Genetics and Breeding of Durum Wheat

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complexity of cereals – Gaboun F., Diria G., Adenike F., Abdelwahd R., Ibriz M., Soulaymani A. ........... 323. Screening durum wheat for heat tolerance – Sissons ...
Proceedings of the International Symposium on

Genetics and Breeding of Durum Wheat

Dedicated to the memory of Gian Tommaso Scarascia Mugnozza

Les opinions, les données et les faits exposés dans ce numéro sont sous la responsabilité des auteurs et n'engagent ni le CIHEAM et la FAO, ni les Pays membres. Opinions, data and information presented in this edition are the sole responsibility of the author(s) and neither CIHEAM and FAO nor the Member Countries accept any liability therefor.

CIHEAM Proceedings of the International Symposium on

Genetics and Breeding of Durum Wheat Scientific Editors: E. Porceddu, A.B. Damania, C.O. Qualset Compilation: E. Porceddu Proceedings of the International Symposium on Genetics and Breeding of Durum Wheat, promoted by the Italian Accademia Nazionale delle Scienze, detta dei XL, in partnership with the Italian National Research Council, the International Center for Agricultural Reasearch in the Dry Areas (ICARDA), the Centro International maize y Trigo (Cimmyt), the Italian National Agency for New technologies, Energy and Sustainable Economic Development (ENEA), Food and Agriculture Organisation of the United Nations (FAO), the Mediterranean Agronomic Institute of Bari – CIHEAM. Rome, Italy May 27-30 2013.

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OPTIONS méditerranéennes

Head of Publication: Cosimo Lacirignola 2014

Series A: Mediterranean Seminars

Centre International de Hautes Etudes Agronomiques Méditerranéennes International Centre for Advanced Mediterranean Agronomic Studies

Number 110

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Comment citer cette publication / How to quote this document : E. Porceddu, A.B. Damania, C.O. Qualset (eds.). Proceedings of the International Symposium on Genetics and Breeding of Durum Wheat - IAM Bari: CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes), 2014 – 63 p. (Série A Mediterranean Seminars, N° 110, Options Méditerranéennes)

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Contents Foreword ..................................................................................................................................................... 7 Preface ......................................................................................................................................................... 9 Symposium Committees ........................................................................................................................ 11 Partners, sponsors and supporting institutions .................................................................................. 13

Opening session Welcome addresses – Chiancone E. ........................................................................................................ 17 Opening address – van Ginkel M. ............................................................................................................ 19 Durum wheat in the MediterraneanăLacirignola C. ............................................................................ 21 Gian Tommaso Scarascia Mugnozza: Rome 1925 - 2011 – Porceddu E. ........................................... 23 Symposium remarks – Porceddu E. .......................................................................................................... 27

Session 1. Origin and evolution of durum wheat Durum wheat evolution-- a genomic analysis – Ben-Abu Y., Tzfadia O., Maoz Y., Kachanovsky D.E., Melamed-Bessudo C., Feldman M., Levy A.A ..................................................... 31

Biodiversity of tetraploid wheats: taxonomy, studying, increasing and preservation – Goncharov N. P. ........................................................................................................................................ 47 Global durum wheat diversity: structure and origin revealed by means of the gliadin markers – Kudryavtsev A.M., Melnikova N.V., Yu Novoselskaya-Dragovich A. ............................................................. 57

Session 2. Genetics resources and durum wheat germplasm enhancement Broadening the genetic bases of durum wheat – David J.L., Tavaud M., Roumet P., Muller M.H., Santoni S., Gautier S., Holtz Y., Ranwez V., Ardisson M., Poux G., Vagne C. .............................. 65

Positive effects on yield-contributing traits associated with Thinopyrum ponticum chromosome segments introgressed into durum wheat – Kuzmanovic’ L., Virili M.E., Gennaro A., Bitti A., Ceoloni C. ..................................................................................................................... 79

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Searching for climate change related traits in plant genetic resources collections using Focused Identification of Germplasm Strategy (FIGS) – Bari A., Street K., Amri A., Nachit M.M., Mackay M., Ouabbou H., Kehel Z., Ghanem M.E., De Pauw E., Nazari K., Alo F., El Bouhssini M., Tsivelikas A., Humeid B. ...................................................................................................... 87

Intra-population variation for agronomic characteristics in the durum wheat landrace “SafraMa’an” (Triticum turgidum L. var. durum) – Al-Tabbal J.A., Duwayri M. .................................... 95 Exploiting landrace genetic diversity for germplasm enhancement in durum wheat breeding in Morocco – Taghouti M., Rhrib K., Gaboun F. ...................................................................... 109 Allelic variation for GS and GOGAT genes in a tetraploid wheat collection – Nigro D., Giancaspro A., Giove S.L., Piarulli L., Marcotuli I., Mangini G., Blanco A. .................................................... 121

Evaluation of a hulled wheat (emmer and spelt) collections – Quaranta F., Belocchi A., Camerini M., Cecchini C., Fornara M., Pucciarmati S., D’Egidio M.G. ........................................................... 127

The strategies to serve and conserve Moroccan durum wheat genetic diversity before it is lost – Ramdani A., Ouabbou H., Nsarellah N., Lhaloui S., Abbad-Andaloussi F., Nachit M.M., Bhavani S., Nazari K., Wanyera R., Ferrahi M., Haddoury J., Udupa S.M. ..................................................... 131

Evolution of durum wheat from Sicilian landraces to improved varieties – Sciacca F., Cambrea M., Licciardello S., Pesce A., Romano E., Spina A., Virzì N., Palumbo M. ....................................... 139

Genetic improvement of durum wheat establishment under fluctuating environmental conditions – Ben-David R., Amram A.,, Nashef K., Peleg Z. ............................................ 147 Variability of total antioxidant capacity among durum wheat genotypes –Taddei F., Ciccoritti R., Cacciatori P., D’Egidio M.G. .................................................................................................... 151 Evolution of durum wheat breeding in Italy – Porceddu E., Blanco A. .............................................. 157

Session 3. Strategies and tools in durum wheat genetics and breeding Developing improved durum wheat germplasm by altering the cytoplasmic genomes – Ghavami, F., Bassi F.M., Burciaga R., Soltani A., Noyszewski A., Michalak De Jimenez M.K., Gu Y.Q., Meinhardt S., Elias E.M., Kianian P.M.A., Mergoum M., Maan S.S., Kianian S.F. ............................. 177

The progeny from the [(T. turgidum X Dasypyrum villosum) amphiploid X Triticum aestivum] hybridization is an effective source of new durum wheat inbred lines – De Pace C., Bizzarri M., Vittori D., Vaccino P., Caceres M.E., Ceccarelli M., Raksegi M., Vida G. .................................... 189

Integrated crop solution as new approach to combine genetics and other innovative inputs in wheat varieties development – Bassi R., Andrè C. .............................................................. 201 Detection of molecular markers associated with yield and yield components in durum wheat (Triticum turgidum L. var. durum) under saline conditions – Dura S., Duwayri M., Nachit M.M. .............................................................................................................. 209

In vitro gynogenesis in some varieties of durum wheat (Triticum durum L.) – Mdarhri Alaoui M., Gaboun F., Cherkaoui S. ............................................................................................... 223

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Exploitation of SNP markers located on wheat 5A chromosome for the study of syntenic relationship with model species – Giancaspro A. , Nigro D., Giove S.L., Zacheo S.A., Simeone R., Piarulli L., Colasuonno P., Blanco A. ............................................................................................................ 229

Efficient callus induction, plantlets regeneration and genetic transformation of durum wheat – Iraqi D., Abdelwahd R., Udupa S.M. .......................................................................... 235

Session 4. Genetics and breeding for durum wheat yield and sustainability Durum wheat adaptation and sustainability: ensuring accurate phenotyping for improving drought tolerance and yield stability – Monneveux P. .............................................. 243 Adaptation of durum wheat to a changing environment – Cattivelli L., Miglietta F., Zaldei A., Rizza F., Mastrangelo A.M., De Vita P., Mazzucotelli E. ................................................................ 279 Durum wheat (T. durum Desf.) vs. bread wheat (T. aestivum L. em.Thell.) in south-east Anatolia, Turkey – Ozberk I., Rajaram S., Ilkhan A., Ozberk F. ........................................ 283 Durum wheat breeding for high yield potential in Egypt – El-Areed SH., Nachit M.M., Hagaras A., El-Sherif S.H., Hamouda M. ................................................................................ 291

Molecular responses to drought and heat stress in durum wheat – Aprile A., Marè C. , Havlickova L., Panna R., Rizza F., Mastrangelo A.M., Borrelli G.M., Rampino P., Cattivelli L., De Bellis L., Perrotta C. ......................................................................................... 295

Durum wheat and local chains: A new strategy to strengthen locally selected genotypes – Carboni G., Dettori M., Goddi G., Mulè P., Satta B., Spanu E. ..................................................................... 301

The n-alkylresorcinols in durum wheat: genotypic and environmental variability – Ciccoritti R., Bellato S., Frate V., Nocente F. ................................................................................................. 307

Application of the international crop information system for retrieval and usage of pedigree and phenotypic data for use in durum research and breeding – Clarke F.R., Clarke J.M., Ruan Y., Lin X., N’Diaye A., Kthiri D., Pozniak C.J., Weibe K., Yates S. ........................................ 315

Durum wheat and climate change: simulation models as a tool to support decisions in targeting genotypes and crop breeding – Dettori M., Cesaraccio C., Motroni A., Spano D., Duce P. .................................................................................................................... 319

The salt tolerance candidate genes family in wheat and its relationship to the phylogenetic complexity of cereals – Gaboun F., Diria G., Adenike F., Abdelwahd R., Ibriz M., Soulaymani A. ........... 323 Screening durum wheat for heat tolerance – Sissons M.J., Emebiri L., Pleming D., Taylor H., Eckermannand P., Collins N.C. .................................................................................................... 339

Durum wheat cultivation and breeding in the Altai Russian region – Rozova M.A., Pokornyak V.P. ....................................................................................................................... 345

Proteomic analyses of the effect of nitrogen assimilation in wheat cultivars under different fertilization regimes – Vita F., Lucarotti V., Salzano A., Scaloni A., Alessio M., Alpi A. ............. 351 QTL mapping of morphological traits associated with drought adaptation in a Iranian mapping population of durum wheat – Zarei L., Farshadfar E., Cheghamirza K., Desiderio F., Cattivell L. .......................................................................................................................... 355

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The history of wheat breeding in Algeria – Benbelkacem A. ............................................................. 363 Avenues for increasing salt tolerance of Tunisian durum wheat cultivars – Chaabane R., Saidi A., Rouissi M., Ben Naceur E., Mejri C., Ben Naceur M’b. ............................................. 371

Durum wheat cultivation and use in the USA with special reference to California – Damania A.B. ............................................................................................................................................. 379

Yield and nitrogen use efficiency as influenced by rates of nitrogen fertilizers of some Tunisian durum wheat cultivars – Ayadi S., Karmous C., Trifa Y., Hammami Z., Rezgui S. .......... 391 Durum wheat grain yield and quality under elevated CO2 : first results of a free air carbon dioxide enrichment (FACE) experiment – Marè C., Mazzucotelli E., Reggiani F., Zaldei A., Miglietta F., Fares C., Rizza F., Cattivelli L., Badeck F.W. ............................................................... 401

Session 5. Genetics and breeding for durum wheat disease and pest resistance Genetic resources for stem rust resistance in cultivated and wild tetraploid wheats – Olivera P.D., Yue J. ...................................................................................................................................... 409

Pyramiding of resistance genes Sr36 and Sr2 in durum wheat background (HI 8498) through marker assisted selection for resistance to stem rust race 117- group pathotypes – Sai Prasad S.V., Singh S.K., Kumar V., Kantwa S.L., Dubey V.G., Ambati D., Prakasha T.L., Mishra A.N. ......... 419

Breeding durum wheat for crown rot tolerance – Kadkol G., Simpfendorfer S., Raju T. ................... 431 Diverse sources of resistance to Indian pathotypes of stem rust and leaf rust in durum wheat – Mishra A.N., Sai Prasad S.V., Shirsekar G.S., Yadav S.R., Kaushal K., Dubey V.G. .......................................... 435

Genetic basis of resistance to leaf rust in tetraploid wheats – Desiderio F., Guerra D., Mastrangelo A.M., Rubiales D., Pasquini M., Simeone R., Blanco A., Cattivelli L., Valè G. ............................ 447

Durum wheat improvement against fungal pathogens by using protein inhibitors of cell wall degrading enzymes – D’Ovidio R., Moscetti I., Janni M., Volpi C., Kalunke M.R., Tundo S., Sella L., Favaron F. ................................................................................................. 453

Pyramiding resistance genes to Fusarium head blight and rusts from Thinopyrum ponticum into durum wheat – Forte P., Kuzmanovic’ L., Virili M.E., Gennaro A., Bitti A., Ceoloni C. ........................ 457 Characterization of sources of resistance to leaf rust in durum wheat germplasm – Goyeau H., Berder J., Lacoudre F., Ammar K., Loladze A., Duchalais L., Goudemand E., Desmouceaux N., André C., Blanc P., Gervais L., Lonnet P., Lefèvre T., Argillier O., Robert O., Lezie A., Poupard B., Olivier A. . 463

Qualitative and quantitative resistance against powdery mildew in wheat – Marone D. , Russo M.A., Laidò G., De Vita P., Papa R., Blanco A., Gadaleta A., Mastrangelo A.M. .................................. 469

Additional genetic factors of resistance to stem rust, leaf rust and powdery mildew from Dasypyrum villosum – De Pace C., Bizzarri M., Pasquini M., Nocente F., Ceccarelli M., Vittori D., Vida G. ................................................................................................................. 477

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Session 6. Genetics and breeding for nutritional and technological quality Improvement of technological and nutritional quality in durum wheat: achievements and perspectives – Lafiandra D., Masci S., Palombieri S., Botticella E., Bovina R., Ferrazzano G., Mantovani P., Massi A., Margiotta B., D’Egidio M.G., Sestili F. ............................. 495

Durum wheat production chain: research, quality and future challenges – Atallah M., Ronchi C., Silvestri M., Ruini I. .................................................................................................. 501

Quality in durum wheat: comparison between landraces and high yielding varieties – Daaloul Bouacha O., Nouaigui S., Daaloul A., Rezgui S. ............................................................................. 505

Breeding and quality of soft-textured durum wheat – Gazza L., Sgrulletta D., Cammerata A., Gazzelloni G., Galassi E., Pogna N. ...................................................................................... 511 Molecular characterization of candidate genes involved in nitrogen metabolism and relationship with the grain protein content of wheat – Gadaleta A., Nigro D., Marcotuli I., Giancaspro A., Blanco A. ...................................................................... 517

Mediterranean durum wheat landraces as a source of variability for quality improvement – Royo C., Nazco R., Peña R.J., Ammar K., Villegas D. .................................................................................... 527

Purple grain colour genes in wheat – Khlestkina E., Shoeva O., Börner A., Gordeeva E. ..................... 533 Biochemical and molecular approaches for the technological quality assessment of durum wheat varieties – Babay E., Hanana M., Mzid R., Haj-Salah H., Ghorbel A., Carrillo J.M., Amara H., Rodriguez-Quijano M. ........................................................................................... 541

Grain quality of durum wheat varieties – Abugalieva A.I. , Morgounov A.Y. ...................................... 549 Phenolic compounds and antioxidant activity in tetraploid wheat – Delvecchio L.N., Taranto F., Mangini G., Blanco A., Pasqualone A. ........................................................................................ 555

Characterization of Phytoene synthase 2 (Psy2) genes in wheat – Colasuonno P., Schiavulli A., Sonnante G., Incerti O., Giove S., Giancaspro A., Zacheo S.A., Gadaleta A. ............................ 565 Evaluation of Triticum durum Desf. germplasm for the improvement of local products – Marzario S., Gioia T., Logozzo G., Spagnoletti Zeuli P.L. .............................................................................. 571

Identification of molecular markers associated with yield and quality traits for Argentinean durum wheat breeding programs – Roncallo P., Echenique V. ..................................... 577 Durum wheat breeding lines with new HMW glutenin subunit combinations selected for bread-making quality – Spina A., Ammar K., Peña R.J., Bentivenga G., Sciacca F., Virzì N., Palumbo M. .......................................................................................... 583

Importance of durum wheat breeding in terms of bulgur in Southeastern Anatolian Region of Turkey – Tedkal S. ........................................................................ 589 Breeding for improved technological quality in winter durum wheat – Vida G., Veisz O. ........... 595

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Session 7. Perspectives in structural and functional genomics A new and “open access” chromosome approach to complex genomes: flow sorting of FISH labeled chromosome in suspension – Giorgi D., Farina A.,Grosso V., Lucretti S. .................. 605 Molecular analysis of a novel DNA transposon in Triticeae – Thiyagarajan K., Cantale C., Porceddu E., Galeffi P. ............................................................................................................... 613

E3 ubiquitin ligases regulating plant stress responses: an overview – Guerra D., Mastrangelo A.M., Cattivelli L., Mazzucotelli E. .......................................................................... 619

Closing session Closing Remarks – Porceddu E. ............................................................................................................... 625

List of Participants ................................................................................................................................... 629

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%UHHGLQJDQGTXDOLW\RIVRIWWH[WXUHG durum wheat Laura Gazza, Daniela Sgrulletta, Alessandro Cammerata, Gloria Gazzelloni, Elena Galassi, Norberto Pogna CRA-QCE, Rome, Italy

Abstract. Puroindolines A (PIN-A) and B (PIN-B) encoded by genes Pina-D1 and Pinb-D1 on chromosome '6DUHWKHSULQFLSDOGHWHUPLQDQWIDFWRUVRINHUQHOKDUGQHVVLQFRPPRQZKHDWDQGH[HUWDVWURQJLPSDFWRQ VHYHUDOTXDOLW\WUDLWVVXFKDV\LHOGDQGJUDQXODULW\RIÀRXUVWDUFKGDPDJHDQGZDWHUDEVRUSWLRQUKHRORJLFDO and baking properties. Effects of grain texture on pastamaking and breadmaking quality were evaluated in soft-textured durum wheat lines (SDLs) as compared with their hard durum sister lines (HDLs). SDLs DFFXPXODWHGSXURLQGROLQHVRQWKHLUVWDUFKJUDQXOHVDQGVKRZHG6.&6YDOXHVVLJQL¿FDQWO\ORZHUWKDQWKRVH RI WKHLU KDUGWH[WXUHG FRXQWHUSDUWV ODFNLQJ 3LQ$ DQG 3LQ%7KH DYHUDJH ÀRXU H[WUDFWLRQ UDWH RI 6'/ ZDV DERXWKLJKHUWKDQWKDWRI+'/,QFUHDVLQJNHUQHOVRIWQHVVVLJQL¿FDQWO\DIIHFWHGUKHRORJLFDOSDUDPHWHUV ZKHUHDVVSDJKHWWLFRRNLQJTXDOLW\ZDVXQDIIHFWHGE\NHUQHOKDUGQHVV/RDIYROXPHH[KLELWHGDLQFUHDVH associated with kernel softening. In order to reduce plant height, soft durum lines with the lowest SKCS LQGH[HVZHUHIXUWKHUFURVVHGZLWKGXUXPZKHDWFY6LPHWRDQG)progeny lines were evaluated in terms of stability for their short height, soft texture and gluten quality. Modulation of kernel texture in durum wheat was obtained as well by transgenic approach by inserting vromindoline genes from oats. Finally soft textured durum wheat were used in crosses with Triticum aestivum with the aim to obtain extra-soft common wheats that may supply breeders with a broader range of kernel texture. Keywords..HUQHOWH[WXUH±'XUXPZKHDWEUHHGLQJ±3DVWDPDNLQJTXDOLW\±%UHDGPDNLQJTXDOLW\.

$PéOioration et TXaOité dX EOé dXr j te[tXre tendre 5ésXPé Les puroindolines A (PIN-A) et B (PIN-B) codées par les gènes Pina-D1 et Pinb-D1 sur le chromosome 5DS sont les principaux facteurs déterminants de la dureté du grain chez le blé commun, et exercent un fort impact sur plusieurs caractères de qualité tels que le rendement et la granularité de la farine, la dégradation de l’amidon et l’absorption d’eau, les propriétés rhéologiques et de cuisson. Les effets de la texture du grain sur la qualité de la production de pktes et de la pani¿cation ont été évalués dans les lignées de blé dur Soft (SDL) par rapport à leurs lignées sœurs Hard (HDL). Les SDL ont accumulé les puroindolines sur leurs granules d’amidon et ont montré des valeurs S.&S signi¿cativement inférieures à celles de leurs homologues Hard sans Pin-A et Pin-B. Le taux d’extraction moyen de farine des SDL était d’environ 23% plus élevé que celui des HDL. L’augmentation de la tendreté du grain a affecté de faoon signi¿cative les paramètres rhéologiques, alors que la qualité de cuisson des spaghettis n’a pas été affectée par la dureté du grain. Le volume du pain a présenté une augmentation de 10% associée au ramollissement de la graine. A¿n de réduire la hauteur des plantes, les lignées de blé dur tendre avec les plus faibles indices de S.&S ont ensuite été croisées avec des blés durs cv. Simeto et 17 lignées de descendance F6 ont été évaluées en termes de stabilité pour leur faible hauteur, la tendreté et la qualité du gluten. La modulation de la texture du grain de blé dur a ainsi été obtenue par une approche transgénique, en insérant des gènes de la vromindoline de l’avoine. (n¿n, le blé dur à grain tendre a été utilisé dans des croisements avec Triticum aestivum dans le but d’obtenir des blés communs extra-tendres qui peuvent fournir aux sélectionneurs une gamme plus large de texture des grains. MotsFOés Texture des grains ± Amélioration du blé dur ± 4ualité de la production des pktes ± 4ualité de la pani¿cation.

I – Introduction Kernel hardness is a main determinant of end product quality because of its strong effects on PLOOLQJFRQGLWLRQVJUDQXODULW\RIÀRXUDQGVWDUFKJUDQXOHLQWHJULW\,QSDUWLFXODUFRPPRQZKHDW

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(Triticum aestivum L.) cultivars can be divided into three endosperm-texture classes based on their average SKCS (Single Kernel Characterization System) values, i.e. soft, medium hard and hard. On the contrary, all durum wheat (T. turgidum ssp durum) cultivars are characterized by DQH[WUDKDUGNHUQHOWH[WXUHZLWK6.&6LQGH[!7KLVH[WUHPHO\KDUGWH[WXUHLVPDLQO\GXHWR the absence of PIN-A and PIN-B, two basic, tryptophan- and cysteine-rich polypeptides encoded E\WZRFORVHO\OLQNHGJHQHVQDPHG3LQD'DQG3LQE'ORFDWHGLQWKHGLVWDOSDUWRIWKHVKRUW DUPRIFKURPRVRPH' 0DWWHUQet al*DXWLHUet al DQGFRQVHTXHQWO\DEVHQWLQ AB-genome durum wheat. Extra-hard durum wheat grain is mainly ground to make semolina for the production of pasta and cous-cous, and in Mediterranean countries it is also used for breads RIDOOW\SHV 4XDJOLD3DOXPERet al %UHHGLQJSURJUDPVKDYHIRFXVHGRQVHOHFWLQJ durum wheat genotypes with superior pastamaking quality because of its primary commercial importance, and selection for baking quality has been applied to a minor extent (Boggini and 3RJQD  3HxD et al  %RJJLQL et al  /LX et al  3DOXPER et al.   7RPDNHDGXUXPEUHDGVHPROLQDLVUHJURXQGWRUHGXFHLWVSDUWLFOHVL]HDQGSURYLGHVXI¿FLHQW starch damage to assure appropriate gassing power during the fermentation process (Quaglia,  %HFDXVHRIWKHH[WUHPHKDUGQHVVRIGXUXPZKHDWJUDLQVHPROLQDUHJULQGLQJFDQUHVXOWLQ excessive starch damage, which alters alveogram and farinogram shapes, and exerts detrimental effects on baking performance (Dexter et al  In order to insert puroindoline genes into durum wheat, Gazza et al  XVHGGXUXPZKHDWOLQH ³&DSSHOOL0´ODFNLQJWKH3KORFXV *LRUJL DVWKHIHPDOHSDUHQWLQDFURVVZLWKWKH' %  VXEVWLWXWLRQOLQHRIGXUXPZKHDWFY/DQJGRQFDUU\LQJZLOGW\SHDOOHOHV3LQD'DDQG3LQE'D7KH UHVXOWLQJVRIWWH[WXUHGSODQWVGHYRLGRIFKURPRVRPH'ZHUHXVHGDVWKHPDOHSDUHQWLQFURVVHV with commercial durum wheat cv. Colosseo (Gazza et al DQGWKUHH)SODQWVHPL]\JRXVDW WKH3LQD'3LQE'ORFXVIURPWKHVHFURVVHVZHUHVHOISROOLQDWHGIRUWKUHHJHQHUDWLRQVWRGHYHORS VL[)OLQHVLHWKUHH6RIW'XUXP/LQHV 6'/ KRPR]\JRXVIRUZLOGW\SHDOOHOHV3LQD'DDQG3LQE 'DDQGWKUHH+DUG'XUXP/LQHV +'/ ODFNLQJWKH3LQD'DQG3LQE'JHQHV Here, soft-textured and hard-textured durum wheat lines are compared for their milling properties, rheological characteristics, pastamaking and breadmaking quality. In addition, in order to reduce SODQWKHLJKWDVHOHFWHG6'/OLQHZDVFURVVHGZLWKGXUXPZKHDWFY6LPHWRDQG)SURJHQ\ lines were evaluated in terms of stability for their short height, soft texture and gluten quality. Modulation of kernel texture was also obtained in transgenic durum wheat cv. Svevo containing vromindolines, two puroindoline-like proteins bound to starch granules, and likely responsible of the extra-soft texture of oat kernels. Production of extra-soft common wheat lines deriving from a cross between SDLs and common wheat is discussed as well.

II – Material and methods '1$ZDVH[WUDFWHGIURPOHDYHVE\WKH&7$%PHWKRG3XURLQGROLQHJHQHVZHUHDPSOL¿HGE\3&5 as described by Gautier et al  665 6LPSOH6HTXHQFH5HSHDW VHTXHQFHVRQFKURPRVRPH 'ZHUHXVHGIRUPLFURVDWHOOLWHPDUNHUFKDUDFWHUL]DWLRQ 6RPHUVet al6RQJet al  6WDUFKERXQGSURWHLQVZHUHH[WUDFWHGZLWKP01D&ODQG YY SURSDQROIURPPJ of air-dried starch granules as described previously (Corona et al $3$*(DWS+RI starch-bound proteins was carried out as described by Corona et al  5HGXFHGHQGRVSHUP proteins were fractionated by SDS-PAGE as described previously (Pogna et al  .HUQHOKDUGQHVVZDVSHUIRUPHGRQNHUQHOVVDPSOHE\WKH3HUWHQ6.&6 6SULQJ¿HOG IL, USA) following the manufacturer’s operating procedure. The instrument was set in a range RIKDUGQHVVYDOXHVEHWZHHQDQG6DPSOHV .J IURPVRIWWH[WXUHGDQGKDUGWH[WXUHG OLQHV ZHUH PLOOHG ZLWK L  WKH 0&. %XKOHU H[SHULPHQWDO PLOO IRU GXUXP ZKHDW LL  WKH 0/8  %XKOHUH[SHULPHQWDOPLOOIRUFRPPRQZKHDWRU LLL WKH%RQD5% %RQD,WDO\ H[SHULPHQWDOPLOO for common wheat.

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The milled samples were analyzed with the Chopin Alveograph (Chopin, Villeneuve La Garenne, )UDQFH  DFFRUGLQJ WR WKH PDQXIDFWXUHU¶V LQVWUXFWLRQV DV PRGL¿HG E\ '¶(JLGLR et al   ,Q DGGLWLRQWKHÀRXUVDPSOHVREWDLQHGZLWKWKH0/8%XKOHUH[SHULPHQWDOPLOOZHUHDQDO\]HG with the Brabender (South Hackensack, NJ) farinograph. Flour and semolina obtained from each soft-textured or hard-textured line with the MCK Buhler experimental mill for durum wheat were FRPELQHGDQGPL[HGZLWKWDSZDWHUWRUHDFKDGRXJKZDWHUFRQWHQWRI IRU6'/V RU IRU +'/V  7KH GRXJK ZDV SURFHVVHG LQWR VSDJKHWWL  PP LQ GLDPHWHU  XVLQJ D ODERUDWRU\ SUHVV$IWHUGU\LQJDWƒ&IRUKVSDJKHWWL J ZHUHFRRNHGDQGHYDOXDWHGIRU¿UPQHVV stickiness and bulkiness by a trained panel of three experts as described by D’Egidio et al.   %UHDGZDVEDNHGDFFRUGLQJWRWKH$$&&0HWKRG%ZLWKPLQRUPRGL¿FDWLRQV &DWWDQHRDQG %RUJKL XVLQJÀRXUVDPSOHVREWDLQHGZLWKWKHPLOOLQJIRUFRPPRQZKHDW/RDIYROXPHZDV determined by rapeseed displacement. All data are the means of at least duplicate determinations. Data were statistically evaluated by Student’s t test or analysis of variance.

III – Results and Discussion 3&5DPSOL¿FDWLRQVRIJHQRPLF'1$ZLWKSULPHUSDLUVVSHFL¿FIRUVHYHQPLFURVDWHOOLWHVORFDWHG RQ '  FKURPRVRPH VXJJHVWHG WKDW 6'/V FRQWDLQ RQO\ D VPDOO '6 IUDJPHQW LQIHULRU WR  F0LQVL]HOLNHO\WUDQVORFDWHGWRKRPRHRORJRXVFKURPRVRPH%68SRQ$3$*(IUDFWLRQDWLRQ soft textured durum wheat lines were found to accumulate PIN-A and PIN-B on the surface of their starch granules in amounts comparable to those observed in soft-textured common wheat FXOWLYDUV$FFXPXODWLRQRISXURLQGROLQHVUHGXFHG6'/VPHDQ6.&6LQGH[HVWRZKLFK are typical of soft-textured common wheat cultivars whereas hard-textured durum wheat lines HDL were similar to durum wheat varieties in lacking both puroindolines. According to SDS-PAGE IUDFWLRQDWLRQ DOO GXUXP ZKHDW OLQHV SURGXFHG LQ WKH SUHVHQW VWXG\ H[KLELWHG /0: JOXWHQLQ subunits, which are associated with superior gluten strength (Pogna et al DQGLQKHULWHG +0:JOXWHQLQVXEXQLWSDLUIURP/DQJGRQ' % VXEVWLWXWLRQOLQH 6'/VUHYHDOHGWKDWJUDLQKDUGQHVVKDVDVWURQJLQÀXHQFHRQVHYHUDOTXDOLW\UHODWHGWUDLWVDWWKH WHWUDSORLGOHYHODVZHOO,QSDUWLFXODUWKHDYHUDJHÀRXUH[WUDFWLRQUDWHRI6'/VZDVDSSUR[LPDWHO\ KLJKHUWKDQWKDWRI+'/VDQGHYHQJUHDWHU DERXW DIWHUPLOOLQJZLWKWKH0&.%XKOHU mill for durum wheat. Grain softness strongly decreased farinograph water absorption and consequently resulted in inferior dough tenacity (P), strength (W) and P/L ratio of SDLs with respect to HDLs. Moreover, the lower starch damage accounts for the higher farinograph dough stability and mixing tolerance of SDL milling products, which likely derives from their lower water absorption. It is noteworthy that the substantial variation in water absorption and rheological properties associated with the contrasting kernel textures of the durum wheat lines did not VLJQL¿FDQWO\DIIHFW¿UPQHVVVWLFNLQHVVDQGEXONLQHVVRIVSDJKHWWL,QDGGLWLRQ+'/VDQG6'/VGLG QRWGLIIHUVLJQL¿FDQWO\IRUWKHLUSDVWDPDNLQJTXDOLW\DVGHWHUPLQHGE\WKHJOREDOTXDOLW\VFRUHDQG were comparable with high-quality durum wheat cultivars grown in Italy. On the other hand, softWH[WXUHGOLQHVVKRZHGDVPDOOEXWVLJQL¿FDQWLQFUHDVHRIWKHEUHDGORDIYROXPH DSSUR[LPDWHO\  FRPSDUHGZLWKWKHLUKDUGWH[WXUHGFRXQWHUSDUWV7KHVHUHVXOWVVXJJHVWWKDWPRGXODWLRQRI kernel hardness in durum wheat does not impair its pasta-making potential, and may improve its baking performance. $VFRPSDUHGZLWK+'/VWKHVRIWWH[WXUHRI6'/VUHVXOWHGLQVLJQL¿FDQWORZHU\HOORZDQGEURZQ LQGH[HVLQERWKÀRXUDQGVHPROLQDIUDFWLRQVREWDLQHGZLWKWKH0&.%XKOHUPLOO7KLVVXJJHVWWKDW FRORUZDVVWURQJO\UHODWHGWRWKHSDUWLFOHVL]HRIWKHPLOOLQJIUDFWLRQV\HOORZQHVVE DQGEURZQQHVV / EHLQJFRQVLVWHQWO\DQGVLJQL¿FDQWO\ORZHULQWKH¿QHUÀRXUDQGVHPROLQDRI6'/V

Proceedings of the International Symposium on Genetics and breeding of durum wheat

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7KHKLJKSODQWKHLJKW !FP RI6'/VDQG+'/VUHVXOWHGLQSDUWLDOORGJLQJDWKDUYHVWLQJ XQWLO  ,QRUGHUWREUHHGVKRUWHUFXOWLYDUVDQ6'/OLQHZDVFURVVHGZLWKGXUXPZKHDWFY6LPHWR Segregation of allele Pina-D1a FRGLQJ IRU ZLOGW\SH 3,1$ ZDV IROORZHG E\ 3&5 DPSOL¿FDWLRQ on single F plants, whereas segregation of texture in F kernels produced by each Fplant was determined by SKCS. Amongst the FUHVXOWLQJSURJHQ\WHQVRIWWH[WXUHGLQGLYLGXDOVFPLQ KHLJKWZHUHVHOHFWHG 7DEOH 0RUHRYHUWKUHHJHQRW\SHV DQG ZHUHIRXQG WRFRQWDLQ+0:JOXWHQLQVXEXQLWSDLU )LJ ZKLFKDUHDVVRFLDWHGWRJOXWHQVWUHQJWK 7DEOH0HDQYDOXHVRISODQWKHLJKW6.&6LQGH[HVDQGVHHGZHLJKWRI)6 individuals of the cross between SDL1 and durum wheat cv Simeto. Genotype                 

Plant height ± SD (cm) “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “

SKCS ± SD “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “

Seed weight±SD (mg) “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “

Figure1. SDS-PAGE fractionation of storage proteins of eight F6OLQHVREWDLQHGIURPWKHFURVV6'/[ Simeto. HMW glutenin subunits pair 7+8 is indicated.

Modulation of kernel texture was obtained in transgenic durum wheat containing vromindolines from oats as well. Two plasmids containing either Vin-2 or Vin-3 genes were used to co-transform durum wheat cv. Svevo by the biolistic method. Vin-2 and Vin-3 from oats (Avena sativa) code IRU YURPLQGROLQHV  DQG  9,1 DQG 9,1  UHVSHFWLYHO\ WZR VWDUFKERXQG SXURLQGROLQHOLNH

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SURWHLQV VKDULQJ D WU\SWRSKDQULFK GRPDLQ RI IRXU WU\SWRSKDQ UHVLGXHV$ WRWDO RI  7 plants DQG7 plants expressing both Vin-2 and Vin-3 genes were grown in growth chambers, and characterized for their genetic structure and kernel texture. :KHQ JURZQ XQGHU FKDPEHU FRQGLWLRQV WKH  7 progeny of two transgenic T individuals with expression of both Vin-2 and Vin-3 were indistinguishable from control cv. Svevo for their morpho-physiological traits and fertility. Transgenic plants produced TNHUQHOVVLJQL¿FDQWO\VRIWHU PHDQ6.&6 “ WKDQWKHLUVLVWHUSODQWVZLWKRXWYURPLQGROLQHWUDQVJHQHV PHDQ6.&6  “7DEOH   As expected, heterozygous T1 plants generated seeds with contrasting KDUGQHVVFKDUDFWHULVWLFVZLWK6.&6YDOXHVUDQJLQJIURPWR7KHVHUHVXOWVVXJJHVWWKDW YURPLQGROLQHV9,1DQG9,1DUHDEOHWRPRGXODWHJUDLQWH[WXUHLQGXUXPZKHDWDVZHOO Soft-textured F lines of durum wheat containing PIN-A and PIN-B were crossed with common wheat cvs Eridano and Bilancia. The hard-textured cv. Eridano contains null allele Pina-D1b and wild-type allele Pinb-D1a, whereas the soft-textured cv. Bilancia has wild type alleles Pina-D1a/ Pinb-D1a. Amongst the F progeny of cv. Eridano, individuals with three doses of puroindolines, i.eRQHRI3,1$DQGWZRRI3,1%ZHUHVHOHFWHGE\3&5DPSOL¿FDWLRQZLWKSULPHUVVSHFL¿FWR Pina-D1b or Pina-D1a )LJ$ DQGIRXQGWRKDYHDPHDQ6.&6YDOXHRI )LJ% 0RUHRYHU the F SURJHQ\ RI FY %LODQFLD VKRZHG D PHDQ 6.&6 YDOXHV RI  ZLWK H[WUDVRIW LQGLYLGXDOV H[KLELWLQJDQ6.&6LQGH[DVORZDV Table 2. SKCS values of kernels produced by 34 T1 plants from two T0 plants of durum wheat cv. Svevo H[SUHVVLQJVin-2 and Vin- Genotype of T1 plants Homozygous Heterozygous Null Control cv. Svevo

No. of T1 plants    

No. of T2 kernels    

Mean SKCS value ± SD “

“ “ “QV

SKCS Range 0LQ0D[        

t-value signi¿cant at P  0.01 with respect to the null T2 progeny ns, t-value not signi¿cant with respect to the null T2 progeny.

)LJXUH  5HVXOWV RI 3&5 DPSOL¿FDWLRQ RI ) 6'/ OLQHV XVLQJ DOOHOH 3LQD'D RU 3LQD 'E VSHFL¿F primer.

Proceedings of the International Symposium on Genetics and breeding of durum wheat

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IV – Conclusions 'XUXPZKHDWOLQHVKRPR]\JRXVIRUDF0WHUPLQDOIUDJPHQWRIFKURPRVRPH'FRQWDLQLQJ WKH 3LQD'D3LQE'D DOOHOHV VKRZHG 6.&6 YDOXHV W\SLFDO RI VRIWWH[WXUHG NHUQHOV 6RIWHQLQJ HIIHFW UHVXOWHG LQ D DERXW  KLJKHU ÀRXU H[WUDFWLRQ UDWHV FRPSDUHG ZLWK KDUGWH[WXUHG OLQHV 6SDJKHWWLFRRNLQJTXDOLW\ZDVXQDIIHFWHGE\NHUQHOKDUGQHVVZKHUHDVORDIYROXPHH[KLELWHGD increase associate with kernel softening. Availability of soft-textured durum wheat genotypes may have important practical and useful implications for breeding multiple-purpose durum wheat (pasta, bread, biscuits and other oven products), and for technological operations of industrial interest.

References Boggini G., Pogna N.E., 1989. The breadmaking quality and storage protein composition of Italian durum wheat. J. Cereal Sci.,SS Boggini G., Tusa P., Pogna N.E., 1995. Breadmaking quality of durum wheat genotypes with some novel glutenin subunit composition. J. Cereal Sci.SS Cattaneo M., Borghi B., 1979. 3ULPL ULVXOWDWL GL SURYH VSHULPHQWDOL GL SDQL¿FD]LRQH VX YDULHWj LWDOLDQH GL frumento tenero. Tecnica Molitoria,  SS Corona V., Gazza L., Boggini G., Pogna N.E., 2001. Variation in friabilin composition as determined by $3$*( IUDFWLRQDWLRQ DQG 3&5 DPSOL¿FDWLRQ DQG LWV UHODWLRQVKLS WR JUDLQ KDUGQHVV LQ EUHDG ZKHDW J. Cereal Sci.,SS D’Egidio M.G., Mariani B.M., Nardi S., Novaro P., Cubadda R., 1990. Chemical and technological variables and their relationships: a predictive equation for pasta cooking quality. Cereal Chemistry,SS± 'H[WHU -( 3UHVWRQ .5 0DUWLQ '* *DQGH (-  The effects of protein content, and starch damage on the physical dough properties and breadmaking quality of Canadian durum wheat. J. Cereal Sci., SS Gautier M.F., Aleman M.E., Guirao A., Marion D., Joudrier P., 1994. Triticum aestivum puroindolines, two basic cysteine-rich seed proteins: cDNA analysis and developmental gene expression. Plant Molecular Biology,SS Gazza L., Niglio A., Mei E., De Stefanis E., Sgrulletta D., Pogna N.E., 2003. Production and characterization of durum wheat (Triticum turgidum ssp. durum) lines containing puroindolines A and B. In: Proceedings of 2nd International Workshop Durum Wheat and Pasta Quality: Recent Achievements and New Trends. 5RPH1RYHPEHUSS Gazza L., Zanella L., Pogna N.E., 2008. Development of durum wheat (Triticum turgidum ssp durum) lines with soft kernel texture by chromosome engineering. In: Proceedings of 11th International Wheat Genetics Symposium.%ULVEDQH4/'$XVWUDOLD9ROSS Giorgi B., 1978. A homoeologous pairing mutant isolated in Triticum durum cv Cappelli. Mutation Breeding Newsletter, SS Liu C.Y., Shepherd K.W., Rathjen A.J., 1996. Improvement of durum wheat pasta-making and breadmaking qualities. Cereal Chemistry,SS Mattern P.Y., Morris R., Schmidt Y.W., Johnson V.A., 1973. Location of genes for kernel properties in the wheat varieties Cheyenne and chromosome substitution lines. In: Proceedings of the Ist International Wheat Genetics Symposium. Sears E.R., Sears L.M.S. (eds), University of Missouri Press, Columbia, 0286$SS Peña R.J., Zarco-Hernandez J., Amaya-Celis A., Mujeeb-Kazi A., 1994. Relationship between chromosome %HQFRGHGJOXWHQLQVXEXQLWFRPSRVLWLRQVDQGEUHDGPDNLQJTXDOLW\FKDUDFWHULVWLFVRIVRPHGXUXPZKHDW (Triticum turgidum) cultivars. J. Cereal Sci.,SS 3RJQD1($XWUDQ-&0HOOLQL)/D¿DQGUD')HLOOHW3&KURPRVRPH%HQFRGHGJOLDGLQVDQG glutenin subunits in durum wheat: genetics and relationship to gluten strength. J. Cereal Sci.,SS Pomeranz Y., Williams P. C., 1990. Wheat hardness: Its genetic, structural, and biochemical background, PHDVXUHPHQWDQGVLJQL¿FDQFH,QAdvances in Cereal Science and TechnologyYRO3RPHUDQ]