Haplogroup I-M253

Haplogroup I-M253, also known as I1, is a Y chromosome haplogroup. The genetic markers confirmed as identifying I-M253 are the SNPs M253,M307.2/P203.2, M450/S109, P30, P40, L64, L75, L80, L81, L118, L121/S62, L123, L124/S64, L125/S65, L157.1, L186, and L187.[6] It is a primary branch of Haplogroup I-M170 (I*).

Haplogroup I1 (M253)
Possible time of origin3,170–4,600[1]-5,070 BP (today's diversification)[2][3] (previously 11,000 BP[4] to 33,000 BP[5]) 27,500 (diversification with I2-FGC77992)[1]
Possible place of originNorthern Europe
AncestorI* (M170)
DescendantsI1a (DF29/S438);
I1b (S249/Z131);
I1c (Y18119/Z17925)
Defining mutationsM253, M307.2/P203.2, M450/S109, P30, P40, L64, L75, L80, L81, L118, L121/S62, L123, L124/S64, L125/S65, L157.1, L186, L187

The haplogroup is believed to have been present among Upper Paleolithic European hunter-gatherers as a minor lineage but due to its absence in pre-Neolithic DNA samples it can't have been very widespread. Neolithic I1 samples are very sparse as well, suggesting a rapid dispersion connected to a founder effect in the Nordic Bronze Age. Today it reaches its peak frequencies in Sweden (52 percent of males in Västra Götaland County) and western Finland (more than 50 percent in Satakunta province).[7] In terms of national averages, I-M253 is found in 38-39% of Swedish males,[8][9][10] 37% of Norwegian males,[11][12][13] 34.8% of Danish males,[14][15][16] and about 28% of Finnish males.[17] Bryan Sykes, in his 2006 book Blood of the Isles, gives the members – and the notional founding patriarch of I1 the name "Odin".[18]

Haplogroup I-M253 is a primary branch of haplogroup I* (I-M170), which has been present in Europe since ancient times. The other primary branch of I* is I-M438, also known as I2.

All known living members descend from a common ancestor 6 times younger than the common ancestor with I2.[1]

Before a reclassification in 2008,[19] the group was known as I1a, a name that has since been reassigned to a primary branch, haplogroup I-DF29. The other primary branches of I1 (M253) are I1b (S249/Z131) and I1c (Y18119/Z17925).

Origins
Map of the early Nordic Bronze Age, where I1 first became prominent. The Nordic Bronze Age is often considered ancestral to the Germanic peoples.

While haplogroup I1 most likely diverged from I* as early as 27,000 years ago in the Gravettian, so far no DNA-studies have been able to locate it in Mesolithic hunter-gatherers. As of November 2020, only 4 ancient DNA samples dating to earlier than the Nordic Bronze Age have been assigned to haplogroup I1:

  • The first is an individual sample labelled BAB5 from Neolithic Hungary.[20]
  • Another is an individual belonging to the middle Neolithic Chasséen culture labelled Cx161. Cx161 had a genetic affinity to other contemporary Neolithic farmers of Europe.[21]
  • Additionally, the third ancient I1 sample is from an individual found in a kurgan burial dating to the late Neolithic Dagger Period in Scandinavia labelled RISE179.[22] RISE179 had a genetic affinity to the populations of the Corded Ware culture and the Unetice culture.[23]
  • The fourth ancient I1 sample predating the Nordic Bronze Age is labelled oll009 and was sequenced in the study titled "The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon".[24] Oll009 is dated to the Scandinavian late Neolithic and was found in a burial in Sweden. Similar to RISE179, he carried a high percentage of Western Steppe-Herder ancestry and had a genetic affinity to the population of the Battle Axe culture and other populations of the Corded Ware horizon.[25]

Despite the high frequency of haplogroup I1 in present-day Scandinavians, I1 is completely absent among Scandinavian Mesolithic DNA samples.[26][27] I1 first appears in Scandinavia during the late Neolithic but does not increase in frequency until the beginning of the Nordic Bronze Age.[28][29][30][23]

Given the nature of ancient DNA samples Cx161 and RISE179, a possible explanation for why haplogroup I1 is thus far absent among Mesolithic and Neolithic hunter-gatherers in Scandinavia is that I1 may have been present as a minor lineage in Central European Neolithic cultures such as the Rössen culture, Globular Amphora culture and the Funnelbeaker culture.[31] It could later have been assimilated by the people of the Corded Ware culture and brought to Scandinavia as a part of the Battle Axe culture.[32][33][34] Given that I1 has not been found in any DNA samples in Scandinavia predating the Battle Axe culture, it is likely that the haplogroup was brought there by the people of that culture.[35][36]

According to a study published in 2010, I-M253 originated between 3,170 and 5,000 years ago, in Chalcolithic Europe.[2] A new study in 2015 estimated the origin as between 3,470 and 5,070 years ago or between 3,180 and 3,760 years ago, using two different techniques.[3]

In 2007, it was suggested that it initially dispersed from the area that is now Denmark.[14] However, Prof. Dr. Kenneth Nordtvedt, Montana State University, regarding the MRCA, in 2009 wrote in a personal message: "We don't know where that man existed, but the greater lower Elbe basin seems like the heartland of I1".

Latest results (Sept. 2019) published by Y-Full suggest I1 (I-M253) was formed 27.500 ybp (95 CI: 29.800 ybp – 25.200 ybp) with TMRCA 4.600 ybp (95 CI: 5.200 ybp – 4.000 ybp). Since the most up-to date calculated estimation of TMRCA of I1 is thought to be around 2750 BC, this likely puts the ancestor of all living I1 men somewhere in Central or Northern Europe around the time that Funnelbeaker and related cultures mingled with offshoots of the Corded Ware culture. The phylogeny of I1 shows the signature of a rapid star-like expansion.[37][38] This suggests that I1 went from being a rare marker to a rather common one in a rapid burst.

A 2014 study in Hungary uncovered remains of two individuals from the Linear Pottery culture, one of whom was found to have carried the M253 SNP which defines Haplogroup I1. This culture is thought to have been present between 7,500 and 6,500 years ago.[39]

Structure

I-M253 (M253, M307.2/P203.2, M450/S109, P30, P40, L64, L75, L80, L81, L118, L121/S62, L123, L124/S64, L125/S65, L157.1, L186, and L187) or I1 [6]

  • I-DF29 (DF29/S438); I1a
    • I-CTS6364 (CTS6364/Z2336); I1a1
      • FGC20030; I1a1a~
        • S4767; I1a1a1~
              • I-M227; I1a1a1a1a
        • A394; I1a1a2~
        • Y11221; I1a1a3~
        • A5338; I1a1a4~
      • CTS10028; I1a1b
        • I-L22 (L22/S142); I1a1b1
          • CTS11651/Z2338; I1a1b1a~
            • I-P109; I1a1b1a1
              • I-Y3662; I1a1b1a1e~
                • I-S14887; I1a1b1a1e2~
                  • I-Y11203; I1a1b1a1e2d~
                    • I-Y29630; I1a1b1a1e2d2~
            • CTS6017; I1a1b1a2
            • I-L205 (L205.1/L939.1/S239.1); I1a1b1a3
            • CTS6868; I1a1b1a4
              • I-Z74; I1a1b1a4a
                • CTS2208; I1a1b1a4a1~
                  • I-L287; I1a1b1a4a1a
                    • I-L258 (L258/S335); I1a1b1a4a1a1
                • I-L813; I1a1b1a4a2
                • FGC12562; I1a1b1a4a3~
          • CTS11603/S4744; I1a1b1b~
                • I-L300 (L300/S241); I1a1b1b1a1
        • FGC10477/Y13056; I1a1b2
        • A8178, A8182, A8200, A8204; I1a1b3~
        • F13534.2/Y17263.2; I1a1b4~
    • I-Z58 (S244/Z58); I1a2
      • I-Z59 (S246/Z59); I1a2a
        • I-Z60 (S337/Z60, S439/Z61, Z62); I1a2a1
          • I-Z140 (Z140, Z141)
            • I-L338
            • I-F2642 (F2642)
          • I-Z73
            • I-L1302
          • I-L573
          • I-L803
        • I-Z382; I1a2a2
      • I-Z138 (S296/Z138, Z139); I1a2b
        • I-Z2541
    • I-Z63 (S243/Z63); I1a3
      • I-BY151; I1a3a
        • I-L849.2; I1a3a1
        • I-BY351; I1a3a2
            • I-CTS10345
              • I-Y10994
            • I-Y7075
        • I-S2078
          • I-S2077
            • I-Y2245 (Y2245/PR683)
              • I-L1237
                • I-FGC9550
              • I-S10360
                • I-S15301
              • I-Y7234
        • I-BY62 (BY62); I1a3a3
  • I-Z131 (Z131/S249); I1b
    • I-CTS6397; I1b1
  • I-Z17943 (Y18119/Z17925, S2304/Z17937); I1c

Historical expansion
Percentage of major Y-DNA haplogroups in Europe; Haplogroup I1 represented by light blue.

Haplogroup I1, as well as subclades of R1b such as R1b-U106 and subclades of R1a such as R1a-Z284, are strongly associated with Germanic peoples and are linked to the proto-Germanic speakers of the Nordic Bronze Age.[40][41] Current DNA research indicates that I1 was close to non-existent in most of Europe outside of Scandinavia and northern Germany before the Migration Period. The expansion of I1 is directly tied to that of the Germanic tribes. Starting around 900 BC, Germanic tribes started moving out of southern Scandinavia and northern Germany into the nearby lands between the Elbe and the Oder. Between 600 and 300 BC another wave of Germanics migrated across the Baltic Sea and settled alongside the Vistula. Germanic migration to that area resulted in the formation of the Wielbark culture, which is associated with the Goths.[42]

I1-Z63 has been traced to the Kowalewko burial site in Poland which dates to the Roman Iron Age. In 2017 Polish researchers could successfully assign YDNA haplogroups to 16 individuals who were buried at the site. Out of these 16 individuals, 8 belonged to I1. In terms of subclades, three belonged to I-Z63, and in particular subclade I-L1237. [43] The Kowalewko archeological site has been associated with the Wielbark culture. Therefore the subclade I-L1237 of I-Z63 may be seen somewhat as a genetic indicator of the Gothic tribe of late antiquity. I1-Z63 has also been found in a burial associated with Goth and Lombard remains in Collegno, Italy.[44][45] The cemetery is dated to the late 6th Century and further suggests that I1-Z63 and downstream subclades are linked to early Medieval Gothic migrations.

In 2015, a DNA study tested the Y-DNA haplogroups of 12 samples dated to 300-400 AD from Saxony-Anhalt in Germany. 8 of them belonged to haplogroup I1. This DNA evidence is in alignment with the historical migrations of Germanic tribes from Scandinavia to central Europe.[46]

Additionally, I1-Z63 was found in the Late Antiquity site Crypta Balbi in Rome, this time with the downstream subclade I-Y7234.[47] Material findings associated with the Lombards have been excavated in Crypta Balbi.

The Pla de l'Horta villa near Girona in Spain is located in close proximity to a necropolis with a series of tombs associated with the Visigoths. The grave goods and the typology of the tombs point to a Visigothic origin of the individuals. A small number of individuals buried at the site were sampled for DNA analysis in a 2019 study. One of the samples belonged to haplogroup I1.[48] This finding is in accordance with the common ancestral origin of the Visigoths and the Ostrogoths.

The Anglo-Saxon settlement of Britain introduced I1 in the British Isles.[49]

During the Viking Age, I-M253 saw another expansion. Norwegian and Danish Vikings brought more I1 to Britain and Ireland, while Swedish Vikings introduced it to Russia and Ukraine and brought more of it to Finland and Estonia.[50]

Geographical distribution

I-M253 is found at its highest density in Northern Europe and other countries that experienced extensive migration from Northern Europe, either in the Migration Period, the Viking Age, or modern times. It is found in all places invaded by the Norse.

During the modern era, significant I-M253 populations have also taken root in immigrant nations and former European colonies such as the United States, Australia and Canada.

Population Sample size I (total) I1 (I-M253) I1a1a (I-M227) Source
Albanians (Tirana)5521.82%=(12/55)3.6%=(2/55)0.0Battaglia et al. 2008
Albanians (North Macedonia)6417.2%=(11/64)4.7%=(3/64)0.0Battaglia et al. 2008
Albanians (Tirana)
Albanians (North Macedonia)		55+64=11919.33%=(23/119)4.2%=(5/119)0.0Battaglia et al. 2008
Kosovo Albanians (Pristina)1147.96%=(9/114)5.31%=(6/114)0.0Pericic et al. 2005
Albanians (Tirana)
Albanians (North Macedonia)
Kosovo Albanians (Pristina)		55+64+114=23313.73%=(32/233)4.72%=(11/233)0.0Pericic et al. 2005
Battaglia et al. 2008
Austria439.32.30.0Underhill et al. 2007
Belarus: Vitbsk100151.00.0Underhill et al. 2007
Belarus: Brest9720.61.00.0Underhill et al. 2007
Bosnia100422.00.0Rootsi et al. 2004
Bulgaria80826.64.30.0Karachanak et al. 2013
Czech Republic4731.98.50.0Underhill et al. 2007
Czech Republic5317.01.90.0Rootsi et al. 2004
Denmark12239.3% (48/122)32.8% (40/122)0.0Underhill et al. 2007
England10419.215.40.0Underhill et al. 2007
Estonia21018.614.80.5Rootsi et al. 2004
Estonia11811.9Lappalainen et al. 2008
Finland (national)28.0Lappalainen et al. 2006
Finland: West23040% (92/230)Lappalainen et al. 2008
Finland: East30619% (58/306)Lappalainen et al. 2008
Finland: Satakunta region50+Lappalainen et al. 20089
France5817.28.61.7Underhill et al. 2007
France1216.716.70.0Cann et al. 2002
France (Low Normandy)4221.411.90.0Rootsi et al. 2004
Germany1252415.20.0Underhill et al. 2007
Greece17115.82.30.0Underhill et al. 2007
Hungary11325.713.30.0Rootsi et al. 2004
Ireland100116.00.0Underhill et al. 2007
Kazan Tatars5313.211.30.0Trofimova 2015
Latvia1133.5Lappalainen et al. 2008
Lithuania1644.9Lappalainen et al. 2008
Netherlands9320.4140.0Underhill et al. 2007
Norway176637% (653/1766)Stenersen et al 2006
Russia (national)162512.50.0Cann et al. 2002
Russia: Pskov13016.95.40.0Underhill et al. 2007
Russia: Kostroma5326.411.30.0Underhill et al. 2007
Russia: Smolensk10312.61.90.0Underhill et al. 2007
Russia: Voronez9619.83.10.0Underhill et al. 2007
Russia: Arkhangelsk14515.87.60.0Underhill et al. 2007
Russia: Cossacks8924.74.50.0Underhill et al. 2007
Russia: Karelians140108.60.0Underhill et al. 2007
Russia: Karelians13215.2Lappalainen et al. 2008
Russia: Vepsa395.12.60.0Underhill et al. 2007
Slovakia7014.34.30.0Rootsi et al. 2004
Slovenia9526.37.40.0Underhill et al. 2007
Sweden (national)16035.6% (57/160)Lappalainen et al. 2008
Sweden (national)38.0Lappalainen et al. 2009
Sweden: Västra Götaland52Lappalainen et al. 2009
Switzerland1447.65.60.0Rootsi et al. 2004
Turkey5235.41.10.0Underhill et al. 2007
Ukraine: Lviv10123.84.90.0Underhill et al. 2007
Ukraine: Ivanovo-Frankiv5621.41.80.0Underhill et al. 2007
Ukraine: Hmelnitz17626.26.10.0Underhill et al. 2007
Ukraine: Cherkasy11428.14.30.0Underhill et al. 2007
Ukraine: Bilhorod5626.85.30.0Underhill et al. 2007
Map showing the distribution of Y chromosomes in a trans section of England and Wales from the paper "Y Chromosome Evidence for Anglo-Saxon Mass Migration". The authors attribute the differences in frequencies of haplogroup I1 to Anglo-Saxon mass migration into England, but not into Wales.

In 2002 a paper was published by Michael E. Weale and colleagues showing genetic evidence for population differences between the English and Welsh populations, including a markedly higher level of Y-DNA haplogroup I1 in England than in Wales. They saw this as convincing evidence of Anglo-Saxon mass invasion of eastern Great Britain from northern Germany and Denmark during the Migration Period.[51] The authors assumed that populations with large proportions of haplogroup I1 originated from northern Germany or southern Scandinavia, particularly Denmark, and that their ancestors had migrated across the North Sea with Anglo-Saxon migrations and Danish Vikings. The main claim by the researchers was

that an Anglo-Saxon immigration event affecting 50–100% of the Central English male gene pool at that time is required. We note, however, that our data do not allow us to distinguish an event that simply added to the indigenous Central English male gene pool from one where indigenous males were displaced elsewhere or one where indigenous males were reduced in number … This study shows that the Welsh border was more of a genetic barrier to Anglo-Saxon Y chromosome gene flow than the North Sea … These results indicate that a political boundary can be more important than a geophysical one in population genetic structuring.

Distribution of Y chromosome haplogroups from Capelli et al. (2003). Haplogroup I-M253 is present in all populations, with higher frequencies in the east and lower frequencies in the west. There appears to be no discrete boundary as observed by Weale et al. (2002)

In 2003 a paper was published by Christian Capelli and colleagues which supported, but modified, the conclusions of Weale and colleagues.[52] This paper, which sampled Great Britain and Ireland on a grid, found a smaller difference between Welsh and English samples, with a gradual decrease in Haplogroup I1 frequency moving westwards in southern Great Britain. The results suggested to the authors that Norwegian Vikings invaders had heavily influenced the northern area of the British Isles, but that both English and mainland Scottish samples all have German/Danish influence.

Prominent members of I-M253

Alexander Hamilton, through genealogy and the testing of his descendants (assuming actual paternity matching his genealogy), has been placed within Y-DNA haplogroup I-M253.[53]

The Varangian Šimon, who was said to have been the founder of the Russian Vorontsov noble family, belonged to haplogroup I1-Y15024.[54][55] Testing by geneticist Peter Sjölund and FamilyTreeDNA showed that the present-day male members of the Vorontsov family still carry this subclade of I1, and downstream subclades.[56][57] Some prominent historical members of the Vorontsov family were Prince Mikhail Semyonovich Vorontsov, a Russian prince and field-marshal, as well as Count Illarion Ivanovich Vorontsov-Dashkov, a general of the Russian Empire.

The Rurikid Prince Sviatopolk the Accursed (son of Vladimir the Great) was found to likely have carried the I1-S2077 subclade of I1-Z63.[58][59][60]

Birger Jarl, 'Duke of Sweden' of the East Geatish House of Bjälbo, founder of Stockholm; his remains were exhumed and tested in 2002 and found to be I-M253.[61] The House of Bjälbo also provided three kings of Norway, and one king of Denmark in the 14th century.

Sting was revealed to belong to haplogroup I1 by the PBS TV series Finding Your Roots.[62]

William Bradford (governor) of the Mayflower, proven through the Mayflower DNA Project[63]

William Brewster (Mayflower passenger) of the Mayflower, proven through the Mayflower DNA Project[63]

General Robert E. Lee belonged to I-M253 based on DNA testing of his descendants as a part of the Lee DNA Project. Other prominent members of the Lee family of Virginia and Maryland were Richard Lee I and Richard Henry Lee. The latter was one of the Founding Fathers of the United States of America.[64]

Robert I of Scotland, commonly known as Robert the Bruce, belonged to haplogroup I1. Descendant testing of Robert, 6th lord of Annandale de Brus, assigned the men of Clan Bruce to I1-Y17395.[65]

The male members of the House of Grimaldi were revealed to carry haplogroup I1 as a part of the Grimaldi Genealogy DNA project.[66] They were lords and princes of Monaco up until 1949. The men of House Grimaldi belong to a Scandinavian subclade of I1, downstream of I1-Y3549.

President Andrew Jackson belonged to haplogroup I1, based on results from the Jackson DNA project and from genealogy.[67][68]

The Russian writer Leo Tolstoy was found to have carried I1. The testing of his male descendant Pyotr Tolstoy revealed that the males of the Tolstoy family carry I1-M253.[69][70]

Snorri Sturluson was found to likely have belonged to haplogroup I1. Y-DNA testing of his presumed descendants revealed an assignment to I-M253. Their results are available on YSearch.org.[71]

The Swedish scientist and theologian Emanuel Swedenborg and other male members of the Swedenborg noble family were found to belong to haplogroup I1-BY229, as a part of the I1-L1302 DNA project by Jakob Norstedt.[72][73]

Siener van Rensburg, Boer patriotic figure and mystic, belonged to haplogroup I1.[74][75]

Björn Wahlroos, Finnish businessman and millionaire, was found to belong to haplogroup I1.[71]

The Finnish mathematician Rolf Nevanlinna belonged to I1-M253 based on the testing of his son Arne Nevanlinna by Geni.com.[76][77][78]

Samuel Morse was found to have carried haplogroup I1 as a part of the Morse DNA project.[79][80][81]

Felix Kjellberg (PewDiePie) was found to belong to haplogroup I1-L22, according to testing by 23andMe.[82]

Swedish actor Björn Andrésen belongs to haplogroup I1-L22 based on the ftDNA and 23andMe tests of one of his first cousins and one uncle on the paternal side as a part of their family research.[83][84][85][86]Their ancestor Johan Andrésen lived on both sides of the Swedish-Norwegian border.[87][88]

As a part of the Pine family DNA project, actor Chris Pine was found to belong to haplogroup I1-A13819.[89][90]

Markers
DNA example: strand 1 differs from strand 2 at a single base pair location (a C >> T polymorphism).

The following are the technical specifications for known I-M253 haplogroup SNP and STR mutations.

Name: M253[91]

Type: SNP
Source: M (Peter Underhill of Stanford University)
Position: ChrY:13532101..13532101 (+ strand)
Position (base pair): 283
Total size (base pairs): 400
Length: 1
ISOGG HG: I1
Primer F (Forward 5′→ 3′): GCAACAATGAGGGTTTTTTTG
Primer R (Reverse 5′→ 3′): CAGCTCCACCTCTATGCAGTTT
YCC HG: I1
Nucleotide alleles change (mutation): C to T

Name: M307[92]

Type: SNP
Source: M (Peter Underhill)
Position: ChrY:21160339..21160339 (+ strand)
Length: 1
ISOGG HG: I1
Primer F: TTATTGGCATTTCAGGAAGTG
Primer R: GGGTGAGGCAGGAAAATAGC
YCC HG: I1
Nucleotide alleles change (mutation): G to A

Name: P30[93]

Type: SNP
Source: PS (Michael Hammer of the University of Arizona and James F. Wilson, at the University of Edinburgh)
Position: ChrY:13006761..13006761 (+ strand)
Length: 1
ISOGG HG: I1
Primer F: GGTGGGCTGTTTGAAAAAGA
Primer R: AGCCAAATACCAGTCGTCAC
YCC HG: I1
Nucleotide alleles change (mutation): G to A
Region: ARSDP

Name: P40[94]

Type: SNP
Source: PS (Michael Hammer and James F. Wilson)
Position: ChrY:12994402..12994402 (+ strand)
Length: 1
ISOGG HG: I1
Primer F: GGAGAAAAGGTGAGAAACC
Primer R: GGACAAGGGGCAGATT
YCC HG: I1
Nucleotide alleles change (mutation): C to T
Region: ARSDP

See also

References
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  3. Batini, Chiara; Hallast, Pille; Zadik, Daniel; Delser, Pierpaolo Maisano; Benazzo, Andrea; Ghirotto, Silvia; Arroyo-Pardo, Eduardo; Cavalleri, Gianpiero L.; De Knijff, Peter; Dupuy, Berit Myhre; Eriksen, Heidi A.; King, Turi E.; De Munain, Adolfo López; López-Parra, Ana M.; Loutradis, Aphrodite; Milasin, Jelena; Novelletto, Andrea; Pamjav, Horolma; Sajantila, Antti; Tolun, Aslıhan; Winney, Bruce; Jobling, Mark A. (2015). "TMRCAs of major haplogroups in Europe estimated using two methods. Large-scale recent expansion of European patrilineages shown by population resequencing : Nature Communications : Nature Publishing Group". Nature Communications. 6: 7152. doi:10.1038/ncomms8152. PMC 4441248. PMID 25988751.
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Sources
Haplogroup I databases
General Y-DNA databases

There are several public access databases featuring I-M253, including:

  1. http://www.ysearch.org/
  2. http://www.yhrd.org/
  3. http://www.yfull.com/tree/I1/
Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]
"Y-chromosomal Adam"
A00 A0-T [χ 3]
A0 A1 [χ 4]
A1a A1b
A1b1 BT
B CT
DE CF
D E C F
F1  F2  F3  GHIJK
G HIJK
IJK H
IJ K
I   J     LT [χ 5]       K2 [χ 6]
L     T    K2a [χ 7]        K2b [χ 8]     K2c     K2d K2e [χ 9]  
K-M2313 [χ 10]     K2b1 [χ 11] P [χ 12]
NO   S [χ 13]  M [χ 14]    P1     P2
N O Q R
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