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The capsid of SV40, an icosahedral virus
The capsid of SV40, an icosahedral virus

Viruses are small infectious agents that can replicate only inside the living cells of an organism. Viruses infect all forms of life, including animals, plants, fungi, bacteria and archaea. They are found in almost every ecosystem on Earth and are the most abundant type of biological entity, with millions of different types, although only about 6,000 viruses have been described in detail. Some viruses cause disease in humans, and others are responsible for economically important diseases of livestock and crops.

Virus particles (known as virions) consist of genetic material, which can be either DNA or RNA, wrapped in a protein coat called the capsid; some viruses also have an outer lipid envelope. The capsid can take simple helical or icosahedral forms, or more complex structures. The average virus is about 1/100 the size of the average bacterium, and most are too small to be seen directly with an optical microscope.

The origins of viruses are unclear: some may have evolved from plasmids, others from bacteria. Viruses are sometimes considered to be a life form, because they carry genetic material, reproduce and evolve through natural selection. However they lack key characteristics (such as cell structure) that are generally considered necessary to count as life. Because they possess some but not all such qualities, viruses have been described as "organisms at the edge of life".

Selected disease

Child with a deformed right leg due to poliomyelitis
Child with a deformed right leg due to poliomyelitis

Polio, also called poliomyelitis or infantile paralysis, was one of the most feared childhood diseases of the 20th century. Poliovirus, the causative agent, only naturally infects humans and spreads via the faecal–oral route. Most infections cause no or minor symptoms. In around 1% of cases, the virus enters the central nervous system, causing aseptic meningitis. There it can preferentially infect and destroy motor neurons, leading in 0.1–0.5% of cases to muscle weakness and acute flaccid paralysis. Spinal polio accounts for nearly 80% of paralytic cases, with asymmetric paralysis of the legs being typical; in a quarter of these cases permanent severe disability results. Bulbar involvement is rare, but in severe cases the virus can prevent breathing by affecting the phrenic nerve, so that patients require mechanical ventilation with an iron lung or similar device.

Depictions in ancient art show that the disease has existed for thousands of years. The virus was an endemic pathogen until the 1880s, when major epidemics began to occur in Europe and later the United States. Polio vaccines were developed in the 1950s and a global eradication campaign started in 1988. The annual incidence of wild-type disease fell from many hundreds of thousands to 22 in 2017, but has since resurged to a few hundreds.

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Ribbon diagram showing T7 RNA polymerase (blue) making messenger RNA (green) based on a DNA template (orange)

Some viruses, such as the T7 bacteriophage, encode their own RNA polymerase, the enzyme that makes messenger RNA based on a DNA template. The T7 enzyme has a single subunit, and is more like chloroplast and mitochondrial enzymes than those of bacteria or the cell.

Credit: Thomas Splettstoesser (25 June 2007)

In the news

Map showing the prevalence of SARS-CoV-2 cases; black: highest prevalence; dark red to pink: decreasing prevalence; grey: no recorded cases or no data
Map showing the prevalence of SARS-CoV-2 cases; black: highest prevalence; dark red to pink: decreasing prevalence; grey: no recorded cases or no data

26 February: In the ongoing pandemic of severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), more than 110 million confirmed cases, including 2.5 million deaths, have been documented globally since the outbreak began in December 2019. WHO

18 February: Seven asymptomatic cases of avian influenza A subtype H5N8, the first documented H5N8 cases in humans, are reported in Astrakhan Oblast, Russia, after more than 100,0000 hens died on a poultry farm in December. WHO

14 February: Seven cases of Ebola virus disease are reported in Gouécké, south-east Guinea. WHO

7 February: A case of Ebola virus disease is detected in North Kivu Province of the Democratic Republic of the Congo. WHO

4 February: An outbreak of Rift Valley fever is ongoing in Kenya, with 32 human cases, including 11 deaths, since the outbreak started in November. WHO

21 November: The US Food and Drug Administration (FDA) gives emergency-use authorisation to casirivimab/imdevimab, a combination monoclonal antibody (mAb) therapy for non-hospitalised people twelve years and over with mild-to-moderate COVID-19, after granting emergency-use authorisation to the single mAb bamlanivimab earlier in the month. FDA 1, 2

18 November: The outbreak of Ebola virus disease in Équateur Province, Democratic Republic of the Congo, which started in June, has been declared over; a total of 130 cases were recorded, with 55 deaths. UN

Selected article

Martinus Beijerinck in his laboratory in 1921
Martinus Beijerinck in his laboratory in 1921

The history of virology is usually considered to begin in the late 19th century, when the first evidence for the existence of viruses came from experiments using filters with pores small enough to retain bacteria. Dmitry Ivanovsky showed in 1892 that sap from a diseased tobacco plant remained infectious despite having been filtered; this agent, later known as tobacco mosaic virus, was the first virus to be demonstrated. In 1898, Friedrich Loeffler and Paul Frosch showed that foot-and-mouth, an animal disease, was caused by a filterable agent. That year, Martinus Beijerinck (pictured) called the filtered infectious substance a "virus" – often considered to mark the beginning of virology.

Bacteriophages, viruses that infect bacteria, were characterised by Frederick Twort and Félix d'Herelle in the early 20th century. In 1926, Thomas Milton Rivers defined viruses as obligate parasites. Viruses were demonstrated to be particles, rather than a fluid, by Wendell Meredith Stanley, and the invention of the electron microscope in 1931 allowed them to be visualised.

Selected outbreak

Map showing Ebola virus disease cases in Guinea, Liberia and Sierra Leone in December 2014

The West African Ebola epidemic was the most widespread outbreak of the disease to date. Beginning in Meliandou in southern Guinea in December 2013, it spread to adjacent Liberia and Sierra Leone, affecting the cities of Conakry and Monrovia, with minor outbreaks in Mali and Nigeria. Cases reached a peak in October 2014 and the epidemic was under control by late 2015, although occasional cases continued to occur into April 2016. Ring vaccination with the then-experimental vaccine rVSV-ZEBOV was trialled in Guinea.

More than 28,000 suspected cases were reported with more than 11,000 deaths; the case fatality rate was around 40% overall and around 58% in hospitalised patients. Early in the epidemic nearly 10% of the dead were healthcare workers. The outbreak left about 17,000 survivors, many of whom reported long-lasting post-recovery symptoms. Extreme poverty, dysfunctional healthcare systems, distrust of government after years of armed conflict, local burial customs of washing the body, the unprecedented spread of Ebola to densely populated cities, and the delay in response of several months all contributed to the failure to control the epidemic.

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Recommended articles

Viruses & Subviral agents: bat virome • elephant endotheliotropic herpesvirus • HIV • introduction to viruses • Playa de Oro virus • poliovirus • prion • rotavirus • virus

Diseases: colony collapse disorder • common cold • croup • dengue fever • gastroenteritis • Guillain–Barré syndrome • hepatitis B • hepatitis C • hepatitis E • herpes simplex • HIV/AIDS • influenza • meningitis • myxomatosis • polio • pneumonia • shingles • smallpox

Epidemiology & Interventions: 2007 Bernard Matthews H5N1 outbreak • Coalition for Epidemic Preparedness Innovations • Disease X • 2009 flu pandemic • HIV/AIDS in Malawi • polio vaccine • Spanish flu • West African Ebola virus epidemic

Virus–Host interactions: antibody • host • immune system • parasitism • RNA interference

Methodology: metagenomics

Social & Media: And the Band Played On • Contagion • "Flu Season" • Frank's Cock • Race Against Time: Searching for Hope in AIDS-Ravaged Africa • social history of viruses • "Steve Burdick" • "The Time Is Now" • "What Lies Below"

People: Brownie Mary • Macfarlane Burnet • Bobbi Campbell • Aniru Conteh • people with hepatitis C • HIV-positive people • Bette Korber • Henrietta Lacks • Linda Laubenstein • Barbara McClintock • poliomyelitis survivors • Joseph Sonnabend • Eli Todd • Ryan White

Selected virus

Electron micrograph of two Epstein–Barr virus particles

Epstein–Barr virus (EBV) (also human herpesvirus 4) is a DNA virus in the Herpesviridae family which infects humans. The virion is around 122–180 nm in diameter. As in all herpesviruses, the nucleocapsid is surrounded by a protein tegument, as well as an envelope. The double-stranded DNA genome is about 172 kb, with 85 genes, making it one of the more complex viruses.

Transmission is via saliva and genital secretions. The virus infects epithelial cells in the pharynx and B cells of the immune system, producing virions by budding. EBV also becomes latent in B cells, possibly in the bone marrow, allowing the infection to persist lifelong. In the latent state, the linear genome is made circular and replicates in the nucleus separately from the host DNA as an episome. Reactivation is poorly understood but is thought to be triggered by the B cell responding to other infections. EBV infection occurs in around 95% of people. Infectious mononucleosis or glandular fever can occur when first infection is delayed until adolescence or adulthood. EBV is associated with some types of cancer, including Burkitt's lymphoma and nasopharyngeal carcinoma. In people with HIV, it can cause hairy leukoplakia and central nervous system lymphomas.

Did you know?

Chicken eyes, showing the appearance in Marek's disease (right) compared with a normal eye (left)
Chicken eyes, showing the appearance in Marek's disease (right) compared with a normal eye (left)

Selected biography

Ali Maow Maalin (1954 – 22 July 2013) was a hospital cook and health worker from Merca, Somalia, who is the last person in the world known to be infected with naturally occurring smallpox. Although he worked in the local smallpox eradication programme, he had not been successfully vaccinated. In October 1977, he was infected with the Variola minor strain of the virus while driving two children with smallpox symptoms to quarantine. He did not experience complications and made a full recovery. An aggressive containment campaign was successful in preventing an outbreak, and smallpox was declared to have been eradicated globally by the World Health Organization (WHO) two years later.

In later life, Maalin volunteered for the successful poliomyelitis eradication campaign in Somalia. He worked for WHO as a local coordinator with responsibility for social mobilisation, and spent several years travelling across Somalia, vaccinating children and educating communities. He encouraged people to be vaccinated by sharing his experiences with smallpox. He died of malaria while carrying out polio vaccinations after the reintroduction of poliovirus to the country in 2013.

In this month

Red ribbon signifying solidarity with people living with HIV/AIDS

5 June 1981: First report of HIV/AIDS (symbol pictured) appeared in medical literature

6 June 1997: Gene silencing in plants shown to be a viral defence mechanism

7–13 June 1962: Donald Caspar and Aaron Klug proposed the quasi-equivalence principle of virus structure

7–13 June 1962: André Lwoff proposed a viral classification scheme based on nature of genome, type of symmetry and presence of envelope

7–13 June 1962: George Hirst proposed that the influenza virus genome is segmented

9 June 1981: The American Society for Virology was founded

13 June 2012: First case of Middle East respiratory syndrome coronavirus (MERS-CoV) occurred in Saudi Arabia

18 June 1981: A vaccine against foot-and-mouth disease was the first genetically engineered vaccine

21 June 1996: Nevirapine approved, first NNRTI for HIV/AIDS

26 June 1993: Clinical trial of hepatitis B virus drug fialuridine terminated; the drug caused several fatalities due to lactic acidosis

28 June 2011: FAO declared rinderpest eradicated

30 June 1985: Ryan White was denied re-admittance to his school after an AIDS diagnosis, in a case that changed public perceptions of the disease

Selected intervention

Influenza vaccine strains are usually cultivated in fertilised chicken eggs.
Influenza vaccine strains are usually cultivated in fertilised chicken eggs.

Influenza vaccines include live attenuated and inactivated forms. Inactivated vaccines contain three or four different viral strains selected by the World Health Organization to cover influenza A H1N1 and H3N2, as well as influenza B, and are usually administered by intramuscular injection. The live attenuated influenza vaccine contains a cold-adapted strain and is given as a nasal spray. Most influenza vaccine strains are cultivated in fertilised chicken eggs (pictured), a technique developed in the 1950s; others are grown in cell cultures, and some vaccines contain recombinant proteins. Annual vaccination is recommended for high-risk groups and, in some countries, for all those over six months. As the influenza virus changes rapidly by antigenic drift, new versions of the vaccine are developed twice a year, which differ in effectiveness depending on how well they match the circulating strains. Despite considerable research effort for decades, no effective universal influenza vaccine has been identified. A 2018 meta-analysis found that vaccination in healthy adults decreased confirmed cases of influenza from about 2.4% to 1.1%. However, the effectiveness is uncertain in those over 65 years old, one of the groups at highest risk of serious complications.

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