potato biology and biotechnology

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POTATO BIOLOGY AND BIOTECHNOLOGY
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Cover images:
The images are credited to the Scottish Crop Research Institute. We are grateful for the
use of the photographs they provided.
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POTATO BIOLOGY AND
BIOTECHNOLOGY
ADVANCES AND
PERSPECTIVES
Edited by
DICK VREUGDENHIL
Laboratory of Plant Physiology
Wageningen University and Research Centre
Wageningen,
The Netherlands
with
JOHN BRADSHAW
CHRISTIANE GEBHARDT
FRANCINE GOVERS
DONALD K.L. MACKERRON
MARK A. TAYLOR
HEATHER A. ROSS
Amsterdam – Boston – Heidelberg – London – New York – Oxford
Paris – San Diego – San Francisco – Singapore – Sydney – Tokyo
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Elsevier
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First edition 2007
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Preface
The potato is the fourth most important food crop in the world after wheat, maize and rice
with 311 million tonnes produced from 19 million hectares at an average fresh weight
yield of 16.4 t/ha in 2003 (FAO statistics), but with a huge range from 2 to 44 t/ha by
country. As well as being a staple food the potato is grown as a vegetable for table
use, is processed into French fries and chips (crisps) and is used for dried products and
starch production. Processing is the fastest growing sector of the world potato economy,
and today, processors are building factories in countries where the potato is primarily
grown as a staple food. In some countries, the potato is still fed to animals but this
use is decreasing. In many countries in Asia, Africa and Central and South America,
there is a need for increased and stable potato production to meet increasing demands for
food from human population growth during a period of environmental (including climate)
change. Potatoes with improved nutritional and health properties are desirable, but the
overriding need is for increased and stable yields to eradicate human hunger and poverty.
In those countries where food security has been achieved, the potato industries are trying
to increase potato usage in an economically and environmentally sustainable way. The
emphasis is on more yield of saleable product at less cost of production, reduced use
of pesticides and fungicides, better use of water and fertilizers and meeting consumer
demands for healthy convenience foods and novel products. These objectives will be met
only through new cultivars, better crop management and utilization of resources, better
post-harvest storage, better control of pests and diseases and a better understanding of the
social, economic and market factors that influence global production and distribution.
Today there is a tremendous opportunity to harness recent advances in potato biology
and biotechnology in these endeavours. We therefore considered it timely to ask a number
of experts to help us review the current state-of-knowledge in all aspects of the potato
crop, from basic science to production, processing and marketing. Therefore, this book

includes a wide variety of chapters, describing potato markets, genetics and genetic
resources, plant growth and development, response to the environment, tuber quality, pests
and diseases, biotechnology and crop management. We gave authors as much freedom
as possible over the content and style of their chapters, consistent with the subject matter
forming a comprehensive and coherent book without unnecessary duplication. We did
our best to help authors make their contributions as readable and free from errors as is
possible in a human enterprise. The idea to compile such a comprehensive volume was
put forward and initiated by our colleagues Howard Davies and Roberto Viola. We are
indebted to them and to many other colleagues for their support, especially Philip Smith
for his proof reading, and to the publishers for their advice and encouragement.
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vi Preface
We hope that the finished product will be of value not only to potato biologists but also
to all those people throughout the world interested in ensuring that the potato continues
to make a major contribution to the feeding of humankind.
Dick Vreugdenhil
John Bradshaw
Christiane Gebhardt
Francine Govers
Donald K.L. MacKerron
Mark A. Taylor
Heather A. Ross
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Acknowledgement
The editors are grateful for the funding of colour prints received from those listed below.
The British Potato Council
UK
SaKa-Ragis Pflanzenzucht GbR

Hamburg, Germany
Böhm-Nordkartoffel Agrarproduktion OHG
Lüneburg, Germany
Intersnack Knabber-Gebäck GmbH & Co. KG
Köln, Germany
Agrico Research BV
Emmeloord, The Netherlands
HZPC Holland BV
Metslawier, The Netherlands
C. Meijer BV
Rilland, The Netherlands
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Contents
Preface v
Acknowledgement vii
List of contributors xxvii
Part I The Markets 1
1 The Fresh Potato Market 3
Iain McGregor
1.1 Introduction and Overview 3
1.2 Production 3
1.3 Supply 7
1.4 Demand 7
1.5 Expenditure and Consumption 12
1.6 The Consumers’ Views 14
1.6.1 When potatoes are consumed 15

1.7 Prices Paid to Producers 18
1.8 Potatoes and the Health Issue 20
1.8.1 Glycaemic indices 22
1.9 Summary, Conclusions and Future Prospects 24
1.9.1 Key points 25
2 Global Markets for Processed Potato Products 27
Michael A. Kirkman
2.1 Introduction 27
2.2 Processed Potato Products 27
2.3 History of Potato Processing 28
2.4 Current Dimensions 29
2.4.1 Global production and consumption 29
2.4.2 Trends 30
2.4.3 Drivers 30
2.5 Potato-Processing Companies and Locations 32
2.6 Potato Supply 33
2.6.1 Supply chain 33
2.6.2 Variety requirements 34
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x Contents
2.7 Potato Cost 37
2.7.1 Theory and practice 37
2.7.2 Contracts 38
2.8 Potato Quality 39
2.8.1 Introduction 39
2.8.2 Tuber shape, size and dry matter composition 39
2.8.3 Blemishing diseases and disorders 40
2.8.4 Sugars and fry colours 41
2.9 Current Issues and Future Development 41

2.9.1 Acrylamide 41
2.9.2 Obesity 42
2.9.3 Nutritional value 43
3 The seed potato market 45
Kees D. van Loon
3.1 Seed Tubers 45
3.2 Seed Market 46
3.2.1 ‘Conventional’ seed tubers 46
3.2.2 Mini-tubers 49
3.2.3 True potato seed 50
3.3 Barriers to Markets in Seed Potatoes 50
3.3.1 Quarantine diseases and pests 50
3.3.2 Non-quarantine diseases and pests 51
3.3.3 Breeder’s rights 51
Part II Genetics and Genetic Resources 53
4 Molecular Taxonomy 55
Ronald G. van den Berg and Mirjam M.J. Jacobs
4.1 Introduction 55
4.2 Taxonomic Background 55
4.2.1 Wild and cultivated potatoes 55
4.2.2 The evolutionary framework 57
4.2.3 Remaining taxonomic problems 58
4.3 Molecular Data 58
4.3.1 Molecular markers applied to tuber-bearing
Solanum spp. 58
4.3.2 Methods of analysis of molecular data sets – phenetic versus
cladistic approaches 59
4.3.3 Application of molecular data to the taxonomy of the
tuber-bearing Solanum spp. 59
4.4 Conclusion 74

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Contents xi
5 Molecular Markers, Maps and Population Genetics 77
Christiane Gebhardt
5.1 Introduction 77
5.2 DNA Marker Types Useful for Potato Genetics 78
5.2.1 Restriction fragment length polymorphism 78
5.2.2 Amplified fragment length polymorphism 80
5.2.3 Simple sequence repeat or microsatellite 80
5.2.4 Cleaved amplified polymorphic sequence, sequence
characterized amplified region and allele-specific
amplification 81
5.2.5 Single-nucleotide polymorphism 81
5.3 Principles of Linkage Map Construction 82
5.4 Molecular Maps of Potato 83
5.5 Comparing the Potato with other Plant Genomes 85
5.6 Population Genetics 86
6 Genetics of Morphological and Tuber Traits 91
Herman J. van Eck
6.1 Introduction 91
6.1.1 The breeder’s perspective 91
6.1.2 What is heritable variation? 91
6.1.3 Morphological and tuber traits discussed in this chapter 92
6.2 Classical Potato Genetics with Molecular Techniques 92
6.2.1 The characteristics of classical genetic analysis 92
6.2.2 The characteristics of molecular genetic analysis 94
6.2.3 Quantitative and qualitative genetic approaches 95
6.3 The Genetics of Morphological Traits 96
6.3.1 Tuber flesh colour 96

6.3.2 Tuber skin and flower colour 99
6.3.3 Tuber shape 100
6.3.4 Eye depth 101
6.3.5 Tuber skin characters 102
6.4 Genetics of Tuber Physiology 103
6.4.1 Tuberization 103
6.4.2 Dormancy, sprouting 104
6.5 Tuber Quality Traits 104
6.5.1 Starch content 104
6.5.2 Discolouration 105
6.5.3 Texture 109
6.5.4 Glycoalkaloids 110
6.5.5 Growing defects (hollow hearts, growth cracks, second growth,
internal heat necrosis) 111
6.5.6 Tuber size uniformity 111
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xii Contents
7 Genetics of Resistance to Pests and Disease 117
Ivan Simko, Shelley Jansky, Sarah Stephenson and David Spooner
7.1 Resistance Screening 117
7.1.1 Field screening 117
7.1.2 Greenhouse screening 119
7.1.3 Laboratory screening 120
7.2 Resistance Genetics in Potato 121
7.2.1 Resistance breeding 121
7.2.2 Resistance genetics based on disease phenotype 127
7.3 Molecular Analysis of Potato Resistance 130
7.3.1 Experimental strategies for gene mapping and cloning 130
7.3.2 Resistance factors mapped in potato 132

7.3.3 Resistance genes cloned and characterized 141
7.3.4 Synteny of resistance loci in Solanaceae 145
7.3.5 Marker-assisted resistance breeding 147
8 Potato-Breeding Strategy 157
John E. Bradshaw
8.1 Introduction 157
8.2 Evolution of the Modern Potato Crop 157
8.3 Potato Breeding and the Need for New Cultivars 158
8.3.1 Potato breeding 158
8.3.2 Need for new cultivars 159
8.3.3 True potato seed 160
8.4 Adaptation to Environments and End Uses 160
8.4.1 Genotype by environment interactions 160
8.4.2 Ideotypes 161
8.5 Germplasm Available 161
8.5.1 Wild species 162
8.5.2 Cultivated species 164
8.6 Introgression of Genes from Wild Species 165
8.6.1 Sexual and somatic hybridization of S. tuberosum with
wild species 165
8.6.2 Molecular-marker-assisted introgression and gene
cloning 166
8.6.3 Base broadening versus introgression 166
8.7 Breeding Cultivars at the Tetraploid Level for Clonal Propagation 167
8.7.1 Parents 167
8.7.2 Early generations 168
8.7.3 Intermediate and later generations 169
8.7.4 Genetic knowledge and molecular-marker-assisted
selection 169
8.8 Breeding Cultivars for TPS 170

8.9 Genetically Modified Potatoes 171
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Contents xiii
8.10 Achieving Durable Disease and Pest Resistance 173
8.11 Conclusions 174
9 Genomics 179
Glenn J. Bryan
9.1 Introduction 179
9.2 Characteristics of the Potato Genome 180
9.3 Gene Isolation 180
9.3.1 Early gene cloning and expression studies 180
9.3.2 Map-based gene isolation 182
9.3.3 Use of candidate gene approaches for gene isolation 182
9.4 Structural Genomic Resources 184
9.4.1 Large-insert genomic libraries 184
9.4.2 Expressed sequence tag resources 184
9.5 Analysis of Potato Gene Expression 187
9.6 Microarrays 189
9.7 Functional Genomic Resources 192
9.7.1 The phenotype gap 192
9.7.2 Transgenic approaches for the study of gene function 193
9.7.3 Transposon tagging 194
9.7.4 Virus-induced gene silencing 194
9.7.5 Activation tagging 196
9.8 Towards a Genome-Wide Physical Map and a Potato Genome
Sequence 197
9.9 Proteomics and Metabolomics 197
9.10 Genomic Databases 199
9.11 Summary 199

10 Potato Cytogenetics 203
Tatjana Gavrilenko
10.1 Introduction 203
10.2 Basic Chromosome Number and Polyploid Complexes 203
10.3 Genome and Species Relationships 204
10.3.1 Genomic designation and relationships of diploid potato
species 204
10.3.2 Genomic nature and relationships in polyploid potato
species 205
10.3.3 Genomic designation and relationships of potato and
non-tuber-bearing species from closely related sections
Etuberosum, Juglandifolium and Lycopersicum 208
10.4 Karyotyping of Potato Species 209
10.4.1 Fluorescent in situ hybridization-based cytogenetic
mapping 209
10.5 Cytogenetics in Potato Improvement 212
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xiv Contents
Part III Plant Growth and Development 217
11 Above-Ground and Below-Ground Plant Development 219
Paul C. Struik
11.1 Introduction 219
11.2 General Morphology 219
11.3 Sprout Development 220
11.4 The Shoot System 221
11.5 The Leaves 222
11.6 The Stolon System 229
11.7 The Tubers 231
11.8 Organs of Sexual Reproduction 232

11.9 Root System 233
11.10 Association Between Development of Above-Ground and
Below-Ground Plant Parts 233
12 Signalling the Induction of Tuber Formation 237
David J. Hannapel
12.1 Introduction 237
12.2 Historical Background 238
12.2.1 Photoregulation 240
12.3 The Role of Growth Regulators in Controlling
Tuberization 242
12.3.1 Gibberellins 242
12.3.2 Cytokinins 243
12.3.3 Lipoxygenase activity and the role of
jasmonates 245
12.4 Gene Activity During Early Tuber
Formation 245
12.5 The Role of Specific Transcription Factors in Tuber
Development 249
12.5.1 A MADS box protein that regulates axillary branching and
affects tuber formation 249
12.5.2 Transcription factors from the TALE superclass 250
12.5.3 Overexpression of POTH1 negatively regulates
GA levels 250
12.5.4 POTH1 protein interacts with seven unique potato BEL
transcription factors 250
12.5.5 Over-expression of POTH1 and StBEL5 produces an
enhanced capacity to form tubers 251
12.5.6 Mechanism for transcription factors in regulating
tuberization 251
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Contents xv
13 Photosynthesis, carbohydrate metabolism and source–sink relations 257
Daniel Hofius and Frederik A.J. Börnke
13.1 Introduction 257
13.2 Photosynthetic Carbon Metabolism 258
13.2.1 CO
2
fixation 258
13.2.2 Carbon partitioning in mesophyll cells 260
13.2.3 Sucrose biosynthesis in source leaves 261
13.3 Starch Metabolism in Source Leaves 265
13.3.1 Starch synthesis within the chloroplast 265
13.3.2 Starch breakdown in leaves 266
13.4 Carbon Export and Long-Distance Transport 268
13.4.1 Pathway from the mesophyll to the
phloem 268
13.4.2 Phloem loading 269
13.4.3 Long-distance transport in the phloem 272
13.5 Carbon Unloading into Sink Organs 273
13.5.1 Symplastic and apoplastic routes of
unloading 273
13.5.2 Phloem unloading in the tuber 275
13.6 Sucrose to Starch Conversion in the Tuber 276
13.6.1 Production of hexose phosphates in the
cytosol 276
13.6.2 Uptake of carbon into the amyloplast 277
13.6.3 Starch synthesis in potato tubers 277
13.7 Source–Sink Regulation by Sugars 279
14 Dormancy and Sprouting 287

Jeffrey C. Suttle
14.1 Introduction 287
14.2 Tuber Dormancy Characteristics 288
14.3 Cell Biology of Dormancy 290
14.4 Gene Expression During Dormancy Transition 293
14.5 Hormonal Regulation of Tuber Dormancy 294
14.5.1 Auxins 294
14.5.2 Abscisic acid 295
14.5.3 Ethylene 299
14.5.4 Gibberellins 300
14.5.5 Cytokinins 301
14.5.6 Other endogenous growth substances 303
14.5.7 Hormonal regulation of tuber dormancy:
an overview 304
14.6 Sprout Growth and Physiological Aging 304
14.7 Conclusions 305
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xvi Contents
15 Molecular Physiology of the Mineral Nutrition of the Potato 311
Marcel Bucher and Jens Kossmann
15.1 Introduction 311
15.2 Nitrogen 313
15.2.1 Nitrogen uptake 313
15.2.2 Nitrogen assimilation 314
15.2.3 Transport of organic N between source and sink 318
15.3 Phosphorus 319
15.3.1 Phosphate uptake 320
15.3.2 Molecular biological analysis of P
i

transport systems 321
15.3.3 P
i
translocation on the whole plant level: long-distance
transport 324
15.4 Conclusion and Outlook 326
Part IV Response to the Environment 331
16 Water Availability and Potato Crop Performance 333
J. Vos and A.J. Haverkort
16.1 Introduction 333
16.2 Determinants and Controls of Water Movement 334
16.2.1 The transport of water in the soil–plant–atmosphere
continuum 334
16.2.2 Plant water relations 336
16.3 Assessing Plant Water Status 338
16.4 Potato Plant Responses to Drought and Biotic Stress 339
16.4.1 Leaf expansion 339
16.4.2 Effect of drought on plant calcium and
13
C
concentrations 340
16.5 Water Use, Leaf Dynamics and Potato Productivity 342
16.5.1 Water-use efficiency in different climates 342
16.5.2 Relative transpiration and leaf dynamics 343
16.5.3 Interactions between drought and biotic stresses 345
16.6 Varietal Differences in Drought Tolerance 347
16.7 Effects of Water Availability on Quality 348
17 Potato crop response to radiation and daylength 353
A.J. Haverkort
17.1 Radiation 353

17.1.1 Development of radiation interception 353
17.1.2 Measurement of radiation interception 356
17.1.3 Environmental effects on interception of solar
radiation 357
17.1.4 Radiation use efficiency 358
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Contents xvii
17.2 Daylength 360
17.2.1 Morphology 360
17.2.2 Tuber initiation 360
17.2.3 Short day sensitivity 363
17.2.4 Earliness 363
18 Responses of the Potato Plant to Temperature 367
Paul C. Struik
18.1 Introduction 367
18.1.1 Background and warnings 367
18.1.2 Reader’s guide 368
18.2 Sprout Growth, Emergence and Crop Establishment 368
18.3 The Shoot System 369
18.3.1 Leaf appearance 370
18.3.2 Final leaf number 370
18.3.3 Leaf growth and leaf size 372
18.3.4 Life span of leaves and specific leaf area 373
18.3.5 Number of stems 374
18.3.6 Stem morphology 374
18.3.7 Stem branching 375
18.4 Stolons 375
18.5 Tubers 377
18.5.1 Tuber induction and tuber initiation 377

18.5.2 Tuber set 378
18.5.3 Tuber bulking 379
18.5.4 Dry matter partitioning to tubers and harvest index 379
18.5.5 Tuber yield 380
18.5.6 Tuber number 380
18.5.7 Tuber size distribution 381
18.5.8 Tuber quality 381
18.5.9 Tuber enzyme activity 381
18.6 Inflorescences and Flowers 382
18.7 Root System 384
18.8 Photosynthesis, Dry Matter Production and Dry Matter Partitioning 384
18.9 Partial Exposure 385
18.10 Effects of Short Periods of Changes in Temperature 386
18.11 Diurnal Temperature Fluctuations 388
18.12 Physiological Behaviour of Seed Tubers 388
18.13 Summary 391
19 Response to the Environment: Carbon Dioxide 395
Ludwig De Temmerman, Karine Vandermeiren and Marcel van Oijen
19.1 Introduction 395
19.2 Effects of Increased CO
2
on Crop Growth and Development 396
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xviii Contents
19.3 Effects of Increased CO
2
on Potato Physiology 397
19.4 Effects of Increased CO
2

on Yield and Quality 400
19.5 Interactions Between Yield and Stresses at Elevated CO
2
405
19.6 Modelling Future Potato Productivity 406
19.6.1 Source-driven potato growth models 407
19.6.2 Source–sink-based potato growth models 408
19.6.3 Applications of potato models to CO
2
-related issues:
towards integrated assessment 409
19.7 Conclusions 409
20 Towards the Development of Salt-Tolerant Potato 415
D.J. Donnelly, S.O. Prasher and R.M. Patel
20.1 Introduction 415
20.2 Salt-Affected Agricultural Lands – Where are They? 416
20.2.1 Is potato grown in salt-affected areas? 416
20.3 Integrated Approach to Cropping Saline Soils 419
20.4 Mechanisms of Salinity Tolerance in Plants 420
20.4.1 What is known of salinity tolerance mechanisms in potato? 420
20.5 Classification of Salinity Tolerance in Potato 421
20.6 Evaluations of Salinity Tolerance in Potato 422
20.6.1 Field and greenhouse evaluations of salinity tolerance in
potato 422
20.6.2 In vitro evaluations of salinity tolerance in potato 423
20.7 Engineering and Cultural Management Practices for Modulation of
Salinity Stress 425
20.7.1 Water management for potato crops under salinity stress 425
20.7.2 Fertiliser management for potato crops under salinity stress 427
20.7.3 Climatic conditions modulate salinity effects on potato 428

20.8 Producing Salinity Tolerant Potato 429
20.8.1 Salinity-tolerant wild and/or primitive potato species 429
20.8.2 Domestication of wild salt-tolerant potato 429
20.8.3 Breeding for increased vigour and yield 430
20.8.4 Obtaining salinity tolerance through cell and tissue culture
techniques 431
20.8.5 Obtaining salinity tolerant potato through genetic engineering 433
20.9 Summary 434
Part V Tuber Quality 439
21 The Harvested Crop 441
Michael Storey
21.1 Introduction 441
21.2 Nutritional Value 442
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Contents xix
21.3 Dry Matter 444
21.3.1 Carbohydrates 444
21.3.2 Protein 448
21.3.3 Vitamins 451
21.3.4 Allergens and anti-nutritionals 452
21.3.5 Glycoalkaloids 453
21.3.6 Other tuber metabolites 454
21.3.7 Minerals 454
21.4 Flesh and Skin Colour 455
21.4.1 Carotenoids 455
21.4.2 Anthocyanins 456
21.5 Greening 458
21.6 Mechanical Damage and Bruising 459
21.6.1 Enzymic browning 460

21.6.2 Structural and cellular changes 461
21.6.3 Field factors and tuber water status 462
21.7 Concluding Comments 466
22 Skin-set, Wound Healing, and Related Defects 471
Edward C. Lulai
22.1 Introduction 471
22.2 Native Periderm and Skin-Set 472
22.2.1 Native periderm formation 472
22.2.2 Skin-set: a part of native periderm maturation 473
22.2.3 Skin-set and native periderm physiology 474
22.2.4 Periderm architecture and skinning injury 476
22.2.5 Cellular changes associated with skin-set 477
22.3 Wound Healing 479
22.3.1 The process of tuber wound healing 479
22.3.2 Induction of suberization 480
22.3.3 Regulation of suberization 481
22.3.4 Environmental effects on suberization 483
22.3.5 Characteristics of the biopolymers that form suberin 483
22.3.6 Suberization: closing layer and wound periderm formation 484
22.3.7 Suberin biosynthesis and structure 485
22.3.8 Suberization and resistance to infection 492
22.4 Related Defects 492
22.4.1 Wound-related tuber defects 492
22.4.2 Shatter bruising and tuber cracking 493
22.4.3 Blackspot and pressure/crush bruising 493
22.4.4 Growth cracks 495
22.4.5 Skinning 495
22.5 Summary 496
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xx Contents
23 Internal Physiological Disorders and Nutritional and Compositional
Factors that Affect Market Quality 501
Joseph R. Sowokinos
23.1 Introduction 501
23.2 General Nature, Incidence and Severity of Internal
Physiological Disorders 502
23.2.1 Calcium nutrition and tuber quality 502
23.2.2 Brown centre and internal brown spot 504
23.2.3 Hollow heart 507
23.2.4 Internal heat necrosis 509
23.2.5 Stem-end discolouration 510
23.2.6 Translucency 512
23.2.7 Mottling 513
23.3 Summary of Internal Physiological Disorders 515
23.4 Compositional and Nutritional Changes Affecting End-Use
Quality 515
23.4.1 Carbohydrates – starch 516
23.4.2 Carbohydrates – sugars 516
23.4.3 Factors affecting RS concentration in stored potatoes 517
23.4.4 Chemical maturity monitoring 518
24 Potato Flavour and Texture 525
Mark A. Taylor, Gordon J. McDougall and Derek Stewart
24.1 Introduction 525
24.2 Potato Flavour 525
24.2.1 Non-volatile components 525
24.2.2 Glycoalkaloids and flavour 527
24.2.3 Volatile compounds 527
24.2.4 Molecular and genetic approaches to the study of potato
flavour 530

24.2.5 Molecular approaches to dissecting key constituents of
tuber flavour 531
24.3 Potato Tuber Texture 532
Part VI Pests and Diseases 541
25 Insect Pests in Potato 543
Edward B. Radcliffe and Abdelaziz Lagnaoui
25.1 Yield and Quality Effects 543
25.1.1 Defoliators 543
25.1.2 Sap feeders 544
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Contents xxi
25.1.3 Pathogen transmission 544
25.1.4 Root and tuber feeding 545
25.2 Insect Pests of Worldwide Importance 545
25.2.1 Aphids 545
25.2.2 Colorado potato beetle 550
25.2.3 Potato tuber moths 552
25.2.4 Leafminers 554
25.3 Regional Pests 556
25.3.1 Leafhoppers 556
25.3.2 Potato psyllid 557
25.3.3 Thrips 557
25.3.4 White grubs 558
25.3.5 Wireworms 558
25.3.6 Ladybird beetles 558
25.3.7 Flea beetles 559
25.3.8 Andean potato weevils 559
25.3.9 Cutworms 560
25.4 Insect Control Tactics 560

25.4.1 Insecticides 561
25.4.2 Host plant resistance 561
25.4.3 Biological control 562
25.5 Conclusions 562
26 The Nematode Parasites of Potato 569
Didier Mugniéry and Mark S. Phillips
26.1 Potato Cyst Nematodes (Globodera Rostochiensis and Globodera
Pallida) 569
26.1.1 Host range 570
26.1.2 Diseases 572
26.1.3 Biology 573
26.1.4 Dormant stage 574
26.2 Root-Knot Nematodes (Meloidogyne SPP.) 575
26.2.1 Disease 576
26.2.2 Biology 576
26.2.3 Spread 578
26.3 The False Root-Knot Nematode Nacobbus Aberrans 578
26.3.1 Host range 578
26.3.2 Disease 579
26.3.3 Biology 579
26.3.4 Spread 579
26.4 Virus Vector Nematodes (Trichodorus SPP.) 580
26.4.1 Disease 581
26.4.2 Biology 581
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xxii Contents
26.5 The Root Lesion Nematodes (Pratylenchus SPP.) 582
26.5.1 Disease 582
26.5.2 Biology 582

26.6 Ditylenchus Destructor and Ditylenchus Dipsaci 582
26.6.1 Host range 583
26.6.2 Disease 583
26.6.3 Biology 584
26.7 Control 584
26.7.1 Prophylaxis 584
26.7.2 Cultural methods 585
26.7.3 Physical methods 586
26.7.4 Chemical treatments 586
26.7.5 Biological methods 587
26.7.6 Resistant varieties 588
26.8 Conclusions 591
27 Bacterial Pathogens of Potato 595
Jan M. van der Wolf and Solke H. De Boer
27.1 Introduction 595
27.2 Pathogen Biology 595
27.2.1 Ralstonia solanacearum 595
27.2.2 Clavibacter michiganensis ssp. sepedonicus 597
27.2.3 Pectolytic erwinias 597
27.2.4 Streptomyces scabies 599
27.3 Pathology 600
27.3.1 Symptoms and factors favouring symptom expression 600
27.3.2 Economic importance 604
27.3.3 Geographic distribution 605
27.4 Ecology 606
27.4.1 Plant colonization 606
27.4.2 Survival 607
27.4.3 Dissemination 608
27.5 Control 610
27.5.1 Use of clean seed 610

27.5.2 Inoculum reduction 612
27.5.3 Agronomic practices 613
27.6 Perspectives 614
28 Viruses: Economical Losses and Biotechnological Potential 619
Jari P.T. Valkonen
28.1 Introduction 619
28.2 Viruses Infecting Potato 619
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28.3 New and Emerging Viruses and their Detection 622
28.3.1 Molecular detection and identification 622
28.3.2 New viruses 623
28.3.3 Emerging viruses 624
28.4 Economic Impact of PVY 626
28.4.1 Mixed infections 628
28.4.2 Impact of primary and secondary infection 628
28.4.3 Costs to seed production 629
28.4.4 Yield loss depending on cultivar resistance and
PVY strain 630
28.4.5 Aetiology and evolutionary perspectives 631
28.5 Infectious cDNA Clones of Potato Viruses and their use as
Biotechnological Tools 632
28.5.1 Use of potato viruses as gene vectors 632
28.5.2 Studies on viral infection cycle using infectious cDNAs of
potato viruses 633
29 Fungal and Fungus-Like Pathogens of Potato 643
Aad J. Termorshuizen
Part VII Biotechnology 667
30 Developments in Transgenic Biology and the Genetic Engineering of

Useful Traits 669
Steve Millam
30.1 Introduction 669
30.2 Genetic Transformation of Potato 670
30.3 Developments in Transgenic Biology 674
30.3.1 Protocol refinements 674
30.3.2 Enhanced or alternative transformation
strategies 675
30.4 The Genetic Engineering of Useful Traits 678
30.4.1 Resistance to major pests and diseases 679
30.4.2 Tuber quality traits 681
30.4.3 Nutritional value 681
30.5 Summary and Future Developments 683
31 Field-Testing of Transgenic Potatoes 687
A.J. Conner
31.1 Introduction 687
31.2 Transgenic Potatoes in the Context of Potato
Breeding 689
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31.3 The Importance of Field-Testing Transgenic Potatoes 691
31.3.1 Field confirmation of transgenic phenotype 691
31.3.2 Occurrence of off-types 692
31.4 The Design of a Field-Testing Programme 695
31.5 Strategies to Reduce the Frequency of Off-Types 697
31.6 Assessment of Biosafety Issues 699
31.7 Conclusions 701
32 Soil-Free Techniques 705
Steve Millam and Sanjeev K. Sharma

32.1 Introduction 705
32.2 Mini-Tuber Production 708
32.3 In Vitro Multiplication Techniques 709
32.3.1 Axillary-bud proliferation 709
32.3.2 Micro-tuber production 710
32.3.3 Somatic embryogenesis 712
32.4 Hydroponics and Aeroponics 714
32.5 Future Prospects 715
Part VIII Crop Management 717
33 Agronomic Practices 719
D.M. Firman and E.J. Allen
33.1 Introduction 719
33.2 Planning and Preparation 719
33.2.1 Market 720
33.2.2 Calendar 721
33.2.3 Seed 722
33.2.4 Site selection 727
33.2.5 Soil analysis 727
33.2.6 Fertiliser 728
33.3 Soil Management 729
33.3.1 Cultivation 729
33.3.2 Control of soil-borne pests and diseases 730
33.3.3 Weed control 730
33.3.4 Irrigation 731
33.4 Crop Establishment and Management 732
33.4.1 Planting 732
33.4.2 Crop protection 733
33.4.3 Covers, mulches, soil amendments and
intercropping 734
33.4.4 Defoliation 734

33.4.5 Harvesting 735