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Genetics Applied to Forestry an Introduction Genetics Applied to Forestry An Introduction Gösta Eriksson Inger Ekberg David Clapham Genetics Applied to Forestry An Introduction Third edition Gösta Eriksson Inger Ekberg David Clapham ISBN 978-91-576-9187-3 1 © 2013 Gösta Eriksson Inger Ekberg David Clapham ISBN 978-91-576-9187-3 Cover photos. Above: A Pinus sylvestris stand in Distribution Central Sweden, photograph Britt Ekberg-Eriks- Department of Plant Biology and Forest VRQ%HORZ$SURIXVHO\ÀRZHULQJTilia cordata Genetics, SLU, Box 7080, 750 07 Uppsala, Sweden tree in Central Sweden, photograph Inger Ekberg. Contact: [email protected] Printing: Elanders Sverige AB 2 Preface This book is a follow-up of An Introduction to Forest Genetics. It is somewhat expanded compared to the book printed in 2007. We were encouraged to ”publish” the revised version of the textbook on the internet. Undergraduate students are the target group as well as graduate students with limited experience of forest genetics. Without the advice and help from Kjell Lännerholm, Björn Nicander, Johan Samuelsson and Hartmut Weichelt the editing would have been more troublesome. We express our sincere thanks to them. A generous grant from Föreningen Skogsträdsförädling, The Tree Breeding Association in Sweden, made this printing grant from Föreningen Skogsträdsförädling, The Tree Breeding possible. A web version of this book may be found under http://vaxt2.vbsg.slu.se/forgen/Forestry_Genetics.pdf Uppsala December 2013 Gösta Eriksson Inger Ekberg David Clapham 3 Content Chapter 1 Chromosome cytology ................................7 Chapter 4 Population genetics Hardy-Weinberg Karyotype....................................................................7 law................................................................................51 Locus, genes, alleles, homozygosity, heterozy- F statistics................................................................. 53 gosity minant and recessive traits......................9 Summary ..................................................................54 Mitosis.............................................................9 Further reading...........................................................54 Meiosis.....................................................................10 Chromosome aberrations..........................................12 Chapter 5 Quantitative genetics....................................55 Development of egg cells and sperm cells.............12 Characteristics of quantitative traits..........................55 Time of meiosis.........................................................14 Quantitative trait locus QTL......................................56 Injuries and irregularities during meiosis.................16 Methods for constructing genetic linkage maps Summary...................................................................16 for QTL...............................................................56 Further reading.........................................................17 Results from detection and mapping of QTL......58 Heritability................................................................59 Chapter 2 Genes, DNA, RNA, molecular evolution, Genotype x environment interaction.........................63 genetic engineering.......................................................19 Inbreeding and heterosis............................................64 DNA structure...........................................................19 Selection differential, selection intensity, and genetic DNA replication........................................................19 gain...........................................................................66 Mutations - changes in DNA....................................20 Genetic correlation....................................................67 :KHUHWR¿QG'1$" Summary ..................................................................68 Where is DNA located in the nucleus of the cell Further reading..........................................................68 D Q G K R Z L V ' 1 $ R U J D Q L ] H G " :KDWLVDJHQH" Chapter 6 Evolution.....................................................69 Conservation of non-genic DNA.............................25 Terminology..............................................................70 The genetic code......................................................26 )DFWRUVLQÀXHQFLQJHYROXWLRQ Regulation of gene activity.......................................28 Natural selection........................................................71 Number of functional genes in plants......................28 The three main types of natural selection...........72 Similar gene and gene order over wide taxonomic Natural selection under severe stress conditions74 families.....................................................................28 Random genetic drift.................................................75 The molecular clock.................................................29 Mutations .................................................................75 Chloroplasts and mitochondria have their own *HQH ÀRZ g e n e t i c s y s t e m s r e s e m b l i n g t h o s e o f b a c t e r i a . 3 0 Phenotypic plasticity..................................................79 The endosymbiotic hypothesis explains the origin :LOOWKHDGDSWHGQHVVHYHUEHSHUIHFW" of organelles..............................................................30 Ecotype and ecocline.................................................80 Interplay between the cell nucleus and the Evolution and global warming..................................83 organelles................................................................30 Coevolution...............................................................85 Genetic linkage maps...............................................30 Speciation.................................................................86 Genetic engineering ................................................31 Allopatric and sympatric speciation....................87 %ULHÀ\ZKDWGRWKHVHPHWKRGVPHDQ" Adaptive landscapes.............................................87 How can genetic engineering be applied to forest Speciation by polyploidy......................................88 WUHHV" The speed of speciation........................................88 Which traits are most amenable to genetic Summary...................................................................88 HQJLQHHULQJ" Further reading..........................................................89 Summary..................................................................42 Further reading.........................................................44 Chapter 7 Genetic variation and provenance research........................................................................91 Chapter 3 Qualitative inheritance..............................45 Genetic structure and how it is estimated..................91 Genetic variation and non-genetic variation............45 Comparison of markers and quantitative traits..........94 Mendelian inheritance..............................................45 Variation among populations in metric traits............96 Gene effects at the biochemical level.......................48 Pinus sylvestris and Picea abies provenance re- Summary..................................................................49 search........................................................................96 Further reading.........................................................49 Provenance research in some other conifers.............102 4 Provenance research in some broadleaved tree Early tests................................................................151 species................................... ..................................106 Progress in breeding................................................154 Adaptation to edaphic conditions............................108 The sustainability of the gain..................................158 Utilization of provenance results.............................109 Summary.................................................................159 Markers...................................................................110 Further reading........................................................159 'DUZLQLDQDQGGRPHVWLF¿WQHVV Postscript..................................................................160 Summary .................................................................115 Further reading........................................................162 Further reading.........................................................116 Chapter 10 Plant production....................................163 Chapter 8 Variation within populations..................117 Summary ................................................................165 Examples of variation among families for various Further reading........................................................165 traits........................................................................117 ,QWHUVSHFL¿FK\EULGV Chapter 11 Forest tree gene conservation...............167 +HULWDELOLWLHVDQGFRHI¿FLHQWVRIDGGLWLYH The three cornerstones of gene conservation..........168 variation..................................................................121 Objectives in gene conservation..............................168 Genetic correlations................................................126 Prime objective...................................................168 Why is there such a large within-population Other
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