H FENETIKH IYZTAZHTON KATOIKftN THI EAAAAAZ

Tjir||u.aTO(; BioXoyiaq tou ApiaxoieXeiou naveTuarrmioD

THE GENETIC COMPOSITIONOF THE INHABITANTS OF GREECE

Constantinos Triantaphv lidisProfessor of Genetics

School of Biology Aristotle University of Thessaloniki

OEIXAAONIKH 1993THESSALONIKI 1993

Introduction to the speech «The gene-tic composition of the inhabitans ofGreece» by Dr Penelope Minasidou-Mavridou, President of the MedicalWomen Association of North Greece

When a few years ago, the Frenchhistorian Dyrosel wrote in one of hisbooks and commented in interviewsthat «the Greeks come from the Turksand that the entry of Greece into theEuropean Community was a mistake»we were all deeply enraged by the-se unfounded and untrue historical re-marks.

Of course we all, not of the fa-culty of Biology, did not know thatapart from historical data, we couldalso prove Dyrosel and some other socalled friends wrong based on scienti-fic research.

We did not know that in theSchool of Biology of the AristotleUniversity of Thessaloniki, the Pro-

fessor of Genetics Mr C. Triantaphyl-lidis and his associates had a scienti-fic publication about our genetic ori-gin.

When we read in the press the ar-ticle-protest of Dr. C. Triantaphylli-

dis, which also referred to more pu-blications from foreign Universitiesdealing with the same subject, likethe one of Professor Piazza and asso-ciates from the University of Torine,we felt proud of this not only serious,but also valuable publication of ouruniversity and we as a MedicalWomen Association of North Greece,concluded that the results of this scie-ntific research be known to all usGreeks and consequently to the wholeworld.

Every Greek, wheever he may be,is interested in discovering his rootsand this not in order to feel moreGreek, of this he is certain, but to beable to refute not only historically butalso scientifically antihellenic opi-nions of some people and others whohave designs on our Greek heritage.

The Medical Women Associationof North Greece would like to thankDr C. Triantaphyllidis who so readilyanswered our request that this speech

be given and that he takes care of thepublication of this leaflet which is asummary of the subject «The geneticcomposition of the inhabitants ofGreece» written in Greek and Englishto be sent all over Greece and even allaround the world, so that everyone re-alises how Greek our genealogy is,genetically speaking as well. TheMedical Women Association of NorthGreece owe a big thanks to the Presi-dent of the Medical Association ofThessaloniki Dr N. Aggelidis, theGeneral Secretary Dr G. Xatzedemosand the board who so eagerly coope-rated in the organization of the speech

which took place in the Chamber ofCommerce on March 3rd 1993 andwho were the granters of the capitalfor the first edition of the leaflet titled« The genetic composition of the inha-bit an ts ofGreece».

We would also like to thank thecolleaque Dr C. Stasinopoulos, theEditor of the Journal of Medicine Ga-linos and Mr Pan. Papageorgiou, Ma-nager Editor of the Journal, for thepublication of the speech in the Jour-nal and the republication of the lea-flet so that it continues to be sent to allthe Libraries of the World and tomany Institutions.

The first inhabitants of Greece,the Prohellens, arrived in Greek terri-tory at about 8.000 to 6.000 BC. La-ter, they developed in Crete and theGreek islands those marvellous civi-lizations which are known as Minoanand Cycladic. The Hellenes (= Gre-eks) arrived on the Greek mainland insuccesive waves during the end of theBronze Age (2.500 to 1.000 BC).Their arrival has been establischedthrough archaelogical excavations,linguistic records and literaly sources.

The next three centuries(1.100-800 BC) saw an increase ofGreece's population, while Hellenic(=Greek) civilization and culture wereadvanced. Meanwhile, important co-lonizing enterprises started, whichspread the Hellenes futher afield from

the Greek mainland. By the 8th cent-ury there were Hellenic settlements inmodern Bulgaria and Rumania, on thecoasts of the Black sea and of Asia Mi-nor. The Greeks also colonized partsof Sicily and Southern Italy, whichcame to be known as Magna Graecia(Great Greece) and many points alongthe French, Spanish and North Africacoasts. This Early colonization wasthe beginning of a process of Heileni-zation of the indigenous populations.In the 4th century Alexander theGreat (356-323 BC) extended greekcolonization and civilization evenfurther, to include much of Egypt andthe Middle East and to reach even In-dia. Colonization and hellenizationcontinued for more than two hundredyears and was maintained during the

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Roman imperial rule. During the By-zantine period Anatolia, with the ma-jor city in the interior of Cappado-chia, was one of the most importantregions with a Greek population out-side of Greece itself.

The study of the inhabitants ofHellas (= Greece) is usually based onhistorical, linguistic and archaeologi-cal records. This article aims to inve-stigate the same subject from a diffe-rent angle, since it is based on Biome-diqal - genetical - data.

It is worth mentioning that it wasin Greece that the first study of bloodgroup frequencies was ever made. Itwas carried out by Professor and MrsHirszfeld (1919) when they were wor-king in Salonika during the FirstWorld War and had the opportunityof studying large numbers of soldiersfrome different countries. Five hun-dred Greeks were included in theirsamples. Since then, and up to thepresent day, a number of investiga-tors, mostly greeks, have studied thefrequency of alleles of genes that codefor blood groups, enzyme and proteinpolymorphisms, among the Greekpeople. Several authors have studiedlocal or isolated Greek populations.This aricle is an attempt to present thegenetic structure of the inhabitants ofmodern Greece, as well as the geneticcomposition of isolated Greek popu-lations for which enough geneticaidata are available. These populationsare the refugees from Asia Minor, theendogamic and nomadic populationsof Sarakatsans and the Gypsies.

The history of a language - itsevolution - can be traced from the

simiiiarities and differences of livinglanguages and their literaly fossils.Such linguistic evidence can be usedto trace the history and relationshipsof populations. In the same way, sha-red genes can be used to trace patternsof genetic relationships among humanpopulations. Because of the abovementioned immigration of Greeks allover the Mediterranean region, itseems interesting to investigate the ge-netic relationships between the inha-bitants of Greece and the inhabitantsof the Mediterranean countries as wellas between the inhabitants of the Me-diterranean countries and the threemajor human reces*.

Previously, the genetic relation-ships between the human populationswere studied mainly by using mor-phological characters. In recent years,however, blood groups and other pro-tein and enzyme polymorphism(1976) have been studied in humanpopulations. The study of bloodgroups and other genetic polymor-phisms for this type of work is moreattractive from, say craniometric data,because of two reasons: their lack ofenvironmental lability and their si-mple genetic determination. Today,for the same purpose, the scientistsare studying the polymorphism of themitochondrial and nuclear genetic

* The data from population genetic studiessupport the notion that there are no humanraces, but only human populations. So, theterm race, in this article, is used only for theconvenience of presentation of the results.

meterial (DNA), as well as the diffe-rences in the order of nucleotides ofparticular segments of nuclear andmitochondrial DNA.

A. The genetic relationship betweenthe inhabitants of Greece and theinhabitants of other Mediterraneancountries

The Mediterranean region is divi-ded into many countries. The popula-tion of this area is quite diverse consi-sting of people of different nationali-ty, language and religion. Hence, it isworthwhile to determine the geneticrelationships among the inhabitantsof the Mediterranean countries as wellas the genetic distances between theinhabitants of 9 Mediterranean coun-tries and the 3 major human races.For that purpose data on at least 16genetic loci from nine Mediterraneancountries were used (1,2).

The genetic markers that wereincluded were blood groups, ie ABO,MNSs, Rhesus, etc, protein loci:ACPH, AK, Hp, etc, from the follo-wing countries: France, Spain, Italy,Greece, Turkey, Libya, Israel, Egypt,Algeria. We also decided to includeBulgaria in the study. The majority ofthe data for Caucasoids and Mongo-loids were taken from US Caucasiansand from Japanese, respectively. Thedata for Negroids were taken fromAfricans as well as from African Ame-ricans.

Using the above mentioned datathe genetic distances among the inha-bitants of the nine Mediterraneancountries were computed. Using these

estimates, a dendrogram (Fig 1) hasbeen constructed. The smallest ge-netic distances (or the greater gene-tic similarity) occur between French- Spaniards, Greeks - Italians, French- Italians and French - Greeks, whe-reas the greatest distances are bet-ween the Algerians and the inha-bitants of the other Mediterraneancountries. The populations of theNorth Mediterranean countries wereclustered together, whereas this wasnot the case for South Mediterrane-ans.

Thus one can conclude that thegenetic distance of the inhabitants ofGreece from the populations of othercountries decreases in the followingorder: Italy, France and then Tur-key. These findings therefore stron-gly contradict the suggestion of theFrench historian Z Dyrozel that theGreeks are descentants of Turks.

B. The genetic distances between theinhabitants of nine Mediterraneancountries and the three major humanraces

Anthropologically, the Mediter-raneans belong to the Caucasoidgroup, although some Negroid andMongoloig admixtures have to be ta-ken into account. The genetic analy-sis demonstrates certain features ofinterest.1. The genetic distance of the popu-

lations of North Mediterraneancountries from Caucasoids decre-ases from the East to the West.

2. The genetic distance from Mongo-

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£Spam

France

Italy

Greece

Turkey

Jews

Libya

Egypt-

Algeria

Fig. 1. Dendogram of the genetic relationships among the inhabitants of nine Mediterraneancountries.

loids increases from the East to theWest for the populations of Northand South Mediterranean coun-tries.

3. The genetic distances betweenBulgarians, Turks, Greeks, Ita-lians, Spaniards, French, Israelisand Caucasians are smaller thanthat between the inhabitants ofeach country and the Negroids.

4. Libyans and Egyptians are closer toNegroids than to Mongoloids andAlgerians are closer to Negroidsthan to Caucasoids. This is proba-bly caused by Negroid immigrationinto these countries.

C. Comparison between the geneticcomposition of the inhabitants ofGreece and Italy

Our results show, that the inhabi-tants of Greece and Italy are gentical-ly very close. It is worth mentioningthat Professor A Piazza et al (3) of the

Dipartimento of Genetica of theUniversity of Torino has reached thesame conclusion. For the reconstru-ction of the genetic history of theinhabitants of Italy his Laboratory hasused many genetic loci (blood groupsand protein polymorphisms), as wellas historic and linguistic records.

By 400 BC the population of Italywas about 4 million people. TheGreek colonies of Sicily alone ac-counted for 1,5 million .people, ofwhich more than 10% (about 200.000)were of Greek origin. To these Greekinhabitants of Sicily another 100.000Greek colonizers in the Italian penin-sula, may be added so that before theRoman period, one in every 10-13inhabitants of Italy was Greek. Even ifthese numbers must be taken withcaution, their order of magnitude mi-ght give an explanation for the remar-kable differentation of Magna Greacia(South Italy and Sicily) from the restof the Italian mainland, as well as for

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the great genetic similarity of the Ita-lian southern regions with the inhabi-tants of modern Greece.

The loci used for such studies aremainly polymorphic. Thus, the cal-culated genetic distances tend to be anoverestimate. Keeping this fact inmind, one can estimate the time sincedivergence between the Greeks, theItalians, the French and the Spa-niards. These populations appear tohave diverged from the Caucasoidline about 8.000 years ago or in themesolithic period. This rough estima-te based on evidence of molecular e-volution aggrees quite well with sug-gestions based on anthropometric me-asurements and archaeological evi-dence that North Mediterranean po-pulations migrated from a commonroot in Central Europe.D. Comparison between the geneticcomposition of the inhabitants ofGreece and Bulgary

The genetic polymorphisms havebeen studied in only one other Balkanpopulation, that of Bulgaria. Histori-cally, the two populations differ con-siderably. The Greeks migrated toSoutheastern Europe at about 1500to 1100 BC. The Bulgarians, a Mon-golic tribe migrated to what is to-day Bulgaria during the 8th centuryAD. There they were absorbed by lo-cal Slav populations.

The estimated genetic distancesbetween the inhabitants of Bulgariaand the four Mediterranean countriesstudied (Turkey, Greece, Italy andFrance) indicate that: The geneticcomposition of the inhabitants of Bul-

garia is closer to that of the inhabi-tants first of Italy, next of Turkey,and then France, whereas the greatestgenetic distance was found betweenthe inhabitants of Greece and Bulga-ria.

The gene frequencies of five poly-morphic systems (AK, PGM, ADA,ACPH, 6-PGD) in the Greek andBulgarian populations were comparedin 1975, by Stamatoyannopoulos et al(4) of the Division of Medical Gene-tics, University of Washington Seat-tle, USA. Only one system, AK, hassimilar frequencies. Thus, among thepolymorphic systems so far compa-red, statistically significant differen-ces were observed in four systems.Therefore, the distribution pattern ofthe gene frequencies differs considera-bly between the two populations.Furthermore, the pattern of gene fre-quencies in the Bulgarian populationresembles to that of Asiatic popula-tions.

The existing data also sheds somelight on the debate between Greek andSlavic historians regarding the ethnicorigin of the population inhabitingGreek Macedonia. On the gene gre-quencies of seven polymorphic sy-stems (AK, PGM, ADA, ACPH,6-PGD, Hp and ABO) studied in theinhabitants of Greek Macedonia andof Bulgaria (4,5) statistically signifi-cant differences between them wereobserved in four systems. These fin-dings thus disprove the theories ofSlav historians who claim that theinhabitants of Greek Macedonia areSlavs and not Greeks.

From the above mentioned pa-

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pers it can be concluded that theTurks and the Bulgarians are geneti-cally closer to the Mongloids amongall the inhabitants of the Mediterrane-an countries. These results supportthe notion of the origin of the Turksand Bulgarians from Central Asia anddistinguish them from the Greeks. Fi-nally, although there has been geneflow from East to West in the Mediter-ranean area, the inhabitants of Greeceare closer to the Italians and French,than to the Turks or to the Bulgarians.Thus, for religious, cultural and lin-guistic reasons there has been only li-mited gene flow between Greeks andTurks or Bulgarians.

E. The genetic composition of Greekrefugees from Asia Minor

At the beginning of the twentiethcentury, some 2.000.000 Greeks wereliving outside Greece in Asia Minor.The presence of this large Greek po-pulation had been established on theWest coasts of Asia Minor by 800 BCand spread gradually northward alongthe Blac Sea coast. These Greek po-pulations maintained their Greek ide-ntity throughout the centuries, evenunder the Otoman rule. They re-mained separated from the neighbo-uring populations by religious andlanguage boundaries. The areas ofPontus and Cappadochia especiallywere centers of Greek culture.

The first world war was followedby the Greek - Turkish war. In 1923 atreaty (Treaty of Lausanne) was si-gned between Greece and Turkey forthe exchange of minorities. Before and

after that Treaty a considerablenumber of Greek Christian refugeesfrom Asia Minor, Pontus, Constanti-nople and other areas emigrated toGreece. Of these, 700.000 settled inMacedonia and West Thrace. About25% of the population of Macedoniawere, in 1928, of immigrant origin. Itis difficult now, more than two gene-rations later, to distinguish the auto-chthonous and immigrant popula-tions.

One of the 1047 communitieswhere refugees were settled was Platy.The village of Platy is situated in themiddle of Greek Macedonia in theprovince of Hemathia. The geneticcomposition of this population wasstudied by Tills et al (6). Of the 1558inhabitants in 1973, 1038 donatedblood for genetic tests. Blood grou-ping tests and electrophoretic analysiswere performed at the laboratory ofBiological Anthropology of the Bri-tish Museum of Natural History atLondon in collaboration with theGreek Red Cross. Tests were carriedout for ten blood group systems (ABO,MNSs, Rhesus, P j , Lutheran, Kell,Duffy, Kidd, Wright and Redin), threeplasma proteins (Haptoglobin, Tran-sferin and Haemoglobin), and 8 redcell enzymes (ACPH, AK, ADA, EsD,G6PD, PGM, 6-PGD and PHI). Thefindings from the Platy sample havebeen compared to those of Greekmainland, Turkey and Bulgaria.

When examining a population,such as the Platy population, systemby system it is difficult to concludewhether the particular population re-sembles either the A or the B popula-

tion. Theoretically, one must simulta-neously examine about ten genetic sy-stems. So, the genetic distance wascalculated (6) using data for the follo-wing 8 genetic systems: ABO, MNSs,Rhesus, Haptoglobin, Acid Phospha-tase, 6-phosphoglucomutase dehydro-genase, Adenyulate Kinase and Ade-nosive deaminase. These were theonly systems for which data wereavailable for all four populations.Using the appropriate mathematics,the genetic distances were calculatedamong the Platy sample, the main-land Greeks, the Turks and the Bulga-rians and the obtained results areshown in Fig 2. As can be seen, thedifferences between the four popula-tions are small, but the Platy sampleappears to be slightly closer in its rela-tionship to the Greeks than to theTurks. Furthermore, the refugees ofPlaty are closer to the Turks, thanto the Bulgarians.

The incidence of carriers of p-Mediterranean and sickle cell anae-mia as well as of the glucose-6-

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phosphate-dehydrogenase deficiencyhave been investigated in the sampleof 410 individuals from Pontus by Si-nakos et al (1975) at the 1st Clinic ofPathology at the Aristotle Universityof Thessaloniki, headed by professorD Valtis. Low frequency of G-6-PDdeficiencies (2%) and p-Mediterrane-an anaemia (3,7%), and only one car-rier of the Hb-S variant were found.The distribution of the moderate p-thalassaemia was heterogenous andcorrelated with the topography of ma-larious areas from where they had im-migrated in the past.

F. The genetic composition of GreekSarakatsans

The nomadic and primative po-pulations in various parts of the worldare experiencing social pressures andchanging their lifestyle by settlingalongside the urban populations. Ho-wever, there are still several popula-tion groups in Europe which providean excellent opportunity to study the

TOYPKIA - TURKEY

BOYAFAPIA - BULGARIA0-28

EAAAAA - GREECE

Fig. 2. Genetic distances among four populations.

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biological effects of such social chan-ges. In this respect, the endogamicpopulation of Sarakatsans is of greatinterest. Sarakatsans are nomadic she-pherds of continental Greece, whotoday number about 80.000 in Gree-ce. They live in the mountainous are-as of Thessaly, Macedonia, Epirus,Thrace and Sterea Hellas. Their sea-sonal migrations have led many ofthem across the border of Bulgaria(15.000), Albania, South Yugoslaviaand Turkey. They all speak solelyGreek, wherever they live (7,8). Theirparticular dialect contains trace ele-ments of ancient Greek. They have re-mained genetically isolated for manycenturies, probably due to the mo-untainous terrain and their culture,pattern of marriage and traditionalway of life. But during the last 20years, they have started to abandontheir nomadic way of life and to settlein villages or cities, changing theiroccupation and marrying outside thegroup. A process of isolation break-down is obvious. The genetic study ofthis group therefore is important.

Chatzimichali (9) gave evidencethat they are a population group whi-ch, in social organization, exemplifycertain prototypical elements ofGreek culture. In recent years Poulia-nos (10) using metrical and morpholo-gical traits suggested that the Saraka-tsans are local upper Paleolithicsurvivors and may be considered asthe most ancient population in Euro-pe. The genetic study of this grouptherefore is equally important as thatof the Basques in Spain and the Lappsin Scandinavia.

Blood samples were collectedfrom 232 unrelated individuals (11).The studied area comprised the wholearea of distribution of Sarakatsans inGreece. Part of the investigation wascarried out at the Department of Ge-netics, Development and MolecularBiology, Aristotle University of Thes-saloniki, Macedonia, Greece and partat the Division of Human Genetics,University of Newcastle Upon Type,UK with the aim of examining theirgenetic structure and estabishing theiraffinity to other populations of Euro-pe,

A random sample of 232 GreekSarakatsans has been studied for 10blood groups, 11 enzymic genetic sy-stems, sickle cell haemoglobin and forcolour blindness. The allelic frequen-cies of the 17 polymorphic loci havebeen compared with those of other sa-mples from the Greek mainland andother European populations. It seemsworthwhile to summarize the resultsof the comparison between the Sara-katsani population and other Greeksamples for the 17 polymorphic ge-nes. In comparison with geographical-ly neighbouring inhabitants, signifi-cant differences were detected in 7 outof 28 possible comparisons. In conc-lusion, the Sarakatsans tend reasona-bly well to resemble the rest of theGreek population.

The comparison with Europeanpopulations indicates that the Saraka-tsans have gene frequencies similar toother Mediterranean and Europeanpopulations. However, the monomor-phism of the Kell system, the low fre-quency of the GLO*1, AK*1 and

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ACP*B alleles and the high frequencyof the GPT*1, EsD*l and ACP*C al-leles, are noteworthy. Furthermore,the Sarakatsans are not a high riskgroup for G6PD dificiency, HbS and(3-thalassemia (12). The low incidenceof G6PD dificiency, abnormal hae-moglobin variants and p-thalassaemiaheterozypotes may be due to their lifein the mountain areas in the past cen-turies.

G. The genetic structure of GreekGypsies

The endogamic and nomadicpopulation of Atzigani, as the GreekGypsies are called, is estimated to be25-30.000 individuals, scatteredthroughout the country. It seems thatthey left India starting from the Xththrough to the Xllth centuries. Theylanded in Greece at the beginning ofthe 14th century. They have their owncustoms and traditions.

Bartsocas et al (13) have studiedthe genetic structure of the GreekGypsies at the Second Department ofPediatrids at the University of Athens.The investigation was carried out in200 individuals in five blood groupsystems (ABO, MN, Rhesus, Kell andDuffy) and two protein systems (hae-moglobin and ceruloplasmin). Fromthe obtained results the authors conc-luded that the ABO, Rhesus, MN andDuffy gene frequencies varied signifi-cantly from the figures obtained forthe Greek population. However, thereseems to be some similarity, especial-ly as concerns the ABO, Rhesus andDuffy blood groups, between the di-

stribution of the gene frequencies ofthe Greek Gypsies and today's inhabi-tants in the Punjab region of India andWestern Pakistan, the supposed re-gions of origin of the Gypsies. Theabove mentioned conclusion was alsosupported from the results of Sinakoset al (14). The Sinakos et al researchwas conducted in 557 individuals.Their research indicated a high fre-quency of G-6-PD dificiency (21,2%)and a considerable one of p-Thalassa-emia (8,6%). These high frequenciesmight be attributed to the fact thatthey have lived for many centuries inregions where malaria was endemic.Alternatively, the above mentionedhight frequencies of hereditary disea-ses, as well as the occurence ofrare hereditary disorders, could be at-tributed to the isolation and inbree-ding of the Gypsies.

Conclusion

The study of the language of ge-nes provides us with clues about theorigin and history of the inhabitants ofGreece. The above mentioned resultsconsistently show the great genetic si-milarity between the inhabitants ofGreece and the inhabitants of Italy.Especially the results indicate stronggenetic affinities between the inhabi-tants of Greece and those of southernparts of Italy. On the other hand, thereare great genetic distances betweenthe inhabitants of Greece and those ofTurkey and even greater betweenGreeks and Bulgarians. Thus, histori-cal facts are backed by genetical data.

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