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Revision of Basques Genetics
The Basques inhabit rugose terrain in the SW extremity of France and adjacent Spain,which is comprised of seven provinces.During the last glacial era the polar front reached the Pyrenees and temperatures are estimated to have been about 15 degrees C below todays average levels.The unfavourable climatic conditions probably fragmented northern Iberian populations and limited gene flow between these groups,which subsequently could have been partially perpetuated by the rugged topography over a portion of the Basque Lands.This premise is compatible with the eight distinct regional Euskera dialects,that are spoken by the Basques [de Yrizar].These lands are called Euskal Herria ,the land of the Euskera speakers. It is their unique,ancient,non-Indo-European, agglutinating language,which identifies the Basques, who have a high frequency of Rh negative blood [ca 25%].No linguistic equivalent to their language has been found.The origin of this tongue is unknown.A proto-type "might" have been introduced to the Basque lands by people,who predated the influx of Neolithic agriculturalists,who "may" have spoken an Indo-European language [speculation].
Preservation of the Basque identity contrasts sharply with the physical,but not genetic, disappearance of the Neanderthals.The sparse scatterings of small Neanderthal groups were not able to cope with the appreciable influxes of Homo sapiens.The Basques were largely concentrated in a relatively rugged region and large tracts of their land was not prime acerage.An indigenous core,which was probably assembled from proximal,warmer,segments of Iberia,after the Younger Dryas,appears to have had sufficient numbers to found the Basques and assimilate subsequent newcomers.The original inhabitants included ancient Iberians and possibly some individuals with Neanderthal genes.Later additions have diluted the frequencies of the founders,but the new populace managed to retain a Basque identity.
A Gonz'ales [2006] analyzed 211 Basque samples utilizing the hypervariable segment of the mtDNA control region and diagnostic RFIP techniques.The four sequences,which were derived from haplogroup [hg] U8 were completely sequenced.Hg U was one of the earliest genetic lineages,that was dispersed from the Middle East to western Europe.Sub-hg U81a,which has a coalescence time of ca 13 +/-5.0Ka,has a 1.0% frequency in Basque lands.The greatest U8a diversity occurs on the Iberia Peninsula,which suggests that a few Basques could be descended from the Gravettians [ibid].The northern Iberian Basque provinces of Guipuzcoa and Viscaya have relatively high frequencies of hg U5.Subgroup U5b has its highest Iberian peak [15.4%] among the Guipuzcoas.U5b is rare among the Basque populations of France.Although hg U8a,which has a mean frequency of 1.3% among the Basques,has a scattered presence to the east in Catalonia,it has not been reported west of the Basque provinces.The high frequency variability of hg U5b and the wide differentiation of U8a among Basque lineages indicates that the Basques have maintained a low frequency of Palaeolithic maternal lineages in their extant female gene pool [O Garcia,2011].Analysis of mtDNA samples, which were collected from 55 males in the Arratia valley and the Goiherri region recorded an 18.2% frequency of hg U.The Adaieta cemetery samples contained four ancient U2e samples,which have not been identified among modern Basques.U2e is common in the Middle East [A Alzuaide,2007]. Analysis of the remains of a ca 8.5Ka old Basque fisherman from the cost of Hondarribia and Pessia suggests,that ca 50% of his diet was comprised of marine food [A Arrizbalagg,2006].
Analysis of 623 Y chromosome samples identified 23 hgs.This study indicated,that the barrier between the Basques and their immediate neighbours was minimal.The Iberian populations had a reduced genetic structure and recent migrations have not totally erased the ancient Iberian Y chromosome patterns [C Flores,2004].A survey of eleven Alu insertion polymorphisms,which are DNA autosomal markers,did not detect appreciable allele frequency variations between the Basques and the adjacent Iberian populations.A genome scan of 650,000 SNPs,marginally differentiated the French Basques from other western Europeans.This study did not provide definitive evidence,that the Basques were a genetic outlier [P Garagnani,2009].Analysis of Alu elements by S Garcia-Obergon [2007] did not sustain a common genetic origin between Basques and north Africans or Caucasians. The Iberian Basques grouped with most European populations.The people in Guipuzcoa province,which is surrounded by other Basque provinces often appears to be the most differentiated Basque group.The population of Ireland has retained a higher indigenous component than the Basques,who were closer to the European migration routes.
The mtDNA genetic study by A Alzuaide [2006] provides information about temporal changes in the Basque female gene pool.The mtDNA of 65 human remains from the ca 550-770 ADE Basque Adaista cemetery were analysed and compared with a large selection of worldwide haplotypes and three prehistoric groups of proto-Basques[?] .The unique haplotypes 5,9 and 11 were not present in the Genbank samples or among the world wide comparison sequences.Haplotypes 7,8,10,12,16 and 17 were deemed to be uncommon in Europe. The other six haplotypes were widely dispersed throughout western Europe.This distribution pattern has similarities to those populations on the Cantabrian fringe [ibid].
Two haplotypes of hg H account for 53% of the Aldaieta samples,which compares to 62.3% for extant Basques.,and 37-44% at the prehistoric sites of Rico Ramos,Langar and Sjapl,which range in age from the Neolithic to the Bronze Age .The percentage of hg H has increased since prehistoric times in Basques Lands.This differs from hg V,which has not been identified at the three prehistoric sites and is represented by a single sample at Aldaieta.The frequency of Hg V ranges from 3-20% among present day Basques.There were two hg K samples at Adaieta.Present day Basques have a 3.6% frequency of hg K.This represents a significant reduction since prehistoric times,when frequencies of hg K varied from 16.7-24.0%.Hg J is generally associated with the introduction of farming to Europe from the Fertile Cresent.At the three prehistoric sites hg J averaged 16.4%,which was reduced to 14.7% at Aldaieta.Extant Basques only have a 2.4% frequency of hg J.The frequency of hgs U5 and U2 have diminished from 16.7 among the prehistoric specimens to 13.7% among modern Basques.No hg W samples were identified among the prehistoric or historic samples,which contrasts with a 1.2% presence among the extant Basques.Hg I was not recorded in the prehistoric or the present day samples [ibid].Hg V has a 20% presence among modern Guipuzcoans [Torroni,1998].
The most variable region of the human genome is the noncoding displacement loop region,which is comprised of about 1120 base pairs.The most polymorphic nucleotide positions within the control region are concentrated in HVS I and II.M Alfonso-Sanchez [2008] utilized both elements to focus on current Basques mtDNA distribution.Samples were collected from 55 males in the Arratia valley and Gaiherri region.When HVS I and II are combined 28 of the 35 mtDNA Basque lineages were unique. Comparisons with 14 European and north African populations indicated that the Basque specimens had the lowest level of variability among the female lineages in the study.The Black Death,influenza epidemics,etc could have eliminated or reduced the numbers of some female lineages.Y chromosome studies have revealed,that Basque paternal lineages exhibited low genetic diversity [ibid].
More than 95% of these Basque mtDNA specimens can be assigned to the European-specific hgs H,J,K,U,V and X,with H dominating 50.9%,U 18.2% and J 14.6%.Hg M,which has a low presence among northern Iberians was not identified in the Basque samples.Hg J has 9.0% frequency in Asturia,8.6% in Galacia and a ca 3.0% average across the Basque Lands.Sub-hgs J1a1 [10.9%] and J2a [3.6%] have "relatively" high presences.Hg J was not reported in the comparative Caucasus samples.The French Basques have appreciably higher frequencies of hg J than their Iberian counterparts.Galacia has a very low hg U frequency,while Saragossa has a significantly higher Hg T content.No African or U8a lineages were identified among these Basque individuals.Hg V was poorly represented,when compared to earlier studies. There are considerable variations in the mtDNA lineage frequencies in different Basque regions and there is a paucity of prehistorical genetuic data that can be used to make direct comparisons between prehistoric and present day mtDNA hg frequencies and distribution patterns in specific areas [ibid].The pre- and historical Basques received external gene flow,which is not always reflected to the same extent in extant Basques populations.S Alonsel [2006] contends that they may not always have been the focus of major population expansions and might not be best representatives of the ancient European gene pool.It appears that hg V was introduced to the Basques at a relatively late date.There are genetic discontinuities between prehistoric/historic Basques and the extant populations [A Alzuaide,2006].
O Gracia et al [2011] sequenced the HVS I and II components for 413 European mtDNA samples and integrated the new data with 5436 specimens from the available literature.The 2011 study revealed.that 236 of the 413 individuals harboured hg H [19 subgroups].A total of 295 different haplotypes were identified.The proportion of different haplotypes was lower [64%] among the Basques than it was in other northern Iberian populations [90%].The Basques appear to have been less susceptible to external influences.However the north African clades M1 and M6 were recorded in the Basque samples.The 13.5% presence of hg T in Viscaya contrasts with the 4.0% in Guipuzcoa and Alva,which emphasizes the variations in mtDNA distribution between the different regions of Euskal Herra and the adjacent Iberian provinces [ibid].The mtDNA hg dispersement patterns among the French Basques varies from their Iberian counterparts [eg;hg J:Richard,2007].
The frequencies of mtDNa hg V are significantly higher in southern Iberia than in the extreme far north of Spain.The presence of hg V in Euskal Herra ranges from 11.7% in Guipuzcoa to 5.9% in Alva.There is no diagnostic evidence that hg V expanded northward from a Basque refugium after the last glacial era.Although the highest frequency of mtDNa hg H has been recorded in northern most Iberia,diversity values are considerably higher in NE Iberia.However,overall H1 has a greater diversity in the Near East than in Iberia.The rare subgroup H1t appears to be restricted to the Iberian peninsula,where H3 is most abundant,but not the most diverse [O Garcia,2011].
C Capelli [2003] reported that the Basques Y chromosome hgs tend to cluster with those of Ireland [Castlerea],Cornwall, and western Wales [eg:Haveford West,Llangfini],which can be attributed to the northward expansion of Iberians from NW coastal communities in Spain after the Younger Dryas [ca 9700 BCE].The close genetic link between the Basques and Castlerea,Ireland,infers minimal "retained admixture" from the expansion of Neolithic male agriculturalists [ibid].
S Oppenheimer [2006] used Y chromosome SRR markers to reconstruct ancient population movements.Hg R was probably introduced to central Europe by the Gravettians from the Eastern Russian plains ca 30Ka ago..Today hg R1b and its 16 descendant lineages have an 86% frequency among the Basques.Some of these lineages expanded north from Iberia refugia after the Younger Dryas.The Y chromosome I1b2 followed a northern Mediterranean coastal route to the west.It contributed 41% to the present day Sardinian male gene pool,but only 6% to the Basques,who should have easily assimilated a relatively low percentage of newcomers.There has been a lower exchange of Y chromosomes between the western Basques and their neighbours,when compared to the degree of admixture to the east,where Basques and Aragon share the Ebro valley [M Iriande, 2003].Basque females were less restricted in their choice of partners [M Brian,2004].Galician G,M, and KM immunoglobian allotype allele frequencies indicate significantly large statistical variations between the Galicians and the Basques.The Guipuzean Basques are the most distinct [R Calderon,2007].
HLA hgs DR3-DQ2.5 and DR7-DO22 are common among the Basques and some west African populations.A29-Cw16-B44 occurs in west Africa,with a peak frequency among the Basques and a peak diversity in the Pasiegas valley.There appears to have been a time lag between the arrival of this genetic lineage in Iberia and its northward expansion,which could infer a slow dispersal through Iberia from north Africa prior to the Moor era.It has an appreciable component in the Pasiegas valley,which could be the site of recombination.This distinct feature suggests ancient admixture. There is A29-Cw16 disequilibrium among the Irish [P Deitiker,2008].
Collation of the available genetic data suggest, that the inhabitants of the present Basques Lands immediately after the end of the Younger Dryas ca 9700 BCE are the distant ancestors of the present day Basques.However subsequent influxes of newcomers over the interim have diluted and changed the original gene pool.Cantabria has closer affinities to the proximal Vizcayan Basque people than to the inhabitants of the other Basque provinces.Most mtDNA studies suggest that the modern Iberians share a common genetic background with other Europeans,but are distinct from Europeans.In the Basque Lands this distinction is even more apparent,because the Basques tended to evolve in discrete genetic clusters,with restricted local expansions. The Basques identity is essentially determined by their unique language. Future genetic studies may contribute to resolving some of the Basques enigmas.
About the Author
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A good mind. A keen sense of knowing what was needed, but wasn't available that technology could provide. Very good record keeping, organizational skills and process. And in his own words - 99% perspiration - Hard work.
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