Fungal biology has an integral role to play in the development of the biotechnology and biomedical sectors. It has become a subject of increasing importance as new fungi and their associated biomolecules are identified. The interaction between fungi and their environment is central to many natural processes that occur in the biosphere. The hosts and habitats of these eukaryotic microorganisms are very diverse; fungi are present in every ecosystem on Earth. The fungal kingdom is equally diverse, consisting of seven different known phyla. Yet detailed knowledge is limited to relatively few species. The relationship between fungi and humans has been characterized by the juxtaposed viewpoints of fungi as infectious agents of much dread and their exploitation as highly versatile systems for a range of economically important biotechnological applications. Understanding the biology of different fungi in diverse ecosystems as well as their interactions with living and non-living is essential to underpin effective and innovative technological developments. This series will provide a detailed compendium of methods and information used to investigate different aspects of mycology, including fungal biology and biochemistry, genetics, phylogenetics, genomics, proteomics, molecular enzymology, and biotechnological applications in a manner that reflects the many recent developments of relevance to researchers and scientists investigating the Kingdom Fungi. Rapid screening techniques based on screening specific regions in the DNA of fungi have been used in species comparison and identification, and are now being extended across fungal phyla. The majorities of fungi are multicellular eukaryotic systems and therefore may be excellent model systems by which to answer fundamental biological questions. A greater understanding of the cell biology of these versatile eukaryotes will underpin efforts to engineer certain fungal species to provide novel cell factories for production of proteins for pharmaceutical applications. Renewed interest in all aspects of the biology and biotechnology of fungi may also enable the development of “one pot” microbial cell factories to meet consumer energy needs in the 21st century. To realize this potential and to truly understand the diversity and biology of these eukaryotes, continued development of scientific tools and techniques is essential. As a professional reference, this series will be very helpful to all people who work with fungi and should be useful both to academic institutions and research teams, as well as to teachers, and graduate and postgraduate students with its information on the continuous developments in fungal biology with the publication of each volume.
More information about this series at http://www.springer.com/series/11224
Bhim Pratap Singh • Lallawmsanga
Ajit Kumar Passari
Editors
Biology of Macrofungi
Editors
Bhim Pratap Singh
Molecular Microbiology and Systematics
Laboratory
Department of Biotechnology
Mizoram University
Aizawl, Mizoram, India
Ajit Kumar Passari
Molecular Microbiology and Systematics
Laboratory
Department of Biotechnology
Mizoram University
Aizawl, Mizoram, India
Lallawmsanga
Molecular Microbiology and Systematics Laboratory
Department of Biotechnology
Mizoram University
Aizawl, Mizoram, India
ISSN 2198-7777
Fungal Biology
ISSN 2198-7785 (electronic)
ISBN 978-3-030-02621-9 ISBN 978-3-030-02622-6 (eBook)
This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
This volume is dedicated to Late Shri Ram Prasad father of Dr. Bhim Pratap Singh, Senior Editor of this volume, for his continuous motivation and encouragement
Late Shri Ram Prasad (1938–2011)
Foreword
It is very important to estimate and document the mushroom (macrofungi) biodiversity as they are well known for the production of bioactive compounds having biotechnological applications and are recognized as supplementary foods due to high nutritional content and medicinal importance. Among the few important features of mushrooms, they can be used as a source of food and medicines besides their key ecological roles. They are also considered as the efficient food for the future as they have high amount of proteins and other nutrients, so they can be used to cope up with the nutrient malnutrition in developing countries. Mushrooms are the only edible source of Vitamin D and considered among the top foods to prevent osteoporosis.
The book volume published by Springer Nature on Biology of Macrofungi has received contributions from academicians and scientists from several countries throughout the world including the United Kingdom, the United States, Brazil, China, Serbia, etc. I believe the knowledge shared by the contributors will be very helpful for the readers working in the field of macrofungi. The book has descriptions about the methods used for the identification of macrofungi and how the macrofungi can be exploited for several applications for sustainable development. The book also describes how to cultivate the important types of mushrooms to generate self-employment.
I am sure that the contents given in this volume are a comprehensive coverage of various important aspects of exploitation of macrofungi and their applications in health and pharmaceutical industries. I congratulate the editors and contributors for bringing this volume on Biology of Macrofungi.
K. R. S. Sambasiva Rao Vice-Chancellor
Mizoram University (A Central University) Aizawl, Mizoram, India
Preface
Mushrooms are often been referred as functional food due to their high nutritional contents. They contain high amount of antioxidants, the chemicals that are required to get rid of free radicals. Free radicals are very dangerous for the body cells as they lead to several deadly diseases including cancer. Mushrooms have high content of selenium, a mineral which is not available by eating most of the fruits and vegetables, and help in detoxifying disease-causing compounds in the body and also play an important role in preventing inflammation. They are high in fiber, potassium, and Vitamin C content which play an important role in preventing cardiovascular health.
The present volume has 19 chapters contributed by the researchers and academicians from several countries including the United Kingdom, the United States, China, Malaysia, Poland, Serbia, Brazil, and India. The volume has covered most of the recent developments in macrofungal biology starting from the latest methods used for the identification of wild macrofungi and several applications of macrofungi for sustainable livelihood.
Editors of the volume express their gratitude to all the contributors for sharing their work in this volume. We are also thankful to Springer Nature publishers for giving us a chance to compile this important volume. We hope that the contents presented in this book will be useful for the readers involved in macrofungi research and all concern.
Mizoram, India
Bhim Pratap Singh Lallawmsanga Ajit Kumar Passari
Acknowledgments
My sincere thanks are extended to all the academicians and scientists who have contributed chapters and happily agreed to share their work on various aspects of macrofungi in this volume. At the same time, I also express my deepest gratitude to my family members, especially my wife (Dr. Garima Singh) and my daughter (Aadita Singh), for their kind support which has prompted me to complete the assignment on time. I am also thankful to the Department of Biotechnology (DBT), New Delhi, Government of India, for supporting us financially in the form of several externally funded projects from time to time and for the establishment of DBT Bioinformatics Centre at Mizoram University which was quite useful during the compilation of the book. I am equally thankful to the Springer Publishing for their full cooperation during the production of the volume. In particular, I am thankful to the series editors, Dr. Vijai Kumar Gupta and Prof. Maria G. Tuohy, for accepting our proposal and providing their full support and encouragements. I am also thankful to the production team of Springer Nature for all their efforts for publishing the volume on time. I admit that it is quite possible that some mistakes may have occurred in the text inadvertently, and I take responsibilities for the mistakes, and please feel free to inform me the same.
I am thankful to Prof. KRS Sambasiva Rao, Vice-Chancellor, Mizoram University, for his endeavor and motivations at all stages of the progress.
Mizoram, India
Bhim Pratap Singh
1 Exploration of Macrofungi in Sub-Tropical Semi-Evergreen Indian Forest Ecosystems .
Lallawmsanga, Ajit Kumar Passari, and Bhim Pratap Singh
2 A Global Overview of Edible Mushrooms .
Malarvizhi Kaliyaperumal, Kezhocuyi Kezo, and Sugantha Gunaseelan
3 Molecular Characterization of Wild Mushrooms: A Paradigm Shift from Morphotyping
Madhusmita Borthakur and S. R. Joshi
4 Antimicrobial and Hepatoprotective Activities of Edible Mushrooms
Jasmina Glamočlija, Marina Kostić, and Marina Soković
5 Mushroom-Mediated Protection from Oxidative Damage to DNA .
John A. Buswell
6 Chemical and Bioactive Profiling of Wild Edible Mushrooms . . . . . . . 129
Katarzyna Sułkowska-Ziaja, Katarzyna Kała, Jan Lazur, and Bożena Muszyńska
7 Biotechnological Requirements for the Commercial Cultivation of Macrofungi: Substrate and Casing Layer
Jaime Carrasco, Maria L. Tello, Margarita Perez, and Gail Preston
8 Role of Mushroom Fungi in Decolourization of Industrial Dyes and Degradation of Agrochemicals
Sachin Gupta, Sudheer K. Annepu, Baby Summuna, Moni Gupta, and Sunil A. Nair
9 Mushrooms: Isolation and Purification of Exopolysaccharides .
Yuxiao Wang, Xiaojun Huang, and Shaoping Nie
177
10
Novel Prospective of Wild Mushroom Polysaccharides as Potential Prebiotics
Yuxiao Wang, Xiaojun Huang, and Shaoping Nie
11 Pharmaceutic Prodigy of Ergosterol and Protein Profile of Ganoderma lucidum .
Anna Goyal and Anu Kalia
12 Application of Wild Macrofungi as Anticancer Therapeutics . .
Peter Chiew Hing Cheong, Chon Seng Tan, and Shin Yee Fung
13 Recent Advances in Cultivation of Edible Mushrooms .
Meena Kapahi
14 Medicinal Mushrooms: Cultivation and Pharmaceutical Impact 287
Peter Chiew Hing Cheong, Chon Seng Tan, and Shin Yee Fung
15 Biological Control of Microbial Pathogens in Edible Mushrooms 305 Gail M. Preston, Jaime Carrasco, Francisco J. Gea, and María J. Navarro
16 Cordycepin: A Biotherapeutic Molecule from Medicinal Mushroom 319 Mohammad Soltani, Roslinda Abd Malek, Nagib A. Elmarzugi, Mohamad Fawzi Mahomoodally, Davin Uy, Ong Mei Leng, and Hesham A. El-Enshasy 17 Biosynthesis of Nanoparticles Using Mushrooms 351 Anu Kalia and Gagandeep Kaur
and
Aparecido Almeida Conceição, Joice Raisa Barbosa Cunha, Vandinelma Oliveira Vieira, Rubén Darío Romero Pelaéz, Simone Mendonça, João Ricardo Moreira Almeida, Eustáquio Souza Dias, Euziclei Gonzaga de Almeida, and Félix Gonçalves de Siqueira
19 Wild Macro-Fungi from Northwest Himalayas: Future Prospects and Challenges 379 Monika Thakur
Contributors
João Ricardo Moreira Almeida Graduate Program Microbial Biology, University of Brasilia, Distrito Federal, Brasília, Brazil
Embrapa Agroenergy, Brasília, Distrito Federal, Brazil
Sudheer K. Annepu ICAR-Directorate of Mushroom Research, Solan, India
Madhusmita Borthakur Microbiology Laboratory, Department of Biotechnology & Bioinformatics, North-Eastern Hill University, Shillong, Meghalaya, India
John A. Buswell Institute of Edible Fungi, Academy of Agricultural Sciences, Shanghai, China
Jaime Carrasco Department of Plant Sciences, University of Oxford, Oxford, UK
Peter Chiew Hing Cheong Medicinal Mushroom Research Group (MMRG), Department of Molecular Medicine, University of Malaya, Kuala Lumpur, Malaysia
Aparecido Almeida Conceição Graduate Program of Biosciences, Federal University of Bahia, Vitória da Conquista, Bahia, Brazil
Embrapa Agroenergy, Distrito Federal, Brasília, Brazil
Joice Raisa Barbosa Cunha Graduate Program Microbial Agricultural, Federal University of Lavras, Lavras, Minas Gerais, Brazil
Embrapa Agroenergy, Distrito Federal, Brasília, Brazil
Eustáquio Souza Dias Graduate Program Microbial Agricultural, Federal University of Lavras, Minas Gerais, Brazil
Euziclei Gonzaga de Almeida Graduate Program of Biotechnology and Biodiversity, Federal University of Mato Grosso, Mato Grosso, Grosso, Cuiabá, Brazil
Félix Gonçalves de Siqueira Graduate Program of Biosciences, Federal University of Bahia,Vitória da Conquista, Bahia, Distrito Federal, Brazil
Contributors
Graduate Program of Biotechnology and Biodiversity, Federal University of Mato Grosso, Mato Grosso, Cuiabá, Brazil
Embrapa Agroenergy, Distrito Federal, Brasília, Brazil
Hesham A. El-Enshasy Institute of Bioproduct Development, Universiti Teknologi Malaysia, Johor Bahru, Johor, Malaysia
School of Chemical Engineering, Faculty of Engineering, Universiti Teknologi Malaysia, Johor Bahru, Johor, Malaysia
City of Scientific Research and Technology Application, New Burg Al Arab, Alexandria, Egypt
Nagib A. Elmarzugi Department of Industrial Pharmacy, Faculty of Pharmacy & Biotechnology Research Center, Tripoli, Libya
Shin Yee Fung Medicinal Mushroom Research Group (MMRG), Department of Molecular Medicine, University of Malaya, Kuala Lumpur, Malaysia
Centre for Natural Products Research and Drug Discovery (CENAR), University of Malaya, Kuala Lumpur, Malaysia
University of Malaya Centre for Proteomics Research (UMCPR), University of Malaya, Kuala Lumpur, Malaysia
Francisco J. Gea Centro de Investigación Experimentación y Servicios del Champiñón (CIES), Quintanar del Rey, Cuenca, Spain
Jasmina Glamočlija Department of Plant Physiology, Institute for Biological Research “Siniša Stanković”, University of Belgrade, Belgrade, Serbia
Anna Goyal Department of Microbiology, College of Basic Sciences and Humanities, Punjab Agricultural University, Ludhiana, Punjab, India
Moni Gupta Division of Biochemistry, Sher-e-Kashmir University of Agricultural Sciences and Technology of Jammu, Jammu, India
Sachin Gupta Division of Plant Pathology, Sher-e-Kashmir University of Agricultural Sciences and Technology of Jammu, Jammu, India
Xiaojun Huang State Key Laboratory of Food Science and Technology, Nanchang University, Nanchang, Jiangxi Province, China
S. R. Joshi Microbiology Laboratory, Department of Biotechnology & Bioinformatics, North-Eastern Hill University, Shillong, Meghalaya, India
Katarzyna Kała Department of Pharmaceutical Botany, Faculty of Pharmacy, Jagiellonian University Medical College, Kraków, Poland
Anu Kalia Electron Microscopy and Nanoscience Laboratory, Department of Soil Science, College of Agriculture, Punjab Agricultural University, Ludhiana, Punjab, India
Malarvizhi Kaliyaperumal Centre for Advanced Studies in Botany, Guindy Campus, University of Madras, Chennai, Tamil Nadu, India
Meena Kapahi Department of Chemistry, Manav Rachna University (MRU), Faridabad, India
Gagandeep Kaur Punjab Agricultural University, Ludhiana, Punjab, India
Kezhocuyi Kezo Centre for Advanced Studies in Botany, Guindy Campus, University of Madras, Chennai, Tamil Nadu, India
Marina Kostić Department of Plant Physiology, Institute for Biological Research “Siniša Stanković”, University of Belgrade, Belgrade, Serbia
Lallawmsanga Molecular Microbiology and Systematics Laboratory, Department of Biotechnology, Mizoram University, Aizawl, Mizoram, India
Jan Lazur Department of Pharmaceutical Botany, Faculty of Pharmacy, Jagiellonian University Medical College, Kraków, Poland
Ong Mei Leng Harita Go Green Sdn Bhd, Johor Bahru, Johor, Malaysia
Mohamad Fawzi Mahomoodally Department of Health Sciences, Faculty of Science, University of Mauritius, Réduit, Mauritius
Roslinda Abd Malek Institute of Bioproduct Development (IBD), Universiti Teknologi Malaysia (UTM), Skudai, Malaysia
Margarita Department of Plant Sciences, University of Oxford, Oxford, UK
Simone Mendonça Embrapa Agroenergy, Distrito Federal, Brasília, Brazil
Bożena Muszyńska Department of Pharmaceutical Botany, Faculty of Pharmacy, Jagiellonian University Medical College, Kraków, Poland
Sunil A. Nair Dr. Y.S. Parmar University of Horticulture and Forestry, Solan, India
María J. Navarro Centro de Investigación Experimentación y Servicios del Champiñón (CIES), Quintanar del Rey, Cuenca, Spain
Shaoping Nie State Key Laboratory of Food Science and Technology, Nanchang University, Nanchang, Jiangxi Province, China
Ajit Kumar Passari Molecular Microbiology and Systematics Laboratory, Department of Biotechnology, Mizoram University, Aizawl, Mizoram, India
Rubén Darío Romero Pelaéz Graduate Program Microbial Biology, University of Brasilia, Distrito Federal, Brasília, Brazil
Embrapa Agroenergy, Distrito Federal, Brasília, Brazil
Margarita Perez Centro Tecnológico de Investigación del Champiñón de La Rioja (CTICH), Autol, Spain
Gail M. Preston Department of Plant Sciences, University of Oxford, Oxford, UK
Bhim Pratap Singh Molecular Microbiology and Systematics Laboratory, Department of Biotechnology, Mizoram University, Aizawl, Mizoram, India
Marina Soković Department of Plant Physiology, Institute for Biological Research “Siniša Stanković”, University of Belgrade, Belgrade, Serbia
Mohammad Soltani Institute of Bioproduct Development (IBD), Universiti Teknologi Malaysia (UTM), Skudai, Malaysia
Gunaseelan Sugantha Centre for Advanced Studies in Botany, Guindy Campus, University of Madras, Chennai, Tamil Nadu, India
Katarzyna Sułkowska-Ziaja Department of Pharmaceutical Botany, Faculty of Pharmacy, Jagiellonian University Medical College, Kraków, Poland
Baby Summuna Sher-e-Kashmir University of Agricultural Sciences and Technology of Jammu, Wadoora, India
Chon Seng Tan Ligno Research Foundation, Balakong Jaya, Selangor, Malaysia
Maria L. Tello Centro Tecnológico de Investigación del Champiñón de La Rioja (CTICH), Autol, Spain
Monika Thakur Amity Institute of Food Technology, Amity University Uttar Pradesh, Noida, India
Davin Uy Research and Innovation Center, Tumnuptek, Chamkarmorn, Phnom Penh, Cambodia
Vandinelma Oliveira Vieira Graduate Program of Biotechnology and Biodiversity, Federal University of Mato Grosso, Cuiabá, Mato, Grosso, Brazil
Embrapa Agroenergy, Brasília, Distrito Federal, Brazil
Yuxiao Wang State Key Laboratory of Food Science and Technology, Nanchang University, Nanchang, Jiangxi Province, China
Chapter 1 Exploration of Macrofungi in Sub-Tropical Semi-Evergreen Indian Forest Ecosystems
Lallawmsanga, Ajit Kumar Passari, and Bhim Pratap Singh
1.1 Introduction
1.1.1 What is a Mushroom?
Mushroom is a term used to define the fruiting body formed by a group of fungi such as Basidiomycetes and Ascomycetes. They are either epigeous or hypogeous large enough to be seen with the naked eye and can be picked by hand (Chang and Miles 1992). The fruiting bodies of fungi (except for bracket) are normally short lived; and their existence varies from one to another. Mushrooms basically have the specific nutritional and ecological requirements for their growth and development; some of them thrive extremely well in soil and some of them utilize degrading plant residues as saprophytes. Many of the mushrooms exist in symbiotic relationship with roots of higher plant species. Due to the differences in their lifestyle, their ecological influence also varies from one to another (Lallawmsanga et al. 2016). These macrofungi appear at different times in a year and some species do not appear every year (Packham et al. 2002). They are a group of fungi that forms relatively conspicuous sporocarp (Ferris et al. 2000). As the tropical regions are considered rich in biodiversity, most of the new species of mushrooms reported in the recent years are from these regions; these newly reported mushrooms are usually associated with the native trees.
The occurrence of mushroom fruiting bodies varies according to different ecological climate (Arora 1991). They are essential for recycling of nutrients, growth and development of saplings in the forest base while some of them cause damage to the trees as they are in parasitic association with them. Humid climate provides
Lallawmsanga · A. K. Passari · B. P. Singh (*)
Molecular Microbiology and Systematics Laboratory, Department of Biotechnology, Mizoram University, Aizawl, Mizoram, India
B. P. Singh et al. (eds.), Biology of Macrofungi, Fungal Biology, https://doi.org/10.1007/978-3-030-02622-6_1
1
more distribution as compared to hot and dry periods (Sibounnavong et al. 2008). Mueller et al. (2007) estimated that there are 53, 000 to 110, 000 mushrooms in the world. However, Hawksworth (2001) estimated the global mushroom is about 140,000 species and approximately 14,000 species of them are known. It is also assumed that the mushrooms which are still unexplored could be of highly beneficial to mankind. Biodiversity research has become more important following the Convention on Biological Diversity in 1992 to understand the structure and function of ecosystems (Gadd 2007). Besides flora and fauna, fungi are considered as one of the most significant organisms in the world due to their important role in ecosystem (Mueller et al. 2004). Fungi have great importance ecologically as well as in industrially. They are found in almost all part of human life and ecosystems (Garibay-Orijel et al. 2009).
There are over 27,000 mushroom species recorded in India which makes it the second largest biotic community after insects (Manoharachary et al. 2005). True fungi belong to kingdom Eukaryota which has four phyla, 103 orders, 484 families and 4979 genera. True fungi consists of monophyletic groups, of which Basidiomycetes alone accounts for 35% of the fungal species described so far. They are of great importance ecologically as well as industrially. Ascomycetes, Basidiomycetes, some Zygomycetes and Glomeromycetes are grouped under hypogeous fungi (Trappe and Castellano 1991). The loss of spore discharge mechanisms in Ascomycetes and Basidiomycetes are the common characteristics observed in the two (Alessandra et al. 2014).
1.1.2 Why do we need to Explore Mushrooms?
Mushrooms are rich in food, medicine and plant growth promotion (Ghate and Sridhar 2015). So they have large number of importance in biodegradation, ecosystems, food industry and pharmacology (Barros et al. 2007). Mushrooms are considered as healthy foods due to their high composition of nutrients like proteins, carbohydrates, vitamins, fatty acids, amino acids and various elements in their fruiting body (Liu et al. 2015; Dimitrijevic et al. 2016). Mushrooms are known as an unlimited source of bioactive compounds and their immense diversity provide a massive prospective in the discovery and development of several drugs (Butler 2004). They are also known to produce the most important kinds of bioactive compounds including lectins, peptidoglucans, phenolic compounds, polysaccharides, steroids, etc., that acquire a broad range of pharmacological properties like antimicrobial (Alves et al. 2013), antioxidant (Ferreira et al. 2009), anti-tumor (Popovic et al. 2013), and therapeutic properties such as counteracting diseases including hypertension, hypercholesterolemia and cancer (Gast et al. 1988). These factors made the large scale production of mushroom became an important industry in several countries (Rafique 1996; Izlam et al. 2009).
Lallawmsanga
Rise in food prices due to high biofuel prices have caused scarcity of food and malnutrition throughout the world. To lessen this, cultivation of mushroom is beneficial and applicable for the poor farmers as it is labor intensive, short-duration for production and land saving (Shah et al. 2004). It could be more applicable in developing countries where malnutrition is one of the biggest problems as mushrooms contain high level of nutrients and its high productivity per unit area (Eswaran and Ramabadran 2000).
1.2 Exploration of Wild Mushroom Diversity
1.2.1
Macroscopic Identification
Mushrooms collected from the selected reserve forests were first identified based on their macroscopic characteristics. The followings are the macroscopic features used for the identification up to genus: attachment, cap size, colour, gills, shape, surface texture and moisture, spacing, stem size, the presence or absence of partial and universal veils. The identification was done by using seven mycological characters such as hymenium type, cap shape, gills, stipe character, color of the spore print, ecological type, and edibility (Krishna et al. 2015).
1.2.2
Molecular Characterization
Several numbers of molecular markers have been employed in the rapid identification of mushrooms in modern biotechnological research (Lian et al. 2008). DNA based techniques have eased the knowledge about microbial diversity from the natural ecosystems (Tuckwell et al. 2005). Mullis and Faloona (1987) have developed DNA-based PCR and taxon specific primers for the identification and study various kinds of fungi. The internal transcribed spacer region has been widely employed for the identification of fungi at species level (Sanchez-Ballesteros et al. 2000). Molecular fingerprinting offer an efficient way to study genetic variations which further helps in breeding programs due to its properties including high levels of detectable polymorphism and independence of environmental parameters (Yin et al. 2014).
Various molecular techniques such as random amplified polymorphic DNA (RAPD), amplified fragment length polymorphism (AFLP) analysis, simple sequence repeat (SSR) analysis, inter-simple sequence repeat (ISSR) analysis have been commonly employed for the genetic characterization of plants, animals, fungi and other organisms (Mei et al. 2014). RAPD analysis is used to reveal genetic
Lallawmsanga et al.
similarity and phylogenetic analysis due to simplicity in its technique. RAPD is furthermore employed for classification and study the genetic diversity in newly discovered species that are with agricultural and industrial importance (Shakeel et al. 2013). The advantages of employing RAPD as a molecular marker is that the DNA sequence information is not required, moreover they require only a small amount of DNA template, no harmful contamination and low-cost technology (Mei et al. 2014). Among the molecular markers, ISSR has gained interest as it is known to be abundant, highly informative, highly polymorphic and extremely reproducible. It has been effectively employed in the molecular identification of plants (Zietkiewicz et al. 1994) and fungi (Tang et al. 2010).
1.3 Biodiversity of Mushrooms in two Ecosystems
Mizoram, a state in North Eastern Region of India is a big bio-prospecting area. It is (21°57' - 24°30' North and 92°15' - 93°26' East) located in the extreme southern part of northeastern India, has a geographical area of 2,108,100 ha (0.6% of India’s geographical area). Mizoram is bounded on the north by Assam and Manipur, on the east and south by Myanmar and on the west by Bangladesh and Tripura. The terrain is hilly and mostly undulating with the average altitude ranging from 500 to 800 m and the maximum reaching 2157 m in the Blue Mountains. It is an important part of the Indo-Burma Biodiversity Hot Spot. Therefore, existence of agriculturally and industrially potential mushroom strains with diverse genetic resources in this region cannot be ruled out. It is well recognized that the diversity of Mushrooms in NorthEastern region, especially in Mizoram remains underexplored. It is important to explore the genetic resources of this region that will be useful for various purposes in the field of industrial, agricultural and pharmaceutical sectors.
1.3.1 Dampa Tiger Reserve
Dampa Tiger Reserve (Mamit District, Mizoram) is the largest wildlife sanctuary in Mizoram. It covers up an area of about 500 sq.km, falls within 23° 23' 15''N – 23° 42' 20''N latitudes and 92° 16' 25''E – 92° 25' 55''E longitudes. Altitude in the forest ranged between 800 to 1100 m above mean sea level. The annual rainfall ranged between 2000 mm to 2500 mm and maximum rainfall occurs in the month of June to August. Pleasant and warm climate prevails in the reserve all through the year with temperate to chill winter during November to January in higher altitudes. The natural vegetation includes Michelia champaca, Dipterocarpus turbinatus, and Terminalia chebula in the lower elevations while the higher elevations are characterized by Castanopsis indica , Dendrocalamus longispathus, Gmelina arborea, Lannea coromandelica, Mesua ferrea, Quercus sp., Schima wallichii and Sterculia
villosa. The reserve hosts several endangered species including Panthera tigris (tiger), Neofelis nebulosa (clouded leopard) and Elephus maximus (Asiatic elephant). It is especially affluent M. assamensis, M. leonina, M. arctoides, Nycticebus bengalensis, Trachypithecus pileatus and T. phayrei, Macaca mulatta.
1.3.2 Murlen National Park
Murlen Nation Park (Champhai District, Mizoram), located in the neighborhood of the Indo-Myanmar boundary and is important for its propinquity to the Chin Hills. It covers up an area of roughly 100 sq. km. It lies within 23° 32' 42'' – 23° 41' 36''N latitudes and 92°13' 12'' – 92°27' 24''E longitudes with altitude ranged from 1000–1600 m. Sub-Montane forest and semi-Evergreen forest are important vegetation that covers of the area. Murlen is a host for variety of rich flora and fauna. The prominent plant species that can be seen includes Arundinaria callosa, Prunus myrica, Rhododendron, Quercus, Betula, Schima wallichai, Arundinaria callosa, Canes, Michelia champaca and different types of orchids. It is one of the most important bird areas (IBM) supporting several threatened species. The Park provides a habitat for mammals Callosciurus pygerythrus, Cervus unicolor, C. erythraeus, Hylobates hoolock, Macaca assamensis, M. arctoides, Martes flavigula, Muntiacus muntjak, Ratufa bicolour, Sus scrofa and Trachypithecus pileatus
1.3.3 Diversity of Macrofungi in Dampa Tiger Reserve and Murlen National Park
The mushroom samples were collected from the abovementioned forests in 2013–2015. The mushroom fruiting bodies were first photographed in their natural habitat before collecting them. The collected fruiting bodies were carefully cleaned to remove soil and other substrates using fine brush. The collected mushroom samples were kept in separate paper bags to avoid mixing and were transferred to the laboratory. The specimens preserved as herbarium were identified with the help of standard literatures (Adhikari 2000).
The assessment of mushroom diversity of the Dampa Tiger Reserve and Murlen National Park (Mizoram, India) was accomplished and we collected 249 mushroom taxa (Fig. 1.1 A-E and Table 1.1) mainly from the genera Amanita, Boletus, Russula, Psathyrella, Schizophyllum, Calocybe, Lentinus, Polyporus, Pleurotus, Trametes and Tricholoma (Fig. 1.2). In total there are 39 mushroom families recorded from the 249 taxa collected (Table 1.2) and the major families are Agaricaceae, Amanitaceae, Boletaceae, Lyophyllaceae, Marasmiaceae, Polyporaceae, Psathyrellaceae, Russulaceae, Schizophyllaceae and Tricholomataceae (Fig. 1.3).
Table 1.1 Wild mushrooms collected from Mizoram, India
Lallawmsanga et al.
Genus No. of individuals
Agaricus 4
Lepiota 1
Lycoperdon 1
Leucoagaricus 1
Amanita 15
Auricularia 4
Bolbitius 3
Boletus 41
Xerocomus 1
Cantharellus 2
Clavaria 1
Clavulinopsis 2
Cortinarius 1
Crepidotus 4
Calocera 1
Entoloma 3
Geoglossum 1
Chlorociboria 1
Helvella 1
Laccaria 4
Hygrocybe 2
Coltrichia 5
Hebeloma 2
Inocybe 1
Leotia 1
Calocybe 8
Termitomyces 2
Marasmius 2
Xeromphalina 4
Hydropus 1
Phlebia 1
Merulius 1
Mycena 5
Favolaschia 1
Phallus 2
Hymenopellis 1
Xerula 3
Mucidula 2
Pleurotus 6
Lentinus 8
Trametes 7
Panus 2
Polyporus 6
Pycnosporus 1
Table 1.1 (continued)
Genus No. of individuals
Coprinellus 3
Coprinopsis 1
Coprinus 1
Psathyrella 12
Lactarius 5
Russula 23
Cookeina 1
Schizophyllum 10
Scleroderma 4
Agrocybe 1
Pholiota 4
Stropharia 1
Thelephora 2
Clitocybe 1
Collybia 1
Lepista 1
Lichenomphalina 1
Tricholoma 6
Flammulina 1
Xylaria 4
Fig. 1.2 Major genera recorded from Dampa Tiger Reserve and Murlen National Park, Mizoram, India
Table 1.2 Family wise distribution of wild mushrooms collected from Mizoram, India
Family No. of individual
Agaricaceae 7
Amanitaceae 15
Auriculariaceae 4
Bolbitiaceae 3
Boletaceae 42
Cantharellaceae 2
Clavariaceae 3
Cortinariaceae 1
Crepidotaceae 4
Dacrymycetaceae 1
Entolomataceae 3
Geoglossaceae 1
Helotiaceae 1
Helvellaceae 1
Hydnangiaceae 4
Hygrophoraceae 2
Hymenochaetaceae 5
Hymenogastraceae 2
Inocybaceae 1
Leotiaceae 1
Lyophyllaceae 10
Marasmiaceae 7
Meruliaceae 2
Mycenaceae 6
Phallaceae 2
Physalacriaceae 6
Pleurotaceae 6
Polyporaceae 24
Psathyrellaceae 17
Russulaceae 28
Sarcoscyphaceae 1
Schizophyllaceae 10
Sclerodermataceae 4
Strophariaceae 6
Thelephoraceae 2
Tricholomataceae 11
Xylariaceae 4
1.4 Future Prospects
Assessment of wild mushroom diversity in the northeast India is essential as it plays a significant role in the socio-economic life of the tribal population. They are imperative for the rural and sub-urban inhabitants through food security and livelihood. They can build important dietary addition through protein and various micronutrients and, together with their medicinal activities. As this region is still underexplored for mushroom biodiversity, there is a very high chance of obtaining mushrooms with high potential in diverse applications.
Acknowledgements The authors are thankful to the Department of Biotechnology, Government of India for DBT sponsored NER-Twinning project (No. BT/320/NE/TBP/2012). We gratefully acknowledge Chief Wildlife Warden, Environment and Forest Department, Government of Mizoram, India. We are grateful for Department of Biotechnology, Government of India for establishing DBT-BIF centre and DBT-State Biotech Hub in the Department of Biotechnology, Mizoram University.
Fig. 1.3 Major families recorded from Dampa Tiger Reserve and Murlen National Park, Mizoram, India
Lallawmsanga et al.
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Chapter 2 A Global Overview of Edible Mushrooms
Malarvizhi Kaliyaperumal, Kezhocuyi Kezo, and Sugantha Gunaseelan
2.1 Introduction
Fungi are one of the most diverse and prominent organisms to inhabit and influence the earth. They are an essential component of ecosystem in recycling the mineral nutrients by acting as agents of decaying. Members of Ascomycota and Basidiomycota , under a precise combination of various abiotic conditions and surrounding flora are known to produce a detectable fruiting body called as “mushrooms” (Stojchev 1995). According to Chang and Miles (1992), “macrofungus are naked to eyes and are able to grow above ground (epigeous) and underground (hypogeous). They might have originated from ancient lineage ca. 400 million years ago and flourished in association with land plants as both saprobes and parasites (Boyce et al. 2007).
Considering the rich magnitude of fungal diversity, total of estimated mushroom diversity available to science is very less (Hawksworth 1991, 2001). While within reported mushrooms, only 50% (7000 species) acquire varying degrees of edibility; ≤ 3000 known species belongs to 31 different genera; ca. 1-10 % are poisonous mushrooms (Miles and Chang 1997). Mushrooms are recognized as rich sources of diverse bioactive principles that make them medically significant as theapeutic agents against pathogens, curing many health disorders and diseases (Wasser and Weis 1999; Lindequist et al. 2005; Ajith and Janardhanan 2007). Nevertheless many edible mushrooms have been integrated with human life since ancient times. Mycophagy is the act of consuming mushrooms. Hay (1887), a well-known British mycologist proposed the exclusive terms “mycophilia” and “mycophobia”. Mycophilic societies refer to the peoples who like and appreciate mushrooms since ancient time. Mycophobic societies comprise of people showing aversion and fear
M. Kaliyaperumal (*) · K. Kezo · S. Gunaseelan
Centre for Advanced Studies in Botany, Guindy Campus, University of Madras, Chennai, Tamil Nadu, India
B. P. Singh et al. (eds.), Biology of Macrofungi, Fungal Biology, https://doi.org/10.1007/978-3-030-02622-6_2
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In shape they resemble a globe or a flask with a narrow mouth, through which the spores are ejected (peronocarpic ascocarps). Different kinds of reproduction—conidia, pycnidia (chiefly with microconidia), chlamydospores, and perithecia—are found in the same species. The various stages in the life-history of these Fungi are so dissimilar, that formally they were considered to be different genera. Ergot furnishes a very good example.
F . 114. A small portion of an ovary attacked with Claviceps purpurea (Sphacelia)
F . 115. An ovary with the conidial stage of Claviceps purpurea (Sphacelia).
This family may be subdivided into 3 sub-families.
Sub-Family 1. Hypocreales.
The perithecia are pale, fleshy, brightly coloured, and generally aggregated on a stroma. Conidia and chlamydospores occur very frequently. Only one order
Order. Hypocreaceæ. In this order the majority are parasites upon Flowering-plants (Nectria, Polystigma, Epichloë, Claviceps); but some are parasites upon Fungi (Hypomyces, Melanospora), or upon insects (Cordyceps).
F . 116.—Claviceps purpurea. A Sclerotium with stromata (cl) (× by 2). B Stroma divided longitudinally to show the perithecia (cp). C A perithecium with the surrounding hyphæ (hy). D An ascus ruptured, with the eight filamentous ascospores emerging.
The most important member of this order is the E (Claviceps purpurea, Figs. 114, 115, 116). This Fungus is found in the flowers of many species of Grasses, especially the Rye, attacking and destroying the ovaries. In the or of the attack, the ovaries are found covered with a white, irregularly folded mycelium (Fig. 114 m, Fig. 115), formed of numerous hyphæ woven together and penetrating the wall of the ovary. From these a number
of hyphæ (Fig. 114 a) project into the air and abstrict from their apices the conidia (b) which serve as reproductive organs. The mycelium also secretes a sticky, stinking fluid (honey-dew) in which the conidia are embedded in great numbers. The honey-dew exudes from the bases of the glumes, and is greedily sought by flies, which thus carry the conidia to other ovaries. In this manner fresh ears are infected, which might escape were the conidia only distributed by the wind. This stage formerly was regarded as an independent Fungus, known as Sphacelia segetum (Fig. 115). On germination, the conidia produce either a new mycelium (Fig. 114 d, c), or new conidia. The or is the one in which the Fungus passes the winter. The mycelium penetrates deeper and deeper into the attacked ovaries, their tissues are destroyed and replaced by the hyphæ, which gradually become more and more felted together. A firm, pseudo-parenchymatous mass of hyphæ is thus formed at the base of the loosely-woven Sphacelia, which is in part transformed into the hard sclerotium, and the remainder thrown off. A dark, hard, poisonous body, longer than the natural grain, is thus formed; these bodies are known as Ergots, and were formerly considered to be a distinct species,—Sclerotium clavus (“Secale cornutum,” Ergot, Fig. 116 A, c). The , described as Claviceps purpurea, is developed in the following spring from the germinating sclerotium, which produces dark-red stromata with short stalks. In the stroma numerous perithecia with asci and ascospores are produced. The latter may infect young flowers of the cereals, in which the disease is then developed as before.
F . 117. Nectria cinnabarina: a branch of Acer pseudoplatanus, with conidial-layers and perithecia (nat. size); b a conidial-layer (Tuberculoria vulgaris); c, a mass of perithecia. (b and c × 8.)
Several species of the genus Nectria, with blood-red perithecia, are found as dangerous parasites, especially N ditissima, which causes “Canker” in the Beech, Ash, and Apple, etc ; N cucurbitula, which appears on Pine-trees, and N cinnabarina (Fig. 117), whose conidial form was formerly named Tubercularia vulgaris. Polystigma rubrum forms shining red spots on the green leaves of Prunus-species. Epichloë typhina is parasitic on the sheaths of Grasses, on which it first forms a white conidial-layer, later on a yellow layer of perithecia. Cordyceps (Chrysalis Fungus, Figs. 118, 119) lives in and destroys insects, and after compassing their death produces the club-formed, generally yellow, stromata,
one part of which bears conidia (Isaria) and another perithecia. C. militaris (Fig. 118) on the chrysalides and caterpillars of moths, is the most common.
The so-called Botrytis bassiana, which produces the disease known by the name of “Muscardine,” in silkworms, is probably a conidial form belonging to Cordyceps.
F 118 Cordyceps militaris I Stromata with conidiophores (Isaria farinosa) II A larva, with stromata, bearing perithecia. III A spore.
F . 119.—Cordyceps robertii on the larva of Hepialus virescens: a stalk of stroma; b perithecia.
Sub-Family 2. Sphæriales.
To this sub-family belong the majority of the Pyrenomycetes. The perithecia are of a firm consistence (tough, leathery, woody or carbonaceous), and of a dark colour. Their covering is quite distinct from the stroma when this structure is present. The stromata are sometimes very large, and may be either cushion-like, crustaceous, upright and club-like, or branched bodies. In general, small,
inconspicuous Fungi, living on dead vegetable matter, sometimes parasites. Free conidiophores and conidiocarps are known in many species, and in several, chlamydospore-like forms of reproduction. Orders 3–18 constitute the Sphæriaceæ of older systematists.
F . 120. Strickeria obducens: a a portion of an Ash-branch with the bark partly thrown off; on the wood are numerous black perithecia (× 20); b longitudinal section through a perithecium; c a spore; d longitudinal section through a pycnidium whose ascospores are being ejected; e portion of the same, with hyphæ and spores
Order 1. Sordariaceæ. Fungi living on dung with fragile perithecia, either aerial or buried in the substratum. The dark brown or black spores have either a mucilaginous envelope (Sordaria, etc.) or mucilaginous appendages (Podospora), by means of which their expulsion and distribution are promoted.
Order 2 Chætomiaceæ. Perithecia fragile, free, bearing on the summit a tuft of hairs Chætomium, on decaying vegetable matter
Orders 3–7. Perithecia scattered or aggregated, situated from the commencement on the surface of the substratum. Stroma wanting.
Order 3. Trichosphæriaceæ. Trichosphæria parasitica (Fig. 121), on Abies alba; Herpotrichia nigra on Picea excelsa and Pinus montana.
F . 121. Trichosphæria parasitica: a a twig of Abies alba, with epiphytic mycelium; b a leaf with mycelium and sporangia (magnified); c a sporangium (× 60); d an ascus with spores (× 550).
Order 4. Melanommaceæ. Rosellinia quercina lives in the roots of 1–3-year-old Oaks, and destroys the plants.
Order 5. Ceratostomaceæ.
Order 6. Amphisphæriaceæ. Strickeria obducens (Fig. 120) has brick-like spores, and lives aggregated on the hard branches of Fraxinus.
Order 7 Lophiostomaceæ.
Order 8. Cucurbitariaceæ. Perithecia tufted, at first embedded, then breaking through, often situated upon an indistinct stroma.
Orders 9–13 The perithecia remain embedded, and are only liberated by the casting off of the covering layers of the substratum Stroma wanting
Order 9. Sphærellaceæ. The species of Sphærella have colourless, bicellular spores. They live upon the leaves of many plants, and develope spherical perithecia upon the fallen leaves
Order 10. Pleosporaceæ. The conidial-forms of Pleospora herbarum and P. vulgaris form a black covering on various plants, known as “smuts.” Venturia ditricha occurs on the underside of dry Birch leaves, and perhaps to this belongs the conidial-form, Fusicladium pirinum, which causes the “Rust spots” on Apples and Pears.
Order 11 Massariaceæ.
Order 12. Clypeosphæriaceæ.
Order 13 Gnomoniaceæ. Perithecia, with peak-like aperture Gnomonia erythrostoma in the leaves of Prunus avium, which turn brown and do not fall in autumn.
Orders 14–18. Stroma generally well developed. The perithecia are embedded in the stroma, but when this is rudimentary, in the substratum.
Order 14. Valsaceæ. Valsa.
Order 15. Diatrypaceæ. Diatrype.
Order 16. Melanconidaceæ.
Order 17 Melogrammataceæ.
Order 18. Xylariaceæ. This order is the most highly developed of the Sphæriales. The stroma arises on the surface of the substratum, which is generally dead or decorticated wood; it is well-developed, crustaceous, hemispherical or upright. In the younger conditions it is covered with a layer of conidia, and later on it bears the perithecia, arranged in a layer immediately beneath its surface. The ascospores are of a dark colour. Often also there are free conidiophores.
F 122 Xylaria hypoxylon (nat size) on a tree stump: a younger, b an older stroma, both of which, with the exception of the black lower portion, are covered with white conidia; n, spot where the perithecia are developed; c an old stroma with upper part fallen off; d, e large branched stromata; k conidia
Hypoxylon and Ustulina have a cushion-like or crustaceous stroma. Xylaria has a club-shaped or branched stroma, often several centimetres high. X. hypoxylon (Fig. 122) and X. polymorpha occur on old tree stumps. Poronia grows on old horse dung, and has a conical stroma.
Sub-Family 3 Dothideales.
The perithecia are always embedded in a black stroma, and are not distinctly separated from it. The accessory forms of reproduction are: conidiophores, conidiocarps, and yeast-like conidia. The majority are parasites. One order.
Order Dothideaceæ. Phyllachora graminis produces scab-like patches on the leaves of the Grasses. Scirrhia rimosa grows on the leaf-sheathes of Phragmites. Rhopographus pteridis on Pteridium aquilinum
Family 4. Hysteriales.
This family, like the following, has hemiangiocarpic ascocarps (apothecia). These are closed in the early stages, but when ripe open in a valvular manner by a longitudinal fissure; they are black, oblong, and often twisted. Some species are parasites, especially upon the Coniferæ.
F . 123.—Lophodermium (Hypoderma) nervisequium: a two leaves of Abies alba seen from above with pycnidia; b a leaf seen from the underside with apothecia; c an ascus with ascospores. (× 500.)
F 124 Three leaves of the Redpine with Lophodermium macrosporum: a under side of the leaves with apothecia; b a leaf from upper side with pycnidia (× about 2 )
F 125 Lophodermium pinastri: a leaves of Pinus sylvestris with apothecia (nat size); b two paraphyses and an ascus with filamentous spores
Order 1. Hysteriaceæ. Hysterium pulicare upon the ruptured bark of many trees.
Order 2. Hypodermaceæ. The species of Lophodermium live upon the leaves of Conifers, and are the cause of their falling off (blight). L. pinastri (Fig. 125), on the leaves of Pinus and Picea; the leaves become red-brown and fall off; at first conidiocarps are formed, and later apothecia; L. nervisequium (Fig. 123), on Abies alba; L. macrosporum (Fig. 124), on Picea excelsa; L. brachysporum, on Pinus strobus.
Order 3 Dichænaceæ.
Order 4. Acrospermaceæ.
Family 5. Discomycetes.
The ascocarps (apothecia) are at first closed, and only open at the time of their ripening, not valvularly, but more or less like a saucer or cup, so that the hymenium lies exposed on their upper surface. In the first three sub-families, and generally also in the fourth, the apothecia are formed inside the substratum. The apothecia are, in contrast to the Pyrenomycetes, light and brightly coloured, and their size varies very much, and may be several centimetres in diameter. Paraphyses are often present between the asci; they often contain colouring matter, and give to the disc its characteristic colour. The tissue on which the asci are borne is known as the hypothecium. The shape and colour of the spores is not so varied as in the Pyrenomycetes. The accessory forms of reproduction are conidia (sometimes of two forms), chlamydospores, and oidia. The family is divided into 5 sub-families.
Sub-Family 1. Phacidiales.
The apothecia are developed in the interior of the substratum, which they break through, and in general dehisce apically. The envelope is tough and black. Hypothecium inconspicuous; hymenium flat.
Order 1. Euphacidiaceæ. Phacidium abietinum, on the leaves of Abies alba. Rhytisma; the pycnidia are found in the summer on the green leaves, while the apothecia are developed on the fallen leaves and dehisce in the following spring. R acerinum causes black spots on the leaves of the Sycamore, and R salicinum on Willows
Order 2. Pseudophacidiaceæ.
Sub-Family 2. Stictidales.
The apothecia when ripe break through the substratum which forms a border round them. Hymenium generally saucer-shaped.
Order 1. Stictidaceæ. Stictis.
Order 2. Ostropaceæ. Ostropa.
Sub-Family 3 Tryblidiales.
The apothecia are embedded in the substratum in the early stages, and then are raised high above it. Hypothecium thick. Hymenium cup-shaped.
Order 1 Tryblidiaceæ. Tryblidium
Order 2. Heterosphæriaceæ. Heterosphæria patella on the dead stalks of Umbellifers.
Sub-Family 4. Dermateales.
The apothecia in the early stages are embedded in the substratum and then break through it, or are from the first situated on the surface of the substratum. Hypothecium thick.
Order 1. Cenangiaceæ. Cenangium.
Order 2 Dermateaceæ. Dermatea
Order 3. Patellariaceæ. Patellea, Biatorella, Patellaria.
Order 4 Caliciaceæ. Calicium, Coniocybe, etc , on the bark of trees
Order 5. Arthoniaceæ. Arthonia on the bark of several trees. Celidium stictarum on the apothecia of Sticta pulmonaria.
Order 6 Bulgariaceæ. Apothecia gelatinous under moist conditions, and horny when dried —Calloria fusarioides; the red apothecia break out in the spring on the dried stalks of Urtica dioica; a gelatinous reproductive form of the Fungus is found before the apothecia, which consists of oidia (formerly described as “Dacryomyces urticæ”). Bulgaria inquinans on the living or fallen trucks of Oaks and Beeches.
F 126 Botrytis cinerea: a slightly magnified; b more highly magnified; c germinating conidium
F 127 Sclerotinia fuckeliania: a sclerotium with conidiophores; b with apothecia; c section through sclerotium and apothecium; d ascus with eight ascospores (× 390 )
Sub-Family 5. Pezizales.
The apothecia are developed on the surface of the substratum and are waxy or fleshy; at the commencement closed, and covered with a saucer- or cup-shaped, seldom flat, hymenium. The
hypothecium is generally well developed. This sub-family is the richest in species of the Discomycetes and contains forms of very different habit. They grow upon dead wood, upon the ground, and upon dung. A few are parasites.
Order 1. Helotiaceæ. Apothecia with waxy envelope of colourless, or yellowish prosenchymatous cells.—Chlorosplenium æruginosum is found on decaying wood (particularly Oak and Birch), to which it gives a green colour Sclerotinia has sclerotia which are developed upon the hostplant and from which, after a period of rest, the long, brown-stalked apothecia arise. S. ciborioides (S. trifoliorum, Fig. 128) is parasitic on Clover; S. sclerotiorum, on Daucus-roots, Phaseolus, etc.; S. baccarum, on the berries of Vaccinium myrtillus; “Botrytis cinerea” is a common parasite and is probably the conidial form of S fuckeliania (Fig 127) Helotium herbarum lives on dry plant stems Dasyscypha willkommii (Fig. 129) produces Larch-canker on the bark of the Larch.
