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Conxita Royo
Miloudi M. Nachit
Natale Di Fonzo
José Luis Araus
Wolfgang H. Pfeiffer
Gustavo A. Slafer
Editors
Durum Wheat Breeding
Current Approaches
and Future Strategies
Volume 1

Pre-publication
REVIEWS, “Thisof comprehensive
book is a very valuable source
information. It
COMMENTARIES, brings together an exceptionally large
number of internationally recognized
EVALUATIONS . . . experts in the field, providing broad
coverage of today’s knowledge as well
as perspectives on the future of durum
wheat breeding. The book covers es-
sential theoretical background but also
introduces in detail national breeding
procedures, achievements, and future
lines. This work will be highly welcomed
by undergraduate and postgraduate
students, cereal scientists, agronomists,
plant breeders, and other players and
experts in the field, all over the world.”

Pirjo Peltonen-Sainio, PhD

Professor, Plant Production Research,
MTT Agrifood Research, Finland
More pre-publication
REVIEWS, COMMENTARIES, EVALUATIONS . . .

urum Wheat Breeding is a com- plored. A feature of this work is the in-
“D prehensive compilation of every-
thing durum wheat researchers need to
depth analysis of durum wheat quality,
spanning the broad range of products
know. It reviews durum wheat from its made from durum wheat and their un-
origins in the eastern Mediterranean to derlying genetic control to the methods
its current position as one of the plant breeders use to improve quality.
world’s most important cereals for hu- All the primary diseases, pests, and
man food consumption. The great stresses that limit durum wheat pro-
duction around the world and the ge-
range of products made from durum netic options available to plant breed-
wheat, the key characteristics needed ers to combat these constraints are
to obtain suitable processing quality, discussed. This book will be an invalu-
and the primary constraints to durum able aid for students, wheat researchers
wheat production are reviewed, as are of any age, and those simply interested
the many different breeding methods in plant breeding. Never before has such
used around the world today. a comprehensive treatise on durum
The book discusses durum wheat wheat breeding been assembled.”
distribution and taxonomy, genetics,
physiology, breeding, and applied and Masa Iwanaga, PhD
strategic research. The fields of mo- Director General,
lecular biology, cytogenetics, statistics, International Maize and Wheat
and crop physiology as they apply to Improvement Center (CIMMYT),
durum wheat improvement are ex- Mexico

Food Products Press®

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Durum Wheat Breeding
Current Approaches
and Future Strategies
Volume 1
 FOOD PRODUCTS PRESS®
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Durum Wheat Breeding
Current Approaches
and Future Strategies
Volume 1

Part I: World Distribution and the Role

of Durum Wheat Breeding
Part II: Genetics and Molecular Aspects
Part III: Physiological Bases of Durum
Wheat Improvement

Conxita Royo
Miloudi M. Nachit
Natale Di Fonzo
José Luis Araus
Wolfgang H. Pfeiffer
Gustavo A. Slafer
Editors

Food Products Press®

An Imprint of The Haworth Press, Inc.
New York • London • Oxford
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Published by

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13904-1580.

© 2005 by The Haworth Press, Inc. All rights reserved. No part of this work may be reproduced or
utilized in any form or by any means, electronic or mechanical, including photocopying, microfilm,
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from the publisher. Printed in the United States of America.

Cover design by Kerry E. Mack.

Cover photos by Javier Goñi, Josep Anton Betbesé, and Wolfgang H. Pfeiffer.

Library of Congress Cataloging-in-Publication Data

Durum wheat breeding : current approaches and future strategies, volumes 1 and 2 / [edited by
Royo, Conxita, Miloudi Nachit, Natale Di Fonzo].
p. cm.
Includes bibliographical references and index.
ISBN 1-56022-333-2 Volumes 1 and 2 (hard : alk. paper)
ISBN 1-56022-966-7 Volume 1 (hard : alk. paper)
ISBN 1-56022-967-5 Volume 2 (hard : alk. paper)
1. Durum wheat—Breeding. 2. Durum wheat—Genetic engineering. I. Royo, C. (Conxita)
II. Nachit, Miloudi. III. Di Fonzo, Natale.

SB191.W5D932 2005
633.1'17—dc22
2004021600
 CONTENTS

VOLUME 1

About the Editors xvii

Contributors xix

List of Abbreviations xxvii

PART I: WORLD DISTRIBUTION AND THE ROLE

OF DURUM WHEAT BREEDING

Chapter 1. Wheat: Its Concept, Evolution, and Taxonomy 3

James Mac Key
Introduction 3
Preadaptation of Wheat and Its Relatives 3
Wild Wheats and Their Spread 8
The Domestication of 2x and 4x Wheats 16
Impact of Polyploidization on Wheat Expansion 26
The Genus Triticum, Concept, and Subdivision 31

Chapter 2. End Products: Present and Future Uses 63

Elias M. Elias
Frank A. Manthey
Introduction 63
Pasta Products 64
Couscous 72
Bulgur 74
Frekeh or Firik 75
Breakfast Cereal 76
Durum Wheat Bread 77
Other Products 80
Conclusion 81
Chapter 3. Importance of Breeding for Further Improving
Durum Wheat Yield 87
Gustavo A. Slafer
Daniel F. Calderini
Introduction 87
Wheat Yield Trends 88
Sources for Yield Improvement in Wheat: Past Trends
and Likely Future 89
PART II: GENETICS AND MOLECULAR ASPECTS
Chapter 4. A Retrospective Analysis of Genetic Diversity
in Durum Wheat Elite Germplasm Based
on Microsatellite Analysis: A Case Study 99
Marco Maccaferri Enrico Porceddu
Maria Corinna Sanguineti Roberto Tuberosa
Paolo Donini
Introduction 99
Gene Pools in Durum Wheat and Their Genetic Diversity 100
Evaluating Genetic Diversity and Genetic Relationships
in Elite Germplasm 104
The Italian Durum Germplasm: A Valuable Breeding
Resource 106
Conclusion 130
Chapter 5. Managing and Collecting Genetic Resources 143
Bent Skovmand
Marilyn L. Warburton
Shawn N. Sullivan
Jacob Lage
Introduction 143
Genetic Resources 144
Genetic Conservation and Erosion 145
Wheat Collection Needs 146
Strategies for Collecting Wheat Landraces and Other
Genetic Resources 148
Passport Data 148
Applying Molecular Techniques to Genetic Resource
Management and Utilization 152
 Legal Issues Affecting Access to Plant Genetic Resources 154
Summary 158
Appendix: Information in a CIMMYT Collection Data
Sheet 159

Chapter 6. The Cytogenetic Contribution to the Analysis

and Manipulation of the Durum Wheat Genome 165
Carla Ceoloni
Marina Pasquini
Rosanna Simeone
Wheat Cytogenetics in the Molecular Era 165
Considerations on Genome Evolution and Differentiation 166
Development and Use of Cytogenetic Stocks in Tetraploid
Wheat 168
The Genetic Control of Intergenomic Relationships 178
Tetraploid Durum Wheat As a Recipient of Alien Gene
Transfers 184
Conclusion 196

Chapter 7. Comparative Genetics of Durum Wheat

and Other Triticeae 209
Ramesh V. Kantety
Ayman A. Diab
Mark E. Sorrells
Introduction 209
Comparative Genetics of the Grasses 211
Conclusion 219

Chapter 8. Genetic Manipulation of Durum Wheat:

Application to Grain Composition and Quality 225
Peter R. Shewry Paola Tosi
Pauline Bell Paul Lazzeri
Natale Di Fonzo Pilar Barcelo
Carmela Lamacchia
Introduction 225
Genetic Modification Technology 226
Manipulation of Grain Composition and Quality 234
Conclusion 246
Chapter 9. Genetic Bases of Resistance to Abiotic Stresses
in Durum Wheat (Triticum turgidum ssp. durum) 255
Anna Maria Mastrangelo Laura Arru
Caterina Marè Natale Di Fonzo
Elisabetta Mazzucotelli Nicola Pecchioni
Enrico Francia Luigi Cattivelli
Introduction 255
Chromosome Regions Involved in Abiotic Stress Tolerance 256
The Molecular Stress Response: Basic Knowledge
from Model Plants 263
Gene Expression Under Drought Stress in Durum Wheat 270
Gene Expression at Low Temperature in Durum Wheat 274
Enhancement of Stress-Response Mechanisms Leads
to Improved Stress Tolerance 276
Genomics Approach to Stress Tolerance in Durum Wheat 279
Chapter 10. Resistance to Diseases 291
Ravi P. Singh Lucy Gilchrist
Julio Huerta-Espino Monique Henry
Guillermo Fuentes Julie M. Nicol
Etienne Duveiller
Introduction 291
The Rusts 291
Bunts and Smuts 297
Foliar Blights: Spot Blotch and Tan Spot 301
Foliar Blight: Septoria tritici Leaf Blotch 304
Fusarium Head Blight 307
Viruses 311
Root Rots 315
Chapter 11. Genetic Basis for Insect Pest Resistance
in Durum Wheat 329
Nasserlehaq Nsarellah Ahmed Amri
Saadia Lhaloui Miloudi M. Nachit
Mustapha El Bouhssini
Introduction 329
Major Insect Pests Affecting Wheat 330
Mechanisms of Resistance to Insects in Plants 332
Genetic Basis of Insect Resistance in Durum Wheat 333
Conclusion 341
Chapter 12. Genetic Bases of Grain Quality 349
Magdalena Ruiz
José Francisco Vázquez
José María Carrillo
Introduction 349
Grain Protein Concentration 349
Genetic Bases of Protein Quality 350
Gliadins and Glutenins 351
Semolina and Pasta Color 363
Milling Quality 364
Starch Quality 365
Molecular Markers and Grain Quality 365

PART III: PHYSIOLOGICAL BASES OF DURUM

WHEAT IMPROVEMENT

Chapter 13. Genetic Improvement Effects on Durum

Wheat Yield Physiology 379
Luis F. García del Moral
Conxita Royo
Gustavo A. Slafer
Introduction 379
Yield Gains Achieved by Breeding in the Twentieth
Century 380
Physiological Determinants of the Yield Gains by Durum
Wheat Breeding 385
Changes in Yield Components 385
Changes in Dry-Matter Production and Partitioning 389
Conclusion 390

Chapter 14. Durum Wheat Ideotypes for Sustainable

Farming in Diversified Environments 397
Paolo Annicchiarico Agata Rascio
Luciano Pecetti Gaetano Boggini
Zina Flagella
Introduction 397
Ideotypes for Unfavorable Environments 400
 Ideotypes for Favorable Environments 408
The Contribution of Physiological Traits to Selection
for Drought Tolerance 412
Chapter 15. Physiological Basis of Yield Potential
in Durum Wheat 425
Philippe Monneveux
Matthew P. Reynolds
Jean Marcel Ribaut
Introduction 425
History of Genetic Progress in Durum Wheat 427
Exploration of Physiological Strategies to Improve Yield
Potential 429
Molecular Approaches for Improving Yield Potential 445
Conclusion 449
Chapter 16. Adaptation to Water Stress: Methodologies
for the Study of the Photosynthetic Response 461
Salvador Nogués
Eduardo A. Tambussi
José Luis Araus
Introduction 461
Methodological Approach: The Case of Durum Wheat 465
Conclusion 477
Chapter 17. Functional Determinants of Grain Quality 483
Mario Augusto Pagnotta
Antonio Blanco
Agata Gadaleta
Clara Fares
Introduction 483
Chemistry of Wheat Proteins 487
Protein Content 492
Nutrition Properties of Wheat 498
Grain Hardness, Whiteness, and Vitreousness 499
Starch and the Waxy Lines 500
Lipids and Lipoproteins 504
Color 505
Enzymatic Activity 509
Ash 511
 VOLUME 2

PART IV: BREEDING STRATEGY AND METHODOLOGY

Chapter 18. Increasing Yield Potential and Stability
in Durum Wheat 531
Wolfgang H. Pfeiffer
Richard M. Trethowan
Karim Ammar
Kenneth D. Sayre
Introduction 531
Yield Potential: Perceptions and Definitions 531
Integrating and Differentiating Factors Related
to Progress in GYP 535
Origins of Durum Breeding for GYP 537
Genetic Progress in GYP of Hallmark CIMMYT Durum
Cultivars 541
Genetic Progress in GYP in Contemporary CIMMYT
Durums 542
Identifying Avenues for Raising GYP 544
Chapter 19. Selection Tools for Improving Yield-
Associated Physiological Traits 563
Conxita Royo Miloudi M. Nachit
Luis F. García del Moral José Luis Araus
Gustavo A. Slafer
The Role of Breeding to Increase Grain Yield 563
Empirical versus Analytical Breeding: An Integrated
Breeding Approach? 564
Accelerating Genetic Progress Through New Selection
Tools 565
Selection Tools for Some Integrative Traits 577
Chapter 20. Selection Strategies for Traits Relevant
for Winter and Facultative Durum Wheat 599
Anatoliy Palamarchuk
Agroecological Characterization of Winter and Facultative
Durum Regions 599
Actual Winter and Facultative Durum Wheat Areas 601
 Winter and Facultative Durum Wheat Production 605
Durum Wheat End Uses 607
Introduction to Durum Wheat and Durum Wheat History 608
Local Populations and Varieties 610
From Local Winter and Facultative Durum Types
to Commercial Varieties 613
Selection Under Optimum Conditions 632

Chapter 21. Selection Strategies and Methodologies

for Biotic Stresses in Durum Wheat 645
Nasserlehaq Nsarellah
Mustapha El Bouhssini
Julie M. Nicol
El-Mostafa Tawfiq
Introduction 645
Biotic Stresses Situation: Diagnosis and Priority Setting 646
Considerations for Strategies and Methodologies Regarding
Biotic Stress Research in Durum Wheat 648
Screening Methodologies 657
Conclusion 659

Chapter 22. Breeding Methodologies and Strategies

for Durum Wheat Quality Improvement 663
Roberto J. Peña
Wolfgang H. Pfeiffer
Introduction 663
Wheat-Based Foods and Grain Quality Requirements 665
Durum Wheat Quality Parameters 674
End-Use Quality in Crop Improvement 685
Chapter 23. Wide Crosses for Durum Wheat Improvement 703
Abdul Mujeeb-Kazi
Introduction 703
The Triticeae Gene Pools: Genetic Diversity
and Distribution 704
Utilization and Practicality of Wide Cross Germplasm 731
 Some Practical Trends for the Future 736
Conclusion 741

Chapter 24. Doubled-Haploid Technique in Durum Wheat

Breeding 745
Juan Ballesteros Carmen García-Llamas
Ana Maria Castillo Antonio Martín
Luis Cistué Maria Pilar Vallés
Introduction 745
Doubled-Haploid Production of Durum Wheat Through
Interspecific Hybridization 746
Durum Wheat Doubled-Haploid Production
by Androgenesis 759

Chapter 25. Management of Genotype ´ Environment

Interactions and Their Implications for Durum
Wheat Breeding 777
Richard M. Trethowan
Jose Crossa
Wolfgang H. Pfeiffer
Introduction 777
Methods for Studying Crossover Genotype ´ Environment
Interaction 779
Multiplicative Models for Assessing Crossover Genotype ´
Environment Interaction Using CIMMYT Durum Wheat
MET Data 784
Implications of Better Understanding GEI in Durum Wheat
Breeding 793
Chapter 26. Design and Analysis of Field Experiments 803
Walter T. Federer
Jose Crossa
Introduction 803
Plot Technique 806
Selecting an Experiment Design 808
Block Experiment Designs 808
 Row-Column Experiment Designs 811
Unreplicated or Screening Experiment Designs 812
Exploratory Model Selection 816
Multisite and/or Multiyear Experiments 818
Parsimonious Experiment Designs 819
Discussion and Summary 819

PART V: APPLIED AND STRATEGIC RESEARCH

IN DURUM BREEDING PROGRAMS

Chapter 27. Durum Wheat Improvement in Italy 825

Natale Di Fonzo Antonio Blanco
Stefano Ravaglia Antonio Troccoli
Enzo DeAmbrogio
Introduction 825
Relevant Breeding Aspects 827
Breeding for Abiotic Stress 848
Breeding for Biotic Stress 852
Breeding for Quality 856
Future Perspective 862

Chapter 28. Durum Wheat Improvement in Spain 883

Conxita Royo
Importance of Durum Wheat in Spain 883
Variety, Structure, and Use of Certified Seed 886
Durum Wheat Breeding in Spain 888
Genetic Resources 896
Quality Improvement 898
Research on Morphophysiological Traits to Improve
Selection Efficiency 899
Overall Genetic Progress 901

Chapter 29. The Romanian Durum Breeding Program 907

Nicolae N. Sãulescu
Durum Wheat in Romania 907
The Environment for Durum Wheat Production in Romania 908
History of Durum Wheat Breeding in Romania 909
Challenges Ahead 913
 Breeding Approaches 916
Conclusion 918

Chapter 30. Durum Wheat Improvement in Canada 921

John M. Clarke
Introduction 921
Production 922
Variety Release 922
Breeding Procedures 924
Improvement of Production Traits 927
Improvement of Quality Traits 929
Future Directions 934

Chapter 31. Durum Wheat Breeding at North Dakota

State University 939
Elias M. Elias
Frank A. Manthey
Introduction 939
Objectives of the Breeding Program 940
Breeding Methods 940
Breeding for Agronomic Traits 947
Breeding for Disease Resistance 949
Breeding for Quality Traits 952

Chapter 32. Durum Wheat Genetic Improvement

in Morocco 963
Nasserlehaq Nsarellah
Ahmed Amri
Introduction 963
Durum Wheat Production in Morocco 963
Characterization of Production Environments
in Morocco 966
Constraints to Durum Wheat Production in Morocco 967
Historical Considerations of Durum Wheat Improvement
in Morocco 969
The Current Durum Wheat Breeding Program 971
Major Achievements 978
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