An integrated DArT-SSR linkage map of durum wheat

An integrated DArT-SSR linkage map of durum wheat

Paola Mantovani AElig Marco Maccaferri AElig Maria Corinna Sanguineti AEligRoberto Tuberosa AElig Ilaria Catizone AElig Peter Wenzl AElig Brent Thomson AEligJason Carling AElig Eric Huttner AElig Enzo DeAmbrogio AElig Andrzej Kilian

Received 20 March 2008 Accepted 25 June 2008 Published online 19 July 2008

Springer Science+Business Media BV 2008

Abstract Genetic mapping in durum wheat (Triti-

cum durum Desf) is constrained by its large genome

and allopolyploid nature We developed a Diversity

Arrays Technology (DArT) platform for durum wheat

to enable efficient and cost-effective mapping and

molecular breeding applications Genomic represen-

tations from 56 durum accessions were used to

assemble a DArT genotyping microarray Microsatel-

lite (SSR) and DArT markers were mapped on a durum

wheat recombinant inbred population (176 lines) The

integrated DArT-SSR map included 554 loci (162

SSRs and 392 DArT markers) and spanned 2022 cM

(5 cMmarker on average) The DArT markers from

durum wheat were positioned in respect to anchor

SSRs and hexaploid wheat DArT markers DArT

markers compared favourably to SSRs to evaluate

genetic relationships among the durum panel with

1315 DArT polymorphisms found across the acces-

sions Combining DArT and SSR platforms provides

an efficient and rapid method of generating linkage

maps in durum wheat

Keywords DArT Durum wheat Linkage map SSR

Abbreviations

DArT Diversity arrays technology

chr Chromosome

cv Cultivar

lsquoC 9 Lrsquo Colosseo 9 Lloyd

ITMI map Ta-SyntheticOpata-BARC map

(Song et al 2005)

lsquoK 9 Srsquo Kofa 9 Svevo

PCR Polymerase chain reaction

RIL Recombinant inbred line

SSR Simple sequence repeat or

microsatellite marker

Introduction

Durum wheat (Triticum turgidum L var durum) is an

allotetraploid species (AABB genome 2n = 4X =

28) mainly cultivated in Mediterranean and in semiarid

areas of the world Even though durum wheat typically

accounts for5ndash8 of the total world wheat produc-

tion (World Grain Statistics wwwigcorguk) its

P Mantovani M Maccaferri M C Sanguineti R Tuberosa I Catizone

Department of Agroenvironmental Sciences

and Technology University of Bologna

Viale Fanin 44 40127 Bologna Italy

I Catizone P Wenzl B Thomson J Carling E Huttner A Kilian (amp)

Diversity Arrays Technology PL and Triticarte Pty Ltd

1 Wilf Crane Cr Yarralumla Canberra ACT 2600

Australia

e-mail akiliandiversityarrayscom

E DeAmbrogio

Societa Produttori Sementi Bologna Research Division

Via Macero 1 40050 Argelato BO Italy

123

Mol Breeding (2008) 22629ndash648

DOI 101007s11032-008-9205-3

importance is related to the fact that durum grain is

mainly used for human consumption However

genetics of important agronomic and quality traits of

durum wheat has been poorly investigated with respect

to other cereals Detailed genetic information on

durum adaptation and grain quality is required as a

basis in breeding programs Molecular markers are

efficient tools to speed up crop improvement (Lang-

ridge 2005 Varshney and Tuberosa 2007) and for the

construction of molecular linkage maps the first step in

the genetic dissection of target traits

Microsatellites (simple sequence repeats SSRs) are

PCR-based markers characterised by a high level of

polymorphism that permits to discriminate among

cultivars and even among closely related wheat

breeding lines (Plaschke et al 1995 Maccaferri et al

2007) In addition to their high polymorphism SSRs

are usually single locus sites an important feature

when dealing with allopolyploid species SSRs have

rapidly become the markers of choice for the con-

struction of genetic maps in wheat (Roder et al 1998

Somers et al 2004 Sourdille et al 2004) up to now

several hundred SSR primer pairs have been developed

for all three genomes of wheat (httpwheatpwusda

govGG2indexshtml) While SSRs are standard PCR-

based markers and can be considered as proven anchor-

markers their suitability for high-throughput mapping

does not favourably compare to the new single

nucleotide polymorphism (SNP)-based genotyping

techniques (Kilian et al 2005) Moreover SSR-mul-

tiplexing requires an extensive additional optimisation

(Hayden et al 2008) Development of massive SNP

resources from the expressed portion of the wheat

genome amenable to fully automated genotyping is a

difficult task for wheat due to its low frequency of

sequence polymorphism and its allopolyploid nature

(Koebner and Summers 2003 Somers et al 2003)

Diversity arrays technology (DArT) for which proof

of concept was first reported by Jaccoud et al (2001)

is becoming increasingly adopted in many species

(most current list of species with DArT technology

developed can be found at wwwdiversityarrayscom)

The technology combines a complexity reduction

method (Wenzl et al 2004) with hybridization-based

polymorphism detection using high-throughput solid-

state platforms and has the potential to generate hun-

dreds of high-quality genomic dominant markers with

a cost- and time-competitive trade-off (Kilian et al

2005)

In contrast to hexaploid bread wheat (Triticum

aestivum L AABBDD genome) for which several

linkage maps have been developed mapping in durum

wheat has received relatively little attention The first

linkage map of durum wheat based on 65 recombinant

inbred lines (RILs) and RFLP markers was reported by

Blanco et al (1998) SSRs from hexaploid wheat

(Roder et al 1998) were subsequently integrated into

this linkage map (Korzun et al 1999) More recently

intra- and inter-specific linkage maps based on RFLPs

SSRs and AFLPs have been developed (Peng et al

2000 Nachit et al 2001 Maccaferri et al 2008)

These studies have shown a generally good conserva-

tion of marker co-linearity as compared to the A and B

genomes of hexaploid wheat

Dedicated DArT genotyping platforms have been

produced for bread wheat (Akbari et al 2006 Semagn

et al 2006) The identification of polymorphic DArT

markers directly obtained from durum wheat by

generating genomic representations from diverse

durum accessions followed by mapping the polymor-

phisms on a durum mapping population and by

genotyping a panel of durum accessions serves as a

basis for broadening the use of DArT markers for

durum wheat This study presents the mapping in

durum wheat of 392 new DArT markers that were

anchored to the wheat map using 162 SSRs DArT

markers were then used to profile a panel of durum

accessions subsequently the genetic relationships

based on DArT markers were compared to those

obtained with highly polymorphic SSRs

Materials and methods

Plant material

A panel of 56 diverse durum wheat accessions was

selected to sample the genetic diversity present in the

cultivated durum gene pools from the most important

durum wheat production areas Additionally the

accessions were chosen in order to explore various

levels of genetic relationships based on previous

results obtained with SSRs (Maccaferri et al 2005)

The panel was assembled with durum cultivars (cvs)

and accessions from (i) Southern Europe (Italy

Spain and France) (ii) the International Maize and

Wheat Improvement Center (CIMMYT) (iii) the

International Center for Agricultural Research in the

630 Mol Breeding (2008) 22629ndash648

123

Dry Areas (ICARDA) and (iv) North-America

(mainly from Canada North Dakota and USA) The

panel included accessions used as parents of mapping

populations recently developed in Italy (Colosseo

Lloyd Svevo and Kofa) and in Australia (Tamaroi

Wollaroi line 139 and line 149) Details on the

accessions are reported in Table 1

A population of 176 F67 recombinant inbred lines

(RILs) were used to generate a combined DArT-SSR-

based linkage map The RILs were obtained by Societa

Produttori Sementi (Bologna Italy) from a cross

between the Italian durum wheat cv Colosseo (Mexarsquos

mutant 9 Creso) and the North American cv Lloyd

(Cando 9 Edmore) Colosseo was selected by Prose-

me Srl (Enna Italy) for high yield potential

resistance to leaf rust and adaptation to climatic

conditions of central Italy Lloyd cv released by

North Dakota State University is a semi-dwarf

photoperiod-sensitive durum wheat and has a genetic

profile consistent with those of the North American

durums (Maccaferri et al 2005 2007 Mantovani

et al 2006)

Molecular analysis

DNA extraction

High-quality genomic DNA was extracted from ca

50 mg of freeze-dried leaf tissue from young leaves

using the cetyl-trimethyl-ammonium bromide (CTAB)

method of Saghai-Maroof et al (1984) The DNA

concentration was adjusted to 20 ngll and DNA

samples were stored at -20C

Diversity arrays technology markers

DArT markers were obtained substantially following

the procedures described for bread wheat by Akbari

et al (2006) A brief description is presented below

with special emphasis on several aspects of DArT

development in durum wheat

Preparation of DArT arrays

Two DArT arrays were assembled in the course of

this study a bread wheat array as described in

Akbari et al (2006) and a dedicated durum array

The durum array was assembled using random clones

derived from a genomic representation composed of

the 56 accessions reported in Table 1 These two

arrays were used to assay according to Akbari et al

(2006) the Colosseo 9 Lloyd (lsquoC 9 Lrsquo) mapping

population and each of the 56 durum accessions

Properly formatted marker names have already been

attributed to the bread wheat DArT-markers (wPt-)

Durum wheat markers are currently being referred to

with their clone ID numbers generated by DArTdb

the Laboratory Information Management (LIMS)

System at Diversity Arrays TechnologyTritcarte

Pty Ltd In the near future they will be named

according to the DArT marker naming convention

For each of these arrays a genomic representation

was generated from a mixture of wheat accessions

using the complexity reduction method described by

Wenzl et al (2004) The procedure involved diges-

tion of 20ndash100 ng of a mixture of DNA samples with

two units of PstI and two units of TaqI (NEB

Beverly MA USA) A PstI adapter (50-CAC GAT

GGA TCC AGT GCA-30 annealed with 50-CTG GAT

CCA TCG TGC A-30) was simultaneously ligated to

the digested DNA with T4 DNA ligase (NEB) A 1 ll

aliquot of the ligation product was used as a template

in 50 ll amplification reactions with DArT-PstI

primer (50-GAT GGA TCC AGT GCA G-30) under

the cycling conditions described by Wenzl et al

(2004)

A library was prepared from the amplification

products essentially as described by Jaccoud et al

(2001) with modifications as in Wenzl et al (2004)

Inserts were amplified from individual clones so that

part of the polylinker region of the cloning vector was

co-amplified (Jaccoud et al 2001) The amplification

reactions were dried at 37C washed with 70 ethanol

and dissolved in a new spotting buffer developed

specifically for Erie Scientific poly-L-lysine micro-

array slides (Wenzl et al in preparation) The

amplification products were printed on poly-L-lysine-

coated slides (Erie Scientific Portsmouth NH USA)

using a MicroGridII arrayer (Biorobotics Cambridge

UK) After printing the slides were denatured by

incubation in hot water (95C) for 2 min and dried by

centrifugation

Genotyping of individual DNA samples

The genomic representations of single wheat acces-

sions were generated with the same complexity

reduction method used to prepare the library spotted

Mol Breeding (2008) 22629ndash648 631

123

Table 1 List of the 56 durum wheat accessions (cultivars and breeding lines) and their origin and registration details

Genotype Code no Registration Pedigree Sourcea

Country Year

Capeiti 8 10 Italy 1940 CappelliEiti 3

Claudio 12 Italy 1998 GraziaCIMMYT line 4

Colosseo 13 Italy 1995 Mexarsquos mutant Creso 4

Creso 14 Italy 1974 Yt 54-N10-B23TC603 Cp B 14 4

Duilio 16 Italy 1984 CappelliAnhingaFlamingo 2

Grazia 18 Italy 1985 M 6800127Valselva 4

Meridiano 34 Italy 1999 SimetoWB881DuilioF21 4

Messapia 35 Italy 1982 MexCraneTito 4

Iride 20 Italy 1996 Altar 84Ares = Ionio 4

Levante 26 Italy 2002 G80PicenoIonio 4

Ofanto 39 Italy 1990 AppuloAdamello 4

Simeto 51 Italy 1988 Capeiti 8Valnova 4

Svevo 52 Italy 1996 CIMMYTrsquos SelectionZenit 4

Saragolla 48 Italy 2002 IridePSBline0114 3

Senatore Cappelli 50 Italy 1930 Strampellirsquo selection from Jennah Khetifa 1

Trinakria 55 Italy 1970 B 14Capeiti 8 4

Valforte 57 Italy 1980 Yt54-N10B2BYLD390 II

145873Cappelli2Yuma

5

Orjaune 42 France 1995 miraduridyn81-04 6

Nefer 37 France 1996 164Keops 4

Neodur 38 France 1987 184-7ValdurEdmore 4

AC Morse 1 Canada 1996 RL 7196D84328 7

AC Pathfinder 2 Canada 1998 DT367WB881 7

Kyle 23 Canada 1984 WakoomaDT320WakoomaDT322 7

Ben 9 US-ND 1996 D8024Monroe 8

Lloyd 31 US-ND 1983 CandoEdmore 8

Maier 33 US-ND 1998 D8193D8335 8

Langdon 25 US-ND 1956 MindumCarletonKhapli3Heiti

StewartMindum Carleton4Stewart

5Carleton

8

Kofa 56 US-AZ 1990 dicoccum alpha pop-85 S-1 9

Reva 47 US-AZ 1990 WWW MSFRS Pop 10

Don Pedro 15 Spain 1990 CARCAUK 12

Altar 84 3 CIMMYT 1984 Ruff lsquolsquoSrsquorsquoFGO lsquolsquoSrsquorsquoMexicali 753SHWAlsquolsquoSrsquorsquo 4

Mexicali 75 36 CIMMYT 1975 61130LeedsJorilsquolsquoSrsquorsquo3GDOVZ469 4

Plata 16 44 CIMMYT 1990 Altar 84Yavaros 79SHWA 4

Rascon2 Tarro 46 CIMMYT 1990 Altar 84CMH82ARancoHUI2AIXKKV5 4

Aghrass 1 4 ICARDA ndash ndash 11

Azeghar 2 7 ICARDA ndash ndash 11

Belikh 2 8 ICARDA 1987 CRSTK 11

Aw12Bit 6 ICARDA ndash CIT105801FGOSCOT 11

Cham 1 = Waha 11 ICARDA 1984 PLCRUF2GTARTTE 11

Gidara 2 17 ICARDA ndash OmrabiSTKJO330365 11


123

on the array These genomic representations were

ten-fold concentrated by precipitation with one

volume of isopropanol and denatured at 95C for

2 min The samples were then labelled with 01 ll of

cy3- or cy5-labelled dUTP and unlabelled random

decamers (Amersham Biosciences Castle Hill NSW

Australia) using the exo-Klenow fragment of Esch-

erichia coli DNA polymerase I (NEB) Labelled

representations also called targets were added to

50 ll of a 5051 mixture of ExpressHyb buffer

(Clontech Mountain View CA USA) 10 gl herring

sperm DNA (Promega Annandale NSW Australia)

and the 6-FAM-labelled polylinker fragment of the

plasmid that was used for library preparation The

polylinker fragment was used as a reference to

determine for each clone the amount of DNA

spotted on the array (Jaccoud et al 2001) The

hybridisation mixtures were denatured hybridised to

microarrays overnight at 65C and slides were

washed according to Jaccoud et al (2001)

Table 1 continued


Country Year

Korifla = Cham 3 22 ICARDA 1987 DS15GEIER 11

Quadalete 45 ICARDA ndash ndash 11

Lahn 24 ICARDA ndash Yavaros 79SHWA 11

Loukos 1 32 ICARDA ndash FGOCITFGO4531 11

Omrabi 5 40 ICARDA 1993 JOHaurani 11

Omruf 2 41 ICARDA ndash OmrabiRUFF 11

Ouaserl 1 43 ICARDA ndash ndash 11

Sebah 49 ICARDA ndash ndash 11

Zeina 1 60 ICARDA ndash SRC_32180 11

Haurani 19 ICARDA ndash Local landrace selection from Syria 11

Jennah Khetifa 21 ICARDA ndash Local landrace selection from North Africa 11

Astrodur 5 Austria 1991 ValdurPandurValgerardo 13

Wollaroi 58ndash59 Australia ndash TAMB-17Kamilaroi 14

Tamaroi 53 Australia ndash ndash 14

Line 139 27ndash28 Australia ndash ndash 14


a Seed sources

1 Ente Nazionale Sementi Elette (ENSE) Milano Italy

2 Societa Italiana Sementi (SIS) Bologna Italy

3 Istituto del Germoplasma Bari Italy

4 Societa Produttori Sementi Bologna (SPB) Bologna

5 Ist Sper Cerealicoltura Sezione di Foggia Foggia Italy

6 Groupe drsquo Etude et de controle des Varietes et des Semences (GEVES) GEVES La Miniere Guyancourt Cedex France

7 Agriculture and Agri-Food Canada Semiarid Prairie Agriculture Research Centre (AAFC SPARC) Swift Current SK Canada

8 North Dakota State University (NDSU) Fargo North Dakota USA

9 Western Plant Breeder (WPB) Bozeman Montana USA

10 World Wide Wheat (WWW) Phoenix Arizona USA

11 ICARDA International Centre for Agricultural Research in the Dry Areas Aleppo Syria

12 UdL-IRTA Institute of Agro-food Research and Technology IRTA and University of Lleida Lleida Spain

13 Probstdorfer Saatzucht Probstdorfer Austria

14 CSIRO Plant Industry Canberra Australia


123

Image analysis and polymorphism scoring

Slides were scanned using Tecan LS300 (Grodig

Salzburg Austria) confocal laser scanner The TIF

images derived from the slide scanning were analysed

using DArTsoft version 73 (Cayla et al in prepara-

tion) a dedicated software package developed at DArT

PL which is available to DArT network members

(wwwdiversityarrayscomdartnetworkhtml) DArT-

soft was used to automatically analyse batches of up to

96 slides to identify and score polymorphic markers

Briefly the relative hybridisation intensity of each

clone on each slide was determined by dividing the

hybridisation signal in the target channel (genomic

representation) by the hybridisation signal in the ref-

erence channel (polylinker) Clones with variable

relative hybridisation intensity across slides were

subjected to fuzzy k-means clustering to convert rela-

tive hybridisation intensities into binary scores

(presence versus absence)

Simple sequence repeat markers

A total of 550 genomic SSR primer pairs were screened

using the two parental lines and a progeny sample of

four lines Markers were prevalently chosen within the

public SSRs (httpwheatpwusdagov) Table 2 pre-

sents the list of the screened SSR markers The

majority of the SSRs used in this study was mapped in a

durum wheat mapping population (249 RILs from the

cross lsquoKofa 9 Svevorsquo Jurman et al unpublished

data) herein indicated as lsquoK 9 Srsquo as well as on the

bread wheat Ta-SSR-2004 consensus SSR map

(Somers et al 2004) and on the Ta-SyntheticOpata-

BARC map (Song et al 2005) hereafter referred to

as ITMI map SSR primer sequences of BARC

CFA CFD DuPW KSUM and WMC primerrsquos

sets are publicly available on the GrainGenes Triti-

ceae database (httpwheatpwusdagov) the primer

sequences of most of the WMS (gwm loci) SSRs are

also catalogued in GrainGenes however for a small

subset (14 out of 65 gwm mapped loci Xgwm783 856

947 1009 1034 1038 1045 1084 1184 1198 1246

1249 1278 1570) the primer sequences of these SSRs

were kindly provided by Dr Martin W Ganal (Trait

Genetics GmbH Am Schwabeplan 1b Gatersleben

Germany) and by Dr Marion Roder (Institut fur

Pflanzengenetik und Kulturpflanzenforschung IPK

Gatersleben Germany) These primers generated SSR

loci that were not previously mapped either in the

Ta-SyntheticOpata-SSR or in the Ta-SSR-2004

SSRs were amplified from 200 ng of genomic

DNA in 25 ll reactions containing 1X PCR buffer

(500 mM potassium chloride and 100 mM TrisndashHCl

at pH 83) 15 mM MgCl2 06 lM of both forward

and reverse primers 016 mM dNTPs and 1 unit of

AmpliTaq DNA Polymerase (Applied Biosystems

Foster City CA USA) PCR amplifications were

performed on a 2720 Perkin-Elmer thermocycler

(Norwalk CT USA) using the following program

94C (3 min)20 cycles of 94C (45 s) 61C

(decreasing by 05C per cycle to a minimum of

51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)

During polymorphism screening the PCR prod-

ucts were separated on a 45 polyacrylamide gel

and visualized by silver-staining (Bassam et al

1991) Most of the polymorphic SSRs were amplified

using 50-labelled forward primers (IR700 or IR800)

and analysed on a 4200 Gene Read IR2 Automated

Genotyper (LI-COR Lincoln NE USA) Typically

SSR reactions were multiplexed in pairs based on

their annealing temperature and amplicon size SSR

markers were used as anchors in map construction

Table 2 SSR markers

screened for polymorphism

between cvs Colosseo and

Lloyd

SSR class Number References

Barc 130 Song et al (2002 2005)

Cfa 30 Sourdille et al (2003) Guyomarcrsquoh et al (2002)

Cfd 20 Sourdille et al (2003) Guyomarcrsquoh et al (2002)

DuPw 5 Eujayl et al (2002)

Ksum 5 Yu et al (2004)

Wmc 175 Gupta et al (2002) httpwheat pw usda govggpagesSSRWMC

Gwm 165 Roder et al (1998) Martin Ganal IPK Gatersleben Germany

EST-SSR 20 Graingenes httpwheat pw usda govITMIEST-SSR


123

and their relative order was compared with the

reference wheat maps

Integrated DArT-SSR linkage map construction

The scores of all polymorphic DArT and SSR markers

were converted into genotype codes (lsquoArsquo lsquoBrsquo) accord-

ing to the scores of the parents heterozygotes were

recorded as missing data EasyMap 01 a program

being developed at Diversity Arrays Technology PL

was used to build a genetic map for the lsquoC 9 Lrsquo RIL

population The program is designed to automate

genetic mapping of BC1 DH and RIL populations

(Wenzl et al in preparation) EasyMap combines pre-

map and post-map quality-filtering steps for both

markers and lines with a suit of algorithms for defining

linkage groups the RECORD algorithm for optimising

marker order and an algorithm to identify potential

genotyping errors with a logarithm-of-odds ratio in

favour of error (LODerror) above a user-provided

threshold (Lincoln and Lander 1992 van Os et al

2005) The program starts by establishing an initial

marker order as if all markers belonged to a single

linkage group Blocks of contiguous markers are then

assigned to different linkage groups based on a

recombination-frequency threshold (REC) and a ten-

sion threshold (TENSE) REC is derived from a user-

defined probability value by modelling the expected

degree of pseudo-linkage between telomere pairs

TENSE is computed by comparing the two-point

Kosambi distance estimate between adjacent markers

with a multi-point estimate computed using a multiple-

regression algorithm (Stam 1993) An initial map was

built using P = 001 (14 chromosomes176 lines REC = 037) TENSE = 12 cM and LODerror = 40

for identifying potential genotyping errors Linkage

groups were assigned to chromosomes based on the

known position of SSR markers This assignment

allowed us to link some chromosome (chr) regions that

at the P = 001 level appeared unlinked The same

data matrix used to construct the integrated SSR-DArT

durum wheat linkage map was also utilised for

segregation distortion analysis by means of JoinMap

v4 (van Ooijen 2006) For each polymorphic marker

the chi-square test was used to identify markers

deviating from the 11 expected segregation markers

showing significant segregation distortion (P B 001)

were classified as skewed

Diversity analysis

Set of accessions

The data matrix containing the 01 scores of the

polymorphic DArT markers found among the durum

accessions was analysed with DARwin 50 software

using the lsquosingle datarsquo option (Perrier et al 2003 Perrier

and Jacquemoud-Collet 2006) Genetic distances were

estimated using the Jaccard dissimilarity index Jac-

cardrsquos dissimilarity index is obtained as follows

J0 frac14 M01 thornM10

M01 thornM10 thornM11

where M11 represents the total number of marker

comparisons (loci being compared) where both

accessions i and j have an attribute of 1 (double

presence of the same allele) M01 represents the total

number of marker comparisons where accession i

has an attribute of 0 and accession j is 1 M10

represents the total number of marker comparisons

where accession i has an attribute of 1 and accession

j is 0

As it can be noted M00 cases are not considered in

the Jaccardrsquos index because of the dominant nature

of the DArT markers that in germplasm collections

of diverse accessions does not allow for the

assumption of allelic identity in the M00 cases

The first two principal coordinates of the resulting

Jaccard matrix were extracted to display the diversity

structure in a two-dimensional plane In addition an

unweighed neighbour-joining tree was built from the

Jaccard matrix and its robustness was assessed by

bootstrapping (resampling no = 1000)

Comparison between marker types

The neighbour-joining tree analysis described in the

previous section was repeated on a subset of 31 durum

accessions that had previously been genotyped with

103 SSR markers (Maccaferri et al 2006) The corre-

sponding SSR dataset was analysed in a similar way

using the lsquoallelic datarsquo option and the lsquosimple-matching

distancersquo to construct an alternative dissimilarity

matrixneighbour-joining tree The dissimilarity index

based on simple matching is suited to SSRs which are

mostly codominantly inherited


123

SM frac14 mn

where m = number of loci being compared with

different allelic attributes between accessions i and j

n = total number of loci being compared excluding

allelic pairs with missing data

Since each high-quality DArT marker represents a

unique locus the two genetic dissimilarity indices

that were herein used for DArT and SSR markers

allowed to evaluate diversity based on the same

concept ie the evaluation of the exact proportion of

loci with dissimilar alleles over the total number of

loci being compared for each accession pair

Mantel (1967) with a permutation matrix strategy

was used to generate statistical significances for

correlation measures of similarity between distance

matrices

The test criterion used is

Z frac14Xn

ifrac141

Xn

jfrac141

AijBij

where Aij and Bij are the off-diagonal elements of the

two genetic dissimilarity matrices (A and B) If the

two matrices show similar relationships then Z should

be higher in comparison to what one would expect by

chance The significance test has been performed by

comparing the observed Z-value with its permutated

distribution Ten-thousand random permutations were

carried out The correlation coefficient r is mono-

tonically related to Z and has the advantage that is

expressed in standardized units

Results

After screening of over 25000 random genomic

wheat clones with a range of durum accessions we

identified 2304 polymorphic durum DArT markers

All these markers can be typed in a single assay on a

cost-effective technology platform The frequency of

markers (approximately 9) is similar to what we

found in hexaploid wheat (Akbari et al 2006)

Importantly all the durum markers can be evaluated

on a single array with approximately 5000 markers

polymorphic in hexaploid wheat (Kilian et al unpub-

lished data) as the method of complexity reduction is

the same (PstITaqI) Below we present the perfor-

mance of the newly developed markers in genetic

mapping and diversity analysis applications

An integrated DArT-SSR linkage map

DArT-SSR map

Among the 550 SSR markers used to screen for

polymorphism between the parental lines (Table 2)

249 (453) were polymorphic One hundred and forty-

five polymorphic SSRs were chosen based on their

known position (Somers et al 2004 Song et al 2005) in

order to ensure fairly good wheat genome coverage and

to avoid closely linked multiple loci These selected

SSRs were genotyped on the entire RIL population 53

specifically amplified the expected single-locus frag-

ment ca 40 amplified one or a few additional mono-

morphic fragments and ca 7 (BARC101 BARC340

BARC353 CFA2163 CFA2164 GWM112 GWM

132 GWM344 GWM443 WMC85 WMC405 WMC

500 and WMC505) amplified from one to three

additional polymorphic fragments leading to a total of

162 SSR loci

Among the 662 polymorphic loci (500 DArT

markers and 162 SSRs) used for assembling the

linkage map 554 loci (392 DArT markers and 162

SSRs) were distributed on 19 linkage groups with gaps

left on chrs 2A 2B 3A and 7A

The final map (Fig 1) spanned a total length of

2022 cM 7B was the longest chromosome

(2214 cM) while the shortest was 4A (880 cM) and

the average chromosome length was 1183 cM The

total number of mapped loci per chromosome ranged

from 12 (chr 5A) to 64 (chr 3B) with an average of

396 loci With regard to the two classes of markers the

number of locichromosome ranged from 1 (chr 5A) to

51 (chr 3B) for the DArT loci and from 7 (chr 4A) to

20 (chr 1B) in the case of SSR loci The marker density

on the map (57 cMmarker on average) varied from

29 to 97 cMmarker on the linkage group assigned to

chr 2BL and chr 5A respectively Map distance

between adjacent markers varied from 03 to 468 cM

and 71 of the intervals (278 out of 391 intervals) were

5 cM There were 19 chr regions with an intermar-

ker distance larger than 20 cM the largest distance

between adjacent markers was observed on the peri-

centromeric portion of chr 3B (468 cM) All these

considerations on average chr length and marker

density disregard the two small linkage groups (25 and

89 cM) assigned to chr 7AL Moreover to calculate

marker density each group of co-segregating markers

was considered as a single marker position to avoid


123

artifacts leading to higher density than the actual the

217 co-segregating markers (206 DArT and 11 SSR

markers) were mapped in 76 groups distributed over all

the chromosomes except for 5A and 5B (Fig 1)

DArT clusters were found in all the durum chro-

mosomes except on 5A where only one DArT marker

was mapped More precisely DArT clustering was

present on the telomeric regions of all chromosomes

except for 4B and on the peri-centromeric portion of

chrs 2B 3B 4B and 6B On the contrary only few SSR

clusters were identified around the centromeric region

of chrs 1B 2A 3A and 6B

Several differences in terms of map length number

and density of markers were observed among homo-

eologous groups Groups 3 and 4 showed the highest

(3586 cM) and shortest (2047 cM) map length

respectively The number of mapped markers was the

highest in group 6 (113 loci) whereas homoeologous

group 5 had the lowest number of markers (30 loci) and

the lowest marker density (91 cMmarker) More

precisely in group 5 the number of SSRs was twice the

number of DArT markers (20 and 10 respectively)

with only one DArT marker mapped on chr 5A and

nine on chr 5B

Map length of genomes A and B was 905 and

1117 cM respectively with 235 markers (163 DArT

and 72 SSR markers) mapped on the A genome and

319 markers (229 DArT and 90 SSR markers) on the

B genome leading to a comparable marker density

(61 and 53 cMmarker respectively)

Finally the 176 RILs of the lsquoC 9 Lrsquo mapping

population had on average 27 plusmn 5 scorable cross-

over events (mean plusmn SD computed by subtracting

potential genotyping errors) with a range of variation

comprised between 12 and 55 The average number

of scorable crossover eventsRIL corresponds to

approximately 2 (191 plusmn 038) crossover events per

chromosome

Segregation distortion

Segregation analysis data indicated that 455 of the

alleles were inherited from Colosseo and 468 from

Lloyd with a residual of missing data (genotypes

scored either missing or heterozygote) of 77

Significant (P 001) segregation distortion was

detected for 265 (147 markers) of the mapped

markers namely 108 DArT markers and 39 SSRs

which correspond to 275 and 240 of the total

DArT and SSR markers used for map construction

respectively The skewed markers occurred in all

chromosomes (Fig 1) except for chrs 5A and 5B the

chromosome with the highest number of skewed

markers (33) was 3B Markers displaying segregation

distortion in favour of Lloyd (82) were more

numerous compared to those with allele ratio in

favour of Colosseo (61) Skewed markers favouring

Lloyd were found on chrs 6A and 7B while those

favouring Colosseo were mapped on chrs 1A 4A 4B

and 6B Additionally chrs 1B 2A 2B 3A 3B and

7A showed skewed markers favouring both Colosseo

and Lloyd These marker loci with distorted segre-

gation were not randomly distributed 130 markers

were clustered in 15 regions on several chromo-

somes nine regions showed segregation distortion in

favour of Colosseo and six other regions had an

excess of alleles from Lloyd Moreover on chrs 1A

2B 3A 3B 7A and 7B the regions with distorted

segregation spanned more than 20 cM each

Map comparison

The position of the 554 DArT and SSR loci mapped in

this study was compared with that already available in

other maps of bread and durum wheat DArT markers

were referred to the bread wheat maps published by

Akbari et al (2006) Semagn et al (2006) and Crossa

et al (2007) while SSRs were referred to the bread

wheat consensus map (Somers et al 2004) and the

ITMI map (Song et al 2005) A total of 229 markers

(98 DArT and 131 SSR markers) out of the 554 mapped

on the lsquoC 9 Lrsquo map were present on one or more of the

already mentioned wheat maps

Ninety-eight DArT markers were reported on at

least one of the maps described by Akbari et al

(2006) Semagn et al (2006) and Crossa et al

(2007) In particular 88 out of 201 DArT markers

that were mapped from the hexaploid wheat array

(wPt-markers) were also present in the integrated

map published by Crossa et al (2007) These DArT

markers were used as anchor markers as in the case of

SSRs None of the wPt-DArT markers located on the

lsquoC 9 Lrsquo chrs 2A 4B 5A and 5B were in common

with those reported by Crossa et al (2007) while

only two wPt-DArT markers on chr 2A were in

common with Akbari et al (2006) Considering the

remaining chromosomes there were on average ca

seven anchor wPt-markers per chromosome


123


123

The map position of most of the SSR loci for the

lsquoC 9 Lrsquo population showed generally good consis-

tency to the reference maps Marker order on ten

chromosomes (2A 2B 3B 4A 4B 5A 5B 6A 7A

and 7B) was in fairly good accordance with the

consensus map SSR order on chr 1A was the same as

in the consensus map except for the markers at the

telomeres where the Xgwm33 and Xgwm136 loci

(telomeric 1AS) were found to be inverted as compared

to reference maps while the interval between Xgwm99

and Xbarc158 (telomeric 1AL) was in agreement only

with the ITMI map Chr 1B showed a good corre-

spondence with the consensus map apart from the

interval Xgwm11ndashXwmc419 where the SSR order was

more similar to that of the ITMI map The SSR loci on

the telomeric region of chr 3A (Xbarc310 Xbarc12

and Xbarc51) while absent on the consensus map

showed similar locations on the ITMI map the position

of the markers mapped to the pericentromeric portion

of chr 3A corresponds quite well with that reported by

Somers et al (2004) Finally several differences with

respect to both reference maps were found for the

interval Xgwm508ndashXgwm193 on chr 6B a detailed

analysis of the recombination frequencies between

pairs of markers within this interval (data not pre-

sented) validated the orientation herein reported

Among all the mapped SSRs 85 have an assigned

physical location (Sourdille et al 2004 Goyal et al

2005 Song et al 2005) The SSRs with physical

location were present on all chromosomes and were

mapped on the designated chromosome arms On the

lsquoC 9 Lrsquo map 31 SSRs were mapped in addition to

those reported by Somers et al (2004) and Song et al

(2005) The chromosomal location of 14 of these

markers is publicly available (httpwheatpwusda

govcgi-bingraingenesbrowsecgiclass=marker)

ten of them were located on the expected chromosome

and four mapped on a different chromosome The

CFA2163 primers amplified two loci one of which

indicated as Xcfa2163a was mapped for the first time

on the lsquoC 9 Lrsquo map (chr 3A) The remainder 16 SSRs

were provided by Dr Martin W Ganal (IPK and Trait

Genetics GmbH Gatersleben Germany) and all

compared fairly well in terms of map position and order

with the lsquoK 9 Srsquo durum wheat map (Jurman et al

unpublished data)

The comparison of the relative genetic distances

between markers in the lsquoC 9 Lrsquo map and the hexaploid

wheat maps evidenced a limited correspondence for

both DArT and SSR markers For example the genetic

interval comprised between the anchor markers

wPt7475 and wPt9075 (chr 6A) and including ten

anchor wPt-markers covered a genetic distance of

207 cM in the hexaploid wheat map of Crossa et al

(2007) as compared to the ca 25 cM in the lsquoC 9 Lrsquo

durum population

Diversity analysis

The panel of 56 durum accessions initially used to

generate the DArT durum clones was profiled with the

durum DArT array used to profile the RIL population

As expected the polymorphic markers that clearly

distinguished two allelic phases (presence and absence

of hybridization to the genomic clones) were more

numerous than those identified in the lsquoC 9 Lrsquo popu-

lation in fact a total of 1315 polymorphic DArT

markers were found among the materials analysed

The hierarchical subdivision (Fig 2a) of the germ-

plasm analysed was in keeping with the pedigree

information detailed in Table 1 The genetic tree

discriminated the accessions adapted to the Mediter-

ranean areas (ie the majority of the accessions in the

upper part of the tree from Meridiano to Zeina) from

those originated from the North American gene pool

which included cvs adapted to northern latitudes bred

in the Great Plains of the USA and Canada and

subsequently in France and in Australia (lower part of

the tree from Lloyd to Wollaroi) This finding was

confirmed by the principal coordinate analysis

(Fig 2b) in fact the first principal coordinate clearly

separated the American accessions on the left side of

the diagram from the Mediterranean accessions

clustered on the right Within the Mediterranean

accessions DArT markers were able to distinguish

subgroups with different origins In the upper part of

Fig 1 Genetic map for the Colosseo 9 Lloyd RIL popula-

tion Map distances (cM) and marker name are shown on the

left and right side of each chromosome respectively SSR

markers are presented in bold font DArT markers in common

between the lsquoC 9 Lrsquo map and the hexaploid maps used as

references are underlined The approximate locations of the

centromers () are deduced from Somers et al (2004) Loci

marked with and exhibit significant distortion from the

expected 11 segregation ratio at P B 001 and P B 0001

respectively Chromosome regions that showed distorted

segregation in favour of Colosseo or Lloyd are indicated with

shaded bars (solid and hatched filled respectively)

b


123

Fig 1 continued


123

the tree (Fig 2a) a relatively homogeneous cluster of

accessions (from Meridiano to Plata 16) included

recent cvs derived from the successful germplasm Jo

AaFg and RuffFgMexicaliShearwater released at

CIMMYT in the lsquo80 s such germplasm is represented

in the dendrogram by the Mexican founder Altar 84

the successful Italian cvs Duilio and Svevo as well as

the cv Lahn obtained at ICARDA All these cvs have

been largely used in modern durum breeding programs

for their high yield potential and yield stability (Giunta

et al 2007) This germplasm can be easily identified

also based on the second principal coordinate

(Fig 2b) cvs related to Altar 84 Duilio Svevo and

Lahn were grouped in the upper part of the principal

coordinate plot Another subgroup mainly included

cvs and advanced materials obtained at ICARDA and

mostly adapted to dryland areas (Fig 2a from Sebah to

Messapia in the centre of the tree) Finally a well-

distinct group of accessions directly related to the

native germplasm from North Africa and west Asia

(from Trinakria to Zeina) was identified

Thirty-one accessions out of the 56 initially con-

sidered were used to compare the information provided

by SSR and DArT markers The Mantel statistic Z was

equal to 1465 and the coefficient of correlation

between the two genetic distance matrices was quite

sizeable (r = 068) Out of 10000 permutations all

showed random Z values observed Z value thus the

one-tail probability P [random Z C observed Z] was

equal to 00002

The good agreement between the two marker

systems was also evident considering the concor-

dance between the hierarchical subdivision generated

by means of the two methods (Fig 3) However it

can be noticed that the hierarchical classification of

relationships obtained with the DArT markers is to be

considered more robust as compared to the analogous

one that was obtained with the SSRs In fact in the

B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)

COLOSSEO (13)CRESO (14)

DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)

JENNAH KHETIFA-TAMGURT (21)

KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)

SENATORE CAPPELLI (50)

SIMETO (51)

SVEVO (52)

TAMAROI (54)TAMAROI (53)

TRINAKRIA (55)

KOFA (56)

VALFORTE (57)

WOOLAROI (59)WOOLAROI (58)

ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A

DArT Jaccard coefficient

-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60

Mediterranean (CIMMYT)

Mediterranean (native)Australian

Mediterranean x North AmericanNorth American

Mediterranean (ICARDA)Mediterranean (other)

Fig 2 Pattern of genetic diversity for a group of 56 accessions

selected to represent the diversity of durum wheat as revealed

by 1315 DArT markers (a) Unweighted neighbour-joining

tree derived from the Jaccard dissimilarity matrix Numbers at

branching points indicate percent bootstrap support of individ-

ual nodes only values [50 are reported (resampling

no = 1000) The two parents (Colosseo and Lloyd) of the

mapping population used for genetic mapping are highlighted

in red Four pairs of technical replicates are highlighted by

coloured genotype namesnumbers (b) The first two factorial

coordinates of a Jaccard dissimilarity matrix (total inertia of

axes 1 and 2 were 159 and 128 respectively) Accessions

are indicated with the corresponding code number (see

Table 1)


123

DArT-derived cluster the number of grouping nodes

with a reliable and high bootstrap support value

(higher than 50) was higher than that observed for

the SSR-derived cluster ie 16 nodes compared to

only four nodes respectively

Discussion


Genome coverage and marker distribution

The lsquoC 9 Lrsquo integrated DArT-SSR linkage map

obtained in the present study has a total length of

2022 cM which corresponds to ca 70 coverage of

the A and B genomes of the bread wheat consensus

map of Somers et al (2004) This percentage was

calculated taking into account only the anchor SSRs

in common between these two maps considering

the presence of additional DArT and SSR loci in

the lsquoC 9 Lrsquo map we estimate a tetraploid genome

(AABB) coverage of ca 77 Although we obtained a

good coverage of the genome gaps of over 50 cM still

remain on chrs 2A and 2B (pericentromeric regions)

3AS and 7AL the presence of large gaps andor chr

regions with low marker density has been described in

several wheat maps (Sourdille et al 2003 Somers

et al 2004 Torada et al 2006) The lsquoC 9 Lrsquo map also

includes several chr regions with inter-marker dis-

tances higher than 20 cM and two regions on chrs 4BS

and 5AL were poorly represented Moreover the short

arm and the peri-centromeric region of chr 4A were

not covered at all which is consistent with other

published bread wheat maps (Paillard et al 2003

Torada et al 2006) In addition Akbari et al (2006)

and Semagn et al (2006) did not report DArT markers

mapping on chr 4AS Gaps and insufficient coverage

of specific lsquoC 9 Lrsquo chr regions could be due to (i)

structural deficiency of polymorphic markers in highly

recombinogenic regions andor limited sequence var-

iation as shown in other maps (Somers et al 2004

Song et al 2005) andor (ii) extended identity by

descent between the parents of the mapping

population

The low density of DArT markers in group 5 was

already reported in hexaploid wheat particularly in

chr 5A In fact Akbari et al (2006) and Semagn et al

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62

SSR (103 markers)DArT (1315 markers)

tneiciffeoc gnihctam-elpmiStneiciffeocdraccaJ

Fig 3 Comparison of neighbour-joining trees obtained with DArT and SSR markers The numbers at branching points indicate

percent bootstrap support of individual nodes only values [50 are reported (resampling no = 1000)


123

(2006) mapped only three DArT markers in chr 5A

over a total of several hundred successfully mapped

DArT markers The under-representation of polymor-

phic fragments from chr group 5 and particularly chr

5A in wheat genomic representations obtained by

using methylation-sensitive restriction enzymes such

as PstI and Sse8387I is confirmed by unpublished

results obtained from AFLP mapping (AP Sorensen

personal communication) It is known that the genomic

representations obtained with PstI reflect the methyl-

ation status of the genomic DNA and produce markers

preferentially mapping in the hypomethylated gene-

rich regions (van Os et al 2006) However hetero-

chromatin content does not seem to cause this under-

representation In fact even if the heterochromatin

content of chr 5B is one of the highest among wheat

chromosomes this does not hold true for chr 5A and it

has been ascertained that gene-rich regions are present

in both chromosomes (Linkiewicz et al 2004)

In the present study the SSR markers were fairly

evenly distributed along the chromosomes due to the

fact that their location was mostly known and the

SSRs were appropriately selected to avoid closely

linked multiple loci In spite of our efforts to evenly

space the SSR loci we identified a few clusters

specifically around the centromere of few chromo-

somes A similar finding has been reported in most

bread and durum wheat mapping studies and has been

attributed to a reduction of recombination in the

proximal regions of chr arms Clustering of DArT

markers was more frequent compared to SSRs This is

not surprising keeping in mind that there was no pre-

selection of DArT markers and that DArT markers

were over three times more abundant than SSRs The

occurrence of DArT clusters near to distal-telomeric

regions of chr arms was observed in other DArT

mapping studies on wheat (Akbari et al 2006

Semagn et al 2006) and barley (Wenzel et al

2004) High-density physical maps of wheat have

revealed that 90 of the genes are confined to gene-

rich regions that represent ca 10 of the genome

interspersed by large blocks of repetitive DNA and

for the most located on distal chromosome portions

these gene-rich regions are characterised by a higher

recombination rate with respect to the proximal

regions (Gill et al 1996a b Faris et al 2000 Sandhu

et al 2001) The clusters of DArT markers herein

discussed matched the gene-rich regions reported in

the wheat gene distribution model proposed by Gill

et al (1996a b) and Sandhu et al (2001) The higher

density of clusters on distal regions could also be

related to the trend of PstI-based markers towards

hypomethylated non-centromeric regions of the

genome (Langridge and Chalmers 1998) Neverthe-

less it is worth noting that the high number of DArT

clusters may also be a consequence of the presence of

redundant clones on the genomic representation

(Semagn et al 2006) As to the distribution of DArT

markers on genomes A and B the higher number of

DArTs mapping on the B genome was also reported in

hexaploid wheat by Semagn et al (2006)

Finally the average number of crossover events per

RIL observed in the lsquoC 9 Lrsquo mapping population is in

line with what has been reported for wheat RIL

populations In the hexaploid wheat ITMI map a

range of 25ndash55 scorable recombinations was observed

across 115 inbred lines with the most frequent

number of recombinations per line equal to 40 (ie

19 recombinations per chromosome Esch et al

2007) Moreover the recombination density per

chromosome found in the lsquoC 9 Lrsquo population is in

line with that expected based on Poissonrsquos models

(Williams et al 2001)


In the lsquoC 9 Lrsquo population we found 265 of

markers with a significant (P 001) segregation

distortion This value is not much different from those

found in previous mapping studies on bread wheat

(Cadalen et al 1997 Paillard et al 2003 Semagn

et al 2006 Singh et al 2007) and durum wheat

(Blanco et al 1998 Nachit et al 2001) Analogously

to what was observed by the above-cited authors

skewed markers were clustered in specific regions on

several chromosomes Various causes can lead to

segregation distortion chromosomal rearrangement

(Faure et al 1993) alleles inducing gametic or

zygotic selection (Xu et al 1997 Lu et al 2002)

parental reproductive differences (Foolad et al 1995)

and the presence of lethal genes (Blanco et al 1998)

are possible sources of deviation In the case of the

lsquoC 9 Lrsquo population the use of RILs excludes the

possibility to attribute the deviation from the expected

segregation ratio to gametophytic selection as

reported for double-haploid progenies (Cadalen et al

1997) However due to the different genetic back-

ground of Colosseo and Lloyd the occurrence of


123

epistatic interactions negatively affecting the fitness

of the progeny should not be excluded

Map comparison

Based on the chromosome position of the anchor

wPt-DArT markers the degree of conservation of

DArT marker order with the hexaploid wheat maps

was high Instead even if the SSR order in the

lsquoC 9 Lrsquo map was generally in accordance with the

reference maps a few differences were observed and

described (see Section lsquolsquoResultsrsquorsquo) These differences

seem acceptable considering that genetic maps pro-

vide only an indication of the relative marker

positions and genetic distances Moreover inconsis-

tency in map position could be explained by the

presence of additional loci in the wheat genome Our

results showed that the co-linearity between DArT

and SSR markers between durum and hexaploid

wheat is conserved notwithstanding a lack of corre-

spondence among the relative genetic distances

Diversity analysis

DArT marker profiling effectively described the

genetic relationships among the accessions in fact

the neighbour-joining tree and the principal coordi-

nate plot clearly distinguished the main gene pools

the accessions came from Origin pedigree records

and genetic relationships among the majority of the

accessions deployed for this study can be found in

previous studies published by Maccaferri et al (2005

2007) and by Mantovani et al (2006)

Based on the SSR data available for 31 out of the

56 durum accessions it was possible to carry out a

comparison of the informativeness and reliability of

the DArT assay versus selected SSR loci characterised

by multi-allelic status (Maccaferri et al 2003 2005)

The results obtained with the DArT markers are in

good agreement with those obtained with highly

informative genomic SSR loci which up to now have

represented the markers of choice to investigate

genetic relationships and to carry out association

mapping studies in wheat (Breseghello and Sorrells

2006 Balfourier et al 2007 Sanguineti et al 2007)

The set of 1315 bi-allelic and polymorphic DArT

markers that was obtained from the hybridization

assay of each accession to the DArT array allowed to

obtain a hierarchical classification of the accessions

(based on relationships) even more precise than that

obtained with a medium number (103) of highly

informative SSR loci This was not a surprising result

and it can be explained based on the following

considerations The number of polymorphic markers

that is now possible to score with the DArT hybrid-

ization assays on wheat germplasm collections is

medium to high obtaining a similar number of

informative data points using the conventional SSR

and AFLP techniques requires a considerably longer

time and higher monetary investment The number of

bi-allelic markers obtained using DArT assay which

is similar to AFLPs obtained with Sse8387-PstIMseI

restriction enzymes should allow the user to obtain

estimates of genetic relationships with a mean coef-

ficient of variation (CV) equal to or lower than 10

Because of the non-linear exponentially decreasing

relationships between the sampling variance of

genetic diversity estimates and the marker sample

size the 10 CV threshold is considered as a good

satisfactory threshold in terms of cost-effectiveness of

markers for evaluation of genetic distances (Tivang

et al 1994)

Using Sse8387MseI derived-AFLP markers to

estimate genetic relationships in durum wheat it was

demonstrated that the 10 threshold in CV sampling

variance could be reached with marker sets including

at least 200 biallelic loci (Maccaferri et al 2007) a

number of markers that is largely exceeded by the

DArT assay SSR markers due to their allelic

hypervariability are very useful for germplasm

characterization and genetic relationships estimates

The use of a limited number of multi-allelic SSRs

provides information on the haplotype genetic pro-

files of the accessions that could be obtained only

with a correspondingly much higher number of bi-

allelic dominant markers (Weir et al 2006) how-

ever this SSR-specific feature when utilized to

generate global genetic diversity estimates implies

that a relatively high number of SSRs have to be used

in order to obtain genetic diversity estimates with a

limited sampling variance In durum wheat Maccaf-

erri et al (2007) estimated that ca 150 genomic SSR

markers on average were needed to obtain genetic

diversity estimates with acceptably low CV values

Therefore DArT markers can be conveniently used

for investigating genetic diversity in durum wheat


123

DArT effectiveness for deployment in QTL

mapping and MAS

To address the cost-effectiveness issues involved with

the DArT technique it can be underlined that the cost

per DArT marker is low due to the highly parallel

nature of genotyping several thousand markers in a

single assay with the cost per marker assay in

commercial service offered by Triticarte PL at around

US$ 002 (or approximately US$ 50 per genotype) The

cost of SSR genotyping (based on a standard 96 well-

PCR assay fluorescent fragment detection and capil-

lary electrophoresis) commonly ranges from a

minimum of one to several US$ per single lane-

electrophoresis run with a multiplex capability of

three markers per run this cost always exceeds that of

DArT per single data points One advantage of SSR

markers is that they can be preselected for polymor-

phism and for an even genome coverage When SNP

marker panels will be available for wheat on high

throughput platforms (eg on Illumina Golden Gate

system) the cost advantage of DArT over alternative

technologies will be reduced However at this time the

Illumina service (httpicomilluminacomproducts

prod_snpilmn) for the few plant species for which

such panels have been developed is still approximately

three times more expensive compared to the similar

marker density DArT service

In order to be broadly applicable DArT markers

have to be effectively transferable between different

mapping populations This requirement has been

clearly satisfied in case of barley where a high-density

integrated map has been developed based on a number

of independent populations sharing a number of

common markers (Wenzl et al 2006) In wheat the

process of integrated map construction was initially

inhibited by lower marker density compared to barley

(due to distribution of similar number of markers

among three homeologous genomes) but the transfer-

ability of markers between mapping populations is

apparent from the available bread wheat DArT map-

ping data (httpwwwtriticartecomaucontentfur

ther_developmenthtml) and from this report With

approximately 200 genetic maps of bread and durum

wheat profiled with the common set of DArT markers

(A Kilian unpublished) the technology becomes

increasingly a reference for other marker types in these

two crops especially because the map position of

DArT markers in durum is in agreement with that

reported in bread wheat

A critical aspect of any genotyping technology is

the ease of access to markers and ability to reproduce

the results to verify data quality DArT markers

reported in this paper can be accessed through

inexpensive available Triticarte service (httpwww

triticartecomau) which processed over 30000

wheat accessions using a similar marker set in the last

2 years For selected set of markers (usually those

linked to traits of interest) any user of Triticarte

service can obtain marker sequences for development

of monoplex assays or data verification When the

discovery process and sequencing of wheat DArT

markers is completed the sequences of all markers

will be reported in scientific publications and at that

stage released to public databases

Conclusions

This study contributed to the development of diver-

sity arrays technology in wheat by creating new

durum-dedicated libraries of clones and arrays in

addition to the existing ones in hexaploid wheat Up

to now we have selected 2304 polymorphic durum

DArT markers that can be typed in a single assay

through a cost-effective technology DArT profiling

proved to be useful to construct a linkage map and to

elucidate the pattern of relatedness among a wide

range of modern wheat accessions from the most

important durum breeding pools Though SSR and

DArT marker systems are characterized by different

information content on a per locus basis it can be

underlined that wheat being a self-pollinating cereal

the use of biallelic dominant markers such as DArT

markers to characterize the genetic stocks usually

deployed in genetic analyses (recombinant inbred

lines and germplasm collections assembled from

inbred materials) does not imply losses of genetic

information The high number of available DArT

markers their cost-effectiveness and relatively high

polymorphism content are ideal characteristics for

both extensive genome-wide screening for QTL

discovery and for fine mapping and positional cloning

of genes and QTLs Additionally the map position of


reported in bread wheat a feature that will facilitate


123

the comparative analysis of results obtained with

these two key crops

Acknowledgments Major financial support for this project

was provided by Australian Grains RampD Corporation (GRDC)

Regione Emilia Romagna (Italy) progetto PRITT Misura 34-A

CEREALAB and the European Union BIOEXPLOIT Integrated

Project contract no 513959 We would like to acknowledge

technical help from a number of colleagues from Diversity

Arrays Technology Pty LtdTriticarte Pty Ltd (Grzegorz

Uszynski Jason Carling Vanessa Caig Ling Xia Damian

Jaccoud Kasia Heller-Uszynska Gosia Aschenbrenner-Kilian)

and from DiSTA University of Bologna (Sandra Stefanelli)

References

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Bassam BJ Anolles GC Gresshoff P (1991) Fast and sensitive

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Breseghello F Sorrells ME (2006) Association mapping of

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genetics105044586

Cadalen T Boeuf C Bernard S Bernard M (1997) An interva-

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Eujayl I Sorrells ME Baum M Wolters P Powell W (2002)

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Faris JD Haen KM Gill BS (2000) Saturation mapping of a

gene-rich recombination hot spot region in wheat

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Faure S Noyer JL Horry JP Bakry F Lanaud C Gonzalez de

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Foolad MR Arulsekar S Becerra V Bliss FA (1995) A genetic

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Gill KS Gill BS Endo TR Boyko EV (1996a) Identification of

high-density mapping of gene-rich regions in chromo-

some group 5 of wheat Genetics 1431001ndash1012

Gill KS Gill BS Endo TR Taylor T (1996b) Identification and



Giunta F Motzo R Pruneddu G (2007) Trends since 1900 in

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Eur J Agron 2712ndash24 doi101016jeja200701009

Goyal A Bandopadhyay R Sourdille P Endo TR Balyan HS

Gupta PK (2005) Physical molecular maps of wheat

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Gupta PK Balyan HS Edwards KJ Isaac P Korzun V Roder

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Leroy P (2002) Genetic mapping of 66 new microsatellite

(SSR) loci in bread wheat Theor Appl Genet 105413ndash

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Guyomarcrsquoh H Sourdille P Edwards KJ Bernard M (2002)

Studies of the transferability of microsatellites derived

from Triticum tauschii to hexaploid wheat and to diploid

related species using amplification hybridization and

sequence comparisons Theor Appl Genet 105736ndash744

Hayden MJ Nguyen TM Waterman A McMichael GL

Chalmers KJ (2008) Application of multiplex-ready PCR

for fluorescence-based SSR genotyping in barley and

wheat Mol Breed doi101007s11032-007-9127-5

Jaccoud D Peng K Feinstein D Kilian A (2001) Diversity

arrays a solid state technology for sequence information

independent genotyping Nucleic Acids Res 29E25 doi

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Kilian A Huttner E Wenzl P Jaccoud D Carling J Caig V

et al (2005) The fast and the cheap SNP and DArT-based

whole genome profiling for crop improvement In

Tuberosa R Phillips RL Gale M (eds) Proceedings of the

international congress in the wake of the double helix

from the green revolution to the gene revolution Avenue

Media Bologna Italy 27ndash31 May 2003 pp 443ndash461

Koebner RM Summers RW (2003) 21st century wheat

breeding plot selection or plate detection Trends Bio-

technol 2159ndash63 doi101016S0167-7799(02)00036-7

Korzun V Roder MS Wendekake K Pasqualone A Lotti C

Ganal MW et al (1999) Integration of dinucleotide

microsatellites from hexaploid bread wheat into a genetic

linkage map of durum wheat Theor Appl Genet 981202ndash

1207 doi101007s001220051185

Langridge P (2005) Molecular breeding of wheat and barley

In Tuberosa R Phillips RL Gale M (eds) Proceedings of

the international congress in the wake of the double helix




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Langridge P Chalmers K (1998) Techniques for marker

development In Proceedings of the 9th international

wheat genet symposium vol 1 Saskatchewan Canada pp

107ndash117

Lincoln SE Lander ES (1992) Systematic detection of errors in

genetic linkage data Genomics 14604ndash610 doi101016

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Linkiewicz AM Qi LL Gill BS Ratnasiri A Echalier B Chao

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ogous group 5 provides insights on gene distribution and

colinearity with rice Genetics 168665ndash676 doi101534

genetics104034835

Lu H Romero-Severson J Bernardo R (2002) Chromosomal

regions associated with segregation distortion in maize

Theor Appl Genet 105622ndash628 doi101007s00122-002-

0970-9

Maccaferri M Sanguineti MC Donini P Tuberosa R (2003)

Microsatellite analysis reveals a progressive widening of

the genetic basis in the elite durum wheat germplasm Theor

Appl Genet 107783ndash797 doi101007s00122-003-1319-8

Maccaferri M Sanguineti MC Noli E Tuberosa R (2005)

Population structure and long-range linkage disequilib-

rium in a durum wheat elite collection Mol Breed

15271ndash290 doi101007s11032-004-7012-z

Maccaferri M Sanguineti MC Natoli V Ortega JAL Salem

MB Bort J et al (2006) A panel of elite accessions of

durum wheat (Triticum durum Desf) suitable for associ-

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Maccaferri M Stefanelli S Rotondo F Tuberosa R Sanguineti

MC (2007) Relationships among durum wheat accessions

I Comparative analysis of SSR AFLP and phenotypic

data Genome 50373ndash384 doi101139G06-151

Maccaferri M Sanguineti MC Corneti S Jose LAO Ben

Salern M Bort J et al (2008) Quantitative trait loci for

grain yield and adaptation of durum wheat (Triticumdurum Desf) across a wide range of water availability

Genetics 178489ndash511 doi101534genetics107077297

Mantel NA (1967) The detection of disease clustering and a

generalized regression approach Cancer Res 27209ndash220

Mantovani P van der Linden G Maccaferri M Sanguineti MC

Tuberosa R (2006) Nucleotide-binding site (NBS) profil-

ing of genetic diversity in durum wheat Genome

491473ndash1480 doi101139G06-100

Nachit MM Elouafi I Pagnotta MA El Saleh A Iacono E

Labhilili M et al (2001) Molecular linkage map for an

intraspecific recombinant inbred population of durum

wheat (Triticum turgidum L var durum) Theor Appl

Genet 102177ndash186 doi101007s001220051633

Paillard S Schnurbusch T Winzeler M Messmer M Sourdille

P Abderhalden O Keller B Schachermayr G (2003) An

integrative genetic linkage map of winter wheat (Triticumaestivum L) Theor Appl Genet 1071235ndash1242

Peng J Korol AB Fahima T Roder MS Ronin YI Li YC et al

(2000) Molecular genetic maps in wild emmer wheat

Triticum dicoccoides genome-wide coverage massive

negative interference and putative quasi-linkage Genome

Res 101509ndash1531 doi101101gr150300

Perrier X Flori A Bonnot F (2003) Data analysis methods In

Hamon P Seguin M Perrier X Glaszmann JC (eds)

Genetic diversity of cultivated tropical plants Enfield

Science Publishers Montpellier pp 43ndash76

Perrier X Jacquemoud-Collet JP (2006) DARwin software

(httpdarwin cirad frdarwin)

Plaschke J Ganal MW Roder MS (1995) Detection of genetic

diversity in closely related bread wheat using microsat-

ellite markers Theor Appl Genet 921078ndash1084

Roder MS Korzun V Wendehake K Plaschke J Tixier MH

Leroy P Ganal MW (1998) A microsatellite map of

wheat Genetics 1492007ndash2023

Saghai-Maroof MA Soliman KM Jorgensen RA Allard RW

(1984) Ribosomal DNA sepacer-length polymorphism in

barley Mendelian inheritance chromosomal location and

population dynamics Proc Natl Acad Sci USA 818014ndash

8019 doi101073pnas81248014

Sandhu D Champoux JA Bondareva SN Gill KS (2001)

Identification and physical localization of useful genes

and markers to major gee-rich region on wheat group 1S

chromosomes Genetics 1571735ndash1747

Sanguineti MC Li S Maccaferri M Corneti S Rotondo F Chiari

T et al (2007) Genetic dissection of seminal root architec-

ture in elite durum wheat germplasm Ann Appl Biol

151291ndash305 doi101111j1744-7348200700198x

Semagn K Bjornstad A Skinnes H Maroy AG Tarkegne Y

William M (2006) Distribution of DArT AFLP and SSRmarkers in a genetic linkage map of a doubled-haploid

hexaploid wheat population Genome 49545ndash555 doi

101139G06-002

Singh K Ghai M Garg M Chhuneja P Kaur P Schnurbusch

T Keller B Dhaliwal HS (2007) An integrated molecular

linkage map of diploid wheat based on a Triticum bo-eoticum x T monococcum RIL population Theor Appl


Somers DJ Kirkpatrick R Moniwa M Walsh A (2003) Mining

single-nucleotide polymorphisms from hexaploid wheat

ESTs Genome 46431ndash437 doi101139g03-027

Somers DJ Isaac P Edwards K (2004) A high-density

microsatellite consensus map for bread wheat (Triticumaestivum L) Theor Appl Genet 1091105ndash1114 doi

101007s00122-004-1740-7

Song QJ Fickus EW Cregan PB (2002) Characterization of

trinucleotide SSR motifs in wheat Theor Appl Genet

104286ndash293

Song QJ Shi JR Singh S Fickus EW Costa JM Lewis J et al

(2005) Development and mapping of microsatellite (SSR)

markers in wheat Theor Appl Genet 110550ndash560 doi

101007s00122-004-1871-x

Sourdille P Cadalen T Guyomarcrsquoh H Snape JW Perretant

MR Charmet G Boeuf C Bernard S Bernard M (2003)

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molecular marker linkage map for the QTL detection of

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538

Sourdille P Singh S Cadalen T Brown-Guedira G Gay G Qi

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the establishment of genetic-physical map relationships in

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412ndash25 doi101007s10142-004-0106-1

Stam P (1993) Construction of integrated genetic linkage maps

by means of a new computer package JoinMap Plant J

3739ndash744

Tivang JG Nienhuis J Smith OS (1994) Estimation of sampling

variance of molecular marker data using the bootstrap


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procedure Theor Appl Genet 89259ndash264 doi101007

BF00225151

Torada A Koike M Mochida K Ogihara Y (2006) SSR-based

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van Ooijen JW (2006) JoinMap 4 software for the calculation

of genetic linkage maps in experimental populations

Kyazma BV Wageningen Netherlands

van Os H Stam P Visser RGF van Eck HJ (2005) RECORD

a novel method for ordering loci on a genetic linkage map


0097-x

van Os H Andrzejewski S Bakker E Barrena I Bryan GJ

Caromel B Ghareeb B Isidore E de Jong W van Koert

P Lefebvre V Milbourne D Ritter E Rouppe van der

Voort JNAM Rousselle-Bourgeois F van Vliet J Waugh

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of a 10 000-marker ultradense genetic recombination map

of potato providing a framework for accelerated gene

isolation and a genomewide physical map Genetics


Varshney RK Tuberosa R (2007) Genomics-assisted crop

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genomics approaches and platforms Springer Dordrecht

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Weir BS Anderson AD Hepler AB (2006) Genetic relatedness

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7771ndash780 doi101038nrg1960

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hofs A et al (2004) Diversity arrays technology (DArT)

for whole-genome profiling of barley Proc Natl Acad Sci

USA 1019915ndash9920 doi101073pnas0401076101

Wenzl P Li H Carling J Zhou M Raman H Paul E et al

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DArT markers to SSR RFLP and STS loci and agricul-

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2164-7-206

Williams RW Gu J Qi S Lu L (2001) The genetic structure of

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1-004618

Xu Y Zhu L Xiao J Huang N McCouch SR (1997) Chromo-

somal regions associated with segregation distortion of

molecular markers in F2 backcross doubled haploid and

recombinant inbred populations in rice (Oryza sativa L)

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(2004) Development and mapping of EST-derived simple

sequence repeat markers for hexaploid wheat Genome

47805ndash818 doi101139g04-057


123

importance is related to the fact that durum grain is

mainly used for human consumption However

genetics of important agronomic and quality traits of

durum wheat has been poorly investigated with respect

to other cereals Detailed genetic information on

durum adaptation and grain quality is required as a

basis in breeding programs Molecular markers are

efficient tools to speed up crop improvement (Lang-

ridge 2005 Varshney and Tuberosa 2007) and for the

construction of molecular linkage maps the first step in

the genetic dissection of target traits

Microsatellites (simple sequence repeats SSRs) are

PCR-based markers characterised by a high level of

polymorphism that permits to discriminate among

cultivars and even among closely related wheat

breeding lines (Plaschke et al 1995 Maccaferri et al

2007) In addition to their high polymorphism SSRs

are usually single locus sites an important feature

when dealing with allopolyploid species SSRs have

rapidly become the markers of choice for the con-

struction of genetic maps in wheat (Roder et al 1998

Somers et al 2004 Sourdille et al 2004) up to now

several hundred SSR primer pairs have been developed

for all three genomes of wheat (httpwheatpwusda

govGG2indexshtml) While SSRs are standard PCR-

based markers and can be considered as proven anchor-

markers their suitability for high-throughput mapping

does not favourably compare to the new single

nucleotide polymorphism (SNP)-based genotyping

techniques (Kilian et al 2005) Moreover SSR-mul-

tiplexing requires an extensive additional optimisation

(Hayden et al 2008) Development of massive SNP

resources from the expressed portion of the wheat

genome amenable to fully automated genotyping is a

difficult task for wheat due to its low frequency of

sequence polymorphism and its allopolyploid nature

(Koebner and Summers 2003 Somers et al 2003)

Diversity arrays technology (DArT) for which proof

of concept was first reported by Jaccoud et al (2001)

is becoming increasingly adopted in many species

(most current list of species with DArT technology

developed can be found at wwwdiversityarrayscom)

The technology combines a complexity reduction

method (Wenzl et al 2004) with hybridization-based

polymorphism detection using high-throughput solid-

state platforms and has the potential to generate hun-

dreds of high-quality genomic dominant markers with

a cost- and time-competitive trade-off (Kilian et al

2005)

In contrast to hexaploid bread wheat (Triticum

aestivum L AABBDD genome) for which several

linkage maps have been developed mapping in durum

wheat has received relatively little attention The first

linkage map of durum wheat based on 65 recombinant

inbred lines (RILs) and RFLP markers was reported by

Blanco et al (1998) SSRs from hexaploid wheat

(Roder et al 1998) were subsequently integrated into

this linkage map (Korzun et al 1999) More recently

intra- and inter-specific linkage maps based on RFLPs

SSRs and AFLPs have been developed (Peng et al

2000 Nachit et al 2001 Maccaferri et al 2008)

These studies have shown a generally good conserva-

tion of marker co-linearity as compared to the A and B

genomes of hexaploid wheat

Dedicated DArT genotyping platforms have been

produced for bread wheat (Akbari et al 2006 Semagn

et al 2006) The identification of polymorphic DArT

markers directly obtained from durum wheat by

generating genomic representations from diverse

durum accessions followed by mapping the polymor-

phisms on a durum mapping population and by

genotyping a panel of durum accessions serves as a

basis for broadening the use of DArT markers for

durum wheat This study presents the mapping in

durum wheat of 392 new DArT markers that were

anchored to the wheat map using 162 SSRs DArT

markers were then used to profile a panel of durum

accessions subsequently the genetic relationships

based on DArT markers were compared to those

obtained with highly polymorphic SSRs

Materials and methods

Plant material

A panel of 56 diverse durum wheat accessions was

selected to sample the genetic diversity present in the

cultivated durum gene pools from the most important

durum wheat production areas Additionally the

accessions were chosen in order to explore various

levels of genetic relationships based on previous

results obtained with SSRs (Maccaferri et al 2005)

The panel was assembled with durum cultivars (cvs)

and accessions from (i) Southern Europe (Italy

Spain and France) (ii) the International Maize and

Wheat Improvement Center (CIMMYT) (iii) the

International Center for Agricultural Research in the


123






















et al 2006)

Molecular analysis

DNA extraction













































(2004)
















centrifugation






123



Country Year


















145873Cappelli2Yuma

5









Maier 33 US-ND 1998 D8193D8335 8



5Carleton

8















123





















Table 1 continued


Country Year

















a Seed sources
















123








































947 1009 1034 1038 1045 1084 1184 1198 1246




















51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)











Table 2 SSR markers



Lloyd


Barc 130 Song et al (2002 2005)









123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123






















et al 2006)

Molecular analysis

DNA extraction













































(2004)
















centrifugation






123



Country Year


















145873Cappelli2Yuma

5









Maier 33 US-ND 1998 D8193D8335 8



5Carleton

8















123





















Table 1 continued


Country Year

















a Seed sources
















123








































947 1009 1034 1038 1045 1084 1184 1198 1246




















51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)











Table 2 SSR markers



Lloyd


Barc 130 Song et al (2002 2005)









123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123



Country Year


















145873Cappelli2Yuma

5









Maier 33 US-ND 1998 D8193D8335 8



5Carleton

8















123





















Table 1 continued


Country Year

















a Seed sources
















123








































947 1009 1034 1038 1045 1084 1184 1198 1246




















51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)











Table 2 SSR markers



Lloyd


Barc 130 Song et al (2002 2005)









123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123





















Table 1 continued


Country Year

















a Seed sources
















123








































947 1009 1034 1038 1045 1084 1184 1198 1246




















51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)











Table 2 SSR markers



Lloyd


Barc 130 Song et al (2002 2005)









123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123








































947 1009 1034 1038 1045 1084 1184 1198 1246




















51C 45 s) 72C (45 s)24 cycles of 94C (45 s)

51C (45 s) 72C (45 s)72C (5 min)











Table 2 SSR markers



Lloyd


Barc 130 Song et al (2002 2005)









123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123














































Diversity analysis

Set of accessions


















j is 0
























123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123

SM frac14 mn















matrices


Z frac14Xn

ifrac141

Xn

jfrac141

AijBij











Results
















DArT-SSR map
















162 SSR loci































123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123
























nine on chr 5B














chromosome






























Map comparison




























123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123














































unpublished data)










durum population

Diversity analysis








































b


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123

Fig 1 continued


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123






























equal to 00002









B

100

ACMORSE (1)

ACPATHFINDER (2)

ALTAR 84 (3)

AGHRASS1 (4)

ASTRODUR

AWL12BIT (6)

AZEGHAR2 (7)

BELIKH2 (8)

BEN (9)

CAPEITI8 (10)

CHAM1 (11)

CLAUDIO (12)


DON PEDRO (15)

DUILIO (16)

GIDARA2 (17)

GRAZIA (18)

HAURANI (19)

IRIDE (20)


KORIFLA (22)

KYLE (23)

LAHN (24)

LANGDON (25)

LEVANTE (26)

LINE139 (28)LINE139 (27)

LINE149 (30)LINE149 (29)

LLOYD (31)

LOUKOS1 (32)

MAIER (33)

MERIDIANO (34)

MESSAPIA (35)

MEXICALI 75 (36)

NEFER (37)

NEODUR (38)

OFANTO (39)

OMRABI 5 (40)

OMRUF2 (41)

ORJAUNE (42)

OUASSEL1 43)

PLATA16 (44)

QUADALETE (45)

RASCON2TARRO (46)

REVA (47)

SARAGOLLA (48)

SEBAH (49)


SIMETO (51)

SVEVO (52)


TRINAKRIA (55)

KOFA (56)

VALFORTE (57)


ZEINA1 (60)

61

100

87

100

96

52

67

100

92

78

100

84

90

75

54

100

63

99

100

100

96

97

89

54

73

65

81

100

65

100

100

62

54

67

99

70

64

68

52

A


-3 -25 -2 -15 -1 -05 05 1 15 2 25 3 35

3

25

2

15

1

05

-05

-1

-15

-2

-25

12

3

4

5

67

8

9

10

11

12

13

14

1516

17

18

19

20

21

22

23

24

2526

27 28

2930

31

32

33

34

35

3637

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

5354

55

56

57

5859

60


















Table 1)


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123






Discussion


































population




0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRO

DUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHNLOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

97

100

100

100

95

99

100

99

100

64

100

96

89

55

51

100

0 01

AGHRASS1

AWL12BIT

AZEGHAR2

CAPEITI8

CHAM1

CLAUDIO

COLOSSEOCRESO

DON PEDRODUILIO

GIDARA2

HAURANI

IRIDE

KORIFLA

LAHN

LOUKOS1

MERIDIANO

MESSAPIA

MEXICALI 75

OFANTO

OMRABI 5

OMRUF2

OUASSEL1

PLATA16

QUADALETE

RASCON2TARRO

REVA

SEBAH

SVEVO

TRINAKRIA

ZEINA1

86

99

58

62






123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123


































































































123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123



Map comparison

















Diversity analysis
















































et al 1994)


























123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123


mapping and MAS































































Conclusions





























123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123


these two key crops











References





0365-4





0517-1







s001220050948



genetics105044586

























87517ndash526 doi101007BF00215098




101007BF00220887













101007s10142-005-0146-1







422













101093nar294e25















1207 doi101007s001220051185







123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123




107ndash117



S0888-7543(05)80158-2





genetics104034835




0970-9








15271ndash290 doi101007s11032-004-7012-z


















491473ndash1480 doi101139G06-100





Genet 102177ndash186 doi101007s001220051633








Res 101509ndash1531 doi101101gr150300

















8019 doi101073pnas81248014








151291ndash305 doi101111j1744-7348200700198x




101139G06-002










101007s00122-004-1740-7



104286ndash293




101007s00122-004-1871-x






538





412ndash25 doi101007s10142-004-0106-1



3739ndash744




123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123


BF00225151




1121042ndash1051 doi101007s00122-006-0206-5







0097-x

















7771ndash780 doi101038nrg1960









2164-7-206




1-004618









47805ndash818 doi101139g04-057


123

Documents

An integrated DArT-SSR linkage map of durum wheat