1

The Mushrooms: an ethnomycological, molecular and nutritional analysis from the perspective of ethnic 1

communities 2

Ethnomycobiology and mushrooms 3

Madhusmita Borthakur, Polashree Khaund , S.R. Joshi* 4

Microbiology Laboratory, Department of Biotechnology & Bioinformatics, 5

North-Eastern Hill University, Shillong 793022, Meghalaya, India 6

Corresponding address: srjoshi2006@yahoo.co.in 7

8

Abstract: 9

Micro-organisms are considered as priceless sources for various important biomolecules having enormous 10

commercial applications in medicine, food industry and bio-anthropology. Biodiversity hotspot regions are 11

considered to be genetic paragon of plant, animal and microbial resources. This mega-biodiversity zone encompasses 12

areas such as the sacred groves which are considered as the abode of deities by ethnic people. These groves are rich 13

in flora and fauna besides the microbial and macrofungal components. Mushrooms are considered as highly 14

nutritious which carry added medicinal value. However, their bioprospection and commercial exploitation is very 15

scarce as compared to other life forms. This may be due to the limited information available on their nutritional and 16

genetic constituents. Proteins and polysaccharide comprise the main component of their dry matter with glycogen, 17

chitin, trehalose, mannitol as the main carbohydrate while n-3 fatty acids are negligible. High proportion of fibre 18

content, less expensive and easy availability has attracted both the interest of researchers and consumers to the 19

science of mushrooms. The ethnomycological, nutritional and molecular aspects of mushrooms has been reviewed in 20

this paper. 21

Key words: Ethnic community, Bioanthropology, Ethnomycology, Mushrooms 22

2

INTRODUCTION 23

Microorganisms are considered to be the richest repository in natural environment [1]. Microbial diversities are 24

found pervasive in every corner of earth. There are as many as 25 biodiversity rich regions of the world [2] and Asia 25

is considered as one of the mega-diversity hotspots which hold 60-70% of the world’s biodiversity [3] including 26

microorganisms. Among various microorganisms, fungi play a vital role in maintaining the sustainability of our 27

mother Earth. The diversity of the fungi was debated for more than 200 years but in last few decades, the DNA 28

sequence has provided a myriad of new characteristics for the analysis of 100 phylum from the kingdom fungi [4, 5]. 29

Fungi are found from the equatorial regions to the poles, from marine ecosystems to fresh water environment [6] 30

surpassed by only bacterial community in their inability to withstand water activity, extreme temperature conditions 31

and carbon sources [5]. Fungal community are considered to be existing in two diverse characteristics - sociological 32

which includes their geographical distribution, community structure, their diversity in various community, habitat 33

specificity and their second aspect of characteristics is their behavioral characteristics which relates to their function 34

in an ecosystem [7]. There are estimates of more than 1.5 million fungi [8] in the world out of which only 98,998 35

species have been described [9]. Mycotechnology has advanced in last five decades for their potential in various 36

organic acids, extracellular enzymes, antibiotics, antioxidant activity, in the field of bioremediation, biome-seed, etc 37

[10, 11]. 38

Malnutrition in terms of deficiency in protein is one of the foremost factors responsible for the morbidity and 39

mortality in under-developed as well as other developing countries in the world [12]. With population explosion, the 40

protein deficiency is expected to become more acute in the coming years and recommending animal protein is 41

beyond the limit for some developing countries. Non-traditional food resources can partly meet the dietary deficit. 42

With the advent of modern molecular biology tools, fungus has been renamed as a ‘cell factory’ for various 43

heterologous proteins [13, 10] which also include human proteins. Mushrooms produce many bioactive proteins and 44

peptides, primarily including lectins, laccasses, fungal immunomodulatory proteins (FIP), antimicrobial/antifungal 45

proteins, ribosome inactivating proteins (RIP), ribonucleases and many of these bioactive compounds have entered 46

into phase I, II and III clinical trials and are successful in treating cancers in world [14]. 47

This review emphasizes mostly on the filamentous fungi, their nutritional perspectives and cytogenetics, since bulk 48

of research has been done in this group and is considered as a alternative food source for sustainable development of 49

a country’s economy. 50

3

MUSHROOMS - THE MACROSCOPIC FUNGI 51

Mushrooms are considered as well-balanced food stuffs and good source of nutrition in the era where whole world 52

suffers from scarcity of food. Geologically, mushrooms existed on earth even before the appearance of human, as 53

evidenced from the fossil records of the lower cretaceous period [15]. The term mushroom was first defined ‘as 54

macrofungi with a defined fruiting body that can be either epigeous or hypogeous’ which can be easily visible to 55

naked eye [16]. Mushrooms belong to basidiomycetes family with stalk and umbrella shape fruiting body bearing 56

either gills or pores although from taxonomical view, some species in ascomycetes family also belong to mushroom. 57

It first appears as a white tiny balls with a cap, also known as pileus and a short stem also known as stipe. The 58

membrane or veil, also known as velum covers the envelope which gets torn off and cap become visible with gills or 59

lamellae radiating down beneath the cap [12]. Several species of mushrooms are known because of its characteristics 60

color, texture and pleasant odour [16](Fig I). It has been estimated that there are more than 140000 mushrooms with 61

only 10% being known [16, 17]. Being from basidiomycetes family, they have compatible haploids commonly the 62

mycelia being composed of filamentous hyphae. Through plasmogamy, haploid compatible mycelia fuse with each 63

other and nuclei migrate into each other’s mycelia and get paired with the resident nuclei. Since karyogamy is 64

delayed, compatible nuclei remain in dikaryotic stage for some time. Often it is seen that mycelium in dikaryotic 65

stage is more vigorous than the monokaryotic stage (the haploid mycelia) and can take up the substrate more easily 66

in which they are growing. Basidiomycota is characterized with the formation of clamp connection. The long lasting 67

dikaryotic period produce basidia, in which a congruent nuclei fuse (karyogamy) to form a diploid cell. Meiosis is 68

followed; by the production of 4 haploid nuclei that transfer into the external part forming basidiospores. However, 69

variation are always observed ; Craterellus spp. often have 6-spored basidia while Sistotrema spp. have 2,4, 6 or 8 70

basidiospores and the common button mushroom, Agaricus bisporous has 1, 2, 3 or 4 spored basidia under some 71

circumstances. Basidiospores are discharged to start up a new haploid mycelia also known as monokaryons. The 72

karyotype of mushrooms has not yet been very clear because of contingency of endomitosis and small size of 73

mushroom chromosomes [2]. The cultivation of mushroom is termed as “non green revolution”[18]. The genome 74

size and chromosome numbers vary in mushrooms when analyzed through various analytic techniques which include 75

Restriction Fragment Length Polymorphism (RFLP) and Randomly Amplified Polymorphic DNA (RAPD). 76

Mushrooms are being consumed over 10 kg yearly in few European countries with a statistical mean of 5.6 kg of 77

cultivated fresh mushrooms per household/year [19]. World- wide mushroom cultivation is dominated by Agaricus 78

4

bisporus [20] which is not only edible but also has been valued as a source of traditional medication by humankind 79

[17]. Considering their nutraceutical and ethnocultural relevance, there is a need of conservation, preservation and 80

characterization of the wild mushrooms from the pristine forests owing to the vigorous destruction of the wild habitat 81

by various anthropogenic activities. The economic prospects of such wild mushrooms which are yet to be identified 82

for their activities including medicinal and antioxidant can assist in their conservation and economic subsistence. 83

SOURCES AND ETHNOMYCOLOGY OF MUSHROOMS 84

In the recent era, the hub for research on mushrooms has turned to undisturbed forests and pristine forests like sacred 85

groves. Sacred groves with a concept of “presiding deity” have been protected by the tribal community. The section 86

contains geographical features along with vegetations and other forms of flora and fauna, demarcated by the 87

indigenous people to keep them in a protected and undisturbed state which have formed an intricate relationship 88

between human and nature Earth [21]. These virgin forests are rich in biodiversity with a huge amount of endemic 89

and endangered species. These forests in recent days have attracted attention for numerous issues including reservoir 90

of threatened species, repositories for various medicinal plants, macrofungi (mushrooms), giant trees, gene bank for 91

economic species, recreation center for urbanites and a huge untapped paradox for researchers and environmentalist 92

[22, 23]. Due to numerous environmental transformations such as climatic variations, human interference such as 93

shifting cultivation and other anthropogenic activities, these forests have dwindled its size leading to the loss of 94

pristine habitats. 95

Sacred groves are protected and largely untrapped with a limited knowledge of the indigenous flora, fauna and 96

microbes. Traditional knowledge on conservation and preservation of its wild habitat are seen among the native tribal 97

communities. The knowledge emerged over the generation for conservation of these groves is as much a resource as 98

a biodiversity. Wild mushrooms (macrofungi) are highly dominant in such groves and only few reports on wild 99

mushrooms are available [24] while their medicinal prospects, antimicrobial, anti-oxidative, trace element prospects 100

are yet to be documented. Tribal Communities are known to collect the wild mushrooms based on their 101

ethnomycological knowledge. Wild poisonous mushrooms are distinguished from their edible counterparts in having 102

a pointed cap, bright colored fruiting body, unpleasant odour, colour of the fruiting body often turns black upon 103

breakage, absence of worms and insects [25]. 104

Ethnomycology is considered as the study of fungi for the development of cultures, religion and mythology. In this 105

respect mushroom received attention for their potential as a hallucinogen, their edibility for the presence of abundant 106

5

bioactive and nutritional compounds and for their toxicity. China have a long tradition of using mushroom since last 107

5000 years due to its therapeutic and consumable purpose but its therapeutic value has also been recognized by many 108

early civilizations including Greeks, Aryan, Romans, Egyptians, Romans and Mexicans. Mushrooms were 109

considered as a “food of God” by Romanians, “strength food” by Greek warriors, “elixir of life” by Chinese, as 110

“Somrus” a decoction of mushroom by Indians [26]. The brown rot fungi, Formitopsis officinalis are recognised as a 111

“Bread of Ghosts” by the indigenous people of Pacific Northwest and use it to mark the grave of shamen. Shamen of 112

Pacific Northwest use the sporophores of Formitopsis officinalis for various rituals and to treat illnesses caused by 113

supernatural powers. They believe the shape of the mushroom sporophore gave mushroom its spirit catching abilities. 114

Apart from Formitopsis, an obligate plant pathogen Exobasidium vaccinii infect stem, leaves and flowers of 115

Ericaceous plant and subsequently form vegetative deformities thus forming a basidia covered gills which are 116

consumed by the indigenous coastal groups of Pacific Northwest and considered them to be berries. The berries are 117

referred to as “Ears of Ghost” and believe in a myth that they are associated with corpse-stealing, child- kidnapping 118

creature [26]. Burk (1983) reported the spiritual and religious use of puffballs mushrooms by the people of North 119

America and they believed that they have the ability to ward off ghosts [27]. Puffballs also received attention in 120

British Columbian Indian spiritual world as they are regarded as “ghost’s make up” or “ground ghost” or “corpse” as 121

reviewed by Nicholson (2009)[26]. In Asia, the mycophilic societies are associated among the indigenous people of 122

North Eastern India, Western ghats and North Western India, China. The local inhabitants collect mushrooms from 123

their neighboring localities, meadows and forests for consumption and sell them to earn revenue for their family 124

during monsoon season when other forest nonwood products are unavailable in the market. Among various species 125

of mushrooms which are commonly consumed by the people are Termitophilous which include T. microcarpus, T. 126

aurantiacus, T. eurhizus, T. clypeatus, T. Tyleranus [28] Few indigenous tribes of North-Eastern corridor of the India 127

such as Khasis of Meghalaya use a traditional technique ‘narsuh’ for cooking mushrooms where they heat the tip of a 128

small iron rod and place it in the middle of the bowl containing cooked mushroom. They believe that heat release 129

from the tip is responsible to destroy or absorb the poisonous harmful substance from the mushrooms [25].Some use 130

other traditional methods. The fruiting bodies are washed and boiled with a few cloves of garlic. If the cloves of 131

garlic turn black in color, they believe that the mushrooms are poisonous in nature. However, if there is no color 132

change then they are safe to eat. 133

CYTOGENETICS OF MUSHROOM 134

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Basidiomycetes contain two types of mycelia, monokaryon (asexual) and dikaryon (sexual) which allow easy 135

exchange of nuclear partners in monokaryotic - dikaryotic and dikaryotic- dikaryotic mycelial fusion [29]. For most 136

cultivable mushroom in the world, Agaricus bisporus researchers have not been able to provide comprehensive 137

information on various expect such as the genetic, physiology, cytology and breeding [30]. Since fungal 138

chromosomes are minute, pulse field gel electrophoresis enables the karyotyping of the various mushroom species 139

[31] showing molecular size and numbers of chromosomes. A study by Sagawa and Nagata (1992) reported the size 140

of six chromosome ranging from 2.1 to 5.7Mbp (million base pair) per chromosome and genomic size to be 20.8Mbp 141

of Pleurotus ostreatus[32]. The size of mushroom chromosomes DNAs range from 2Mb to 30Mbp. Fruiting bodies 142

normally develop on dikaryon. Karyotypic analysis of the mushroom fruiting bodies can be revealed by spreading 143

the chromosome at meiotic prophase stage. Karyogamic in basidiomycetes occur directly before meiosis where intra-144

tetrad fertilization occur, that is the nuclei of one tetrad unite in pairs but do not fuse [33] which normally takes place 145

in the gills of the fruiting bodies which is followed by meiosis (maturation of basidia, an autonomous endotrophic 146

process) and by production of meiotic basidiospores. A report by Larraya et al (1999) suggest that chromosomal 147

length polymorphism are most common in basidiomycetes and are analyzed by electrophoretic karyotyping, however 148

two approaches have been proposed for polymorphism to occur, by increasing the copy number of ribosomal DNA 149

segment or subtelomeric region of DNA and by mitotic and meiotic recombination process [34]. From various 150

studies it has been reported that both monokaryon and dikaryon can form multicellular resting bodies or sclerotia and 151

various mitotic spores, the tiny uninucleate aerial oidia and large thick walled chlamydospores within submerged 152

mycelium [35, 36, 37]. In a study by Summerbell et al (1989) it has been reported that one spore receives two non-153

sister nuclei, i.e. the progeny is formed of two different nuclei in first mitotic division [38]. However, in 154

basidiomycetes, mating is supported by population genetics evidence of recombination and also from intravarietal 155

hybrids from the environment [39]. Spore production can be examined by analyzing molecular markers of progeny 156

that are derived from a single basidium. A report suggest that in Cryptococcus neoformans basidium, there occurs a 157

single meiosis and other multiple rounds of mitosis to generate nuclei in the basidium [40]. Spores remain in 158

aneuploid state and do not germinate rather remain dormant. Chains are formed from heterogenous population of 159

spores of different genotypes due to nuclear movements which occurs in basidium [41]. Nuclei can remain fixed in 160

the basidia from which the spores bud or can move after every mitosis which can get influenced by environmental 161

factors including temperature, light, humidity and nutrients. Light conditions determine the formation of hyphal 162

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knots or oidiophores with oidia and also determine whether hyphal knots will mature into sclerotia or fruiting body. It 163

induces hymenium differentiation and finally the karyogamy. Among genes specifically expressed within fruiting 164

body for hyphal aggregation are β-galactoside binding lectins [29]. Variation in temperature also determines the 165

formation of fruiting body. Lower temperature favors the formation of fruiting body initial and lower temperature 166

favors the formation of oidia [36, 42, 43, 44, 45, and 46]. Fruiting body needs high humidity whereas sclerotia and 167

chlamydospores will appear at low humidity. In later half of the development after 48 h, light is required for the 168

induction of karyogamy during DNA synthesis prior to the formation of nuclear fusion [43, 47, 48, 49, 50, 51, 52, 169

53] while stipe elongation and maturation of cap takes place parallelly. For the formation of fruiting bodies, around 170

20 to 37 0C temperature is necessary but prior to the meiotic DNA replication, a two hours dark period at 37

0C is 171

utmost important. The ratio of carbon and nitrogen also determine the initiation of fruiting body formation and 172

sclerotium maturation, whereas fruiting body initiation is inhibited by glucose and glucose analogues [54, 55, 56, 57, 173

and 58]. 174

TAXONOMICAL STUDIES TO ANALYZE VARIOUS STRAINS 175

The study of mushrooms to distinguish among different strains is perhaps inadequate based on morphological 176

features. For the genetic analysis, it is essential to express both the alleles in heterozygous which is possible by 177

allozyme analysis (recognition of codominant phenotype) which involves movement and separation of enzymes in 178

gel. Electrophoretic mobility of the allozyme is directly correlated with the genotypic variations among different 179

strains within a single species [16]. Among the species identified by multilocus enzyme electrophoresis are wild 180

mushroom including Agaricus bisporus, Agaricus campestris, Volvariella volvacea, Lentinula edodes, Pleurotus spp. 181

and Morchella spp. 182

Arbitrarily primed polymerase chain reaction, a marker based upon DNA probe technique is also advantageous 183

process to distinguish between different strains of mushrooms over allozyme analysis. The amplicons are revealed by 184

electrophoresis and are characterized by direct sequencing. 185

Edible and poisonous mushrooms are characterized by mycologists based on certain palynological, sporological and 186

morphological trait, but very few reports are available on their molecular aspects. In the genus Psilocybe, P. 187

semilanceata and P. cubensis produce hallucinogenic effect while P. merdaria and P. montana donot produce such 188

effect [59]. Few toxic mushrooms belonging to the genus Lepiota are identical to that of the edible mushroom of 189

genus Macrolepiota. Similarly, Inocybe asteropora can be misidentified as Armillaris spp.[60]. Rapid molecular tools 190

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including molecular markers, PCR analysis would support the identification of poisonous nature of mushrooms. 191

Various toxins including amanitoxins, phallotoxins, monomethylhydrazine, orellanine, muscarine, ibotenic acid, 192

muscimol, coprine, psilocybin, psilocin are confined to various wild macrofungi [61]. An interesting poisonous 193

mushroom is Gomphus floccosus. Consumption of this mushroom in the northern United States and in Canada has 194

been reported to cause severe gastrointestinal disorders [62]. The consumption of this poisonous mushroom by the 195

ethnic population of Meghalaya without any apparent harm may be hypothesized to reduced toxicity of the specific 196

local strain available in the region, unique indigenous cooking practices and methodology or even to genetic 197

resistance to the mushroom toxin developed among the local population [25]. Poisonous and edible ones are grouped 198

and named as toadstool by English speakers and “jhasmuka” or “lu” among the Mayans [63]. Epis et al (2010) 199

developed species specific molecular marker to identify the poisonous nature of four different mushrooms including 200

Amanita phalloides, Lepiota cristata, Lepiota brunneoincarnata, Inocybe asterospora based on the ITS1-5S rDNA-201

ITS2 sequences which are highly conserved among the fungi [60]. Lee et al (2000) developed primers 202

complementary to ITS1 locus to differentiate between hallucinogen mushroom Panaeolus and Psilocybe based on 203

the length of the PCR product [64]. 204

Identification of novel molecular markers and rapid PCR protocol can be an approachable strategy to differentiate the 205

hallucinogen from edible counterpart. 206

In a study on Amanita mushrooms it was found that amatoxins, the lethal constituents of these poisonous mushrooms, 207

are bicyclic octapeptides [65]. Two genes in Amanita bisporigera, AMA1 and PHA1, directly encode α-amanitin, an 208

amatoxin, and the related bicyclic heptapeptide phallacidin, a phallotoxin, indicating that these compounds are 209

synthesized on ribosomes and not by nonribosomal peptide synthetases. AMA1 and PHA1 are present in other toxic 210

species of Amanita section Phalloidae but are absent from nontoxic species in other sections. The genomes of 211

Amanita bisporigera and Amanita phalloides contain multiple sequences related to AMA1 and PHA1. In another such 212

study by Qing et al (2012), to establish a standard DNA barcode for poisonous species of Amanita in China, three 213

candidate markers, the large subunit nuclear ribosomal RNA (nLSU), the internal transcribed spacer (ITS), and the 214

translation elongation factor 1 alpha (tef1α) were tested using the eukaryotic general primers for their feasibility as 215

barcodes to identify seven species of lethal fungi and two species of edible ones which can easily be confused with 216

the lethal ones known from China [66]. This study showed that the tef1α and nLSU can be proposed as 217

supplementary barcodes for the genus Amanita, while ITS can be used as a primary barcode marker considering that 218

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the ITS region may become a universal barcode marker for the fungal kingdom. 219

In the study on the wild mushrooms which are consumed by the ethnic tribal population of India, it was found that 220

the differentiation between the poisonous and edible varieties made strictly based on the morphology and molecular 221

characterization often do not bestow apparent results. Molecular characterization being confined to one conserved 222

locus, ITS1 and ITS4 only identify various macrofungi rather than discriminating among edible and closely 223

associated poisonous mushroom (Fig II, Fig III). An alternative DNA barcode based on mitochondrial COI gene 224

allows correct identification of only 20-70% of fungal species [67, 68]. In a study on wild edible mushrooms, 225

morphological features and ITS markers have been proposed as barcode for mushroom identification [69]. So, a hunt 226

for a novel primer for rapid differentiation among the poisonous and edible one are in progress to discriminate the 227

two varieties to a greater extent. 228

MUSHROOMS AS DIETARY SUPPLEMENTS 229

Mushrooms have been used as a food item since time immemorial [70]. Mushrooms are considered as the world’s 230

largest untapped resources of dietary supplements. Their consumption increased due to the presence of high amount 231

of protein content and trace mineral [71, 72]. In comparison with other eatables including fruit, vegetables and 232

legumes, mushroom or basidiomycetes can prove better as a rich source of certain dietary factors that can be 233

beneficial to human health and growth [73]. They are considered as a healthy food which contains essential fatty 234

acids, low in the amount of calories and high in the concentration of the protein, fats and minerals [5]. As wild 235

mushrooms are considered to have more amount of protein content than commercially available mushroom, several 236

reports reveal that they are used as a diet to combat various diseases [15]. For a developing country like India, they 237

can serve as a good substitute for protein malnutrition [20]. Mushrooms with fleshy fruiting bodies can be used as a 238

good source of biopharmaceutical product as they contain polysaccharides and triterpenoids [70]. Various studies 239

have shown that after cooking, dried mushrooms show highest level of nutritional value due to insufficient 240

rehydration [74]. In general mushrooms contain 90% water and 10% dry wight [75]. In a study by Orgundana and 241

Fagade (1981), mushrooms are reported to contain about 16.5% dry matter, the approximate composition of crude 242

protein is 14.6%, crude fibre is 7.4%, and composition of fat and oil is around 4.48% [76]. Though mushrooms are 243

commercially used as a source of nutritional supplement, they have an adverse effect in the health of human as they 244

are also considered as a source of human poisoning and are associated with various carcinogenesis while 245

experimenting in animal [73]. According to a study by Chang (1990), only 2,000 mushrooms are considered edible of 246

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which 20 are commercially cultivated and 4 to 5 are cultivated industrially [77]. Among various higher species of 247

basidiomycetes, Agaricaceae are widely consumed worldwide. Many polysaccharide and protein based 248

polysaccharide isolated from Agaricaceae have anti-tumour activity and specific or non specific immune response 249

activity [36, 37, 42, and 74]. The dietary fibre of mushroom is due to the polysaccharides and triterpenoids contained 250

in them. The cell wall of mushroom contains a mixture of fibrillar and matrix components which comprises chitin, 251

polymer of β 1- 4 linkages N-acetyl-glucosamine and polysaccharides of 1-3 linkage of β-D-glucans and mannans 252

respectively. Mushrooms bear mainly the water insoluble dietary fibers which include both the chitin and β-glucans. 253

Under unfavorable environmental conditions, mushrooms regenerate a special structure known to be a sclerotia 254

which are compact mycelium structure bearing chitin and β-glucans with β 1-3 backbone and 1- 6 linked side 255

branches and due to this they are considered to be a novel source of dietary component as human enzyme cannot 256

digest them [70]. 257

PROTEIN COMPOSITION 258

Mushrooms being rich in the protein component comprise mostly of the essential amino acid component lysin, 259

histidine, arginine, threonine and tryptophan that are commonly not present in cereals. The component of lysin has 260

been reported as a most abundant essential amino acid and methionine and tryptophan being the least abundant 261

amino acid. The content of free amino acid is low which consist of only 1% of the dry weight of mushroom, thus 262

limiting the nutritional level but they participate in the taste of mushroom [19]. The protein component of mushroom 263

depends on its growth substratum, the level of nitrogen and its location, the size of the pileus, varies among species 264

and its harvesting time [78]. Immature primordia contain more amount of protein than mature primordia as reported 265

in Agaricus bisporus and Pleurotus spp. [79]. The composition does not change during oven drying of mushroom in 266

40 degree centigrade but significant reduction in its composition can be visualized upon boiling of fresh mushroom 267

(19). Protein production efficiency for mushroom is about twice that of cabbage and asparagus, four times as of 268

orange, 12 times as of apples [77]. On the basis of its dry weight it contains 65% protein as compared to 25.2% in 269

milk, 20% for pork meat and 4% for beef , 39.1% in soybean, 7.3% in rice, 13.2% in wheat [77, 80]. An early study 270

by Rose (1937) had compared the presence of all essential amino acid for the growth of mammalian cells, which 271

revealed that the composition of amino acid in mushroom is adequate except in phenylalanine and methionine[81]. 272

The palatable taste of the mushroom is enhanced by the presence of aspartic and glutamic acid which are 273

monosodium glutamate-like (MSG-like) components [82]. It has been reported by Bauer-Petrovska (2001) that mean 274

11

level of protein fraction such as albumins, prolamines, globulins, glutenins, glutenin-like material and prolamine-like 275

material are likely to be 24.8, 5.7, 11.5, 11.5, 7.4 and 5.3% respectively [83]. Proteins and peptides produced by 276

mushrooms possess interesting biological activities such as lectin, antimicrobial protein, laccases, ribonucleases, 277

ribosome inactivating protein and fungal immunomodulatory protein [14]. 278

A wide range of bioactive compounds have been isolated from mushrooms which include protein- polysaccharide 279

complexes, proteins, polysaccharides, etc [84, 85]. Among those the most specific are lectin protein, which are 280

glycoproteins that can bind to cell surface carbohydrates with their ability for cell agglutination. They exhibit 281

antiproliferative activity towards tumor cell lines specifically to human leukemic T cells, breast cancer MCF7 cell, 282

hepatoma Hep G2 cells and can recognise human blood group A determinant carbohydrates [14]. The sacred 283

mushroom Reishi (Ganoderma spp.) contain a polysaccharide, β-glucan which stimulate or modulate the level of 284

immune system by activating T cells and macrophages as well as activating immunoglobin levels which can combat 285

and respond against foreign cells [86]. Crystal structure of lectin protein was first determined by Cioci et al (2006) 286

with a regular seven-bladed beta-propeller fold with N-terminal region being stuck into the central cavity around 287

pseudo 7-fold axis [87]. Ribosome inactivating protein enzymes which eliminate one or more adenosine residue from 288

rRNA inactivates ribosomes, inhibit proliferation of HIV- 1 reverse transcriptase activity [88]. 289

Another isolated bioactive compound from mushroom is laccase, which is a multicopper oxidase with its 290

implication in inhibiting HIV-1 reverse transcriptase, proliferation of hepatoma HepG2 cells and MCF 7 tumour cell 291

[89]. Fungal immunomodulating proteins was isolated from various mushrooms which include Ganoderma sp., 292

Russula paludosa, Pleurotus citrinopileatus, Grifola frondosa, Antrodia camphorata [90, 91, 92, 93, 94, 95, 96, 97] 293

with various immunomodulating, antitumour and antiviral activity. A novel Se-containing protein Se-GL-P (36 kDa) 294

isolated from Ganoderma lucidum using ammonium sulphate precipitation, indicated the incorporation of selenium 295

in the protein in the form of selenocysteine and selenomethionine having a positive response towards inhibiting the 296

multiplication of tumour cells [91]. Various other proteins have been isolated which include trichogin protein from 297

Tricholoma giganteum with their antifungal activity against Mycospaerella arachidicola, Fusarium oxysporum and 298

Psylla piricola [98]. Chen et al (2009) isolated a glycoprotein PCP-3A from the fresh fruiting body of oyster 299

mushroom which showed a positive antitumour activity [94]. A report by Chang et al (2007) assessed the utilization 300

of two proteins, Agaricus bisporus lectin (ABL) from Agaricus bisporus and immunomodulating Agaricus polytricha 301

protein (APP) from Auricularia polytricha which showed good stability after thermal, freezing, acid, alkali and 302

12

dehydration treatments, thus indicating the proteins to be stable immune-stimulant for food, health and 303

pharmaceutical applications [99]. 14-3-3 protein with molecular mass of 28.8kDa was isolated from Sparassia crispa 304

and Hericium erinaceum which can be a potential candidate protein for phylogenetic relationship analysis among 305

mushroom species because of its molecular biological feature such as similar pI, molecular mass and slightly 306

differing amino acid sequence [100]. 307

LIPID COMPOSITION 308

Fatty acids are considered to play a major role in immune system functioning and balancing the hormonal level [101]. 309

In basidiomycetes, lipid content ranges from 0.6 to 18.4% (w/w) [102]. The fatty acid profile of mushroom is 310

relatively higher in basidiomycetes which ranges from saturated palmitic acid (16: 0) to oleic acid (18:1, Δ9c) and 311

linoleic acid (18:2 Δ9c, 12c) that are present in the membrane of basidiomycetes [102, 103]. Mushrooms are mostly 312

dominated by the presence of unsaturated fatty acids. However, low proportion of oleic acid is characterized for the 313

mushroom species Agaricus spp. and Cantharellus cibarius [19]. Boletus species are known for their higher 314

concentration of monoenic acid. The concentration of oleic acid are higher in B. edulis, B. piperatus, B. subglabripes, 315

B. erythropus, B. subtomentosus and B. variipes as reported by Hanus et al (2008)[101]. From the above mentioned 316

Boletus species, the polar lipids, including phospholipids and betaine, showed 317

diacylglyceryltrimethylhomoserine(DGTS) and phosphotidylcholine (PC) as a major polar lipid of varying 318

concentration of 72-93% from the total amount of polar lipids. Neutral lipid content ranges from 0.7 to 9.4% and the 319

range of polar lipid content in case of Agaricus sp. ranges from 2.4 to 11.8% [102]. The quantity of polar lipids 320

account for more than 50% of total lipids. Unsaturated fatty acids account for an average of 74.4 % of the total fatty 321

acid. The ratios of unsaturated to saturated fatty acid (U:S) ranges from 1.61 for Lycoperdon pyriforme to 5.36 for 322

Boletus edulis. However, few unusual fatty acids, cyclopropane FA, 11,12 methyleneoctadecanoic acids are found in 323

nonpolar lipid fraction of many basidiomycetes [103]. Davidoff and Kom (1963) reported the presence of isomer cis-324

11 of heptadecenoic acid in the basidiomycetes(104). Elaidic acid (18:1 D9t), a common fatty acid in milk fat and in 325

the tissues of ruminant animals and occasionally in seed oils are found in few basidiomycetes as reported by Pfeuffer 326

and Schrezenmeir (2006)[105]. From human nutritional point, long chain of polyunsaturated linoleic acid and α-327

linolenic acid are important for basal metabolism in human and mushrooms are known to contain all these additional 328

value. Overall, the low value of lipid content in basidiomycetes is due to low proportion of desirable n-3 fatty acid. 329

CARBOHYDRATES AND FIBER CONTENT 330

13

The polysaccharide of mushroom is glycogen unlike that of starch as in plants. Mannitol, glucose and α, α- 331

trehalose(alphaD-glucopyranosyl-(1-1)alpha-Glucopyranoside) are main representatives of monosaccharides. 332

Carbohydrates usually account for the prevailing component of fruiting bodies. The content of carbohydrate on dry 333

weight basis ranges from 47.38 to 63.40g/100g in various basidiomycetes including Agaricus bisporus, Lentinus 334

edodes, Pleurotus ostreatus, Pleurotus pulmonarius [106]. However, the contents of trehalose and glucose in the 335

fruiting body of mushrooms are low, in order of 100 per gram of dry matter. In Agaricus bisporus, the synthesis of 336

mannitol is mediated by NADPH-dependent mannitol dehydrogenase using fructose as its substrate. It functions as 337

an osmolyte, which get accumulated at higher concentration in the fruiting body while after sporulation its amount 338

decreases drastically. Trehalose also serves as reserve carbohydrate in Agaricus bisporus which get synthesized in 339

mycelium and later translocated to fruiting body [107]. Mannitol concentration also varies widely among 340

basidiomycetes. The concentration is found to be of 1.0, 6.5 and 13.7% for dry matter of Tricholoma portentosum, 341

Agaricus arvensis and Lactarius deliciosus respectively [108] and 0.8, 0.2, 11.7 and 13.9% of dry matter of Lepista 342

Nuda, Lycoperdon perlatum, Ramaria botrytis and Cantharellus cibarius respectively [19]. The amount of trehalose 343

and mannitol get reduced considerably while boiling the fresh mushrooms, while freezing and drying result in only 344

limited losses [108]. In Cantharallus cibarius, trehalose and arginine are the most important compounds for carbon 345

assimilation [108]. Chitin is a water-insoluble structural polysaccharide that accounts for up to 80–90% of dry matter 346

in cell walls of mushroom. Contents of dry matter as reported from eight samples of Boletus spp. ranged from 6.8–347

10.2% [74]. Chitin being indigestible for humans apparently decreases the availability of other mushroom 348

components. 349

Information on dietary fibre content in wild growing mushrooms has been very limited. Boletus spp. is reported to 350

contain 4.2–9.2% and 22.4–31.2% of dry matter for soluble and insoluble fibres respectively for oyster mushroom 351

and Agariocus bisporus [109, 70]. Cheung (1997) determined hemicelluloses and pectic substances in 352

basidiomycetes [110]. Information on chitin and fibre changes during different preservation and cooking remedy has 353

been lacking. Great attention has recently been given to fiber content of the mushroom, the β-glucans which act as a 354

health-promoting factor [86]. A recent study reported that two mushroom carbohydrates inhibit breast cancer cell 355

growth by enhancing the immune function [111]. 356

MINERAL COMPOSITION 357

Wild mushrooms are assumed to accumulate a large amount of both macro and micro-minerals. Mineral content 358

14

varies among different species of basidiomycetes. A research on 11 different mushroom species showed the level of 359

potassium concentration in Tricholoma saponaceum to be around 39.8 mg/g, which is lower in case of Laccaria 360

laccata and Candida rugosa being 30.2 mg/g and 28.9 mg/g [112]. Selenium content in the tubes and gills of 361

mushrooms are quite high which act as an antioxidant. It is also required for the biosynthesis of selenoenzymes 362

which include glutathione peroxidase, thioredoxin reductases, Selenoprotein W and Selenoprotein P, iodothyronine 363

5’-deiodases. Selenium content in mushrooms ranges from approximately 5μg/g in Lycoperdon spp. to about 200μg/g 364

in Albatrellus pes-caprae. King Bolete(Boletus edulis) is considered to contain an average concentration of 365

approximately 20μg/g of selenium whereas Pinewood King Bolete(Boletus pinicola) contain selenium on an average 366

of 40μg/g based on its dry weight. The fruiting body of Lurid Bolete(Boletus luridus), Lepista luscina and Parasol 367

Mushroom (Macrolepiota procera) contained, respectively, 49, 91 and 47 of selenium in crude mushroom[113]. 368

Potassium is not distributed evenly within fruiting bodies, the concentration seems to be higher in cap followed by 369

stipe, spore forming part and finally to spores. It seems to be higher by 20 to 40 folds in the fruiting bodies than the 370

remaining part [114]. Followed by potassium, the second most abundant major element in mushroom fruiting bodies 371

is phosphorus which ranges from 5 to 10 g per kg of dry matter [19]. A report shows high amount of calcium 372

concentration of 1.600 mg/g in Craterellus tubaeformis followed by 10.50 mg/g in Laccaria Laccatta [112]. Sodium 373

concentration is relatively less in mushroom species due to which they have been considered good for patient with 374

hypertension which ranges from 100 to 400 mg per kg of dry matter [19]. Few mushrooms including Formitopsis 375

officinalis offer chlorine in the form of coumarine which include 6-chloro-4-phenyl-2H-chromen-2-one and ethyl 6-376

chloro-2-oxo-4-phenyl-2H-chromen-3-carboxylate [115]. As per the report of Rudawska and Leski (2005), total 377

sulphur content per kg of dry matter in Amanita rubescens and Xerocomus chrysenteron is found to be in between 378

900 and 4400 mg per kg dry matter respectively [116]. Mushrooms are considered to contain high amount of trace 379

elements including cadmium, mercury, lead, copper, antimony in their fruiting bodies [117]. Agaricus spp. 380

accumulates a high level of mercury and cadmium upto a range of 130 mg of cadmium per kg of dry weight of 381

fruiting body [118]. Other mercury accumulators include Lycoperdon perlatum, Lepista spp. and Macrolepiota spp. 382

[118]. Chromium and nickel concentration is higher in Lactarius deliciosus with around 4.51 mg per kg of dry 383

weight and 9.9 mg per kg of dry weight respectively in Tricholoma terreum [119]. Mushrooms are also known as 384

zinc accumulator. Lactarius spp. is known for its higher concentration of zinc accumulation which ranges from 385

55.7mg/kg to 158mg/kg on the basis of its dry weight. The gills of Clitocybe alexandri are considered to contain high 386

15

amount of copper which ranges from 26.3 to 95.9mg/kg on its dry weight basis. The concentration of iron seems to 387

be slightly higher in Volvariella speciosa which ranges from 220 to 7162 mg/kg [120]. 388

During Chernobyl accident, mushrooms fruiting bodies (radiotrophic fungi) were seen to accumulate huge amount of 389

toxic heavy metal [121]. It has been reported that natural radionuclide 40

K accumulates highly in mushroom and 390

accumulation factor ranges from 20 to 40 [114,122]. However, Boletus group are considered to contain high amount 391

of selenium content [19]. Heavy metal radiocaesium are also found to a greater extent in mushrooms of coniferous 392

forest then deciduous forest [114, 123]. 393

CONCLUSION 394

Various wild macrofungi are traditionally used as a delicacy and source of medicine since time immemorial. 395

However, large sections of their population are still unknown due to fragmentary and poor information about their 396

bioavailability and toxicity. Furthermore, the use of morphological and molecular identification based on ITS 397

sequences can to some extent be used in terms of their ecological perspective. 398

n-3 fatty acid concentration is low in mushroom which contributes negligibly in its nutritional profile but high 399

proportion of insoluble fibre encompassing chitin, β-glucan and other polysaccharides seems to be highly valuable 400

both in nutritional and pharmacological aspects. With the high concentration of unsaturated fatty acid, they prove to 401

be beneficial for basal metabolism in human. The concentration of potassium and selenium is high in the tubes and 402

gills of mushrooms for which they can be used as a source of antioxidant. Few fruiting bodies can accumulate quite a 403

large concentration of trace elements and radioactive components which make them vulnerable to consume. However, 404

a large section of poisonous mushrooms resemble their non-toxic counterparts. So, a need of a barcode to 405

differentiate their toxic nature will further develop their biological and therapeutic application and open vistas for 406

their bio-prospection and use for human benefits. 407

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LEGENDS TO FIGURES 653

Fig I Wild macrofungi : (MS1)Formitopsis spp., (MS2)Russula spp., (MS3) Mycena spp., (MS4)Hypholoma spp., 654

(MS5)Trichaptum spp., (MS6)Panus spp., (MS7)Galerina spp., (MS8)Lactarius spp. 655

Fig II Phylogenetic tree generated for edible macrofungi 656

Fig III Phylogenetic tree generated of inedible macrofungi 657

25

TABLE I. Essential amino acid per 100 grams of proteins of mushroom [82, 83, 124]

Amino acid Mycoprotein (g) Egg (g) Milk (g) Wheat (g)

Arginine....................... 6.7 6.4 - -

Histidine...................... 0.35- 2.1 2.1 0.09 0.32

Lysine.......................... 0.83- 5.0 7.2 0.26 0.30

Tryptophan.................. 0.16- 0.9 1.5 0.05 0.18

Phenylalanine.............. 0.49- 2.0 6.3 0.16 0.68

Methionine.................. 0.21- 1.26 4.1 0.08 0.22

Threonine.................... 0.55- 4.2 4.9 0.15 0.37

Leucine....................... 0.86- 4.4 9.2 0.32 0.93

Isoleucine................... 0.52- 5.8 8.0 0.20 0.53

Valine.......................... 0.62- 4.65 7.3 0.22 0.59

TABLE II. Nutritional composition in mushroom fruiting body g/100g (% of dry matter)

Species Crude protein Lipid Carbohydrates Fibre Reference

Agaricus bisporus 26.3- 56.3 1.8-2.7 37.5-59.96 - [50, 51]

Agaricus arvensis 56.3 2.7 37.5 - [50]

Amanita rubescens 26-31.9 7.2 - 27.5 30.6-62.2 - [50, 84, 85]

Armillariella mellea 21.9 1.8-6.08 70 - [50, 84]

Auricularia polytrica 7.7 0.8 87.6 - [51]

B. Aereus 26.9 2.1 34 17.0 [86]

Boletus edulis 26.5-28.7 2.8-4.1 30.6- 65.4 15.3 [50, 86]

B. speciosus 28.1 2.9 28.6 21.0 [86]

Cantharellus cibarius 34.17- 53.7 1.40- 2.9 31.9-57 - [50, 84]

C. aureus 14.1 4.0 61.5 5.2 [86]

C. Cornucopioides 50.10 5.89 34 - [84]

Flaninnilbia velutipes 17.6 1.9 73.1 - [51]

H. repandum 34.14 8.80 55 - [84]

Lactarius deliciosus 20.2- 29.8 2.2-4.0 5.1-25 36.8 [50, 51]

Lactarius hatsudake 15.3 1.0 38.2 31.8 [51]

Lactarius volemus 17.6- 87.57 3.98-6.7 15- 64 40.0 [84, 51]

Lentinula edodes 17.1 1.9 30.2 39.4 [51]

Lentinula modes 17.5 8.0 67.5 - [86]

Lepista nuda 19.8-59.4 1.8-9 20.3-71.0 - [50]

Lycoperdon perlatum 17.2- 44.93 0.4 -10.58 42- 50.4 - [50, 84]

L. Crocipodium 29.3 1.0 12.8 37.9 [86]

Macrolepiota procera 23.9 2.3 68.4 - [50]

Pleurotus ostreatus 10.5 1.6 81.8 - [51]

Ramaria botrytis 39.0 1.4 50.8 - [50]

R. Flava 35.55 5.20 65 - [84]

R. virescens 28.3 1.5 13.4 32.8 [86]

S. Imbricatus 27.45 8.85 57 - [84]

Suillus granulatus 16.5 4.0 74.3 - [50]

S. aspratus 12.0 2.8 64.6 5.1 [86]

Tricholoma flavovirens 18.1 2.0 37.0 - [50]

Tricholoma portentosum 19.6-30.5 5.5-5.8 34.6 30.1 [87]

Tricholoma terreum 20.1 6.6 31.1 30.1 [87]

Trcinclla fuciforinis 4.6 0.2 94.8 - [51]

T. Matsutake 14.3 5.0 36.7 29.1 [86]

Volvariella volvacea 29.3 5.7 60.0 - [51]

TABLE III. Accumulation of various trace elements by the mushroom fruiting bodies [78, 79, 88, 89, 90]

Element Content Accumulators

Antimony 0.05-0.15 Suillus sp., Laccaria amethystina, Amanita rubescens

Arsenic < 1 Laccaria amethystina, L. Laccata, Agaricus sp., Lepista nuda

Lycoperdon perlatum

Beryllium < 0.01 Boletus edulis, Lactarius trivialis

Cadmium 0.15-30.0 Agaricus sp. (group flavescentes), Clitocybe alexandri,

Clitocybe flaccida, Lepista inversa, Volvariella speciosa,

Amanita soliteria

Caesium 3-12

Chromium 0.1-2 Agaricus sp., Macrolepiota procera, Lacterius deliciosus

Cobalt <0.1- 7.49 Agaricus arvensis, Clitocybe alexandri, Clitocybe flaccida,

Lactarius sanguifluus

Copper 10-95.9 Agaricus sp., Macrolepiota procera, M. Rhacodes,

Clitocybe alexandri

Gold < 0.02 Boletus edulis, Morchella esculenta

Iron 30-644 Suillus variegatus, Suillus luteus, Hygrophoropsis aurantiaca,

Clitocybe alexandri

Lead 0.09-6.88 Macrolepiota rhacodes, M. procera, Lycoperdon perlatum,

Agaricus sp., Lepista nuda, Volvariella speciosa, Lentinus

tigrinus, Amanita soliteria, Lactarius deliceus, Russula delica

Manganese 4.67-60 Agaricus sp., Clitocybe alexandri

Mercury < 0.5-5 Agaricus sp.,Macrolepiota procera, Lepista nuda,

Macrolepiota rhacodes

Nickel 0.4-5.72 Laccaria amathystina, Leccinum sp., Clitocybe alexandri

Selenium 1-5 Boletus edulis, B. penicola, B. aestivalis, Albatrellus pes-

caprae, Xerocomus bodius

Silver 0.2-3 Amanita strobilisformis, Agaricus sp., Boletus edulis,

Lycoperdon perlatum

Strontium 0.22-0.67 Lactarius trivialis, Boletus edulis

Thallium < 0.25

Zinc 30-158 Suillus variegatus, Suillus luteus, Lycoperdon perlatum,

Lactarius semisanguifluus