.[24]
The inner layer consists of the shell protein, λ3, as well as the inner clamp protein σ2, which is thought to play a role in the structural stabilization of the icosahedron.[25] The outer layer is thought to play a stabilizing role for the inner capsid and is made up of μ1(Outer Capsid protein) as well as σ3 (Outer Clamp Protein). Additionally, the outer layer has fibre proteins (σ1) that mediate viral attachment and entry into the host.
PRV is a turreted reovirus, exhibiting a turret protein (λ2) on the five-fold axes of its inner capsid icosahedron.[10][26] The homologous proteins for L2 in MRV and ARV have both guanyltransferase and methyltransferase activity. Although not entirely conserved, the active regions of (λ2) exhibit fidelity to MRV and ARV sequences, suggesting λ2 is the turret protein and that it plays a similar role in the 5' capping of transcribed viral mRNA.
Discovery
PRV was first identified in 2010 via high throughput DNA sequencing and bioinformatics approaches which determined the virus was present in maricultured Atlantic salmon affected by Heart and Skeletal Muscle Inflammation (HSMI), and while control of the spread of PRV was advised, a causal relationship between PRV and HSMI was not demonstrated at that time.[1]
In 2012, several wild fish species off the coast of Norway tested positive for PRV indicating that PRV is present in wild populations, however the majority of sample tests yielded negative results indicating low prevalence of PRV off the Norwegian coast.[27]
There has been speculation that PRV 1a was introduced to the West Coast of North America from a North Atlantic source. PRV 1a was introduced to Chile from North Pacific and North Atlantic sources[21]
Distribution
PRV has been found to have a nearly worldwide distribution with various studies detecting the presence of PRV in fisheries in the Atlantic off the coast of the UK, Ireland, and Norway, as well as in both the Atlantic and Pacific Coasts off of North and South America. PRV has been detected in pacific farmed and wild fish species as far north as Alaska, and as far south as Chile.[8][21][28]
PRV-1 is less common in wild salmon than it is in farmed salmon, and while wild Sea Trout tend to exhibit PRV-3 infection, the same can not be said for wild Atlantic Salmon populations.[21]
Interaction with Host
Infection Dynamics
The kinetics of PRV infection have been identified into three distinct phases:[21][29]
- early entry and systemic dissemination
- peak systemic replication and
- long-term, high-load viral persistence
Transmission
Disease
PRV and HSMI
While Norwegian strains of PRV have led to HSMI outbreaks, it's uncertain if HSMI develops as a result of infection with North American strains of PRV as various salmon and trout have been found to have detectable PRV infections despite an apparent absence of any HSMI in associated waters.[30]
Research in Norway has indicated that exposing fish to injected PRV from diseased fish will lead to HSMI, but this causal relationship seems to be more difficult to demonstrate with PRV variants found off the west coast of Canada.[30]
Disease Prevention
HSMI is one of the most significant diseases in Atlantic salmon aquaculture in Norway. In efforts to mitigate this disease, two studies have been done, using vaccines that target PRV-1. One study uses a formalin-killed virus preparation strategy, and the other study uses DNA that expresses non-structural proteins. While both vaccines seem to have shown moderate protection against HSMI, neither of them could prevent the infection of PRV-1. There has also been attempts to create a strain of Atlantic Salmon that was ‘HSMI-resistant’ and while these fish seem to be somewhat resistant to HSMI, they are not resistant against PRV. Another form of treatment was to use specific feed formulations that would reduce the effects of HSMI and while this was successful, the PRV infection still remained.[21]
Currently, there are no strategies that seem to be able to eliminate PRV once it has been introduced to a population of naïve fish. The best form of prevention so far has been egg-surface disinfection in farmed salmon to limit the spread of PRV and while it is possible to remove PRV from a confined aquaculture system with intense molecular testing and disinfection, the salmon may become infected with PRV once exposed to saltwater environments.[21]
Prevalence in Aquaculture
PRV has been found to be most prevalent in Europe, and the Americas with relatively few reports in Asia. The prevalence of PRV varies greatly depending on the region and host species. PRV surveillance occurs most significantly in North America and Norway[21]
Europe:
PRV-1 has been present in Norwegian farmed Atlantic Salmon since roughly 1988; within the rest of Europe, PRV is suspected to be common, but reports in some countries remain limited. Iceland has reported finding high PRV-1 prevalence in its Atlantic salmon. In 2018, PRV-3 was reported at Brown Trout and Rainbow trout farms in Denmark and Germany.[21]
North America:
PRV seems to exhibit high infectivity in North America, with a single infected fish often resulting in infection of a farms entire fish stock. PRV-1 tends to be initially detected following the return of farmed salmon from seawater. It’s likely that PRV-1 has been present in North America for decades, or possibly longer.[21]
South America:
In Chile, farmed Atlantic salmon show high rates of PRV-1 infection, while wild fish populations show low rates of infection. Wild Coho Salmon do however show high rates of PRV-3 infection. Rainbow trout show high rates of infection with PRV-1.[21]
Asia:
Within Asia, PRV-2 has only been found in Japan and it has been linked to Erythrocytic Inclusion Body Syndrome (EIBS) in farmed Coho Salmon.[21]
Host range:
PRV has been found in a range of aquacultured/wild host species such as:
- Cutthroat Trout (Oncorhynchus clarkii)[30]
- Chinook Salmon (Oncorhynchus tshawytscha)[30]
- Sockeye Salmon (Oncorhynchus nerka)[30]
- Steelhead Trout (Oncorhynchus mykiss)[30]
- Coho Salmon (Oncorhynchus kisutch)[30]
- Chum Salmon (Oncorhynchus keta)[30]
- Pink Salmon (Oncorhynchus gorbusca)[30]
- Atlantic Salmon (Salmo salar)[30]
- Sea-Trout (Salmo trutta)[30]
- Great Silver Smelt (Argentina silus)[30]
- Atlantic Horse Mackerel (Trachurus trachurus)[30]
- Atlantic Herring (Clupea harengus)[30]
- Capelin (Mallotus villosus)[30]
References
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{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b c Takano T, Nawata A, Sakai T, Matsuyama T, Ito T, Kurita J, et al. (2016-10-27). "Full-Genome Sequencing and Confirmation of the Causative Agent of Erythrocytic Inclusion Body Syndrome in Coho Salmon Identifies a New Type of Piscine Orthoreovirus". PloS One. 11 (10): e0165424. doi:10.1371/journal.pone.0165424. PMC 5082797. PMID 27788206.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b Garseth ÅH, Ekrem T, Biering E (2013-12-11). "Phylogenetic evidence of long distance dispersal and transmission of piscine reovirus (PRV) between farmed and wild Atlantic salmon". PloS One. 8 (12): e82202. doi:10.1371/journal.pone.0082202. PMID 24349221.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b Cartagena J, Jiménez C, Spencer E (November 2020). "Detection of Piscine orthoreoviruses (PRV-1b AND PRV-3a) in farmed Coho salmon with jaundice syndrome from Chile". Aquaculture. 528: 735480. doi:10.1016/j.aquaculture.2020.735480. ISSN 0044-8486.
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{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Haatveit HM, Nyman IB, Markussen T, Wessel Ø, Dahle MK, Rimstad E (January 2016). "The non-structural protein μNS of piscine orthoreovirus (PRV) forms viral factory-like structures". Veterinary Research. 47 (1): 5. doi:10.1186/s13567-015-0302-0. PMC 4705589. PMID 26743679.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b Kuehn R, Stoeckle BC, Young M, Popp L, Taeubert JE, Pfaffl MW, Geist J (2018-10-22). "Identification of a piscine reovirus-related pathogen in proliferative darkening syndrome (PDS) infected brown trout (Salmo trutta fario) using a next-generation technology detection pipeline". PloS One. 13 (10): e0206164. doi:10.1371/journal.pone.0206164. PMC 6197672. PMID 30346982.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b c Godoy M, Medina DA, Suarez R, Valenzuela S, Romero J, Kibenge M, et al. (January 2021). "Extensive Phylogenetic Analysis of Piscine Orthoreovirus Genomic Sequences Shows the Robustness of Subgenotype Classification". Pathogens. 10 (1): 41. doi:10.3390/pathogens10010041. PMID 33430212.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b c Godoy MG, Kibenge MJ, Wang Y, Suarez R, Leiva C, Vallejos F, Kibenge FS (June 2016). "First description of clinical presentation of piscine orthoreovirus (PRV) infections in salmonid aquaculture in Chile and identification of a second genotype (Genotype II) of PRV". Virology Journal. 13 (1): 98. doi:10.1186/s12985-016-0554-y. PMC 4906990. PMID 27296722.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b c Kibenge FS (February 2019). "Emerging viruses in aquaculture". Current Opinion in Virology. Emerging viruses: intraspecies transmission • Viral Immunology. 34: 97–103. doi:10.1016/j.coviro.2018.12.008. PMID 30711892.
- ^ Di Cicco E, Ferguson HW, Kaukinen KH, Schulze AD, Li S, Tabata A, et al. (2018-10-01). "The same strain of Piscine orthoreovirus (PRV-1) is involved in the development of different, but related, diseases in Atlantic and Pacific Salmon in British Columbia". FACETS. 3 (1): 599–641. doi:10.1139/facets-2018-0008. ISSN 2371-1671.
- ^ a b c d e f g h i j k l Polinski MP, Vendramin N, Cuenca A, Garver KA (November 2020). "Piscine orthoreovirus: Biology and distribution in farmed and wild fish". Journal of Fish Diseases. 43 (11): 1331–1352. doi:10.1111/jfd.13228. PMID 32935367.
- ^ Vendramin N, Kannimuthu D, Olsen AB, Cuenca A, Teige LH, Wessel Ø, et al. (February 2019). "Piscine orthoreovirus subtype 3 (PRV-3) causes heart inflammation in rainbow trout (Oncorhynchus mykiss)". Veterinary Research. 50 (1): 14. doi:10.1186/s13567-019-0632-4. PMID 30777130.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ a b Dhamotharan K, Vendramin N, Markussen T, Wessel Ø, Cuenca A, Nyman IB, et al. (April 2018). "Molecular and Antigenic Characterization of Piscine orthoreovirus (PRV) from Rainbow Trout (Oncorhynchus mykiss)". Viruses. 10 (4). doi:10.3390/v10040170. PMC 5923464. PMID 29614838.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Wessel Ø, Braaen S, Alarcon M, Haatveit H, Roos N, Markussen T, et al. (2017-08-25). "Infection with purified Piscine orthoreovirus demonstrates a causal relationship with heart and skeletal muscle inflammation in Atlantic salmon". PloS One. 12 (8): e0183781. doi:10.1371/journal.pone.0183781. PMID 28841684.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Zhou ZH, Zhang H, Jakana J, Lu XY, Zhang JQ (June 2003). "Cytoplasmic polyhedrosis virus structure at 8 A by electron cryomicroscopy: structural basis of capsid stability and mRNA processing regulation". Structure. 11 (6): 651–63. doi:10.1016/s0969-2126(03)00091-1. PMID 12791254.
- ^ Miyazaki N, Uehara-Ichiki T, Xing L, Bergman L, Higashiura A, Nakagawa A, et al. (November 2008). "Structural evolution of reoviridae revealed by oryzavirus in acquiring the second capsid shell". Journal of Virology. 82 (22): 11344–53. doi:10.1128/JVI.02375-07. PMC 2573255. PMID 18787002.
- ^ Wiik-Nielsen CR, Løvoll M, Sandlund N, Faller R, Wiik-Nielsen J, Bang Jensen B (January 2012). "First detection of piscine reovirus (PRV) in marine fish species". Diseases of Aquatic Organisms. 97 (3): 255–8. doi:10.3354/dao02425. PMID 22422096.
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{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Polinski MP, Marty GD, Snyman HN, Garver KA (March 2019). "Piscine orthoreovirus demonstrates high infectivity but low virulence in Atlantic salmon of Pacific Canada". Scientific Reports. 9 (1): 3297. doi:10.1038/s41598-019-40025-7. PMID 30867461.
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