Table Of Content

PrinciplesofPlantGeneticsandBreeding Companion website The book is accompanied by a companion resources site: www.wiley.com/go/acquaah/plantgeneticsandbreeding With figures and tables from the book. Principles of Plant Genetics and Breeding Second Edition George Acquaah Bowie State University, Maryland, USA Thiseditionfirstpublished2012#2012byJohnWiley&Sons,Ltd Wiley-BlackwellisanimprintofJohnWiley&Sons,formedbythemergerofWiley’sglobalScientific,Technicaland MedicalbusinesswithBlackwellPublishing. Registeredoffice: JohnWiley&Sons,Ltd,TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK Editorialoffices: 9600GarsingtonRoad,Oxford,OX42DQ,UK TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 111RiverStreet,Hoboken,NJ07030-5774,USA Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyforpermissionto reusethecopyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell. TherightoftheauthortobeidentifiedastheauthorofthisworkhasbeenassertedinaccordancewiththeUKCopyright, DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inanyformor byanymeans,electronic,mechanical,photocopying,recordingorotherwise,exceptaspermittedbytheUKCopyright, DesignsandPatentsAct1988,withoutthepriorpermissionofthepublisher. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesandproduct namesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheirrespectiveowners.The publisherisnotassociatedwithanyproductorvendormentionedinthisbook.Thispublicationisdesignedtoprovide accurateandauthoritativeinformationinregardtothesubjectmattercovered.Itissoldontheunderstandingthatthe publisherisnotengagedinrenderingprofessionalservices.Ifprofessionaladviceorotherexpertassistanceisrequired,the servicesofacompetentprofessionalshouldbesought. LibraryofCongressCataloging-in-PublicationData Acquaah,George. Principlesofplantgeneticsandbreeding/GeorgeAcquaah.—2nded. p.cm. Includesbibliographicalreferencesandindex. ISBN978-0-470-66476-6(cloth)—ISBN978-0-470-66475-9(pbk.) 1. Plant breeding. 2. Plantgenetics. I. Title. SB123.A3342012 631.5’2—dc23 2012010941 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprintmaynotbeavailablein electronicbooks. CoverdesignbyDanJubb. Setin10/12pt,Galliard-RomanbyThomsonDigital,Noida,India FirstImpression2012 Dedication Tomywife,Theresa,withloveandappreciationforuncommoncharacter. Contents Preface, ix Acknowledgements, xi Industryhighlightsboxes, xiii Industryhighlightsboxes:authors, xv Section1 Overviewandhistoricalperspectives, 1 1 Introduction, 3 2 Historyofplantbreeding, 22 Section2 Populationandquantitativegeneticprinciples, 41 3 Introductiontoconceptsofpopulationgenetics, 43 4 Introductiontoquantitativegenetics, 63 Section3 Reproductivesystems, 95 5 Introductiontoreproductionandautogamy, 97 6 Allogamy, 121 7 Hybridization, 131 8 Clonalpropagationandinvitroculture, 146 Section4 Germplasmforbreeding, 171 9 Variation:types,originandscale, 173 10 Plantdomestication, 185 11 Plantgeneticresources, 199 Section5 Breedingobjectives, 227 12 Yieldandmorphologicaltraits, 229 13 Qualitytraits, 246 14 Breedingforresistancetodiseasesandinsectpests, 260 15 Breedingforresistancetoabioticstresses, 280 Section6 Selectionmethods, 301 16 Breedingself-pollinatedspecies, 303 17 Breedingcross-pollinatedspecies, 337 18 Breedinghybridcultivars, 355 19 Breedingclonallypropagatedspecies, 374 Section7 Molecularbreeding, 383 20 Molecularmarkers, 385 21 Mappingofgenes, 402 22 Markerassistedselection, 424 23 Mutagenesisinplantbreeding, 436 24 Polyploidyinplantbreeding, 452 25 Moleculargeneticmodificationsandgenome-widegenetics, 470 viii CONTENTS Section8 Marketingandsocietalissuesinbreeding, 489 26 Performanceevaluationforcropcultivarrelease, 491 27 Seedcertificationandcommercialseedrelease, 511 28 Regulatoryandlegalissues, 523 29 Value-drivenconceptsandsocialconcerns, 543 30 Internationalplantbreedingefforts, 556 Section9 Breedingselectedcrops, 575 31 Breedingwheat, 577 32 Breedingcorn, 591 33 Breedingrice, 606 34 Breedingsorghum, 617 35 Breedingsoybean, 629 36 Breedingpeanut, 639 37 Breedingpotato, 647 38 Breedingcotton, 657 39 Breedingtomato, 667 40 Breedingcucumber, 679 41 Breedingroses, 682 Supplementarychapters:reviewofgeneticstatisticalprinciples, 689 1 Plantcellularorganizationandgeneticstructure:anoverview, 691 2 Commonstatisticalmethodsinplantbreeding, 707 Glossaryof terms, 726 Appendix1:Conversionrates, 731 Index, 732 Companion website The book is accompanied by a companion resources site: www.wiley.com/go/acquaah/plantgeneticsandbreeding With figures and tables from the book. Preface ThesecondeditionofPrinciplesofPlantGeneticsand on variation and its genetic basis, while Chapter 10 Breeding represents a thoroughly overhauled version focuses on domestication of plant species. The dis- oftheprecedingedition,followingrecommendations cussion includes the dependence of plant breeding and suggestions from users and reviewers. The major on heritable variation. Finally, Chapter 11 addresses changes in the new edition include restructuring and the matter of plant genetic resources used in plant reordering the chapters to follow more closely with breeding. It includes a discussion of how germ- howplantbreedingisdoneinpractice,andexpanding plasm is collected and managed for long term use themoleculargeneticscomponent.Also,thebasicsci- by breeders. enceinformationhasbeenreduced.Twoofthechap- SectionVisdevotedtodiscussing common breed- ters in the first edition have been transferred to the ing objectives pursued by plant breeders. Thediscus- back of the textbook as supplementary material, so it sions include the genetic basis of those traits and the may be referred to by users only as needed. In this implication in their breeding. Chapter 12 focuses on way, students and users who already have a back- breeding for increased yield and improving morpho- ground in genetics will not feel obligated to study logical traits that enhance crop productivity. In the thosechaptersbeforeadvancingtomoreplantbreed- ensuing chapter, 13, breeding for selected quality ingrelatedtopics.Afeatureofthefirsteditionthatis traits is the focus of discussion. Breeding for disease retained and expanded in the second edition is the and pest resistance is a major breeding objective in inclusionofcontributionsonselectedtopicsbyindus- most crops. This is the subject of Chapter 14, while try professionals. The book is copiously illustrated to Chapter15isdevotedtoissuespertainingtobreeding facilitateteachingandlearningofthetopics. for resistance or tolerance to selected abiotic factors, Thebook isorganizedinto nine sections. SectionI suchassalttolerance. isanoverviewandhistoricalperspectiveofplantbreed- The topics of Section VI focus on selection or ing.Chapter1providesanintroductiontothefieldof breedingmethods.Inthissection,breedingmethods plant breeding, describing its importance to society for autogamous species is the subject of Chapter 16, whileChapter2provideshistoricalperspectives,high- while Chapter 17 is devoted to breeding allogamous lightingthecontributionsbyresearcherstoknowledge species. Chapter 18 concerns the selection methods inthefield.ThetwochaptersinSectionIIaredevoted usedforbreedinghybridcultivarswhilethelastchap- to discussing pertinent population and quantitative ter in the section, 19, is devoted to discussing the genetic concepts, to assist the reader in better under- breeding methods used for clonally propagated spe- standingthepracticesofplantbreeders. cies. The discussions in these chapters provide the Section III, reproductive systems, comprises four advantages and disadvantages of each method, and chapters.Chapter5,autogamy,andChapter6,allog- includealternativeapproaches. amy,focusonreproductiveandgenetic issuesasthey Molecular breeding is the subject of Section VII, pertaintoself-pollinatedandcross-pollinatedspecies, which received the most overhaul. The concept of respectively. Chapter 7 is devoted to discussing the markers and various commonly used molecular genetic issues associated with crossing plants to markers in plant breeding are discussed in detail in reorganize the genetic matrix, while Chapter 8 ends Chapter20,includingtheiradvantagesanddisadvan- the section with a discussion ofissuesassociatedwith tages,aswellthecostandeaseofapplicationinbreed- clonalpropagation. ing.Chapter21isdevotedtodiscussingthemapping Section IV, deals with germplasm for breeding. It of genes and the importance of such maps in is impossible to conduct plant breeding without the plant breeding. Marker assisted selection (MAS) as a proper germplasm. Chapter 9 in this section focuses methodoffacilitatingplantbreedingisthesubjectof x PREFACE Chapter 22. Chapter 23 focuses on the use of muta- confronting breeders on the international plant genesisforinducingvariabilityforcropimprovement. breedingscene. The discussions include the types of mutagens com- Thelastsection,IX,isdevotedtodiscussionofthe monly used in crop improvement, and the success of breeding of selected crops. This section includes dis- thisapproachtobreeding.Manyimportant cropspe- cussionsonthegeneticsofselectedcropplants,germ- cies are polyploids. The methods used for improving plasm used, and breeding methods used for their polyploidsarediscussedinChapter24.Thelastchap- improvement. Professional highlights are provided terinthissection,25,addressesthesubjectofmolec- forthesechapters. ular genetic modifications, including the role of Anefforthasbeenmadetoorganizethisbooksuch genetic engineering in plant improvement. Also, in that the sequence of discussion of topics follows this chapter, the contemporary subject of genome- closely the sequence in conducting a plant breeding widegeneticsisintroduced. project. A plant breeding course, at the minimum, is Section VIII is titled marketing and societal issues usually an upper level course at the undergraduate in plant breeding. In Chapter 26, the reader is level.Itisassumedthatastudenttakingaplantbreed- exposed to the process of preparing a cultivar for ing course would have received prior instruction in releasetofarmersforuse.Commercialseedproducers the basic biology, including genetics, botany, and ensure the quality of theirproducts throughthe seed physiology. A review of basic genetic principles is certificationprocess,asdescribedinChapter27.Plant helpful to better understanding the material in breeders protect their products through securing this book and a plant breeding course in general. legal protection, the subject of Chapter 28. The last Sometimes, some of this basic material is reviewed as two chapters, 29 and 30, end the section with dis- appropriate. In addition, some of the underlying cussions on social concerns that arise from the science is presented in the supplementary chapters of applications of biotechnological tools, and issues thebook.