Geschichte der Virologie

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[[File:TobaccoMosaicVirus.jpg|thumb|right|[[Electron microscope|Electron micrograph]] of the rod-shaped particles of [[tobacco mosaic virus]] that are too small to be seen using a light microscope]]

The '''history of virology''' – the scientific study of viruses and the infections they cause – began in the closing years of the 19th century. Although [[Louis Pasteur]] and [[Edward Jenner]] developed the first [[vaccine]]s to protect against viral infections, they did not know that viruses existed. The first evidence of the existence of viruses came from experiments with filters that had pores small enough to retain bacteria. In 1892, [[Dmitry Ivanovsky]] used one of these filters to show that sap from a diseased [[tobacco plant]] remained infectious to healthy tobacco plants despite having been filtered. [[Martinus Beijerinck]] called the filtered, infectious substance a "virus" and this discovery is considered to be the beginning of [[virology]]. The subsequent discovery and partial characterization of [[bacteriophage]]s by [[Felix d'Herelle]] further catalyzed the field, and by the early 20th century many viruses were discovered.

==Pioneers==
[[File:Mwb in lab.JPG|thumb|alt=An old, bespectacled man wearing a suit and sitting at a bench by a large window. The bench is covered with small bottles and test tubes. On the wall behind him is a large old-fashioned clock below frick u which are four small enclosed shelves on which sit many neatly labelled bottles.|[[Martinus Beijerinck]] in his laboratory in 1921]]

Despite his other successes, [[Louis Pasteur]] (1822–1895) was unable to find a causative agent for [[rabies]] and speculated about a pathogen too small to be detected using a microscope.<ref>{{cite journal |author=Bordenave G |title=Louis Pasteur (1822–1895) |journal=Microbes and Infection / Institut Pasteur |volume=5 |issue=6 |pages=553–60 |date=May 2003 |pmid=12758285 |doi=10.1016/S1286-4579(03)00075-3}}</ref> In 1884, the French [[microbiologist]] [[Charles Chamberland]] (1851–1931) invented a filter – known today as the [[Chamberland filter]] – that had pores smaller than bacteria. Thus, he could pass a solution containing bacteria through the filter and completely remove them from the solution.<ref name ="Shors 76–77">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |pages=76–77 |isbn=0-7637-2932-9}}</ref>

In 1876, [[Adolf Mayer]], who directed the Agricultural Experimental Station in [[Wageningen]] was the first to show that what he called "Tobacco Mosaic Disease" was infectious, he thought that it was caused by either a toxin or a very small bacterium. Later, in 1892, the Russian biologist [[Dmitry Ivanovsky]] (1864–1920) used a Chamberland filter to study what is now known as the [[tobacco mosaic virus]]. His experiments showed that crushed leaf extracts from infected tobacco plants remain infectious after filtration. Ivanovsky suggested the infection might be caused by a [[toxin]] produced by bacteria, but did not pursue the idea.<ref name="Collier 3">{{cite book |author1=Sussman, Max |author2=Topley, W. W. C. |author3=Wilson, Graham K. |author4=Collier, L. H. |author5=Balows, Albert |title=Topley & Wilson's microbiology and microbial infections |publisher=Arnold |location=London |year=1998 |page=3 |isbn=0-340-66316-2}}</ref>

In 1898, the Dutch microbiologist [[Martinus Beijerinck]] (1851–1931), a microbiology teacher at the Agricultural School in [[Wageningen]] repeated experiments by [[Adolf Mayer]] and became convinced that filtrate contained a new form of infectious agent.<ref name="Dimmock 4–5">{{cite book |author1=Leppard, Keith |author2=Nigel Dimmock |author3=Easton, Andrew |title=Introduction to Modern Virology |publisher=Blackwell Publishing Limited |location= |year=2007 |pages=4–5 |isbn=1-4051-3645-6 }}</ref> He observed that the agent multiplied only in cells that were dividing and he called it a ''[[contagium vivum fluidum]]'' (soluble living germ) and re-introduced the word ''virus''.<ref name="Collier 3"/> Beijerinck maintained that viruses were liquid in nature, a theory later discredited by the American biochemist and virologist [[Wendell Meredith Stanley]] (1904–1971), who proved that they were in fact, particles.<ref name="Collier 3"/> In the same year [[Friedrich Loeffler]] (1852–1915) and [[Paul Frosch]] (1860–1928) passed the first animal virus through a similar filter and discovered the cause of [[foot-and-mouth disease]].<ref>{{cite book |author=Fenner F. |editor=Mahy B. W. J. and Van Regenmortal M. H. V.|title=Desk Encyclopedia of General Virology |edition= 1|publisher=Academic Press |location=Oxford, UK |year=2009 |page=15|isbn=0-12-375146-2}}</ref>

In 1881, [[Carlos Finlay]] (1833–1915), a Cuban physician, first conducted and published research that indicated that mosquitoes were carrying the cause of yellow fever,<ref name="pmid2684378">{{cite journal |author=Chiong MA |title=Dr. Carlos Finlay and yellow fever |journal=Canadian Medical Association Journal |volume=141 |issue=11 |page=1126 |date=December 1989 |pmid=2684378 |pmc=1451274}}</ref> a theory proved in 1900 by commission headed by [[Walter Reed]] (1851–1902). During 1901 and 1902, [[William Crawford Gorgas]] (1854–1920) organised the destruction of the mosquitoes' breeding habitats in Cuba, which dramatically reduced the prevalence of the disease.<ref name="pmid11482006">{{cite journal |doi=10.1353/pbm.2001.0051 |author=Litsios S |title=William Crawford Gorgas (1854–1920) |journal=Perspectives in Biology and Medicine |volume=44 |issue=3 |pages=368–78 |year=2001 |pmid=11482006}}</ref> Gorgas later organised the elimination of the mosquitoes from Panama, which allowed the [[Panama Canal]] to be opened in 1914.<ref name="pmid2673502">{{cite journal |author=Patterson R |title=Dr. William Gorgas and his war with the mosquito |journal=Canadian Medical Association Journal |volume=141 |issue=6 |pages=596–7, 599 |date=September 1989 |pmid=2673502 |pmc=1451363}}</ref> The virus was finally isolated by [[Max Theiler]] (1899–1972) in 1932 who went on to develop a successful vaccine.<ref name="pmid20589188">{{cite journal |author=Frierson JG |title=The yellow fever vaccine: a history |journal=Yale Journal of Biology and Medicine |volume=83 |issue=2 |pages=77–85 |date=June 2010 |pmid=20589188 |pmc=2892770}}</ref>

By 1928 enough was known about viruses to enable the publication of ''Filterable Viruses'', a collection of essays covering all known viruses edited by [[Thomas Milton Rivers]] (1888–1962). Rivers, a survivor of [[typhoid fever]] contracted at the age of twelve, went on to have a distinguished career in virology. In 1926, he was invited to speak at a meeting organised by the Society of American Bacteriology where he said for the first time, "Viruses appear to be obligate parasites in the sense that their reproduction is dependent on living cells."<ref>[http://books.nap.edu/html/biomems/trivers.pdf Frank L. Horsfall Jnr.(1965) "Thomas Milton Rivers (1888–1962)—A biographical memoir" ''The National Academy of Sciences'' Washington D.C. Retrieved 3 December 2010].</ref>

From the 1950s to the 1960s, [[Chester M. Southam]], a prominent virologist, injected malignant [[HeLa]] cells into cancer patients, healthy individuals, and prison inmates from the [[Ohio Penitentiary]] in order to observe if cancer could be transmitted.<ref name="Broadway Paperbacks">{{cite book|last1=Skloot|first1=Rebecca|title=The Immortal Life of Henrietta Lacks|date=2010|publisher=Broadway Paperbacks|location=New York|pages=128–135}}</ref> He was also examining if one could become immune to cancer by developing an acquired immune response in hopes of creating a vaccine for cancer.<ref name="Broadway Paperbacks"/>

The notion that viruses were particles was not considered unnatural and fitted in nicely with the [[Germ theory of disease|germ theory]]. It is assumed that Dr. J. Buist of Edinburgh was the first person to see virus particles in 1886, when he reported seeing "micrococci" in vaccine lymph, though he had probably observed clumps of [[vaccinia]].<ref>*In 1887, Buist visualised one of the largest, Vaccinia virus, by optical microscopy after staining it. Vaccinia was not known to be a virus at that time. (Buist J.B.(1887) ''Vaccinia and Variola: a study of their life history'' Churchill, London)</ref> In the years that followed, as optical microscopes were improved "inclusion bodies" were seen in many virus-infected cells, but these aggregates of virus particles were still too small to reveal any detailed structure. It was not until the invention of [[electron microscopy|the electron microscope]] in 1931 by the German engineers [[Ernst Ruska]] (1906–1988) and [[Max Knoll]] (1887–1969),<ref>From ''Nobel Lectures, Physics 1981–1990'', (1993) Editor-in-Charge Tore Frängsmyr, Editor Gösta Ekspång, World Scientific Publishing Co., Singapore.</ref> that virus particles, especially [[bacteriophage]]s, were shown to have complex structures. The sizes of viruses determined using this new microscope fitted in well with those estimated by filtration experiments. Viruses were expected to be small, but the range of sizes came as a surprise. Some were only a little smaller than the smallest known bacteria, and the smaller viruses were of similar sizes to complex organic molecules.<ref>{{cite book |author1=Carr, N. G. |author2=Mahy, B. W. J. |author3=Pattison, J. R. |author4=Kelly, D. P. |title=The microbe 1984: Thirty-sixth Symposium of the Society for General Microbiology, held at the University of Warwick, April 1984 |publisher=Published for the Society for General Microbiology [by] Cambridge University Press |location=Cambridge |year=1984 |page=4 |isbn=0-521-26056-6}}</ref>

In 1935, Wendell Stanley examined the tobacco mosaic virus and found it was mostly made of protein.<ref>{{cite journal |vauthors=Stanley WM, Loring HS | year = 1936 | title = The isolation of crystalline tobacco mosaic virus protein from diseased tomato plants | url = | journal = Science | volume = 83 | issue = 2143| page = 85 | pmid = 17756690 | doi=10.1126/science.83.2143.85|bibcode = 1936Sci....83...85S }}</ref> In 1939, Stanley and [[Max Lauffer]] (1914) separated the virus into protein and [[nucleic acid]],<ref>{{cite journal |vauthors=Stanley WM, Lauffer MA | year = 1939 | title = Disintegration of tobacco mosaic virus in urea solutions | url = | journal = Science | volume = 89 | issue = 2311| pages = 345–347 | pmid = 17788438 | doi=10.1126/science.89.2311.345|bibcode = 1939Sci....89..345S }}</ref> which was shown by Stanley's postdoctoral fellow Hubert S. Loring to be specifically [[RNA]].<ref>{{cite journal |vauthors=Loring HS | year = 1939 | title = Properties and hydrolytic products of nucleic acid from tobacco mosaic virus | url = http://www.jbc.org/content/130/1/251.short | journal = Journal of Biological Chemistry | volume = 130 | issue=1 | pages = 251–258 }}</ref> The discovery of RNA in the particles was important because in 1928, [[Fred Griffith]] (c.1879–1941) provided the first evidence that its "cousin", [[DNA]], formed [[genes]].<ref>{{cite book |author=Burton E. Tropp |title=Molecular Biology: Genes to Proteins. Burton E. Tropp |publisher=Jones & Bartlett Publishers |location=Sudbury, Massachusetts |year=2007 |page=12 |isbn=0-7637-5963-5}}</ref>

In Pasteur's day, and for many years after his death, the word "virus" was used to describe any cause of infectious disease. Many [[bacteriologist]]s soon discovered the cause of numerous infections. However, some infections remained, many of them horrendous, for which no bacterial cause could be found. These agents were invisible and could only be grown in living animals. The discovery of viruses was the key that unlocked the door that withheld the secrets of the cause of these mysterious infections. And, although [[Koch's postulates]] could not be fulfilled for many of these infections, this did not stop the pioneer virologists from looking for viruses in infections for which no other cause could be found.<ref>{{cite book |author1=Carr, N. G. |author2=Mahy, B. W. J. |author3=Pattison, J. R. |author4=Kelly, D. P. |title=The microbe 1984: Thirty-sixth Symposium of the Society for General Microbiology, held at the University of Warwick, April 1984 |publisher=Published for the Society for General Microbiology [by] Cambridge University Press |location=Cambridge |year=1984 |page=3|isbn=0-521-26056-6 }}</ref>

==Bacteriophages==
{{Main article|Bacteriophage}}
[[File:Phage S-PM2.png|thumb|left|Bacteriophage]]

===Discovery===
Bacteriophages are the viruses that infect and replicate in bacteria. They were discovered in the early 20th century, by the English bacteriologist [[Frederick Twort]] (1877–1950).<ref name="Shors 589 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |page=589 |isbn=0-7637-2932-9}}</ref> But before this time, in 1896, the bacteriologist [[Ernest Hanbury Hankin]] (1865–1939) reported that something in the waters of the [[River Ganges]] could kill ''[[Vibrio cholerae]]'' – the cause of [[cholera]]. Whatever it was in the water could be passed through filters that remove bacteria but was destroyed by boiling.<ref name = "Ackerman p3">{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |page=3|isbn=0-12-375146-2 }}</ref> Twort discovered the action of bacteriophages on [[staphylococcus|staphylococci]] bacteria. He noticed that when grown on nutrient agar some colonies of the bacteria became watery or "glassy". He collected some of these watery colonies and passed them through a Chamberland filter to remove the bacteria and discovered that when the filtrate was added to fresh cultures of bacteria, they in turn became watery.<ref name="Shors 589 "/> He proposed that the agent might be "an amoeba, an ultramicroscopic virus, a living protoplasm, or an enzyme with the power of growth".<ref name = "Ackerman p3"/>

Félix d'Herelle (1873–1949) was a mainly self-taught French-Canadian microbiologist. In 1917 he discovered that "an invisible antagonist", when added to bacteria on [[agar]], would produce areas of dead bacteria.<ref name="Shors 589 "/> The antagonist, now known to be a bacteriophage could pass through a Chamberland filter. He accurately diluted a suspension of these viruses and discovered that the highest dilutions (lowest virus concentrations), rather than killing all the bacteria, formed discrete areas of dead organisms. Counting these areas and multiplying by the dilution factor allowed him to calculate the number of viruses in the original suspension.<ref name="D'Herelle F 2007">{{cite journal | pmid = 17855060 | doi=10.1016/j.resmic.2007.07.005 | volume=158 | issue=7 |date=September 2007 | pages=553–4 | author=D'Herelle F | title = On an invisible microbe antagonistic toward dysenteric bacilli: brief note by Mr. F. D'Herelle, presented by Mr. Roux☆ | journal = Research in Microbiology}}</ref> He realised that he had discovered a new form of virus and later coined the term "bacteriophage".<ref name="Ackermann H-W 2009 4">{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |page=4|isbn=0-12-375146-2 }}</ref><ref>"The antagonistic microbe can never be cultivated in media in the absence of the dysentery bacillus. It does not attack heat-killed dysentery bacilli, but is cultivated perfectly in a suspension of washed cells in physiological saline. This indicates that the anti dysentery microbe is an obligate bacteriophage".
Felix d'Herelle (1917) ''An invisible microbe that is antagonistic to the dysentery bacillus'' (1917) [http://202.114.65.51/fzjx/wsw/wswfzjs/pdf/1917p157.pdf Comptes rendus Acad. Sci. Paris Retrieved on 2 December 2010]</ref>
Between 1918 and 1921 d'Herelle discovered different types of bacteriophages that could infect several other species of bacteria including ''Vibrio cholerae''.<ref>{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |page=4 Table 1|isbn=0-12-375146-2 }}</ref> Bacteriophages were heralded as a potential treatment for diseases such as [[typhoid]] and [[cholera]], but their promise was forgotten with the development of [[penicillin]].<ref name="Ackermann H-W 2009 4"/> Since the early 1970s, bacteria have continued to develop resistance to [[antibiotic]]s such as [[penicillin]], and this has led to a renewed interest in the use of [[phage therapy|bacteriophages to treat serious infections]].<ref name="Shors 591 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |page=591|isbn=0-7637-2932-9}}</ref>

===Early research 1920–1940===
D'Herelle travelled widely to promote the use of bacteriophages in the treatment of bacterial infections. In 1928, he became professor of biology at [[Yale]] and founded several research institutes.<ref name="Shors 590 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |page=590|isbn=0-7637-2932-9}}</ref> He was convinced that bacteriophages were viruses despite opposition from established bacteriologists such as the Nobel Prize winner [[Jules Bordet]] (1870–1961). Bordet argued that bacteriophages were not viruses but just [[enzyme]]s released from [[lysogenic cycle|"lysogenic"]] bacteria. He said "the invisible world of d'Herelle does not exist".<ref>Quoted in: {{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |page=4|isbn=0-12-375146-2 }}</ref> But in the 1930s, the proof that bacteriophages were viruses was provided by [[Christopher Andrewes]] (1896–1988) and others. They showed that these viruses differed in size and in their chemical and [[serology|serological]] properties.
In 1940, the first [[electron microscope|electron micrograph]] of a bacteriophage was published and this silenced sceptics who had argued that bacteriophages were relatively simple enzymes and not viruses.<ref>{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |pages=3–5|isbn=0-12-375146-2 }}</ref>
Numerous other types of bacteriophages were quickly discovered and were shown to infect bacteria wherever they are found. But this early research was interrupted by [[World War II]]. Even d'Herelle was silenced. Despite his Canadian citizenship, he was interned by the [[Vichy France|Vichy Government]] until the end of the war.<ref>{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |page=5|isbn=0-12-375146-2 }}</ref>

===Modern era===
Knowledge of bacteriophages increased in the 1940s following the formation of the [[Phage group|Phage Group]] by scientists throughout the US. Among the members were [[Max Delbrück]] (1906–1981) who founded a course on bacteriophages at [[Cold Spring Harbor Laboratory]].<ref name="Shors 591 "/> Other key members of the Phage Group included [[Salvador Luria]] (1912–1991) and [[Alfred Hershey]] (1908–1997). During the 1950s, [[Hershey–Chase experiment|Hershey and Chase]] made important discoveries on the replication of DNA during their studies on a bacteriophage called [[Enterobacteria phage T2|T2]]. Together with Delbruck they were jointly awarded the 1969 Nobel Prize in Physiology or Medicine "for their discoveries concerning the replication mechanism and the genetic structure of viruses".<ref>[http://nobelprize.org/nobel_prizes/medicine/laureates/1969/ Nobel Organisation]</ref> Since then, the study of bacteriophages has provided insights into the switching on and off of genes, and a useful mechanism for introducing foreign genes into bacteria and many other fundamental mechanisms of [[molecular biology]].<ref>{{cite book |author=Ackermann H-W |title=Desk Encyclopedia of General Virology |publisher=Academic Press |location=Oxford |year=2009 |pages=5–10 Table 1|isbn=0-12-375146-2 }}</ref>

==Plant viruses==
In 1882, [[Adolf Mayer]] (1843–1942) described a condition of tobacco plants, which he called "mosaic disease" ("mozaïkziekte"). The diseased plants had [[variegation|variegated]] leaves that were [[mottle]]d.<ref>Mayer A (1882) Over de moza¨ıkziekte van de tabak: voorloopige mededeeling. Tijdschr
Landbouwkunde Groningen 2: 359–364 (In German)</ref> He excluded the possibility of a fungal infection and could not detect any bacterium and speculated that a "soluble, enzyme-like infectious principle was involved".<ref name="pmid16732421">Quoted in: {{cite journal |vauthors=van der Want JP, Dijkstra J |title=A history of plant virology |journal=Archives of Virology|volume=151 |issue=8 |pages=1467–98 |date=August 2006 |pmid=16732421 |doi=10.1007/s00705-006-0782-3}}</ref> He did not pursue his idea any further, and it was the filtration experiments of Ivanovsky and Beijerinck that suggested the cause was a previously unrecognised infectious agent. After tobacco mosaic was recognized as a virus disease, virus infections of many other plants were discovered.<ref name="pmid16732421"/>

The importance of tobacco mosaic virus in the history of viruses cannot be overstated. It was the first virus to be discovered, and the first to be [[crystal]]lised and its structure shown in detail. The first [[X-ray diffraction]] pictures of the crystallised virus were obtained by Bernal and Fankuchen in 1941. On the basis of her pictures, [[Rosalind Franklin]] discovered the full structure of the virus in 1955.<ref name="pmid18702397">{{cite journal|vauthors=Creager AN, Morgan GJ |title=After the double helix: Rosalind Franklin's research on Tobacco mosaic virus|journal=Isis
|volume=99|issue=2|pages=239–72|date=June 2008|pmid=18702397|doi=10.1086/588626}}</ref> In the same year, [[Heinz Fraenkel-Conrat]] and [[Robley Williams]] showed that purified tobacco mosaic virus RNA and its [[capsid|coat protein]] can assemble by themselves to form functional viruses, suggesting that this simple mechanism was probably the means through which viruses were created within their host cells.<ref name="Dimmock 12 ">{{cite book |author1=Leppard, Keith |author2=Nigel Dimmock |author3=Easton, Andrew |title=Introduction to Modern Virology |publisher=Blackwell Publishing Limited |location= |year=2007 |page=12 |isbn=1-4051-3645-6 }}</ref>

By 1935 many plant diseases were thought to be caused by viruses. In 1922, [[John Kunkel Small]] (1869–1938) discovered that insects could act as [[vector (epidemiology)|vectors]] and transmit virus to plants. In the following decade many diseases of plants were shown to be caused by viruses that were carried by insects and in 1939, [[Francis Holmes (virologist)|Francis Holmes]], a pioneer in plant virology,<ref name="pmid11718380">{{cite journal |vauthors=Pennazio S, Roggero P, Conti M |title=A history of plant virology. Mendelian genetics and resistance of plants to viruses |journal=New Microbiology |volume=24 |issue=4 |pages=409–24 |date=October 2001 |pmid=11718380 }}</ref> described 129 viruses that caused disease of plants.<ref name="Shors 563 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |page=563 |isbn=0-7637-2932-9}}</ref> Modern, intensive agriculture provides a rich environment for many plant viruses. In 1948, in Kansas, US, 7% of the wheat crop was destroyed by [[wheat streak mosaic virus]]. The virus was spread by mites called ''[[Aceria tulipae]]''.<ref>D. Hansing, C. O. Johnston, L. E. Melchers and H. Fellows (1949) "Kansas Phytopathological Notes: 1948" ''Transactions of the Kansas Academy of Science'' (1903–) Vol. 52, No. 3, pp. 363–369 [https://www.jstor.org/stable/3625805 Stable URL Retrieved on 13 December 2010]</ref>

In 1970, the Russian plant virologist [[Joseph Atabekov]] discovered that many plant viruses only infect a single species of host plant.<ref name="pmid11718380"/> The [[International Committee on Taxonomy of Viruses]] now recognises over 900 plant viruses.<ref name="Shors 564 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |page=564 |isbn=0-7637-2932-9}}</ref>

==20th century==
By the end of the 19th century, viruses were defined in terms of their [[infectivity]], their ability to be filtered, and their requirement for living hosts. Up until this time, viruses had only been grown in plants and animals, but in 1906, [[Ross Granville Harrison]] (1870–1959) invented a method for growing [[Tissue (biology)|tissue]] in [[lymph]],<ref>[http://books.nap.edu/html/biomems/rharrison.pdf J. S. Nicholas, ''Ross Granville Harrison 1870—1959 A Biographical Memoir'', National Academy of Sciences, 1961, Retrieved on December 4, 2010]</ref> and, in 1913, E Steinhardt, C Israeli, and RA Lambert used this method to grow [[vaccinia]] virus in fragments of guinea pig corneal tissue.<ref>Steinhardt, E; Israeli, C; and Lambert, R.A. (1913) "Studies on the cultivation of the virus of vaccinia" ''J. Inf Dis. 13, 294–300</ref> In 1928, HB and MC Maitland grew vaccinia virus in suspensions of minced hens' kidneys.<ref name="pmid13475780">{{cite journal |doi=10.1017/S0022172400037268 |vauthors=Maitland HB, Magrath DI |title=The growth in vitro of vaccinia virus in chick embryo chorio-allantoic membrane, minced embryo and cell suspensions |journal=The Journal of Hygiene |volume=55 |issue=3 |pages=347–60 |date=September 1957 |pmid=13475780 |pmc=2217967}}</ref> Their method was not widely adopted until the 1950s, when [[poliovirus]] was grown on a large scale for vaccine production.<ref name="Collier 4">{{cite book |author1=Sussman, Max |author2=Topley, W. W. C. |author3=Wilson, Graham K. |author4=Collier, L. H. |author5=Balows, Albert |title=Topley & Wilson's microbiology and microbial infections |publisher=Arnold |location=London |year=1998 |page=4 |isbn=0-340-66316-2}}</ref> In 1941–42, [[George Hirst (virologist)|George Hirst]] (1909–94) developed assays based on [[hemagglutination|haemagglutination]] to quantify a wide range of viruses as well as virus-specific antibodies in serum.<ref name=Joklik>Wolfgang Joklik|Joklik WK. (1999) When two is better than one: Thoughts on three decades of interaction between ''Virology'' and the ''Journal of Virology''. ''[[Journal of Virology|J Virol]]'' 73: 3520–3523 ([https://www.ncbi.nlm.nih.gov/pmc/articles/PMC104123/ text])</ref><ref name=Schlesinger>{{cite journal |vauthors=Schlesinger RW, Granoff A | year = 1994 | title = George K. Hirst (1909–1994) | url = | journal = [[Virology (journal)|Virology]] | volume = 200 | issue = 2| page = 327 | doi=10.1006/viro.1994.1196}}</ref>

===Influenza===
[[File:Influenza Pandemic Masked Typist.jpg|thumb|right|A woman working during the 1918–1919 influenza epidemic. The face mask probably afforded minimal protection.]]
{{Main article|Influenza}}

Although the [[influenza virus]] that caused the [[1918 flu pandemic|1918–1919]] influenza pandemic was not discovered until the 1930s, the descriptions of the disease and subsequent research has proved it was to blame.<ref name="Shors 238-344 ">{{cite book |author=Teri Shors |title=Understanding Viruses |publisher=Jones & Bartlett Publishers |location=Sudbury, Mass |year=2008 |pages= 238–344|isbn=0-7637-2932-9}}</ref>
The pandemic killed 40–50 million people in less than a year,<ref>{{cite book |author=Oldstone MBA |title=Viruses, Plagues, and History: Past, Present and Future |publisher=Oxford University Press, USA |year=2009 |page=306 |isbn=0-19-532731-4}}</ref> but the proof that it was caused by a virus was not obtained until 1933.<ref name="pmid15081510">{{cite journal |author=Cunha BA |title=Influenza: historical aspects of epidemics and pandemics |journal=Infectious Disease Clinics of North America |volume=18 |issue=1 |pages=141–55 |date=March 2004 |pmid=15081510 |doi=10.1016/S0891-5520(03)00095-3}}</ref> ''[[Haemophilus influenzae]]'' is an opportunistic bacterium which commonly follows influenza infections; this led the eminent German bacteriologist [[Richard Friedrich Johannes Pfeiffer|Richard Pfeiffer]] (1858–1945) to incorrectly conclude that this bacterium was the cause of influenza.<ref>{{cite book |author=Oldstone MBA |title=Viruses, Plagues, and History: Past, Present and Future |publisher=Oxford University Press, USA |year=2009 |page=315 |isbn=0-19-532731-4}}</ref> A major breakthrough came in 1931, when the American pathologist [[Ernest William Goodpasture]] grew influenza and several other viruses in fertilised chickens' eggs.<ref>{{cite journal |vauthors=Goodpasture EW, Woodruff AM, Buddingh GJ | year = 1931 | title = The cultivation of vaccine and other viruses in the chorioallantoic membrane of chick embryos | url = | journal = Science | volume = 74 | issue = 1919| pages = 371–372 | pmid = 17810781 | doi=10.1126/science.74.1919.371|bibcode = 1931Sci....74..371G }}</ref> Hirst identified an enzymic activity associated with the virus particle, later characterised as the [[viral neuraminidase|neuraminidase]], the first demonstration that viruses could contain enzymes. [[Frank Macfarlane Burnet]] showed in the early 1950s that the virus recombines at high frequencies, and Hirst later deduced that it has a segmented genome.<ref name=Kilbourne>Kilbourne ED. (1975) Presentation of the Academy Medal to George K. Hirst, M.D. ''Bulletin of the New York Academy of Medicine'' 51: 1133–1136 ([https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1749565/pdf/bullnyacadmed00166-0013.pdf pdf])</ref>

===Poliomyelitis===
{{Main article|Poliomyelitis}}

In 1949, [[John F. Enders]] (1897–1985) [[Thomas Huckle Weller|Thomas Weller]] (1915–2008), and [[Frederick Robbins]] (1916–2003) grew polio virus for the first time in cultured human embryo cells, the first virus to be grown without using solid animal tissue or eggs. Infections by poliovirus most often cause the mildest of symptoms. This was not known until the virus was isolated in cultured cells and many people were shown to have had mild infections that did not lead to poliomyelitis. But, unlike other viral infections, the incidence of polio – the rarer severe form of the infection – increased in the 20th century and reached a peak around 1952. The invention of a [[cell culture]] system for growing the virus enabled [[Jonas Salk]] (1914–1995) to make an effective [[polio vaccine]].<ref>{{cite journal | author = Rosen FS | year = 2004 | title = Isolation of poliovirus—John Enders and the Nobel Prize | url = | journal = New England Journal of Medicine | volume = 351 | issue = 15| pages = 1481–83 | pmid = 15470207 | doi=10.1056/NEJMp048202}}</ref>

===Epstein–Barr virus===
[[Denis Parsons Burkitt]] (1911–1993) was born in Enniskillen, County Fermanagh, Ireland. He was the first to describe a type of cancer that now bears his name [[Burkitt's lymphoma]]. This type of cancer was endemic in equatorial Africa and was the commonest malignancy of children in the early 1960s.<ref name="pmid19620863">{{cite journal |author=Magrath I |title=Lessons from clinical trials in African Burkitt lymphoma |journal=Current Opinion in Oncology |volume=21 |issue=5 |pages=462–8 |date=September 2009 |pmid=19620863 |doi=10.1097/CCO.0b013e32832f3dcd}}</ref> In an attempt to find a cause for the cancer, Burkitt sent cells from the tumour to [[Anthony Epstein]] (b. 1921) a British virologist, who along with [[Yvonne Barr]] and [[Bert Achong]] (1928–1996), and after many failures, discovered viruses that resembled herpes virus in the fluid that surrounded the cells. The virus was later shown to be a previously unrecognised herpes virus, which is now called [[Epstein–Barr virus]].<ref>{{cite book |last= Epstein |first= M. Anthony |authorlink= M. Anthony Epstein |editor1-last=Robertson |editor1-first=Earl S. |title= Epstein-Barr Virus |url= https://books.google.com/books?id=TRO-wXto8hcC |accessdate= 18 September 2010 |year= 2005 |publisher= Cromwell Press |location= Trowbridge |isbn= 1-904455-03-4 |pages=1–14 |chapter= 1. The origins of EBV research: discovery and characterization of the virus }}</ref> Surprisingly, Epstein–Barr virus is a very common but relatively mild infection of Europeans. Why it can cause such a devastating illness in Africans is not fully understood, but reduced immunity to virus caused by [[malaria]] might be to blame.<ref name="pmid19165855">{{cite journal |author=Bornkamm GW |title=Epstein-Barr virus and the pathogenesis of Burkitt's lymphoma: more questions than answers |journal=International Journal of Cancer. Journal International Du Cancer |volume=124 |issue=8 |pages=1745–55 |date=April 2009 |pmid=19165855 |doi=10.1002/ijc.24223}}</ref> Epstein–Barr virus is important in the history of viruses for being the first virus shown to cause cancer in humans.<ref name="pmid16083776">{{cite journal |author=Thorley-Lawson DA |title=EBV the prototypical human tumor virus—just how bad is it? |journal=The Journal of Allergy and Clinical Immunology |volume=116 |issue=2 |pages=251–61; quiz 262 |date=August 2005 |pmid=16083776 |doi=10.1016/j.jaci.2005.05.038 }}</ref>

===Late 20th century===
[[File:Rotavirus Reconstruction.jpg|thumb|right|150px|A [[rotavirus]] particle]]

The second half of the 20th century was the golden age of virus discovery and most of the 2,000 recognised species of animal, plant, and bacterial viruses were discovered during these years.<ref name="pmid18446425">{{cite journal |author=Norrby E |title=Nobel Prizes and the emerging virus concept |journal=Archives of Virology |volume=153 |issue=6 |pages=1109–23 |year=2008 |pmid=18446425 |doi=10.1007/s00705-008-0088-8 }}</ref><ref>[http://talk.ictvonline.org/media/p/633.aspx Frederick A Murphy (2008) "Discoverers and Discoveries", ''International Committee on Taxonomy of Viruses'']</ref> In 1946, [[Bovine virus diarrhea]] was discovered,<ref name="pmid20995890">{{cite journal |vauthors=Olafson P, MacCallum AD, Fox FH |title=An apparently new transmissible disease of cattle |journal=The Cornell Veterinarian |volume=36 |issue= |pages=205–13 |date=July 1946 |pmid=20995890}}</ref> which is still possibly the commonest pathogen of cattle throughout the world<ref name="pmid20197026">{{cite journal |vauthors=Peterhans E, Bachofen C, Stalder H, Schweizer M |title=Cytopathic bovine viral diarrhea viruses (BVDV): emerging pestiviruses doomed to extinction |journal=Veterinary Research |volume=41 |issue=6 |page=44 |year=2010 |pmid=20197026 |pmc=2850149 |doi=10.1051/vetres/2010016}}</ref> and in 1957, [[Arterivirus|equine arterivirus]] was discovered.<ref name="pmid13397177">{{cite journal |vauthors=Bryans JT, Crowe ME, Doll ER, McCollum WH |title=Isolation of a filterable agent causing arteritis of horses and abortion by mares; its differentiation from the equine abortion (influenza) virus |journal=The Cornell Veterinarian |volume=47 |issue=1 |pages=3–41 |date=January 1957 |pmid=13397177}}</ref> In the 1950s, improvements in virus isolation and detection methods resulted in the discovery of several important human viruses including [[Varicella zoster virus]],<ref name="pmid8545033">{{cite journal |author=Weller TH |title=Varicella-zoster virus: History, perspectives, and evolving concerns |journal=Neurology |volume=45 |issue=12 Suppl 8 |pages=S9–10 |date=December 1995 |pmid=8545033 |doi=10.1212/wnl.45.12_suppl_8.s9}}</ref> the [[paramyxovirus]]es,<ref name="pmid15522442">{{cite journal |vauthors=Schmidt AC, Johnson TR, Openshaw PJ, Braciale TJ, Falsey AR, Anderson LJ, Wertz GW, Groothuis JR, Prince GA, Melero JA, Graham BS |title=Respiratory syncytial virus and other pneumoviruses: a review of the international symposium—RSV 2003 |journal=Virus Research |volume=106 |issue=1 |pages=1–13 |date=November 2004 |pmid=15522442 |doi=10.1016/j.virusres.2004.06.008}}</ref> – which include [[measles]] virus,<ref name="pmid19203111">{{cite journal |doi=10.1007/978-3-540-70617-5_10 |vauthors=Griffin DE, Pan CH |title=Measles: old vaccines, new vaccines |journal=Current Topics in Microbiology and Immunology |volume=330 |issue= |pages=191–212 |year=2009 |pmid=19203111|series=Current Topics in Microbiology and Immunology |isbn=978-3-540-70616-8 }}</ref> and [[respiratory syncytial virus]]<ref name="pmid15522442"/> – and the [[rhinovirus]]es that cause the [[common cold]].<ref name="pmid3039038">{{cite journal |author=Tyrrell DA |title=The common cold—my favourite infection. The eighteenth Majority Stephenson memorial lecture |journal=The Journal of General Virology |volume=68 |issue= 8|pages=2053–61 |date=August 1987 |pmid=3039038 |doi=10.1099/0022-1317-68-8-2053}}</ref> In the 1960s more viruses were discovered. In 1963, the [[Hepatitis B|hepatitis B virus]] was discovered by [[Baruch Blumberg]] (b. 1925),<ref name="pmid18298788">{{cite journal |author=Zetterström R |title=Nobel Prize to Baruch Blumberg for the discovery of the aetiology of hepatitis B |journal=Acta Paediatrica (Oslo, Norway : 1992) |volume=97 |issue=3 |pages=384–7 |date=March 2008 |pmid=18298788 |doi=10.1111/j.1651-2227.2008.00669.x }}</ref> and in 1965, [[Howard Temin]] (1934–1994) described the first [[retrovirus]].<ref name="pmid9001412">{{cite journal |author=Yoshida M |title=Howard Temin memorial lectureship. Molecular biology of HTLV-1: deregulation of host cell gene expression and cell cycle |journal=Leukemia : Official Journal of the Leukemia Society of America, Leukemia Research Fund, U.K |volume=11 |issue=1 |pages=14–5 |date=January 1997 |pmid=9001412 |doi=10.1038/sj.leu.2400545}}</ref> [[Reverse transcriptase]], the key enzyme that retroviruses use to [[translation (biology)|translate]] their RNA into DNA, was first described in 1970, independently by Howard Temin and [[David Baltimore]] (b. 1938).<ref name="pmid4348509">{{cite journal|vauthors=Temin HM, Baltimore D |title=RNA-directed DNA synthesis and RNA tumor viruses|journal=Advances in Virus Research
|volume=17|pages=129–86|year=1972|pmid=4348509
|doi=10.1016/S0065-3527(08)60749-6|series=Advances in Virus Research|isbn=9780120398171}}</ref> This was important to the development of [[antiviral drug]]s – a key turning-point in the history of viral infections.<ref name="pmid20018391">{{cite journal |author=Broder S |title=The development of antiretroviral therapy and its impact on the HIV-1/AIDS pandemic |journal=Antiviral Research |volume=85 |issue=1 |pages=1–18 |date=January 2010 |pmid=20018391 |pmc=2815149 |doi=10.1016/j.antiviral.2009.10.002}}</ref> In 1983 [[Luc Montagnier]] (b. 1932) and his team at the [[Pasteur Institute]] in France, first isolated the retrovirus now called HIV.<ref name="pmid6189183">{{cite journal |vauthors=Barré-Sinoussi F, Chermann JC, Rey F, Nugeyre MT, Chamaret S, Gruest J, Dauguet C, Axler-Blin C, Vézinet-Brun F, Rouzioux C, Rozenbaum W, Montagnier L |title=Isolation of a T-lymphotropic retrovirus from a patient at risk for acquired immune deficiency syndrome (AIDS) |journal=Science |volume=220 |issue=4599 |pages=868–71 |date=May 1983 |pmid=6189183 |doi=10.1126/science.6189183 |bibcode=1983Sci...220..868B}}</ref> In 1989 [[Michael Houghton (virologist)|Michael Houghton]]'s team at [[Chiron Corporation]] discovered [[Hepatitis C]].<ref name="pmid19781804">{{cite journal |author=Houghton M |title=The long and winding road leading to the identification of the hepatitis C virus |journal=Journal of Hepatology |volume=51 |issue=5 |pages=939–48 |date=November 2009 |pmid=19781804 |doi=10.1016/j.jhep.2009.08.004 |url=http://www.journal-of-hepatology.eu/article/S0168-8278%2809%2900535-2/fulltext}}</ref>
New viruses and strains of viruses were discovered in every decade of the second half of the 20th century. These discoveries have continued in the 21st century as new viral diseases such as [[Severe acute respiratory syndrome|SARS]]<ref name="pmid21116811">{{cite journal |vauthors=Peiris JS, Poon LL |title=Detection of SARS Coronavirus |journal=Methods in Molecular Biology (Clifton, N.J.) |volume=665 |issue= |pages=369–82 |year=2011 |pmid=21116811 |doi=10.1007/978-1-60761-817-1_20|series=Methods in Molecular Biology |isbn=978-1-60761-816-4 }}</ref> and [[nipah virus]]<ref name="pmid11334748">{{cite journal |vauthors=Field H, Young P, Yob JM, Mills J, Hall L, Mackenzie J |title=The natural history of Hendra and Nipah viruses |journal=Microbes and Infection / Institut Pasteur |volume=3 |issue=4 |pages=307–14 |date=April 2001 |pmid=11334748 |doi=10.1016/S1286-4579(01)01384-3}}</ref> have emerged. Despite scientists' achievements over the past one hundred years, viruses continue to pose new threats and challenges.<ref>{{cite book |author=Mahy BWJ |title=Desk Encyclopedia of Human and Medical Virology |publisher=Academic Press |location=Boston |year=2009 |pages=583–587 |isbn=0-12-375147-0}}</ref>

{| class ="wikitable collapsible collapsed"
|-
|+ '''Some of the many viruses discovered in the 20th century'''
! scope="col" | Year
! scope="col" | Virus
! scope="col" | References
|-
! scope="row" | 1908
| [[poliovirus]]
|<ref name="pmid20683737">{{cite journal |author=Skern T |title=100 years poliovirus: from discovery to eradication. A meeting report |journal=Archives of Virology |volume=155 |issue=9 |pages=1371–81 |date=September 2010 |pmid=20683737 |doi=10.1007/s00705-010-0778-x}}</ref>
|-
! scope="row" | 1911
| [[Rous sarcoma virus]]
|<ref name="pmid20503720">{{cite journal |author=Becsei-Kilborn E |title=Scientific discovery and scientific reputation: the reception of Peyton Rous' discovery of the chicken sarcoma virus |journal=Journal of the History of Biology |volume=43 |issue=1 |pages=111–57 |year=2010 |pmid=20503720 |doi= 10.1007/s10739-008-9171-y|url=}}</ref>
|-
! scope="row" | 1915
| bacteriophage of staphylococci
|<ref name="Shors 589 "/>
|-
! scope="row" | 1917
| bacteriophage of shigellae
|<ref name="Shors 589 "/>
|-
! scope="row" | 1918
| bacteriophage of salmonellae
|<ref name="D'Herelle F 2007"/>
|-
! scope="row" | 1927
| [[yellow fever]] virus
|<ref name="pmid20513550">{{cite journal |vauthors=Gardner CL, Ryman KD |title=Yellow fever: a reemerging threat |journal=Clinics in Laboratory Medicine |volume=30 |issue=1 |pages=237–60 |date=March 2010 |pmid=20513550 |pmc=4349381 |doi=10.1016/j.cll.2010.01.001 |url=}}</ref>
|-
! scope="row" | 1930
| [[western equine encephalitis virus]]
|<ref name="pmid19775836">{{cite journal |vauthors=Zacks MA, Paessler S |title=Encephalitic alphaviruses |journal=Veterinary Microbiology |volume=140 |issue=3–4 |pages=281–6 |date=January 2010 |pmid=19775836 |pmc=2814892 |doi=10.1016/j.vetmic.2009.08.023 |url=}}</ref>
|-
! scope="row" | 1933
| [[eastern equine encephalitis virus]]
|<ref name="pmid19775836"/>
|-
! scope="row" | 1934
| [[mumps virus]]
|<ref name="pmid19870227">{{cite journal |doi=10.1084/jem.59.1.1 |vauthors=Johnson CD, Goodpasture EW |title=An investigation of the etiology of mumps|journal=The Journal of Experimental Medicine |volume=59 |issue=1 |pages=1–19 |date=January 1934 |pmid=19870227 |pmc=2132344}}</ref>
|-
! scope="row" | 1935
| [[Japanese encephalitis]] virus
|<ref name="pmid20132860">{{cite journal |vauthors=Misra UK, Kalita J |title=Overview: Japanese encephalitis |journal=Progress in Neurobiology |volume=91 |issue=2 |pages=108–20 |date=June 2010 |pmid=20132860 |doi=10.1016/j.pneurobio.2010.01.008 |url=}}</ref>
|-
! scope="row" | 1943
| [[Dengue]] virus
|<ref name="pmid20513545">{{cite journal |author=Ross TM |title=Dengue virus |journal=Clinics in Laboratory Medicine |volume=30 |issue=1 |pages=149–60 |date=March 2010 |pmid=20513545 |doi=10.1016/j.cll.2009.10.007 |url=}}</ref>
|-
! scope="row" | 1949
| [[enterovirus]]es
|<ref name="pmid8024744">{{cite journal |author=Melnick JL |title=The discovery of the enteroviruses and the classification of poliovirus among them |journal=Biologicals : Journal of the International Association of Biological Standardization |volume=21 |issue=4 |pages=305–9 |date=December 1993 |pmid=8024744 |doi=10.1006/biol.1993.1088 |url=}}</ref>
|-
! scope="row" | 1952
| [[Varicella zoster virus]]
|<ref name="pmid8545033"/>
|-
! scope="row" | 1953
| [[adenovirus]]
|<ref name="isbn0-7817-6060-7">{{cite book |author1=Martin, Malcolm A. |author2=Knipe, David M. |author3=Fields, Bernard N. |author4=Howley, Peter M. |author5=Griffin, Diane |author6=Lamb, Robert |title=Fields' virology |publisher=Wolters Kluwer Health/Lippincott Williams & Wilkins |location=Philadelphia |year=2007 |page=2395 |isbn=0-7817-6060-7}}</ref>
|-
! scope="row" | 1954
| [[measles virus]]
|<ref name="pmid19203111"/>
|-
! scope="row" | 1956
| [[paramyxovirus]]es, [[rhinovirus]]
|<ref name="pmid15522442"/><ref name="pmid3039038"/>
|-
! scope="row" | 1958
| [[monkeypox]]
|<ref name="pmid1331540">{{cite journal |vauthors=Douglass N, Dumbell K |title=Independent evolution of monkeypox and variola viruses |journal=Journal of Virology |volume=66 |issue=12 |pages=7565–7 |date=December 1992 |pmid=1331540 |pmc=240470}}</ref>
|-
! scope="row" | 1962
| [[rubella virus]]
|<ref name="pmid3890105">{{cite journal |author=Cooper LZ |title=The history and medical consequences of rubella |journal=Reviews of Infectious Diseases |volume=7 Suppl 1 |issue= |pages=S2–10 |year=1985 |pmid=3890105 |doi=10.1093/clinids/7.supplement_1.s2}}</ref>
|-
! scope="row" | 1963
| [[hepatitis B virus]]
|<ref name="pmid16190102">{{cite journal |author=Yap SF |title=Hepatitis B: review of development from the discovery of the "Australia Antigen" to end of the twentieth Century |journal=The Malaysian Journal of Pathology |volume=26 |issue=1 |pages=1–12 |date=June 2004 |pmid=16190102}}</ref>
|-
! scope="row" | 1964
| [[Epstein–Barr virus]]
|<ref name="pmid4288580">{{cite journal |vauthors=Epstein MA, Achong BG, Barr YM, Zajac B, Henle G, Henle W |title=Morphological and virological investigations on cultured Burkitt tumor lymphoblasts (strain Raji) |journal=Journal of the National Cancer Institute |volume=37 |issue=4 |pages=547–59 |date=October 1966 |pmid=4288580}}</ref>
|-
! scope="row" | 1965
| [[retrovirus]]es
|<ref name="pmid15682876">{{cite journal |doi=10.1017/S1464793104006505 |author=Karpas A |title=Human retroviruses in leukaemia and AIDS: reflections on their discovery, biology and epidemiology |journal=Biological Reviews of the Cambridge Philosophical Society |volume=79 |issue=4 |pages=911–33 |date=November 2004 |pmid=15682876}}</ref>
|-
! scope="row" | 1966
| [[Lassa fever]] virus
|<ref name="pmid16802617">{{cite journal |author=Curtis N |title=Viral haemorrhagic fevers caused by Lassa, Ebola and Marburg viruses |journal=Advances in Experimental Medicine and Biology |volume=582 |issue= |pages=35–44 |year=2006 |pmid=16802617 |doi=10.1007/0-387-33026-7_4 |series=Advances in Experimental Medicine and Biology |isbn=978-0-387-31783-0 }}</ref>
|-
! scope="row" | 1967
| [[Marburg]] virus
|<ref name="pmid20513546">{{cite journal |vauthors=Hartman AL, Towner JS, Nichol ST |title=Ebola and marburg hemorrhagic fever |journal=Clinics in Laboratory Medicine |volume=30 |issue=1 |pages=161–77 |date=March 2010 |pmid=20513546 |doi=10.1016/j.cll.2009.12.001}}</ref>
|-
! scope="row" | 1972
| [[norovirus]]
|<ref name="pmid10804141">{{cite journal |author=Kapikian AZ |title=The discovery of the 27-nm Norwalk virus: an historic perspective |journal=The Journal of Infectious Diseases |volume=181 Suppl 2 |issue= |pages=S295–302 |date=May 2000 |pmid=10804141 |doi=10.1086/315584}}</ref>
|-
! scope="row" | 1973
| [[rotavirus]], [[hepatitis A virus]]
|<ref name="pmid186236">{{cite journal |author=Bishop RF, Cameron DJ, Barnes GL, Holmes IH, Ruck BJ |title=The aetiology of diarrhoea in newborn infants |journal=Ciba Foundation Symposium |volume= |issue=42 |pages=223–36 |year=1976 |pmid=186236 |doi=10.1002/9780470720240.ch13|series=Novartis Foundation Symposia |isbn=9780470720240 }}</ref><ref name="pmid6307916">{{cite journal |doi=10.1159/000149367 |vauthors=Gust ID, Coulepis AG, Feinstone SM, Locarnini SA, Moritsugu Y, Najera R, Siegl G |title=Taxonomic classification of hepatitis A virus |journal=Intervirology |volume=20 |issue=1 |pages=1–7 |year=1983 |pmid=6307916}}</ref>
|-
! scope="row" | 1975
| [[parvovirus B19]]
|<ref name="pmid6117755">{{cite journal |author=Cossart Y |title=Parvovirus B19 finds a disease |journal=Lancet |volume=2 |issue=8253 |pages=988–9 |date=October 1981 |pmid=6117755 |doi=10.1016/S0140-6736(81)91185-5}}</ref>
|-
! scope="row" | 1976
| [[Ebola]] virus
|<ref name="pmid21084112">{{cite journal |vauthors=Feldmann H, Geisbert TW |title=Ebola haemorrhagic fever |journal=Lancet |volume= 377|issue= 9768|pages= 849–862|date=November 2010 |pmid=21084112 |doi=10.1016/S0140-6736(10)60667-8 |url= |pmc=3406178}}</ref>
|-
! scope="row" | 1980
| [[human T-lymphotropic virus 1]]
|<ref name="pmid16155599">{{cite journal |author=Gallo RC |title=History of the discoveries of the first human retroviruses: HTLV-1 and HTLV-2 |journal=Oncogene |volume=24 |issue=39 |pages=5926–30 |date=September 2005 |pmid=16155599 |doi=10.1038/sj.onc.1208980}}</ref>
|-
! scope="row" | 1982
| [[human T-lymphotropic virus 2]]
|<ref name="pmid16155599"/>
|-
! scope="row" | 1983
| [[HIV]]
|<ref name="pmid20152474">{{cite journal |author=Montagnier L |title=25 years after HIV discovery: prospects for cure and vaccine |journal=Virology |volume=397 |issue=2 |pages=248–54 |date=February 2010 |pmid=20152474 |doi=10.1016/j.virol.2009.10.045}}</ref>
|-
! scope="row" | 1986
| [[human herpesvirus 6]]
|<ref name="pmid15653828">{{cite journal |vauthors=De Bolle L, Naesens L, De Clercq E |title=Update on human herpesvirus 6 biology, clinical features, and therapy |journal=Clinical Microbiology Reviews |volume=18 |issue=1 |pages=217–45 |date=January 2005 |pmid=15653828 |pmc=544175 |doi=10.1128/CMR.18.1.217-245.2005}}</ref>
|-
! scope="row" |1989
| [[hepatitis C virus]]
|<ref name="pmid2523562">{{cite journal |vauthors=Choo QL, Kuo G, Weiner AJ, Overby LR, Bradley DW, Houghton M |title=Isolation of a cDNA clone derived from a blood-borne non-A, non-B viral hepatitis genome |journal=Science |volume=244 |issue=4902 |pages=359–62 |date=April 1989 |pmid=2523562 |doi=10.1126/science.2523562|bibcode = 1989Sci...244..359C }}</ref>
|-
! scope="row" | 1990
| [[hepatitis E virus]], [[Human herpesvirus 7]]
|<ref name="pmid20335188">{{cite journal |vauthors=Bihl F, Negro F |title=Hepatitis E virus: a zoonosis adapting to humans |journal=The Journal of Antimicrobial Chemotherapy |volume=65 |issue=5 |pages=817–21 |date=May 2010 |pmid=20335188 |doi=10.1093/jac/dkq085}}</ref>
|-
! scope="row" | 1994
| [[henipavirus]]
|<ref name="pmid18511217">{{cite journal |author=Wild TF |title=Henipaviruses: a new family of emerging Paramyxoviruses |journal=Pathologie-biologie |volume=57 |issue=2 |pages=188–96 |date=March 2009 |pmid=18511217 |doi=10.1016/j.patbio.2008.04.006 }}</ref>
|-
! scope="row" | 1997
| ''[[Anelloviridae]]''
|<ref name="pmid19230554">{{cite journal |doi=10.1007/978-3-540-70972-5_1 |author=Okamoto H |title=History of discoveries and pathogenicity of TT viruses |journal=Current Topics in Microbiology and Immunology |volume=331 |issue= |pages=1–20 |year=2009 |pmid=19230554|series=Current Topics in Microbiology and Immunology |isbn=978-3-540-70971-8 }}</ref>
|-
|}

==See also==
{{Portal bar|History of science|Medicine|Viruses}}
*[[List of viruses]]

==References==
{{Reflist|colwidth=30em}}

==Further reading==
* {{cite book|title=The Virus: A History of the Concept|author=Hughes, Sally Smith|publisher=Heinemann|place=London|year=1977|isbn=0882021680}}

{{Virus topics}}
{{Baltimore (virus classification)}}
{{History of medicine}}

[[Category:Virology|*Virology, History of]]
[[Category:History of science and technology in the Netherlands]]
[[Category:Martinus Beijerinck]]

Wikipedia - Benutzerbeiträge [de]

Geschichte der Virologie