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Haplogroup N is a human mitochondrial DNA (mtDNA) clade. A macrohaplogroup, its descendant lineages are distributed across many continents. Like its sibling macrohaplogroup M, macrohaplogroup N is a descendant of the haplogroup L3.

Haplogroup N
World map of prehistoric human migrations.jpg
Possible time of origin~71,000 YBP[1][1]
Possible place of originAsia[2][3][4][5][6] or East Africa[7][8]
AncestorL3
DescendantsN1'5, N2, N8, N9, N10, N11, N13, N14, N21, N22, A, I, O, R, S, X, Y, W
Defining mutations8701, 9540, 10398, 10873, 15301[9]

All mtDNA haplogroups found outside of Africa are descendants of either haplogroup N or its sibling haplogroup M. M and N are the signature maternal haplogroups that define the theory of the recent African origin of modern humans and subsequent early human migrations around the world. The global distribution of haplogroups N and M indicates that there was likely at least one major prehistoric migration of humans out of Africa, with both N and M later evolving outside the continent.[5]

Contents

OriginsEdit

 
Suggested routes of the initial settlement of Europe based on mtDNA haplogroups M and N, Metspalu et al. 2004. A major population split near the Persian Gulf would explain the ubiquity of Haplogroup N and the absence of Haplogroup M in West Eurasia

There is widespread agreement in the scientific community concerning the African ancestry of haplogroup L3 (haplogroup N's parent clade).[10] However, whether or not the mutations which define haplogroup N itself first occurred within Asia or Africa has been a subject for ongoing discussion and study.[10]

The out of Africa hypothesis has gained generalized consensus. However, many specific questions remain unsettled. To know whether the two M and N macrohaplogroups that colonized Eurasia were already present in Africa before the exit is puzzling.

Torroni et al. 2006 state that Haplogroups M, N and R occurred somewhere between East Africa and the Persian Gulf.[11]

Also related to the origins of haplogroup N is whether ancestral haplogroups M, N and R were part of the same migration out of Africa, or whether Haplogroup N left Africa via the Northern route through the Levant, and M left Africa via Horn of Africa. This theory was suggested because haplogroup N is by far the predominant haplogroup in Western Eurasia, and haplogroup M is absent in Western Eurasia, but is predominant in India and is common in regions East of India. However, the mitochondrial DNA variation in isolated "relict" populations in southeast Asia and among Indigenous Australians supports the view that there was only a single dispersal from Africa. Southeast Asian populations and Indigenous Australians all possess deep rooted clades of both haplogroups M and N.[12] The distribution of the earliest branches within haplogroups M, N, and R across Eurasia and Oceania therefore supports a three-founder-mtDNA scenario and a single migration route out of Africa.[13] These findings also highlight the importance of Indian subcontinent in the early genetic history of human settlement and expansion.[14]

Asian origin hypothesisEdit

The hypothesis of Asia as the place of origin of haplogroup N is supported by the following:

  1. Haplogroup N is found in all parts of the world but has low frequencies in Sub-Saharan Africa. According to a number of studies, the presence of Haplogroup N in Africa is most likely the result of back migration from Eurasia.[4]
  2. The oldest clades of macrohaplogroup N are found in Asia and Australia.
  3. It would be paradoxical that haplogroup N had traveled all the distance to Australia or New World yet failed to affect other populations within Africa besides North Africans and Horn Africans.
  4. N1 is the only sub-clade of haplogroup N that has been observed in Africa. However N1a is the only one in East Africa: this haplogroup is even younger and is not restricted to Africa, N1a has also been detected in Southern Siberia and was found in a 2,500-year-old Scytho-Siberian burial in the Altai region.[15]
  5. The mitochondrial DNA variation in isolated "relict" populations in southeast Asia supports the view that there was only a single dispersal from Africa.[12] The distribution of the earliest branches within haplogroups M, N, and R across Eurasia and Oceania provides additional evidence for a three-founder-mtDNA scenario and a single migration route out of Africa.[13] These findings also highlight the importance of Indian subcontinent in the early genetic history of human settlement and expansion.[14] Therefore, N's history is similar to M and R which have their most probable origin in South Asia.

African origin hypothesisEdit

According to Toomas Kivisild "the lack of L3 lineages other than M and N in India and among non-African mitochondria in general suggests that the earliest migration(s) of modern humans already carried these two mtDNA ancestors, via a departure route over the Horn of Africa.[7]

DistributionEdit

Haplogroup N is derived from the ancestral L3 macrohaplogroup, which represents the migration discussed in the theory of the recent African origin of modern humans. Haplogroup N is the ancestral haplogroup to almost all clades today distributed in Europe and Oceania, as well as many found in Asia and the Americas. It is believed to have arisen at a similar time to haplogroup M. Haplogroup N subclades like haplogroup U6 are also found at high to low frequencies in northwest and northeast Africa due to a back migration from Europe or Asia during the Paleolithic ca. 46,000 ybp, the estimated age of the basal U6* clade.[16]

The haplogroup N descendant lineage U6 has been found among Iberomaurusian specimens at the Taforalt site, which date from the Epipaleolithic.[17] The N1b subclade has been observed in an individual belonging to the Mesolithic Natufian culture.[18] Additionally, haplogroup N has been found among ancient Egyptian mummies excavated at the Abusir el-Meleq archaeological site in Middle Egypt, which date from the Pre-Ptolemaic/late New Kingdom, Ptolemaic, and Roman periods.[19]

The N1 subclade has also been found in various fossils that were analysed for ancient DNA, including specimens associated with the Starčevo (N1a1a1, Alsónyék-Bátaszék, Mérnöki telep, 1/3 or 33%), Linearbandkeramik (N1a1a1a3, Szemely-Hegyes, 1/1 or 100%; N1a1b/N1a1a3/N1a1a1a2/N1a1a1/N1a1a1a, Halberstadt-Sonntagsfeld, 6/22 or ~27%), Alföld Linear Pottery (N1a1a1, Hejőkürt-Lidl, 1/2 or 50%), Transdanubian Late Neolithic (N1a1a1a, Apc-Berekalja, 1/1 or 100%), Protoboleráz (N1a1a1a3, Abony, Turjányos-dűlő, 1/4 or 25%), and Iberia Early Neolithic cultures (N1a1a1, Els Trocs, 1/4 or 25%).[20]

In popular scienceEdit

In the book The Real Eve, Stephen Oppenheimer refers to haplogroup N as "Nasreen" as haplogroup N may have arisen near the Persian Gulf. In his popular book The Seven Daughters of Eve, Bryan Sykes named the originator of this mtDNA haplogroup "Naomi".

Subgroups distributionEdit

Haplogroup N's derived clades include the macro-haplogroup R and its descendants, and haplogroups A, I, S, W, X, and Y.

Rare unclassified haplogroup N* has been found among fossils belonging to the Cardial and Epicardial culture (Cardium pottery) and the Pre-Pottery Neolithic B.[21] A rare unclassified form of N has been also been reported in modern Algeria[22].

  • Haplogroup N1'5
  • Haplogroup N2
    • Haplogroup N2a – small clade found in West Europe.[27]
    • Haplogroup W[28] – found in Western Eurasia and South Asia[29]
  • Haplogroup N8 – found in China.[30]
  • Haplogroup N9 – found in Far East.[23] [TMRCA 45,709.7 ± 7,931.5 ybp; CI=95%[31]]
    • Haplogroup N9a – Thailand/Laos, China, Vietnam [TMRCA 17,520.4 ± 4,389.8 ybp; CI=95%[31]]
      • Haplogroup N9a1'3 [TMRCA 15,007.4 ± 6,060.1 ybp; CI=95%[31]]
        • Haplogroup N9a1 - Chinese (Hakka in Taiwan, etc.), She, Tu, Uyghur, Tuvan, Mongolia, Khamnigan,[32] Korean,[32] Japan [TMRCA 9,200 (95% CI 7,100 <-> 11,600) ybp[33]]
          • Haplogroup N9a1a - Chinese (Sichuan, Zhanjiang, etc.) [TMRCA 7,300 (95% CI 3,800 <-> 12,800) ybp[33]]
          • Haplogroup N9a1b - Kyrgyz (Tashkurgan)[34][33]
          • Haplogroup N9a1c - Vietnam (Tay people),[35] northern Thailand (Khon Mueang),[36] northeastern Thailand (Lao Isan)[36]
        • Haplogroup N9a3 - China [TMRCA 11,500 (95% CI 7,500 <-> 16,800) ybp[33]]
          • Haplogroup N9a3a - Japan, Korean (Seoul), Taiwan (incl. Paiwan), Thailand/Laos, China, Kyrgyz (Tashkurgan), Uyghur, Buryat, Chechen Republic, Russian (Belgorod), Belarus, Poland, Czech (West Bohemia) [TMRCA 8,280.9 ± 5,124.4 ybp; CI=95%[31]]
      • Haplogroup N9a2'4'5'11 [TMRCA 15,305.4 ± 4,022.6 ybp; CI=95%[31]]
        • Haplogroup N9a2 - Japan, China (Barghut in Hulunbuir, Uyghur, etc.) [TMRCA 10,700 (95% CI 8,200 <-> 13,800) ybp[33]]
          • Haplogroup N9a2a - Japan, Korea, Uyghur [TMRCA 8,100 (95% CI 6,500 <-> 10,000) ybp[33]]
            • Haplogroup N9a2a1 - Japan [TMRCA 4,200 (95% CI 1,850 <-> 8,400) ybp[33]]
            • Haplogroup N9a2a2 - Japan, Volga-Ural region (Tatar)[32] [TMRCA 5,700 (95% CI 3,500 <-> 8,900) ybp[33]]
            • Haplogroup N9a2a3 - Japan, Hulun-Buir region (Barghut)[32] [TMRCA 4,700 (95% CI 2,400 <-> 8,400) ybp[33]]
            • Haplogroup N9a2a4 - Japan [TMRCA 2,800 (95% CI 600 <-> 7,900) ybp[33]]
          • Haplogroup N9a2b - China
          • Haplogroup N9a2c [TMRCA 7,200 (95% CI 3,600 <-> 12,700) ybp[33]]
            • Haplogroup N9a2c* - Japan
            • Haplogroup N9a2c1 - Japan, Uyghur [TMRCA 3,700 (95% CI 2,000 <-> 6,100) ybp[33]]
          • Haplogroup N9a2d - Japan [TMRCA 4,900 (95% CI 1,650 <-> 11,400) ybp[33]]
          • Haplogroup N9a2e - China
        • Haplogroup N9a4 [TMRCA 7,900 (95% CI 3,900 <-> 14,300) ybp[33]]
          • Haplogroup N9a4a - Japan [TMRCA 4,400 (95% CI 1,500 <-> 10,200) ybp[33]]
          • Haplogroup N9a4b [TMRCA 5,700 (95% CI 2,400 <-> 11,400) ybp[33]]
            • Haplogroup N9a4b* - Japan
            • Haplogroup N9a4b1 - China (Minnan in Taiwan, etc.)
            • Haplogroup N9a4b2 - China
        • Haplogroup N9a5 [TMRCA 8,700 (95% CI 4,700 <-> 15,000) ybp[33]]
          • Haplogroup N9a5* - Korea
          • Haplogroup N9a5a - Japan
          • Haplogroup N9a5b - Japan [TMRCA 5,300 (95% CI 1,150 <-> 15,300) ybp[33]]
        • Haplogroup N9a6 - Thailand/Laos, Sumatra [TMRCA 11,972.5 ± 5,491.7 ybp; CI=95%[31]]
          • Haplogroup N9a6a - Cambodia (Khmer), Malaysia (Bidayuh, Jehai, Temuan, Kensiu), Sumatra, Sundanese
          • Haplogroup N9a6b - Malaysia (Seletar)
        • Haplogroup N9a7 - Japan
        • Haplogroup N9a8 - Japan, Buryat
        • Haplogroup N9a9 - Tubalar, Kyrgyz, Ukraine
        • Haplogroup N9a10 - Thailand, Laos (incl. Hmong), Vietnam (Tay Nung), China (incl. Han in Chongqing)
          • Haplogroup N9a10a - Taiwan (Ami)
            • Haplogroup N9a10a1 - Chinese (Suzhou)
            • Haplogroup N9a10a2 - Philippines (Ivatan), Taiwan (Ami)
              • Haplogroup N9a10a2a - Taiwan (Atayal, Tsou)
          • Haplogroup N9a10b - China
        • Haplogroup N9a11 - Taiwan (Hakka, Minnan), Thailand/Laos
    • Haplogroup N9b – Japan, Udegey, Nanai [TMRCA 14,885.6 ± 4,092.5 ybp; CI=95%[31]]
      • Haplogroup N9b1 - Japan [TMRCA 11,859.3 ± 3,760.2 ybp; CI=95%[31]]
        • Haplogroup N9b1a - Japan [TMRCA 10,645.2 ± 3,690.3 ybp; CI=95%[31]]
        • Haplogroup N9b1b - Japan [TMRCA 2,746.5 ± 2,947.0 ybp; CI=95%[31]]
        • Haplogroup N9b1c - Japan [TMRCA 6,987.8 ± 4,967.0 ybp; CI=95%[31]]
          • Haplogroup N9b1c1 - Japan
      • Haplogroup N9b2 - Japan [TMRCA 13,369.7 ± 4,110.0 ybp; CI=95%[31]]
        • Haplogroup N9b2a - Japan
      • Haplogroup N9b3 - Japan [TMRCA 7,629.8 ± 6,007.6 ybp; CI=95%[31]]
      • Haplogroup N9b4 - Japan, Ulchi
    • Haplogroup Y[37] – found especially among Nivkhs, Ulchs, Nanais, Negidals, Ainus, and the population of Nias Island, with a moderate frequency among other Tungusic peoples, Koreans, Mongols, Koryaks, Itelmens, Chinese, Japanese, Tajiks, Island Southeast Asians (including Taiwanese aborigines), and some Turkic peoples[15] [TMRCA 24,576.4 ± 7,083.2 ybp; CI=95%[31]]
      • Haplogroup Y1 - Korea, Uyghur, Khamnigan, Taiwan (Minnan), Poland, Slovakia, Czech Republic [TMRCA 14,689.5 ± 5,264.3 ybp; CI=95%[31]]
        • Haplogroup Y1a - Nivkh, Ulchi, Hezhen, Udegey, Even, Zabaikal Buryat, Mongolian, Daur, Han, Tibet [TMRCA 7,467.5 ± 5,526.7 ybp; CI=95%[31]]
          • Haplogroup Y1a1 - Uyghur, Kyrgyz, Yakut, Buryat, Hezhen, Udegey, Taimyr Evenk
          • Haplogroup Y1a2 - Koryak, Kamchatka Even
        • Haplogroup Y1b - Volga Tatar [TMRCA 9,222.8 ± 4,967.0 ybp; CI=95%[31]]
          • Haplogroup Y1b1 - Chinese, Japanese, Korea, Russia
      • Haplogroup Y2 - Chinese, Japanese, Korean, Khamnigan, South Africa (Cape Coloured) [TMRCA 7,279.3 ± 2,894.5 ybp; CI=95%[31]]
        • Haplogroup Y2a - Taiwan (Atayal, Saisiyat, Tsou), Philippines, Brunei, Indonesia, Malaysia, Hawaii, USA (Hispanic), Spain, Ireland [TMRCA 4,929.5 ± 2,789.6 ybp; CI=95%[31]]
        • Haplogroup Y2b - Japan, Buryat [TMRCA 1,741.8 ± 3,454.2 ybp; CI=95%[31]]
  • Haplogroup N10 – found in China (Han from Shanghai, Jiangsu, Fujian, Guangdong, and Yunnan, Hani and Yi from Yunnan, She from Guizhou, Uzbek from Xinjiang) and Southeast Asia (Thailand, Indonesia, Vietnam, Malaysia).[30]
  • Haplogroup N11 – found in China (Han from Yunnan, Sichuan, and Hubei, Tibetan from Xizang, Dongxiang from Gansu, Oroqen from Inner Mongolia) and the Philippines.[30][38]
  • Haplogroup O or N12- found among indigenous Australians and the Floresians of Indonesia.
  • Haplogroup N13 – indigenous Australians[39]
  • Haplogroup N14 – indigenous Australians[40]
  • Haplogroup N21 – In ethnic Malays from Malaysia and Indonesia.[41]
  • Haplogroup N22 – Southeast Asia, Bangladesh, India, Japan[42]
  • Haplogroup A[43] – found in Central and East Asia, as well as among Native Americans.
  • Haplogroup S[44] – extended among indigenous Australians
  • Haplogroup X[45] – found most often in Western Eurasia, but also present in the Americas.[23]
    • Haplogroup X1 – found primarily in North Africa as well as in some populations of the Levant, notably among the Druze
    • Haplogroup X2 – found in Western Eurasia, Siberia and among Native Americans
  • Haplogroup R[46] – a very extended and diversified macro-haplogroup.

SubcladesEdit

TreeEdit

This phylogenetic tree of haplogroup N subclades is based on the paper by Mannis van Oven and Manfred Kayser Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation[9] and subsequent published research.

  • N
    • N1'5
      • N1
        • N1a'c'd'e'I
          • N1a'd'e'I
            • N1a'e'I
              • N1a
                • N1a1
                  • N1a1a
              • N1e'I
                • I
                • N1e
            • N1d
          • N1c
        • N1b
          • N1b1
            • N1b1a
            • N1b1b
            • N1b1c
              • N1b1d
          • N1b2
      • N5
    • N2
      • N2a
      • W
    • N9
      • N9a
        • N9a1'3
          • N9a1
          • N9a3
        • N9a2'4'5
          • N9a2
            • N9a2a'b
              • N9a2a
              • N9a2b
            • N9a2c
            • N9a2d
          • N9a4
          • N9a5
        • N9a6
          • N9a6a
      • N9b
        • N9b1
          • N9b1a
          • N9b1b
          • N9b1c
            • N9b1c1
        • N9b2
        • N9b3
      • Y
    • N13
    • N14
    • N21
    • N22
    • A
    • O
      • O1
    • S
    • X
    • R

See alsoEdit

Phylogenetic tree of human mitochondrial DNA (mtDNA) haplogroups

  Mitochondrial Eve (L)    
L0 L1–6  
L1 L2   L3     L4 L5 L6
M N  
CZ D E G Q   O A S R   I W X Y
C Z B F R0   pre-JT   P   U
HV JT K
H V J T

ReferencesEdit

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