Contents...1 Introduction 149 2 Factors affecting tissue culture techniques 153 3 Tissue culture techniques 155 4 Future trends 165 5 References 165 5 Advances in molecular breeding

© Burleigh Dodds Science Publishing Limited, 2020. All rights reserved.

Contents

Series list xReviewers to acknowledge xviIntroduction xvii

Part 1 Physiology and breeding

1 Environmental physiology of ornamental crops 3John Erwin, University of Maryland, USA

1 Introduction 3

2 Propagation: sexual propagation 3

3 Propagation: asexual propagation 12

4 Vegetative development 21

5 Increasing plant mass 26

6 Reproductive development 28

7 Dormancy 49

8 References 50

2 Exploiting the genetic diversity of ornamentals 71Yoo Gyeong Park, Gyeongsang National University, Republic of Korea; Young Hoon Park, Pusan National University, Republic of Korea; Abinaya Manivannan, National Institute of Horticultural and Herbal Science, Republic of Korea; Prabhakaran Soundararajan, National Institute of Agricultural Science, Republic of Korea; and Byoung Ryong Jeong, Gyeongsang National University, Republic of Korea

1 Introduction 71

2 Management of ornamental germplasm to exploit genetic diversity:

collection strategy 72

3 In situ conservation methods 73

4 Ex situ conservation 75

5 Management of plant genetic resources in the Republic of Korea

and in Japan 79


vi Contents

6 Management of plant genetic resources in the United States 82

7 Management of plant genetic resources in Canada 84

8 Management of plant genetic resources in Europe 88

9 Utilizing ornamental germplasm to exploit genetic diversity in roses 94

10 Utilizing ornamental germplasm to exploit genetic diversity in cacti 100

11 Conclusions 104

12 References 109

3 Advances in conventional breeding techniques for ornamentals 119Traud Winkelmann and Philipp Braun, Leibniz Universität Hannover, Germany; and Emmy Dhooghe and Johan van Huylenbroeck, Flanders Research Institute for Agriculture, Fisheries and Food, Belgium

1 Introduction 119

2 Current common breeding techniques in ornamentals 120

3 Conventional breeding for sustainable production of ornamentals 126

4 Conclusion and future trends 138

5 References 138

4 Advances in tissue culture techniques for ornamental plant propagation 149G. R. Rout, Odisha University of Agriculture and Technology, India; and S. Mohan Jain, University of Helsinki, Finland

1 Introduction 149

2 Factors affecting tissue culture techniques 153

3 Tissue culture techniques 155

4 Future trends 165

5 References 165

5 Advances in molecular breeding of ornamentals 189Neil O. Anderson, University of Minnesota, USA

1 Introduction 189

2 Flowers as study organisms for early genetic research 190

3 Flower breeding in the nineteenth and twentieth centuries 191

4 Early use of biotechnology in marker-assisted breeding 194

5 Flower color 198

6 Flowering genes 201

7 Genetically engineered flowers (GMOs and GEOs) 202

8 Genome sequencing 211

9 Gene editing (CRISPR-Cas9) 217

10 Conclusion 218


viiContents

11 Future trends in research 219

12 Where to look for further information 219

13 References 220

6 The use of gene-editing techniques in breeding improved ornamentals 231Bruno Trevenzoli Favero, Josefine Nymark Hegelund and Henrik Lütken, University of Copenhagen, Denmark

1 Introduction 231

2 Applications in horticulture: a case study on Campanula 237

3 Prospective uses and regulation of ornamental plants 245

4 Conclusion 246

5 Acknowledgements 246


7 References 247

7 Advances in abiotic stress-resistant varieties of ornamentals 255Qiansheng Li and Mengmeng Gu, Texas A&M AgriLife Extension Service, USA

1 Introduction 255

2 Improving resistance to low-temperature stress 257

3 Improving resistance to high-temperature stress 260

4 Improving resistance to drought 261

5 Improving resistance to waterlogging 264

6 Improving tolerance to saline conditions 265

7 Improving tolerance to poor soil conditions 268

8 Future trends and conclusion 269


10 References 270

8 Improving nutrient management in the cultivation of ornamental plants in greenhouse, container and field production 279John Majsztrik, Clemson University, USA and James S. Owen Jr., U.S. Department of Agriculture - Agricultural Research Service, USA

1 Introduction 279

2 Irrigation and water quality 281

3 Types of production systems 284

4 Mineral nutrients 289

5 Fertilizers 291

6 Fertilizer delivery 293


viii Contents

7 Soils and substrates 295

8 Environmental impacts of runoff 296

9 Future trends and conclusion 296


11 References 297

Part 2 Cultivation techniques

9 Advances in irrigation practices and technology in ornamental cultivation 305John D. Lea-Cox, University of Maryland, USA

1 Introduction 305

2 Ornamental production systems and water use 306

3 Key challenges for improving irrigation efficiency: systems

design and maintenance 311

4 Sensor-based technologies for irrigation scheduling 314

5 Irrigation data management and decision support systems 319

6 Developing a universal irrigation protocol: a case study 324

7 Future trends 327


9 References 328

10 Advances in protected cultivation of ornamentals 335James E. Faust, Clemson University, USA

1 Introduction 335

2 Light-emitting diode (LED) lighting systems 337

3 Manipulation of plant growth and development 346

4 Sustainable practices and renewable inputs 354

5 Case study: Propagation of ornamentals in a vertical propagation area 359

6 Conclusion and future trends 360


8 References 363

11 Diseases affecting ornamental geophytes and their control 367Gary A. Chastagner and Andrea R. Garfinkel, Washington State University, USA

1 Introduction 367

2 Biotic and abiotic diseases 370

3 General disease management principles 385

4 Approaches to reduce chemical fungicide usage 392

5 Managing fungicide resistance 394


ix

6 Botrytis infection of ornamental geophytes 395

7 Diversity of Botrytis species 399

8 Botrytis as an endophyte 401

9 Conclusions 402


11 References 404

Index 415

Contents


IntroductionOrnamental plants are plants grown for decorative purposes such as gardening and landscape design and include cut flowers, bulbs, potted plants, shrubs and trees. Like other crops, ornamentals face challenges such as biotic and abiotic stresses as well as the need to develop more sustainable, ‘climate-smart’ methods of cultivation. This collection reviews the rich range of research addressing these challenges. Part 1 discusses advances in understanding plant physiology, genetic diversity and breeding techniques. Chapters cover recent research on how plants respond to abiotic stress, ways of exploiting genetic diversity to improve target traits, advances in both conventional and marker-assisted breeding techniques, as well as their use to produce abiotic stress-resistant varieties. Part 2 surveys advances in cultivation techniques in such areas as nutrition, irrigation, protected cultivation and post-harvest storage. The book also includes chapters on developments in integrated disease and pest management.

Part 1 Physiology and BreedingAs Chapter 1 points out, growers produce hundreds of species from all over the world to market often at a specific time of year. To achieve this, growers must have a sophisticated understanding of the environmental physiology of plant growth and development. This chapter reviews key research on the physiology of sexual and asexual propagation, vegetative and reproductive development, photosynthesis and dormancy.

Chapter 2 discusses ways of exploiting genetic diversity in ornamentals. This chapter reviews collecting, conserving and utilizing ornamental germplasm. Topics include collection strategies, in situ and ex situ conservation methods. The chapter also summarises the management of ornamental genetic resources in key collections around the world. It concludes with a case study on utilizing ornamental germplasm to exploit genetic diversity in roses.

Chapter 3 reviews developments in breeding programs using conventional techniques such as classical cross breeding and selection, polyploidization and interspecific hybridization. The review summarises advances in key breeding techniques in ornamental crops and discusses their application in such areas as: drought tolerance, low temperature tolerance, elevated pH tolerance, resistance and tolerance to biotic stress, nutrient use efficiency and postharvest life.

Advances in tissue culture techniques for ornamental plant propagation is the focus for Chapter 4. It reviews research on key techniques and factors affecting their effectiveness. Techniques include: clonal propagation and


Introductionxviii

organogenesis, embryo rescue, thin cell layer (TCL) technology, somatic embryogenesis, the use of tissue culture in mutagenesis and mutation breeding, slow-growth conservation techniques, bioreactor/liquid culture, plant improvement through genetic engineering and the use of molecular markers in tissue culture.

Chapter 5 reviews advances in molecular breeding of ornamental plants such as marker-assisted selection. To illustrate developments it includes examples of breeding to improve flower colour and flowering. The chapter also reviews developments in genetic modification using the examples of carnations, roses and chrysanthemums. The chapter also discusses trends in genome sequencing and gene editing. Building on the previous chapter, Chapter 6 examines how innovation in breeding has increased with the advent of genome editing and its application in horticulture. These techniques use nucleases to guide site-directed mutagenesis such as the Clustered Regularly Interspaced Short Palindromic Repeats nuclease-associated systems (CRISPR). To illustrate the potential of the technique, the chapter focuses on the example of reducing ethylene sensitivity in the potted plant Campanula portenschlagiana.

Recently there has been more focus in ornamental breeding on the importance of tolerance to abiotic stresses, in response to more extreme weather events related to climate change such as drought and flooding. The final chapter in Part 1, Chapter 7, reviews ways of improving resistance to a wide range of stresses, including: low and high temperatures, drought, waterlogging and saline conditions as well as poor soil quality.

Part 2 Cultivation techniquesChapter 8 reviews nutrient management in different production systems for ornamentals. Proper nutrition in containerized and field grown plants is essential for maximized growth and profitability. The chapter discusses the use of macro- and micro-nutrients along with substrate or soil amendments to ensure that plants remain healthy and continue to grow optimally in both mineral soils and soilless substrates. The chapter reviews irrigation and water quality, fertilizer types and usage, the effect of soil and substrates on fertilizers, and the impact of production systems on fertilizer longevity.

Two issues will likely dominate irrigation and water management for ornamental cultivation in the coming years, namely the decreasing availability of fresh water for agricultural use and environmental concerns that will increasingly potentially contaminate the runoff of water from commercial operations. Chapter 9 discusses how both issues are the subject of increasing scrutiny and regulation in the US and Europe, and how many operations are addressing these issues through an array of better management practices. It discusses ways of increasing the precision of irrigation systems and how we can


Introduction xix

better time the frequency and duration of irrigation events, not only to optimize plant growth but also to limit leaching and runoff.

Chapter 10 addresses key developments in protected cultivation systems for ornamentals. The advent of LED technology has created new and unique opportunities for growing plants, while vertical propagation systems and the ability to precisely control the light spectra have the potential to revolutionize the industry in the coming years. Labour issues have also spurred the engineering of automated and robotic systems to improve production efficiency. The drive to reduce the industry’s dependency on synthetic pesticides, non-renewable resources and plastics has also led to the exploration of alternative production techniques and materials. This chapter explores these recent advances as well as future directions for this dynamic industry.

The final chapter in the book addresses the key challenge of diseases affecting ornamentals. It focuses on the example of geophytes which represent a significant segment of the world floriculture industry, particularly in cut flower production. Chapter 11 reviews the main biotic and abiotic diseases affecting geophytes and summarizes current best practice in integrated disease management to reduce fungicide use to address the problem of resistance. To illustrate integrated disease management in practice, it focusses on Botrytis infection of ornamental geophytes.

Part 1Physiology and breeding

http://dx.doi.org/10.19103/AS.2020.0066.01© Burleigh Dodds Science Publishing Limited, 2020. All rights reserved.

Chapter 1Environmental physiology of ornamental cropsJohn Erwin, University of Maryland, USA

1 Introduction 2 Propagation: sexual propagation 3 Propagation: asexual propagation 4 Vegetative development 5 Increasing plant mass 6 Reproductive development 7 Dormancy 8 References

1 IntroductionOrnamental horticulture is one of the most challenging sectors of agriculture. Growers produce hundreds of species from all over the world to market at a specific time of a year or for a holiday. To achieve this, growers must have a sophisticated understanding of the environmental physiology of plant growth and development including propagation and vegetative and reproductive development. These issues will be reviewed in this chapter along with identifying key areas that are gaps in our knowledge that have practical implications. Useful background on this important topic can be found in Dole and Wilkins (2004) and Acquaah (2008).

2 Propagation: sexual propagationThe degree of sophistication of seed production, storage, sowing, and germination technologies increased greatly with the advent of the ‘plug’ or seedling production industry in the 1980s. Seed producers developed better seed harvesting, maturing, and storage procedures that increased the rates of germination success and uniformity. Simultaneously, the seedling production industries installed seed storage facilities, environmentally controlled germination chambers and automated seeders and irrigation booms to ensure

Environmental physiology of ornamental crops Environmental physiology of ornamental crops

Environmental physiology of ornamental crops4


uniform and precise environmental and moisture conditions to maximize germination success. Yet seed and seedling producers still have challenges, often around providing optimal conditions for germination. Current knowledge about seed physiology and factors affecting dormancy, germination, and propagation is reviewed in Copeland and McDonald (2001), Basra (2006), Bewley et al. (2013), Geneve (2013), Finch-Savage and Bassel (2016), and Davies et al. (2017).

2.1 Seed storage

Improper temperature and/or humidity during seed storage can reduce the percentage of germination, uniformity in germination, and early growth. In general, optimal seed storage conditions for many species indigenous to temperate climates include low humidity and temperatures above 2°C. A general ‘rule of thumb’ used in the USA is that % humidity + temperature (oF) should be <70. Put in another way, seed longevity doubles when storage temperatures decrease to 5–6°C, and seed longevity doubles when seed moisture content is reduced to 1–2.5% (Roberts, 1991).

Seed viability over time varies with species. Seed viability of Aquilegia, Callistephus, Catharanthus, Centauria, Cleome, Consolida, Delphinium, Linum, Lunaria, Phlox, Salvia, Torenia, Verbena, and Viola decreases after the first year (Dole and Wilkins, 1999); in addition, the vigor of germinated seedlings decreases as the seed age increases (Corbineau and Come, 1991). Species in the genera Anemone, Asparagus, Callistephus, Calceolaria, Delphinium, Dahlia, Gloxinia, Phlox, Salivia, and Viola are generally viable for only 1–2 years. In contrast seed of species in the Begonia, Dendranthema, Centaurea, Cyclamen, and Tropaeolum genera can remain viable for 3–15 years (Corbineau and Come, 1991).

2.2 Temperature and germination

Delivering the optimal temperature for seed germination is still one of the greatest challenges in commercial production; this is particularly the case if growers are germinating seed in a greenhouse. In general, seedling producers manage air temperatures to maintain a media temperature of 22–24°C to maximize germination (Erwin, personal observation); this temperature range is optimal for many ornamental plant species (Table 1). In contrast to most species, foxglove (Digitalis purpurea), Phlox, and sweet pea (Lathyrus splendens), seed germination is optimal at 16–18°C. Vinca (Catharanthus roseus) and tomato (Lycopersicon esculentum) seed germination is optimal at 24–27°C.

Fluctuating temperatures can stimulate the germination of some species (Aragino, 1981). This can be the case with species from areas with large day/


Environmental physiology of ornamental crops 5

Table 1 Optimal temperature, light, and moisture requirements for seed germination of different ornamental crop species

Species Temperature Light Cover Moisture Storage

Abutilon 22–24 – Yes – –Achillea 18–21 Light No – –Agastache 15–21 – Yes – –Ageratum houstonianum 24–25 – No M MAmmi majus 22–24 Light – – –Anchusa capensis 20–21 Light – – –Anemone coronaria 18–21 – Yes – SAngelonia 22–24 Light No – –Antirrhinum majus 18–21 Light No M MAquilegia caerulea 21–24 – Yes – –Asclepias 21–24 – Yes – –Asclepias tuberosa 18–24 – – – –Asparagus densiflorus 29 day/24 night Dark Yes M SAstilbe x arendsii 21–24 – No – –Astrophytum myriostigma 20–25 – – – –Bacopa 20–23 Light No – –Begonia ‘Dragon Wing’ 24–25 Light No – –Begonia x hiemalis 24–26 Light – – –Begonia semperflorens 24–27 Light No W SBegonia tuberhybrida 24–26 Light No W SBellis perennis 21–24 LightBrachycome iberidofolia 21–22 Light – M –Browallia speciosa 24–25 – No M SCalceolaria herbeohybrida 18–21 Light – M –Calendula officinalis 20–21 Dark Yes D –Callistephus chinensis 18–21 – Yes D SCampanula isophylla 15–18 Light No – –Canna x generalis 21–24 Dark – M –Catharantus roseus 25–26 Dark Yes W SCelosia argentea 24–25 – No W MCentaurea americana 21–24 – Lightly M –Clarkia amoena 20–22 Dark – – –Cleome hassleriana 26 day/21 night Light No MCobea scandens 21 Cover

lightly– – –

Cordyline indivisa 22–26 Light Lightly M –

(Continued)




Coreopsis grandiflora 18–24 Light No D –Cosmos bipinnatus 18–21 – Yes D –

Crossandra infundibuliformis 26 day/21 night – Yes – –Cuphea platycentra 21 – Cover lightly M –Cyclamen persicum 18–20 Dark – W MCynoglossum amabile 18–21 Dark – – –Dahlia x hybrida 20–21 – Yes D MDelphinium grandiflorum 24–26/18–21 – Yes – SDendranthema x grandiflorum 15–21 Light – M –Dianthus barbaratus 21–24 – Cover lightly M MDianthus chinensis 18–20 – Cover lightly M MDiascia 18–21 – Yes – –Dichondra 22–24 – Yes – –Digitalis purpurea 21–24 Light No – –Dolichos lablab 21–22 Dark – – –Echinacea purpurea 20–22 – Yes – –Eschscholzia californica 21–22 – – – –Euphorbia marginata 15–20 Dark – – –Eustoma grandiflorum 20–25 Light Lightly – –Exacum affine 22–25 – Cover lightly M –Freesia spp. 15–20 Dark Yes – –Gaillardia x grandiflora 21–24 Light No M –Gaura lindheimeri 21–22 – Yes – –Gazania rigens 20–21 Dark Yes D –Gerbera jamesonii 20–22 Light Yes M –Gomphrena globosa 22–24 – Yes M –Helianthus annuus 18–25 – Yes M –Helichrysum bracteatum 21–24 – Yes M –Heliotrope 24–26 – – M –Hibiscus moscheutos 21–26 – Yes – –Hypoestes phyllostachya 21–24 – No M –Iberis amara 15–18 Light Cover lightly W –Impatiens balsamina 22–25 Light – W SImpatiens walleriana 22–24 Light No W SImpatiens hawkeri 22–25 Light No W SLathyrus splendens 13–20 – Yes – L

Table 1 (Continued)


Environmental physiology of ornamental crops 7


Lavendula angustifolia 18–24 – Yes M –Lavatera trimestris 22 Dark Yes – –Leucanthemum lacustre 20–25 – Cover lightly M LLimonium sinuatum 21–25 Dark – M –Linum perenne 21–25 – Cover lightly – –Lisianthus 20–22 Light No M MLobelia erinus 21–24 Light No M MLobularia maritima 20–22 Light – M MLupinus polyphyllus 18–21 – Yes M –Mathiola incana 17–21 – Yes – LMelampodium divaricatum 18–20 – Cover lightly M –Mimosa pudica 26 Dark Yes – –Mimulus x hybridus 15–21 Light No M –Mirabilis jalapa 22 Dark Yes – –Nemesia strumosa 20–23 – Yes – –Nemophila menziesii 18 Dark Lightly – –Nicotiana alata 21–24 Light No M –Nierembergia hippomanica var. violacea 22–24 – Cover lightly M –Nigella damascene 18–21 – Cover lightly – –Opuntia spp. 25–35 – – – –Osteospermum 21–24 – Yes – –Papaver nudicaule 15–18 Light No M –Papaver orientale 18–24 Light No M –Pelargonium x hortorum 21–24 – Yes W MPelargonium peltatum 21–24 – Yes W MPentas 23–25 – No M –Pericallis x hybrida 20–24 Light – M –Petunia x hybrida 24–26 Light No M MPhlox drummondi 18–20 Dark Yes D SPlatycodon grandifloras 20–22 Light No M –Portulaca grandiflora 25–26 – No M MPrimula vulgaris 18–20 Light Lightly W –Ranunculus asiaticus 15–20 – Yes M –Rudbeckia fulgida 28–31 – Cover lightly M –Rudbeckia hirta 20–21 – Yes M –Salpiglossus sinuate 21–22 Dark No – –

(Continued)



night temperature (DT/NT) fluctuations such as deserts and mountainous regions (Tudela-Isanta et al., 2018). For instance, Rumex retroflexus will not germinate when maintained at constant 20°C or 30°C, but 100% germination occurs when temperature is fluctuated between 20°C and 30°C diurnally (Toole et al., 1955). Cucumis anguria germination is inhibited by light, but it will germinate in the light if temperatures are fluctuated between 15°C and 35°C diurnally (Felippe, 1980). Rumex obtusifolius requires light to germinate, but it will germinate in the dark if temperature is fluctuated between 15°C and 35°C (Toole et al., 1955). Interestingly, recent research on cacti suggest that many species germinate better with constant 20–25°C rather than with fluctuating temperatures; authors suggest this may encourage germination under plants or in crevices where survival may be maximized (Lindow-Lopez et al., 2018).

Interestingly, short-term low- or high-temperature pulses can stimulate germination in some cases (Takaki et al., 1981). Cucumis imbibed at 25°C


Salvia coccinea 24–26 Dark Yes W SSalvia farinacea 21–25 – Yes W SSalvia splendens 21–25 – Yes W SScabiosa atropurpurea 21–24 Light No – –Senecio cineraria 22–24 – No M MSinningia speciosa 18–21 Light No – –Solenostemon scutellarioides 22–24 – Yes M MStenocereus griseus 24–25 Yes No – –Streptocarpus x hybridus 21 Light No – –Tagetes erecta 21–22 – Yes M MTagetes patula 21–22 – Yes M MThunbergia alata 21–24 – Yes M –Tithonia 20–21 – Yes – –Torenia fournieri 22–24 Light No M –Tropaeolum majus 18–21 Dark Yes M –Verbena x hybrida 24–26 – Yes D MVerbena bonariensis 24 – Cover lightly D MViola tricolor 18–24 – Yes W SViola wittrockiana 20–21 – Cover lightly M SZinnia angustifolia 20–22 – Yes D LZinnia elegans 20–22 – Yes D L

Source: developed from Styer and Koranski (1997), Dole and Wilkins (1999), Ball Horticultural Germination Fact Sheet, Pimiento-Barrios and del Castillo (2002), Erwin, personal observation.

Table 1 (Continued)


Index

ABA. see Abscisic acid (ABA)Abiotic disorders 385Abiotic stress-resistant varieties 255–257

future trends 269–270 improving resistance to

drought 261–264 high-temperature stress 260–261 low-temperature stress 257–260 waterlogging 264–265

improving tolerance to poor soil conditions 268–269 saline conditions 265–268

Abscisic acid (ABA) 9, 127, 201, 263Activated charcoal 155ADT. see Average daily temperature (ADT)AFLPs. see Amplified fragment length

polymorphisms (AFLPs)Agenda 21, 74Agrobacterium tumefaciens 202Agrobiodiversity conservation 74Aizoaceae (midday flowers) 128, 129Alba (R. alba) 95Altered phloem development (APL) 43Ambiphotoperiodic day plants 39Amplicons 244–245Amplified fragment length polymorphisms

(AFLPs) 163, 197Anthirrhinum majus 31Anthurium 159Apical dominance 21APL. see Altered phloem development (APL)AP-PCR. see Arbitrarily primed PCR (AP-PCR)Applause 209, 210Arabidopsis sp. 127

A. thaliana 130Arbitrarily primed PCR (AP-PCR) 163Argentina 246Ariocarpus fissuratus 103Asclepias curassavica 31

Automated irrigation systems 314, 318Autophagy 269–270Average daily temperature (ADT) 25Axillary meristems 21

Bacterial and pathogens 377Begonia sp.

B. gracilis 158 B. x hiemalis 40

Benzimidazole fungicides 394Benzyladenine 351Betula verrucosa J.F. Ehrh. tissue 29Biological controls 354–355Bio-pots 358Blossfeldia liliputiana 102Blue light 343Botanical Institute at Karlsruhe Institute of

Technology 92Botrytis species 355–356, 369, 392

diversity of 399–401 as endophyte 401–402 infection of ornamental geophytes

395–399BoWaS: Botrytis Warning System 393BRANCHED1 22Brassinosteroids 264Brazil 246Broadcast application, of fertilizer 294Bryophyllum 19Buddleja sp.

B.davidii 124 B.globosa 124

Buffering capacity 281Bulb mold 377Burbank, Luther 192, 194

Cactaceae 100Cactus Research Institute (CRI) 104–108Calonectria pseudonaviculata 135


Index416

CAM. see Crassulacean acid metabolism (CAM)

Camellia japonica var. decumbens 259Campanula sp.

case study 237–239 agrobacterium tumefaciens

transformation 243 expression cassette creation

242–243 gRNAs designing 241–242 molecular analysis 243–245 outcrossing 245 plant transformation and

regeneration 243 precise genome editing targets

239–241 C. portenschlagiana 241–245

CAM plant 102, 103Canada’s germplasm system 85Canada’s Plant Germplasm System 84Canadian Clonal Genebank 85, 87Canadian germplasm conservation 87Canadian Plant Germplasm System 86CAPS. see Cleaved amplified polymorphic

sequences (CAPS)Capsicum annuum 202Carbohydrate 154Carotenoid cleavage dioxygenase

(CmCCD4a) 217Cas9. see CRISPR-associated

endonuclease-9 (Cas9); CRISPR-associated protein 9 (Cas9)

CBD. see Convention on biological diversity (CBD)

Center for Genetic Resources (CGN), Netherlands 89

Central Experimental Farm in Ottawa 84Centrifolia (R. centrifolia) 95CFL. see Compact fluorescent luminaires

(CFL)CGN. see Center for Genetic Resources

(CGN), NetherlandsChelation 291Chile 246Chilling tolerance 259China 336Chlorophyll fluorescence (Fv/Fm) 131, 134Choline oxidase (CodA) 259Chromosome doubling 124Chrysanthemum sp. 158, 159

C. cinerariaefolium 161 C. x morifolium Ramat 29

Cleaved amplified polymorphic sequences (CAPS) 196

Clustered regularly interspaced short palindromic repeats (CRISPR) 231–235, 245

CmDREB6 gene 261Cmeil2 gene 240, 241CNTBio. see National Biosafety Technical

Commission (CNTBio)CodA. see Choline oxidase (CodA)Colchicine 124Compact fluorescent luminaires (CFL) 345Constitutive Triple Response factor 1

(CTR1) 238Controlled-release fertilizer (CRF) 285,

292–293Conventional breeding techniques

current common breeding techniques conventional cross breeding and

selection 120–121 interspecific and intergeneric

hybridization 121–123 polyploidization 123–126

future trends 138 overview of 119–120 for sustainable production

compact growing plants 136 drought tolerance 126–130 elevated pH tolerance 132–133 low temperature tolerance 130–132 nutrient use efficiency 135 postharvest life 136–137 resistance and tolerance to biotic

stressors 133–135Conventional freezers 83Convention on biological diversity

(CBD) 74Coreopsis sp.

C. grandiflora 4 C. grandiflorum 259

Correns, C.E. 191Crassulacean acid metabolism (CAM) 100CRF. see Controlled-release fertilizer (CRF)CRI. see Cactus Research Institute (CRI)CRISPR. see Clustered regularly interspaced

short palindromic repeats (CRISPR)CRISPR-associated endonuclease-9

(Cas9) 217–219CRISPR-associated protein 9 (Cas9)

233–235, 245Crop evapotranspiration approaches

316–318


Index 417

Crop wild relatives (CWRs) 91Cryogenic preservation techniques 84Cryopreservation 87, 160C. silverstrii 104CTR. see Constitutive Triple Response factor

1 (CTR1)Cucumis sp. 8, 25

C. anguria germination 8Cutting propagation 12CWRs. see Crop wild relatives (CWRs)Cyclamen persicum 158, 199, 202Cyclical lighting 345Cytokinins 21

DAF. see DNA amplification fingerprinting (DAF)

Dagmar Hastrup 259Daily crop water use (ETc) 317Daily light integral (DLI) 31, 347–348Damask (R. damasenca) 95DAPI-stained Delosperma 126Day-neutral plants (DNPs) 39Dendrobium 159Devernalization 45de Vries, Hugo Marie 191DFR. see Dihydroflavonol 4-reductaseDianthus caryophyllus 205Dihydroflavonol 4-reductase (DFR) 200,

210DIP. see Dropping temperatures (DIP)Diseases, affecting ornamental

geophytes 367–369 approaches to reduce fungicide

usage 392–394 biotic and abiotic 370–385 fungicide resistance management

394–395 general disease management

principles 385, 388–392 see also Botrytis species

Disease triangle concept 389Division 12DLI. see Daily light integral (DLI)DNA

cloning technology 79 markers 98 sequencing methods 159

DNA amplification fingerprinting (DAF) 163DNPs. see Day-neutral plants (DNPs)Donor explants 154Double stranded breaks (DSB) 232, 233Dracaena deremensis 155

Dropping temperatures (DIP) 23Drought

avoidance 127 escape 127 tolerance 127

DSB. see Double stranded breaks (DSB)Dutch botanic gardens 89Dutch Botanic Gardens Collections

Foundation 89–90Dutch National Plant Collection 89

Easily available water (EAW) 322, 324EAW. see Easily available water (EAW)Echeveria leaf cuttings 19ECPGR. see European Cooperative

Programme for Plant Genetic Resources (ECPGR)

E. leninghausii 104El Salvador 336Enrichment index 93Environmental physiology

asexual propagation 12, 14 cutting propagation 14–18 knowledge gaps 20–21 plant growth regulators and

rooting 19 rooting and moisture 18–19 rooting environment irradiance

15, 18 rooting photoperiod 15–18 rooting temperature 15 tissue culture (TC) 20

dormancy 49–50 increasing plant mass

carbon partitioning 27 knowledge gaps 27 photosynthesis 26–27

overview 3 reproductive development

autonomous flowering pathway 28–38

chemical promotion of flowering 48 juvenility vs. maturity 28 knowledge gaps 49 photoperiodic flowering pathway 33,

38–44 stress induction of flowering 47 temperature inhibition of

flowering 48–49 vernalization flowering pathway

44–47 sexual propagation 3–4


Index418

factors impacting seed germination 11

knowledge gaps 12 light and germination 5–8, 9–10 moisture and germination 5–8, 10 seed dormancy 11 seedling storage 11–13 seed storage 4 temperature and germination 4–8

vegetative development 21–22 knowledge gaps 26 leaf expansion 25 plant development rate 25–26 plant stem elongation 22–25

Epipactis 159Escallonia 124, 136Eschscholzia californica 39ESTs. see Expressed sequence tags (ESTs)ETc. see Daily crop water use (ETc)Ethephon 351–352Ethiopia 336Ethylene insensitivity 237–239Euphorbia Pulcherrima 158EURISCO. see European Plant Genetic

Resources Search Catalogue (EURISCO)

European Cooperative Programme for Plant Genetic Resources (ECPGR) 88

European Plant Genetic Resources Search Catalogue (EURISCO) 88, 89, 91

Eustoma grandiflora 202Expressed sequence tags (ESTs) 216

Facultative irradiance (FI) 31FAO. see Food and Agriculture Organization

(FAO)Far-red light 344Father of American Ornamental Breeding.

see Burbank, LutherFCM. see Flow cytometry (FCM)Federal Central Genebank of the Leibniz

Institute of Plant Genetics and Crop Plant Research (IPK) 91

FI. see Facultative irradiance (FI)Ficus benjamina 155Field production 288–289FLC. see Flowering locus C (FLC)Floral

evocation 28 induction 28

Floribunda rose 95 ‘Arunima’ 156

Florigene ‘Moon™’ carnation 205 ‘Moondust™’ 207

Flow cytometry (FCM) 125, 126Flower

development 28 dormancy 28 initiation 28

Flowering locus C (FLC) 47Flowering locus T (FT) 43Fludioxonil 392, 393Flutolanil 393Food and Agriculture Organization

(FAO) 85, 87FRAC. see Fungicide Resistance Action

Committee (FRAC)Fragaria chiloensis 86Fredericton Research Centre 85Freezing stress 257–258French Genetic Resources Office 93FT. see Flowering locus T (FT)FT-interacting protein 1 (FTIP1) 43Fungi 370Fungicide

application of 391 resistance management 355–356

Fungicide Resistance Action Committee (FRAC) 395

GA. see Gibberellin mutantsGallica (R. galllica) 95G. baladianum 104GBS. see Genotyping-by-sequencing (GBS)Gemini viruses, as vectors 235, 237Genebank for Tobacco 91Genebank Management System (GMS) 81Gene editing techniques 231–232

background 232–234 horticulture applications 237–245 ornamental plants prospective uses and

regulation 245–246 in plants, precise 234–237

Genetically engineered flowers (GEOs) 202–206

Genetic diversity exploitation in cacti, ornamental germplasm utilization

cacti-renowned ornamentals 101–102 distribution and diversity 101 overview of 100 in Republic of Korea 103–108 utilizing to cope with global climate

change 102–103


Index 419

ex situ conservation 75–76 botanical gardens 77–78 DNA storage 79 field genebanks 77 pollen storage 76–77 seed storage 76 in United Kingdom 92–93 in vitro conservation 77

ornamental germplasm management of 72–73

plant genetic resources management in Canada 84–88 in Europe 88–89 in France 93–94 in Germany 90–91 in Japan 81 in Korea 79–82 in Netherlands 89–90 reporting and monitoring system,

in situ conservation 91–92 in United States 82–84 wild grape Vitis vinifera L. ssp.

sylvestris C.C. Gmel., securing viability of 92

in roses, ornamental germplasm utilization

breeding aspect, important traits from 96–97

conventional breeding limitations 97–98

early genetic improvement 95–96 genetic modification (GM) 99–100 modern hybrids 96 molecular marker techniques 98–99 taxonomy and classification 94–95

in situ conservation methods 73–74 home gardens 75 on-farm conservation 74–75 protected areas 74

Genetic fingerprinting 194Genetic improvement, via genetic

transformation 162–163Genome-wide association studies

(GWAS) 215–217Genotypes 153Genotyping-by-sequencing (GBS) 216GEOs. see Genetically engineered flowers

(GEOs)German Genebank for Crop Wild

Relatives 91German Genebank for Fruit Crops 91German Genebank for Grapevine 91

German Genebank for Ornamentals 91Gibberellin (GA) mutants 22Gladiolus and Caladium 158Glochids 100GMS. see Genebank Management System

(GMS)Greenhouse production 287–288Green light 343–344Guide RNAs (gRNAs) 241–242GWAS. see Genome-wide association

studies (GWAS)Gymnocalycium mihannovichii 104

Hardy–Weinberg equilibrium 191Hatiora 101HDR. see Homology directed repair (HDR)Heat delay 40Heat shock genes (HSGs) 261Helianthus germination 11Helleborus 122Hibiscus syriacus

‘DVPAzurri’ 124 ‘Oiseau Bleu’ 124

High-pressure sodium (HPS) luminaire 337, 345

Homology directed repair (HDR) 232, 233HPS. see High-pressure sodium (HPS)

luminaireHSGs. see Heat shock genes (HSGs)Hybrid rose cv. Landora 157Hybrid tea rose 95Hydrangea 123

IBA. see Indole-3-butyric acid (IBA)IFGs. see Irrigation functional groups (IFGs)Impatiens walleriana 199Incandescent bulbs (INC) 344–345India 336Indole-3-butyric acid (IBA) 19, 352INKARHO® rootstocks 133Integrated pest management (IPM) 385,

388, 389International Plant Genetic Resources

Institute (IPGRI) 87International policy framework 74International Treaty on Plant Genetic

Resources for Food and Agriculture (ITPGRFA) 90, 92, 94

Inter-simple sequence repeat (ISSR) 98, 163Intra-canopy lighting 339In vitro

plant cell culture technique 151


Index420

plant cultivation 160 propagation 152–153

via meristem culture 150–151 via somatic embryogenesis 158

techniques 72IPGRI. see International Plant Genetic

Resources Institute (IPGRI)IPM. see Integrated pest management (IPM)Iris sp.

I. halophila 266 I. lactea var. chinensis 266

Irradiance 15 indifferent 31

Irrigation functional groups (IFGs) 317Irrigation practices and technology

305–306 data management and decision support

systems 319–320 precision accuracy, and

repeatability 320 sensor reliability and cost 320–321 transforming information into

knowledge 321–323 variability harnessing for reliability

improvement 321 variability sources 321

future trends 327–328 ornamental production systems and

water use 306–311 protocol case study 324–327 sensor-based technologies for irrigation

scheduling 314–318 system maintenance 313–314 systems design 312–313

ISSR. see Inter-simple sequence repeat (ISSR)ITPGRFA. see International Treaty on Plant

Genetic Resources for Food and Agriculture (ITPGRFA)

Jasmonic acid (JA) 135, 263, 264John Cabot 259

Kalanchoe blossfeldiana 25, 136K-IBA. see Potassium salt form of IBA (K-IBA)

Lampranthus (Aizoaceae) 129Lavandula sp.

L. angustifolia 124 L. × intermedia 124 L. latifolia 124

LDPs. see Long-day plants (LDPs)LEDs. see Light-emitting diodes (LEDs)

Leptonycteris curasoae 101Light-emitting diodes (LEDs) 10Lime tolerance 132, 133Liquid nitrogen 84Lobivia 104Lolium temulentum plants 43Long-day plants (LDPs) 39, 42

MAD. see Maximum allowable depletion (MAD)

Malaxis 159Malaysian oil palm 77Male germ units (MGU) 126Mammillaria 101Managing Irrigation and Nutrition through

Distributed Sensing (MINDS) 318Mancozeb 392Marker-assisted selection (MAS) 197Maryland Water Quality Improvement Act

(1998) 310MAS. see Marker-assisted selection (MAS)Maximum allowable depletion (MAD)

3241-MCP. see 1-Methylcyclopropene (1-MCP)Mendel, Gregor 1901-Methylcyclopropene (1-MCP) 252Methyltransferase (METI) 47MGU. see Male germ unitsmicroRNAs (miRNAs) 264Mildew resistance locus o (MLO) 135Millennium Seed Bank 93MINDS. see Managing Irrigation and

Nutrition through Distributed Sensing (MINDS)

miRNAs. see MicroRNAs (miRNAs)Mitotic polyploidization 123MLO. see Mildew resistance locus o (MLO)MLS. see Multilateral system (MLS)Molecular breeding advances

flower breeding, nineteenth and twentieth

centuries 191–194 color 198–201 as study organisms, early genetic

research 190–191 flowering genes 201 future trends 219 gene editing (CRISPR-Cas9) 217–218 genetically engineered flowers

(GEOs) 202–206 genetically modified carnations

205–209


Index 421

genetically modified chrysanthemums 211–213

genetically modified roses 208–211 genome sequencing 211–215

genome-wide association studies (GWAS) 215–217

marker-assisted breeding, biotechnology use 194–200

overview 189–190MORN-motif repeat protein regulating

flowering 1 (mrf1) 44Moss rose (R. centrifolia moscosa) 95mrf1. see MORN-motif repeat protein

regulating flowering 1 (mrf1)MtDREB1C gene 260Multi-crop passport descriptor list 91Multilateral system (MLS) 90Multi-nodal system 85, 87

NAA. see Napthalacetic acid (NAA)NAC. see National Agrobiodiversity

Center (NAC)Nannongxuefeng 265Napthalacetic acid (NAA) 19NARO. see National Agriculture and Food

Research Organization (NARO)NAS. see National Institute of Agricultural

Sciences (NAS)National Agriculture and Food Research

Organization (NARO) 81National Agrobiodiversity Center (NAC)

79, 80National Biosafety Technical Commission

(CNTBio) 246National Center for Genetic Resources

Preservation (NCGRP) 83National Horticulture and Herbal Research

Institute (NHHRI) 104–108National Institute of Agricultural Sciences

(NAS) 79, 80National Institute of Horticultural and Herbal

Science (NIHHS) 104National Plant Germplasm System

(NPGS) 82, 83National Resource Conservation Service

(NRCS) 310N. capa 104NCGRP. see National Center for Genetic

Resources Preservation (NCGRP)Nematodes 377, 384–385Neobuxbaumia tetetzo 101NHEJ. see Non-homologous end-joining

(NHEJ)

NHHRI. see National Horticulture and Herbal Research Institute (NHHRI)

NIAS Genebank 81Nicaragua 336Night interruption lighting integral 345–346NIHHS. see National Institute of Horticultural

and Herbal Science (NIHHS)1992 CBD 74Nitrogen

fertilizer 135 use efficiency 135

Non-homologous end-joining (NHEJ) 232, 233

Non-obligate-vernalization (NV) 201Nopalea species 102NPGS. see National Plant Germplasm System

(NPGS)NRCS. see National Resource Conservation

Service (NRCS)Numerous genera 84Nutrient management 279–280

fertilizer 291–293 delivery 293–295

future trends 296–297 irrigation and water quality 281–284 mineral nutrients 289–291 production system types 284–289 runoff environmental impacts 296 soils and substrates 295–296

NV. see Non-obligate-vernalization (NV)

Oomycetes 370Open air nursery 284–286OPGC. see Ornamental Plant Germplasm

Center (OPGC)Opuntia sp. 102

genera 101 O. ficus-indica 103

Opuntioid cacti 100Ornamental

breeders 128 horticulture 3

Ornamental Plant Germplasm Center (OPGC) 83, 84

Osmolytes 264Osteospermum fruticosum 137Overhead irrigation 307

interception efficiency 285

Pachycereus pringlei 102PAGE. see Polyacrylamide gel

electrophoresis (PAGE)Panther 259


Index422

Paraguay 246PAW. see Plant available water (PAW)PCR. see Polymerase chain reaction (PCR)Peat 356–357PeCHYR1 263Perlite 356Petunia 269–270Petunia hybrida 124PGRC. see Plant Gene Resources of Canada

(PGRC)PGRDEU 90, 91PGRs. see Plant genetic resources (PGRs);

Plant growth regulators (PGRs)Phalaenopsis sp. 159

P. equestris 215Phosphorus 290Photon capture 339Photoperiodic lighting 337, 344–346Photosynthetic photon efficiency (PPE)

337, 339Photosynthetic photon flux density

(PPFD) 337, 362pH tolerance 132Phytoplasmas 377Pistacia lentiscus 266Plant available water (PAW) 322Plant genebanks 72Plant Gene Resources of Canada

(PGRC) 84, 87 seed genebank 85

Plant genetic resources (PGRs) 72, 81, 89–91, 93, 94

Plant growth regulators (PGRs) 288, 326, 350–354

Plant Heritage’s National Plant Collections 93

Plant Heritage’s Threatened Plants Project 93

Plant Quarantine Act (1919–1926) 389Plant Wizard of Santa Rosa 194PlGPAT gene 260Polyacrylamide gel electrophoresis

(PAGE) 194Polymerase chain reaction (PCR) 79Polyploid 264

Lonicera japonica 130Positive displacement injectors 294Potassium 290Potassium salt form of IBA (K-IBA) 352Pot-in-pot production 286–287PPE. see Photosynthetic photon efficiency

(PPE)

PPFD. see Photosynthetic photon flux density (PPFD)

Prezygotic barriers 123Protected cultivation of ornamentals

335–337 future trends 360–362 light-emitting diode (LED) lighting

systems 337–346 plant growth and development

manipulation 346–347, 350–354 cutting production and propagation

systems 348–350 light management 347–348

sustainable practices and renewable inputs 354–359

in vertical propagation area 359–360

Qinglu 265QTL. see Quantitative trait loci (QTL)Quantitative trait loci (QTL) 131

mapping 99

Radio frequency identification technology (RFID) 361

Random amplified microsatellite polymorphisms (RAMPO) 195

Random fragment length polymorphisms (RFLPs) 196

Randomly amplified polymorphic DNA (RAPDs) 163, 196, 197

RAW. see Readily available water (RAW)RDA. see Rural Development Administration

(RDA)Readily available water (RAW) 322, 324Recycled tail-water 283Recycled water, from reservoirs 284Recycling plastics 358–359Red light 344Red Missile 259Relative water content (RWC) 129Renewable energy 357–358Renewable growing media 356Restriction fragment length polymorphisms

(RFLPs) 163RFID. see Radio frequency identification

technology (RFID)RFLPs. see Random fragment length

polymorphisms (RFLPs); Restriction fragment length polymorphisms (RFLPs)

RGSI. see Rose genome sequencing initiative (RGSI)


Index 423

Rhipsalis baccifera 101Rhizobium rhizogenes 136, 137Rhododendron 132, 155Rhynchostylis 159Ricinus communis 213Robotic-arm machines 350Rockwool 356Root cutting 15Rosa sp.

genus 94, 97 R. chinensis 99 R. × hybrida 95, 96, 98 R. multiflora 155 R. rugosa 259

Rose breeding technology 95Rose genome sequencing initiative

(RGSI) 99Rose rosette disease (RRD) 97Rose rosette virus (RRV) 97RRD. see Rose rosette disease (RRD)R. rouletii Correvon 96RRV. see Rose rosette virus (RRV)Rumex sp.

R. obtusifolius 8 R. retroflexus 8

Rural Development Administration (RDA) 79

RWC. see Relative water content (RWC)

SA. see Salicylic acid (SA)Saintpaulia ionantha cv. Benjamin 158Salicylic acid (SA) 263Salsa Yellow 259Sanitizers 390SAR. see Sodium adsorption ratio (SAR)Saskatoon Research Centre 85SbSKIP. see Sorghum bicolor Sloan-

Kettering retrovirus interacting protein (SbSKIP)

SCAR. see Sequence characterized amplified regions (SCAR)

Schlumbergera sp. 101 S. truncata 25

ScoT markers 98SDPs. see Short-day plants (SDPs)Sedum species 262Segmental allopolyploids 94Selenicereus grandifloras 102Sensorweb™ 317, 324Sequence characterized amplified regions

(SCAR) 98, 196

Sequence specific nucleases (SSN) 232Sequence-tagged sites (STSs) 163Sesuvium portulacastrum 266sgRNA. see Single-guide RNA (sgRNA)Short-day plants (SDPs) 39Shrub roses 86Side-dress applications 294Side sediment channel 309Single-guide RNA (sgRNA) 234, 235Single-strand conformation polymorphism

(SSCP) 196SL mutants 22Slow release fertilizer (SRF) 292, 293SLs. see Strigolactones (SLs)sMTA. see Standard material transfer

agreement (sMTA)Sodium adsorption ratio (SAR) 282Sodium hypochlorite (NaOCl) 154Soil-moisture approaches 314–316Soil pH 268Soleil d’Or 96Sole-source lighting 341–342Somatic polyploidization 123Sorghum bicolor Sloan-Kettering

retrovirus interacting protein (SbSKIP) 264

Spathiphyllum 124, 129, 155Spectral responses 342–344SQUAMOSA promoter binding-like (SPL)

transcription factors 33SRF. see Slow release fertilizer (SRF)SSCP. see Single-strand conformation

polymorphism (SSCP)SSN. see Sequence specific nucleases

(SSN)Standard material transfer agreement

(sMTA) 90Star Bright 259Sticking cuttings 349–350Stock plants 14Strigolactones (SLs) 21STSs. see Sequence-tagged sites (STSs)Succulent

Drosanthemum 128, 129 plants 262

Supplemental lighting 337–341Surface

disinfectants 154 sterilization 154 waters 283

Synthetic auxins 19


Index424

TALENs. see Transcription activator-like effector nucleases (TALENs)

TC. see Tissue culture (TC)Tender shoots 20Tensiometers 315Thermomorphogenesis 23Tissue culture (TC) 20

techniques 87 bioreactor/liquid culture 161 clonal propagation and

organogenesis 155–156 embryo rescue 156 factors affecting 153–155 future trends 165 molecular marker uses 163–165 overview of 149–153 plant improvement through genetic

engineering 161–163 slow-growth conservation

techniques 160–161 somatic embryogenesis 157–159 thin cell layer 156–157 use in mutagenesis and mutation

breeding 159–160Tolumnia-related genus, Oncidium 159Top-dressing fertilizers 294–295Transcription activator-like effector nucleases

(TALENs) 232Trifloxystrobin 393TSF. see Twin sister of FT (TSF)Tulips, field production of 369Twin sister of FT (TSF) 43

Urease inhibitors 292Uruguay 246

USDA-ARS. see U.S. Department of Agriculture-Agricultural Research Service (USDA-ARS)

USDA’s Cooperative State Research, Education, and Extension Service 83

USDA's State Agricultural Experiment Stations and Land Grant Universities 83

U.S. Department of Agriculture-Agricultural Research Service (USDA-ARS) 82

Venturi-type injectors 293–294Vernalization 44Vietnam 336Viruses 370, 372, 376Volumetric water content (VWC) 316,

321, 324

Water buffering capacity (WBC) 322Water molds. see OomycetesWater pH 281Water-soluble fertilizers 292WBC. see Water buffering capacity (WBC)Wetlands 80Whole genome sequencing (WGS) 211,

213Wood

fiber 356, 357 fuel 357–358

Woody Landscape Plant Germplasm Repository at the National Arboretum 83

Zinc-finger nucleases (ZFNs) 232

Documents

Contents...1 Introduction 149 2 Factors affecting tissue culture techniques 153 3 Tissue culture techniques 155 4 Future trends 165 5 References 165 5 Advances in molecular breeding