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Index
Foreword
Preface
Contents
About the Editors
1 Classification of Cannabis sativa L. in Relation to Agricultural, Biotechnological, Medical and Recreational Utilization
Abstract
1.1 Introduction
1.2 Legal Classification
1.3 Pharmacological Classification
1.4 Folk Classification: “Hemp” Versus “Marijuana”
1.5 Ancient Phytogeography
1.6 Four Utilitarian Classes of Cannabis
1.6.1 Wild Plants
1.6.1.1 “Ditchweed”
1.6.1.2 Primitive Versus Secondary (Ruderal) Wildness
1.6.1.3 Adaptive Morphological and Anatomical Differences Between Wild and Domesticated Cannabis sativa
1.6.1.4 Adaptive Physiological Differences Between Wild and Domesticated Plants
1.6.2 Fiber Plants
1.6.2.1 Historical Review
1.6.2.2 Architecture and Anatomy
1.6.2.3 Physiology
1.6.2.4 Cannabinoid Profile
1.6.2.5 Economic Status and Potential
1.6.3 Oilseed Plants
1.6.3.1 Historical Review
1.6.3.2 Architecture and Anatomy
1.6.3.3 Physiology
1.6.3.4 Cannabinoid Profile
1.6.3.5 Economic Status and Potential
1.6.4 Intoxicant Plants
1.6.4.1 “Marijuana”
1.6.4.2 “Bracts”
1.6.4.3 “Buds”
1.6.4.4 Historical Review
1.6.4.5 Architecture and Anatomy
1.6.4.6 Physiology
1.6.4.7 Cannabinoid Profile
1.6.4.8 Economic Status and Potential
1.7 Genetic Groupings in Cannabis
1.8 Gene Flow Among Domesticates and Ruderal Populations
1.9 Sources of Taxonomic Disagreement with Particular Reference to Cannabis
1.9.1 Classification Difficulties Due to Hybridization and Typological Thinking
1.9.2 Taxonomic Splitting and Rank Inflation
1.9.3 Classification Difficulties Due to Obliteration of Populations by Humans
1.10 Classification of Domesticated Plants with Special Reference to Cannabis
1.10.1 Defining “Domestication”
1.10.2 Taxonomic Difficulties with Marijuana Strain Names
1.10.3 Why Classifying Domesticated Plants like Wild Plants Is Misleading
1.10.4 Possible Relationships of Wild and Domesticated Plants
1.11 Alternative Classification Systems for Cannabis
1.11.1 Classification of Cannabis Assemblages as Conventional Taxa
1.11.2 Classification of Cannabis Assemblages as “Groups” Under the Cultivated Plant Code
1.11.3 Classification of Cannabis Assemblages as Non-formal Groups
1.12 How Many Species of Cannabis Merit Recognition?
1.13 A Rationale for Emphasizing the Principal Selected Character Complexes in Classifying Cannabis
1.14 A Practical and Natural Taxonomy for Cannabis
1.15 Conclusions
Acknowledgements
References
2 History of Cannabis as Medicine: Nineteenth Century Irish Physicians and Correlations of Their Observations to Modern Research
Abstract
2.1 Introduction
2.1.1 Ancient Antecedents: The Celts, the Lacnunga, and Irish Hemp
2.2 William Brooke O’Shaughnessy and Indian Hemp
2.3 Michael Donovan and Neuropathic Pain
2.4 Corrigan and Chorea
2.5 Fleetwood Churchill and Uterine Hemorrhage
2.6 Richard Greene and the Scourge of Migraine
2.7 Conclusion: Contemporary Cannabis Research in Ireland
Acknowledgements
References
3 Cannabis sativa L.: Botany and Horticulture
Abstract
3.1 Introduction
3.2 The Plant Cannabis and Cannabinoids
3.3 Cannabis Horticulture
3.3.1 Indoor Cultivation
3.3.1.1 Grow Room Environment
3.3.1.2 Propagation Through Seeds
3.3.1.3 Vegetative Propagation
3.3.1.4 Hydroponics Propagation
3.3.1.5 Micropropagation
3.3.2 Outdoor Cultivation
3.3.2.1 Harvesting
3.3.2.2 Handling, Drying, Processing and Storage
Acknowledgements
References
4 Cannabis sativa and Cannabis indica versus “Sativa” and “Indica”
Abstract
4.1 Introduction
4.2 Formal Botanical Nomenclature: C. sativa
4.3 Formal Botanical Nomenclature: C. indica
4.4 The Slide from Formal to Vernacular
4.5 The Hybridization Impasse
4.6 Cannabinoids
4.7 Terpenoid Studies
4.8 Genetic Studies
4.9 Conclusions
Acknowledgements
References
5 Morpho-Anatomy of Marijuana (Cannabis sativa L.)
Abstract
5.1 Introduction
5.2 Morphology
5.3 Trichomes
5.3.1 Glandular Trichomes
5.3.2 Non-glandular Trichomes
5.4 Anatomy
5.5 Conclusion
Acknowledgements
References
6 Chemical and Morphological Phenotypes in Breeding of Cannabis sativa L.
Abstract
6.1 Introduction
6.2 Cannabinoids
6.2.1 Cannabinoid Quantity
6.2.2 Cannabinoid Quality
6.2.3 Cannabinoid Genetics
6.3 Essential Oil
6.4 Hemp Seed Oil and Protein
6.5 Bast Fiber
6.6 Part II: Current Breeding Status
6.6.1 Fiber Production
6.6.1.1 Cannabinoid Content
6.6.1.2 Seed Production
6.7 Conclusions
Acknowledgements
References
7 Natural Cannabinoids of Cannabis and Methods of Analysis
Abstract
7.1 Introduction
7.2 Phytocannabinoids
7.2.1 Historical Overview of the Isolation and Structure Elucidation of (-)-Δ9-Trans-Tetrahydrocannabinol (Δ9-THC) Type Cannabinoids
7.3 Δ9-THC Pharmaceutical Preparation
7.4 Stability of Δ9-THC
7.5 Analysis of Cannabinoids in Cannabis and Cannabis Products
7.5.1 Gas Chromatography (GC)
7.5.2 High-Performance Liquid Chromatography (HPLC)
7.5.3 Ultra-Performance Liquid Chromatography (UPLC)
7.5.4 High Performance Thin Layer Chromatography (HPTLC)
Acknowledgements
References
8 Cannabinoids: Biosynthesis and Biotechnological Applications
Abstract
8.1 Introduction
8.2 Cannabinoid Biosynthesis
8.2.1 Tetrahydrocannabinolic Acid Synthase
8.2.1.1 Biochemical Properties
8.2.1.2 Localization and Possible Physiological Function
8.2.2 Cannabidiolic Acid Synthase
8.2.2.1 Biochemical Properties
8.2.2.2 Cannabidiolic Acid Synthase, the Chemotype-Determining Enzyme
8.2.3 Cannabichromenic Acid Synthase
8.2.4 Geranyl Pyrophosphate:Olivetolate Geranyltransferase
8.2.5 Olivetol Synthase, a Polyketide Synthase for OLA Biosynthesis
8.2.6 Olivetolic Acid Cyclase, the First Plant Polyketide Cyclase
8.3 Biotechnological Cannabinoid Production
8.3.1 Cannabinoid Production in C. sativa
8.3.2 Cannabinoid Production in a Heterologous Host
8.3.3 Production of Unnatural Cannabinoids by Reengineering Cannabinoid Biosynthetic Genes
8.4 Conclusions
Acknowledgements
References
9 The Pharmacology and Therapeutic Potential of Plant Cannabinoids
Abstract
9.1 Introduction
9.2 A Brief Overview of the Cannabinoid Receptors
9.3 The in Vitro Pharmacological Effects of Certain Plant-Derived Cannabinoids
9.3.1 Δ9-Tetrahydrocannabinol
9.3.2 Cannabidiol
9.3.3 Δ9-Tetrahydrocannabivarin
9.3.4 Cannabigerol
9.3.5 Cannabichromene
9.4 Potential and Approved Therapeutic Uses of Plant Cannabinoids
9.4.1 Multiple Sclerosis
9.4.2 Nausea and Vomiting
9.4.3 Cancer
9.4.4 Pain
9.4.5 Schizophrenia
9.5 Conclusions
References
10 Cannabinoid CB2 Receptor Mechanism of Cannabis sativa L.
Abstract
10.1 Introduction
10.2 Sub-type CB2 Cannabinoid Receptor Gene, Gene Structure, Regulation and Variation
10.3 Diversification of CBR Genes, Expression and Function in Human, Rhesus Monkey, Dog, Rat and Mouse
10.4 Genetic Manipulation of CB2 Cannabinoid Receptors
10.5 Central Nervous System (CNS) Effects and Distribution of CB2Rs
10.6 Frame Work on the Molecular Basis for the Therapeutic Potential of Cannabinoids
10.7 The Role of Endocannabinoids in Psychiatric, Neuroinflammatory and Neurodegenerative Disorders
10.8 CB2Rs as Potential Therapeutic Target in Neurological and Mental Diseases
10.9 Conclusion
Acknowledgements
References
11 Cannabidiol as a Treatment for Seizures, Convulsions and Epilepsy
Abstract
11.1 Introduction
11.2 In Vitro Pharmacology of Cannabidiol and the Modulation of Neuronal Excitability
11.2.1 Actions of Cannabidiol in the Endocannabinoid System
11.2.2 Other Neuronal Mechanisms of Action of Cannabidiol
11.3 In Vivo Preclinical and Clinical Studies of Cannabidiol as an Antiepileptic Agent
11.4 Conclusion
Acknowledgements
References
12 Allergenicity to Cannabis sativa L. and Methods to Assess Personal Exposure
Abstract
12.1 Introduction
12.2 Scenarios of C. sativa Exposures
12.2.1 Personal Exposure
12.2.2 Occupational Hemp Exposures
12.2.3 Environmental Exposures
12.3 Allergens of Cannabis sativa and Cross-Reactivity
12.4 Biotechnological Advances in Diagnosis of Allergic Sensitization to C. sativa
12.5 Treatment of Allergic Exposure to Cannabis sativa
12.6 Additional Comments
12.7 Conclusion
Acknowledgements
References
13 Micropropagation of Cannabis sativa L.—An Update
Abstract
13.1 Introduction
13.2 Strategies Used for the Propagation of Cannabis Sativa L.
13.2.1 Conventional Propagation
13.2.2 In Vitro Propagation
13.2.3 Callus Production
13.2.4 Agrobacterium Mediated Transformation
13.2.5 Regeneration
13.2.6 Germplasm Conservation
13.3 Conclusions
Acknowledgements
References
14 Hairy Root Culture as a Biotechnological Tool in C. sativa
Abstract
14.1 Introduction
14.2 Agrobacterium Transformation of Hemp: Establishment and Applications of Transformed Root Cultures
14.2.1 Axenic Growth of Hemp Seedlings and Infection Protocol
14.2.2 Hemp Responses to Agrobacterium
14.2.3 Wild-Type Transformed Tissues
14.2.4 RolABC, rolA, rolB and rolC Transgenic Roots
14.2.5 Secondary Metabolites in C. sativa Transformed Root Cultures
14.3 Callusing Responses and Regeneration Ability of Hemp Hairy Roots
14.4 Conclusion
Acknowledgements
References
15 Genomics and Molecular Markers in Cannabis sativa L.
Abstract
15.1 Introduction
15.2 Molecular Markers for the Study of Variability and Genetic Structure of Cannabis sativa
15.2.1 Early Molecular Markers (RAPDs, RFLPs, AFLPs, Microsatellites (SSRs), ISSRs): The Forensic Issue and Estimates of Genetic Variability in Cannabis
15.2.2 Late Sequence-Based Molecular Markers (EST-SSR and SNPss): Broadening Knowledge on Genetic Variability in Cannabis
15.2.3 Genetic Maps
15.3 The Search for Markers Linked to Sex in Dioecious Hemp and to Monoecy in C. sativa
15.4 Genomics and Marker-Assisted Selection for Chemotype
15.4.1 Premise: The Genetics of Chemotype
15.4.2 Molecular Markers Linked to Chemotype
15.4.3 Single Nucleotide Polymorphisms in Chemotype-Determining Genes
15.5 Conclusions
References
16 The Role of Agrobacterium-Mediated and Other Gene-Transfer Technologies in Cannabis Research and Product Development
Abstract
16.1 Introduction
16.2 Plant Genetic Transformation Using Agrobacterium Species
16.2.1 Agrobacterium-Mediated Stable Transformation
16.2.2 Transient Gene Expression by Agroinfiltration
16.2.3 Hairy Root Transformation Using Agrobacterium rhizogenes
16.3 Genetic Transformation of Cannabis sativa
16.3.1 Plant Regeneration
16.4 Stable and Transient Heterologous Expression of Cannabinoid Genes
16.4.1 Localization of Pathway Intermediates
16.4.1.1 Tetrahydrocannabinolic Acid Synthase is Secreted into the Storage Cavity of Glandular Trichomes
16.4.1.2 Hexanoyl-CoA Synthetase (CsAAE1) is Localized to the Cell Cytoplasm
16.4.2 Enzyme Kinetics: Expression and Purification of Recombinant Cannabis Enzymes in Heterologous Hosts
16.4.2.1 Characterization of the Tetrahydrocannabinolic Acid Synthase Reaction Mechanism
16.4.2.2 Identification of Enzymes Involved in the Biosynthesis of Olivetolic Acid
16.4.3 Metabolic Engineering of the Cannabinoid Pathway in Heterologous Hosts
16.4.3.1 Over-Expression of Tetrahydrocannabinolic Acid Synthase in Tobacco Hairy Root Cultures
16.4.3.2 Over-Expression of Tetrahydrocannabinolic Acid Synthase in Yeast Cultures
16.4.3.3 Metabolic Engineering of the Cannabinoid Pathway for Commercialization
16.5 Future Opportunities with Cannabis
Acknowledgements
References
17 Induction of Polyploidy and Its Effect on Cannabis sativa L.
Abstract
17.1 Introduction
17.1.1 Different Methods to Increase Plant Secondary Metabolites
17.1.2 Polyploidy
17.1.3 Polyploidy in Cannabis
17.2 Methods
17.2.1 Tetraploid Induction in Cannabis
17.2.2 Apical Tip Meristem Treatment of the Seedlings
17.2.3 Detection of Tetraploid Plants Following Colchicine Application to Seedling Apical Tips
17.2.4 Preliminary Morphological Screening for Putative Tetraploids
17.2.5 Ploidy Level Determination by Flow Cytometry
17.2.6 Analysis of Morphological Characteristics
17.2.7 Phytochemical Measurement
17.3 Results
17.3.1 Survival and Growth of Colchicine-Treated Shoot Tips
17.3.2 Induction of Tetraploidy in Cannabis by Colchicine Application to Seedling Apical Meristem
17.3.3 Ploidy Level Determination by Fow Cytometry
17.3.4 Morphological Characteristics
17.4 Conclusion
References
18 Classical and Molecular Cytogenetics of Cannabis Sativa L.
Abstract
18.1 Introduction
18.2 Classical Cytogenetic Study of Hemp Chromosomes
18.3 Molecular Cytogenetics in Hemp
18.4 Conclusion
Acknowledgements
References
19 Assessing Genetic Diversity in Cannabis sativa Using Molecular Approaches
Abstract
19.1 Introduction
19.2 Origins of Genetic Diversity in Cannabis
19.3 Flower Induction in Cannabis
19.4 Mating Systems in Cannabis
19.5 Genetics of Cannabis
19.6 Assessing Genetic Diversity in Plants
19.7 Random Amplified Polymorphic DNA (RAPD) Analysis of Cannabis
19.8 Inter Sequence Simple Repeat (ISSR) Analysis of Cannabis
19.9 Amplified Fragment Length Polymorphism (AFLP) Analysis
19.10 Short Tandem Repeat (STR) and SSR Analysis
19.11 Sequence Characterized Amplified Regions (SCAR) Analysis of Cannabis
19.12 Single-Nucleotide Polymorphism (SNP) and Genotype-by-Sequencing (GSB) Analysis of Cannabis
19.13 Markers to Assess the Sex Phenotype in Cannabis
19.14 Future Directions
Acknowledgements
References
20 Cannabis Endophytes and Their Application in Breeding and Physiological Fitness
Abstract
20.1 Introduction
20.2 Diversity and Quantification of Endophytes Harbored in Cannabis Sativa
20.3 Strategies for Discovering Potential Cannabis Endophytes Aiding in Chemical Defense of Host Plants
20.4 Screening of Endophytic Microorganisms Living in Cannabis Plants Conferring Plant Adaptation and Growth
20.5 Investigation of Cannabis Endophytes Employing Antivirulence Strategy for Attenuation of Virulence Factors
20.6 Strategies for Discovering Endophytic Microbial Community Harbored in Plants with Similar Secondary Metabolites: Radula marginata as an Example
20.7 Conclusion
Acknowledgements
References
21 Chemical and Physical Elicitation for Enhanced Cannabinoid Production in Cannabis
Abstract
21.1 Introduction
21.2 Stress Response
21.3 Nutrient Deficiency
21.4 Drought Stress
21.5 Temperature
21.6 Photo-Radiation
21.7 Heavy Metals
21.8 Wounding
21.9 Pathogens
21.10 Hormones
21.11 Future Role of Elicitation in Development of Cannabis
21.12 Conclusion
Acknowledgements
References
22 Contaminants of Concern in Cannabis: Microbes, Heavy Metals and Pesticides
Abstract
22.1 Introduction
22.2 Microbial Contaminants
22.3 Microbial Testing
22.4 Microbial Harm Reduction
22.5 Janis Face—Endophytes
22.6 Heavy Metals and Radionucleotides
22.7 Janis Face—Bioremediation
22.8 Pesticide Residues
Acknowledgements
References
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