HARTMANN AND KESTER’S PLANT PROPAGATION

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A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage iFront Matter MUSTcomprise 16 pages.suggest settinghighlighted text sameas book subtitle hereand on title page. Pls.advise.HARTMANN ANDKESTER’S PLANTPROPAGATIONPRINCIPLES AND PRACTICES

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A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage iiiHARTMANN ANDKESTER’S PLANTPROPAGATIONPRINCIPLES AND PRACTICESEIGHTH EDITIONHudson T. Hartmann, PhDUniversity of California, DavisDale E. Kester, PhDUniversity of California, DavisFred T. Davies, Jr., PhDTexas A&M University College StationRobert L. Geneve, PhDUniversity of Kentucky, LexingtonPrentice HallBoston Columbus Indianapolis New York San Francisco Upper Saddle RiverAmsterdam Cape Town Dubai London Madrid Milan Munich Paris Montreal TorontoDelhi Mexico City São Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage ivEditorial Director: Vernon AnthonyAcquisitions Editor: William LawrensenEditorial Assistant: Lara DimmickDirector of Marketing: David GesellSenior Marketing Coordinator: Alicia WozniakMarketing Assistant: Les RobertsAssociate Managing Editor: Alexandrina Benedicto WolfInhouse Production Liaison: Alicia RitcheyOperations Specialist: Laura WeaverArt Director: Diane ErnsbergerCover Designer: Anne DemarinisCover Art: Robert L. GeneveLead Media Project Manager: Karen BretzFull-Service Project Management: Douglas Bell/PreMediaGlobalComposition: PreMediaGlobalPrinter/Binder: Courier/KendallvilleCover Printer: Lehigh-Phoenix ColorText Font:Credits and acknowledgments borrowed from other sources and reproduced, with permission, in this textbook appear onappropriate page within text (or on page xx).Copyright 2011, 2002, 1997, 1990, 1983 Pearson Education, Inc., Publishing as Prentice Hall, One Lake Street, UpperSaddle River, NJ 07458. All rights reserved. Manufactured in the United States of America. This publication is protected bycopyright, and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, ortransmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permission(s)to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, Prentice Hall,One Lake Street, Upper Saddle River, NJ 07458.Many of the designations by manufacturers and seller to distinguish their products are claimed as trademarks. Where thosedesignations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initialcaps or all caps.Library of Congress Cataloging-in-Publication DataDavies, Fred T., 1949Hartmann and Kester’s plant propagation : principles and practice / Fred T. Davies, Robert L. Geneve, Dale E. Kester.—8th ed.p. cm.Revision of: 7th ed. 2002.Includes bibliographical references and index.ISBN 978-0-13-501449-3 (alk. paper)1. Plant propagation. I. Geneve, R. L. II. Kester, Dale E. III. Hartmann, Hudson Thomas, 1914IV. Title. V. Title: Plant propagation.SB119.P55 2011631.5’3—dc22201001251810 9 8 7 6 5 4 3 2 1Paper boundLoose leafISBN 10:ISBN 13:ISBN 10:ISBN 0-13-505441-3

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage vPUB/AUS: Pls advise on how to fit dedication one one page.Reduce size of photo and wrap text an option, but will still not run back overmatter.Please advise.Dedications7 July 2009The eighth edition of Plant Propagation is dedicated toDr. Dale Emmert Kester, Professor Emeritus for theUniversity of California, Davis. Dale passed away onNovember 21, 2003.His lifelong interest in horticulture led Dale toenroll as a horticulture student at Iowa State Universityin Ames, Iowa in 1941. His college career was interrupted in 1943 when Dale joined the war effort as a USAir Force P-51 Mustang pilot. As a World War II pilot,he escorted bombers on 28 missions over Italy andCentral Europe. Dale met his future wife, DaphneDougherty, while he was stationed in Baton Rouge,Louisiana. Daphne was a USO dancer at the time.Following the war, he returned to Iowa State Universityand completed his horticulture degree in 1947.Dale was the first PhD graduate from theUniversity of California, Davis Pomology Departmentfollowing the war. His dissertation concerned embryoculture of peaches. In 1951, he was offered an AssistantProfessor position in the Department of Pomology atUC Davis where his work was to focus on almond production and breeding. This was the position he wouldhold until his retirement 40 years later in 1991. Hetaught undergraduate plant propagation and pomologycourses. Early in his career, he partnered with Dr. HudsonHartmann to publish the first edition of “PlantPropagation—Principles and Practices” in 1959.Along with Hudson Hartmann and others, Dalewas a founding member of the Western Region of theInternational Plant Propagators’ Society. He served thatorganization as Vice-President, program chair in 1996and President in 1997. Dale received the Curtis J. AlleyAward in 1999 for his lifetime service to the InternationalPlant Propagators’ Society. In 2002, shortly before hisdeath, he received the society’s highest award, theInternational Award of Honor. With this award, he wasrecognized for “his long-standing reputation as a dedicated teacher of students interested in plant propagation, his service to the International Plant Propagators’Society and especially, for his seminal textbook onplant propagation used the world over.”Dale was a longtime member of the AmericanSociety for Horticultural Science and was recognized asa Fellow in 1977. He served as the first chair of thePropagation Working Group and received the StarkAward in 1980. In 1998, he was the Spenser AmbroseBeach Lecturer at Iowa State University. He publishedover 120 research papers in journals and conferenceproceedings. His research efforts in almond led tonumerous root stock introductions, as well as the causefor noninfectious bud failure in almond.Dale Kester was one of the most internationallyrecognized horticulturists of his generation, but remaineda very unpretentious man. He was easy-going, goodhumored and appeared more impressed with his colleagues’ achievements than his own. Dale was a mentor,role model, and a friend. He will be greatly missed bythe horticultural community.The seventh edition of Plant Propagation wasdedicated to Dr. Hudson T. Hartmann. Dr. Hartmanndied March 2, 1994 just as plans for the sixth editionwere getting underway. He is remembered as a dedicated, hard-working, conscientious scientist, teacher,and human being. He conceived of the writing of thistext about 1955 and asked the second author, Dr. DaleE. Kester, to join him. Dr. Hartmann taught PlantPropagation at the University of California at Davisfrom 1945 to his retirement in 1980. His research inpropagation involved early studies on hormones, mistpropagation, and other aspects of cutting propagation

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage viOVERMATTERparticularly as they applied to fruit trees. He was also aspecialist in olive research and development, attaining aworldwide reputation for this crop.One of his primary accomplishments was hisactivity with the International Plant PropagationSociety. He became a member in 1953 and then wasinstrumental in initiating the Western Region of theSociety in 1960. He served as Western Region Editorfor the Society from 1960 to 1993, serving also asInternational Editor from 1970 until 1991. During hiscareer he published many scientific papers and populararticles. As well as the present text, he was senior authorof Plant Science: Growth, Development and Utilizationof Cultivated Plants, first edition (1981), second edition(1988) published by Prentice Hall.Dr. Hartmann was a member of the AmericanSociety for Horticultural Science, becoming a Fellow in1974. As an undergraduate he was a member ofGamma Sigma Delta and Alpha Zeta. He receivedmany awards, including the Charles G. WoodburyAward (1960), Joseph H. Gourley Award (1962), andStark Brothers Award (1964) from ASHS. TheAmerican Association of Nurserymen awarded him itsNorman J. Coleman Award (1970), The CaliforniaAssociation of Nurserymen presented him with itsResearch award (1977), and Pi Alpha Xi made him anhonorary member (1981). The Western Region IPPSawarded Dr. Hartmann its Merit award (1979),Honorary Membership (1983), and established theHudson T. Hartmann Western Region Research Grantin his honor. The International IPPS Board ofDirectors awarded him the International Award ofHonor in 1990.Dr. Hartmann was a close personal friend, a collaborator who made working together a pleasure, and arespected peer whose guidance and insight are missed.

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage viibrief contentsPreface xiiAbout the Authors xivAcknowledgements xvpart oneGeneral Aspects of Propagation123How Plant Propagation Evolved in Human SocietyBiology of Plant Propagation 14The Propagation Environment 492part twoSeed Propagation45678Seed Development 110Principles and Practices of Seed Selection 140Techniques of Seed Production and Handling 162Principles of Propagation from Seeds 200Techniques of Propagation by Seed 250part threeVegetative Propagation910111213141516Principles of Propagation by Cuttings 280Techniques of Propagation by Cuttings 344Principles of Grafting and Budding 415Techniques of Grafting 464Techniques of Budding 512Layering and Its Natural Modifications 537Propagation by Specialized Stems and Roots 561Principles and Practices of Clonal Selection 594part fourCell and Tissue Culture Propagation1718Principles of Tissue Culture and MicropropagationTechniques for Micropropagation 699644part fivePropagation of Selected Plant Species192021Propagation Methods and Rootstocks for Fruit and Nut Species 728Propagation of Ornamental Trees, Shrubs, and Woody Vines 774Propagation of Selected Annuals and Herbaceous Perennials Usedas Ornamentals 840Appendix 000Glossary 000Index 000vii

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage viiicontentsPreface xiiAbout the Authors xivAcknowledgements xvpart oneEnvironmental Factors Affecting Propagation 50General Aspects of Propagation 1Physical Structures for Managing the PropagationEnvironment 541How Plant Propagation Evolved inHuman Society 2Introduction 2Learning Objectives 2Stages of Agricultural Development 3Organization of Human Societies 4Exploration, Science, and Learning 5The Development of Nurseries 8The Modern Plant Propagation Industry 12Discussion Items 12References 122Biology of Plant Propagation 14Introduction 14Learning Objectives 14Biological Life Cycles in Plants 14Taxonomy 18Legal Protection of Cultivars 21Genetic Basis for Plant Propagation 21Genetic Inheritance 27Gene Structure and Activity 30Plant Hormones and Plant Development 38Discussion Items 45References 453The Propagation Environment 49Introduction 49Learning Objectives 49viiiContainers for Propagating and Growing YoungLiner Plants 70Management of Media and Nutrition in Propagationand Liner Production 77Management of Microclimatic Conditions inPropagation and Liner Production 85Biotic Factors—Pathogen and Pest Management inPlant Propagation 90Post-Propagation Care of Liners 100Discussion Items 102References 103part twoSeed Propagation 1094Seed Development 110Introduction 110Learning Objectives 110Reproductive Life Cycles of Vascular Plants 110Characteristics of a Seed 112Reproductive Parts of the Flower 117Relationship Between Flower and SeedParts 118Stages of Seed Development 122Unusual Types of Seed Development 130Plant Hormones and Seed Development 133Ripening and Dissemination 136Discussion Items 137References 137

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage ixcontents5Principles and Practicesof Seed Selection 140ixLearning Objectives 250Seedling Production Systems 250Discussion Items 276References 276Introduction 140Learning Objectives 140part threeBreeding Systems 140Vegetative Propagation 279Categories of Seed-Propagated Cultivars andSpecies 147Control of Genetic Variability DuringSeed Production 150Seed Production Systems 153Discussion Items 159References 1596Techniques of Seed Productionand Handling 162Introduction 162Learning Objectives 162Sources For Seeds 162Harvesting and Processing Seeds 166Seed Testing 175Seed Treatments to Improve Germination 184Seed Storage 189Discussion Items 195References 1957Principles of Propagation from Seeds 200Introduction 200Learning Objectives 200The Germination Process 200Dormancy: Regulation of Germination 218Kinds of Primary Seed Dormancy 220Secondary Dormancy 235Dormancy Control by Plant Hormones 236Discussion Items 240References 2408Techniques of Propagation by Seed 250Introduction 2509Principles of Propagation by Cuttings 280Introduction 280Learning Objectives 280Descriptive Observations of Adventitious Rootand Bud (and Shoot) Formation 281Correlative Effects: How Hormonal ControlAffects Adventitious Root and Bud (and Shoot)Formation 293The Biochemical Basis for Adventitious RootFormation 299Molecular/Biotechnological Advances in AsexualPropagation 304Management and Manipulation of Adventitious Rootand Shoot Formation 305Management of Stock Plants to Maximize CuttingPropagation 307Treatment of Cuttings 318Environmental Manipulation of Cuttings 323Discussion Items 331References 33210Techniques of Propagation byCuttings 344Introduction 344Learning Objectives 344Types of Cuttings 344Sources of Cutting Material 363Rooting Media 367Wounding 373Treating Cuttings with Auxins 373Preventative Disease Control 381Environmental Conditions for RootingLeafy Cuttings 383

A01 DAVI4493 08 SE FM.qxdx8/14/101:22 AMPage xcontentsPreparing the Propagation Bed, Bench, Rooting Flats,and Containers, and Inserting the Cuttings 393Preventing Operation Problems with Mist and FogPropagation Systems 395Management Practices 396Care of Cuttings During Rooting 401Hardening-Off and Post-Propagation Care 403Handling Field-Propagated Plants 406Container-Grown Plants and Alternative FieldProduction Systems 409Discussion Items 409References 4091113Techniques of Budding 512Introduction 512Learning Objectives 512Importance and Utilization of Budding 512Rootstocks for Budding 513Time of Budding—Summer, Spring, or June 513Types of Budding 519Top-Budding (Topworking) 532Double-Working by Budding 533Microbudding 534Discussion Items 535References 536Principles of Grafting and Budding 415Introduction 415Learning Objectives 41514Terminology 417Layering and Its NaturalModifications 537Seedling and Clonal Rootstock Systems 419Introduction 537Reasons for Grafting and Budding 419Learning Objectives 537Natural Grafting 424Reasons for Layering Success 537Formation of the Graft Union 425Management of Plants During Layering 539Graft Union Formation in T- and Chip Budding 432Procedures in Layering 539Factors Influencing Graft Union Success 433Plant Modifications Resulting inNatural Layering 551Discussion Items 558References 558The History of Grafting 415Genetic Limits of Grafting 439Graft Incompatibility 441Scion-Rootstock (Shoot-Root) Relationships 450Discussion Items 457References 4571215Propagation by Specialized Stemsand Roots 561Techniques of Grafting 464Introduction 561Introduction 464Learning Objectives 561Learning Objectives 464Bulbs 563Requirements for Successful Grafting 464Corms 577Types of Grafts 465Tubers 579Production Processes of Graftage 491Tuberous Roots and Stems 581Aftercare of Grafted Plants 502Rhizomes 584Field, Bench, and MiscellaneousGrafting Systems 504Discussion Items 509References 509Pseudobulbs 587Discussion Items 590References 590

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage xicontents16Uses for Micropropagation 699Disadvantages of Micropropagation 701Principles and Practices of ClonalSelection 594General Laboratory Facilities and Procedures 702Introduction 594Stage I—Establishment 713Learning Objectives 594Stage II—Shoot Multiplication 716History 594Stage III—Root Formation 717Using Clones as Cultivars 595Stage IV—Acclimatization to GreenhouseConditions 718Discussion Items 724References 724Origin of Clones as Cultivars 597Phenotypic Variations Within Clones 601Patterns of Genetic Chimeras Within Clones 603Management of Phase Variation During VegetativePropagation 613Pathogens and Plant Propagation 619Selection and Management ofPropagation Sources 623Propagation Sources and Their Management 630Discussion Items 635References 636part fourCell and Tissue Culture Propagation 64317Principles of Tissue Cultureand Micropropagation 644Introduction 644Learning Objectives 644A Brief History of Tissue Cultureand Micropropagation 644Micropropagation Procedures 712part fivePropagation of Selected PlantSpecies 72719Propagation Methods and Rootstocksfor Fruit and Nut Species 728Introduction 728References 76620Propagation of Ornamental Trees, Shrubs,and Woody Vines 774Introduction 774References 82521Control of the Tissue Culture Environment(119, 209, 229) 679Propagation of Selected Annualsand Herbaceous Perennials Used asOrnamentals 840Special Problems Encountered by In VitroCulture 681Introduction 840References 869Variation in Micropropagated Plants 684Discussion Items 687References 687Subject Index 000Plant Index, Scientific Names 000Plant Index, Common Names 000Types of Tissue Culture Systems 64918Techniques for Micropropagation 699Introduction 699Learning Objectives 699xi

A01 DAVI4493 08 SE FM.qxd8/14/101:22 AMPage xiiPrefaceThe eighth edition of Plant Propagation: Principles and Practices continues the legacyof updating the ever-changing principles and practices associated with plant propagation, but it is also the first edition with expanded color figures throughout the text.This is an exciting prospect that the co-authors hope will enhance student learning.Some 90% or more of the images and illustrations are either new or enhanced.The eighth edition is published a half-century after the initial printing of PlantPropagation: Principles and Practices in 1959, but still continues the tradition of presenting paired chapters where the principles underlying the science of propagationalternate with the technical practices and skills utilized for commercial plant propagation. As with previous editions, the amount of material between editions hasincreased at an incredible rate and many aspects of growth and development haveexpanded beyond the wildest forecasts in 1959. We have tried to integrate the mostcurrent commercial techniques and understanding of the biology of propagation intocurrent chapters. We have substantially updated the references and sections on“Getting More in Depth on the Subject” to help the reader delve deeper into thesesubjects than the general scope of this textbook.As in previous editions, the book is organized into four basic parts. The initialthree chapters are general chapters meant to support general aspects of propagationincluding a historical perspective, basic plant biology concepts and the impact andcontrol of the environment as it affects propagation and nursery practices. Chapter 2has been significantly revised to reflect the significant progress in plant hormone biology and the molecular advances in plant growth and development. We hope that itserves as background support for understanding the concepts described in thePrinciples chapters, and provides a foundation for students to pursue these fascinatingsubjects in the literature. Chapter 3 continues the integration of concepts and application to control the propagation environment, which is of major importance in commercial propagation. The latest engineering, computerization, and mechanizationsystems for propagation are included. The next two sections describe seed and vegetative propagation, respectively. Each revised section provides a chapter on the conceptsbehind genetic selection for either sexual or clonal plants, and then specific chaptersfor the principles and practices. The final section is an upda

1 How Plant Propagation Evolved in Human Society 2 2 Biology of Plant Propagation 14 3 The Propagation Environment 49. part two. Seed Propagation. 4 Seed Development 110 5 Principles and Practices of Seed Selection 140 6 Techniques of Seed Production and Handling 162 7 Principles of Propagati

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