VEGETABLE GRAFTING Principles and Practices This book is dedicated to the memory of our friend Prof. Dr Jung-Myung Lee who was a pioneer in the field of vegetable grafting and gave a substantial contribution to the first chapter of this book. VEGETABLE GRAFTING Principles and Practices Giuseppe Colla Department of Agricultural and Forestry Sciences University of Tuscia Italy Francisco Pérez-Alfocea Department of Plant Nutrition Centro de Edafología y Biología Aplicada del Segura (CEBAS) Consejo Superior de Investigaciones Científicas (CSIC) Campus Universitario de Espinardo Spain Dietmar Schwarz Leibniz Institute of Vegetable and Ornamental Crops Germany CABI is a trading name of CAB International CABI CABI Nosworthy Way 745 Atlantic Avenue Wallingford 8th Floor Oxfordshire OX10 8DE Boston, MA 02111 UK USA Tel: +44 (0)1491 832111 Tel: +1 (617)682-9015 Fax: +44 (0)1491 833508 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org CAB International, 2017. © 2017 by CAB International. Vegetable Grafting: Principles and Practices is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Names: Colla, Giuseppe, 1972- | Pérez-Alfocea, F. (Francisco) | Schwarz, Dietmar, 1956- Title: Vegetable grafting : principles and practices / [edited by] Giuseppe Colla, Department of Agricultural and Forestry Sciences, University of Tuscia, Viterbo, Italy, Francisco Pérez-Alfocea, Department of Plant Nutrition, CEBAS-CSIC, Murcia, Spain, Dietmar Schwarz, Leibniz Institute of Vegetable and Ornamental Crops, Grossbeeren, Germany. Description: Wallingford, Oxfordshire, UK : CAB International, [2017] | Includes bibliographical references and index. Identifiers: LCCN 2016057959 | ISBN 9781780648972 (hbk : alk. paper) Subjects: LCSH: Vegetables. | Grafting. | Rootstocks. Classification: LCC SB324.7 .V43 2017 | DDC 635--dc23 LC record available at https://lccn.loc.gov/2016057959 ISBN-13: 978 1 78639 058 5 Commissioning editor: Rachael Russell Editorial assistant: Emma McCann Production editor: Shankari Wilford Typeset by SPi, Pondicherry, India Printed and bound in the UK by CPI Group (UK) Ltd, Croydon, CR0 4YY, UK Contents Contributors xi Preface xv Acknowledgements xvii 1 Introduction to Vegetable Grafting 1 Zhilong Bie, Muhammad Azher Nawaz, Yuan Huang, Jung-Myung Lee and Giuseppe Colla 1.1. Importance and Use of Vegetable Grafting 1 1.1.1. Historical perspective 1 1.1.2. Purpose and scope 2 1.1.3. Growing areas and plantlet production 5 1.2. The Process of Vegetable Grafting 6 1.2.1. Selection of rootstock and scion cultivars 6 1.2.2. Overview of grafting methods 7 1.2.3. Preference of grafting method for different species 12 1.2.4. Post-graft healing environment 12 1.3. Problems Associated with Vegetable Grafting 14 1.4. Conclusions 15 References 15 2 Genetic Resources for Rootstock Breeding 22 Maria Belen Pico, Andrew J. Thompson, Carmina Gisbert, Halit Yetis¸ir and Penelope J. Bebeli 2.1. Genetic Diversity 22 2.1.1. Diversity in the Cucurbitaceae family 22 2.1.2. Diversity in the Solanaceae family 24 2.2. Gene Bank Collections 27 2.2.1. Cucurbitaceae 27 2.2.2. Solanaceae 31 v vi Contents 2.3. Current Usage of Genetic Material in Rootstocks 36 2.3.1. Rootstocks for cucurbit production 36 2.3.2. Rootstocks for production of solanaceous crops 44 2.4. Germplasm Collections and Grafting in Other Plant Families 54 2.4.1. Cynara grafting 54 2.4.2. Phaseolus grafting 54 2.5. Conclusions 55 Acknowledgements 55 References 55 3 Rootstock Breeding: Current Practices and Future Technologies 70 Andrew J. Thompson, Maria Belen Pico, Halit Yetişir, Roni Cohen and Penelope J. Bebeli 3.1. Introduction 70 3.2. Stacking Traits: Meiosis or Grafting or Both? 70 3.3. Developing Stable Core Collections of Germplasm for Breeding 73 3.4. Deploying Genetic Diversity for Rootstocks 74 3.4.1. General principles 74 3.4.2. Use of Cucurbita F1 hybrids 75 3.4.3. Use of Solanum F1 hybrids 76 3.4.4. Interspecific hybrids and hybridization barriers 76 3.5. Grafting as a Tool For Genetic Hybridization and Chimera Production 77 3.5.1. Genetic hybridization: transfer of nuclear and organellar DNA between cells of the graft union 78 3.5.2. Use of grafting to generate chimeras 79 3.6. Selection of Improved Rootstocks 80 3.6.1. Phenotypic selection 80 3.6.2. Marker-assisted selection 82 3.7. Transgenic Rootstocks 84 3.8. Rootstock Registration and Commercialization 85 Acknowledgements 85 References 85 4 Rootstock-scion Signalling: Key Factors Mediating Scion Performance 94 Jan Henk Venema, Francesco Giuffrida, Ivan Paponov, Alfonso Albacete, Francisco Pérez-Alfocea and Ian C. Dodd 4.1. Introduction 94 4.2. Current Knowledge of Ionic and Chemical Signalling Between Rootstock and Scion 95 4.2.1. Ionic signalling 95 4.2.2. Plant hormone signalling 100 4.2.3. Metabolite profile of the xylem sap: xylomics 111 Contents vii 4.2.4. Physical signalling 115 4.2.5. Proteins 116 4.2.6. Small RNAs 117 4.3. Conclusions 117 References 118 5 Physiological and Molecular Mechanisms Underlying Graft Compatibility 132 Ana Pina, Sarah Cookson, Angeles Calatayud, Alessandra Trinchera and Pilar Errea 5.1. Introduction 132 5.2. Anatomical and Physiological Steps During Graft Union Development 133 5.2.1. Graft establishment between compatible and incompatible combinations 133 5.2.2. Translocation between grafted partners 137 5.3. Role of Secondary Metabolites at the Interface in Graft Incompatibility 138 5.4. Cell-to-cell Communication Between Graft Partners 141 5.4.1. Plant growth regulator and graft union formation 141 5.4.2. Cell-to-cell communication at the graft interface 142 5.5. Understanding the Molecular Mechanisms Involved in Graft Union Formation and Compatibility. 143 5.5.1. Genes differentially expressed during graft union formation 143 5.5.2. Genes differentially expressed between compatible and incompatible graft combinations 145 5.6. Methods for Examining Graft Union Development and Compatibility 146 5.6.1. In vitro techniques 146 5.6.2. Histological studies 147 5.6.3. Chlorophyll fluorescence imaging as a diagnostic technique 148 5.7. Conclusions 148 References 149 6 Grafting as Agrotechnology for Reducing Disease Damage 155 Roni Cohen, Aviv Dombrovsky and Frank J. Louws 6.1. Introduction 155 6.2. First Step: Managing Diseases in the Nursery 156 6.2.1. Tobamovirus management: grafted cucurbits and cucumber green mottle mosaic virus: an example of risk and a solution 158 6.2.2. Bacterial canker management: grafted tomatoes and an old nemesis 159 6.3. Disease Spread from the Nursery to the Field, the Example of Powdery Mildew of Watermelons 160 viii Contents 6.4. Intra- and Interspecific Grafting and their Relationship to Diseases 160 6.5. Biotic or Abiotic Stress? Different Responses of Grafted Plants to Environmental Conditions: the Case of ‘Physiological Wilt’ 161 6.6. Response of Grafted Plants to Nematodes 163 6.7. Commercial Rootstocks and Unknown Genetics 164 6.8. Different Mechanisms Involved in Disease Resistance Induced by Grafting 164 6.9. Conclusions 167 References 167 7 Grafting as a Tool for Tolerance of Abiotic Stress 171 Youssef Rouphael, Jan Henk Venema, Menahem Edelstein, Dimitrios Savvas, Giuseppe Colla, Georgia Ntatsi, Meni Ben-Hur, Pradeep Kumar and Dietmar Schwarz 7.1. Introduction 171 7.2. Temperature Stress 172 7.2.1. Diminishing the temperature constraints for vegetable production 172 7.2.2. Contribution of rootstocks to improved low- and high-temperature tolerance 174 7.2.3. Rootstock selection for improved temperature-stress tolerance 178 7.2.4. Cold- and heat-tolerant Cucurbitaceae and Solanaceae rootstocks 179 7.3. Salinity Stress 182 7.4. Nutrient Stress 187 7.4.1. Excessive nutrient availability 187 7.4.2. Deficient nutrient availability 188 7.5. Stress Induced by Metalloids and Heavy Metals 190 7.5.1. Boron 190 7.5.2. Heavy metals 193 7.6. Stress by Adverse Soil pH 197 7.7. Drought and Flood Stresses 199 7.7.1. Drought 199 7.7.2. Flooding and water logging 201 7.8. Conclusions 202 Acknowledgements 203 References 203 8 Quality of Grafted Vegetables 216 Cherubino Leonardi, Marios C. Kyriacou, Carmina Gisbert, Gölgen B. Oztekin, Isabel Mourão and Youssef Rouphael 8.1. What is Quality? 216 8.2. Rootstock Effects on Fruit Quality 217 8.2.1. Appearance 217 8.2.2. Texture 221 Contents ix 8.2.3. Organoleptic compounds and relation to sensory properties 223 8.2.4. Health-promoting substances 227 8.2.5. Contaminants 230 8.3. Effects of Grafting on Ripening and Postharvest Behaviour 231 8.4. Biophysiological Processes Affecting Fruit Quality 232 8.5. Conclusions 235 References 237 9 Practical Applications and Speciality Crops 245 Amnon Koren, Eyal Klein, J. Anja Dieleman, Jan Janse, Youssef Rouphael, Giuseppe Colla and Isabel Mourão 9.1. Establishment of Grafted Transplant under Mediterranean Climate Conditions 245 9.1.1. Factors affecting the establishment of grafted plants 246 9.1.2. Abiotic stress 251 9.1.3. Biotic stress 254 9.2. Recommendations for the Use of Grafted Plants in Greenhouses. The Case of the Netherlands 255 9.2.1. The grafting process 256 9.2.2. Cultivation system of grafted plants 256 9.2.3. Start of cultivation 257 9.2.4. Later phases in cultivation 258 9.3. Role of Grafting in Speciality Crops 258 9.3.1. Globe artichoke 258 9.3.2. Green bean 260 9.4. Conclusions and Future Perspective on Vegetable Grafting 263 Acknowledgements 263 References 263 Index 271 Contributors Alfonso Albacete, Department of Plant Nutrition, Centro de Edafología y Biología Aplicada del Segura (CEBAS), Consejo Superior de Investigaciones Científicas (CSIC), Campus Universitario de Espinardo, Murcia 30100, Spain. E-mail: [email protected] Muhammad Azher Nawaz, College of Horticulture and Forestry Sciences, Huazhong Agricultural University/Key Laboratory of Horticultural Plant Biology, Ministry of Education, Wuhan, PR China; Department of Horticulture, University College of Agriculture, University of Sargodha, Sargodha, Pakistan.