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Collection Highlights

Developments in Fungal Biology and Applied Mycology 1st

Edition Tulasi Satyanarayana

Neotropical Social Wasps: Basic and applied aspects Fabio

Prezoto

Physiology and Genetics Selected Basic and Applied Aspects

2nd Edition Timm Anke

Systemic Lupus Erythematosus: Basic, Applied and Clinical

Aspects 2nd Edition George Tsokos (Editor)
Thermophilic fungi : basic concepts and biotechnological
applications First Edition Salar

Basic and Applied Phytoplankton Biology Perumal Santhanam

Recent Trends in Human and Animal Mycology Karuna Singh

The Heparins: Basic and Clinical Aspects 1st Edition David

Green

Textbook of Medical Mycology 4th Edition Chander

Tulasi Satyanarayana
Sunil Kumar Deshmukh
Mukund V. Deshpande Editors

Advancing
Frontiers in
Mycology &
Mycotechnology
Basic and Applied Aspects of Fungi
Advancing Frontiers in Mycology
& Mycotechnology
Tulasi Satyanarayana
Sunil Kumar Deshmukh
Mukund V. Deshpande
Editors

Advancing Frontiers
in Mycology
& Mycotechnology
Basic and Applied Aspects of Fungi
Editors
Tulasi Satyanarayana Sunil Kumar Deshmukh
Division of Biological Science and Biotech & Management of
Engineering Bioresources Div
Netaji Subhas University of Technology The Energy and Resources Institute
New Delhi, Delhi, India New Delhi, Delhi, India

Mukund V. Deshpande
Division of Biological Sciences
CSIR-National Chemical Laboratory
Pune, Maharashtra, India

ISBN 978-981-13-9348-8    ISBN 978-981-13-9349-5 (eBook)

https://doi.org/10.1007/978-981-13-9349-5

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Foreword

The term ‘mycology’ dates back to the eighteenth century, and in its formative
years, the study of fungi was guided by reputed botanists like C. H. Persoon,
E.M. Fries and A. deBary. During the nineteenth and twentieth centuries, phenom-
enal progress in classical mycology took place, including descriptive morphology,
identification, classification and taxonomy of fungi primarily, due to the efforts of
botanists, despite the fact that fungi are not phylogenetically related to plants. The
reason for this essentially stems from the devastating fungal diseases in plants, nota-
bly the crop plants, causing serious problems in agriculture. There was an urgent
need to understand the aetiology of the destructive fungal diseases and devise suit-
able control measures to save the crops. As an example may be mentioned the stud-
ies in India by Prof. M.J. Narasimhan (1930) who collected and studied diverse
heterothallic strains of the destructive Phytophthora1 infection on areca nut palms
and devising fungicidal treatment schedules for controlling the disease in the rain-
forest areas of Karnataka (Mysore) state.
During the early years, traditional mycological knowledge facilitated the (a)
development of various oriental fermented foods, such as miso, tempeh and sake,
based on strains of fungi such as Aspergillus oryzae, Rhizopus oligosporus and oth-
ers (Hesseltine 1983), (b) differentiation of edible and poisonous mushrooms and
(c) understanding the aetiology of ergot poisoning that caused serious convulsions
due to consumption of the ergot sclerotia contaminating the grain which is con-
sumed as food.
During the twentieth century, several milestone advances in biological sciences
based on mycological studies can be recognized. As examples, we may mention the
following:

(a) The Nobel Prize-winning (1958) studies on Neurospora by G.W. Beadle and
E.L. Tatum which resulted in their discovery of how genes act by regulating
definite chemical events that in turn affected the development in organisms.
Based on ingenious selection of mutants deficient in the synthesis of single
growth factors, Beadle and Tatum concluded that mutations to genes affected

1
Recently, the genera Phytophthora and Pythium have been placed in Kingdom Chromista or
Kingdom Straminipila, distinct from Kingdom Fungi (Ho 2018).

v
vi Foreword

the enzymes of organisms. It is noteworthy that these studies provided the early
link between genetics and the upcoming new scientific field of molecular
biology.
(b) The discovery of truly anaerobic chytrids present in the rumen of herbivores
and playing a significant role in the digestion of the cellulosic feed is a mile-
stone discovery of the twentieth century. Classified under Neocallimastigaceae,
these chytrids with chitinous cell walls are truly anaerobes lacking mitochon-
dria, cytochromes and other biochemical features of the oxidative phosphoryla-
tion pathway. The group possesses organelles (hydrogenosomes) for a major
part of anaerobic energy metabolism (Nagpal et al. 2009).

Taxonomy and classification of fungi have been central to the mycological stud-
ies, and this has led to a well-established database for understanding fungal biodi-
versity. Classical taxonomy relied upon the morphological features of the spore
forms. With the advent of biochemical and molecular data becoming available, sev-
eral conceptual changes in fungal taxonomy have arisen. While giving an opportu-
nity to gain newer knowledge, these developments also pose several challenges and
opinion divides in their interpretation and implementation.
Classical taxonomy classified all the zoospore-forming fungi under Phycomycetes
considering their monophyletic origin from algae. Recent advances in biochemical
knowledge have established that the Chytridiomycetes2 with chitinous cell walls,
despite being zoospore-forming, are distinct from the rest of the zoospore-forming
fungi with β-glucan in their cell walls, and these are grouped together under the
Oomycota (as opposed to the Eomycota under which the Zygomycetes, Ascomycetes
and Basidiomycetes are classified). The term Phycomycetes as originally adapted is
no longer valid and, hence, abandoned from present-day mycological literature.
DNA sequencing and bar-coding of fungal species are major developments
which offer challenges to accepted morphology-based taxonomy. The nuclear ribo-
somal internal transcribed spacer (ITS) is widely used as a DNA bar-coding marker
to characterise the diversity and composition of fungal communities. Since the
1990s, the ITS region has been extensively used in molecular methods as well as
ecological studies on fungi.
Estimates of fungal biodiversity have undergone sharp changes following envi-
ronmental DNA sequencing of fungal ITS reads. Currently, there are around 81,000
accepted fungal names out of a total of up to 5.1 million species (Blackwell 2011).
Environmental DNA sequencing has accumulated more than one million fungal ITS
reads. Whether ‘sequence-based voucher-less fungi’ (in the absence of actual speci-
mens) should be given a formal nomenclature is widely debated, and the opinions
of expert mycologists are sharply divided on this point with reference to the estima-
tion of biodiversity of fungi in the natural environment. Several informative reviews
on DNA bar-coding as well as sequence-based fungal nomenclature are published
and may be consulted for a more complete understanding of the problem and its

2
Now chytrids are grouped under Chytridiomycota (N. P. Money, in The Fungi (Third Edition),
2016)
Foreword vii

perspectives (Shenoy et al. 2007; Begerow et al. 2010; Das and Deb 2015; Hibbett
et al. 2016; Hongsanen et al. 2018).
Conventionally, taxonomic principles related to fungi are debated and decisions
taken under the authority of the International Code of Botanical Nomenclature
(ICBN) held once in 6 years during the International Botanical Congress. Recently,
the ICBN has been renamed as International Code of Nomenclature for algae, fungi
and plants. In the Melbourne ICBN held in 2011, a momentous decision was taken
to abolish Article 59 which permitted the use of dual nomenclature for pleomorphic
fungi. In the morphology-based taxonomy, anamorphs (asexual forms) and teleo-
morphs (sexual forms) of fungi discovered and described independently could
receive different but valid names as ruled by Article 59 of the ICBN. In the present
era of phylogenetic molecular analysis, co-ordination of anamorphic and teleomor-
phic elements of a fungus can, in principle, be unequivocally established. Molecular
mycologists are strongly in favour of the unification of the fungal nomenclature and
achieve the ‘one fungus-one name’ ideal. However, there are several practical hur-
dles, and several specialist committees are in serious debates to elucidate the practi-
cal feasibility of implementing the same (Gams 2014).
Simultaneous with the focused attention on fungi as troublesome pathogens of
plants, animals and humans, beneficial attributes of fungi as sources of valuable
metabolites were also realized. Taka-Diastase from A. oryzae and citric acid from
Aspergillus niger are examples of fungal-based processes of manufacture in the late
nineteenth and early twentieth centuries. With the discovery of penicillin and the
dawn of the antibiotic era, the decades of the twentieth century became an era for
screening diverse microorganisms including fungi for discovering novel bioactive
molecules and producing them by fermentation. Several industrially useful enzymes,
hormones, vitamins and growth factors of fungal origin spearheaded the fermenta-
tion technology development. Fungi and their metabolites were also employed in
the biotransformation of organic molecules (e.g. steroids).
The beneficial attributes of fungi far outweighed their earlier negative image, and
this led to a very healthy and positive attitude towards fungi which deserved to be
studied for deriving beneficial products from their metabolism. The term ‘biodiver-
sity’ acquired a newer dimension in terms of its relevance to applied microbiologi-
cal and biotechnological research. An immense focus on the physiological,
biochemical and molecular aspects of mycodiversity became imminent for biopro-
cess development based on novel fungal strains.
Heterologous gene expression and production of mammalian proteins in fungal
hosts have been successfully carried out for chymosin (cheese manufacture enzyme)
in A. niger var. awamori and insulin in Komagataella (Pichia) pastoris.
Spectacular advances in both mycology and mycotechnology during the twenti-
eth century have added new dimensions to mycological research, necessitating
newer approaches to achieve success in this field at present. A different mindset and
training have become necessary in order to carry out meaningful research projects
viii Foreword

in the twenty-first century. Some of the salient features of this new preparation may
be defined as follows:

(a) Knowledge-based exploration of fungal biodiversity in different ecosystems by

mycologists is an important aspect. Understanding the type of habitats in which
specific types of fungal populations would be naturally enriched will be very
helpful. For example, screening for thermophiles in self-heating composts,
bagasse piles or sun-heated soils would be very useful. The utilization of special
traits in specific groups of fungi is also very beneficial, such as the overlaying
of agar plates with moistened detritus to allow colonies to develop from forcibly
‘shot’ spores in the case of saprophytic Entomophthorales like Conidiobolus
and Basidiobolus as well as yeasts like Sporobolomyces. Adopting selective iso-
lation techniques including the use of specific inhibitory chemicals or antibiot-
ics has significantly enabled the slower growing and rarer populations to be
isolated on agar plates overcoming competition from the rapidly growing fungal
populations. The topic of selective isolation is a vast one and has been reviewed
(Srinivasan 2004, 2008) which may be consulted.
(b) Apart from taxonomic studies, in this biotechnology era, mycologists have the
responsibility of conserving their discoveries in pure culture through optimiza-
tion of culture preservation methods suited for their specific strains. They have
to ensure both morphological and genetic stability of their cultures under the
laboratory conditions for prolonged periods. Repeated and frequent subcultures
on sugar-rich media, like potato dextrose agar, often adversely affect sporula-
tion, particularly in strains associated with decomposing plant litter. While
freeze-drying (lyophilisation) is widely accepted as a safe method of long-term
conservation, it must be verified in regard to its suitability for the strains under
study. It is experienced that some fungi (e.g. Conidiobolus) failed to survive
lyophilisation, while many of the lignocellulose-degrading hyphomycetes
showed reduced sporulation upon revival from lyophilized cultures. After in-­
depth studies to standardize the optimum conservation method(s), mycologists
in India should deposit their cultures in the national fungal germplasm banks
such as the National Fungal Culture Collection of India (NFCCI), Agharkar
Research Institute, Pune; Microbial Type Culture Collection (MTCC), CSIR-­
Institute of Microbial Technology, Chandigarh; National Centre for Microbial
Resource (NCMR), National Centre for Cell Science, Pune; or the National
Collection of Industrial Microorganisms (NCIM), CSIR-National Chemical
Laboratory, Pune. While depositing the cultures, it is mandatory that the full
information on the optimal methods of conservation for the strain(s) is given to
the curators of the culture collections to follow for successful conservation.
(c) When exploring fungal diversity for novel metabolites, it may be necessary to
pay attention right from the isolation stage, keeping in mind the purpose of the
strain screening. I shall illustrate this point taking the case of ‘endophytic fungi’
and exploration of their potential to produce the plant metabolite by fermenta-
tion. Widespread interest in exploring endophyte fungal populations of diverse
medicinal plants has arisen after the report from Prof. Gary Strobel’s laboratory
Foreword ix

in the USA that an endophyte designated Taxomyces andreanae produced the

valuable anticancer metabolite taxol by fermentation (Stierle et al. 1993).
Presently, diverse endophytic fungi have been isolated and maintained in pure
culture in several laboratories with the purpose of studying their potential to
produce the valuable plant metabolites by fermentation. A point that deserves to
be seriously given consideration is as follows:

The endophytic fungus has imbibed the genes from the medicinal plant through
constant association. In the laboratory, culturing it on routine sugar-rich media
while facilitating satisfactory growth, there is no selection pressure for the growing
fungus to conserve these imbibed genes, and over several subcultures on routine
media may even be lost. An alternate suggestion which may be worth considering is
to plan the culture isolation of the endophyte on a relatively simple nutrient medium
fortified with some of the key chemical intermediate components involved in the
biosynthesis of the specific plant metabolite. It is possible on such a medium, the
metabolic genes for the plant metabolite are conserved better, and these may be
subjected to mutation and gene amplification through molecular techniques, leading
to strains which will be able to produce the metabolite at commercially viable lev-
els. Presently, the levels of production observed are very low and insignificant, and
much more studies are warranted before practical and viable technologies for
endophyte-­based plant metabolites would become a reality.
Recent years have witnessed increased emphasis on the beneficial attributes of
fungi, recognizing their role in the natural ecosystems and as the source of several
metabolites of value for humankind. Rambold et al. (2013) in an article, suggesting
that mycology should be recognized as a major field in biology, stated ‘Given the
ecological and economic relevance of fungi...Mycology is insufficiently recognized
as a major field of science’. In the theme for the International Mycological Congress
held at Puerto Rico in July 2018, mycological discoveries for a better world and
how fungi contributed to the health of society and of ecosystem were given empha-
sis. Clearly, one can perceive the changing face of mycology from ‘fungi as destruc-
tive agents to be despised to fungi as true benefactors of humanity to be adored’.
In order to harness the full potential of fungi for mankind’s benefit, mycologists
must identify themselves as part of multidisciplinary teams exploring fungi for
novel metabolites. They should broaden their horizons of interest and acquire
knowledge in physiology, biochemistry and molecular biology of fungi. Familiarity
with natural product chemistry and bioprocess engineering for mould metabolites
including mould morphogenesis in fermentors related to growth conditions for
maximum yields are desirable.
Multidisciplinary collaboration to achieve success in technology is a must, and
the status of mycology in the twenty-first century is meaningfully reflected in the
following quote from Kreger (2003):
We all realise that most scientists in the future will be part of multi-disciplinary research
teams…the shift is causing a way scientists need to train … although they must still be an
expert in their specialty, they must also become conversant with techniques that once
seemed beyond their domain … they also need to recognize where their knowledge ends and
where they should seek the help from others….
x Foreword

It gives me pleasure in writing the foreword for this book, Advancing Frontiers
in Mycology and Mycotechnology: Basic and Applied Aspects of Fungi. The invited
chapters included in this book cover most of the aspects of fungal biology and
applied aspects of fungi. I wish to congratulate the editors and all the contributors
for their efforts in bringing out this book. I sincerely hope and wish that the book
will be useful for students, scholars, scientists and teachers of biology, microbiol-
ogy and biotechnology.

Ex-Head, Biochemical Sciences Division

M. C. Srinivasan
CSIR-NCL
Pune, Maharashtra, India
R.H.17, Planet Millennium
Pune, Maharashtra, India

References
Begerow D, Nilsson H, Unterseher M, Maier W (2010) Current state and perspectives of fungal
DNA bar coding and rapid identification procedures. Appl Microbiol Biotechnol 87:99–108
Blackwell M (2011) The fungi: 1, 2, 3 … 5.1 million species ? Am J Bot 98: 426–438
Das S, Deb B (2015) DNA bar coding of fungi using ribosomal ITS marker for genetic diversity
analysis: a review. Int J Pure Appl Biosci 3:160–167
Gams W (2014) A new nomenclature for fungi. Mycol Iranica 1:1–5
Hesseltine CW (1983) Microbiology of oriental fermented foods. Annu Rev Microbiol 37: 575–601
Hibbett D, Abarenkov K, Kõljalg U, Öpik M, Vhai B, Cole J, Wang Q, Crous P, Robert V, Helgason
T, Herr JR, Kirk P, Lueschow S, O’Donnell K, Nilsson RH, Oono R, Schoch C, Smyth C,
Walker DM, Porras-Alfaro A, Taylor JW, Geiser DM (2016) Sequence based classification and
identification of fungi. Mycologia 108:1049–1068
Ho HH (2018) The taxonomy and biology of Phytophthora and Pythium. J Bacteriol Mycol Open
Access 6(1):40–45. https://doi.org/10.15406/jbmoa.2018.06.00174
Hongsanen S, Jeewon R, Purahong W, Xie N, Liu J-K, Jayawardana RS, Ekanayaka AH,
Dissanayake A, Raspe O, Hyde KD, Stadler M, Pers D (2018) Can we use environmental DNA
as holotypes? Fungal Divers 92:1–30.
Kreeger C (2003) The learning curve. Nat Biotechnol 21:951–952
Nagpal R, Puniya AK, Griffith GW, Goel G, Puniya M, Sehgal JP, Singh K (2009) Anaerobic
rumen fungi: potential and applications. Chapter 17, 375–393. In: Khachatourians GG, Arora
DK, Rajendran TP, Srivastava AK (eds), Agriculturally important micro-organisms, volume 1,
Academic World International
Narasimhan MJ (1930) Studies in the genus Phytophthora X Heterothallic strains of Phytophthora.
Phytopathology 20:201–214
Rambold G, Stadler M, Begerow D (2013) Mycology should be recognised as a field in biology at
an eye level with other major disciplines – a memorandum. Mycol Prog 12:455–463
Shenoy BD, Jeewon R, Hyde KD (2007) Impact of DNA sequence-data on the taxonomy of ana-
morphic fungi. Fungal Divers 26:1–54
Srinivasan MC (2004) Selective isolation of fungi, Chapter 2. In: Practical mycology for industrial
biotechnologists. Tata McGraw-Hill Publishing co. Ltd., New Delhi, p 26–46
Foreword xi

Srinivasan MC (2008) Unconventional techniques to explore fungal diversity for industrial enzyme
technology. In: Sridhar et al (ed) Novel techniques and ideas in mycology. Fungal Diversity
Research Series 26:261–270
Srinivasan MC (2014) Role of mycology and mycologists in an era of industrial biotechnology.
Kavaka 43:3–5
Stierle A, Strobel G, Stierle D (1993) Taxol and taxane production by Taxomyces andreanae, an
endophytic fungus of Pacific yew. Science 260:214–216
Preface

Fungi are members of the group of eukaryotic organisms, which include microbes
like yeasts, moulds and mushrooms. They are classified as kingdom Fungi and more
closely related to animals than to plants, therefore, placed with the animals in the
monophyletic group. Analyses using molecular phylogenetics support a monophy-
letic origin of fungi. They are heterotrophs, as they acquire their food by absorbing
dissolved molecules, typically by secreting digestive enzymes into their environ-
ment. The origin of fungi can be traced to single-celled marine ancestors in the
Mesoproterozoic era more than one billion years ago. Fungi have since then con-
quered not only land but also almost every potential habitat and substrate. Although
all fungi are heterotrophs, the fungal kingdom comprises a wide range of life strate-
gies ranging from saprotrophy through mutualism to parasitism. The fungal king-
dom encompasses an enormous diversity of taxa with varied ecologies, life cycle
strategies and morphologies ranging from unicellular aquatic chytrids and yeasts to
large mushrooms. However, very little is known about the true diversity of kingdom
Fungi, which had been estimated at five million species. Fungi are immensely
diverse, with 144,000 species named and classified so far at a current rate of around
2,000 per year. Of these, over 8,000 species are known to be detrimental to plants,
and at least 300 can be pathogenic to human beings.
Our fascination for the fungal kingdom is a natural and ancient one based on the
following: (i) the roles of fungi in the production of a variety of foods and beverages
and even as a source of food themselves; (ii) their global ecological impact, espe-
cially as the cause of devastating infections of humans and other animals and of
plants, including many crops grown around the world; and (iii) their roles as funda-
mental model systems in genetics and biological research.
Since the pioneering eighteenth- and nineteenth-century taxonomical works of
Carl Linnaeus, Christiaan Hendrik Persoon and Elias Magnus Fries, fungi have
been classified based on their morphological, physiological and biochemical char-
acteristics. Advances in molecular genetics have opened the way for DNA analysis
to be incorporated into taxonomy, which has sometimes challenged the historical
groupings based on morphology and other traits. Phylogenetic studies published in
the last decade have helped to reshape the classification within kingdom Fungi,
which has been divided into seven phyla (Microsporidia, Chytridiomycota,
Blastocladiomycota, Neocallimastigomycota, Glomeromycota, Ascomycota and
Basidiomycota).

xiii
xiv Preface

Along with bacteria, fungi are the major decomposers in most terrestrial and
some aquatic ecosystems; therefore, they play an essential role in nutrient cycling,
especially as saprobes and symbionts, degrading organic matter to inorganic mole-
cules, which can then re-enter anabolic metabolic pathways in plants and other
organisms. Certain fungi, in particular white-rot fungi, can degrade insecticides,
herbicides, pentachlorophenol, creosote, coal tars and heavy fuels and turn them
into carbon dioxide, water and basic elements. Fungi have been shown to mineralize
uranium oxides, suggesting that they may have application in the bioremediation of
radioactively polluted sites.
The human use of fungi for food preparation and other purposes is extensive and
has a long history. Mushroom farming and mushroom gathering are large industries
in several countries. The global commercial mushroom market was ~US$35 billion
in 2015 and is anticipated to grow to as much as ~US$60 billion by 2021.
Ethnomycology is the study of the historical uses and sociological impact of fungi.
Fungi have the capacity to produce an enormous range of natural products with
antimicrobial or other biological activities; therefore, many species have long been
used or are being developed for industrial production of antibiotics, vitamins and
anticancer and cholesterol-lowering drugs. Very recently, methods have been devel-
oped for genetic engineering of fungi that enable metabolic engineering of fungal
species. For example, genetic modification of yeast species, which are easy to grow
at fast rates in large fermentation vessels, has opened up ways of pharmaceutical
production that are potentially more efficient than production by the original source
organisms.
Several pivotal discoveries in biology were made by researchers using fungi as
model organisms. Indeed, seven Nobel prizes have been awarded to scientists study-
ing yeasts and moulds as model organisms that explain fundamental aspects of cell
biology: from Alexander Fleming, Ernst Chain and Howard Florey in 1945 for the
discovery of penicillin from Penicillium notatum; to George Beadle and Edward
Tatum in 1958 for their ‘one gene-one enzyme’ hypothesis in Neurospora crassa; to
Paul Nurse and Leland Hartwell in 2001 for cell division and cancer in
Schizosaccharomyces pombe and Saccharomyces cerevisiae; to Roger Kornberg in
2006 for eukaryotic gene transcription in S. cerevisiae; to Jack W. Szostak (shared)
in 2009 for chromosome telomeres in S. cerevisiae; to Randy Schekman in 2013
(shared) for machinery regulating vesicle traffic in S. cerevisiae; and to Yoshinori
Ohsumi in 2016 for autophagy in S. cerevisiae. Other important model fungi, which
includes Aspergillus nidulans; Candida albicans, a dimorphic, opportunistic human
pathogen; Magnaporthe grisea, a plant pathogen; and Pichia pastoris, a yeast
widely used for eukaryotic protein production, have more recently emerged, which
address specific biological questions relevant to medicine, plant pathology and
industrial uses.
In fact, a fungal species was the first eukaryotic organism to have its genome
completely sequenced (the model budding yeast S. cerevisiae). Advances in genet-
ics and cell biology have contributed to provide a detailed view of how the genome
contributes to the functions of the cell and of the organism. Together, these advances
in genomics and genetics provide a ‘blueprint’ for how these species operate and
Preface xv

have evolved at a cellular level, and consequently, they offer a wealth of knowledge
about how representative species in the fungal kingdom function and the diversity
that lies within. This diversity spans from the most basic way that a fungal cell is
organized, either as a yeast or as a filamentous hypha, to the myriad ways these spe-
cies interact with their environment, from aquatic basal fungi (Chytridiomycota,
Cryptomycota) to fungi that are associated with plants and were critical for their
emergence from the oceans and colonization of the planet to fungi that are patho-
gens of plants or animals. This diversity also extends to the biological behaviour and
cell biology of fungi, including fungi that can sense light and those that have evolved
to be insensitive to light (blind), the modes of sexual reproduction including hetero-
thallism and homothallism, the loss and retention of RNAi pathways, the replace-
ment of regional centromeres by point centromeres and the retention of flagella in
basal fungi versus their loss in fungal branches that evolved the ability to be aerially
dispersed.
This book entitled Advancing Frontiers in Mycology and Mycotechnology: Basic
and Applied Aspects of Fungi is aimed at reviewing major recent developments in
understanding the biology and potential biotechnological applications of fungi.
Parts I, II, III and IV focus on the basic aspects of fungi; their role in environmental
sustainability; their interactions with humans, plants and animals; and bioprospects,
respectively. Part I includes chapters on the diversity of fungi from different envi-
ronments, growth, morphogenesis and genetics and conservation and taxonomy.
Part II deals with the role of fungi in environmental sustainability (biodegradation
and bioremediation) and in nanobiotechnology. The interactions of fungi with
plants, animals and humans, such as mycorrhizal association, endophytism, human
mycoses and mycotoxins, are covered in Part III. The bioprospecting aspects of
entomo- and myco-pathogens, industrial enzymes and secondary metabolites in
healthcare are dealt with in Part IV.
We wish to thank all the contributors for readily accepting our invitation and
submitting their well-written chapters in their areas of specialization within the
stipulated period. We sincerely hope and wish that this book will be useful to stu-
dents, scholars, teachers and scientists in the broad areas of microbiology, life sci-
ences and biotechnology. Thanks are also due to Springer Nature for publishing the
book.

New Delhi, Delhi, India Tulasi Satyanarayana

New Delhi, Delhi, India  Sunil K. Deshmukh
Pune, Maharashtra, India  Mukund V. Deshpande
Contents

Part I Basic Aspects of Fungi

1 The Mystical World of Mushrooms........................................................ 3
V. K. Bhalerao, A. P. Gaikwad, C. D. Deokar,
and K. S. Raghuwanshi
2 The Developmental History of Ustilago maydis:
A Saprophytic Yeast, a Mycelial Fungus,
Mushroom-Like, and a Smut.................................................................. 49
José Ruiz-Herrera, José L. Cabrera-Ponce, Claudia León-­Ramírez,
Fernando Pérez-Rodríguez, Mayela Salazar-­Chávez,
Alejandro Sánchez-Arreguín, and John Vélez-Haro
3 Biochemical and Molecular Aspects of Dimorphism in Fungi............. 69
Ejaj K. Pathan, Vandana Ghormade, Redeemson Panmei,
and Mukund V. Deshpande
4 Diversity, Ecology and Utilization of Soil Fungi:
Indian Scenario........................................................................................ 95
C. Manoharachary and D. Nagaraju
5 Diversity and Bioprospecting of Yeasts from Extreme
Environments............................................................................................ 117
Shiv Mohan Singh, Nitin Adhapure, and Rohit Sharma
6 Marine Fungal Ecology in the Molecular Era....................................... 143
V. Venkateswara Sarma and Rajesh Jeewon
7 Saccharomyces cerevisiae: Oscillatory Orchestration
of Growth.................................................................................................. 181
David Lloyd
8 Neurospora Genetic Backgrounds Differ in Meiotic Silencing
by Unpaired DNA (MSUD) Strength: Implications
for Dp-­Mediated Suppression of Repeat-Induced Point
Mutation (RIP)......................................................................................... 215
Durgadas P. Kasbekar

xvii
xviii Contents

Part II Environmental Sustainability

9 Fungal Bioremediation: A Step Towards Cleaner Environment......... 229
Darshan M. Rudakiya, Archana Tripathi, Shilpa Gupte,
and Akshaya Gupte
10 Exploring Fungi-Associated Lignocellulose Degradation:
Secretomic and Proteomic Approaches.................................................. 251
Akshay Shankar, Shruti Ahlawat, and Krishna Kant Sharma
11 Fungi the Crucial Contributors for Nanotechnology:
A Green Chemistry Perspective.............................................................. 279
Aliesha Moudgil and Bhushan P. Chaudhari

Part III Interactions with Plants, Animals and Humans

12 Recent Developments in Ectomycorrhizal Research............................ 301
Tanveer Kaur and M. Sudhakara Reddy
13 Rumen Microbiome and Plant Secondary Metabolites (PSM):
Inhibition of Methanogenesis and Improving
Nutrient Utilization.................................................................................. 325
D. N. Kamra and B. Singh
14 Class B-Trichothecene Profiles of Fusarium Species as Causal
Agents of Head Blight.............................................................................. 347
Emre Yörük and Tapani Yli-Mattila
15 Aflatoxin and Ochratoxin A Detection:
Traditional and Current Methods.......................................................... 377
Shraddha Rahi, Priyanka Choudhari, and Vandana Ghormade
16 The Explosion of Brazilian Endophytic Fungal Diversity:
Taxonomy and Biotechnological Potentials........................................... 405
Jadson Diogo Pereira Bezerra, Leticia Francisca da Silva,
and Cristina Maria de Souza-Motta
17 Arbuscular Mycorrhizal Fungi in Alleviation of Cold
Stress in Plants......................................................................................... 435
Thokchom Sarda Devi, Samta Gupta, and Rupam Kapoor
18 Challenges in Invasive Fungal Disease................................................... 457
Arunaloke Chakrabarti and Shreya Singh
19 Diversity of Endophytic Fungi and Their Role in Artificial
Agarwood Production in Aquilaria Tree................................................ 479
Hemraj Chhipa and Sunil K. Deshmukh
Contents xix

Part IV Bioprospects
20 Bioprospecting of Fungal Entomo- and Myco-Pathogens.................... 497
E. K. Pathan, A. V. Patil, and M. V. Deshpande
21 Fungal Enzymes: Sources and Biotechnological Applications............. 515
Naveen Kango, Uttam Kumar Jana, and Ritumbhara Choukade
22 Fungi in Hypogean Environment: Bioprospection Perspective........... 539
S. R. Joshi and Upashna Chettri
23 Secondary Metabolites of Mushrooms: A Potential Source
for Anticancer Therapeutics with Translational Opportunities.......... 563
Sudeshna Nandi, Rimpa Sikder, and Krishnendu Acharya
24 Modulation of Fungal Metabolome by Biotic Stress............................ 599
Geane Pereira de Oliveira, Bruna de Almeida Martins,
Matheus Thomaz Nogueira Silva Lima,
and Jacqueline Aparecida Takahashi
25 Marine Fungi as a Potential Source of Future Cosmeceuticals........... 627
Shivankar Agrawal, Sunil K. Deshmukh, and Colin J. Barrow

Index.................................................................................................................. 671
About the Editors and Contributors

Editors

Prof. T. Satyanarayana is a UGC-BSR Faculty Fellow at the Division of Biological

Sciences & Engineering, Netaji Subhas University of Technology, New Delhi, after
superannuating from the Department of Microbiology, University of Delhi South
Campus, New Delhi, as Professor and Head, in June 2016. He has over 270 scien-
tific papers and reviews, 8 edited books and 2 patents to his credit. He is a fellow of
National Academy of Agricultural Sciences (NAAS), Association of Microbiologists
of India (AMI), Biotech Research Society (I), Mycological Society of India (MSI)
and Telengana Academy of Sciences. He is a recipient of Dr. Manjrekar award of
AMI, Dr. Agnihotrudu award of MSI and Malaviya Memorial award of BRSI. He
has over 40 years of research and teaching experience and has mentored 30 scholars
for Ph.D. He was the President of AMI and MSI. His research efforts have been
focused on understanding the diversity and applications of yeasts, thermophilic
fungi and bacteria and their enzymes, metagenomics, carbon sequestration employ-
ing extremophilic bacterial carbonic anhydrases and bioethanol production from
lignocellulosic substrates using enzyme cocktails.

Dr. Sunil Kumar Deshmukh received his Ph.D. in Mycology from Dr. H.S. Gour
University, Sagar (M.P.) in 1983. The veteran industrial mycologist spent a substan-
tial part of his career at Hoechst Marion Roussel Limited [now Sanofi India Limited],
Mumbai and Piramal Enterprises Limited, Mumbai in drug discovery. He has to his
credit, 8 patents, 120 publications and 9 books on various aspects of Fungi and natu-
ral products of microbial origin. He is the past president of the Mycological Society
of India. He is a fellow of Mycological Society of India (MSI), the Association of
Biotechnology and Pharmacy and the Society for Applied Biotechnology. He is cur-
rently Fellow at Nano-Biotechnology Centre, TERI, New Delhi, and Adjunct
Associate Professor in Deakin University, Australia, working towards the develop-
ment of natural food colors, antioxidants and biostimulants through nanotechnology
intervention.

xxi
xxii About the Editors and Contributors

Dr. Mukund V. Deshpande obtained his PhD in 1982 in Biochemistry and D.Sc. in
Microbiology of the University of Pune in 1994. His extensive work in the area of
fungal biology, especially, fungal differentiation earned him D.Sc. He has worked
extensively on the use of fungi and fungal products in Biotechnology. Dr. Deshpande
successfully completed more than 35 research projects funded by national and inter-
national funding agencies, such as Indo-Swiss Collaboration in Biotechnology
(ISCB) programme of Department of Biotechnology (DBT), New Delhi and Swiss
Development Cooperation (SDC), Berne, Switzerland on development of mycoin-
secticide, Indo-Belarus programme of DBT on biopesticides, Indo-Mexico pro-
gramme of Department of Science and Technology, New Delhi and CONACYT on
fungal dimorphism. Dr. Deshpande is an elected fellow of the Maharashtra Academy
of Sciences (FMASc, 1994) and the Society for Biocontrol Advancement (FSBA,
2010). He is also a recipient of the Department of Biotechnology Overseas (Short-
term) Associateship (1995), and Commonwealth Science Council Fellowship
(1998). He has to his credit more than 140 research papers, reviews and chapters, 8
patents, 6 books and a number of popular articles. He has his own start-up
Greenvention Biotech located in Urli-Kanchan, Pune for the translational activities
in Agricultural Biotechnology.

Contributors

Krishnendu Acharya Molecular and Applied Mycology and Plant Pathology

Laboratory, Department of Botany, University of Calcutta, Kolkata, West Bengal,
India
Nitin Adhapure Department of Biotechnology and Microbiology, Vivekanand
Arts, Sardar Dalip Singh Commerce and Science College, Aurangabad, Maharashtra,
India
Shivankar Agrawal TERI-Deakin Nano Biotechnology Centre, Biotechnology
and Management of Bioresources Division, The Energy and Resources Institute,
New Delhi, India
Centre for Chemistry and Biotechnology (CCB), School of Life and Environmental
Sciences, Deakin University, Waurn Ponds, VIC, Australia
Indian Council of Medical Research (ICMR), Delhi, India
Shruti Ahlawat Department of Microbiology, Maharshi Dayanand University,
Rohtak, Haryana, India
Colin J. Barrow Centre for Chemistry and Biotechnology (CCB), School of Life
and Environmental Sciences, Deakin University, Waurn Ponds, VIC, Australia
Jadson Diogo Pereira Bezerra Departamento de Micologia Prof. Chaves Batista,
Programa de Pós-Graduação em Biologia de Fungos (PPG-BF), CB, Universidade
Federal de Pernambuco, Recife, PE, Brazil
About the Editors and Contributors xxiii

V. K. Bhalerao AICRP on Fruits, Mahatma Phule Krishi Vidyapeeth Rahuri,

Ahmednagar, Maharashtra, India
José L. Cabrera-Ponce Departamento de Ingeniería Genética, Unidad Irapuato,
Centro de Investigación y de Estudios Avanzados del IPN, Irapuato, Guanajuato,
México
Arunaloke Chakrabarti Department of Medical Microbiology, PGIMER,
Chandigarh, India
Bhushan P. Chaudhari Biochemical Sciences Division, CSIR-National Chemical
Laboratory, Pune, Maharashtra, India
Upashna Chettri Microbiology Laboratory, Department of Biotechnology and
Bioinformatics, North-Eastern Hill University, Shillong, Meghalaya, India
Hemraj Chhipa College of Horticulture and Forestry, Agriculture University
Kota, Jhalawar, Rajasthan, India
Priyanka Choudhari Nanobioscience, Agharkar Research Institute, Pune,
Maharashtra, India
Ritumbhara Choukade Department of Microbiology, Dr. Harisingh Gour
Vishwavidyalaya, Sagar, Madhya Pradesh, India
Bruna de Almeida Martins Department of Chemistry, Exact Sciences Institute,
Universidade Federal de Minas Gerais, Belo Horizonte-MG, Brazil
Geane Pereira de Oliveira Department of Chemistry, Exact Sciences Institute,
Universidade Federal de Minas Gerais, Belo Horizonte-MG, Brazil
Cristina Maria de Souza-Motta Departamento de Micologia Prof. Chaves
Batista, Programa de Pós-Graduação em Biologia de Fungos (PPG-BF), CB,
Universidade Federal de Pernambuco, Recife, PE, Brazil
C. D. Deokar Department of Plant Pathology, Mahatma Phule Krishi Vidyapeeth,
Ahmednagar, India
Sunil K. Deshmukh Biotech & Management of Bioresources Div, The Energy and
Resources Institute, New Delhi, Delhi, India
Mukund V. Deshpande Division of Biological Sciences, CSIR-National Chemical
Laboratory, Pune, Maharashtra, India
Thokchom Sarda Devi Department of Botany, University of Delhi, New Delhi,
India
A. P. Gaikwad AICRP on Mushroom, College of Agriculture, Pune, India
Vandana Ghormade Nanobioscience, Agharkar Research Institute, Pune,
Maharashtra, India
Samta Gupta Department of Botany, University of Delhi, New Delhi, Delhi, India
xxiv About the Editors and Contributors

Akshaya Gupte Department of Microbiology, N V Patel College of Pure &

Applied Sciences, Anand, Gujarat, India
Shilpa Gupte Department of Microbiology, Ashok & Rita Patel Institute of
Integrated Study & Research in Biotechnology and Allied Sciences, Anand, Gujarat,
India
Uttam Kumar Jana Department of Microbiology, Dr. Harisingh Gour
Vishwavidyalaya, Sagar, Madhya Pradesh, India
Rajesh Jeewon Department of Health Sciences, Faculty of Science, University of
Mauritius, Reduit, Mauritius
S. R. Joshi Microbiology Laboratory, Department of Biotechnology and
Bioinformatics, North-Eastern Hill University, Shillong, Meghalaya, India
D. N. Kamra Animal Nutrition Division, ICAR-Indian Veterinary Research
Institute, Izatnagar, Uttar Pradesh, India
Naveen Kango Department of Microbiology, Dr. Harisingh Gour Vishwavidyalaya,
Sagar, Madhya Pradesh, India
Rupam Kapoor Department of Botany, University of Delhi, New Delhi, India
Durgadas P. Kasbekar Centre for DNA Fingerprinting and Diagnostics,
Hyderabad, Telangana, India
Tanveer Kaur Department of Biotechnology, Thapar Institute of Engineering &
Technology, Patiala, Punjab, India
Claudia León-Ramírez Departamento de Ingeniería Genética, Unidad Irapuato,
Centro de Investigación y de Estudios Avanzados del IPN, Irapuato, Guanajuato,
México
Matheus Thomaz Nogueira Silva Lima Department of Food Science, Faculty of
Pharmacy, Universidade Federal de Minas Gerais, Belo Horizonte-MG, Brazil
David Lloyd School of Biosciences, Cardiff University, Cardiff, Wales, UK
C. Manoharachary Mycology and Molecular Plant Pathology Laboratory,
Department of Botany, Osmania University, Hyderabad, Telangana, India
Aliesha Moudgil Biochemical Sciences Division, CSIR-National Chemical
Laboratory, Pune, Maharashtra, India
D. Nagaraju Department of Botany, Government Degree College, Warangal,
Telangana, India
Sudeshna Nandi Molecular and Applied Mycology and Plant Pathology
Laboratory, Department of Botany, University of Calcutta, Kolkata, West Bengal,
India
Redeemson Panmei Biochemical Sciences Division, CSIR-National Chemical
Laboratory, Pune, Maharashtra, India
About the Editors and Contributors xxv

Ejaj K. Pathan Biochemical Sciences Division, CSIR-National Chemical

Laboratory, Pune, Maharashtra, India
A. V. Patil Biochemical Sciences Division, CSIR-National Chemical Laboratory,
Pune, Maharashtra, India
Fernando Pérez-Rodríguez Departamento de Ingeniería Genética, Unidad
Irapuato, Centro de Investigación y de Estudios Avanzados del IPN, Irapuato,
Guanajuato, México
K. S. Raghuwanshi Department of Plant Pathology, Mahatma Phule Krishi
Vidyapeeth, Ahmednagar, India
Shraddha Rahi Nanobioscience, Agharkar Research Institute, Pune, Maharashtra,
India
M. Sudhakara Reddy Department of Biotechnology, Thapar Institute of
Engineering & Technology, Patiala, Punjab, India
Darshan M. Rudakiya Department of Microbiology, N V Patel College of Pure &
Applied Sciences, Anand, Gujarat, India
José Ruiz-Herrera Departamento de Ingeniería Genética, Unidad Irapuato,
Centro de Investigación y de Estudios Avanzados del IPN, Irapuato, Guanajuato,
México
Mayela Salazar-Chávez Departamento de Ingeniería Genética, Unidad Irapuato,
Centro de Investigación y de Estudios Avanzados del IPN, Irapuato, Guanajuato,
México
Alejandro Sánchez-Arreguín Departamento de Ingeniería Genética, Unidad
Irapuato, Centro de Investigación y de Estudios Avanzados del IPN, Irapuato,
Guanajuato, México
V. Venkateswara Sarma Department of Biotechnology, Pondicherry University,
Kalapet, Pondicherry, India
Akshay Shankar Department of Microbiology, Maharshi Dayanand University,
Rohtak, Haryana, India
Krishna Kant Sharma Department of Microbiology, Maharshi Dayanand
University, Rohtak, Haryana, India
Rohit Sharma National Centre for Microbial Resource (NCMR), National Centre
for Cell Science (NCCS), Pune, Maharashtra, India
Rimpa Sikder Molecular and Applied Mycology and Plant Pathology Laboratory,
Department of Botany, University of Calcutta, Kolkata, West Bengal, India
Leticia Francisca da Silva Departamento de Micologia Prof. Chaves Batista,
Programa de Pós-Graduação em Biologia de Fungos (PPG-BF), CB, Universidade
Federal de Pernambuco, Recife, PE, Brazil
xxvi About the Editors and Contributors

B. Singh ICAR-Indian Veterinary Research Institute, Palampur, Himachal Pradesh,

India
Shiv Mohan Singh Banaras Hindu University (BHU), Varanasi, Uttar Pradesh,
India
Shreya Singh Department of Medical Microbiology, PGIMER, Chandigarh, India
Jacqueline Aparecida Takahashi Department of Chemistry, Exact Sciences
Institute, Universidade Federal de Minas Gerais, Belo Horizonte-MG, Brazil
Archana Tripathi Department of Microbiology, N V Patel College of Pure &
Applied Sciences, Anand, Gujarat, India
John Vélez-Haro Departamento de Ingeniería Genética, Unidad Irapuato, Centro
de Investigación y de Estudios Avanzados del IPN, Irapuato, Guanajuato, México
Tapani Yli-Mattila Molecular Plant Biology, Department of Biochemistry,
University of Turku, Turku, Finland
Emre Yörük Department of Molecular Biology and Genetics, Faculty of Arts and
Sciences, Istanbul Yeni Yuzyil University, Istanbul, Turkey
 Part I
Basic Aspects of Fungi
The Mystical World of Mushrooms
1
V. K. Bhalerao, A. P. Gaikwad, C. D. Deokar,
and K. S. Raghuwanshi

Abstract
The mushrooms have existed approximately 130 million years ago, i.e., long
before human beings evolved on this planet as per the fossil records. Earlier in
Sanskrit mushroom is known as “Ksuonpa.” In Hindi, mushroom is known as
“Khumbi.” Since long mushrooms are worshiped and also considered as divine.
In nature, mushrooms have not only been a source of food for man and other
animals but also have contributed an important role in the cycling of carbon and
other elements through the breakdown of lignocellulolytic plant residues and
animal dung which serves as the substrates for these saprophytic fungi. The his-
torical records of intentionally cultivated mushrooms estimated that the first
mushroom cultivation was started in 600 AD. In the last 20 years, much progress
has been made in the field of mechanization of mushroom cultivation, i.e.,
manure turners, spawning, filling and casing of trays, mechanical harvesting, and
polythene bag method of cultivation. In India, cultivation of edible mushrooms is
of very recent origin, though methods of cultivation of them were known for
many years. The research on different aspects, viz., production, productivity,
spawn production, strain improvement, post-harvest technology disease, and
pest management, was attempted by several scientists. All mushrooms belong to
the group of fungi, a group very distinct from plants, animals, and bacteria. Most
fungi have plant-like cells but miss the most important features of plants. The
known number species of fungi was about 69,000 till 1990, while it was conser-
vatively estimated that 1.5 million species of fungi actually existed in nature.

V. K. Bhalerao (*)
AICRP on Fruits, Mahatma Phule Krishi Vidyapeeth, Rahuri,
Ahmednagar, Maharashtra, India
A. P. Gaikwad
AICRP on Mushroom, College of Agriculture, Pune, India
C. D. Deokar · K. S. Raghuwanshi
Department of Plant Pathology, Mahatma Phule Krishi Vidyapeeth, Ahmednagar, India

© Springer Nature Singapore Pte Ltd. 2019 3

T. Satyanarayana et al. (eds.), Advancing Frontiers in Mycology & Mycotechnology,
https://doi.org/10.1007/978-981-13-9349-5_1
4 V. K. Bhalerao et al.

Mushrooms are cultivated throughout the world. About 200 species of more
than 2000 edible fungi are widely adopted for human consumption. Out of these,
only 10–12 species are commercially cultivated since the past few decades in
India due to technical advancement. Mushrooms have the capacity to produce
highest proteins per unit area and time by utilizing vertical space which is hun-
dred times more than the traditional agriculture and animal husbandry. This hi-­
tech horticulture venture can reduce the pressure on cultivated land to meet the
food shortages all over the world. More than 100 countries are engaged in mush-
room farming today which is increasing at an annual rate of 6–7% per annum.
The very high levels of mechanization and automation were achieved in mush-
room farming in developed countries of Europe and America. As per FAO Stat,
the present world production of mushrooms is around 3.5 million tonnes, which
is more than 25 million tonnes (estimated) as per claims of Chinese Association
of Edible Fungi.The mushrooms can be cultivated under varied climatic condi-
tions. Some of the economically important mushrooms cultivated all over the
world under temperate, subtropical, and tropical conditions are Agaricus bispo-
rus, Lentinula edodes, Flammulina velutipes, Agaricus bitorquis, Pleurotus spp.,
Auricularia spp., Agrocybe aegerita, Volvariella spp., Calocybe indica,
Ganoderma lucidum, etc.
Mushrooms are rich source of proteins, carbohydrates, valuable salts, and
vitamins in diet of human being. Mushroom fungus has the ability to secrete a
wide variety of hydrolyzing and oxidizing enzymes which have potential for
biotechnological applications. More than 100 medicinal edible mushrooms have
been identified. However, important medicinal mushrooms are Lentinus edodes
(shiitake mushroom), Ganoderma lucidum (reishi mushroom), Grifola frondosa
(maitake mushroom), Pleurotus ostreatus (oyster mushroom), Agaricus bisporus
(button mushroom), Coriolus versicolor (PSK), Boletus edulis, Tremella fucifor-
mis, Auricularia polytricha, Hericium erinaceus, and Cordyceps sinensis.

Keywords
Mushrooms · Oyster mushroom · Agaricus sp. · Medicinal values · Cultivation ·
Commercial production · Nutritional resource

1.1 Introduction

The mushrooms have existed approximately 130 million years ago, i.e., long before
human beings evolved on this planet as per the fossil records. The origin of the term
“Mushroom” has different views. In Latin, “Fungo” means to flourish. The Greek
term mushroom was derived from the word “Sphonggos” or “Sphoggos” which
means “Sponge” and referred to the sponge-like structure of some species. In
French, the term mushroom was derived from the word “Mousseron” (muceron),
“Mousse,” or “Moss.”
1 The Mystical World of Mushrooms 5

About three and half millennia ago, the Greek hero Perseus founded Mycenae
city after a legendary mushroom. “Mycenae” is derived from Greek word “mycol-
ogy” (i.e., mykes – mushroom + logos – discourse). Earlier in Sanskrit, mushroom
is known as “Ksuonpa.” In Hindi, mushroom is known as “Khumbi.” The other
words for fleshy capped fungi are Chatra, Kukurmutta, Kavaka, Bhumi kavak, and
Bhustrana. Aryans during migration into the Indian subcontinent around in
1500 B.C. carried with them an intoxicating drink “Soma” which they used in their
religious rites. According to Wasson) (1969), the “Soma” in Rig Veda refers to
Amanita muscaria.
Since long mushrooms are worshiped and also considered as divine. There are
lots of superstitions about the mushrooms. According to the ancient Indian, Greek,
and Roman myths, mushrooms sprang from a stroke of lightning. The Indians in
Mexico believed that mushrooms are sacred because they are borne of the sexual
intercourse between a bolt of lightning and the earth. The mushroom-shaped stone
carvings have also been found in Central America and the highlands of Guatemala.
These objects resemble very closely to the fruit body of Amanita muscaria.
In nature, mushrooms have not only been a resource of food for man and other
animals but also have an important role in the cycling of carbon and other elements
through the breakdown of lignocellulolytic plant residues and animal dung which
serves as the substrates for these saprophytic fungi. In this way, mushroom species
as agents of decay helps in keeping the environment clean.
The mushrooms were long appreciated because of their flavor, texture, and
medicinal or tonic attributes. The recognition, that mushrooms are nutritionally a
very good food is much more recent.
The popularity of mushroom is desired from the following highly preferred char-
acteristics of food:

1. Significant taste and flavor.

2. Nutritious, not only because they contain high level of protein containing signifi-
cant amounts of lysine and methionine (which are generally low in plants),
fibers, minerals, and vitamins but also for what they do not have (high calories,
sodium, fat, and cholesterol).
3. Can be easily stored by processing, drying, pickling, and canning which allow
maximum storage. Mushrooms appeal to different people in different ways.
Mushrooms are objects of beauty for artists. Mushrooms are possible source of
new drugs for medical people. Architects have constructed minerals, temples,
and cupola column in its shape. Jewelers have made expensive pieces of mush-
room design. Designers have reproduced the mushroom design on fabrics.
6 V. K. Bhalerao et al.

1.2 History of Mushroom Research and Cultivation

1.2.1 International Developments

The historical records of intentionally cultivated mushrooms are shown in Table 1.1.
The first artificial cultivation of Auricularia auricula mushroom was estimated
around 600 AD in China on wood logs and the same time other wood rotting mush-
rooms, such as Flammulina velutipes (800 AD) and Lentinula edodes (1000 AD).
In Europe, the first cultivated fungi, the mushroom, was introduced in the seven-
teenth century. Agaricus bisporus was not cultivated until 1600 AD which is the
leading mushrooms. The Pleurotus ostreatus was first cultivated in the USA, and
several other species of Pleurotus were initially cultivated in India.
The first record of the cultivation of mushrooms was during 1638–1715, i.e., the
reign of Louis XIV. Treschow shortly before 1700 suggested a method of growing
A. bisporus in hotbeds under heat. Tourneforte, a French man in 1707, published
description and know-how of growing mushrooms which is similar to the present
methods.
Van Griensven (1988) reviewed the historical events in mushroom cultivation,
whereas a detailed history was elaborated by Shu-ting and Miles Philip. The
­cultivation of mushrooms aboveground is originated in Sweden. Bahl (2002) reviewed
that Lundberg described mushroom growing in greenhouses in year 1754 whereas,
the French started growing mushrooms underground in the quarries around Paris on
horse manure in 1800 and Callow reported shelf bed method in a peculiar type of
house for mushroom cultivation in 1831.
By the end of the nineteenth century, French mycologists Matrochot and
Constantin (1894) discovered the cause of mushroom disease La mole (Mycogone
perniciosa Magnus) and started sulfur fumigation. They were able to germinate
spores for obtaining sterile spawn. Miss Fergussen (1902) of Corne described
detailed method for germinating spores.
Duggar (1905), an American, perfected a method of making pure culture spawn
from mushroom tissue. Lambert (1929) introduced and marketed pure spore culture
bottled spawn in the USA. The cultivation of mushroom aboveground is originated
in Sweden. Sinden (1938) patented his grain spawn process. Thomas, Austin, and
Jary worked on pests of mushrooms. Ware, Glasscock, and Bewley in the UK and
Lambert and Beach in the USA worked on diseases and competitive molds of
mushrooms.
In 1945, MGA (Mushroom Growers Association of England and Ireland), MRA
(Mushroom Research Association Ltd.), and MGP (Midlands Group of Publication)
were formed and established. In the same year, an organization called Mushroom
Growers Association of Great Britain and North Ireland was established. This asso-
ciation publishes a mushroom journal, which is supplied to the 50 nations.
In 1946, the first mushroom research station was founded in England with Dr.
R.L. Edwards as its first director. The mushroom research at this station is mainly
devoted to Agaricus bisporus and involves all aspects of this single species. In 1954,
1 The Mystical World of Mushrooms 7

Table 1.1 Historical evidence of commonly cultivated mushrooms (Miles and Chang 1997)
Species Record first cultivated (est.)
Agaricus bisporus 1600 (Atkins 1979)
Agaricus bitorquis 1961
Agrocybe cylindracea 1950
Amanita caesarea 1984
Armillaria mellea 1983
Auricularia auricula 600
Coprinus comatus 1984
Dictyophora duplicata 1982
Flammulina velutipes 800
Ganoderma spp. 1621
Gloestereum incarnatum 1989
Grifola frondosus 1983
Hericium coralloides 1984
Hericium erinaceus 1960
Hohenbuehelia serotina 1982
Hypsizygus marmoreus 1973
Lentinus edodes 1000
Lentinus tigrinus 1988
Lyophyllum ulmarium 1987 (Wang and Zhang 1987)
Morchella spp. 1986
Oudemansiella radicata 1982
Pholiota nameko 1958
Pleurotus citrinopileatus 1981
Pleurotus cystidiosus 1969
Pleurotus ferulae 1958
Pleurotus flabellatus 1962 (Bano and Srinvatava 1962)
Pleurotus Florida 1958
Pleurotus ostreatus 1900
Pleurotus sajor-caju 1974 (Jandaik 1974)
Poria cocos 1232
Sparassis crispa 1985
Tremella fuciformis 1800
Tremella mesenterica 1985
Tricholoma gambosum 1991
Tricholoma lobayense 1990
Tricholoma mongolicum 1991
Volvariella volvacea 1700

the mushroom research station was transferred along with three staff members and
government grants to the new glasshouse crop research station at Littlehampton,
Sussex.
In 1959 at Horst in Holland, a fine center on mushroom was opened. In the later
years of the eighteenth century, French growers discovered the usefulness of gyp-
sum powder to prevent greasiness in compost. Bewley and Lambert worked on
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