简介
Ferns existing today represent a genetic inheritance of great value as they include species of ancient vascular plants, which have direct connection with the crucial steps done in the past for settling life on Earth. Their life cycle is an example of alternation of generations, in which both a multicellular diploid organism and a multicellular haploid organism occur and give rise to the other. Spore, gametophyte and sporophyte have been studied either for basic and practical purposes. In the first case, they offer us simple and easy to grow experimental systems to investigate on plant development, and secondly, they are culture by their ornamental appeal, their environmental benefit or as source of metabolites. This timely volume brings a selection of chapters, each one composed by experts in their respective fields. The chapters included cover a broad range from the knowledge of its biology and contribution to understanding of plant development, useful protocols for propagation and conservation purposes, genetic variability as well as environmental and theurapeutical applications. This wide spectrum of the contributions gives to reader a rapid idea of the enormous potential of this plant group. The originality of this book is to expose the most recent tendencies in their investigation, which is far from the traditional perspective usually followed. The collected articles in this volume incorporate most of novel techniques used nowadays routinely to resolve traditional questions. Ashwani Kumar Professor, Department of Botany, University of Rajasthan, Jaipur. The author鈥檚 repertoire of published works spreads across 150 research articles in various national and international journals. He has an experience of over three decades in his field of research, namely, tissue culture and biochemistry, being awarded with prestigeous V. Puri Medal in 2008 for his services to advancement of Botany. Helena Fern谩ndez Associate Professor, Department of Biology of Organisms and Systems, Oviedo University, Spain. Her research focuses on micropropagation and reproduction in ferns during the two last decades. In 2002 she obtained the award 鈥淩am贸n y Cajal鈥?by The Ministry of Science and Technology, being engaged to the Oviedo University as tenurer, full time, in the Area of Plant Physiology since then. M. Angeles Revilla Plant Physioloy Assistant Professor in the Biology Faculty at the Oviedo University (Spain) since 1987. Twenty years experience in plant tissue culture. She has also worked in cryopreservation and genetic stability for the last ten years, mainly in the development of protocols for in vitro shoot apices in agronomic species.
目录
Preface 5
Preface 7
About the Book 9
Contents 11
Contributors 15
Chapter 1: Introduction 21
References 28
Part I: Contribution of Ferns to Understanding of Plant Development 29
Chapter 2: Cellular, Molecular, and Genetic Changes During the Development of Ceratopteris richardii Gametophytes 30
2.1 Introduction 30
2.2 Regulation of Differentiation in Multicellular Gametophytes 31
2.3 Genes and Genomic Studies of C. richardii Spores 34
2.4 Spores as a Tool for the Study of Cellular Gravity Response 36
2.5 Resources Available and on the Horizon for Use in C. richardii Research 40
References 41
Chapter 3: Laboratory-Induced Apogamy and Apospory in Ceratopteris richardii 44
3.1 Introduction 44
3.2 Alternation of Generations 44
3.3 Apogamy and Apospory in Ferns 47
3.4 Comparison of Apogamy and Apospory in Ferns with Apomixis in Angiosperms 48
3.5 Induction of Apogamy and Apospory in C. richardii 51
References 53
Chapter 4: Sexual Reproduction in Ferns 56
4.1 Introduction 56
4.2 Looking for New Experimental Systems 57
4.3 Sexual Reproduction in Pteridophyta 58
4.3.1 Antheridiogens 58
4.3.2 Plant Growth Regulators and Sex Determination in Blechnum spicant L 59
4.4 New Tools to Study the Molecular Basis of Sex Determination 61
4.5 Sexuality in the Laboratory and in Nature 64
References 65
Chapter 5: Gibberellic Acid and Ethylene Control Male Sex Determination and Development of Anemia phyllitidis Gametophytes 68
5.1 Main Aspects of A. phyllitidis Gametophyte Development 68
5.2 Spore Germination, Growth, and Differentiation of A. phyllitidis Gametophytes 70
5.3 The Antheridiogens \u2013 Epigenetic Aspects of Development of A. phyllitidis Gametophytes 72
5.4 Precocious, GA3-induced, Antheridia Formation and the \u201cThree-zonal Model\u201d of Structure and Function in Development of A. phyllitidis Gametophytes 73
5.5 ACCF and DDG Disturb GA3-Induced Antheridiogenesis and Development of A. phyllitidis Gametophytes 74
5.6 How Does Ethylene Participate in Development and Male Sex Determination in A. phyllitidis gametophytes? 75
5.7 Regulation of Antheridia Formation 76
5.8 Regulation of Cell Growth and Cell Division Cycleduring GA3-induced Antheridiogenesis 76
5.9 A. phyllitidis Gametophytes Produce ACC 79
5.10 Main Aspects of Antheridiogenesis 79
References 82
Chapter 6: The Sporophytes of Seed-Free Vascular Plants \u2013 Major Vegetative Developmental Features and Molecular Genetic Pathways 85
6.1 Introduction 85
6.2 Six Sporophyte Body Plans 87
6.3 Embryogeny 90
6.4 Apical Meristem Structure 92
6.5 Branching 93
6.6 Radial Patterning of Sporophyte Axes 95
6.7 Leaf Development 97
6.8 Developmental Genes 100
6.8.1 KNOX and ARP Genes 102
6.8.2 HD-ZIP Genes 104
6.8.3 MIKC-type MADS-box Genes 105
6.8.4 AP2 and ANT Genes 106
6.8.5 FLORICAULA/LEAFY 106
6.8.6 Other Genes 106
6.8.7 MicroRNA Regulation of Genes 107
6.9 Conclusion 108
References 109
Part II: Propagation, Conservation and Control of Genetic Variability in Ferns 113
Chapter 7: From Spore to Sporophyte: How to Proceed In Vitro 114
7.1 Introduction 114
7.2 Spore Culture 115
7.3 Gametophyte Culture: Nutritional and Environmental Conditions 118
7.4 Gametophyte Multiplication 119
7.4.1 By Natural Means 119
7.4.2 Homogenised Cultures 119
7.5 Sporophyte Formation 120
7.5.1 Sexual Reproduction 120
7.5.2 Apogamy 121
7.6 Sporophyte Multiplication 121
References 124
Chapter 8: In Vitro Regeneration Systems of Platycerium 128
8.1 Introduction 128
8.2 In Vitro Regeneration Systems of Sporophytic Tissue 130
8.2.1 Direct Shoot Organogenesis 131
8.2.1.1 Shoot and Rhizome Culture 131
8.2.1.2 Leaf Culture 133
8.2.1.3 Culture of Bud Scales 134
8.2.1.4 Homogenization of Sporophytes 136
8.2.2 Indirect Shoot Organogenesis 136
8.2.3 Apospory and Apogamy 137
8.3 In Vitro Cultures of Platycerium in Developmental and Physiological Studies 138
8.4 Conclusion 139
References 140
Chapter 9: Stipule Propagation in Five Marattioid Species Native to Taiwan (Marattiaceae; Pteridophyte) 143
9.1 Introduction 143
9.2 Materials and Methods 144
9.3 Results 145
9.4 Discussion 148
References 150
Chapter 10: Tree Ferns Biotechnology: From Spores to Sporophytes 151
10.1 Introduction 151
10.2 Media Most Often Used 153
10.3 Plant Growth Hormones 154
10.4 Origin of Spore 154
10.5 Spore Sterilization 154
10.6 Spore Germination 155
10.7 Gametophyte Growth and Development 156
10.8 Gametophyte Multiplication 157
10.9 Sexual Determination of Gametophyte 158
10.10 Sporophyte Production 159
10.11 Conclusion 160
References 161
Chapter 11: In Vitro Propagation of Rare and Endangered Serpentine Fern Species 164
11.1 Introduction 164
11.2 Materials and Methods 165
11.2.1 Initiation of Culture 165
11.2.2 Growth of Gametophytes 166
11.2.3 Gene Bank of Prothalli 166
11.2.4 Culture of Sporophytes 167
11.2.5 Regeneration of Shoot Buds 167
11.2.6 Acclimation of Plants Ex Vitro 167
11.2.7 Measurements of Stomata Cells and Spores Size 167
11.3 Results and Discussion 168
11.3.1 Sporophyte Formation 172
11.3.2 Sporophytes Multiplication 172
11.3.3 Regeneration from Sporophytes 174
11.3.4 Plant Preparation Before Transfer to Greenhouse 175
11.3.5 Comparison of Plants from Natural Stands and Originated In Vitro 176
References 178
Chapter 12: Conservation of Fern Spores 180
12.1 Introduction 180
12.2 Background: Fern Spore Longevity and Storage 180
12.3 Water Content of Fern Spores and Storage Stability 181
12.4 Fern Spores: Orthodox or Recalcitrant Storage Behavior? 183
12.5 Conclusions 184
References 184
Chapter 13: Exploration of Cryo-methods to Preserve Tree and Herbaceous Fern Gametophytes 188
13.1 Introduction 188
13.1.1 Fern Gametophytes as Germplasm for Genebanks 188
13.1.2 Current Status of Gametophyte Cryopreservation 189
13.1.3 Brief Summary of Habitat for Fern Species in This Study 190
13.2 Materials and Methods 192
13.2.1 Plant Material 192
13.2.2 Cryoprotection Procedures 193
13.2.3 Thawing of Gametophytes 193
13.2.4 Survival Assessments 194
13.3 Results 194
13.3.1 Gametophyte Survival Following Various Cryo-methods 194
13.3.2 Preculture Effects on Detectable Damage 196
13.3.3 Recovery of Gametophyte in Culture and Sporophyte Production 198
13.4 Discussion 200
13.4.1 Cryoprotection and Gametophyte Survival 202
13.4.2 Autofluorescence as a Simple Methods Assessment of Gametophyte Viability 203
13.4.3 Cellular Damage to Gametophytes and Totipotency 203
13.4.4 Desiccation and Cold Tolerance and Cryopreservability 204
13.5 Conclusions 204
References 205
Chapter 14: Pteridophyte Spores Viability 208
14.1 Concept of Spore Viability \u2013 Biological Importance 208
14.2 Viability Variation Among Species 210
14.3 Factors Affecting Viability 210
14.3.1 Genotype 211
14.3.2 Age 211
14.3.3 Temperature 213
14.4 Physiology of Fern Spores Viability 214
14.5 Viability Detection Techniques 215
14.6 Viability and Conservation 217
References 217
Chapter 15: Microsatellites: A Powerful Genetic Marker for Fern Research 221
15.1 Genetic Molecular Markers 221
15.2 Microsatellites 223
15.2.1 What and Where Are They? 223
15.2.2 How to Use Them? 225
15.3 Microsatellites in Ferns 227
15.3.1 Previous Microsatellite Works on Ferns 227
15.3.2 A Particular Study Case: Dryopteris aemula 228
15.4 Advantages and Disadvantages of Microsatellites for Fern Studies 229
15.4.1 Why to Use microsatellites? 229
15.4.2 Problems and Limitations 230
15.5 Future Prospects 231
15.5.1 Potential Applications 231
15.5.2 Next-Generation DNA Sequencing 231
15.6 Conclusion 231
References 232
Chapter 16: Diversity in Natural Fern Populations: Dominant Markers as Genetic Tools 235
16.1 Introduction 235
16.2 Why Choose a Dominant Marker? 236
16.3 Measuring Molecular Diversity in Complex Genomes 237
16.4 Critical Steps in AFLP Protocols 239
16.5 Measure of Genetic Diversity Using Dominant Markers in Natural Fern Populations 240
16.6 Perspectives: Dominant Markers as an Effective Way to Develop Specific Markers 244
16.7 Conclusion 245
References 246
Part III: Environmental Biotechnology:Ecotoxicology and Bioremediation in Ferns 249
Chapter 17: Mitochondrial Activity of Fern Spores for the Evaluation of Acute Toxicity in Higher Plant Development 250
17.1 Introduction 250
17.2 Characteristics of Toxicity Tests 251
17.3 Current Challenges of Ecotoxicology 251
17.4 Ferns and Ecotoxicology 252
17.5 Candidate Ferns for Toxicity Testing 255
17.6 Application of Fern Spores to Toxicity Testing 257
17.7 Conclusion 259
References 260
Chapter 18: Chronic Phytotoxicity in Gametophytes: DNA as Biomarker of Growth and Chlorophyll Autofluorescence as Biomarker of Cell Function 261
18.1 Introduction 261
18.2 Current State of Phytotoxicity Testing 262
18.3 Limitations of Traditional Acute Toxicity Tests 263
18.4 Limitations of Traditional Terrestrial Plant Toxicity Tests 264
18.5 Usefulness of Ferns in Phytotoxicity Testing 264
18.6 DNA as a Measurement of Cellular Proliferation 265
18.7 Chlorophyll a Autofluorescence as Surrogate of Plant Physiological State 266
18.8 Applications of the Bioassay of Chronic Toxicity Based on Fern Spores 267
18.9 Protocols 269
18.9.1 Toxicity Tests 269
18.9.1.1 Plant Material 269
18.9.1.2 Chronic Toxicity Bioassay 270
DNA Quantification 270
Chlorophyll Quantification 270
18.10 Conclusions 270
References 271
Chapter 19: Arsenic Hyperaccumulator Fern Pteris vittata: Utilities for Arsenic Phytoremediation and Plant Biotechnology 273
19.1 Arsenic \u2013 A Toxic Metalloid Widespread in the Environment 273
19.2 Discovery of Arsenic Hyperaccumulation Trait in Ferns 274
19.3 Biological Roles for Arsenic Hyperaccumulation 275
19.4 Arsenate Uptake 276
19.5 Arsenate Reduction 276
19.6 Arsenite Uptake 277
19.7 Arsenic Transport within the Plant 277
19.8 Oxidative Stress Tolerance 277
19.9 Utilities of Brake Fern for Phytoremediation 278
19.10 Brake Fern as a Source of Genes for Biotechnology 278
References 278
Chapter 20: Aerobiology of Pteridophyta Spores: Preliminary Results and Applications 282
20.1 Introduction 282
20.2 Methodology in Aerobiology Studies 283
20.3 Aerobiological Notes of Pteridophyta 283
20.4 Perspectives and Applications 289
References 290
Part IV: Therapeutical/Medicinal Applications 293
Chapter 21: Studies on Folk Medicinal Fern: An Example of \u201cGu-Sui-Bu\u201d 294
21.1 Introduction 294
21.2 Comparative Analysis of Different Phenolic Compounds and Their Activities in Six Different \u201cGu-Sui-Bu\u201d 296
21.2.1 Antioxidant Activity 296
21.2.2 Antioxidant Activity by ABTS Assay 296
21.2.3 Antioxidant Activity by Dot-Blot and DPPH Staining 296
21.2.4 Determination of \u201cReducing Power\u201d 298
21.2.5 Scavenging Activity Against DPPH Radical 299
21.2.6 Total Polyphenols Content (Flavonoids, Flavonols, Condensed Tannin, and Proanthocyanidin) Determination in \u201cGu-Sui-Bu\u201d 300
21.2.6.1 Flavonoids Content 302
21.2.6.2 Flavonols Content 302
21.2.6.3 Condensed Tannins/Proanthocyanidin Content 302
21.3 In Vitro Studies on D. fortunei 303
21.3.1 Effects of MS Basal Medium, Sucrose Concentration, and Sugars on Germination of Spores and Development of Gametophytes 303
21.3.2 Effects of pH on Spore Germination, Development of Gametophytes, and Reproductive Organs 304
21.3.3 Effects of Light Spectra on Spore Germination, Development of Gametophytes, and Reproductive Organs 305
21.4 Conclusion 310
References 311
Chapter 22: Ecdysteroids in Ferns: Distribution, Diversity, Biosynthesis, and Functions 314
22.1 Introduction 314
22.2 Distribution and Diversity 314
22.3 Biosynthesis 319
22.4 Regulation of Ecdysteroid Biosynthesisand Accumulation 322
22.5 Function(s) 323
References 324
Chapter 23: Ferns: From Traditional Uses to Pharmaceutical Development, Chemical Identification of Active Principles 329
23.1 Introduction 329
23.2 Traditional Medicine Uses of Ferns 329
23.3 Fern Bioactive Components 338
23.3.1 Terpenoids 339
23.3.2 Triterpenoids 339
23.3.3 Diterpenoids 339
23.3.4 Sesquiterpenoids 344
23.3.5 Phenolic Compounds 346
23.3.6 Flavonoids 347
23.3.7 Alkaloids 347
23.4 Pharmaceutical Development 348
23.4.1 Huperzine A 348
23.4.2 Pharmaceutical Preparations from Polypodium leucotomos 349
References 349
Chapter 24: Functional Activities of Ferns for Human Health 355
24.1 Introduction 355
24.2 Natural Antioxidant 356
24.3 Foods 359
24.4 Natural Antimicrobial Agents 360
24.5 Cosmetic Ingredient 362
24.6 Air Purifier 362
24.7 The Future of Ferns and Fern Allies 363
References 364
Chapter 25: Toxicological and Medicinal Aspects of the Most Frequent Fern Species, Pteridium aquilinum (L.) Kuhn 368
25.1 Introduction 368
25.1.1 Chemical Aspects of the Plant 369
25.2 Toxicological Aspects 369
25.2.1 Compounds of Anti-thiamine Character 369
25.2.2 Cyanogen Glycosides 370
25.2.3 Compounds of Illudane Skeleton 370
25.2.3.1 Ptaquiloside 370
25.2.3.2 Pterosins 372
25.2.3.3 Other Chemical Components 372
25.3 Effects on Animals 373
25.3.1 Non-ruminant Animals 373
25.3.1.1 Swine 373
25.3.1.2 Rabbit 373
25.3.1.3 Guinea Pig 374
25.3.1.4 Rat and Mice 374
25.3.2 Ruminant Animals 374
25.3.2.1 Cattle 374
The Bovine Enzootic Haemature 374
Haemorrhagic Syndromes (Acute Haemorrhagic Disease) 375
Upper Alimentary Carcinoma 375
25.3.2.2 Sheep 376
25.4 Effects in Humans 376
25.5 Working Mechanism of Ptaquiloside 376
25.6 Presence of Ptaquiloside in Soil and Water 377
25.7 Medicinal Uses of P. aquilinum 378
25.8 Conclusions 378
References 379
Index 383
Preface 7
About the Book 9
Contents 11
Contributors 15
Chapter 1: Introduction 21
References 28
Part I: Contribution of Ferns to Understanding of Plant Development 29
Chapter 2: Cellular, Molecular, and Genetic Changes During the Development of Ceratopteris richardii Gametophytes 30
2.1 Introduction 30
2.2 Regulation of Differentiation in Multicellular Gametophytes 31
2.3 Genes and Genomic Studies of C. richardii Spores 34
2.4 Spores as a Tool for the Study of Cellular Gravity Response 36
2.5 Resources Available and on the Horizon for Use in C. richardii Research 40
References 41
Chapter 3: Laboratory-Induced Apogamy and Apospory in Ceratopteris richardii 44
3.1 Introduction 44
3.2 Alternation of Generations 44
3.3 Apogamy and Apospory in Ferns 47
3.4 Comparison of Apogamy and Apospory in Ferns with Apomixis in Angiosperms 48
3.5 Induction of Apogamy and Apospory in C. richardii 51
References 53
Chapter 4: Sexual Reproduction in Ferns 56
4.1 Introduction 56
4.2 Looking for New Experimental Systems 57
4.3 Sexual Reproduction in Pteridophyta 58
4.3.1 Antheridiogens 58
4.3.2 Plant Growth Regulators and Sex Determination in Blechnum spicant L 59
4.4 New Tools to Study the Molecular Basis of Sex Determination 61
4.5 Sexuality in the Laboratory and in Nature 64
References 65
Chapter 5: Gibberellic Acid and Ethylene Control Male Sex Determination and Development of Anemia phyllitidis Gametophytes 68
5.1 Main Aspects of A. phyllitidis Gametophyte Development 68
5.2 Spore Germination, Growth, and Differentiation of A. phyllitidis Gametophytes 70
5.3 The Antheridiogens \u2013 Epigenetic Aspects of Development of A. phyllitidis Gametophytes 72
5.4 Precocious, GA3-induced, Antheridia Formation and the \u201cThree-zonal Model\u201d of Structure and Function in Development of A. phyllitidis Gametophytes 73
5.5 ACCF and DDG Disturb GA3-Induced Antheridiogenesis and Development of A. phyllitidis Gametophytes 74
5.6 How Does Ethylene Participate in Development and Male Sex Determination in A. phyllitidis gametophytes? 75
5.7 Regulation of Antheridia Formation 76
5.8 Regulation of Cell Growth and Cell Division Cycleduring GA3-induced Antheridiogenesis 76
5.9 A. phyllitidis Gametophytes Produce ACC 79
5.10 Main Aspects of Antheridiogenesis 79
References 82
Chapter 6: The Sporophytes of Seed-Free Vascular Plants \u2013 Major Vegetative Developmental Features and Molecular Genetic Pathways 85
6.1 Introduction 85
6.2 Six Sporophyte Body Plans 87
6.3 Embryogeny 90
6.4 Apical Meristem Structure 92
6.5 Branching 93
6.6 Radial Patterning of Sporophyte Axes 95
6.7 Leaf Development 97
6.8 Developmental Genes 100
6.8.1 KNOX and ARP Genes 102
6.8.2 HD-ZIP Genes 104
6.8.3 MIKC-type MADS-box Genes 105
6.8.4 AP2 and ANT Genes 106
6.8.5 FLORICAULA/LEAFY 106
6.8.6 Other Genes 106
6.8.7 MicroRNA Regulation of Genes 107
6.9 Conclusion 108
References 109
Part II: Propagation, Conservation and Control of Genetic Variability in Ferns 113
Chapter 7: From Spore to Sporophyte: How to Proceed In Vitro 114
7.1 Introduction 114
7.2 Spore Culture 115
7.3 Gametophyte Culture: Nutritional and Environmental Conditions 118
7.4 Gametophyte Multiplication 119
7.4.1 By Natural Means 119
7.4.2 Homogenised Cultures 119
7.5 Sporophyte Formation 120
7.5.1 Sexual Reproduction 120
7.5.2 Apogamy 121
7.6 Sporophyte Multiplication 121
References 124
Chapter 8: In Vitro Regeneration Systems of Platycerium 128
8.1 Introduction 128
8.2 In Vitro Regeneration Systems of Sporophytic Tissue 130
8.2.1 Direct Shoot Organogenesis 131
8.2.1.1 Shoot and Rhizome Culture 131
8.2.1.2 Leaf Culture 133
8.2.1.3 Culture of Bud Scales 134
8.2.1.4 Homogenization of Sporophytes 136
8.2.2 Indirect Shoot Organogenesis 136
8.2.3 Apospory and Apogamy 137
8.3 In Vitro Cultures of Platycerium in Developmental and Physiological Studies 138
8.4 Conclusion 139
References 140
Chapter 9: Stipule Propagation in Five Marattioid Species Native to Taiwan (Marattiaceae; Pteridophyte) 143
9.1 Introduction 143
9.2 Materials and Methods 144
9.3 Results 145
9.4 Discussion 148
References 150
Chapter 10: Tree Ferns Biotechnology: From Spores to Sporophytes 151
10.1 Introduction 151
10.2 Media Most Often Used 153
10.3 Plant Growth Hormones 154
10.4 Origin of Spore 154
10.5 Spore Sterilization 154
10.6 Spore Germination 155
10.7 Gametophyte Growth and Development 156
10.8 Gametophyte Multiplication 157
10.9 Sexual Determination of Gametophyte 158
10.10 Sporophyte Production 159
10.11 Conclusion 160
References 161
Chapter 11: In Vitro Propagation of Rare and Endangered Serpentine Fern Species 164
11.1 Introduction 164
11.2 Materials and Methods 165
11.2.1 Initiation of Culture 165
11.2.2 Growth of Gametophytes 166
11.2.3 Gene Bank of Prothalli 166
11.2.4 Culture of Sporophytes 167
11.2.5 Regeneration of Shoot Buds 167
11.2.6 Acclimation of Plants Ex Vitro 167
11.2.7 Measurements of Stomata Cells and Spores Size 167
11.3 Results and Discussion 168
11.3.1 Sporophyte Formation 172
11.3.2 Sporophytes Multiplication 172
11.3.3 Regeneration from Sporophytes 174
11.3.4 Plant Preparation Before Transfer to Greenhouse 175
11.3.5 Comparison of Plants from Natural Stands and Originated In Vitro 176
References 178
Chapter 12: Conservation of Fern Spores 180
12.1 Introduction 180
12.2 Background: Fern Spore Longevity and Storage 180
12.3 Water Content of Fern Spores and Storage Stability 181
12.4 Fern Spores: Orthodox or Recalcitrant Storage Behavior? 183
12.5 Conclusions 184
References 184
Chapter 13: Exploration of Cryo-methods to Preserve Tree and Herbaceous Fern Gametophytes 188
13.1 Introduction 188
13.1.1 Fern Gametophytes as Germplasm for Genebanks 188
13.1.2 Current Status of Gametophyte Cryopreservation 189
13.1.3 Brief Summary of Habitat for Fern Species in This Study 190
13.2 Materials and Methods 192
13.2.1 Plant Material 192
13.2.2 Cryoprotection Procedures 193
13.2.3 Thawing of Gametophytes 193
13.2.4 Survival Assessments 194
13.3 Results 194
13.3.1 Gametophyte Survival Following Various Cryo-methods 194
13.3.2 Preculture Effects on Detectable Damage 196
13.3.3 Recovery of Gametophyte in Culture and Sporophyte Production 198
13.4 Discussion 200
13.4.1 Cryoprotection and Gametophyte Survival 202
13.4.2 Autofluorescence as a Simple Methods Assessment of Gametophyte Viability 203
13.4.3 Cellular Damage to Gametophytes and Totipotency 203
13.4.4 Desiccation and Cold Tolerance and Cryopreservability 204
13.5 Conclusions 204
References 205
Chapter 14: Pteridophyte Spores Viability 208
14.1 Concept of Spore Viability \u2013 Biological Importance 208
14.2 Viability Variation Among Species 210
14.3 Factors Affecting Viability 210
14.3.1 Genotype 211
14.3.2 Age 211
14.3.3 Temperature 213
14.4 Physiology of Fern Spores Viability 214
14.5 Viability Detection Techniques 215
14.6 Viability and Conservation 217
References 217
Chapter 15: Microsatellites: A Powerful Genetic Marker for Fern Research 221
15.1 Genetic Molecular Markers 221
15.2 Microsatellites 223
15.2.1 What and Where Are They? 223
15.2.2 How to Use Them? 225
15.3 Microsatellites in Ferns 227
15.3.1 Previous Microsatellite Works on Ferns 227
15.3.2 A Particular Study Case: Dryopteris aemula 228
15.4 Advantages and Disadvantages of Microsatellites for Fern Studies 229
15.4.1 Why to Use microsatellites? 229
15.4.2 Problems and Limitations 230
15.5 Future Prospects 231
15.5.1 Potential Applications 231
15.5.2 Next-Generation DNA Sequencing 231
15.6 Conclusion 231
References 232
Chapter 16: Diversity in Natural Fern Populations: Dominant Markers as Genetic Tools 235
16.1 Introduction 235
16.2 Why Choose a Dominant Marker? 236
16.3 Measuring Molecular Diversity in Complex Genomes 237
16.4 Critical Steps in AFLP Protocols 239
16.5 Measure of Genetic Diversity Using Dominant Markers in Natural Fern Populations 240
16.6 Perspectives: Dominant Markers as an Effective Way to Develop Specific Markers 244
16.7 Conclusion 245
References 246
Part III: Environmental Biotechnology:Ecotoxicology and Bioremediation in Ferns 249
Chapter 17: Mitochondrial Activity of Fern Spores for the Evaluation of Acute Toxicity in Higher Plant Development 250
17.1 Introduction 250
17.2 Characteristics of Toxicity Tests 251
17.3 Current Challenges of Ecotoxicology 251
17.4 Ferns and Ecotoxicology 252
17.5 Candidate Ferns for Toxicity Testing 255
17.6 Application of Fern Spores to Toxicity Testing 257
17.7 Conclusion 259
References 260
Chapter 18: Chronic Phytotoxicity in Gametophytes: DNA as Biomarker of Growth and Chlorophyll Autofluorescence as Biomarker of Cell Function 261
18.1 Introduction 261
18.2 Current State of Phytotoxicity Testing 262
18.3 Limitations of Traditional Acute Toxicity Tests 263
18.4 Limitations of Traditional Terrestrial Plant Toxicity Tests 264
18.5 Usefulness of Ferns in Phytotoxicity Testing 264
18.6 DNA as a Measurement of Cellular Proliferation 265
18.7 Chlorophyll a Autofluorescence as Surrogate of Plant Physiological State 266
18.8 Applications of the Bioassay of Chronic Toxicity Based on Fern Spores 267
18.9 Protocols 269
18.9.1 Toxicity Tests 269
18.9.1.1 Plant Material 269
18.9.1.2 Chronic Toxicity Bioassay 270
DNA Quantification 270
Chlorophyll Quantification 270
18.10 Conclusions 270
References 271
Chapter 19: Arsenic Hyperaccumulator Fern Pteris vittata: Utilities for Arsenic Phytoremediation and Plant Biotechnology 273
19.1 Arsenic \u2013 A Toxic Metalloid Widespread in the Environment 273
19.2 Discovery of Arsenic Hyperaccumulation Trait in Ferns 274
19.3 Biological Roles for Arsenic Hyperaccumulation 275
19.4 Arsenate Uptake 276
19.5 Arsenate Reduction 276
19.6 Arsenite Uptake 277
19.7 Arsenic Transport within the Plant 277
19.8 Oxidative Stress Tolerance 277
19.9 Utilities of Brake Fern for Phytoremediation 278
19.10 Brake Fern as a Source of Genes for Biotechnology 278
References 278
Chapter 20: Aerobiology of Pteridophyta Spores: Preliminary Results and Applications 282
20.1 Introduction 282
20.2 Methodology in Aerobiology Studies 283
20.3 Aerobiological Notes of Pteridophyta 283
20.4 Perspectives and Applications 289
References 290
Part IV: Therapeutical/Medicinal Applications 293
Chapter 21: Studies on Folk Medicinal Fern: An Example of \u201cGu-Sui-Bu\u201d 294
21.1 Introduction 294
21.2 Comparative Analysis of Different Phenolic Compounds and Their Activities in Six Different \u201cGu-Sui-Bu\u201d 296
21.2.1 Antioxidant Activity 296
21.2.2 Antioxidant Activity by ABTS Assay 296
21.2.3 Antioxidant Activity by Dot-Blot and DPPH Staining 296
21.2.4 Determination of \u201cReducing Power\u201d 298
21.2.5 Scavenging Activity Against DPPH Radical 299
21.2.6 Total Polyphenols Content (Flavonoids, Flavonols, Condensed Tannin, and Proanthocyanidin) Determination in \u201cGu-Sui-Bu\u201d 300
21.2.6.1 Flavonoids Content 302
21.2.6.2 Flavonols Content 302
21.2.6.3 Condensed Tannins/Proanthocyanidin Content 302
21.3 In Vitro Studies on D. fortunei 303
21.3.1 Effects of MS Basal Medium, Sucrose Concentration, and Sugars on Germination of Spores and Development of Gametophytes 303
21.3.2 Effects of pH on Spore Germination, Development of Gametophytes, and Reproductive Organs 304
21.3.3 Effects of Light Spectra on Spore Germination, Development of Gametophytes, and Reproductive Organs 305
21.4 Conclusion 310
References 311
Chapter 22: Ecdysteroids in Ferns: Distribution, Diversity, Biosynthesis, and Functions 314
22.1 Introduction 314
22.2 Distribution and Diversity 314
22.3 Biosynthesis 319
22.4 Regulation of Ecdysteroid Biosynthesisand Accumulation 322
22.5 Function(s) 323
References 324
Chapter 23: Ferns: From Traditional Uses to Pharmaceutical Development, Chemical Identification of Active Principles 329
23.1 Introduction 329
23.2 Traditional Medicine Uses of Ferns 329
23.3 Fern Bioactive Components 338
23.3.1 Terpenoids 339
23.3.2 Triterpenoids 339
23.3.3 Diterpenoids 339
23.3.4 Sesquiterpenoids 344
23.3.5 Phenolic Compounds 346
23.3.6 Flavonoids 347
23.3.7 Alkaloids 347
23.4 Pharmaceutical Development 348
23.4.1 Huperzine A 348
23.4.2 Pharmaceutical Preparations from Polypodium leucotomos 349
References 349
Chapter 24: Functional Activities of Ferns for Human Health 355
24.1 Introduction 355
24.2 Natural Antioxidant 356
24.3 Foods 359
24.4 Natural Antimicrobial Agents 360
24.5 Cosmetic Ingredient 362
24.6 Air Purifier 362
24.7 The Future of Ferns and Fern Allies 363
References 364
Chapter 25: Toxicological and Medicinal Aspects of the Most Frequent Fern Species, Pteridium aquilinum (L.) Kuhn 368
25.1 Introduction 368
25.1.1 Chemical Aspects of the Plant 369
25.2 Toxicological Aspects 369
25.2.1 Compounds of Anti-thiamine Character 369
25.2.2 Cyanogen Glycosides 370
25.2.3 Compounds of Illudane Skeleton 370
25.2.3.1 Ptaquiloside 370
25.2.3.2 Pterosins 372
25.2.3.3 Other Chemical Components 372
25.3 Effects on Animals 373
25.3.1 Non-ruminant Animals 373
25.3.1.1 Swine 373
25.3.1.2 Rabbit 373
25.3.1.3 Guinea Pig 374
25.3.1.4 Rat and Mice 374
25.3.2 Ruminant Animals 374
25.3.2.1 Cattle 374
The Bovine Enzootic Haemature 374
Haemorrhagic Syndromes (Acute Haemorrhagic Disease) 375
Upper Alimentary Carcinoma 375
25.3.2.2 Sheep 376
25.4 Effects in Humans 376
25.5 Working Mechanism of Ptaquiloside 376
25.6 Presence of Ptaquiloside in Soil and Water 377
25.7 Medicinal Uses of P. aquilinum 378
25.8 Conclusions 378
References 379
Index 383
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