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The Origin of Palestinians and Their Genetic Relatedness With Other Mediterranean Populations
Antonio Arnaiz-Villena, Nagah Elaiwa, Carlos Silvera, Ahmed Rostom, Juan Moscoso, Eduardo Gómez-Casado, Luis Allende, Pilar Varela, and Jorge Martínez-Laso
ABSTRACT: The genetic profile of Palestinians has, for the first time, been studied by using human leukocyte antigen (HLA) gene variability and haplotypes. The comparison with other Mediterranean populations by using neighbor-joining dendrograms and correspondence analyses reveal that Palestinians are genetically very close to Jews and other Middle East populations, including Turks (Anatolians), Lebanese, Egyptians, Armenians and Iranians. Archaeologic and genetic data support that both Jews and Palestinians came from the ancient Canaanites, who extensively mixed with Egyptians, Mesopotamian and Anatolian peoples in ancient times. Thus, PalestinianJewish rivalry is based in cultural and religious, but not in genetic, differences. The relatively close relatedness of both Jews and Palestinians to western Mediterranean populations reflects the continuous circum-Mediterranean cultural and gene flow that have occurred in prehistoric and historic times. This flow overtly contradicts the demic diffusion model of western Mediterranean populations substitution by agriculturalists coming from the Middle East in the Mesolithic-Neolithic transition. Human Immunology 62, 889-900 (2001). ã American Society for Histocompatibility and Immunogenetics, 2001. Published by Elsevier Sciece Inc. KEYWORDS: Palestinians; Macedonians; Greeks; Mediterraneans; Berbers; Moroccans; Albanians; Turks; Iranians; Egyptians; Jews; Lebanese.
INTRODUCTION The highly polymorphic human leukocyte antigen (HLA) system has been validated as useful for distinguishing and/or relating populations (and individuals) in many research studies since the first International HLA Anthropology Workshop  and in all the subsequent International Workshops. HLA gene frequencies correlate with geographically related populations. The existence or absence of gene flow among neighbouring ethnic groups may be assessed with the study of HLA
This work is dedicated to all Palestinians and Jews who are suffering war. From de Department of Immunology and Molecular Biology (A.A.-V., C.S, J.M., E.G.-C., L.A., P.V., J.M.-L.), H. 12 de Octubre, Universidad Complutense, Madrid, Spain, and Laboratories & Blood Bank (N.E., A.R.), El-Shifa Hospital, Gaza, Palestine. Address reprint request to Antonio Arnaiz-Villena, Departamento de Inmunología y Biología Molecular, H. 12 de Octubre, Universidad Complutense, Carretera Andalucía. 28041 Madrid, Spain; Tel: +34 (91) 390 8315; Fax: +34 (91) 390 8399; E-mail: email@example.com, Web address: http://chopo.pntic.mec.es/biolmol. Received May 16, 2001; accepted June 15, 2001.
frequencies and the corresponding genetic distances and haplotypes [2, 3]. Most ancient recorded inhabitants of Palestine are named Canaanites (3rd millennium BC or more ancient). They became urbanized and lived in city-states, one of which was Jericho. Palestine’s location at the center of routes linking three continents made it the meeting place for religious and cultural influences from Egypt, Syria, Mesopotamia, and Anatolia. During the second millennium BC, Egyptian hegemony and Canaanite autonomy were constantly challenged by such ethnically diverse invaders as the Amorites, Hittites, and Hurrians from Anatolia and the East. These invaders, however, were defeated by the Egyptians and absorbed by the Canaanites, who at that time may have numbered about 200,000. Egyptian power began to weaken, and new invaders or autochthonous people appeared or made themselves noticeable . The Israelites, a confederation of Hebrew tribes, finally defeated most of the Canaanites
Human Immunology 62, 889-900 (2001) ã American Society for Histocompatibility and Immunogenetics, 2001 Published by Elsevier Science Inc
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Nazareth and modern Tel-Aviv are also indicated . FIGURE 2 United Nations agreement for Palestine partition in 1947. a new iron technology probably was taken de novo by some autochthonous Canaanite tribes that acquired superiority. the Palestinians are nowadays thought to come from the Egyptian garrisons that were abandoned to their own fate on the Canaan land by 1200 years BC (Figure 1) and had to manage to construct or reinforce or rebuilt some ancient Canaanite city-states. The “sea people” contributed to the fall of the Anatolian Hittites and other Middle East people by 1200 BC and apparently seriously threatened Egypt . By 1000 BC. left to their own fate according to archaeological records . Jerusalem.890 A. it is doubtful that big amounts of people entered nowadays Anatolia and Palestine. The Philistines have been included among the “sea people” invaders. FIGURE 1 Canaan in ancient times. Philistines had established an independent state on the southern coast of Palestine and also controlled the Canaanite town of Jerusalem. Palestinian citystates may come from the remains of Egyptian garrisons. Israelites could also stem from autochthonous Canaanite tribes that were agglutinated by a group of people led by Moses to fight against other Canaanites. who was killed by King David according to the Bible. the Palestine. Gath may be placed more to the south because it has not been recognized in modern times. Ash- . and after warring with Philistines and other Canaanites. an Israelite state was founded by king Saul . including Philistines and finally set up ancient Israel . Palestinians (Philistines) city-states by 1100-1000 BC (black squares) Jericho was an ancient Canaanite city-state. However. together with the old autochthonous tribes . (1125 BC) but found the struggle with the Philistines (Palestinians) more difficult. Palestinians held five important city-states when the fighting with Israel began (after 1200 BC): Gaza. the ancient Palestinians might have come from Crete or its empire . Arnaiz-Villena et al. Goliath. In fact. Otherwise. come from this city. the Bible quotes that Palestinians may have come from Crete .
000.000.9]). 5. Nowadays.HLA Genes in Palestinians 891 FIGURE 3 Location of the studied populations and map of the present day Israel-Palestine. After several regional wars. Israel has taken more space and sized Jerusalem. Israel´s self-proclaimed independent in 1948 and started a war against Muslim Palestinians and other Muslim neighbouring countries. Palestinians were about 5.000 by 1947 when Palestine population amounted 2. Immigration rapidly increased the number of Jews (who had been several times led to Diaspora. most western European countries.000 [9-11]. Three kind of Palestinians. The present situation (April 2001) is unstable.000 at the beginning of the last decade. according to their status. deported. Ashod. to the European crusaders and finally to the Ottoman-Empire  since the XVI century. and massacred by ancient Iranians and Romans. The United Nations plan for Palestine partition in 1947 is illustrated in Figure 2 . They won several battles and the time that they were in control of all Canaan. They probably had an iron technology higher than Israelites. The Palestinian Autonomous Government has a limited rule in the Gaza Strip and West (Jordan) Bank. because they did not let Israelites to work as blacksmiths when they were ruling over Israelites [see Bible starting in Samuel 13 and 19. to Egyptian Muslim Mammeluks. Gaza became an important Christian center  and afterwards an important Islamic center because Palestine was converted to Islam by Arabic troops and priests by 700 AD . kelon.7].000 Jews present in Palestine by 1936 and 600.000 were Jewish. Palestine had 750.000. they might reach 7. In 1918 British led mixed Arab-British troops seized the region.000 inhabitants in 1919  and only 70. expelled. Ekron and Gath (Figure 1). may be defined as follows: (1) the inhabitants under an autonomous Palestinian government (about 30%) that controls very little of the West Jordan Bank . There were 400. Later.000 inhabitants. west to the Jordan river (about 100 years?) has been enough to name the land as Palestine until after World War II . as illustrated in Figure 3. Alexander the Great  surrendered Gaza after a long siege about 333 BC. Ancient Canaan (Palestine in Middle Age and modern times) has sequentially belonged to the Roman-Byzantine empire. and finally Hitler [8.
892 A. -B. In summary. Statistical analysis was performed with Arlequin v1.10]).) (inhabited by more than 1. Gaza) were used for HLA genotyping and phylogenetic calculations. and the Gaza strip is now a relatively or totally (when the ongoing conflict aggravates) secluded area. and (2) explain the historic basis of the present day Middle East conflict between Palestinians and other Muslim countries with Israelite Jews. Hardy-Weinberg equilibrium and the linkage disequilibrium between two alleles at two different loci. Arnaiz-Villena et al.000. Belgium). They speak the Palestinian-Arab dialect . The aim of the present study is to examine the genetic relationships of Palestinians with their neighbours (particularly the Jews) and other Mediterranean populations in order to: (1) discover the Palestinian origins. who live within Israel (37%).000 Palestinians). b KZ= Kafar Zubian (town).000 Palestinians. Iraq and other Mediterranean. 3. El-Shifa Hospital. (See Ref. . -B. Syria (12%) and Lebanon (13%). -DRB1 and -DQB1 allele frequencies. and (3) the Israel Palestinians. (2) the . coming from other parts of Palestine.000 Palestinians). MATERIAL AND METHODS Population samples Samples from 165 unrelated Palestinians in Gaza (Laboratories and Blood Bank. [9. but there are also Druze and Christian minorities . (2) the Diaspora Palestinians (occurred after 1947).500. c NS= Niha el Shouff (town). see Imanishi et al. the most frequent complete haplotypes were deduced following a methodology used in the 11th International Histocompatibility Workshop: (1) the 2. and live either in concentration camps or are scattered in Jordan (38%). and 4 HLA loci haplotype frequencies [3. This sample may be considered as representative of the Palestinian population because many people have been forced to live in the Gaza strip. Linkage disequilibrium (D´. All other populations used for comparisons are detailed in Table 1 and Figure 3. HLA Genotyping. and -DQB1 allele DNA sequencing was only done when indirect DNA typing (reverse dot-blot) yielded ambiguous results . TABLE 1 Identification number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 a Populations used for the present study Population Palestinians Berbers (Souss) Moroccan Jews Ashkenazi Jews Non-Ashkenazi Jews Lebanese KZ b Lebanese NS c Italians French Spaniards Portuguese Spanish-Basques Sardinians Cretans Macedonians Algerians Moroccans Number a 165 98 94 80 80 93 59 284 179 176 228 80 91 135 172 102 98 Reference Present study 37 38 8 8 3 3 16 16 27 18 27 16 19 20 17 28 Identification number 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Population Egyptians (Siwa) Armenians Turks Iranians Senegalese Bushmen (San) South African Blacks South American Blacks Greeks (Aegean) Greeks (Attica) Greeks (Cyprus) Oromo Amhara Fulani Rimaibe Mossi Japanese Number a 101 105 228 100 192 77 86 59 85 96 101 83 98 38 39 42 493 Reference 3 16 21 39 16 16 16 16 3 3 3 3 3 3 3 3 16 Number of individuals analysed for each population. this program calculated HLA-A. Saudi Arabia and Kuwait gather about 500. Egypt. ) and its level of significance (p) for 2 x 2 comparisons were determined using the formulae of Mattiuz and co-workers  and the 11th International Histocompatibility Workshop methodology .. but controls more in the Gaza strip (about 1. HLA-A.16-21]. In addition. DNA Sequencing and Statistics Generic HLA class I (A and B) and high resolution HLA class II (DRB1 and DQB1) genotyping was performed using a reverse dot-blot technique with the Automated Innolipa system (Innogenetics N.V. Most of Palestinians profess Muslim religion. European and American countries have more reduced communities. All were selected in order that their ancestors (eight grandparents) had a Palestinian origin. also named LD. who have refugee status (about 40%. where Palestinians have to lived mixed with Jewish colonist in their theoretically own territories.1 software kindly provided by Excoffier and Slatkin . Zwijndrecht. -DRB1.
4 and 5). Armenians and Egyptians. B*44 B*49. Cretans (using DRB1-DQB1) or Algerians (using DRB1) (see Table 3) and no discontinuity is observed between eastern and western Mediterranean populations reflecting the genetic similarity among all these populations. one class I allele obtained by generic DNA typing may contain several class I alleles. Correspondence analysis comprises a geometric technique that may be used for displaying a global view of the relationships among populations according to HLA (or other) allele frequencies. -B. Phylogenetic trees (dendrograms) were constructed with the allelic frequencies by applying the Neighbor-Joining (NJ) method  with the genetic distances between populations (DA. Turks.02 computer program (. B*50. Jews [Morocco]. A*03. 31 different DRB1 alleles were found and only seven had frequencies higher than 5% (DRB1*0301. is also present in North African Caucasians.28].16]. These three types of analysis were performed because some of the populations used for comparison lacked HLA-A and -B data (Berbers [from Souss. (2) with DRB1 data. Agadir area]. Cretans. The Palestinian population reveals the closest genetic distance with Jews (generic DR-DQ typing).psych. the reference tables used were those of the 11th and 12th International HLA Workshops [3. *0403. Greeks [Cyprus]. Lebanese [NS and KZ]. North African and Jewish populations [8. A*32. In order to compare allelic and haplotype HLA frequencies with other populations. see Table 1). A*30. whereas this is not the case for most DRB1 alleles at present. Seventeen HLA-A and 26 different HLA-B alleles were observed in the Palestinian population.HLA Genes in Palestinians 893 haplotypes previously described in other populations [3. RESULTS Characteristic HLA Allele Frequencies of the Palestinian Population Compared With Other Mediterraneans The expected and observed gene allelic frequencies for HLA-A. Populations are grouped into three main branches: the first one groups both eastern (including Palestinians. and (3) with generic (low resolution) DR-DQ data. A*02. These results are confirmed using DR and DQ generic typings (see Table 3 and Figure 6) which were used in . and B*51) and these are characteristic of Mediterranean populations. Table 2 illustrates the HLA allele frequencies found in the Palestinian population. A three-dimensional correspondence analysis and its bidimensional representation was carried out using the VISTA v5.29]. DRB1*0302 (allele frequency 3. see also Table 1. 3. it should be pointed out that class I generic typing tends to homogenize the comparisons based on DRB1 high resolution typing . http:/ forrest. Seven HLA-A and seven HLA-B alleles had frequencies higher than 5% (A*01.unc.3%). or high resolution HLA-DQ data (Greeks [Attica]. With regard to the HLA class II alleles. Jews [non-Ashkenazi].16]. see Table 1).1921. Analyses using DRB1 and DQB1 conjointly were made but are not illustrated because only a few populations could be used and the results are concordant with the DRB1 analysis. but they could be analyzed conjointly taking into account only either DRB1 or generic DR and DQ frequencies (Table 3. The second branch is formed by African Negroid populations and Japanese (Mediterranean outgroups) and the third one includes Greek and Ethiopian/sub-Saharan populations.27.17. Figure 4 depicts an HLA class II “high-resolution” (DRB1) NJ tree.28. Greeks [AtticaAegean]. B*18. particularly from eastern Mediterranean regions [17. Jews [Ashkenazi]. Finally. These partially HLA-typed populations should have been ignored. and (3) haplotypes that were assigned if they appeared in two or more individuals and the alternative haplotype was well defined. Iranians. Jews. *1101. B*41. which is probably a more informative and discriminating methodology. characteristic of African Black populations. B*35. It is evidenced that Palestinians/Greeks distance is high and confirms the different genetic background of the Greeks. *1303) being also characteristic of eastern Mediterranean populations. 2. DRB1*0403 is present in high frequency in Lebanese . DRB1*0406 is also present in North Africans populations [17. see Table 1). Sardinians are included in the first group). *0701. Lebanese) and western Mediterranean populations (Europeans and North Africans. A*23.28].25].edu). -DRB1 and -DQB1 loci do not significantly differ and the population sample is in Hardy-Weinberg equilibrium. Macedonians. *1104. This distribution is also confirmed in the correspondence analysis (Figure 5): the three groups are clearly delimited and an smooth West to East Mediterranean gradient is shown. Besides. ) and using DISPAN software containing the programs GNKDST and TREEVIEW [24. *0406. In particular. A*24. DQ allele frequencies reflect the DRB1 locus allele distribution due to the strong linkage disequilibrium between these two loci. Three types of analyses were carried out to compare Palestinian HLA frequencies with other Mediterranean population frequencies: (1) with DRB1-DQB1 data. Figsures 1. Jews and Lebanese [8.20]. or only generic HLA-DR and – DQ data were available (Portuguese.17]. This methodology is based on the allelic frequency variance among populations (similar to the classical principal components methodology) and on the display of a statistical projection of the differences. who have received a substantial sub-Saharan gene flow [3.
2 3.6 Alleles DQB1*0201 and 0202 were all assigned as DQB1*02. HLA-A.4 1.8 0. order to include other Mediterranean populations (Iranians.2 3.4 20.5 1.8 6.1 0.0 0. Other analyses and genetic distances confirm these results (Table 3.3 12.6 3.5 20.6 2. see Table 1).5 Alleles B*53 B*55 B*56 B*57 B*58 HLA-DRB1* 0101 0102 0301 0302 0309 0401 0402 0403 0404 0405 0406 0408 0701 0804 0901 1001 1101 1102 1103 1104 1201 1301 1302 1303 1305 1334 1401 1501 1502 1503 1601 Allele frequencies % 2. Alleles HLA-A A*01 A*02 A*03 A*11 A*23 A*24 A*26 A*29 A*30 A*31 A*32 A*33 A*66 A*68 A*69 A*74 A*80 HLA-B B*07 B*08 B*13 B*14(B65) B*15(B62) B*18 B*27 B*35 B*37 B*38 B*39 B*40(B60) B*41 B*42 B*44 B*45 B*47 B*49 B*50 B*51 B*52 Allele frequencies % 12. Egyptians and Anatolians (Turks ) further support an autochthonous Canaanite/Middle East origin for both Palestinians and Jews.9 7.4 5.7 0.9 4.6 3. Figures 4 and 5).4 1.3 1. Turks (old Anatolians). -DRB1.8 1.0 3.3 2.8 1.4 Alleles HLA-DQB1* 02 0301 0302 0303 0402 0501 0502 0503 0601 0602 0603 0604 Allele frequencies % 20. Cretans.9 26.8 11.5 10.5 6.9 10.3 1.3 1.6 1. Armenians. -DRB1. and -DQB1 Linkage Disequilibria in Palestinians Extended HLA haplotypes were defined in Palestinians and compared with those previously reported in other populations (Table 4).3 7.3 1. A DR-DQ neighbour-joining tree (not shown) maintains the west and east Mediterranean groups and also the group formed by Greeks and sub-Saharan populations. -B.3 9.3 3. Armenians. TABLE 2 HLA-A.4 9.5 6.9 0. On the other hand.0 3.7 2. Arnaiz-Villena et al. HLA-A-B and DRB1*-DQB1* two-loci linkage disequilibrium data (not shown) show that the most .3 0.7 0.8 0.6 1.9 3.4 4. Turks and non-Ashkenazi Jews are the populations closest to the Palestinians.1 0.8 4. genetic distances obtained by using DR-DQ generic typing allele frequencies (Table 3) illustate that Ashkenazi Jews.5 0.9 5.2 3. and -DQB1 allele frequencies in the Palestinian population.9 5.9 0.2 6.9 3. Jews and Lebanese are illustrated specifically to cluster with Palestinians.9 3.8 2. Armenians.3 0.4 4.3 9. Number in brackets indicates the serologic antigen most probably corresponding to the genetic allele obtained. Iranians. The close relatedness of Palestinians (Table 3 first column.5 7.4 1.6 0.6 2.2 2.3 0.3 10.9 1.3 1.2 3.1 2.7 17.4 8. followed by the other Mediterraneans populations.8 2.894 A.5 1.2 0.6 3. Egyptians and Turks. Figure 6) to Iranians.3 3.3 0.2 1.6 4. Armenians.6 2. -B.
A*33-B*14. HF: 1.8.25 2.38 11.37 3.36 38.85 HLA-DRB1 (DA) Algerians Cretans Non-Ashkenazi Jews Moroccans Lebanese-KZ Spaniards Moroccan Jews Italians Lebanese-NS Ashkenazi Jews French Macedonians Berbers (Souss) Sardinians South African Blacks Spanish-Basques Greeks (Attica) Oromo Senegalese Amhara Greeks (Cyprus) Japanese Greeks (Aegean) San (Bushmen) Mossi Fulani Rimaibe 12. However.33 12. The HLA-A-B-DR-DQ extended haplotypes found in the Palestinian population (Table 4) reflect common characteristics with the other “older” Mediterranean population mainly from eastern Mediterraneans and North Africans (see footnote to Table 4).2.09 14. Africa and Asia. Babylonian and Roman-induced Diaspora.36 48. B*18-DRB1*1104.HLA Genes in Palestinians 895 TABLE 3 Genetic distances (DA) between the Palestinians and other populations (x 102) obtained by using generic HLADR-DQ. HLA-DR-DQ (DA) Ashkenazi Jews Iranians Cretans Armenians Turks Non-Ashkenazi Jews Spaniards Portuguese Algerians Lebanese-KZ Macedonians French Moroccans Moroccan Jews Egyptians Sardinians Italians Berbers (Souss) South American Blacks South African Blacks Senegalese Spanish-Basques San (Bushmen) Oromo Amhara Greeks (Attica) Japanese Mossi Fulani Rimaibe 1.73 53. HF: 1. B*35-DRB1*1001.55 11.83 34. A*2-B*51. A*2-B*41. Jews wrote the Bible. NJ trees and correspondence analyses. but that only tells the Jewish view . A*23-B*44.65 7.72 23. and high-resolution HLA-DRB1 and HLA-DRB1-DQB1 allele frequencies (see Table 1 for identification of populations).5). to establish a more objective history of Middle East and particularly ancient Canaan [6.62 27.71 8.96 24.32 220.127.116.11 39.82 14.99 19. HF: 2. B*8-DRB1*0301.48 13. HF: 2. HF: 2.50 15.04 4. HF: 2. B*35-DRB1*0403.2. B*49-DRB1*0403.61 2. B*35-DRB1*1104.00 9.34 4. These haplotype results are concordant with those ob- tained by the allele frequency analyses (genetic distances.63 4. The combination A*69-B*49 (HF: 2.62 15.01 30.33 11.50 9. HF: 2. like Jews .28 17. HF: 1.51 4. DISCUSSION Palestinians and Jews The genetic identity of Ashkenazi and non-Ashkenazi Jews who now lives in Israel has already been reported . Jews started to come back to Palestine during the 19th and 20th centuries . drove Jews to many parts of Europe.15 13.52 6.66 25.93 14.48 40. HF: 3.27 34.95 10. B*50-DRB1*0701.53 11.82 20. respectively. B*14DRB1*0102.08 3. haplotype Frequency (HF): 5.53 12.6.47 12. a religious and historical book that is a continuous source of historical Middle East facts.3. such as archaeology. HF: 2.28 3.25 57.78 25.81 13.24 15.64 5.95 43.37 15. which occurred in 587 BC and 70 AD.25 3.33 64. HF: 18.104.22.168 15.3. A*24-B*35. HF: 3.12 3. see above).67 46. B*35-DRB1*1101.27 9.37 HLA-DRB1-DQB1 (DA) Cretans Moroccans Spaniards Non-Ashkenazi Jews Lebanese-KZ Algerians Moroccan Jews Ashkenazi Jews French Italians Macedonians Lebanese-NS Berbers (Souss) Sardinians Spanish-Basques South African Blacks Japanese Senegalese San (Bushmen) 9.45 3. etc.5.41 36.83 33.7) has not been found in any of the populations tested and it is included in an extended haplotype (A*69-B*49-DRB1*0403-DQB1*0302. religion and close communities have kept Jews relatively isolated from the inhabitants of the countries that hosted them during this long period of time.27 32.92 frequent combinations are characteristic of Mediterranean (western and eastern) populations (A*1-B*35.56 42.9]. . B*49-DRB1*1001. It is now necessary to rely on other sources.32 12.36 58.54 17. see below) not previously described.67 6.39 20. HF: 1.61 5. HF: 3.0.92 17.1.0. A*24-B*18. linguistics.70 12.1.60 16. HF: 2.5.17 3. HF:1.
Bootstrap values from 1000 replicates are illustrated. The present day concept based in archaeology is that most original Palestinians were already in Canaan and some tribes were agglutinated by Egyptian garrisons. Data from other populations were from references detailed in Table 1. left to their own fate in Canaan . FIGURE 4 Neighbor-Joining dendrogram demonstrating relatedness between Palestinians and other populations. Palestinians appeared in the Bible as coming from Crete or its empire . but the input of one “elite” coming from Crete may not .896 A. Genetic distances between populations (DA) were calculated by using HLA-DRB1 (high resolution). Arnaiz-Villena et al.
Italians and Basques (Table 3. Palestinians named their leaders or princes as “seren” (Basque. ancient stock.HLA Genes in Palestinians 897 FIGURE 5 Correspondence analysis depicting a global view of the relationship between Mediterraneans and Palestinians according to HLA allele frequencies in three dimensions (bidimensional representation). ). Turks and Armenians are probably the closest relatives to Palestinians and this favors the hypothesis that most of the HLA Palestinian genetic background comes from the Middle East (ancient Canaan. The western and eastern Mediteranean populations are intermingled in this case. Figures 4. Iranians.: ancient Canaanites. . DRB1 genetic distances (Table 1) are probably the most reliable ones due to the higher polymorphism detected in this locus. ). French. Both Jews and Palestinians share a very similar HLA genetic pool (Table 3. American. i. 5 and 6) that support a common ancient Canaanite origin. Moses) as described in the Bible . it also has artificially divided peoples both coming from ancient Canaanites (Jews and Palestinians). Egyptians. The typical Philistine crest-hut already appeared in the Cretan Phaistos Disk (1600 BC) and in the Ramses III-Medinet Habu temple. Egypt (1200 BC. Zar = old man. Cretans. Also.e. A difficult problem has now been created between two communities that are close genetic relatives. and 6). it is clear that they spoke a language different to Arab or Jewish in ancient times and only a few words have been preserved. The study of this and other words suggests that they spoke a Dene-Caucasian language like other Mediterranean populations [30. which has also caused a massive displacement of Palestinians and wars. Cretans. Lebanese. and also to Algerians. the origin of the long-lasting Jewish-Palestinian hostility is the fight for land in ancient times. Macedonians and Sardinians. HLA-DRB1 allele frequencies data. Figures 4. Jews. Spaniards. this is compatible with an input of foreign leaders and their groups (Abraham. it supports the long-standing prehistoric and historic circum-Mediteranean gene flow . be discarded. Therefore. the bulk of Jewish people probably came from ancient autochthonous Canaanites . Palestinians and Other Middle East and European People Palestinians are close to Egyptians. together with the massive European. Regarding Palestinian population identity. 5. Asian and African Jews settlements in the area. The Eurocentric confusion “Arab = Muslim” has also lowered the Palestinian identity by identifying the country were Mohammed was born (Saudi Arabia) with the Muslim religion. en = the most important) . Religious and cultural have enhanced the conflict in the last centuries. Iranians.31].
FIGURE 6 Correspondence analysis depicting a global view of the relationship among Palestinians. This could have occurred when hyperarid Saharan condition become established and large-scale migration occurred in all directions out from the desert. By 1500-1200 BC the Greek presence was very scarce in Canaan. The Cretan Aegean Sea empire was destroyed and continued by the Mycaenians. according to archaeologic records .898 A. This is why Greeks are Mediterranean outliers in all kind of analyses [19-21. the most ancient Greek Pelasgian substratum would come from a Negroid stock. In fact. HLA-DR and DQ (low resolution) allele frequencies data. Cretans and Greeks The Biblical origin for Palestinians (Crete) cannot be disregarded  because an “elite” group could have joined to Canaanite proto-Palestinian tribes and made themselves noticeable. Anatolia and Egypt . East Mediterraneans. An explanation of the Egypt-to-Greece migration may be that a densely populated Sahara (before 5000 BC) may have contained an admixture of Negroid and Caucasoid populations. this is supported by the ancient Palestinians high war technology and the many confrontations with the Jews after 1500 BC [6. A more likely explanation is that at an undetermined time during Egyptian pharaonic times a Black dynasty with their followers were expelled and went towards Greece where they settled [20. according to Amelie Kuhrt .20]. Arnaiz-Villena et al. and some of the Negroid populations may have migrated by chance or unknown causes towards present day Greece [19. It is very unlikely that a massive immigration of Palestinians came from Crete . the “Mycaenian” Greeks attacked Crete by 1450 BC after rendering tributes to Cretans by a relatively long period.28]. Greeks are found to have a substantial HLA gene flow from sub-Saharan Ethiopian and Black people [3. Also. Canaan had received gene and cultural flow from Mesopotamia. 30]. Once an African input to the ancient Greek genetic pool is established. ancient Greeks believed that their religion and culture came from Egypt . West Mediterraneans. Palestinians. Greeks and subSaharian populations and Blacks according to HLA allele frequencies in three dimensions (bidimensional representation) .7]. In this case. it remains to be determined what the . Egyptian garrisons in Canaan abandoned to their fate by the Egyptian Kingdom weakness may have catalyzed the union of some Canaanite tribes to become the historical Palestinians. This African genetic and cultural input was documented by Herodotus  who states that the daughters of Danaus (who were black) came from Egypt in great numbers to settle in Greece.34-36].
Corell A.4).3). Other low frequency haplotypes present in Palestinians are also shared with Mediterraneans and central Europeans (A*01-B*08-DRB1*0301-DQB1*02. Vol 1. Surney: Encyclopaedia Britannica International Ltd. Spaniards (1. French (HF: 1.5) (10turk). Editions La Dècouverte. Martínez-Laso J.9%). Also found in Germans (HF: 1.7).9). HF:0.1%) and Greeks (4. Comparisons were made by using references [3. Fernandez V. HF: 0.0).1) and Caucasoid Australians (HF: 3.0%). HF:0. 11. Eurasiatics (A*02-B*08-DRB1*0301-DQB1*02. 3. Beck’sche Verlagsbuchhandlung.8 1. Regueiro JR: Primary immunodeficiency caused by mutations in the gene . Paris. This complete haplotype has not been found in any of the populations tested.7). African Blacks (A*02-B*42-DRB1*0302. Biblia de Jerusalén. A*02-B*49-DRB1*1001DQB1*0501. 1997.0). Timón M.5%).3/14/98). Armenians (HF: 5. Also. Sellier J. i. Also. Tissue Antigens 47: 63.2 1.0) and Portuguese (HF: 2.9 0. Copenhagen. 1996. and French (HF: 1. Turks (1. Lonjou C: Allele and Haplotype frequencies for HLA loci in various ethnic groups. 10. Sellier A: Atlas des Peuples d’Orient. Haplotypes HF (%) 2. Timorese (HF: 2.6.1).6. Munksgaard.9 0. This haplotype has been found in Albanians (3.4 1.1%) and Italians (0. Gojorobi T: Genetic relationships among various human populations indicated by MHC polymorphisms. Mediterraneans (A*02-B*35-DRB1*1104-DQB1*0301.6) haplotype has been found. Gonem A: The encyclopaedia of the peoples of the world. Aizawa M. 1996. -B. 2001. k. 2000.6).9). Oxford. Vol II. Dausset J. In Tsuji K.6). A*03-B*07-DRB1*1501-DQB1*0602. Kuhrt A: The ancient Near East (3000-330 BC). 4. Extended haplotype (generic typing) has been found in Turks (HF: 0.2 1.1). 9. Jews (HF: 6. Madrid. Barcelona. 143-5. Pérez-Aciego P.2). Wakisaka A. Kienitz FK: Volker imm schatten.2).2 1. Haplotype previously described as Mediterranean. Albanians (HF: 3. South West Asians (A*30-B*49-DRB1*1001-DQB1*0501. REFERENCES 1.1) and Spaniards (HF: 1. Spaniards (HF: 1. d. Sardinians (HF: 4. Nei for providing the GNKDST and TREEVIEW programs found on the DISPAN software. Desclee de Brouwer (ed): Bilbao.1). HF: 0.2 0.8 1. l. 627-32. Encyclopaedia Britannica (1999) Sutton. In Charron D 6. Italians (2.1%). We are grateful to Alberto Garcia for his help with art design work on the computer. Greeks (HF: 1. -DRB1. Spaniards (HF: 1. HF:0. g. ACKNOWLEDGMENTS (ed): Genetic diversity of HLA. 20]. Italians (HF: 6. Austrians (HF: 4.7%). 12. 665-820. 1972. Arnaiz-Villena A: HLA DR and DQ polymorphism in Ashkenazi and non-Ashkenazi Jews: comparison with other Mediterraneans. Hungarians (HF: 3. j.6.1).8). We also thank M. Arnaiz-Villena A. Haplotype found in Armenians (2. 1998.2). The reason why a subSaharan admixture is not seen in Crete is unclear but may be related to the influential and strong Minoan empire.9).0). 1992. Germans (HF: 1. however partial and generic typing B49-DR4 is present in French (HF: 1. Editorial Critica. A*23-B*35-DRB1*1001-DQB1*0501. EDK. Martinez-Quiles N.27-29] cultural importance of this input is for constructing the classical Hellenistic culture.1%). Alvarez M.0).3). Akal Editions. HF:0. Anuario geopolítico. Morales P.3). This work was supported in part by grants from the Spanish Ministry of Education (PM95-57 and PM96-21) and the Madrid Regional Government (06/70/97 and 8. a. Spain. Vol 1. Partially (B35-DR14) found in Italians (HF: 3.5 1.2) and Italians (HF: 0. 1981. 5. Extended haplotype found in Mexican Mestizos (HF: 2. Die gegenspieter der Griechen und Römer. Indians (HF: 3.6). 1993. Functional and Medical Implications.5 1.4) and Italians (HF: 1. A*02-B*35-DRB1*0403-DQB1*0302. Gómez-Casado E. h. Gazit E. Paris. Germans (HF: 3. b and e. Sasazuki T (eds): HLA 1991. Present in Austrians (HF: 2. 2000.9 0. probably autochthonous A*02B*50-DRB1*0402-DQB1*0302 (HF:0. Munchen. El Estado del Mundo. 7.7). Partially (B35-DR4) found in Jews (HF: 6.6). HF:0.6). Clayton J. Oxford University Press. HF: 0. Mediterranean-Europeans. 8.8. Yugoslavs (3. 2.9 Possible origin Palestinian/ Mediterranean Mediterranean/ Central European Central-South-Eurasian Mediterranean/ Central European Palestinian Mediterranean/ Central European Mediterranean/ Central European Mediterranean/ European Eurasiatic Mediterranean Eastern Mediterranean Mediterranean Mediterranean A*69-B*49-DRB1*0403-DQB1*0302a A*01-B*35-DRB1*1104-DQB1*0301b A*24-B*18-DRB1*1104-DQB1*0301c A*03-B*35-DRB1*0701-DQB1*02d A*02-B*41-DRB1*0701-DQB1*02 A*01-B*35-DRB1*1101-DQB1*0301e A*24-B*35-DRB1*1101-DQB1*0301f A*23-B*44-DRB1*0701-DQB1*02g A*02-B*50-DRB1*0701-DQB1*02h A*02-B*35-DRB1*1401-DQB1*0503i A*24-B*35-DRB1*0403-DQB1*0302j A*02-B*18-DRB1*1104-DQB1*0301k A*33-B*14-DRB1*0102-DQB1*0501l HF: Haplotype frequency. f. Publishing House Ltd. Manchu (HF: 2. the partial haplotype B35-DR11-DQ7 is present in South African Negroid (HF: 2.6). Jerusalem.6). Imanishi T. which hindered foreigners establishment if the African invasion occurred in Minoan times [19. HF:0. and -DQB1 extended haplotypes in the Palestinian population and their possible origin.0).4).16-21. c.6. Martín-Villa JM. Greeks (HF: 2. Colombani J: Histocompatibility Testing 1972.HLA Genes in Palestinians 899 TABLE 4 Most frequent HLA-A. Present in Mongolians (HF: 3. Martin-Villa JM.
Martinez-Laso J. 30. 2001. Arnaiz-Villena A. Arnaiz-Villena A. Paris. Anthropology and Linguistics. N Eng J Med 327:529. Moscoso J. Iliakis P. Arnaiz-Villena A. Un estudio genético y lingüístico. Nei M. Bann CM: A Visual Statistics system. Guillén J. 1995. Arnaiz-Villena A. -C and -DR. Akaza T. Gomez-Casado E. 26. 35. 1999. Rajalingam R. 28. Longás J. Mol Biol Evol 12:921. 1989. Excoffier L. Tissue Antigens 53:213. 18. Tissue Antigens 57:308. 32. Akaza T. 1995. -B. Saitou N. Brautbar C: Comparative analysis of HLA polymorphism at the serologic and molecular level in Moroccan and Ashkenazi Jews. 2000. Pacho A. Vol I. Gredos. Functional and Medical Implications. del Moral P. Caillat-Zucman S. . 353-363. Nei M: Genetic distances between populations. Allende L. Arnaiz-Villena A. In Charron D (ed): Genetic diversity of HLA. EDK. McEnemy L. and Algerian studied by HLA allelic frequencies and haplotypes. Tissue Antigens 46: 104. Tissue Antigens 57:118. Anthropology. Nelken D. Gómez-Casado E. Alonso-Garcia A. Akhayat O. Moscoso J. New York. In Arnaiz-Villena A (ed): Prehistoric Iberia: Genetics. 19. Gazit E: 12th International Histocompatibility Workshop: Anthropology SWAS regional report. Gomez-Casado E. Alonso-Garcia J: The Usko-Mediterranean languages. 1999. Piazza A. 1992. 27. Kluwer Academic-Plenum Publishers. Gojorobi T: Estimation of allele and haplotype frequencies for HLA and complement loci. Iberia: Population genetics. Pacho A. 15. 1983. 23. Editorial Complutense SA. Editorial Complutense SA. Histocompatibility Testing 1970. New York. Oxford. Funcional and Medical Implications. 24. Arnaiz-Villena et al. Breda-Coimbra H: Relatedness among Basques. Arnaiz-Villena A. Safirman C. Nei M: Analysis of gene diversity in subdivided populations. Madrid. 13. Grollemberg LH: The Penguin shorter Atlas of the Bible. Singal DP: Genetic diversity of HLA in the populations of India. Bias W. Martínez-Laso J. 1992. 1972. Tokunaga K. Cuadrado E. Mattiuz PL. Trapaga J. 1998. 1997. Tajima F. Martínez-Laso J: The origin of Cretan population as determined by characterization of HLA alleles. Editorial Complutense SA. Portuguese. 2000. Longas J. Ceppelini R. Allende L. 1973. Gene frequency data. Kandil M. Sfyridaki K. Mehra NK. Aizawa M. De Juan D. Agadir area. Álvarez M. Sasazuki T (eds): HLA 1991. Beaurain G. Proc Natl Acad Sci USA 70:3321. Ed. Am Nat: 106:283. Spaniards. 38 Izaabel H. Slatkin M: Maximum-likelihood estimation of molecular haplotype frequencies in a diploid population. ArnaizVillena A: HLA genes in Arabic-speaking Moroccans: close relatedness to Berbers and Iberians. Arnaiz-Villena A. Immunogenetics 47:37. Sherman L. Hum Biol 71:725. Vol 1. Kanga U. Arnaiz-Villena A. Alonso-García J: Minoicos. Tissue Antigens: 45:237. Paris. Arnaiz-Villena A: The contribution of the HLA-A. Martínez-Laso J. Madrid. Alonso-García J: The correlation between languages and genes: the uskoMediterranean peoples. Martinez-Laso J: HLA genes in Macedonians and the sub-Saharan origin of the Greeks. Pollack MS. Vol 1. Varela P. Carin M. Silvera C. Alonso-García J: Egipcios. 14. Garchon HJ. In Stine RA & Fox J (eds): Statistical Computing Environments for Social Researchers. 314-20. 1997. 1065-220. Hum Immunol 43:259. Amirzargar A. Wodmer WF. In Tsuji K. Martinez-Laso J: HLA alleles and haplotypes in the Turkish population: relatedness to Kurds. Mol Biol Evol 4:406. 39. Munksgaard. New York. Tateno Y: Accuracy of estimated phylogenetic trees from molecular data II. Armenians and other Mediterraneans. Agarwal S. J Mol Evol 19:153. Silvera C. Martinez-Laso J. Tissue Antigens 55:239. Ihde D. Friedmann A. Santos P. 1970 16. Tissue Antigens 51: 106. Sri Lanka and Iran. 207-36. Benmamar D. 40. Gomez-Casado E. Díaz-Campos N. Sage publications. 1987. Oxford: Oxford University Press. Martínez-Quiles N. Bach JF. Imanishi T. Battat S. 37. 34. Imanishi T. Madrid. Arnaiz-Villena A. 1998. Roitberg-Tambur A. Middleton D. In Charron D (ed): Genetic diversity of HLA. Bereberes. Zamora J. 2001. Oxford University Press. 1995. Tokunaga K. Penguin Books Ltd. Nei M: The neighbor-joining method: a new method for reconstructing phylogenetic trees. Vol 1. 76-9. In Tsuji K. 1997. Martinez-Laso J. Cullen C. González-Hevilla M. Gojobori T: Allele and haplotype frequencies for HLA and complement loci in various ethnic groups. and linguistics.900 A. Witt CS. 22. 1995. Bendikuze N. Varela P. 1996. Silvera-Redondo C. 25. Arnaiz-Villena A. Dimitroski K. 1978. Human Immunol (This issue) 31. Kimura A. Copenhagen. 193-206. Silvera-Redondo C. Sasazuki T (eds): HLA 1991. Relatedness to Spaniards and Basques. Kimura A. Matsouka C. Alonso-García J: El origen de los Vascos y otros pueblos Mediterraneos. Martinez-Laso J: HLA allele and haplotype frequencies in Algerians. 21. 36. 2000. González-Hevilla M. GomezCasado E. 17. Cretenses y Vascos. Aizawa M. Gomez-Casado E. EDK. Young FW. 33. encoding the CD3-g subunit of the T-lymphocyte g receptor. DíazCampos N. 20. 1999. Herodotus: History. Martinho A. GarcíaGómez A. Sanchez-Mazas A: HLA class II DNA polymorphism in a Moroccan population from the Souss. -DQ DNA typing to the study of the origins of Spaniards and Basques. Gómez-Casado E. 1992. Arnaiz-Villena A. 29. Madrid. Guanches y Vascos.
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