History of HIV/AIDS: Difference between revisions

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Similar researches were undertaken with [[SIV]] strains collected from several wild [[sooty mangabey]] (''Cercocebus atys atys'') (SIVsmm) communities of the West African nations of [[Sierra Leone]], [[Liberia]], and [[Ivory Coast]]. The resulting phylogenetic analyses show that the viruses most closely related to the two strains of HIV-2 which spread considerably in humans (HIV-2 groups A and B) are the SIVsmm found in the sooty mangabeys of the Tai forest, in western [[Ivory Coast]].<ref name=Santiago2005/>
Similar researches were undertaken with [[SIV]] strains collected from several wild [[sooty mangabey]] (''Cercocebus atys atys'') (SIVsmm) communities of the West African nations of [[Sierra Leone]], [[Liberia]], and [[Ivory Coast]]. The resulting phylogenetic analyses show that the viruses most closely related to the two strains of HIV-2 which spread considerably in humans (HIV-2 groups A and B) are the SIVsmm found in the sooty mangabeys of the Tai forest, in western [[Ivory Coast]].<ref name=Santiago2005/>


There are six known additional HIV-2 groups, each having been found in just one person. They all seem to derive from independent transmissions from [[sooty mangabey]]s to humans. Groups C and D have been found in two people from [[Liberia]], groups E and F have been discovered in two people from [[Sierra Leone]], and groups G and H have been detected in two people from the [[Ivory Coast]]. These HIV-2 strains are probably dead end infections, and each of them is most closely related to SIVsmm strains from [[sooty mangabeys]] living in the same country where the human infection was found.<ref name=Santiago2005/><ref name=Hooper2000/>
There are six known additional HIV-2 groups, each having been found in just one person. They all seem to derive from independent transmissions from [[sooty mangabey]]s to humans. Groups C and D have been found in two people from [[Liberia]], groups E and F have been discovered in two people from [[Sierra Leone]], and groups G and H have been detected in two people from the [[Ivory Coast]]. These HIV-2 strains are probably dead end infections, and each of them is most closely related to SIVsmm strains from [[sooty mangabeys]] living in the same country where the human infection was found.<ref name=Marx2001>{{cite journal |author=Marx PA, Alcabes PG, Drucker E |title=Serial human passage of simian immunodeficiency virus by unsterile injections and the emergence of epidemic human immunodeficiency virus in Africa |journal=Philos. Trans. R. Soc. Lond., B, Biol. Sci. |volume=356 |issue=1410 |pages=911–20 |year=2001 |month=June |pmid=11405938 |pmc=1088484 |doi=10.1098/rstb.2001.0867 |url=http://rstb.royalsocietypublishing.org/content/356/1410/911.abstract}}</ref><ref name=Santiago2005/><ref name=Hooper2000/>


Molecular datation studies suggest that both the epidemic groups (A and B) started to spread among humans between 1924 and 1959.<ref name=Lemey2003>{{cite journal |author=Lemey P. Pybus OG, Wang B, Saksena NK, Salemi M, Vandamme AM |title=Tracing the origin and history of the HIV-2 epidemic |journal=Proc Natl Acad Sci U S A |volume=100 |issue=11 |pages=6588–92 |year=2003 |pmid=164491 |doi=10.1073/pnas.0936469100 |url=http://www.pnas.org/content/100/11/6588.full}}</ref>
Molecular datation studies suggest that both the epidemic groups (A and B) started to spread among humans between 1924 and 1959.<ref name=Lemey2003>{{cite journal |author=Lemey P, Pybus OG, Wang B, Saksena NK, Salemi M, Vandamme AM |title=Tracing the origin and history of the HIV-2 epidemic |journal=Proc Natl Acad Sci U S A |volume=100 |issue=11 |pages=6588–92 |year=2003 |pmid=164491 |doi=10.1073/pnas.0936469100 |url=http://www.pnas.org/content/100/11/6588.full}}</ref><ref name=Wertheim>{{cite journal |author=Wertheim JO, Worobey M |title=Dating the Age of the SIV Lineages That Gave Rise to HIV-1 and HIV-2 |journal=PLoS Comput Biol |volume=5 |issue=5 |pages=e1000377 |year=2009 |pmid=2669881 |doi=10.1371/journal.pcbi.1000377 |url=http://www.ploscompbiol.org/article/info%3Adoi%2F10.1371%2Fjournal.pcbi.1000377}}</ref>


=== Bushmeat practice as the most plausible cause ===
=== Bushmeat practice as the most plausible cause ===

Revision as of 06:20, 24 November 2010

False-color scanning electron micrograph of HIV-1 budding from cultured lymphocyte.

HIV, the virus that causes AIDS, originated in non-human primates in Sub-Saharan Africa and was transferred to humans during the late 19th or early 20th century.

Two types of HIV infect humans: HIV-1 and HIV-2. HIV-1 is more virulent, is more easily transmitted and is the cause of the vast majority of HIV infections globally.[1] The pandemic strain of HIV-1 is closely related to a virus found in the chimpanzees of West Central Africa (subspecies Pan troglodytes troglodytes), and molecular phylogenetics indicates that HIV-1 started to spread in humans sometime between 1884 and 1924 in equatorial Africa.[2] HIV-2 is less transmittable and is largely confined to West Africa, along with its closest relative, a virus of the sooty mangabey (Cercocebus atys atys), an Old World monkey inhabiting southern Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, and western Ivory Coast.[1][3]

Transmission from non-human to human populations

Most HIV researchers agree that HIV evolved at some point from the closely related Simian immunodeficiency virus (SIV), and that SIV or HIV (post mutation) was transferred from non-human primates to humans in the recent past (as a type of zoonosis). Research in this area is conducted using molecular phylogenetics, comparing viral genomic sequences to determine relatedness.

HIV-1 transmission from chimpanzees and gorillas to humans

Scientists generally accept that the known strains (or groups) of HIV-1 are most closely related to the simian immunodeficiency viruses endemic in wild ape populations of West Central African forests. Particularly, each of the known HIV-1 strains is either closely related to the SIV that infects the chimpanzee subspecies Pan troglodytes troglodytes (SIVcpz), or to the SIV that infects Western lowland gorillas (Gorilla gorilla gorilla), called SIVgor.[4][5][6].[7][8][9] The pandemic HIV-1 strain (group M or Main) and a very rare strain only found in a few Cameroonian people (group N) are clearly derived from SIVcpz strains endemic in Pan troglodytes troglodytes chimpanzee populations living in Cameroon.[4] Another very rare HIV-1 strain (group P) is clearly derived from SIVgor strains of the same country.[7] Finally, the primate ancestor of HIV-1 group O, a strain infecting tens of thousands of people mostly from Cameroon but also from neighboring countries, is still uncertain, but there is evidence that it is either SIVcpz or SIVgor.[6] The pandemic HIV-1 group M is most closely related to the SIVcpz collected from the southeastern rain forests of Cameroon (modern East Province) near the Sangha River.[4] Thus, this region is presumably where the virus first was transmitted from chimpanzees to humans. However, reviews of the epidemiological evidence of early HIV-1 infection in stored blood samples, and of old cases of AIDS in Central Africa have led many scientists to believe that HIV-1 group M early human epicenter was probably not in Cameroon, but rather farther south in the Democratic Republic of the Congo, more probably in its capital city, Kinshasa.[4][10][11][12]

Using HIV-1 sequences preserved in human biological samples along with estimates of viral mutation rates, scientists calculate that the jump from chimpanzee to human probably happened during the late 19th or early 20th century, a time of rapid urbanisation and colonisation in equatorial Africa. Exactly when the zoonosis occurred is not known. Some estimates suggest that HIV-1 (group M) entered the human population in the early 20th century, probably between 1915 and 1941.[13][14][15] A study published in 2008, analyzing viral sequences recovered from a recently-discovered biopsy made in Kinshasa, in 1960, along with previously-known sequences, suggested a common ancestor between 1884 and 1924.[2][16]

Genetic recombination had earlier been thought to "seriously confound" such phylogenetic analysis, but later "work has suggested that recombination is not likely to systematically bias [results]", although recombination is "expected to increase variance".[2] The results of phlyogenetics study supported the later work and indicated that HIV evolves "fairly reliably".[2][17]

HIV-2 transmission from sooty mangabeys to humans

Similar researches were undertaken with SIV strains collected from several wild sooty mangabey (Cercocebus atys atys) (SIVsmm) communities of the West African nations of Sierra Leone, Liberia, and Ivory Coast. The resulting phylogenetic analyses show that the viruses most closely related to the two strains of HIV-2 which spread considerably in humans (HIV-2 groups A and B) are the SIVsmm found in the sooty mangabeys of the Tai forest, in western Ivory Coast.[3]

There are six known additional HIV-2 groups, each having been found in just one person. They all seem to derive from independent transmissions from sooty mangabeys to humans. Groups C and D have been found in two people from Liberia, groups E and F have been discovered in two people from Sierra Leone, and groups G and H have been detected in two people from the Ivory Coast. These HIV-2 strains are probably dead end infections, and each of them is most closely related to SIVsmm strains from sooty mangabeys living in the same country where the human infection was found.[18][3][12]

Molecular datation studies suggest that both the epidemic groups (A and B) started to spread among humans between 1924 and 1959.[19][20]

Bushmeat practice as the most plausible cause

According to the 'Hunter Theory', the "simplest and most plausible explanation for the cross-species transmission"[8] of SIV or HIV (post mutation), the virus was transmitted from a chimpanzee to a human when a bushmeat hunter was bitten or cut while hunting or butchering an animal. The resulting exposure of the hunter to blood or other bodily fluids of the chimpanzee could have resulted in SIV infection.[21] How the SIV virus would have transformed into HIV after possible infection of the hunter from the chimpanzee is still a matter of debate, although natural selection would favor any virions capable of adjusting so that they could live, infect and reproduce in the T cells of a human host.

Method of spread

Zoonosis (transfer of a pathogen from non-human animals to humans) and subsequent spread of the pathogen between humans, requires the following conditions:

  1. a human population;
  2. a nearby population of a host animal;
  3. an infectious pathogen in the host animal that can spread from animal to human;
  4. interaction between the species to transmit enough of the pathogen to humans to establish a human foothold, which could have taken millions of individual exposures;
  5. ability of the pathogen to spread from human to human (perhaps acquired by mutation);
  6. some method allowing the pathogen to disperse widely, preventing the infection from "burning out" by either killing off its human hosts or provoking immunity in a local population of humans.

Conditions that facilitated the establishment and spread of infection in human populations may have included the reuse of needles for injectable antibiotics, antimalarial drugs and vaccines.[22] Another cause may have been the increase in bushmeat hunting and immune suppression as a result of the harsh conditions, forced labor and displacement associated with colonialism, particularly in French Equatorial Africa.[23] A third theory blames rapid urbanisation that brought infected people into close contact with others, along with commerce, and urban red light districts that provided opportunities for further geographical spread.[16][22]

SIV in non-human primates tends to cause a non-fatal disease. Comparison of the gene sequence of SIV with HIV should therefore give us information about the factors necessary to cause disease in humans. The factors that determine the virulence of HIV as compared to most SIVs are only now being elucidated. Non-human SIVs contain a nef gene that down-regulates CD3, CD4, and MHC class I expression; most non-human SIVs therefore do not induce immunodeficiency; the HIV nef gene however has lost its ability to down-regulate CD3, which results in the immune activation and apoptosis that is characteristic of chronic HIV infection.[24]

History of known cases and spread

1955–1957: David Carr

The oldest documented possible case of the then-unknown syndrome was thought to have been detected in 1959, when David Carr, a 25-year-old British printer who had served in the Royal Navy between 1955 and 1957 (but apparently not in Africa) sought help at the Royal Infirmary of Manchester, England. He reported to have been suffering from puzzling symptoms, among them purplish skin lesions, for nearly two years. His condition had taken a turn for worse during Christmas 1958, when he started suffering from shortness of breath, extreme fatigue, rapid weight loss, night sweats and high fever. The doctors thought he might be suffering from tuberculosis and, even though they found no evidence of bacterial infection, they treated him for tuberculosis just to be safe, to no avail. The printer continued to weaken and he died shortly after in August 1959. His autopsy revealed evidence of two unusual infections, cytomegalovirus and Pneumocystis carinii pneumonia (PCP, later, when redetermined as P. jirovecii, renamed Pneumocystis pneumonia), very rare at the time but now commonly associated with AIDS patients. His case had puzzled his doctors, who preserved tissue samples from him and for years retained some interest in solving the mystery.

Sir Robert Platt, then president of the Royal College of Physicians, wrote in the printer's hospital chart that he wondered "If we are in for a new wave of virus disease now that the bacterial illnesses are so nearly conquered". It was only 31 years later, after the AIDS pandemic had become well-known and widespread, that they decided to perform HIV-tests on the preserved tissues of the printer, which initially turned out a positive result. While the case was reported in the July 7, 1990 issue of the British medical journal The Lancet, their claim was retracted in a letter in the January 20, 1996 issue, in which they reported that the tissue sample had become contaminated in the laboratory.[25][26][27]

1959: Congolese man

One of the earliest documented HIV-1 infections was discovered in a preserved blood sample taken in 1959 from a man from Leopoldville, Belgian Congo (now Kinshasa, Democratic Republic of the Congo).[28] However, it is unknown whether this anonymous person ever developed AIDS and died of its complications.[28]

1960: Congolese woman

A second early documented HIV-1 infection was discovered in a preserved lymph node biopsy sample taken in 1960 from a woman from Leopoldville, Belgian Congo.[2]

1969: Robert R.

Main article: Robert R.

In 1969, a 15-year-old African-American male known to medicine as Robert R. died at the St. Louis City Hospital from aggressive Kaposi's sarcoma. AIDS was suspected as early as 1984, and in 1987, researchers at Tulane University School of Medicine confirmed this, finding HIV-1 in his preserved blood and tissues. The doctors who worked on his case at the time suspected he was a prostitute, though the patient did not discuss his sexual history with them in detail.[29][30][31][32][33]

1969: Arvid Noe

Main article: Arvid Noe

In 1976, a Norwegian sailor, with the alias name Arvid Noe, his wife, and his nine-year-old daughter died of AIDS. The sailor had first presented symptoms in 1969, eight years after he first spent time in ports along the West African coastline. A gonorrhea infection during his first African voyage shows he was sexually active at this time. Tissue samples from the sailor and his wife were tested in 1988 and found to contain HIV-1 (Group O).[34][35][36]

Spread to the western hemisphere

HIV-1 strains are thought to have arrived in the United States from Haiti in the late 1960s or early 1970s.[37] HIV-1 is believed to have arrived in Haiti from central Africa, possibly through professional contacts with the Democratic Republic of the Congo.[38] The current consensus is that HIV was introduced to North America by a Haitian immigrant who contracted it while working in the Democratic Republic of the Congo in the late 1960s, or from another person who worked there during that time.[39]

Because of the long incubation period of HIV (up to a decade or longer) before symptoms of AIDS appear, and because of the initially low incidence, AIDS was not noticed at first. By the time the first reported cases of AIDS were found in large United States cities, the prevalence of HIV infection in some communities had passed 5%.[40] Worldwide, HIV infection has spread from urban to rural areas, and has appeared in regions such as China and India.

Canadian flight attendant theory

Main article: Gaëtan Dugas

A Canadian airline steward named Gaëtan Dugas was referred to as "Patient 0" in an early AIDS study by Dr. William Darrow of the Centers for Disease Control. Many people consider Dugas to be responsible for bringing HIV to North America. This is considered inaccurate, as HIV had spread long before Dugas began his career. This rumor may have started with Randy Shilts' 1987 book And the Band Played On (and the movie based on it, in which Dugas is referred to as AIDS' Patient Zero), but neither the book nor the movie state that he had been the first to bring the virus to North America. He was called "Patient Zero" because at least 40 of the 248 people known to be infected by AIDS in 1983 had had sex with him, or with someone who had sexual intercourse with him.

1981–2: From GRID to AIDS

The AIDS epidemic officially began on June 5, 1981, when the U.S. Centers for Disease Control and Prevention in its Morbidity and Mortality Weekly Report newsletter reported unusual clusters of Pneumocystis pneumonia (PCP) caused by a form of Pneumocystis carinii (now recognized as a distinct species Pneumocystis jirovecii) in five homosexual men in Los Angeles.[41]

Over the next 18 months, more PCP clusters were discovered among otherwise healthy men in cities throughout the country, along with other opportunistic diseases (such as Kaposi's sarcoma[42] and persistent, generalized lymphadenopathy[43]), common in immunosuppressed patients.

In June 1982, a report of a group of cases amongst gay men in Southern California suggested that a sexually transmitted infectious agent might be the etiological agent,[44] and the syndrome was initially termed "GRID", or gay-related immune deficiency.[45]

Health authorities soon realized that nearly half of the people identified with the syndrome were not homosexual men. The same opportunistic infections were also reported among hemophiliacs,[46] heterosexual intravenous drug users, and Haitian immigrants.[47]

By August 1982, the disease was being referred to by its new CDC-coined name: Acquired Immune Deficiency Syndrome (AIDS).[48]

Identification of the virus

May 1983: LAV

In May 1983, doctors from Dr. Luc Montagnier's team at the Pasteur Institute in France reported that they had isolated a new retrovirus from lymphoid ganglions that they believed was the cause of AIDS.[49] The virus was later named lymphadenopathy-associated virus (LAV) and a sample was sent to the U.S. Centers for Disease Control, which was later passed to the National Cancer Institute (NCI).[50][49]

May 1984: HTLV-III

In May 1984 a team led by Robert Gallo of the United States confirmed the discovery of the virus, but they renamed it human T lymphotropic virus type III (HTLV-III).[51]

Jan 1985: both found to be the same

In January 1985 a number of more detailed reports were published concerning LAV and HTLV-III, and by March it was clear that the viruses were the same, were from the same source, and were the etiological agent of AIDS.[52][53]

May 1986: the name HIV

In May 1986, the International Committee on Taxonomy of Viruses ruled that both names should be dropped and a new name, HIV (Human Immunodeficiency Virus), be used.[54]

Genetic studies

According to a 2008 Proceedings of the National Academy of Sciences study, a team lead by Robert Shafer at Stanford University School of Medicine has discovered that the Gray Mouse Lemur has an endogenous lentivirus (the genus to which HIV belongs) in its genetic makeup. This suggests that lentiviruses have existed for at least 14 million years, much longer than the currently known existence of HIV. In addition, the time frame falls into place when Madagascar was still yet connected to what is now the African continent; the said lemurs later developed immunity to the virus strain and survived an era when the lentivirus was widespread among other mammalia. The study is being hailed as crucial, because it fills the blanks in the origin of the virus, as well as in its evolution, and may be important in the development of new antiviral drugs.[55][56]

In 2010, researchers reported that SIV had infected monkeys in Bioko for at least 32,000 years. Previously it was thought that SIV infection in monkeys had happened over the past few hundred years.[57] Scientists estimated that it would take a similar amount of time before humans adapted naturally to HIV infection in the way monkeys in Africa have adapted to SIV and not suffer any harm from the infection.[58]

Alternative hypotheses

Main article: AIDS origins opposed to scientific consensus

Several alternative hypotheses for the origin of AIDS have been proposed. AIDS denialism argues that HIV or AIDS does not exist or that AIDS is not caused by HIV; some of its proponents believe that AIDS is caused by lifestyle, including sexuality or drug use. Some conspiracy theories allege that HIV was created in a bioweapons laboratory, perhaps as an agent of genocide or an accident. These hypotheses have been rejected by scientific consensus.

See also

Notes

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  2. ^ a b c d e Worobey M, Gemmel M, Teuwen DE; et al. (2008). "Direct evidence of extensive diversity of HIV-1 in Kinshasa by 1960". Nature. 455 (7213): 661–4. doi:10.1038/nature07390. PMID 18833279. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  3. ^ a b c Santiago ML, Range F, Keele BF, Li Y, Bailes E; et al. (2005). "Simian Immunodeficiency Virus Infection in Free-Ranging Sooty Mangabeys (Cercocebus atys atys) from the Taï Forest, Côte d'Ivoire: Implications for the Origin of Epidemic Human Immunodeficiency Virus Type 2". J. Virol. 79 (19): 12515–27. doi:10.1128/JVI.79.19.12515-12527.2005. PMID 1211554. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  4. ^ a b c d Keele, B. F., van Heuverswyn, F., Li, Y. Y., Bailes, E., Takehisa, J., Santiago, M. L., Bibollet-Ruche, F., Chen, Y., Wain, L. V., Liegois, F., Loul, S., Mpoudi Ngole, E., Bienvenue, Y., Delaporte, E., Brookfield, J. F. Y., Sharp, P. M., Shaw, G. M., Peeters, M., and Hahn, B. H. (2006). "Chimpanzee Reservoirs of Pandemic and Nonpandemic HIV-1". Science. Online 2006-05-25 (5786): 523. doi:10.1126/science.1126531. PMC 2442710. PMID 16728595.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  5. ^ "HIV's ancestry traced to wild chimps in Cameroon". USA Today. 2006-05-25. Retrieved 2010-05-20.
  6. ^ a b Van Heuverswyn F, Li Y, Neel C, Bailes E, Keele BF; et al. (2006). "Human immunodeficiency viruses: SIV infection in wild gorillas". Nature. 444 (7116): 164. doi:10.1038/444164a. PMID 17093443. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  7. ^ a b Plantier JC, Leoz M, Dickerson JE, De Oliveira F, Cordonnier F; et al. (2009). "A new human immunodeficiency virus derived from gorillas". Nature Med. 15 (8): 871–72. doi:10.1038/nm.2016. PMID 19648927. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  8. ^ a b Sharp PM, Bailes E, Chaudhuri RR, Rodenburg CM, Santiago MO, Hahn BH (2001). "The origins of acquired immune deficiency syndrome viruses: where and when?" (PDF). Philos Trans R Soc Lond B Biol Sci. 356 (1410): 867–76. doi:10.1098/rstb.2001.0863. PMC 1088480. PMID 11405934.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Takebe Y, Uenishi R, Li X (2008). "Global molecular epidemiology of HIV: understanding the genesis of AIDS pandemic". Adv Pharmacol. 56: 1–25. doi:10.1016/S1054-3589(07)56001-1. PMID 18086407.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ^ Gao, F., Bailes, E., Robertson, D. L., Chen, Y., Rodenburg, C. M., Michael, S. F., Cummins, L. B., Arthur, L. O., Peeters, M., Shaw, G. M., Sharp, P. M., and Hahn, B. H. (1999). "Origin of HIV-1 in the Chimpanzee Pan troglodytes troglodytes". Nature. 397 (6718): 436–441. doi:10.1038/17130. PMID 9989410.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ Sousa JDd, Müller V, Lemey P, Vandamme AM (2010). "High GUD Incidence in the Early 20th Century Created a Particularly Permissive Time Window for the Origin and Initial Spread of Epidemic HIV Strains". PLoS ONE. 5 (4): e9936. doi:10.1371/journal.pone.0009936. PMID 2848574.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  12. ^ a b Hooper, Edward (2000) The river : a journey to the source of HIV and AIDS Boston, MA : Little, Brown and Co ISBN 0-316-37261-7 9780316372619
  13. ^ Salemi M, Strimmer K, Hall WW, Duffy M, Delaporte E, Mboup S, Peeters M, Vandamme AM (2000). "Dating the common ancestor of SIVcpz and HIV-1 group M and the origin of HIV-1 subtypes using a new method to uncover clock-like molecular evolution". The FASEB journal. 15 (2): 276–78. doi:10.1096/fj.00-0449fje. PMID 11156935.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  14. ^ Korber B, Muldoon M, Theiler J; et al. (2000). "Timing the ancestor of the HIV-1 pandemic strains". Science. 288 (5472): 1789–96. doi:10.1126/science.288.5472.1789. PMID 10846155. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  15. ^ http://evolve.zoo.ox.ac.uk/papers/Lemey%20et%20al%20(2004)%20Genetics.pdf
  16. ^ a b "AIDS virus leapt the species barrier early last century: study" Breitbart, October 1, 2008. Accessed October 2, 2008.
  17. ^ Colonial clue to the rise of HIV. BBC News. Retrieved 20-1-2009.
  18. ^ Marx PA, Alcabes PG, Drucker E (2001). "Serial human passage of simian immunodeficiency virus by unsterile injections and the emergence of epidemic human immunodeficiency virus in Africa". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 356 (1410): 911–20. doi:10.1098/rstb.2001.0867. PMC 1088484. PMID 11405938. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  19. ^ Lemey P, Pybus OG, Wang B, Saksena NK, Salemi M, Vandamme AM (2003). "Tracing the origin and history of the HIV-2 epidemic". Proc Natl Acad Sci U S A. 100 (11): 6588–92. doi:10.1073/pnas.0936469100. PMID 164491.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  20. ^ Wertheim JO, Worobey M (2009). "Dating the Age of the SIV Lineages That Gave Rise to HIV-1 and HIV-2". PLoS Comput Biol. 5 (5): e1000377. doi:10.1371/journal.pcbi.1000377. PMID 2669881.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  21. ^ Annabel Kanabus & Sarah Allen. Updated by Bonita de Boer (2007). "The Origins of HIV & the First Cases of AIDS". AVERT (an international HIV and AIDS charity based in the UK). Retrieved 2007-02-28.
  22. ^ a b Donald G. McNeil, Jr. (September 16, 2010). "Precursor to H.I.V. Was in Monkeys for Millennia". New York Times. Retrieved 2010-09-17. Dr. Marx believes that the crucial event was the introduction into Africa of millions of inexpensive, mass-produced syringes in the 1950s. ... suspect that the growth of colonial cities is to blame. Before 1910, no Central African town had more than 10,000 people. But urban migration rose, increasing sexual contacts and leading to red-light districts. {{cite news}}: Cite has empty unknown parameter: |coauthors= (help)
  23. ^ Chitnis A, Rawls D, Moore J (2000). "Origin of HIV type 1 in colonial French Equatorial Africa?" (PDF). AIDS Res. Hum. Retroviruses. 16 (1): 5–8. doi:10.1089/088922200309548. PMID 10628811. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  24. ^ Schindler M, Münch J, Kutsch O; et al. (2006). "Nef-mediated suppression of T cell activation was lost in a lentiviral lineage that gave rise to HIV-1". Cell. 125 (6): 1055–67. doi:10.1016/j.cell.2006.04.033. PMID 16777597. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  25. ^ Corbitt G, Bailey AS, Williams G (1990). "HIV infection in Manchester, 1959". Lancet. 336 (8706): 51. doi:10.1016/0140-6736(90)91561-N. PMID 1973229. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  26. ^ Altman, Lawrence K. (1990-07-24). "THE DOCTOR'S WORLD; Puzzle of Sailor's Death Solved After 31 Years: The Answer Is AIDS". The New York Times. Retrieved 2010-05-20.
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