Copyrighted Material MUSHROOMS AS FUNCTIONAL FOODS Edited by Peter C.K.Cheung WILEY Copyrighted Material
■CONTENTS Foreword Preface Acknowledgments Contributors 1 Overview of Mushroom Cultivation and Utilization as Functional foods 1 Shu-Ting Chang 1..Introduction 1.2.What Are Mushrooms? 3 1.2.1.Definition of a Mushroom 3 1.2.2.Ecological Classification of Mushrooms 4 1.2.3.Identification of Mushrooms 4 1.3.Concept of Mushroom Biology and Applied Mushroom Biology 6 1.3.1.Mushroom Biology 6 1.3.2.Applied Mushroom Biology 7 1.3.3.Impact of Applied Mushroom Biology 9 1.3.3.1.Nongreen Revolution 9 1.3.3.2.Mushroom Bioremediation 11 1.4.Mushroom Cultivation 11 1.4.1.Major Phases of Mushroom Cultivation 1.4.2.Cultivation of Several Selected Mushrooms 1.4.2.1.Cultivation of it Agaricus 1.4.2.2.Cultivation of Lentinula edodes 14 1.4.2.3.Cultivation of Pleurotus saior-cait 1.4.2.4.Cultivation of Volvariella 17 1.4.2.5.Cultivation of Agaricus brasiliensis 18 vi
CONTENTS Foreword xv Preface xvii Acknowledgments xix Contributors xxi 1 Overview of Mushroom Cultivation and Utilization as Functional Foods 1 Shu-Ting Chang 1.1. Introduction 1 1.2. What Are Mushrooms? 3 1.2.1. Definition of a Mushroom 3 1.2.2. Ecological Classification of Mushrooms 4 1.2.3. Identification of Mushrooms 4 1.3. Concept of Mushroom Biology and Applied Mushroom Biology 6 1.3.1. Mushroom Biology 6 1.3.2. Applied Mushroom Biology 7 1.3.3. Impact of Applied Mushroom Biology 9 1.3.3.1. Nongreen Revolution 9 1.3.3.2. Mushroom Bioremediation 11 1.4. Mushroom Cultivation 11 1.4.1. Major Phases of Mushroom Cultivation 12 1.4.2. Cultivation of Several Selected Mushrooms 13 1.4.2.1. Cultivation of it Agaricus 14 1.4.2.2. Cultivation of Lentinula edodes 14 1.4.2.3. Cultivation of Pleurotus sajor-caju 17 1.4.2.4. Cultivation of Volvariella 17 1.4.2.5. Cultivation of Agaricus brasiliensis 18 vii
vill CONTENTS 1.4.2.6.Cultivation of Ganoderma lucidum 1.4.3.Utilization of Mushroom Germplasm 1.5.World Mushroom Production 21 1.6.Mushroom Biotechnology 1.6.1.Nutritional and Medicinal Value of Mushrooms 1.6.2.Nutriceuticals and Dietary Supplements 1.7.Development of World Mushroom Industry Movements 1.7.1.International Movement for Edible Mushrooms 1.7.2.International Movement for Medicinal Mushrooms 1.7.3.International Movement for Wild Mushrooms 1.8.Concluding Remarks References 29 2 Molecular Analysis and Genomic Studies of Shiitake Mushroom Lentin 2.1.Introduction 2.2.Isolation of Genes 2.2.1.Growth 2.2.1.1.Substrate-Utilizing Genes 36 2.2.2.Development 3 2.2.2.1.Mating-Type Genes 2..2.Genes Differentially Expressed in Dikarvotic mycelium 哪 2.2.2.3.Genes for Initial Fruiting Bodies/Primordium Formation 学 2.2.2.4.Genes for Mature Fruiting Bodies Formation 2.2.3.Physiological Processes in Lentinula edodes 2.2.3.1.Signal Transduction 2.2.3.2.Energy Production 2.2.3.3.Structural Proteins in Development 2.3.Molecular Genetics 2.3.1.Generation of Markers 2.3.2.Typing/Fingerprinting 2.3.3.Genetic Mapping 50
viii CONTENTS 1.4.2.6. Cultivation of Ganoderma lucidum 19 1.4.3. Utilization of Mushroom Germplasm 20 1.5. World Mushroom Production 21 1.6. Mushroom Biotechnology 23 1.6.1. Nutritional and Medicinal Value of Mushrooms 23 1.6.2. Nutriceuticals and Dietary Supplements 24 1.7. Development of World Mushroom Industry Movements 25 1.7.1. International Movement for Edible Mushrooms 26 1.7.2. International Movement for Medicinal Mushrooms 27 1.7.3. International Movement for Wild Mushrooms 27 1.8. Concluding Remarks 28 References 29 2 Molecular Analysis and Genomic Studies of Shiitake Mushroom Lentinula edodes 35 Hoi-Shan Kwan and Winnie W. Y. Chum 2.1. Introduction 35 2.2. Isolation of Genes 36 2.2.1. Growth 36 2.2.1.1. Substrate-Utilizing Genes 36 2.2.2. Development 37 2.2.2.1. Mating-Type Genes 38 2.2.2.2. Genes Differentially Expressed in Dikaryotic Mycelium 38 2.2.2.3. Genes for Initial Fruiting Bodies/Primordium Formation 38 2.2.2.4. Genes for Mature Fruiting Bodies Formation 44 2.2.3. Physiological Processes in Lentinula edodes 47 2.2.3.1. Signal Transduction 47 2.2.3.2. Energy Production 47 2.2.3.3. Structural Proteins in Development 48 2.3. Molecular Genetics 48 2.3.1. Generation of Markers 49 2.3.2. Typing/Fingerprinting 50 2.3.3. Genetic Mapping 50
CONTENTS 2.4.Functional Genomic Approaches for Gene Expression Analysis 0 2.4.1.Differential Display:RAP-PCR 2.4.2.cDNA Representation Difference Analysis 2.4.3.SAGE and LongSAGE 52 2.4.3.1.SAGE Profiles:Mycelium to Primordium 9 2.4.3.2.SAGE Profiles:Fruiting Bodies 2.4.4.cDNA Microarray 2.4.5.Expressed Sequence Tag 246 Yeast Two -Hybrid Sys 2.47 双 2.5.Transcriptional Regulation 2.5.1.Transcriptional Factors 2.5.2.Promoter Analysis 2.6.Transformation 2.6.1.Transformation Methods 66 2.6.1.1.PEG-Mediated Transformation 2.6.1.2.Restriction Enzyme-Mediated Integration 2.6.1.3.0 thers 58 2.6.2.Lentinula edodes Genes Used in Transformation 59 2.7.1.Postharvest Studies 2.7.2.StressResponses 2.7.2.1.Studies of Temperature Stress in Mushrooms 2.7.2.2.Studies of Molecular Chaperones in Fungi 2.7.3.Lignocellulose Degradation 2.7.4.Meiosis % 2.8.Conclusion 6 References 6 3 Nutritional Value and Health Benefits of Mushrooms 个 Peter C.K.Cheung 3.1.Introduction 3.2.Wild and Cultivated Edible Mushrooms 之 3.3.Production of Cultivated Mushrooms 12
CONTENTS ix 2.4. Functional Genomic Approaches for Gene Expression Analysis 50 2.4.1. Differential Display: RAP-PCR 51 2.4.2. cDNA Representation Difference Analysis 52 2.4.3. SAGE and LongSAGE 52 2.4.3.1. SAGE Profiles: Mycelium to Primordium 53 2.4.3.2. SAGE Profiles: Fruiting Bodies 53 2.4.4. cDNA Microarray 53 2.4.5. Expressed Sequence Tag 54 2.4.6. Yeast Two-Hybrid System 54 2.4.7. Sequencing-by-Synthesis Approach (454 Life Science) 54 2.5. Transcriptional Regulation 55 2.5.1. Transcriptional Factors 55 2.5.2. Promoter Analysis 55 2.6. Transformation 56 2.6.1. Transformation Methods 56 2.6.1.1. PEG-Mediated Transformation 56 2.6.1.2. Restriction Enzyme–Mediated Integration 57 2.6.1.3. Others 58 2.6.2. Lentinula edodes Genes Used in Transformation 58 2.7. Process Analysis 59 2.7.1. Postharvest Studies 59 2.7.2. Stress Responses 59 2.7.2.1. Studies of Temperature Stress in Mushrooms 59 2.7.2.2. Studies of Molecular Chaperones in Fungi 59 2.7.3. Lignocellulose Degradation 60 2.7.4. Meiosis 60 2.8. Conclusion 61 References 61 3 Nutritional Value and Health Benefits of Mushrooms 71 Peter C. K. Cheung 3.1. Introduction 71 3.2. Wild and Cultivated Edible Mushrooms 72 3.3. Production of Cultivated Mushrooms 72
X CONTENTS 3.4.Nutritional Composition 3.4.1.Con ntional Edible Mushrooms 3.4.1.1.Moisture 373 3.4.1.2.Protein and Amino Acids 3.4.1.3.Fat 3.4.1.4.Ash and Minerals 341 5 Vitamins 3.4.1.6.Dietary Fiber 3.4.1.7.Carbohydrates 3.4.1.8.Energy 3.4.1.9.0h omponents 18 3.5.Newly Cultivated/Nonconventional Mushrooms 7 3.6.Nutritional Evaluation 3.6.1.General Aspects 3.6.2 Biological Methods for Nutritional Evaluation 3.6.3.Mushroom Protein Ouality 87 3.7.Health Benefits of Edible Mushrooms 3.7. General Aspects 3.7.2 Antioxidants in Mushrooms 999 3721AmeRmnnt ive Acti 89 3.7.2.2 ation of M ushroom Phenolic 91 37.2.3 Biosy is of Pher Compounds 93 3.7.3.Hypochole rooms 3.7.4.Hypoglyc Mush 97 3.8.Conclusion 90 References 4 Sclerotia:Em g Functional Food Derived from Mushrooms 111 Ka-Hing Won and Peter C.K.Cheung 4.1.Introduction 111 4.2.Concepts of Mushroom Sclerotia 112 4.3.Ontogeny of Sclerotia 112 4.3.1.Morphological Aspects 4.3.2.Physiological Aspects
x CONTENTS 3.4. Nutritional Composition 73 3.4.1. Conventional Edible Mushrooms 73 3.4.1.1. Moisture 73 3.4.1.2. Protein and Amino Acids 74 3.4.1.3. Fat 75 3.4.1.4. Ash and Minerals 75 3.4.1.5. Vitamins 76 3.4.1.6. Dietary Fiber 77 3.4.1.7. Carbohydrates 78 3.4.1.8. Energy 78 3.4.1.9. Other Components 78 3.5. Newly Cultivated/Nonconventional Mushrooms 79 3.6. Nutritional Evaluation 80 3.6.1. General Aspects 80 3.6.2. Biological Methods for Nutritional Evaluation 80 3.6.3. Mushroom Protein Quality 87 3.7. Health Benefits of Edible Mushrooms 89 3.7.1. General Aspects 89 3.7.2. Antioxidants in Mushrooms 89 3.7.2.1. Bioactive Components and Their Antioxidative Activities 89 3.7.2.2. Characterization of Mushroom Phenolic Antioxidants 91 3.7.2.3. Biosynthesis of Phenolic Compounds from Mushrooms or Fungi 93 3.7.3. Hypocholesterolemic Effect of Mushrooms 94 3.7.4. Hypoglycemic Effect of Mushrooms 97 3.8. Conclusion 99 References 99 4 Sclerotia: Emerging Functional Food Derived from Mushrooms 111 Ka-Hing Wong and Peter C. K. Cheung 4.1. Introduction 111 4.2. Concepts of Mushroom Sclerotia 112 4.3. Ontogeny of Sclerotia 112 4.3.1. Morphological Aspects 112 4.3.2. Physiological Aspects 114
CONTENTS xi 4.3.2.1.Translocation 114 4.3.2.2.Exudation 115 4.4.Structure of Sclerotia 115 4.4.1.Rind 115 4.4.2.Cortex 116 4.4.3.Medulla 117 4.5.Cultivation of Mushroom Sclerotia 117 4.5.1.Sclerotia of Pleurotus tber-regium (Fries)Singer 118 4.5.2.Sclerotia of Polyporus rhinocerus Cooke 119 45anaemSnRBG 4.6. 121 4.6.1.Biochemical Components of Mushroom Sclerotia 46 11 Cell walls 121 4.6.1..Extracellular Matrix 122 122 4.6.2 hroom Sclerotia 4.6.2 224 ional Properties 126 4.7.Biopharmacological Values of Mushroom Sclerotia of P tuber-regium,P.rhinocerus,and W.cocos 128 4.7.1.In Vitro Mineral Binding Capacity 4.7.2.In Vitro Fermentability 4.7.3.In Vivo Ca and Mg Absorption 131 4.7.4.Antitumor and Immunomodulatory Activities 132 4.8.Conclusion 134 References 134 5 Antitumor and Imm odulatory activities of Mushroom Polysaccharides 147 Vincent E.C.Oo 5.1.Introduction 147 5.2.Antitumor Polysaccharides from Mushrooms (Higher Fungi) 149
CONTENTS xi 4.3.2.1. Translocation 114 4.3.2.2. Exudation 115 4.4. Structure of Sclerotia 115 4.4.1. Rind 115 4.4.2. Cortex 116 4.4.3. Medulla 117 4.5. Cultivation of Mushroom Sclerotia 117 4.5.1. Sclerotia of Pleurotus tuber-regium (Fries) Singer 118 4.5.2. Sclerotia of Polyporus rhinocerus Cooke 119 4.5.3. Sclerotia of Wolfiporia cocos (Schw.) Ryv. Et Gilbn [Poria cocos (Schw.) Wolf] 120 4.6. Biochemical, Nutritional, and Technological Characteristics of Mushroom Sclerotia 121 4.6.1. Biochemical Components of Mushroom Sclerotia 121 4.6.1.1. Cell Walls 121 4.6.1.2. Extracellular Matrix 122 4.6.1.3. Cytoplasmic Reserves 122 4.6.2. Nutritional Evaluation of Mushroom Sclerotia 123 4.6.2.1. Proximate Composition 123 4.6.2.2. Sclerotial Dietary Fiber 124 4.6.3. Physicochemical and Functional Properties of Mushroom Sclerotial DF 126 4.7. Biopharmacological Values of Mushroom Sclerotia of P. tuber-regium, P. rhinocerus, and W. cocos 128 4.7.1. In Vitro Mineral Binding Capacity 128 4.7.2. In Vitro Fermentability 129 4.7.3. In Vivo Ca and Mg Absorption 131 4.7.4. Antitumor and Immunomodulatory Activities 132 4.8. Conclusion 134 References 134 5 Antitumor and Immunomodulatory Activities of Mushroom Polysaccharides 147 Vincent E. C. Ooi 5.1. Introduction 147 5.2. Antitumor Polysaccharides from Mushrooms (Higher Fungi) 149
xil CONTENTS 5.3.Mechanisms of Antitumor Action of Mushroom Polysaccharides 153 5a1Asaotcrrcakaiat 5.3.2.Immunomodulatior 83 on Macrophages and S 163 5.3.2.2 NK C lushroom Polysaccharides 167 5.32.3. 168 5.3.2.4.Effects of Mushroom Polysaccharides on Hematopoietic Stem Cells 170 5.3.3.Antimetastasis 171 5.3.4.Antiangiogenesis 172 5.4.Structure and Antitumor Activity Relationship of 5.4.1.Effect of Molecular Mass 72 5.4.2.Impact of Branching Configuration 5.4.3.Relationship of Antitumor Activity and Conformation 175 5.4.4.Improvement of Antitumor Activity by Chemical Modifications 176 5.5.Conclusions 178 References 179 6 ues of Mush 199 6.1.Introduction 6.2.Legal and Reg y Issues of Int Controlling Dietary S Mushrooms in Different Countries 202 6.2.1.World Health Organization Guidelines 202 62 2 Codex Alimentarius 202 6.2.3.United States 203 6.2.4.European Union 6.2.5.Canada 0 626 ralia and New Zealand Japa Israe
xii CONTENTS 5.3. Mechanisms of Antitumor Action of Mushroom Polysaccharides 153 5.3.1. Antiproliferation of Cancer Cells and Induction of Apoptosis 153 5.3.2. Immunomodulation 161 5.3.2.1. Effects of Mushroom Polysaccharides on Macrophages and Spleen Cells 163 5.3.2.2. Effects of Mushroom Polysaccharides on NK Cells 167 5.3.2.3. Effects of Mushroom Polysaccharides on DCs 168 5.3.2.4. Effects of Mushroom Polysaccharides on Hematopoietic Stem Cells 170 5.3.3. Antimetastasis 171 5.3.4. Antiangiogenesis 172 5.4. Structure and Antitumor Activity Relationship of Polysaccharides 173 5.4.1. Effect of Molecular Mass 174 5.4.2. Impact of Branching Configuration 174 5.4.3. Relationship of Antitumor Activity and Conformation 175 5.4.4. Improvement of Antitumor Activity by Chemical Modifications 176 5.5. Conclusions 178 References 179 6 Regulatory Issues of Mushrooms as Functional Foods and Dietary Supplements: Safety and Efficacy 199 Solomon P. Wasser and Eden Akavia 6.1. Introduction 199 6.2. Legal and Regulatory Issues of Introducing and Controlling Dietary Supplements from Medicinal Mushrooms in Different Countries 202 6.2.1. World Health Organization Guidelines 202 6.2.2. Codex Alimentarius 202 6.2.3. United States 203 6.2.4. European Union 208 6.2.5. Canada 210 6.2.6. Australia and New Zealand 212 6.2.7. Japan 213 6.2.8. Israel 215
CONTENTS XI 6.3.Safety and Diversity of Dietary Supplement Types from Culinary-Medicinal Mushrooms 216 efor Obtaining 220 6.5.Experiences of Seven Countries in Consolidating Their Food Safety Systems 220 6.6.Summary References Index 227
CONTENTS xiii 6.3. Safety and Diversity of Dietary Supplement Types from Culinary–Medicinal Mushrooms 216 6.4. Submerged Culturing as Best Technique for Obtaining Consistent and Safe Mushroom Products 220 6.5. Experiences of Seven Countries in Consolidating Their Food Safety Systems 220 6.6. Summary 221 References 221 Index 227
FOREWORD It has been over twenty years since the concept of"functional foods"was first as 2 analysis of foods a ter nutne onsumers are nov tion The aso demand mor detailed information about food factors in order to obtain appropriate functional food products. like Ch na and apan,mushre oms ha e been co try and the nutritiona d Fo y alone have b en more thar as functional food ava The curren book of Professor C.K.Cheung,Mushrooms as Functional Foods,is a timely and well welcomed book for scientists and students working in functional food res archs onal values and health ber efits of mushrooms.this book also introduces emen molecular analysis and functional genomics to the study of mushroom.Health benefits of mushrooms such as. ive,nypo are d ubstar ith ies and this topic been treated extensively ina separate chapter.Included also isa unique and useful thper edge for the devel- opment of mushrooms into functional foods. CHI-TANG Ho Rutgers University
FOREWORD It has been over twenty years since the concept of “functional foods” was first introduced as a factor in the analysis of foods after nutrients. Consumers are now deeply interested in food bioactives that provide beneficial effects to humans in terms of health promotion and disease risk reduction. They also demand more detailed information about food factors in order to obtain appropriate functional food products. In Asian countries, like China and Japan, mushrooms have been collected and cultivated for hundred of years. They have a long history of use for their health promotion benefits. In recent years reports on the chemistry, and the nutritional and functional properties of mushroom have been overwhelming. In the Journal of Agricultural and Food Chemistry alone there have been more than 300 articles related to mushrooms published since 1990. However, there is no in-depth comprehensive reference book of mushrooms as functional food available. The current book of Professor C. K. Cheung, Mushrooms as Functional Foods, is a timely and well welcomed book for scientists and students working in functional food research. Besides covering the agricultural production, nutritional values, and health benefits of mushrooms, this book also introduces emerging molecular analysis and functional genomics to the study of mushroom. Health benefits of mushrooms, such as, antioxidative, hypocholesterolemic, and hypoglycemic effects are discussed in depth. Polysaccharides are the best known and potent mushroom-derived substances with immunomodulating and antitumor activities and this topic has been treated extensively in a separate chapter. Included also is a unique and useful chapter on regulatory issues of mushrooms as functional foods in different countries. Scientists and students who research mushrooms will certainly benefit from reading this comprehensive monograph to gain in-depth knowledge for the development of mushrooms into functional foods. CHI-TANG HO Rutgers University xv
PREFACE Mushrooms have been known for their nutritional and culinary values as well as viewed asonddsmdicesby humansforagesmothey can be considered ds which can provid ath benefits beyon and healing p the Ganoderma,Shiitake mushroom,and Cordyceps for the general public.How. e比feraure of rence books h the basic and applied aspects of mushrooms as This book is an integration of the recent research conducted on the biologi- cal and chemical aspects of mushrooms when being utilized as a functional food Topics that are covere in this n the agricultural production o from of to the tional ffectof the unconventional form of the mushroom(scou)and from stic actions of the physiological benefits and pharmacologa s to the regu of th biologists:food manufacturers:students majoring in food science.nutrition,biol ogy,and bi chemistry,to n ho are int PETER C.K.CHEUNG The Chinese University of Hong Kong
PREFACE Mushrooms have been known for their nutritional and culinary values as well as viewed as tonics and used as medicines by humans for ages. In modern terms, they can be considered as functional foods which can provide health benefits beyond the traditional nutrients they contain. There are monographs that cover the medicinal and healing properties of some individual traditional mushrooms and fungi such as the Ganoderma, Shiitake mushroom, and Cordyceps for the general public. However, there are very few in-depth and up-to-date comprehensive reference books in the scientific literature of both the basic and applied aspects of mushrooms as functional foods. This book is an integration of the recent research conducted on the biological and chemical aspects of mushrooms when being utilized as a functional food. Topics that are covered in this book range from the agricultural production of mushrooms to the use of molecular biological techniques like functional genomics, from nutritional values of newly cultivated mushroom species to the multifunctional effects of the unconventional form of the mushroom (sclerotium), and from the mechanistic actions of the physiological benefits and pharmacological properties of bioactive components in mushrooms to the regulations of their uses as functional foods and dietary supplements in different parts of the world. This comprehensive book should serve as a reference for scientists; chemists; biologists; food manufacturers; students majoring in food science, nutrition, biology, and biochemistry, to name a few; and all those who are interested in obtaining a stronger background in the development of mushrooms and edible fungi into functional foods. PETER C. K. CHEUNG The Chinese University of Hong Kong xvii