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Conservation and Management of Forest Genetic Resources in Europe Thomas Geburek & Jozef Turok / editors Publishers

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Conservation and Management of ForestGenetic Resources in Europe

Thomas Geburek & Jozef Turok / editors

Publishers

Table of contents

Preface iTable of contents iiiList of contributors xixList of reviewers xxiii

Chapter 1: Introduction 1

Conservation and sustainable management of forest genetic resources inEurope - an introduction (Th. Geburek & }. Turok) 3

Introduction 3Why should we care? 4What is a forest genetic resource? 5What is forest genetic conservation? 6Outlook 8References 8

Chapter 2: Forest biodiversity - the policy, legal and economicaspects 11

Biodiversity in national forest and environmental policy (M. Pregernig) .. 13Introduction 13Definitions of biodiversity 14Biodiversity as a public good 14(Economic) values of biodiversity 16Policy instruments for realising privately appropriable values 16

Assigning well-defined property rights 16Certification and eco-labelling schemes 19

Policy instruments for realising public values 19Regulatory instruments 19Financial instruments 20

Integrated approaches to biodiversity policy 21

Th. Geburek &]. Turok (eds.)Conservation and Management of Forest Genetic Resources in EuropeArbora Publishers, Zvolen, 2005, iii-xvii, ISBN 80-967088-1-3 iii

'Markets plus': applying a mix of policy instruments 21Sectoral integration 22Integration across hierarchies: polycentric governance regimes 23Deliberative and participatory processes 24

International and European policy initiatives for ensuring biological diversityin forests ."...".' 25

International level 25Pan-European level 27European Union level 28

Conclusions 29References 29

Look at Pan-European Forest Policy: the Ministerial Conference on theProtection of Forests in Europe and the ' Living Forest Summit'(P. Mayer & A. Buck) 33

Introduction 33Characteristics of the MCPFE 33History of the MCPFE 35

Strasbourg 1990 - Initiating cross-border mechanisms for the protection offorests in Europe 35Helsinki 1993 - A commitment to sustainable forest management in Europe .. 36Lisbon 1998 - Recognising the multiple roles of forests 37Implementing the Ministerial Decisions - The MCPFE Work Programme 38

The Living Forest Summit in 2003 39Vienna Living Forest Summit Declaration 39Five Vienna Resolutions 40New Work Programme 41

Conclusions 42Internet Source for further information 42References : 42

Forest genetic resources in the international context: processes, agree-ments and programmes (Ch. Palmberg-Lerche, ]. Turok & P. Sigaud) 45

Introduction 45Global processes and agreements 46

Global agreements resulting from UNCED 46The Forest Principles 46The Convention on Biological Diversity 46The United Nations Framework Convention on Climate Change 48The United Nations Convention to Combat Desertification 48

The international forest policy dialogue 49The Inter-Governmental Panel, the Inter-Governmental Forum and theUnited Nations Forum on Forests 49Criteria and Indicators for Sustainable Forest Management 49

IV

The Ministerial Conferences on the Protection of Forests in Europe and thePan-European Biological and Landscape Diversity Strategy 50

International plant genetic resources policies 51The International Undertaking and the International Treaty on Plant Ge-netic Resources for Food and Agriculture 51The Global Plan of Action for the Conservation and Use of Plant GeneticResources for Food and Agriculture 52

Brief review of relevant programmes of some international organizations andinstitutions 53The Food and Agriculture Organization of the United Nations 54

Mandate 54FAO's Forest Genetic Resources Programme 54The Panel of Experts on Forest Gene Resources 54

The United Nations Educational, Scientific and Cultural Organization 55Mandate 55The Man and the Biosphere Programme 56Biosphere Reserves 56

The United Nations Environment Programme 56Mandate 56UNEP and the International Environmental Conventions 57The UNEP World Conservation Monitoring Centre 57

The United Nations Development Programme 57The International Tropical Timber Organization 57The World Bank 58The Global Environment Facility 58The Consultative Group on International Agricultural Research 58

The Centre for International Forestry Research 59The World Agroforestry Center 60The International Plant Genetic Resources Institute 60

The International Union of Forestry Research Organizations 61The World Conservation Union and the World Wide Fund for Nature 62

Regional forest genetic resources programmes, workshops and networks 62European Forest Genetic Resources Programme . 63

Background 63Management and organization 63Long-term conservation strategies 64Technical guidelines 65Information exchange and information standards 65Publications 65Exchange of genetic materials 66Strengths of regional networking collaboration 66

Regional networks 67National policies and programmes 68Conclusions 69References 70

International legislation with implications on the exchange of forest ge-netic resources (L. Ackzell & J. Turok) 75

Introduction 75International trade with forest reproductive material 76

Schemes on the transfer of forest reproductive material in EU and OECD 76Influences of market regulations 77Definitions and categories 77National authority tasks 78Differences between EC and OECD schemes 79Genetically modified FRM 79Conservation of forest genetic resources 80

International access and benefit sharing 80Bilateral transactions under the Convention on Biological Diversity 80Multilateral system for plant genetic resources for food and agriculture 81Material transfer agreements -. 82

National legislation 83European Union legislation 84Conclusions 85References 85

Economic approaches for valuing forest genetic resources (P. Elsasser) 89Introduction: Resource allocation as a decision problem 89Some theoretical background of environmental valuation 91What type of object might a valuation address? 93Motivations for protecting genetic diversity as a base for valuation 94Methods for analysing economic values of forest genetic resources 96

Market prices for genetic resources 96Indirect methods to analyse revealed preferences ."' 97Direct methods to analyse stated preferences in simulated markets 98Bias sources and reliability of stated preferences .. ; 99Information problems 101Lexicographic preferences 101Embedding effects 102

Some extensions of the CVM approach 103Integrating attitude - behaviour research 103Deliberative and participatory approaches 104

Conclusions 105References 106

Forest resources in Europe and their condition (M. Lorenz, R. Fischer& V. Mues) I l l

Introduction I l lForest area and its changes in Europe 112Status and development of forest condition in Europe 113

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Forest damage and its causes 113The monitoring system of UNECE and EU 115

Monitoring results 116Spatial and temporal variation of defoliation 116

Spatial variation of defoliation 116Temporal variation of defoliation 118

Deposition and critical load excess of nitrogen and acidity 123Nitrogen 123Acidity 124

Conclusions 124References 125

Chapter 3: Genetics in forest tree populations 127

Phenotypic and genetic variation (H. H. Hattemer) 129Introduction 129Phenotypic variation 129

Morphological traits 131Physiological traits 132

Genetically controlled phenotypic variation 134What are genes? 134Mendelian laws of inheritance and the concept of genetic markers 134Estimation of phenotypic components 136How to quantify genetic control? 141

Significance of genetic variation for adaptive traits 142Conclusions 143Acknowledgement 144References 144

Genetic structures and population sizes (H. H. Hattemer) 149Introduction 149What is a population? 149Panmictic equilibrium in a population 150

One single gene locus 151Several gene loci 154

Selection equilibrium in a population 154Impact of the reproduction system on genetic structures 156Inbreeding 159

Co-ancestry in regular pedigrees 159Mating system in an effectively large population 160Combined effect of genetic drift and inbreeding in a population 161

Population sizes 162Effective population size 162Concept of the metapopulation 165

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Types of rarity in plants 166Conclusions 167Acknowledgement ; 168References 168

Sexual reproduction in forest trees (Th. Geburek) 171Introduction 171Reproductive cycle 172Pollen and egg development 172Asexual embryogenesis 173Sexual systems 173Sex function and structure 176Incompatability 177

Prezygotic mechanisms 177Postzygotic mechanisms 179

Pollination and pollen movement 181Flowering 186Mating 187Natural seed dispersal (migration) 188

Abundance and frequency of natural seeding 190Natural seed ripening and seed fall 191

Conclusions 192References 193

Evolution and evolutionary factors, adaptation, adaptability (G. Eriksson) 199Introduction - population concepts 199Evolutionary factors 199

Sources of genetic and genotypic variation: mutation and recombination . . . . 200The role of natural selection 201Non-directed (spontaneous) processes: random drift, gene flow 202Phenotypic plasticity 203Synthesis - 204

What are adaptation, adaptability and adaptedness? 205Concepts 205How do we safeguard the potential for adaptation? . .'.• 207How do we estimate the potential for adaptation? 208

Conclusions 209Acknowledgement 210References 210

Quantitative genetics (G. Jansson) 213Introduction 213From a single-locus trait to polygenic inheritance 213

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Average effect of a gene and breeding value 215Phenotypic and genotypic values 216

Partitioning of variance 217Coefficient of additive and genotypic variation 217

Estimation of variance components 219Heritability 221General and specific combining ability 223Genetic and genotypic gain 225Genetic correlation 228Genotype by environment interaction 231Conclusions 232References 233

Measuring genetic variation within and among populations at markerloci (E. Gillet, D. Gomory & L. Paule) 237

Introduction 237From phenotype to marker locus - a review of terminology 239The allelic complement of deme 241

Sources of new alleles 242Mutation 242Gene flow 242

Loss of alleles 242Genetic drift 242Selection 243

Measuring the number of alleles in a deme 243Allelic multiplicity 243Detecting alleles 244Allelic richness 245

Genotypic and allelic structures 246Frequency distributions .; 246Causes of change in allelic and genotypic structures 248

Mutation 248Genetic drift 249Gene flow 249

Natural selection 250Mating system 252Colonization 253Artificial selection 253

Measuring genetic variation within demes 255Diversity as an effective number of types 256Diversity as the heterogeneity of demes 257Heterozygosity as genetic variation within individuals 258

Measuring genetic distance between two demes 260Genetic distance between demes 260Genetic distance between individuals 263

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Graphical representation of distance matrices 264Measuring genetic differences among several demes 264

Hierarchical analysis of allelic variation among demes 264Subpopulation differentiation 266Differentiation among individuals 269

Conclusions 270Acknowledgements 270References 270

Provenance research: evaluating the spatial pattern of genetic variation(A. O. Konig) 275

Introduction 275Objectives of provenance research 277Brief overview of genecological studies 279Some examples of provenance research 280

Coniferous species 281Pinus sylvestris, Scots pine 281

Results from the IUFRO and Russian experiments 282Swedish experiments 283North American experiments 284

Picea abies, Norway spruce 284Results 286

Pseudotsuga menziesii, Douglas fir 290Results :..... 291

Broad-leaved species 293Fagus sylvatica, European beech 293

Results 294Selected other species 295

Factors determining and shaping genetic variation 297Glacial and postglacial history ; ; 297Environmental factors 298Genetic variation and fitness of provenances 302The problem of discontinuous variation 304

Provenance research and genetic conservation 304Planning, establishment and evaluation of provenance tests 306

Seed collections 307Nursery experiments and the establishment of field trials 307Evaluation 310

Some additional aspects and problems 313Sampling 313Nursery versus field experiment results 315Problems of design and competition 317Establishment and silvicultural treatment 317Joint evaluations 318Classification of intraspecific variation 319

What use can be made of the results of provenance research experiments? 320Conclusions 323Acknowledgement 325References 325

Chapter 4: Genetic techniques and their application in conserv-ation and management of forest genetic resources 335

Molecular markers for characterizing diversity in forest trees (G. G.Vendramin & O. H. Hansen) 337

Introduction 337Genetic markers: characteristics, advantages and disadvantages 338

Isoenzymes 340RFLPs (Restriction Fragment Length Polymorphism) 341VNTRs (Variable Number of Tandem Repeats) - Minisatellites 344Microsatellites 347RAPD Markers 351AFLPs (Amplified Restriction Fragment Polymorphism) 353SSCPs (Single Stranded Conformation Polymorphism), SNPs (Single Nucleo-tide Polymorphism) 355

Choice of marker 358Conclusions ; !. 360References 363

Forest genomics and new molecular genetic approaches to measuringand conserving adaptive genetic diversity in forest trees (K. V.Krutovsky & D. B. Neale) 369

Introduction •. 369Traditional methods to measure adaptive genetic diversity 369

Field experiments 369Molecular genetic markers 370How forest genetic conservation can benefit from new achievements in genomics 373

Introduction to genomics 373Structural genomics 373Functional genomics 376Comparative genomics 376Associative genomics 376Statistical genomics 378DNA sequencing of entire genomes : . . . . 379Gene discovery and expressed sequence tag polymorphisms (ESTPs) 381Physical and genetic mapping of the whole genome using numerous geneticmarkers 381Analysis of genetic control of complex adaptive traits via quantitative trait loci(QTL) mapping 383

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Candidate gene mapping of adaptive genes 384Comparative mapping of adaptive genes 385

Bioinformatics and genomic databases 386Conclusions 387Acknowledgements 388References 388

Selection of target species and sampling for genetic resources in absenceof genetic knowledge (G. Eriksson) 391

Introduction 391Target species 391

Grouping of species 393Capturing adaptedness 393

Evolutionary factors 394Random mating populations 395Selective environmental neighbourhoods 396Gene flow among different selective environmental neighbourhoods 397Phenotypic plasticity 398

Differentiation in species with large random mating populations 399Species with assumed large random mating populations 401Species with non-random mating populations 405Species with assumed non-random mating populations 406Conclusions 407Acknowledgement 408References 408

On the appropriate size of forest genetic resources (H. H. Hattemer) 413Introduction 413Establishment of a resource to be conserved ex situ 413

The risk of loss of alleles at a single gene locus 414Loss of alleles at one of several gene loci 417

Population size and the decay of heterozygosity 419Concept of the minimum viable population 421

Definition and applications 421Dynamics of species viability 423

Extinctions 424Demographic stochasticity 426Environmental stochasticity including catastrophes 426Genetic stochasticity 426

Approaching adequate size and structure of genetic resources 429Conclusions 431Acknowledgements 432References 432

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Chapter 5: The role of biodiversity in forest ecosystems andfor sustainability 435

Genetic diversity in forest trees - its importance and potential humanimpact (Th. Geburek) 437

Introduction 437Forest ecosystem stability and its genetic component 438Potential threats to genetic diversity 440

Deforestation and fragmentation 440Exploitation 443Habitat alteration 445

Today's forestry 448Natural regeneration 448Artificial regeneration 450

Artificial regeneration: origin of forest reproductive material 450Artificial regeneration: reduction of genetic diversity in commercialseed lots 454Artificial regeneration: reduction of genetic diversity in clonal forestry... 454Artificial regeneration: mode of establishment (planting vs. seeding) . . . . 455Artificial regeneration: plant processing in nursery 456

Selective thinning in traditional forestry 456Conclusions 457References 458

Expected climate instability and its consequences for conservation offorest genetic resources (Cs. Mdtyds) 465

Longevity of trees and adaptation to environmental changes 465Life strategy options to mitigate effects of long-term environmental changes 466Analysis of response to environmental change 468Predicting effects of climate change \ 469Climate change scenarios 471

Species and gene migration constraints and the need for human interference . 472Priorities and need for action 474Conclusions 475References 476

Host-pathogen interaction in forest ecosystems (B. R. Stephan & Th.Geburek) 477

Introduction 477Host-pathogen interaction in stable forest ecosystems 478Host-pathogen interactions in disturbed forest ecosystems 482

Example 1: needle cast disease of Douglas fir (Pseudotsuga menziesii [Mirb.]Franco) .483Example 2: blister rust of five-needle pines 485

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Example 3: chestnut blight 487Example 4: Dutch elm disease and plane tree cancer 488

Effects of host-pathogen relationships on ecosystems 490Conclusions 490References 491

Chapter 6: In situ and ex situ conservation methods and techniques497

National Forest Inventories: how can they contribute to forest geneticconservation? (Th. Geburek & K. Schadauer) 499

Introduction 499Indicators of genetic erosion 500What can be realistically done through NFIs 501

Species distribution 501Subspecific entities 502Natural regeneration 504..Proportion of natural versus artificial regeneration 506Data collection for specific genes 506Socio-economic and political aspects 507Forest history and management regime 508

Outlook: Remote sensing 508Conclusions 509References 510

Protected areas in Europe and their importance for conservation (G. Koch)513

Introduction 513Protected forest areas ; 514

Protected forests without forest management: Strict Forest Reserves (SFRs) .. 516Protected forests without forest management: National Parks 517Protected forests with forest management: Genetic Reserve Forests (GRFs) .. 517Protected forests with forest management: Biosphere Reserves 519

Network of forest protection areas 521Socio-economic and political factors affecting reserve networks 523Research recommendations 523

Design of reserves 524Reserve size 525Buffer zones 526Sample plot design 526Visitors' access 528

Conclusions 529References 530

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In situ conservation methods (P. Rotach) 535Introduction 535A systematic approach to develop in situ conservation programmes 537

Collection of relevant information 537Selection of target species and setting of priorities 541Establishment of basic conservation method 545Identification and selection of populations to be conserved 548Definition of conservation objectives and specific targets 554Definition of management guidelines 555Establishment of a monitoring system 559

Conclusions 562References 562

Ex situ conservation methods (T. Skreppa) 567Introduction 567Ex situ conservation stands 568

Sampling the source population 569Site selection and plantation size 570Establishment 570Management 571Regeneration 571Effects of establishing several populations 572Ex situ conservation stands of introduced species (exotics) 572

Conservation seed orchards and clone collections 573Ex situ conservation populations in tree breeding 574

Genetic variability in breeding populations 575Clonal archives 576Seed orchards 577Genetic test plantations in tree breeding 578Ex situ conservation in research plantations 579

Ex situ conservation in arboreta or botanical gardens I 580Conclusions 580References 581

Multiple Population Breeding System as a method to conserve geneticvariation (G. Eriksson) 585

Introduction 585The functional roles of populations 585What is MPBS? 586Sampling of existing genetic variation and adaptedness 587Effective population size 588Adaptive management of genetic conservation subpopulations 590MPBS and conservation of associated species 593Selection and evolution in various directions 593

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Links to tree breeding programmes 595Conclusions 595Acknowledgements 596References 596

Seed and pollen storage: European focus (R. Klumpp) 601Introduction 601Seeds, pollen or single genes? 602Species' status 602

Reproductive biology 602Mating system 603The efficiency of collecting 604The efficiency of managing 605

Pollen storage 606Longevity and storage of seeds 607Current practices and case studies 610

Current practices with temperate and boreal species 610Current practices with Mediterranean species 612

Problems with recalcitrant and intermediate species 612Cryopreservation 616Conclusions 617References 618

Micro- and macropropagation of forest trees (E. Wilhelm) 623Introduction 623Micropropagation methods 624

Organogenesis 625Somatic embryogenesis 627Haploid embryogenesis in trees 629Long-term storage of germplasm 631Somaclonal variation 634Field evaluation and performance of micropropagated trees 635

Macrovegetative propagation methods 636Grafting and budding 636Cutting propagation 638

Topophysis 642Conclusions 643References 644

How can silvicultural management contribute to genetic conservation?(Th. Geburek & F. Miiller) 651

Introduction 651Common silvicultural types 652

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Sprout forests 652Coppice 652Coppice with standards 653

High forests 653Silvicultural practices relying on artificial regeneration 653

Forest Management system: clear-cutting with artificial regeneration 653Selection of seeds stands 654Selection of trees to be harvested 655Seed and plant processing 656Extraction of wild seedlings 657Planting 657Artificial seeding 658

Silvicultural practices relying on natural regeneration 658Naturally occurring forest types 658

Subalpine larch-stone pine forest 658Subalpine and montane spruce forests 658Boreal spruce forest 659Boreal pine forest 659Lowland and submontane beech forest 659Mixed beech-fir-spruce forests 659Mixed oak-hornbeam forests 660Mediterranean and submediterranean mixed oak forest 660

General genetic effects 660Forest management systems 661

Forest management system: seed-tree stand (clear-cutting with reser-vation of standards) 661Forest management system: regular shelterwood 662Forest management system: strip shelterwood and wedge shelterwood .. 663Forest management system: irregular (group) shelterwood 663Forest management sytem: selection forest ('Plenterwald') 664Forest management systems: close-to-nature forestry - the concept ofcontinuos forest cover ('Dauerwald') \ 665

Conclusions 665References 666

Species index 671Subject index 677

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