| Haplogroup G |
|---|
| Possible time of origin |
35,700 YBP[1] |
|---|
| Possible place of origin |
East Asia |
|---|
| Ancestor |
M12'G |
|---|
| Descendants |
G1, G2, G3, G4 |
|---|
| Defining mutations |
709, 4833, 5108[2] |
|---|
In human mitochondrial genetics, Haplogroup G is a human mitochondrial DNA (mtDNA) haplogroup.
Origin
Haplogroup G is a descendant of haplogroup M. Haplogroup G is divided into subclades G1, G2, G3, and G4.
Distribution
It is an East Asian haplogroup.[3] Today, haplogroup G is found at its highest frequency in indigenous populations of the lands surrounding the Sea of Okhotsk.[4][5] Haplogroup G is one of the most common mtDNA haplogroups among modern Ainu, Japanese, Mongol, and Tibetan people (as well as among people of the prehistoric Jōmon culture in Hokkaidō). It is also found at a lower frequency among many other populations of East Asia, Central Asia, Bangladesh, Sri Lanka and Nepal.[6][7][8][9] However, unlike other mitochondrial DNA haplogroups typical of populations of northeastern Asia, such as haplogroup A, haplogroup C, and haplogroup D, haplogroup G has not been found among indigenous peoples of the Americas.
Table of Frequencies of MtDNA Haplogroup G
| Population |
Frequency |
Count |
Source |
Subtypes |
|---|
| Itelmen | 0.681 | 47 | Starikovskaya 2005 | G1=32 |
| Koryak | 0.419 | 155 | Starikovskaya 2005 | G1=65 |
| Chuvantsi (Markovo, Chukotka) | 0.281 | 32 | Volodko 2008 | G1=9 |
| Negidal | 0.273 | 33 | Starikovskaya 2005 | G1=9 |
| Tharu (Chitwan, Nepal) | 0.233 | 133 | Fornarino 2009 | G2a=19, G(xG2a)=12 |
| Kazakh (Uzbekistan/Kyrgyzstan) | 0.200 | 20 | Comas 2004 | G2a=4 |
| Ainu | 0.196 | 51 | Satou 2009 | G1=8, G2=2 |
| Tibetan (Lhasa, Tibet) | 0.182 | 44 | Ji 2012 | G2a=3, G3b=3, G2(xG2a)=2 |
| Mongolian (Ulan Bator) | 0.170 | 47 | Jin 2009 | G2a=5, G(xG1a, G2, G3)=2, G3=1 |
| Korean (Arun Banner) | 0.167 | 48 | Kong 2003 | G2(xG2a)=3, G2a=3, G1a=1, G3=1 |
| Tibetan (Nyingchi, Tibet) | 0.167 | 24 | Ji 2012 | G=4 |
| Oirat Mongol (Xinjiang) | 0.163 | 49 | Yao 2004 | G2a=3, G2(xG2a)=3, G(xG1a, G2, G3)=2 |
| Tibetan (Shannan, Tibet) | 0.158 | 19 | Ji 2012 | G=3 |
| Yukaghir (Lower Kolyma-Indigirka) | 0.146 | 82 | Volodko 2008 | G1=12 |
| Kyrgyz (Talas) | 0.146 | 48 | Yao 2004 | G2a=7 |
| Tibetan (Shannan, Tibet) | 0.145 | 55 | Ji 2012 | G2a=4, G2(xG2a)=3, G3b=1 |
| Uyghur (Xinjiang) | 0.128 | 47 | Yao 2004 | G2a=5, G3=1 |
| Tharu (Morang, Nepal) | 0.125 | 40 | Fornarino 2009 | G2a=4, G(xG2a)=1 |
| Japanese (Gifu) | 0.116 | 1617 | Fuku 2007 | G=188 |
| Ulch | 0.115 | 87 | Starikovskaya 2005 | G1=9, G2=1 |
| Oroqen (Oroqen Autonomous Banner) | 0.114 | 44 | Kong 2003 | G(xG1a, G2, G3)=5 |
| Tibetan (Qinghai) | 0.107 | 56 | Wen 2004 | G(xG2, G3)=2, G2a=2, G2(xG2a)=1, G3=1 |
| Mongolian (Ulan Bator) | 0.106 | 47 | Derenko 2007 | G2a=4, G1=1 |
| Tuvan | 0.105 | 95 | Starikovskaya 2005 | G2=6, G3=4 |
| Huatou Yao (Fangcheng, Guangxi) | 0.105 | 19 | Wen 2005 | G2=2 |
| Japanese | 0.104 | 211 | Maruyama 2003 | G4a=12, G2a=6, G4b=2, G2(xG2a)=1, G(xG2, G4a, G4b)=1 |
| Tibetan (Chamdo, Tibet) | 0.103 | 29 | Ji 2012 | G3b=2, G2a=1 |
| Tibetan (Shigatse, Tibet) | 0.103 | 29 | Ji 2012 | G2a=2, G2(xG2a)=1 |
| Korean (South Korea) | 0.103 | 185 | Jin 2009 | G2(xG2a)=7, G2a=6, G3=4, G1a=1, G(xG1a, G2, G3)=1 |
| Japanese (Tokyo) | 0.102 | 118 | Zheng 2011 | G=12 |
| Khamnigan (Buryatia) | 0.101 | 99 | Derenko 2007 | G2a=9, G3=1 |
| Han (Beijing) | 0.100 | 40 | Jin 2009 | G2a=2, G2(xG2a)=1, G(xG1a, G2, G3)=1 |
| Manchurian | 0.100 | 40 | Jin 2009 | G1a=3, G2a=1 |
| Tu Yao (Hezhou, Guangxi) | 0.098 | 41 | Wen 2005 | G2=4 |
| Japanese (Tōkai) | 0.096 | 282 | Umetsu 2005 | G1a=13, G(xG1a, G1b)=12, G1b=2 |
| Even (Eveno-Bytantaysky & Momsky) | 0.095 | 105 | Fedorova 2013 | G1b=9, G2a(xG2a5)=1 |
| Barghut (Hulunbuir) | 0.094 | 149 | Derenko 2012 | G2=13, G3=1 |
| Chukchi | 0.091 | 66 | Starikovskaya 2005 | G1=6 |
| Xiban Yao (Fangcheng, Guangxi) | 0.091 | 11 | Wen 2005 | G2=1 |
| Daur (Evenk Autonomous Banner) | 0.089 | 45 | Kong 2003 | G1a=2, G2a=2 |
| Hui (Xinjiang) | 0.089 | 45 | Yao 2004 | G2a=2, G1a=1, G(xG1a, G2, G3)=1 |
| Japanese (Hokkaidō) | 0.088 | 217 | Asari 2007 | G1a=11, G(xG1a, G1b)=7, G1b=1 |
| Evenk (New Barag Left Banner) | 0.085 | 47 | Kong 2003 | G(xG1a, G2, G3)=4 |
| Pumi (Ninglang, Yunnan) | 0.083 | 36 | Wen 2004 | G(xG2, G3)=2, G3=1 |
| Kalmyk (Kalmykia) | 0.082 | 110 | Derenko 2007 | G2a=7, G1=1, G(xG1, G2a, G3)=1 |
| Buryat | 0.080 | 25 | Starikovskaya 2005 | G2=1, G3=1 |
| Buryat | 0.079 | 126 | Kong 2003 | G2a=8, G2(xG2a)=2 |
| Korean (South Korea) | 0.079 | 203 | Umetsu 2005 | G1a=9, G(xG1a, G1b)=7 |
| Bai (Dali, Yunnan) | 0.074 | 68 | Wen 2004 | G2(xG2a)=5 |
| Dargin (Dagestan) | 0.071 | 28 | Marchani 2008 | G=2 |
| Uzbek (Xinjiang) | 0.069 | 58 | Yao 2004 | G2a=2, G3=1, G(xG1a, G2, G3)=1 |
| Chinese (Shenyang, Liaoning) | 0.069 | 160 | Umetsu 2005 | G(xG1a, G1b)=8, G1a=3 |
| Korean (South Korea) | 0.068 | 103 | Derenko 2007 | G2a=3, G1=2, G3=2 |
| Korean (Seoul National University Hospital) | 0.068 | 633 | Fuku 2007 | G=43 |
| Yakut (northern Yakutia) | 0.068 | 148 | Fedorova 2013 | G2a5=6, G2a(xG2a5)=2, G1b=2 |
| Chukchi (Anadyr) | 0.067 | 15 | Derenko 2007 | G1=1 |
| Naxi (Lijiang, Yunnan) | 0.067 | 45 | Wen 2004 | G(xG2, G3)=3 |
| Tujia (Yongshun, Hunan) | 0.067 | 30 | Wen 2004 | G(xG2, G3)=1, G2(xG2a)=1 |
| Tuvinian | 0.067 | 105 | Derenko 2007 | G2a=4, G1=2, G3=1 |
| Gelao (Daozhen County, Guizhou) | 0.065 | 31 | Li 2007 | G2a=2 |
| Mien (Shangsi, Guangxi) | 0.063 | 32 | Wen 2005 | G2=2 |
| Korean (South Korea) | 0.061 | 261 | Kim 2008 | G(xG2)=11, G2=5 |
| Mansi | 0.061 | 98 | Starikovskaya 2005 | G2=6 |
| Japanese (Miyazaki) | 0.060 | 100 | Uchiyama 2007 | G4a=2, G1a=1, G1b=1, G2a1(xG2a1a)=1, G2a1a=1 |
| Han (Beijing Normal University) | 0.058 | 121 | Zheng 2011 | G=7 |
| Tibetan (Zhongdian, Yunnan) | 0.057 | 35 | Wen 2004 | G3=2 |
| Kazakh (Xinjiang) | 0.057 | 53 | Yao 2004 | G1a=1, G2a=1, G(xG1a, G2, G3)=1 |
| Altai Kizhi | 0.056 | 90 | Derenko 2007 | G1=4, G2a=1 |
| Tibetan (Nyingchi, Tibet) | 0.056 | 54 | Ji 2012 | G2(xG2a)=1, G2a=1, G3b=1 |
| Han (Denver, Colorado) | 0.055 | 73 | Zheng 2011 | G=4 |
| Kazakh (Kazakhstan) | 0.055 | 55 | Yao 2004 | G2a=3 |
| Japanese (Tōhoku) | 0.054 | 336 | Umetsu 2005 | G1a=13, G(xG1a, G1b)=5 |
| Nivkh (northern Sakhalin) | 0.054 | 56 | Starikovskaya 2005 | G1=3 |
| Karakalpak (Uzbekistan/Kyrgyzstan) | 0.050 | 20 | Comas 2004 | G2a=1 |
| Kim Mun (Malipo, Yunnan) | 0.050 | 40 | Wen 2005 | G2=2 |
| Tajik (Uzbekistan/Kyrgyzstan) | 0.050 | 20 | Comas 2004 | G2a=1 |
| Uzbek (Uzbekistan/Kyrgyzstan) | 0.050 | 40 | Comas 2004 | G2a=2 |
| Yi (Shuangbai, Yunnan) | 0.050 | 40 | Wen 2004 | G(xG2, G3)=1, G2(xG2a)=1 |
| Orok (Sakhalin) | 0.049 | 61 | Bermisheva 2005 | G=3 |
| Gelao (Daozhen County, Guizhou) | 0.049 | 102 | Liu 2011 | G(xG2, G3)=4, G2a1=1 |
| Yakut (vicinity of Yakutsk) | 0.049 | 164 | Fedorova 2013 | G2a(xG2a5)=6, G2a5=2 |
| Hmong (Jishou, Hunan) | 0.049 | 103 | Wen 2005 | G3=2, G(xG2, G3)=2, G2=1 |
| Vietnamese | 0.048 | 42 | Jin 2009 | G1a=1, G3=1 |
| Japanese (northern Kyūshū) | 0.047 | 256 | Umetsu 2005 | G(xG1a, G1b)=9, G1a=3 |
| Tujia (western Hunan) | 0.047 | 64 | Wen 2004 | G(xG2, G3)=1, G2(xG2a)=1, G3=1 |
| Tajik (Tajikistan) | 0.045 | 44 | Derenko 2007 | G2a=1, G3=1 |
| Yukaghir (Verkhnekolymsky & Nizhnekolymsky) | 0.045 | 22 | Fedorova 2013 | G1b=1 |
| Hazara (North West Frontier Province & Balochistan) | 0.043 | 23 | Quintana-Murci 2004 | G=1 |
| Mongol (New Barag Left Banner) | 0.042 | 48 | Kong 2003 | G2(xG2a)=2 |
| Evenk (Krasnoyarsk) | 0.041 | 73 | Derenko 2007 | G2a=2, G1=1 |
| Aini (Xishuangbanna, Yunnan) | 0.040 | 50 | Wen 2004 | G2a=2 |
| Korean (northern China) | 0.039 | 51 | Jin 2009 | G2a=1, G2(xG2a)=1 |
| Kumik (Dagestan) | 0.038 | 26 | Marchani 2008 | G=1 |
| Lanten Yao (Tianlin, Guangxi) | 0.038 | 26 | Wen 2005 | G2=1 |
| Yakut (Vilyuy River basin) | 0.036 | 111 | Fedorova 2013 | G2a(xG2a5)=2, G2a5=1, G1b=1 |
| Dong (Tianzhu County, Guizhou) | 0.036 | 28 | Li 2007 | G(xG1a, G2)=1 |
| Cun (Hainan) | 0.033 | 30 | Peng 2011 | G=1 |
| Nu (Gongshan, Yunnan) | 0.033 | 30 | Wen 2004 | G(xG2, G3)=1 |
| Lingao (Hainan) | 0.032 | 31 | Peng 2011 | G=1 |
| Yi (Luxi, Yunnan) | 0.032 | 31 | Wen 2004 | G(xG2, G3)=1 |
| Pan Yao (Tianlin, Guangxi) | 0.031 | 32 | Wen 2005 | G(xG2, G3)=1 |
| Nogai (Dagestan) | 0.030 | 33 | Marchani 2008 | G=1 |
| Han (Southwest China; pool of 44 Sichuan, 34 Chongqing, 33 Yunnan, & 26 Guizhou) | 0.029 | 137 | Ji 2012 | G1=3, G2=1 |
| Han (southern California) | 0.028 | 390 | Ji 2012 | G=11 |
| Telenghit (Altai Republic) | 0.028 | 71 | Derenko 2007 | G2a=2 |
| Yakut (Yakutia) | 0.028 | 36 | Derenko 2007 | G2a=1 |
| Hmong (Wenshan, Yunnan) | 0.026 | 39 | Wen 2005 | G(xG2, G3)=1 |
| Yakut | 0.026 | 117 | Kong 2003 | G2a=2, G1a=1 |
| Evenk (Ust-Maysky, Oleneksky, and Zhigansky) | 0.024 | 125 | Fedorova 2013 | G1b=2, G2a(xG2a5)=1 |
| Uzbek (Surkhandarya, Uzbekistan) | 0.024 | 42 | Quintana-Murci 2004 | G=1 |
| Evenk (Buryatia) | 0.022 | 45 | Derenko 2007 | G3=1 |
| Taiwanese (Taipei, Taiwan) | 0.022 | 91 | Umetsu 2005 | G(xG1a, G1b)=2 |
| Han (Taiwan) | 0.021 | 1117 | Ji 2012 | G=24 |
| Han (Xinjiang) | 0.021 | 47 | Yao 2004 | G2a=1 |
| Kyrgyz (Sary-Tash) | 0.021 | 47 | Yao 2004 | G2a=1 |
| Hindu (New Delhi) | 0.021 | 48 | Fornarino 2009 | G(xG2a)=1 |
| Kazakh (Kosh-Agachsky, Altai Republic) | 0.020 | 98 | Derenko 2012 | G2=1, G3=1 |
| Turkish (Anatolia, Turkey) | 0.020 | 50 | Quintana-Murci 2004 | G=1 |
| Khanty | 0.019 | 106 | Pimenoff 2008 | G2=2 |
| Uyghur (Kazakhstan) | 0.018 | 55 | Yao 2004 | G2(xG2a)=1 |
| Khakassian (Khakassia) | 0.018 | 57 | Derenko 2007 | G3=1 |
| Mansi | 0.016 | 63 | Pimenoff 2008 | G2=1 |
| Okinawa | 0.015 | 326 | Umetsu 2005 | G(xG1a, G1b)=3, G1a=2 |
| Persian (eastern Iran) | 0.012 | 82 | Derenko 2007 | G2a=1 |
| Pakistani (Karachi, Pakistan) | 0.010 | 100 | Quintana-Murci 2004 | G=1 |
| Li (Hainan) | 0.009 | 346 | Peng 2011 | G=3 |
| Dolgan (Anabarsky, Volochanka, Ust-Avam, and Dudinka) | 0.006 | 154 | Fedorova 2013 | G1b=1 |
| Cham (Bình Thuận, Vietnam) | 0.006 | 168 | Peng 2010 | G=1 |
| Taiwan aborigines | 0.002 | 640 | Peng 2011 | G=1 |
| Dingban Yao (Mengla, Yunnan) | 0.000 | 10 | Wen 2005 | - |
| Yukaghir (Upper Kolyma) | 0.000 | 18 | Volodko 2008 | - |
| Filipino (Palawan) | 0.000 | 20 | Scholes 2011 | - |
| Yi (Hezhang County, Guizhou) | 0.000 | 20 | Li 2007 | - |
| Hindu (Chitwan, Nepal) | 0.000 | 24 | Fornarino 2009 | - |
| Guoshan Yao (Jianghua, Hunan) | 0.000 | 24 | Wen 2005 | - |
| Bunu (Dahua & Tianlin, Guangxi) | 0.000 | 25 | Wen 2005 | - |
| Kurd (northwestern Iran) | 0.000 | 25 | Derenko 2007 | - |
| Iu Mien (Mengla, Yunnan) | 0.000 | 27 | Wen 2005 | - |
| Andhra Pradesh (tribal) | 0.000 | 29 | Fornarino 2009 | - |
| Tujia (Yanhe County, Guizhou) | 0.000 | 29 | Li 2007 | - |
| Batak (Palawan) | 0.000 | 31 | Scholes 2011 | - |
| Wuzhou Yao (Fuchuan, Guangxi) | 0.000 | 31 | Wen 2005 | - |
| Bapai Yao (Liannan, Guangdong) | 0.000 | 35 | Wen 2005 | - |
| Tibetan (Nagchu, Tibet) | 0.000 | 35 | Ji 2012 | - |
| Aleut (Commander Islands) | 0.000 | 36 | Volodko 2008 | - |
| Eskimo (Sireniki) | 0.000 | 37 | Volodko 2008 | - |
| Eskimo (Naukan) | 0.000 | 39 | Volodko 2008 | - |
| Nganasan | 0.000 | 39 | Volodko 2008 | - |
| Thai | 0.000 | 40 | Jin 2009 | - |
| Lowland Yao (Fuchuan, Guangxi) | 0.000 | 42 | Wen 2005 | - |
| Eskimo (Chaplin) | 0.000 | 50 | Volodko 2008 | - |
| Teleut (Kemerovo) | 0.000 | 53 | Derenko 2007 | - |
| Han (Hunan & Fujian) | 0.000 | 55 | Zheng 2011 | - |
| Saami (Finland) | 0.000 | 69 | Tambets 2004 | - |
| Shor (Kemerovo) | 0.000 | 82 | Derenko 2007 | - |
| Eskimo (Canada) | 0.000 | 96 | Volodko 2008 | - |
| Saami (Sweden) | 0.000 | 98 | Tambets 2004 | - |
| Aleut (Aleutian Islands) | 0.000 | 163 | Volodko 2008 | - |
| Saami (Norway) | 0.000 | 278 | Tambets 2004 | - |
| Eskimo (Greenland) | 0.000 | 385 | Volodko 2008 | - |
|
Subclades
Subclade G2 is the most widely distributed, being found with low frequency in many populations all the way from western Siberia (Mansi, Khanty) to Japan (Japanese, Ainu) and from Iran (Persian) to South Central China (Hmong and Tujia in Hunan and Mien in Guangxi). G2 (and especially its subclade G2a) is notably frequent among many Mongolic- or Turkic-speaking populations of northern East Asia and Central Asia. G2a also has been found with high frequency in some samples of Tharus from southern Nepal.[10][11]
Subclade G1 is almost completely responsible for the high frequency of haplogroup G in populations located around the Sea of Okhotsk (Itelmen, Koryak, Negidal, Ulch, Ainu, Chukchi, Nivkh, etc.). G1 in Luoravetlans (Koryak & Chukchi) is essentially G1b, and this subclade is also found with generally low frequency in populations of Yakutia to the west (Evens, Yukaghirs, Evenks, Yakuts, Dolgans) as well as in Japan.[12] G1a has been found in samples from China (Daur, Hui, Kazakh, Korean, Manchu, and a sample of the general population of the city of Shenyang), Japan, Korea, Vietnam, and Siberia (Yakut). G1c has been found in China and Korea.
Subclade G3 is relatively rare. It has been found mainly among Koreans,[13] Tibetans, and presently Turkic- or Mongolic-speaking populations in southern Siberia and vicinity, and occasionally among Evenks in Buryatia, Japanese, Pumi, Tajiks, Hmong and Tujia in western Hunan, and Vietnamese.
Subclade G4 has been found in Japan.
Tree
This phylogenetic tree of haplogroup G subclades is based on the paper by Mannis van Oven and Manfred Kayser Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation[2] and subsequent published research.
See also
References
- ↑ Soares, Pedro; Luca Ermini; Noel Thomson; Maru Mormina; Teresa Rito; Arne Röhl; Antonio Salas; Stephen Oppenheimer; Vincent Macaulay; Martin B. Richards (4 June 2009). "Supplemental Data Correcting for Purifying Selection: An Improved Human Mitochondrial Molecular Clock". The American Journal of Human Genetics. 84 (6): 82–93. doi:10.1016/j.ajhg.2009.05.001. PMC 2694979
. PMID 19500773. Retrieved 2009-08-13. - 1 2 van Oven, Mannis; Manfred Kayser (13 Oct 2008). "Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation". Human Mutation. 30 (2): E386–E394. doi:10.1002/humu.20921. PMID 18853457. Retrieved 2009-05-20.
- ↑ Haplogroup G.
- ↑ mtDNA Haplogroup Testing
- ↑ Natalia V. Volodko, Elena B. Starikovskaya, Ilya O. Mazunin et al., "Mitochondrial Genome Diversity in Arctic Siberians, with Particular Reference to the Evolutionary History of Beringia and Pleistocenic Peopling of the Americas", The American Journal of Human Genetics 82, 1084–1100, May 2008. DOI 10.1016/j.ajhg.2008.03.019.
- ↑ Kazuo Umetsu, Masashi Tanaka, Isao Yuasa et al., "Multiplex amplified product-length polymorphism analysis of 36 mitochondrial single-nucleotide polymorphisms for haplogrouping of East Asian populations", Electrophoresis (2005), 26, 91–98. DOI 10.1002/elps.200406129
- ↑ Fuku, Noriyuki; Soo Park, Kyong; Yamada, Yoshiji; et al. (2007). "Mitochondrial Haplogroup N9a Confers Resistance against Type 2 Diabetes in Asians". Am. J. Hum. Genet. 80: 407–415. doi:10.1086/512202.
- ↑ Asari, M; et al. (2007). "Utility of haplogroup determination for forensic mtDNA analysis in the Japanese population". Leg Med. 9: 237–240. doi:10.1016/j.legalmed.2007.01.007.
- ↑ http://www.nature.com/jhg/journal/v59/n1/full/jhg2013112a.html
- ↑ Fornarino, Simona; Pala, Maria; Battaglia, Vincenza; et al. (2009). "Mitochondrial and Y-chromosome diversity of the Tharus (Nepal): a reservoir of genetic variation". BMC Evolutionary Biology. 2009 (9): 154. doi:10.1186/1471-2148-9-154. PMC 2720951. PMID 19573232.
- ↑ Pimenoff, Ville N; Comas, David; Palo, Jukka U; et al. (2008). "Northwest Siberian Khanty and Mansi in the junction of West and East Eurasian gene pools as revealed by uniparental markers". European Journal of Human Genetics. 16: 1254–1264. doi:10.1038/ejhg.2008.101. PMID 18506205.
- ↑ Uchiyama, Taketo; Hisazumi, Rinnosuke; Shimizu, Kenshi; et al. (2007). "Mitochondrial DNA Sequence Variation and Phylogenetic Analysis in Japanese Individuals from Miyazaki Prefecture". Japanese Journal of Forensic Science and Technology. 12 (1): 83–96. doi:10.3408/jafst.12.83.
- ↑ Jin, H-J; Tyler-Smith, C; Kim, W (2009). "The Peopling of Korea Revealed by Analyses of Mitochondrial DNA and Y-Chromosomal Markers". PLoS ONE. 4 (1): e4210. doi:10.1371/journal.pone.0004210. PMC 2615218. PMID 19148289.
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