L. Lasić et al.: Population-Genetic Studies in , Coll.Coll. Antropol. Antropol. 40 40 (2016) (2016) 2: 2: 145–149 145–149 Review

HHistoricalistorical OverviewOverview ofof thethe HumanHuman Population-Population- GGeneticenetic StudiesStudies inin BBosniaosnia andand Herzegovina:Herzegovina: SSmallmall Country,Country, GGreatreat DDiversityiversity

LLejlaejla LasiLasić, JasminaJasmina HindijaHindija Čaakar,kar, GGabrijelaabrijela RRadosavljeviadosavljević, BBelmaelma KKalamujialamujić aandnd NarisNaris PojskiPojskić

University of Sarajevo, Institute for Genetic Engineering and Biotechnology, Sarajevo, Bosnia and Herzegovina

AABSTRACTB S T R A C T

Modern Bosnia and Herzegovina is a multinational and multi-religious country, situated in the western part of the Balkan Peninsula in South-eastern Europe. According to recent archaeological fi ndings, Bosnia and Herzegovina has been occupied by modern humans since the Palaeolithic period. The structure of Bosnia-Herzegovina’s human populations is very complex and specifi c, due to which it is interesting for various population-genetic surveys. The population of Bos- nia and Herzegovina has been the focus of bio-anthropological and population genetics studies since the 19th century. The fi rst known bio-anthropological analyses of Bosnia-Herzegovina population were primarily based on the observation of some phenotypic traits. Later examinations included cytogenetic and DNA based molecular markers. The results of all studies which have been done up to date showed no accented genetic difference among the populations (based on geo- graphical regions) with quite high diversity within them. Human population of Bosnia and Herzegovina is closely related to other populations in the Balkans. However, there are still many interesting features hidden within the existing diver- sity of local human populations that are still waiting to be discovered and described.

Key words: human populations, population genetic studies, molecular markers, Bosnia and Herzegovina

IIntroductionntroduction

Due to its position and complex historical events, mod- Greeks to their northern neighbours, but it is not clear if ern Bosnia and Herzegovina is multi-ethnic and multi- they had been linguistically and culturally homogeneous religious country with lots of smaller, isolated populations. or ever collectively regarded themselves as »Illyrians«4. It has always been the place where different civilisations Even though the origin of Illyrians has been discussed would collide, which resulted in numerous migrations and through history, it is believed that Illyrian tribes had been constant mixing between various populations. All of this settled continuously in this area between the Bronze and makes it very interesting area for population-genetics succeeding Iron Age5. studies. From the end of 3rd century BC Romans began to Recent archaeological studies from northern Bosnia emerge in this area1. They haven’t conquered Illyrians at indicate that fi rst populations had inhabited this area in once, but rather it was the beginning of one hundred year Palaeolithic period, ~100 000 years ago and they belonged long war, thence on, this area was controlled by Romans to Homo neanderthalensia1. The earliest credible evidence for more than half a millenium6. Although the conquered of Homo sapiens in Europe is dating ~45 000 years ago. natives belonged to one category and Roman settlers to They apparently represent a population movement into other1, during that time Latin speaking settlers from all the Balkans during a warm climate interval2. For the next over the Empire settled among the Illyrians and Roman period, which lasted for tens of thousands of years, there soldiers were encouraged to retire in the region7. was no long absence of life in this area1. The ancient Bosnia and Herzegovina populations were In the early Bronze Age around 4th century BC, this introduced to Christian faith rather early. The earliest region was populated by the Illyrians, which established literature data on the presence of new faith in this area probably the fi rst known civilization in this part of Eu- are dating back to the 4th century AD. It is believed that rope3. The name »Illyrians« was given by the ancient the Slavs received the Christianity from indigenous, rath-

Received for publication June 21, 2016

149 L. Lasić et al.: Population-Genetic Studies in Bosnia and Herzegovina, Coll. Antropol. 40 (2016) 2: 145–149 er than foreign missionaries1. Following history, including little fi nger, digital index, tongue rolling, distal and prox- two mayor events along with the number of minor his- imal extensibility of the thumb36. In general, in the group torical episodes, laid the foundations for what is contem- of local populations studied, relatively low levels of ge- porary structure of B&H human population. The fi rst one netic heterogeneity and differential genetic specifi city is the large migration of people during 6th and 7th centu- were found. The effects of genetic drift and the high degree ries, which led to the fi nal collapse of Roman Empire and of propagational isolation were considered the most pos- moved different Gothic and Slavic clans into the area. The sible causes of such fi ndings. second one, which greatly infl uenced the region of today Research on phenotypic systems has continued into the Bosnia and Herzegovina, is the expansion of Ottoman 21st century. Genetic structure of both pre-war and post- Empire into this part of Balkan in the fi fteen century and war populations in Bosnia and Herzegovina were com- so caused numerous migrations during the following half pared based on the analysis of ten qualitative traits. Large 8 of millennium . migration has been taken as a turning point and their Last century was not less interesting regarding his- effects on the structure of the small populations were ob- torical events in this area. Two world wars and war in served41. Bosnia and Herzegovina between 1992 and 1995 launched Simultaneous monitoring of genetic diversity of a large many migrations including those within today’s country number of carefully selected phenotypic characteristics in 9 border . population-genetic studies may result in signifi cant infor- The historical events left a big mark on the structure mation about the micro-changes in the genetic structure of today B&H population and made it attractive for popu- of observed populations42. lation-genetic surveys. This paper gives a short historical review of previously published studies with new data of the latest population-genetic surveys of different levels MMolecularolecular geneticgenetic markersmarkers usedused fforor hhumanuman and approaches. ppopulationopulation geneticgenetic sstudiestudies inin B&HB&H

After a pause which took place during 1992–2000, re- PPreviousrevious bbio-anthropologicalio-anthropological sstudiestudies inin B&HB&H search process on local human populations in B&H with- hhumanuman ppopulationopulation in the context of its demographic structure, ethnic and cultural milieu has been continued. Revitalisation and The fi rst known bio-anthropological analyses of the modernisation of the capacities for population-genetic re- population conducted by the Austro-Hungarian army doc- search enabled the application of highly sophisticated ge- tor can be considered an initial phase in determining ge- netic markers for detection of human population variation. netic structure10. In 1887, Austro-Hungarian doctor Him- The main objectives of those studies were to determine the mel examined the specifi c phenotypic characteristics diversity of local human populations, especially isolated (average height, weight, eye colour, hair colour, and other ones, as well as three main ethnic groups (Bosnian Bos- anthropological – body measures) in a population of Bos- niaks, Bosnian Serbs and Bosnian Croats). 11 nian soldiers, members of the Austrian army . Later, The fi rst molecular genetic analysis of Bosnian popula- Weisbach (1895, 1905) conducted tests of phenotypic tion included an analysis of autosomal STR loci as well as markers in the population of Bosnia and Herzegovina and analysis of HV1 and HV2 region of mitochondrial DNA in was the fi rst to make a comparison of the observed pheno- isolated populations of rural areas43–47. The results of typic systems within the three constituent nations of Bos- these studies indicated that genetic variation is not dimin- 12,13 nia and Herzegovina . However, the fi rst exact results ished in these isolated populations and analysis of genet- of human population-genetic studies were obtained in ic distance obtained from DNA typing showed consistency 1930’s using analysis of frequencies of blood with initial hypothesis based on ecological, geographical, 14,15 group systems . social and cultural factors. th During the 20 century, in the period 1960–1980, in- In 2005, Marjanovic et al. published a paper on 28 Y- tensive efforts were made to study the diversity of morpho- chromosome biallelic markers within 256 males (90 logically complex phenotypic systems of qualitative and Croats, 81 Serbs and 85 Bosniaks) from Bosnia and quantitative traits within the local, regional and ethni- Herzegovina, with the aim of providing new clues about cally defi ned human populations in Bosnia and Herze- their origin and the ancient events of fl ow which have 16–33 govina . infl uenced this area located in the heart of Europe48. High Period between 1980 and 1990 was the time of intense frequencies of hg I-P37 were observed in all three ethnic research34–40. More sophisticated studying approach has groups. The highest frequencies were observed in the been employed, such as the analysis of genealogy and ge- population of Croats (~71%) and could be partially attrib- netic distance of 7100 included subjects. Data on overall uted to genetic drift and founding events on this ethnic 15 morphological, biochemical and physiological markers group. High frequencies were also observed in the Bos- were obtained: ABO blood groups, ABH antigen secretion, niaks (~44%) and Serbs (~31%). This shows that different PTC tasting, red colour vision, green colour vision, ear ethnic groups in Bosnia and Herzegovina share a large lobe type, fi ssured tongue, chin dimple, midphalangeal subset of their paternal lineages, affected by a major de- hairiness, nail form, position of distal phalange of the mographic event, the post-LGM expansion. A population

150 L. Lasić et al.: Population-Genetic Studies in Bosnia and Herzegovina, Coll. Antropol. 40 (2016) 2: 145–149 with a high frequency of I-P37 from one of the refuges, logroups: H, I, J, K, T, U, V, W or X. The most frequent located possibly in the Balkans, played a great role in the haplogroup in the analysed population was the haplogroup peopling of Bosnia and Herzegovina and surrounding ar- H (52.65%) which, due to its increased frequency, repre- eas. sents a marking haplogroup of the population of North- Y STR marker analysis was also conducted on 100 un- eastern Bosnia. The results of intergroup genetic analysis related male individuals from different regions of Bosnia showed that Bosnian-Herzegovinian population is geneti- and Herzegovina. Distribution of frequencies for 12 cally closer to previously studied populations of Herzego- Y-chromosomal STR was analyzed. Eighy-one different vinians (part of Bosnia and Herzegovina), Slovenians and Y-STR haplotypes were detected: 69 of them were unique, Croats in relation to other neighbouring populations lo- 7 appeared twice, 4 appeared three and only 1 fi ve times49. cated in South-eastern Europe. This study also suggests Population genetic research of representative sample that population genetic structure of Tuzla region is domi- of Bosnians and Herzegovinians were continued in 200650. nated by mutations that are classifi ed as »Palaeolithic«. Allelic frequencies for the 15 STR loci included in the Pow- These mutations were probably brought to the area of erPlex 16 kit were obtained from a multi-ethnic sample of North-eastern Bosnia through waves of prehistoric and 100 unrelated individuals born in Bosnia and Herzegovi- historic migrations, but the impact of any pre-Neolithic, na. The authors compared B&H data with data obtained Neolithic or some »later« migrations, with a slightly lower from geographically closer European populations in order contribution to the genetic structure of this population, 54 to put molecular genetic diversity of B&H into regional also can-not be neglected . and European context. In 2014, population genetic study focusing on mtDNA 55 Also, complex studies involving the application of var- haplogroups in Bosniaks was done . This study included ious molecular genetic markers to study the genetic struc- 227 samples from different geographical regions across ture of human populations in Bosnia and Herzegovina Bosnia and Herzegovina (Eastern Bosnia, Middle Bosnia, have been performed51. In order to investigate concordance Central Bosnia, Sarajevo region, Central Herzegovina, of different genetic markers, Pojskic et al. used 10 Alu loci, Eastern Herzegovina, and Krajina). This research has 15 autosomal STR loci and 6 blood groups (genotyped us- shown somewhat different frequency distributions of the ing PCR-RFLP and PCR-SSP methods). Also, polymor- observed haplogroups in the analysed regions. Statisti- phism of 10 phenotypic traits have been estimated within cally signifi cant difference was not recorded, meaning 300 individuals in human populations placed in 10 regions that the geographical distribution has no differentiation across Bosnia and Herzegovina. The main goal was to character in the population of Bosniaks in Bosnia and compare validity of human population genetic analyses Herzegovina. Since no genetic differences between the implementing different type of genetic markers. Modest seven observed regions (populations) were obeserved, the correlation between results of population genetic analyses conclusion is that the distribution of mtDNA haplogroups based on different markers has been detected, especially is not geographically conditioned. This research has con- between biallelic markers. Strong and clear correlation fi rmed the assumption based on the previous human between biallelic and multialleic markers has not been population genetic studies in Bosnia and Herzegovina that detected52. the Bosnian population (hence the Bosniak population) is Extensive study based on molecular-genetic typing of specifi c, relatively heterogeneous, without a signifi cant various blood group systems had been done in 201053. Six differentiation from the characteristics of the European blood group systems (Rh, MN, Duffy, Kidd, Kell and Lu- gene pool. theran) were investigated among three ethnical groups as Primorac56 analysed 180 samples from unrelated vol- well as ten regional subpopulations of Bosnia and Herze- unteers in Herzegovina. Populations were divided into govina, with some of these blood group systems typed for three regions according to geographical characteristics: the fi rst time in the population of Bosnia and Herzegovi- Central Herzegovina with Neum, Eastern Herzegovina na. This study showed that there is no accented differen- and Western Herzegovina, as well as three ethnic groups: tiation between geographically defi ned populations in Bosniaks, Bosnian Croats and Bosnian Serbs. Total of Bosnia and Herzegovina, and there are no signifi cant ge- nine Western-Eurasian haplogroups (i.e. H, I, J, K, T, U, netic differences between the three main ethnic groups. V, W and X) were evenly distributed in three geographical These results are in agreement with results of previous regions. In relation to ethnic groups, Bosniaks displayed studies, which indicates that three ethnical subpopula- a signifi cant increase in the frequency of haplogroup H tions of Bosnia and Herzegovina belong to the same gene while Bosnian Serbs showed signifi cantly increased fre- pool and have the same origin. quency of haplogroup U. The difference in the frequency The mitochondrial DNA (mtDNA) polymorphisms, de- can be explained as the result of genetic drift and in- termined in terms of mtDNA haplogroups, were analysed creased emigration of the population in this area. His- in many studies of B&H populations. One of these studies torically, this difference can also be explained by the mild included a total sample of 245 unrelated individuals from climate which enabled survival of the tribes after end of the area of North-eastern Bosnia (also known as Tuzla the last Ice Age and by an interesting history of Herze- region). The study revealed that 95.51% of the analyzed govina which was independent until the arrival of the Ot- individuals belonged to the typical Western-Eurasian hap- toman Empire. The frequencies of haplogroups in total

151 L. Lasić et al.: Population-Genetic Studies in Bosnia and Herzegovina, Coll. Antropol. 40 (2016) 2: 145–149

Herzegovinian population were in accordance with Euro- studied Western Balkan populations revealed their ge- pean average56. netic closeness regardless of a genetic system inspected, In 2013, Pojskic et al. published a paper on Alu poly- in particular among the Slavic speakers. In a more gen- morphisms in 10 geographical regions across Bosnia and eral perspective these results reveal clear genetic continu- Herzegovina (Krajina, Posavina, North-Eastern Bosnia, ity between the Near Eastern and European populations, Central Bosnia, Middle Bosnia, Western Herzegovina, lending further credence to extensive, likely multiple and Sarajevo Region, Eastern Bosnia, Central Herzegovina, possibly bidirectional ancient gene fl ows between the Near 58 Eastern Herzegovina). The sample comprised of 278 Bos- East and Europe, cutting through the Balkans . niaks, 134 Bosnian Croats and 94 Bosnian Serbs. Conclu- In this respect, Bosnia and Herzegovina represents a sion of this study was that the results are in agreement unique region and source of genetic information discov- with most previous human population studies in Bosnia ered previously, but also, yet to be discovered in future and Herzegovina, indicating that the three populations of studies. Bosnia and Herzegovina have the same genetic back- ground. Also, there was no statistically signifi cant dif- ferentiation among regional populations, pointing to the CConclusiononclusion absence of geographic infl uence. Therefore, authors con- Due to the complex historical background and migra- cluded that the whole population of Bosnia and Herze- tion events in Balkan region, overall B&H population govina is homogeneous considering Alu polymorphisms shows quiet high degree of diversity between its regional and there is no scientifi c basis for its genetic stratifi ca- subpopulations, considering specifi c genetic characteris- 57 tion . tics of isolated populations. There is no major genetic dif- Another great study involving the investigation of poly- ference between three main ethnic groups in Bosnia and morphisms of autosomal SNP markers in human popula- Herzegovina (Bosniaks, Bosnian Serbs, and Bosnian Cro- tions of the Western Balkan Peninsula was published in ats) indicating that they represent the same gene pool. 201458. In this study, the genetic variation of 660000 au- Comparison of B&H population with neighbouring and tosomal SNPs, mitochondrial DNA markers and non-re- other European populations shows no signifi cant genetic combining part of Y chromosome was assessed in three difference between them. These fi ndings clearly suggest ethnic groups from Bosnia and Herzegovina (Bosniaks, that all three ethnic groups and total population of B&H Bosnian Croats and Bosnian Serbs) and four other Slavic- belong to the European gene pool. This conclusion sug- speaking Western Balkan populations: Serbians, Croa- gests that any differences between three main ethnic tians, Macedonians from the republic of Macedonia, Mon- groups in Bosnia and Herzegovina do not have the ge- tenegrins, and Albanian-speaking Kosovars. Comparison netic background but are primarily of social and cultural of the variation within autosomal and haploid data sets of manner.

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L. Lasić

University of Sarajevo, Institute for Genetic Engineering and Biotechnology, Zmaja od Bosne 8, 71000 Sarajevo, Bosnia and Herzegovina e-mail: [email protected]

PPOVIJESNIOVIJESNI PPREGLEDREGLED GGENETSKIHENETSKIH IISTRAŽIVANJASTRAŽIVANJA PPOPULACIJEOPULACIJE BBOSNEOSNE I HHERCEGOVINE:ERCEGOVINE: MMALAALA ZZEMLJA,EMLJA, VVELIKAELIKA RRAZNOLIKOSTAZNOLIKOST

SSAŽETAKA Ž E T A K

Moderna Bosna i Hercegovina je višenacionalna i multi-religiozna zemlja, nalazi se na zapadnom dijelu balkanskog poluotoka u jugoistočnoj Europi. Prema posljednjim arheološkim nalazima, Bosna i Hercegovina je okupirana od strane modernih ljudi od razdoblja paleolitika. Struktura ljudske populacije Bosne i Hercegovine je vrlo složena i specifi čna, zbog čega je zanimljiva za razna populacijsko-genetička istraživanja. Stanovništvo Bosne i Hercegovine je u fokusu bio- antropoloških i populacijskih (genetika) studija još od 19. stoljeća. Prva poznata bio-antropološka analiza bosansko- hercegovačkog tanovništva prvenstveno se temeljila na promatranju nekih fenotipskih osobina. Kasniji pregledi su uključili citogenetske i DNA molekularnih markera u istraživanja. Rezultati svih studija koje su do sada napravljene do sada nisu pokazale značajnu genetsku razliku među populacijama (na temelju geografskih regija) s vrlo visokom raznolikošću unutar njih. Ljudsko stanovništvo Bosne i Hercegovine je usko povezano s drugim populacijama na Bal- kanu. Međutim, još uvijek postoje mnoge zanimljive značajke skrivene unutar postojeće raznolikosti lokalnih popu- lacija koje još uvijek čekaju da budu otkrivene i opisane.

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