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This article is about the virus related to smallpox vaccines. For the plant genus, see Vaccinium.
Not to be confused with vaccine.

Vaccinia virus (VACV or VV) is a large, complex, enveloped virus belonging to the poxvirus family. It has a linear, double-stranded DNA genome approximately 190 kbp in length, which encodes approximately 250 genes. The dimensions of the virion are roughly 360 × 270 × 250 nm, with a mass of approximately 5–10 fg. The vaccinia virus is the source of the modern smallpox vaccine, which the World Health Organisation used to eradicate smallpox in a global vaccination campaign in 1958–1977. Although smallpox no longer exists in the wild, vaccinia virus is still studied widely by scientists as a tool for gene therapy and genetic engineering.

Vaccinia
SpecialtyVirology
TypesProgressive vaccinia

Smallpox had been an endemic human disease that had a 30% fatality rate. In 1796, the British doctor Edward Jenner proved that an infection with the relatively mild cowpox virus would also confer immunity to the deadly smallpox. Jenner referred to cowpox as variolae vaccinae (smallpox of the cow). However, the origins of the smallpox vaccine became murky over time, especially after Louis Pasteur developed laboratory techniques for creating vaccines in the 19th century. Allan Watt Downie demonstrated in 1939 that the modern smallpox vaccine was serologically distinct from cowpox, and vaccinia was subsequently recognized as a separate viral species. Whole-genome sequencing has revealed that vaccinia is most closely related to horsepox, and the cowpox strains found in Great Britain are the least closely related to vaccinia.

Contents

In addition to the morbidity of uncomplicated primary vaccination, transfer of infection to other sites by scratching, and post-vaccinial encephalitis, other complications of vaccinia infections may be divided into the following types:: 391

Vaccinia virus is closely related to the virus that causes cowpox; historically the two were often considered to be one and the same. The precise origin of vaccinia virus is unknown due to the lack of record-keeping, as the virus was repeatedly cultivated and passaged in research laboratories for many decades. The most common notion is that vaccinia virus, cowpox virus, and variola virus (the causative agent of smallpox) were all derived from a common ancestral virus. There is also speculation that vaccinia virus was originally isolated from horses, and analysis of DNA from an early (1902) sample of smallpox vaccine showed that it was 99.7% similar to horsepox virus.

Poxviruses are unique among DNA viruses because they replicate only in the cytoplasm of the host cell, outside of the nucleus. Therefore, the large genome is required for encoding various enzymes and proteins involved in viral DNA replication and gene transcription. During its replication cycle, VV produces four infectious forms which differ in their outer membranes: intracellular mature virion (IMV), the intracellular enveloped virion (IEV), the cell-associated enveloped virion (CEV) and the extracellular enveloped virion (EEV). Although the issue remains contentious, the prevailing view is that the IMV consists of a single lipoprotein membrane, while the CEV and EEV are both surrounded by two membrane layers and the IEV has three envelopes. The IMV is the most abundant infectious form and is thought to be responsible for spread between hosts. On the other hand, the CEV is believed to play a role in cell-to-cell spread and the EEV is thought to be important for long range dissemination within the host organism.[citation needed]

Vaccinia virus is able to undergo multiplicity reactivation (MR). MR is the process by which two, or more, virus genomes containing otherwise lethal damage interact within an infected cell to form a viable virus genome. Abel found that vaccinia viruses exposed to doses of UV light sufficient to prevent progeny formation when single virus particles infected host chick embryo cells, could still produce viable progeny viruses when host cells were infected by two or more of these inactivated viruses; that is, MR could occur. Kim and Sharp demonstrated MR of vaccinia virus after treatment with UV-light, nitrogen mustard, and X-rays or gamma rays. Michod et al. reviewed numerous examples of MR in different viruses, and suggested that MR is a common form of sexual interaction in viruses that provides the advantage of recombinational repair of genome damages.[additional citation(s) needed]

Vaccinia contains within its genome genes for several proteins that give the virus resistance to interferons:

  • K3L (P18378) is a protein with homology to the protein eukaryotic initiation factor 2 (eIF-2alpha). K3L protein inhibits the action of PKR, an activator of interferons.
  • E3L (P21605) is another protein encoded by Vaccinia. E3L also inhibits PKR activation; and is also able to bind to double stranded RNA.
  • B18R is a protein which serves as an interferon inhibitor in one of Moderna's technologies.
Site of a vaccinia injection, several days later.

Vaccinia virus infection is typically very mild and often does not cause symptoms in healthy individuals, although it may cause rash and fever. Immune responses generated from a vaccinia virus infection protects the person against a lethal smallpox infection. For this reason, vaccinia virus was, and still is, being used as a live-virus vaccine against smallpox. Unlike vaccines that use weakened forms of the virus being vaccinated against, the vaccinia virus vaccine cannot cause a smallpox infection because it does not contain the smallpox virus. However, certain complications and/or vaccine adverse effects occasionally arise. The chance of this happening is significantly increased in people who are immunocompromised. Approximately 1 to 2 people out of every 1 million people vaccinated could die as a result of life-threatening reactions to the vaccination.

Currently[when?], the vaccine is only administered to health care workers or research personnel who have a high risk of contracting the variola virus, and to the military personnel of the United States. Due to the threat of smallpox bioterrorism, there is a possibility the vaccine may have to be widely administered again in the future. Therefore, scientists are currently[when?] developing novel vaccine strategies against smallpox which are safer and much faster to deploy during a bioterrorism event.

On September 1, 2007, the U.S. Food and Drug Administration (FDA) licensed a new vaccine ACAM2000 against smallpox which can be produced quickly upon need. Manufactured by Sanofi Pasteur, the U.S. Centers for Disease Control and Prevention stockpiled 192.5 million doses of the new vaccine (see list of common strains below).

A new smallpox vaccine, Imvanex, which is based on the Modified vaccinia Ankara strain, was approved by the European Medicines Agency in 2013.

Vaccinia is also used in recombinant vaccines, as a vector for expression of foreign genes within a host, in order to generate an immune response. Other poxviruses are also used as live recombinant vaccines.

The original vaccine for smallpox, and the origin of the idea of vaccination, was Cowpox, described by Edward Jenner in 1798. The Latin term used for Cowpox was Variolae vaccinae, Jenner's own translation of "smallpox of the cow". That term lent its name to the whole idea of vaccination. When it was realized that the virus used in smallpox vaccination was not, or was no longer, the same as cowpox virus, the name 'vaccinia' was used for the virus in smallpox vaccine. (See OED.) Vaccine potency and efficacy prior to the invention of refrigerated methods of transportation was unreliable. The vaccine would be rendered impotent by heat and sunlight, and the method of drying samples on quills and shipping them to countries in need often resulted in an inactive vaccine. Another method employed was the "arm to arm" method. This involved vaccinating an individual then transferring it to another as soon as the infectious pustule forms, then to another, etc. This method was used as a form of living transportation of the vaccine, and usually employed orphans as carriers. However, this method was problematic due to the possibility of spreading other blood diseases, such as hepatitis and syphilis, as was the case in 1861, when 41 Italian children contracted syphilis after being vaccinated by the "arm to arm" method. Henry Austin Martin introduced a method for vaccine production from calves.

In 1913, E. Steinhardt, C. Israeli, and R. A. Lambert grew vaccinia virus in fragments of pig corneal tissue culture.

A paper published in 1915 by Fredrick W. Twort, a student of Willian Bulloch, is considered to be the beginning of modern phage research. He was attempting to grow vaccinia virus on agar media in the absence of living cells when he noted that many colonies of contaminating micrococci grew up and appeared mucoid, watery or glassy, and this transformation could be induced in other colonies by inoculation of the fresh colony with material from the watery colony. Using a microscope, he observed that bacteria had degenerated into small granules that stained red with Giemsa stain. He concluded that "...it [the agent of transformation] might almost be considered as an acute infectious disease of micrococci."

In 1939 Allan Watt Downie showed that the smallpox vaccines being used in the 20th century and cowpox virus were not the same, but were immunologically related.

In March 2007, a 2-year-old Indiana boy and his mother contracted a life-threatening vaccinia infection from the boy's father. The boy developed the telltale rash over 80 percent of his body after coming into close contact with his father, who was vaccinated for smallpox before being deployed overseas by the United States Army. The United States military resumed smallpox vaccinations in 2002. The child acquired the infection due to eczema, which is a known risk factor for vaccinia infection. The boy was treated with intravenous immunoglobulin, cidofovir, and Tecovirimat (ST-246), a (then) experimental drug developed by SIGA Technologies. On April 19, 2007, he was sent home with no after effects except for possible scarring of the skin.

In 2010, the Centers for Disease Control and Prevention (CDC) reported that a woman in Washington had contracted vaccinia virus infection after digital vaginal contact with her boyfriend, a military member who had recently been vaccinated for smallpox. The woman had a history of childhood eczema, but she had not been symptomatic as an adult. The CDC indicated that it was aware of four similar cases in the preceding 12 months of vaccinia infection after sexual contact with a recent military vaccinee. Further cases—also in patients with a history of eczema—occurred in 2012.

This is a list of some of the well-characterized vaccinia strains used for research and vaccination.[citation needed]

  • Lister (also known as Elstree): the English vaccine strain used by Leslie Collier to develop heat stable vaccine in powdered form. Used as the basis for vaccine production during the World Health Organization Smallpox Eradication Campaign (SEC)
  • Dryvax (also known as "Wyeth"): the vaccine strain previously used in the United States, produced by Wyeth. Used in the SEC, it was replaced in 2008 by ACAM2000 (see below), produced by Acambis. It was produced as preparations of calf lymph which was freeze-dried and treated with antibiotics.
  • EM63; Russian strain used in the SEC
  • ACAM2000: The current strain in use in the US, produced by Acambis. ACAM2000 was derived from a clone of a Dryvax virus by plaque purification. It is produced in cultures of Vero cells.
  • Modified vaccinia Ankara (also known as MVA): a highly attenuated (not virulent) strain created by passaging vaccinia virus several hundred times in chicken embryo fibroblasts. Unlike some other vaccinia strains it does not make immunodeficient mice sick and therefore may be safer to use in humans who have weaker immune systems due to being very young, very old, having HIV/AIDS, etc.
  • LC16m8: an attenuated strain developed and currently used in Japan
  • CV-1: an attenuated strain developed in the United States and used there in the late 1960s- 1970s
  • Western Reserve
  • Copenhagen
  • Connaught Laboratories (Canada)
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Vaccinia Article Talk Language Watch Edit This article is about the virus related to smallpox vaccines For the plant genus see Vaccinium Not to be confused with vaccine Vaccinia virus VACV or VV is a large complex enveloped virus belonging to the poxvirus family 2 It has a linear double stranded DNA genome approximately 190 kbp in length which encodes approximately 250 genes The dimensions of the virion are roughly 360 270 250 nm with a mass of approximately 5 10 fg 3 The vaccinia virus is the source of the modern smallpox vaccine which the World Health Organisation used to eradicate smallpox in a global vaccination campaign in 1958 1977 Although smallpox no longer exists in the wild vaccinia virus is still studied widely by scientists as a tool for gene therapy and genetic engineering Vaccinia virusA TEM micrograph of Vaccinia virus virionsVirus classification unranked VirusRealm VaridnaviriaKingdom BamfordviraePhylum NucleocytoviricotaClass PokkesviricetesOrder ChitoviralesFamily PoxviridaeGenus OrthopoxvirusSpecies Vaccinia virusMember viruses 1 Buffalopox virus Cantagalo virus Rabbitpox virus Utrecht Vaccinia virus Ankara Vaccinia virus Copenhagen Vaccinia virus WRVacciniaSpecialtyVirologyTypesProgressive vaccinia Smallpox had been an endemic human disease that had a 30 fatality rate In 1796 the British doctor Edward Jenner proved that an infection with the relatively mild cowpox virus would also confer immunity to the deadly smallpox Jenner referred to cowpox as variolae vaccinae smallpox of the cow However the origins of the smallpox vaccine became murky over time 4 especially after Louis Pasteur developed laboratory techniques for creating vaccines in the 19th century Allan Watt Downie demonstrated in 1939 that the modern smallpox vaccine was serologically distinct from cowpox 5 and vaccinia was subsequently recognized as a separate viral species Whole genome sequencing has revealed that vaccinia is most closely related to horsepox and the cowpox strains found in Great Britain are the least closely related to vaccinia 6 Contents 1 Classification of vaccinia infections 2 Origin 3 Virology 4 Multiplicity reactivation 5 Host resistance 6 Use as a vaccine 7 History 8 Recent cases 9 Common strains 10 References 11 Further reading 12 External linksClassification of vaccinia infections EditIn addition to the morbidity of uncomplicated primary vaccination transfer of infection to other sites by scratching and post vaccinial encephalitis other complications of vaccinia infections may be divided into the following types 7 391 Generalized vaccinia Eczema vaccinatum Progressive vaccinia vaccinia gangrenosum vaccinia necrosum Roseola vacciniaOrigin EditVaccinia virus is closely related to the virus that causes cowpox historically the two were often considered to be one and the same 8 The precise origin of vaccinia virus is unknown due to the lack of record keeping as the virus was repeatedly cultivated and passaged in research laboratories for many decades 9 The most common notion is that vaccinia virus cowpox virus and variola virus the causative agent of smallpox were all derived from a common ancestral virus There is also speculation that vaccinia virus was originally isolated from horses 8 and analysis of DNA from an early 1902 sample of smallpox vaccine showed that it was 99 7 similar to horsepox virus 10 Virology EditPoxviruses are unique among DNA viruses because they replicate only in the cytoplasm of the host cell outside of the nucleus 11 Therefore the large genome is required for encoding various enzymes and proteins involved in viral DNA replication and gene transcription During its replication cycle VV produces four infectious forms which differ in their outer membranes intracellular mature virion IMV the intracellular enveloped virion IEV the cell associated enveloped virion CEV and the extracellular enveloped virion EEV 12 Although the issue remains contentious the prevailing view is that the IMV consists of a single lipoprotein membrane while the CEV and EEV are both surrounded by two membrane layers and the IEV has three envelopes The IMV is the most abundant infectious form and is thought to be responsible for spread between hosts On the other hand the CEV is believed to play a role in cell to cell spread and the EEV is thought to be important for long range dissemination within the host organism citation needed Multiplicity reactivation EditVaccinia virus is able to undergo multiplicity reactivation MR 13 MR is the process by which two or more virus genomes containing otherwise lethal damage interact within an infected cell to form a viable virus genome Abel 13 found that vaccinia viruses exposed to doses of UV light sufficient to prevent progeny formation when single virus particles infected host chick embryo cells could still produce viable progeny viruses when host cells were infected by two or more of these inactivated viruses that is MR could occur Kim and Sharp demonstrated MR of vaccinia virus after treatment with UV light 14 nitrogen mustard 15 and X rays or gamma rays 16 Michod et al 17 reviewed numerous examples of MR in different viruses and suggested that MR is a common form of sexual interaction in viruses that provides the advantage of recombinational repair of genome damages additional citation s needed Host resistance EditVaccinia contains within its genome genes for several proteins that give the virus resistance to interferons K3L P18378 is a protein with homology to the protein eukaryotic initiation factor 2 eIF 2alpha K3L protein inhibits the action of PKR an activator of interferons 18 E3L P21605 is another protein encoded by Vaccinia E3L also inhibits PKR activation and is also able to bind to double stranded RNA 18 B18R is a protein which serves as an interferon inhibitor in one of Moderna s technologies 19 Use as a vaccine Edit Site of a vaccinia injection several days later Vaccinia virus infection is typically very mild and often does not cause symptoms in healthy individuals although it may cause rash and fever Immune responses generated from a vaccinia virus infection protects the person against a lethal smallpox infection For this reason vaccinia virus was and still is being used as a live virus vaccine against smallpox Unlike vaccines that use weakened forms of the virus being vaccinated against the vaccinia virus vaccine cannot cause a smallpox infection because it does not contain the smallpox virus However certain complications and or vaccine adverse effects occasionally arise The chance of this happening is significantly increased in people who are immunocompromised Approximately 1 to 2 people out of every 1 million people vaccinated could die as a result of life threatening reactions to the vaccination 20 Currently when the vaccine is only administered to health care workers or research personnel who have a high risk of contracting the variola virus and to the military personnel of the United States Due to the threat of smallpox bioterrorism there is a possibility the vaccine may have to be widely administered again in the future Therefore scientists are currently when developing novel vaccine strategies against smallpox which are safer and much faster to deploy during a bioterrorism event On September 1 2007 the U S Food and Drug Administration FDA licensed a new vaccine ACAM2000 against smallpox which can be produced quickly upon need Manufactured by Sanofi Pasteur the U S Centers for Disease Control and Prevention stockpiled 192 5 million doses of the new vaccine see list of common strains below 21 A new smallpox vaccine Imvanex which is based on the Modified vaccinia Ankara strain was approved by the European Medicines Agency in 2013 22 Vaccinia is also used in recombinant vaccines as a vector for expression of foreign genes within a host in order to generate an immune response Other poxviruses are also used as live recombinant vaccines 23 History EditThe original vaccine for smallpox and the origin of the idea of vaccination was Cowpox described by Edward Jenner in 1798 The Latin term used for Cowpox was Variolae vaccinae Jenner s own translation of smallpox of the cow That term lent its name to the whole idea of vaccination 24 When it was realized that the virus used in smallpox vaccination was not or was no longer the same as cowpox virus the name vaccinia was used for the virus in smallpox vaccine See OED Vaccine potency and efficacy prior to the invention of refrigerated methods of transportation was unreliable The vaccine would be rendered impotent by heat and sunlight and the method of drying samples on quills and shipping them to countries in need often resulted in an inactive vaccine Another method employed was the arm to arm method This involved vaccinating an individual then transferring it to another as soon as the infectious pustule forms then to another etc This method was used as a form of living transportation of the vaccine and usually employed orphans as carriers However this method was problematic due to the possibility of spreading other blood diseases such as hepatitis and syphilis as was the case in 1861 when 41 Italian children contracted syphilis after being vaccinated by the arm to arm method 25 Henry Austin Martin introduced a method for vaccine production from calves 26 In 1913 E Steinhardt C Israeli and R A Lambert grew vaccinia virus in fragments of pig corneal tissue culture 27 A paper published in 1915 by Fredrick W Twort a student of Willian Bulloch is considered to be the beginning of modern phage research He was attempting to grow vaccinia virus on agar media in the absence of living cells when he noted that many colonies of contaminating micrococci grew up and appeared mucoid watery or glassy and this transformation could be induced in other colonies by inoculation of the fresh colony with material from the watery colony Using a microscope he observed that bacteria had degenerated into small granules that stained red with Giemsa stain He concluded that it the agent of transformation might almost be considered as an acute infectious disease of micrococci 28 In 1939 Allan Watt Downie showed that the smallpox vaccines being used in the 20th century and cowpox virus were not the same but were immunologically related 5 29 Recent cases EditIn March 2007 a 2 year old Indiana boy and his mother contracted a life threatening vaccinia infection from the boy s father 30 The boy developed the telltale rash over 80 percent of his body after coming into close contact with his father who was vaccinated for smallpox before being deployed overseas by the United States Army The United States military resumed smallpox vaccinations in 2002 The child acquired the infection due to eczema which is a known risk factor for vaccinia infection The boy was treated with intravenous immunoglobulin cidofovir and Tecovirimat ST 246 a then experimental drug developed by SIGA Technologies 31 On April 19 2007 he was sent home with no after effects except for possible scarring of the skin 30 In 2010 the Centers for Disease Control and Prevention CDC reported that a woman in Washington had contracted vaccinia virus infection after digital vaginal contact with her boyfriend a military member who had recently been vaccinated for smallpox The woman had a history of childhood eczema but she had not been symptomatic as an adult The CDC indicated that it was aware of four similar cases in the preceding 12 months of vaccinia infection after sexual contact with a recent military vaccinee 32 Further cases also in patients with a history of eczema occurred in 2012 33 Common strains EditThis is a list of some of the well characterized vaccinia strains used for research and vaccination citation needed Lister also known as Elstree the English vaccine strain used by Leslie Collier to develop heat stable vaccine in powdered form Used as the basis for vaccine production during the World Health Organization Smallpox Eradication Campaign SEC Dryvax also known as Wyeth the vaccine strain previously used in the United States produced by Wyeth Used in the SEC it was replaced in 2008 34 by ACAM2000 see below produced by Acambis It was produced as preparations of calf lymph which was freeze dried and treated with antibiotics EM63 Russian strain used in the SEC ACAM2000 The current strain in use in the US produced by Acambis ACAM2000 was derived from a clone of a Dryvax virus by plaque purification It is produced in cultures of Vero cells Modified vaccinia Ankara also known as MVA a highly attenuated not virulent strain created by passaging vaccinia virus several hundred times in chicken embryo fibroblasts Unlike some other vaccinia strains it does not make immunodeficient mice sick and therefore may be safer to use in humans who have weaker immune systems due to being very young very old having HIV AIDS etc LC16m8 an attenuated strain developed and currently used in Japan CV 1 an attenuated strain developed in the United States and used there in the late 1960s 1970s Western Reserve Copenhagen Connaught Laboratories Canada References Edit ICTV 9th Report 2011 Poxviridae International Committee on Taxonomy of Viruses ICTV Retrieved 17 December 2018 Ryan KJ Ray CG eds 2004 Sherris Medical Microbiology 4th ed McGraw Hill ISBN 978 0 8385 8529 0 Johnson L Gupta A K Ghafoor A Akin D Bashir R 2006 Characterization of vaccinia virus particles using microscale silicon cantilever resonators and atomic force microscopy Sensors and Actuators B Chemical 115 1 189 197 doi 10 1016 j snb 2005 08 047 Baxby Derrick 1981 Jenner s Smallpox Vaccine The Riddle of Vaccinia Virus and Its Origin Heinemann Educational Books ISBN 978 0 435 54057 9 a b Downie AW 1939 The Immunological Relationship of the Virus of Spontaneous Cowpox to Vaccinia Virus British Journal of Experimental Pathology 20 2 158 176 PMC 2065307 Carroll Darin S Emerson Ginny L Li Yu Sammons Scott Olson Victoria Frace Michael Nakazawa Yoshinori Czerny Claus Peter Tryland Morten Kolodziejek Jolanta Nowotny Norbert Olsen Rasmussen Melissa Khristova Marina Govil Dhwani Karem Kevin Damon Inger K Meyer Hermann 8 August 2011 Chasing Jenner s Vaccine Revisiting Cowpox Virus Classification PLOS ONE 6 8 e23086 Bibcode 2011PLoSO 623086C doi 10 1371 journal pone 0023086 ISSN 1932 6203 PMC 3152555 PMID 21858000 James William D Berger Timothy G et al 2006 Andrews Diseases of the Skin clinical Dermatology Saunders Elsevier ISBN 978 0 7216 2921 6 a b Huygelen C 1996 Jenner s cowpox vaccine in light of current vaccinology Verh K Acad Geneeskd Belg in Dutch 58 5 479 536 discussion 537 538 PMID 9027132 Henderson DA Moss B 1999 1988 Smallpox and Vaccinia In Plotkin SA Orenstein WA eds Vaccines 3rd ed Philadelphia Pennsylvania WB Saunders ISBN 978 0 7216 7443 8 Schrick Livia Tausch Simon H Dabrowski P Wojciech Damaso Clarissa R Esparza Jose Nitsche Andreas 2017 An Early American Smallpox Vaccine Based on Horsepox New England Journal of Medicine 377 15 1491 1492 doi 10 1056 NEJMc1707600 PMID 29020595 Tolonen N Doglio L Schleich S Krijnse Locker J 1 July 2001 Vaccinia Virus DNA Replication Occurs in Endoplasmic Reticulum enclosed Cytoplasmic Mini Nuclei Mol Biol Cell 12 7 2031 46 doi 10 1091 mbc 12 7 2031 PMC 55651 PMID 11452001 Smith GL Vanderplasschen A Law M 1 December 2002 The formation and function of extracellular enveloped Vaccinia virus J Gen Virol 83 Pt 12 2915 31 doi 10 1099 0022 1317 83 12 2915 PMID 12466468 a b ABEL P August 1962 Multiplicity reactivation and marker rescue with vaccinia virus Virology 17 4 511 9 doi 10 1016 0042 6822 62 90150 2 PMID 13858909 Sharp DG Kim KS July 1966 Multiplicity reactivation and radiation survival of aggregated vaccinia virus Calculation of plaque titer based on MR and particle aggregation seen in the electron microscope Virology 29 3 359 66 doi 10 1016 0042 6822 66 90211 X PMID 5922451 Kim KS Sharp DG February 1967 Multiplicity reactivation of vaccinia virus particles treated with nitrogen mustard J Virol 1 1 45 9 doi 10 1128 JVI 1 1 45 49 1967 PMC 375503 PMID 5623957 Kim KS Sharp DG January 1968 Multiplicity reactivation of gamma and x irradiated Vaccinia virus in L cells Radiat Res 33 1 30 6 Bibcode 1968RadR 33 30K doi 10 2307 3572239 JSTOR 3572239 PMID 5634978 Michod RE Bernstein H Nedelcu AM 2008 Adaptive value of sex in microbial pathogens Infect Genet Evol 8 3 267 285 doi 10 1016 j meegid 2008 01 002 PMID 18295550 a b Davies MV Chang HW Jacobs BL Kaufman RJ 1 March 1993 The E3L and K3L vaccinia virus gene products stimulate translation through inhibition of the double stranded RNA dependent protein kinase by different mechanisms J Virol 67 3 1688 1692 doi 10 1128 JVI 67 3 1688 1692 1993 PMC 237544 PMID 8094759 Warren Luigi Manos Philip D Ahfeldt Tim Loh Yuin Han Li Hu Lau Frank Ebina Wataru Mandal Pankaj K Smith Zachary D Meissner Alexander Daley George Q Brack Andrew S Collins James J Cowan Chad Schlaeger Thorsten M Rossi Derrick J 2010 Highly Efficient Reprogramming to Pluripotency and Directed Differentiation of Human Cells with Synthetic Modified mRNA Cell Stem Cell 7 5 618 630 doi 10 1016 j stem 2010 08 012 PMC 3656821 PMID 20888316 Side Effects of Smallpox Vaccination Smallpox CDC 2017 07 12 Heilprin John 1 September 2007 FDA approves new smallpox vaccine Houston Chronicle AP Retrieved 25 May 2018 European public assessment report summary Imvanex 2018 09 17 Vanderplasschen A Pastoret P P December 2003 The Uses of Poxviruses as Vectors Current Gene Therapy 3 6 583 595 doi 10 2174 1566523034578168 PMID 14683453 Baxby D 1999 Edward Jenner s Inquiry a bicentenary analysis Vaccine 17 4 301 307 doi 10 1016 S0264 410X 98 00207 2 PMID 9987167 Tucker Jonathan B Scourge The Once and Future Threat of Smallpox New York Grove Atlantic Inc 2001 Esparza Jose Lederman Seth Nitsche Andreas Damaso Clarissa R 2020 06 19 Early smallpox vaccine manufacturing in the United States Introduction of the animal vaccine in 1870 establishment of vaccine farms and the beginnings of the vaccine industry Vaccine 38 30 4773 4779 doi 10 1016 j vaccine 2020 05 037 ISSN 0264 410X PMC 7294234 PMID 32473878 Steinhardt E Israeli C Lambert RA September 1913 Studies on the cultivation of the virus of vaccinia J Inf Dis 13 2 294 300 doi 10 1093 infdis 13 2 294 JSTOR 30073371 Phages their role in bacterial pathogenesis and biotechnology Waldor Matthew K Friedman David I Adhya Sankar Lal Washington D C ASM Press 2005 ISBN 1 55581 307 0 OCLC 57557385 a href wiki Template Cite book title Template Cite book cite book a CS1 maint others link Tyrrell D A J McCarthy K 1990 Allan Watt Downie September 1901 26 January 1988 Biographical Memoirs of Fellows of the Royal Society 35 98 112 doi 10 1098 rsbm 1990 0004 PMID 11622284 a b Centers for Disease Control and Prevention CDC 2007 Household transmission of vaccinia virus from contact with a military smallpox vaccinee Illinois and Indiana 2007 Morbidity and Mortality Weekly Report 56 19 478 81 PMID 17510612 SIGA s Smallpox Drug Candidate Administered to Critically Ill Human Patient Press release SIGA Technologies 2007 03 17 Retrieved 2018 07 20 Centers for Disease Control and Prevention CDC 2010 Vaccinia Virus Infection After Sexual Contact with a Military Smallpox Vaccinee Washington 2010 Morbidity and Mortality Weekly Report 59 25 773 75 PMID 20592687 Centers for Disease Control and Prevention CDC March 2013 Secondary and tertiary transmission of vaccinia virus after sexual contact with a smallpox vaccinee San Diego California 2012 Morbidity and Mortality Weekly Report 62 8 145 7 PMC 4604863 PMID 23446513 Notice to Readers Newly Licensed Smallpox Vaccine to Replace Old Smallpox Vaccine MMWR Morb Mortal Wkly Rep 57 8 207 8 February 29 2008 Further reading EditGubser C Hue S Kellam P Smith GL January 2004 Poxvirus genomes a phylogenetic analysis J Gen Virol 85 1 105 17 doi 10 1099 vir 0 19565 0 PMID 14718625 Centers for Disease Control and Prevention CDC 2007 Vulvar vaccinia infection after sexual contact with a military smallpox vaccinee Alaska 2006 MMWR Morb Mortal Wkly Rep 56 17 417 9 PMID 17476203 Al Ali S Baldanta S Fernandez Escobar M Guerra S 2016 Use of Reporter Genes in the Generation of Vaccinia Virus Derived Vectors Viruses 8 5 134 doi 10 3390 v8050134 PMC 4885089 PMID 27213433 Rubins K H Hensley L E Bell G W Wang C Lefkowitz E J Brown P O Relman D A 2008 Comparative analysis of viral gene expression programs during poxvirus infection A transcriptional map of the vaccinia and monkeypox genomes PLOS ONE 3 7 e2628 Bibcode 2008PLoSO 3 2628R doi 10 1371 journal pone 0002628 PMC 2440811 PMID 18612436 Vaccinia virus complete genome National Center for Biotechnology Information Retrieved 2007 07 25 Condit RC Moussatche N Traktman P The Vaccinia Virion 3D Tour Retrieved 2007 07 26 Smallpox Emergency Preparedness amp Response Centers for Disease Control and Prevention Archived from the original on 2007 08 13 Retrieved 2007 07 26 External links EditVirus Pathogen Database and Analysis Resource ViPR Poxviridae Retrieved from https en wikipedia org w index php title Vaccinia amp oldid 1092777275, wikipedia, wiki, book,

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