ISSN 1022-7954, Russian Journal of Genetics, 2009, Vol. 45, No. 9, pp. 1082–1091. © Pleiades Publishing, Inc., 2009.Original Russian Text © O.S. Dedkova, E.D. Badaeva, A.V. Amosova, S.P. Martynov, V.V. Ruanet, O.P. Mitrofanova, V.A. Pukhal’skiy, 2009, published in Genetika, 2009, Vol. 45,No. 9, pp. 1234–1243.

1082

INTRODUCTION

Emmer

Triticum

dico

ÒÒ

um

(Schrank) Schuebl. isone of the most ancient cultivated plants. It is believedthat

T. dicoccum

was domesticated in the area of mod-ern Turkey approximately 10000 years ago and spreadfrom this area both eastward and westward, occupyingthe total Europe by the 4th or 5th millennium B.C.

Hulled emmer is considered to enter Europe viathree paths during its distribution history. One started insouthern Levant or southern Turkey and went throughAnatolia and Greece, where emmer was cultivated incoastal plains and islands [1–3]. By the 5th millenniumB.C., emmer reached Italy, southern France, and Spainand became a dominating crop there [4]. According toPliny, Latium people cultivated nothing but emmeronly 300 years before Rome was founded (second halfof the 5th century B.C.) (Pliny XVIII:11). Remains ofemmer grains were found in archeological sites ofSpain [5].

The second path of emmer distribution went fromthe Fertile Crescent toward the Rhine Valley throughthe Balkans and the Danube Valley (early 5th millen-nium B.C.). From these regions, emmer entered Cen-tral, West, and North Europe. Emmer reached the Neth-erlands in the 4th millennium B.C. and the UnitedKingdom and Scandinavia in the 3rd millennium B.C.[3]. Via these paths, emmer was brought to Europe as amajor component of a mixture with other wheat spe-

cies. This mixture probably contained einkorn wheat ina substantial proportion and naked tetraploid andhexaploid wheats were secondary components, whilelacking hulled hexaploid wheat [4, 6, 7].

The third, far more recent and relatively small flowreached Central Europe through Transcaucasia and theCaucasus (late 5th millennium B.C.) and southern Rus-sia (early and mid-4th millennium B.C.). This flowincluded mostly naked hexaploid wheat and only aminor proportion of hulled wheat. Hulled emmer wasthe most important crop in Europe up to the 1st millen-nium B.C. and came to be replaced by naked tetraploidwheat

T. durum

in the early 1st millennium B.C. [8].The current region of hulled emmer cultivation in

Europe is restricted to Balkan highlands (Bulgaria andformer Yugoslavia), Czechia, Slovakia, Italy, andSpain. Many emmer accessions are maintained ingenetic collections in the United States, Japan, Italy,and Russia [5, 9].

Our previous study of the intraspecific diversity of

T.dicoccum

by chromosome C-banding showed thatEuropean emmer accessions are clearly subdividedaccording to their geographical regions and that theT7A:5B translocation in widespread in this group [10].A subdivision of emmer accessions into eastern andwestern groups was observed in an analysis of the dis-tribution of hybrid necrosis genes [11]. The objective ofthis work was to further study European emmer with a

Diversity and the Origin of the European Population of

Triticum dicoccum

(Schrank) Schuebl. As Revealed by Chromosome Analysis

O. S. Dedkova

a

, E. D. Badaeva

b

, A. V. Amosova

b

, S. P. Martynov

c

, V. V. Ruanet

a

, O. P. Mitrofanova

c

, and V. A. Pukhal’skiy

a

a

Vavilov Institute of General Genetics, Russian Academy of Sciences, Moscow, 119991 Russia;e-mail: dediog@mail.ru

b

Engelhardt Institute of Molecular Biology, Russian Academy of Sciences, Moscow, 119991 Russia

c

All-Russia Institute of Plant Industry, St. Petersburg, 190000 Russia

Received May 15, 2008

Abstract

—Cluster analysis of the

Triticum dicoccum

chromosome passports by artificial neural networks andUPGMA divided the European

T. dicoccum

population into two groups, West European and Volga–Balkan. TheWest European

T. dicoccum

accessions displayed a predominance of the marker translocation T7A:5B (67% ofthe accessions), which was also found in a few accessions from other countries (Turkey, Iran, and northernAfrica), and were similar in chromosome C-banding patterns. The Volga–Balkan

T. dicoccum

accessions dif-fered in the C-banding patterns of some chromosomes from the West European accessions, thus probably orig-inating from another founder population. It was assumed that the

T. dicoccum

accessions carrying the T7A:5Btranslocation had a common origin and that the wild

T. dicoccum

population of the Middle East (Syria and Leb-anon) contributed to the origin of West European

T. dicoccum.

DOI:

10.1134/S1022795409090099

PLANT GENETICS

RUSSIAN JOURNAL OF GENETICS

Vol. 45

No. 9

2009

DIVERSITY AND THE ORIGIN OF THE EUROPEAN POPULATION 1083

larger sample of accessions from Western and EasternEurope by standard statistical tests and artificial neuralnetworks. In addition, genomic in situ hybridization(GISH) and in situ hybridization with the B genome-specific repetitive sequence pSc119.2 and a 5S rDNAprobe were carried out to determine the exact structureof the translocation between chromosomes 7A and 5B.

MATERIALS AND METHODS

To estimate the relationships among European

T. dicoccum

accessions, the chromosome passports of53 accessions were analyzed by statistical tests(unweighted pair group with mathematical average,UPGMA) and artificial neural networks (ANNs,Kohonen neural networks). Of the 53 accessions,44 were from European countries and nine were fromother countries (Turkey, Iran, and Algeria) and includedfour accessions carrying the 7A:5B translocation(Table 1). Five accessions of the Turkish origin with thenormal karyotypes were included in the analysisbecause the territory of modern Turkey is considered tobe a region where tetraploid wheat was domesticated inthe Neolithic [1, 2, 12–14].

A total of 26 accessions were obtained from the col-lection of the State Research Center of the All-RussiaInstitute of Plant Industry (IPI, Russia); 28 accessionswere from the National Institute for AgriculturalResearch (INRA, Clermont–Ferrand, France).

Chromosome preparation and C-banding were per-formed according to a published protocol [15]. Prepa-rations were examined under a Leitz Wetzlar micro-scope, and selected metaphase plates were photo-graphed (immersion objective Apochromat

100

×

) witha Leica DFC 280 digital FireWire Color Camera Sys-tem for Analysis and Documentation. Images were pro-cessed using the Adobe Photoshop package, v. 7.0.Chromosomes were classified according to a standardgenetic nomenclature [16, 17].

Chromosome passports were constructed by com-paring the C-banding patterns of the accessions understudy with a generalized species ideogram of wheatchromosomes [18, 19]. To describe an accession, dataon its heterochromatin bands (129 positions in total)were input into an Excel spreadsheet.

The resulting table was subjected to cluster analysis,which was performed using the NTSYS 2.02c package[20]. Hierarchic cluster analysis was carried out byUPGMA. A similarity measure was calculated by atwo-step procedure. At the first step, a data matrix withdimensions

n

×

m

, where

n

= 53 is the number of theaccessions and

m

= 129 is the number of heterochroma-tin bands, was used to obtain a square matrix of stan-dardized Euclidean distances with dimensions

n

×

n

.The resulting matrix was used as a new data matrix. Theestimated distances from an accession to all otheraccessions were used as traits in this case. Then, we cal-culated a matrix (

n

×

n

) of the coefficients of correlation

Table 1.

Triticum dicoccum

accessions and their origins

No.. Original country Collec-tion

Acces-sion

Rearrange-ment

1 Iran IPI k-7146 T7A:5B2 Turkey IPI k-14380

none

3

"

IPI k-20967

"

4

"

IPI k-20968

"

5

"

IPI k-20989 T7A:5B6

"

IPI k-20993

none

7

"

IPI k-21007

"

8 Russia (Gor'kovskaya oblast)

IPI k-30728

"

9 Russia (Chuvashia) IPI k-24482 Inv3B10 Bulgaria IPI k-14236

none

11

"

INRA 27089*

"

12

"

INRA 27090*

"

13 Yugoslavia IPI k-23036

"

14

"

IPI k-38904

"

15

"

IPI k-38917

"

16

"

IPI k-40030

"

17

"

INRA 27119*

"

18

"

INRA 27126*

"

19 Hungary INRA 26654 T7A:4B20 Germany INRA 26639

none

21

"

IPI k-18623 T7A:5B22

"

IPI k-21433

none

23

"

IPI k-21961

"

24

"

INRA 26642* T7A:5B + + T6A:1B-3

25

"

INRA 27123*

none

26

"

INRA 27124* T7A:5B27

"

INRA 27125* HÂÚ28 Netherlands IPI k-35890 T7A:5B29 Italy INRA 27095

none

30

"

INRA 27096

"

31

"

IPI k-21416 T7A:5B32

"

INRA 27088* T2A:1B33

"

INRA 27097* T7A:5B34 Poland IPI k-14999 T3B:6B-235 France INRA 6807 T7A:5B36

"

INRA 898

"

37

"

INRA 900

none

38

"

IPI k-21582

"

39

"

INRA 26647* T7A:5B40

"

INRA 26658*

none

41

"

INRA 26659* T7A:5B42

"

INRA 26648*

"

43

"

INRA 26657*

"

44 Czechoslovakia INRA 26651

"

44 Switzerland INRA 12946 Inv(T5B:7A)46

"

INRA 23757 T7A:5B47 Sweden IPI

k

-36527

"

48 Spain INRA 27076

"

49

"

IPI k-20410

"

50

"

IPI k-20541

"51 " IPI k-20638 "52 Algeria INRA 26893 "53 " INRA 26894 "

Note: Chromosome rearrangements were classified according toBadaeva et al. [27].

* Accessions that we had not studied before.

1084

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DEDKOVA et al.

between the Euclidean distances and used this matrixfor cluster analysis.

To analyze the results by ANNs, we used the ExcelNeural Package and the freely available NeuroPro 0.25program [25].

DNA probes. We used two DNA sequences. Theclone pSc119.2 was a 120-bp DNA fragment isolatedfrom Secale cereale [22] and subcloned in McIntyre[23]. The clone pTa794 was a 410-bp BamHI fragmentof wheat 5S rDNA, which was cloned in pBR322 [24].

In situ hybridization. GISH was carried out by animproved protocol [25]. Hybridization was performedwith biotinylated (biotin-16-dUTP) T. boeoticumgenomic DNA, and fragmented Aegilops speltoidesgenomic DNA was used to prevent cross-hybridization.The cloned sequences were hybridized to chromo-somes according to a standard method [26].

Preparations were examined under an OlympusBX61 microscope and photographed using a Cool SnapCCD camera (United States). The chromosome imageswere processed using Adobe Photoshop software(v. 7.0).

RESULTS

Our sample included 29 T. dicoccum accessionsexamined previously [10] and 15 new accessions fromthe INRA collection, France (asterisked in Table 1),which represented both Volga–Balkan (four acces-sions) and European (11 accessions) groups. The newaccessions from Bulgaria and former Yugoslavia hadnormal karyotypes without overt chromosome rear-rangements; their karyotypes were similar in chromo-some C-banding patterns to the Volga–Balkan T. dicoc-cum accessions examined previously. The total sampleincluded 33 accessions from West European countries,12 accessions belonging to the Volg–Balkan group, and8 accessions of Asian and north African origins.

Among the 11 new European accessions, seven car-ried the marker translocation T7A:5B. The accessionINRA 26642 had not only T7A:5B, but also an addi-tional centromeric translocation between chromosomes6A and 1B (T6A:1B) (Table 1, Fig. 1, a). The Italianaccession INRA 27088 displayed a new single translo-cation between chromosomes 2A and 1B with break-points in the short arm of chromosome 1B and the longarm of chromosome 2A (Table 1, Fig. 1, i).

Thus, examination of a larger sample of European T.dicoccum accessions confirmed that T7A:5B tends topredominate in Western Europe. The translocation wasfound in 22 accessions, which account for 50% of thetotal sample of European hulled emmer accessions. Atthe same time, T7A:5B was detected in Turkish, Ira-nian, and two Algerian accessions. The translocationwas reported for wild tetraploid wheat T. dicoccoidesfrom Middle East countries [27]. We did not detect the7A:5B translocation in the European T. dicoccumaccessions of the Volga–Balkan group.

The structure of the rearranged chromosomes wasstudied by GISH in two T. dicoccum accessions, k-7146and k-20989. The results showed that the translocationwas reciprocal and that the chromosome structureswere the same in both accessions. The fragment trans-ferred from the long arm of chromosome 7A to theshort arm of chromosome 5B accounted approximatelyfor a half of the arm length (Figs. 2c, 2d). The fragmenttransferred from the short arm of chromosome 5B tochromosome 7A was extremely short, less than one-fifth of the arm length. Thus, the translocation wasassociated with an exchange of chromosome regionssubstantially differing in size.

To localize the breakpoints, we performed in situhybridization with two probes, pSc119.2 and pTa794(5S rDNA). The 5S rRNA genes are known to mark theshort arms of the chromosomes from homeologygroups 1 and 5 in wheat [28], while pSc119.2 producestwo, large telomeric and minor interstitial, distinct sig-nals in the short arm of chromosome 5B and does notinteract with chromosome 7A [29]. Chromosome 5B ofthe two translocation-containing accessions lacked alarge telomeric site of the pSc119.2 probe, an intenselystained 5S rDNA locus was in the center of the shortarm, and a minor pSc119.2 site was proximal to the 5SrDNA locus (Figs. 2a, 2b). At the same time, one of theA-genome chromosomes had a large pSc119.2 signal,which is normally absent. Thus, the breakpoint of chro-mosome 5B was distal of the minor pSc119.2 site andthe 5S rRNA gene locus, while the large pSc119.2 sitewas transferred to chromosome 7A (Fig. 2a). Theregion located distal of the 5S rDNA locus in the rear-ranged arm of chromosome 5B and corresponding tothe transposed fragment of chromosome 7A accountedfor approximately half of the arm length (Fig. 2b).Thus, the result nearly coincided with the GISH find-ings.

The structural similarity revealed for the rearrangedchromosomes with T7A:5B by C-banding [27] and insitu hybridization and the general similarity of the T.dicoccum karyotypes containing this translocationposes a question as to a common origin of these acces-sions. To study this issue, it was expedient to analyzethe within-species diversity of European T. dicoccumby statistical methods. As such methods, UPGMA andKohonen neural networks were chosen.

In the analysis of Kohonen neural networks, binarydivision at the first step produced two clusters (Fig. 3).The first cluster (cluster A) included almost all (ten) ofthe Volga–Balkan accessions, five Turkish accessionswith the normal karyotype, and four European acces-sions from Germany (three) and France (one). Theaccession INRA 26658 from France substantially dif-fered in chromosome C-banding patterns from theother emmer accessions. In addition, the accession dis-played a complete inversion of the pericentric hetero-chromatin complex of chromosome 4B. The inversionwas not found in tetraploid wheat species. Two of the

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DIVERSITY AND THE ORIGIN OF THE EUROPEAN POPULATION 1085

three German accessions of cluster A had an extremelysmall band in the long arm of chromosome 4A, as char-acteristic of Volga–Balkan accessions, and clusteredtogether with the Bulgarian accession k-14236 on thedendrogram.

The second cluster (B) included all European andnon-European accessions carrying the 7A:5B markertranslocation (26 accessions), five European accessions(one from Germany, two from Italy, and two fromFrance) with normal karyotypes, one Bulgarian acces-

(a) (b)d fb c e g h la

1

2

3

4

5

6

7

T7AS.7AL-5BS

T6AS.1BL

T5BL.5BS-7AL

T2AS.2AL-1BS

T1BS.6AL

T1BL.1BS-2AL

Fig. 1. Karyotypes of the emmer accessions from the INRA collection: a, 26642 (Germany); b, 27097 (Italy); c, 26659 (France);d, 27124 (Germany); e, 27123 (Germany); f, 27125 (Germany); g, 26658 (France); h, 27126 (Yugoslavia, before 1990); and i,(27088 Italy).

d fb c e g h la

1086

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DEDKOVA et al.

sion, and one Russian accession. It was not surprisingthat ANN analysis placed all emmer accessions withthe 7A:5B translocation in one cluster, since the acces-sions were similar not only in having the translocation(information on the translocation was not included inthe database), but also in the distribution of C-bandsthrough other chromosomes. The Bulgarian emmeraccession INRA 27090, which appeared in cluster B,substantially differed from the other Bulgarian acces-sions.

At the next step, we used UPGMA to estimate thegenetic relationships among European T. dicoccumaccessions. The resulting dendrogram included threeclusters: A, B, and C, which contained 33, 3, and 6accessions, respectively (Fig. 4). The accession 26658-FRA was not assigned to any cluster. Cluster Aincluded 33 accessions: all of the 26 accessions carry-ing the marker translocation (22 from Europe and 7from other countries) and six accessions lacking thetranslocation, which mostly originated from Western

Europe. Cluster B included three accessions of differ-ent origins. Cluster C included 16 accessions: nine ofthe Volga–Balkan group, five from Turkey, and threeWest European emmer accessions without the 7A:5Bmarker translocation. Interestingly, cluster B and C cor-responded in composition to cluster B isolated byKohonen neural networks.

To check the reliability of the clustering, we calcu-lated the mean values of the within- and between-clus-ter similarity indices (Table 2). It is clear that thewithin-cluster similarity should be significantly higherthan the mean between-cluster similarity index. Thewithin- and between-cluster similarity indices werecompared by Student’s t-test.

The mean weighted coefficient of correlationbetween Euclidean distances within the clusters wasRw = 0.29, the standard error was Sxw = 0.01.

The mean weighted coefficient of correlationbetween Euclidean distances between the clusters wasRb = –0.11, the standard error was Sxb = 0.01.

(c) (d)

(a) (b)10 ÏÍÏT7AS.7AL-5BS

1B

4A

2B3B

1A

6B

1B

4B2B

7B

4B6B

5A 5A

1A

4A7B

3B

1B

1B

1A

T5BL.5BS-7AL

T5BL.5BS-7ALT7AS.7AL-5BS

4A

4A

T7AS.7AL-5BS

T7AS.7AL-5BS

T5BL.5BS-7AL

T5BL.5BS-7AL

5A

5A

1A

T7AS.7AL-5BS

T7AS.7AL-5BS

4A T5BL.5BS-7ALT5BL.5BS-7AL

T5BL.5BS-7AL

T7AS.7AL-5BS

T7AS.7AL-5BS

T5BL.5BS-7AL

4A

Fig. 2. Examination of the structure of the 7A:5B translocation by in situ hybridization. (a, b) FISH on the T. dicoccum k-7146 chro-mosomes was performed with the DNA probes (a) pSc199.2 and (b) pTa794. (c, d) GISH was performed on the chromosomes ofthe T. dicoccum accessions (c) k-20989 and (d) k-7146.

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DIVERSITY AND THE ORIGIN OF THE EUROPEAN POPULATION 1087

Student’s t-test for a comparison of the within- andbetween-cluster distances was

tEb Ew–

Sxw2 Sxb

2+------------------------- 0.29 0.11–( )–

0.01----------------------------------- 40.0.= = =

The results showed that the coefficients of correla-tion between clusters were significantly lower than thecoefficients of correlation within clusters. The differ-ence was significant at P < 0.0001, suggesting the reli-ability of the clustering.

20993-TUR20967-TUR20968-TUR23036-YUG27088-ITA27119-YUG14380-TUR21007-TUR21433-DEU14236-BGR27125-DEU27126-YUG40030-YUG38917-YUG24482-RUS38904-YUG14999-POL21961-DEU26658-FRA27089-BGR36527-SWE*6807-FRA*900-FRA26651-CSK*23757-CHE*12946-CHE*898-FRA*26654-HUN*26639-DEU*26893-DZA*26894-DZA*27090-BGR27095-ITA27096-ITA30728-RUS21582-FRA*27123-DEU*35890-NLD*26647-FRA*26657-FRA*20638-ESP*26642-DEU*26648-FRA*20989-TUR*27076-ESP*26659-FRA*27097-ITA*27124-DEU*18623-DEU*20541-ESP*21416-ITA*7146-IRN*20410-ESP*

Ä

Ç

20993-TUR20967-TUR20968-TUR23036-YUG27088-ITA27119-YUG14380-TUR21007-TUR21433-DEU14236-BGR27125-DEU27126-YUG40030-YUG38917-YUG24482-RUS38904-YUG14999-POL21961-DEU26658-FRA27089-BGR36527-SWE*6807-FRA*900-FRA26651-CSK*23757-CHE*12946-CHE*898-FRA*26654-HUN*26639-DEU*26893-DZA*26894-DZA*27090-BGR27095-ITA27096-ITA30728-RUS21582-FRA27123-DEU35890-NLD*26647-FRA*26657-FRA*20638-ESP*26642-DEU*26648-FRA*20989-TUR*27076-ESP*26659-FRA*27097-ITA*27124-DEU*18623-DEU*20541-ESP*21416-ITA*7146-IRN*20410-ESP*

Ä

Ç

Fig. 3. Dendrogram of the T. dicoccum accessions carrying the 7A:5B translocation and originating from Europe, Turkey, and othergeographical regions. The dendrogram was constructed using artificial neural networks (Kohonen networks). The accessions withthe translocation are asterisked.

1088

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DEDKOVA et al.

The identical compositions of the clusters isolatedby UPGMA and a Kohonen neural network justify theisolation of two cytogenetically different groups ofEuropean emmer accessions on the basis of chromo-

some C-banding patterns. The results coincide with themorphological and botanical division of emmer into theVolga–Balkan and European groups. It is logical toassume that the division of European emmer into thetwo clusters reflects different origins of the populationsexamined.

The ancestors of the European group most likelyincluded plant that contained the 7A:5B translocation.All of the accessions carrying this translocation aresimilar in karyotype C-banding patterns regardless oftheir origin [10]. Thus, the presence of the translocationreflects a founder effect, rather than resulting frommore recent spontaneous hybridizations. It seemsunlikely that the translocation arose via several inde-pendent events. The distribution of the 7A:5B translo-cation through Europe (Fig. 5) agrees with the histori-cal, archeological, and genetic data that indicate thepaths of a spreading of cultivated emmer [1, 3, 4,11, 30].

The broad distribution of the 7A:5B translocation incultivated emmer populations suggests that the translo-

20993-TUR

20967-TUR20968b-TUR

23036-YUG

27088-ITA27119-YUG

14380-TUR

21007-TUR

21433-DEU

14236-BGR

27125-DEU

27126-YUG

40030-YUG38917-YUG

24482-RUS

38904-YUG

14999-POL21961-DEU

26658-FRA

27089-BGR

36527-SWE*

6807-FRA*

900-FRA26651-CSK*

23757-CHE*

12946-CHE*

898-FRA*

26654-HUN*

26639-DEU*

26893-DZA*

26894-DZA*27090-BGR

27095-ITA27096-ITA

30728-RUS

21582-FRA

27123-DEU

35890-NLD*

26647-FRA*

26657-FRA*

20638-ESP*

26642-DEU*

26648-FRA*

20989-TUR*

26659-FRA*

27097-ITA*

27124-DEU*

18623-DEU*20541-ESP*

21416-ITA*20410-ESP*

7146-IRN*

–0.14 0.11 0.36 0.61 0.86

27076-ESP*

A

B

C

Fig. 4. Dendrogram of the T. dicoccum accessions carrying the 7A:5B translocation and originating from Europe, Turkey, and othergeographical regions. The dendrogram was constructed by UPGMA clustering.

Table 2. Matrix of within- and between-cluster similarityindices (coefficients of correlation between Euclidean dis-tances)

Cluste A B C

A (n = 33) 0.29 0.02 0.15

B (n = 3) 0.40 0.00

C (n = 16) 0.29

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DIVERSITY AND THE ORIGIN OF THE EUROPEAN POPULATION 1089

cation belongs to the most ancient chromosome rear-rangements in tetraploid wheat. The question arises asto at which evolutionary step and in what species thetranslocation arose initially. It is possible that T7A:5Bfirst arose in wild emmer T. dicoccoides and then wastransmitted to the cultivated species during domestica-tion or via natural hybridization of local cultivatedemmer populations with the wild species that carriedthis rearrangement. It cannot also be excluded that thetranslocation arose in T. dicoccum at early steps of itsdomestication.

We examined several wild emmer populations fromTurkey (where emmer cultivation probably started) anddid not found any form with the translocation. How-ever, several genotypes with the 7A:5B translocationwere detected in the accessions originating from theMiddle East [27]. These findings agree with the pub-lished data that Middle East populations of wild emmerexerted a substantial effect on the T. dicoccum genepool of the Mediterranean countries [1]. It is possible toassume on these grounds that emmer appeared inEurope via an intricate path rather than directly fromTurkey. Initially, cultivated emmer populations might

migrate from the domestication center southward, tothe Middle East, where they spontaneously hybridizedwith wild emmer populations carrying the 7A:5B trans-location. In view of its apparent adaptive significance,the translocation might be fixed in T. dicoccum andmight spread more recently in the first emmer migra-tion wave to the Mediterranean countries and, then, tonorthward through Europe. To better understand thepaths of wheat distribution through Europe, it is neces-sary to study the cultivated emmer populations ofGreece and adjacent islands.

All of the Turkish accessions with the normal kary-otype clustered together with the Volga–Balkan group.In a larger T. dicoccum sample including hulled emmeraccessions from Ethiopia, India, and Morocco, Turkishaccessions are most similar to the same Europeanemmer accessions as in the dendrogram [10]. This find-ing indicates that this group of T. dicoccum accessionsprobably originates from Turkish emmer lacking themarker translocation. It is clear that these populationscorrespond to a second wave of wheat migrationthrough Europe, which originated from a founder pop-

Sw

eden

German

y

Russia

Poland

France

Spain

Morocc

o

Algeria

ò‚ÂȈ.Italy

CzechiaSlovakia

Hungary

YugoslaviaBulgaria

Turkey

Iran

Saudi Arabia

Ethiopia

India

Translocation 7A:6Bwas not found

Translocation 7A:5Bwas found in at least one accession

Fig. 5. Distribution of the T. dicoccum accessions carrying the 7A:5B translocation.

1090

RUSSIAN JOURNAL OF GENETICS Vol. 45 No. 9 2009

DEDKOVA et al.

ulation other than the ancestor of West Europeanemmer.

Accessions with and without the translocation werefound in Italy, France, and Germany. A possible expla-nation is that normal karyotypes were as fit as translo-cation-containing ones under the given environmentalconditions or that emmer was repeatedly introduced inthese regions later.

ACKNOWLEDGMENTS

We are grateful to J. Koenig (INRA, France) for thematerial for this work.

This work was supported by the Russian Foundationfor Basic Research (project no. 08-04-00302) and theprogram “Fundamental Basis for Managing BiologicalResources” of the Russian Academy of Sciences.

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