The Biology and Cultivation of Edible Mushrooms

CONTRIBUTORS

J. ANGELI-PAPA

IKUO ARITA

S. T. CHANG

P. C. CHEN

S. CHENG

KENNETH W. COCHRAN

E LI V. CRISAN

J. R. DELCAIRE

J. DELMAS

R. L. EDWARDS

G. EGER

J. EYME

GERDA FRITSCHE

G. GRAMSS

W. A. HAYES

M. S. HO

H. H. HOU

TATSUZIRO ITO

S. C. JONG

D. S. KIM

M. KOMATSU

R. H. KURTZMAN, JR.

CARLENE A. RAPER

ANNE SANDS

ALEXANDER H. SMITH

K. SZUDYGA

K. TOKIMOTO

Y. TOMINAGA

H. TONOMURA

C. C. TU

P. J. C. VEDDER

F. ZADRAZIL

THE BIOLOGY AND

CULTIVATION OF

EDIBLE MUSHROOMS

EDITE D B Y

S.r. Chang

Department of Biology

The Chinese University of Hong Kong

Shatin, N. T., Hong Kong

W. A. Hayes

Department of Biological Sciences

University of Aston

Birmingham, England

ACADEMIC PRESS New York San Francisco London 1978

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ACADEMI C PRESS , INC . (LONDON ) LTD .

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Library of Congress Cataloging in Publication Data

Main entry under title:

The Biology and cultivation of edible mushrooms.

Includes bibliographies and index.

1. Mushroom culture. 2. Mushrooms, Edible.

I. Chang, Shu-ting, Date II. Hayes, W. A.

SB353.B47 635\8 77-6591

ISBN 0-12-168050- 9

PRINTED IN THE UNITED STATES OF AMERICA

List of Contributors

Numbers in parentheses indicate the pages on which the authors* contributions begin.

J. ANGELI-PAPA (53), Station de Recherches sur les Champignons, LN.-

R.A., 33140 Pont de la Maye, France

IKUO ARITA (475), The Tottori Mycological Institute, Japan Kinoko Re￾search Centre Foundation, Tottori, Japan

S. T. CHANG (35, 573), Department of Biology, The Chinese University

of Hong Kong, Shatin, N. T., Hong Kong

P. C. CHEN (629), Department of Plant Pathology and Entomology,

National Taiwan University, Taipei, Taiwan

S. CHENG (605), Department of Horticulture, National Taiwan Uni￾versity, Taipei, Taiwan

KENNETH W . COCHRAN (169), Department of Epidemiology, School of

Public Health, and Department of Pharmacology, Medical School,

University of Michigan; Ann Arbor, Michigan 48109

E L I V. CRISAN* (137), Department of Food Science and Technology,

University of California, Davis, Cahfomia 95616

J. R. DELCAIRE (727), 8 Rue des Dardanelles, 75017 Paris, France

J. DELMAS (251, 645, 699), Station de Recherches sur les Champignons,

I.N.R.A., Centre de Recherches de Bordeaux, Domaine de la Grande

Ferrado, 33140 Pont de la Maye, France

R. L. EDWARDS (299), 2 Landway, Bearsted, Maidstone, Kent ME14 4BD,

England

G. EGER (497), Institut fόr Pharmazeutische, Technologie der Universitδt

Marburg, D-3550 Marburg, West Germany

J. EYM E (53), Universite de Bordeaux I, Laboratoire de Botanique,

33405 Talence, France

GERDA FRITSCHE (239, 371), Proefstation voor de Champignoneultuur,

* Present address: 690 Alvarado Avenue, Apartment 13, Davis, Cahfomia 95616.

XV

XVI List of Contributors

Peelheideweg 1, Horst-America 5960-AA, Postbus 6042, The Nether￾lands

G. GRAMSS (423), 69 Jena-Winzerla, Grenzstrasse 28, German Demo￾cratic Republic

W. A. HAYES (191, 219), School of Biological Sciences, The University

of Aston, Birmingham B4 7ET, England

M. S. Ho (337), Taiwan Provmcial Farmer's Association, Taichung,

Taiwan

H. H. Hou (629), Department of Plant Pathology and Entomology,

National Taiwan University, Taipei, Taiwan

TATSUZIRO ITO (461), Mushroom Research Institute of Japan, Kiryu,

Japan

S. C. JONG (119), Mycology Department, American Type Culture Col￾lection, Rockville, Maryland 20852

D. S. KIM (345), Department of Apphed Mycology and Mushrooms,

Institute of Agricultural Sciences, OflBce of Rural Development,

Suweon, Japan

M. KOMATSU (445), The Tottori Mycological Institute, Japan Kinoko

Research Centre Foundation, Tottori, Japan

R. H. KURTZMAN, JR. (393), Western Regional Research Laboratory,

Agricultural Research Service, U.S. Department of Agriculture,

Berkeley, California 94710

CARLENE A. RAPER (83, 365), The Biological Laboratories, Harvard Uni￾versity, Cambridge, Massachusetts 02138

ANNE SANDS (137), Division of Environmental Studies, University of

California, Davis, California 95616

ALEXANDER H. SMITH (3), University of Michigan Herbarium, North

University Building, Ann Arbor, Michigan 48109

K. SZUDYGA (559), Research Institute of Vegetable Crops, 96-100 Skier￾niewice, Poland

K. TOKIMOTO (445), The Tottori Mycological Institute, Japan Kinoko

Research Centre Foundation, Tottori, Japan

Y. TOMINAGA (683), Genetics Institute, Hiroshima Agricultural College,

Higashi Hiroshima City, 724, Japan

H. TONOMURA (409), Mori Mushroom Research Institute, Kiryu, Japan

C. C. Tu (605), Department of Plant Pathology, Taiwan Agricultural Re￾search Institute, Taipei, Taiwan

P. J. C. VEDDER (377), Director Mushroom Growers Training Centre,

Westerholstraat 2, 5961 BJ Horst (L) , Holland

F. ZADRAZIL (521), Institut fόr Bodenbiologie, FAL, Bundesallee 50, 3300

Braunschweig, Federal Republic of Germany

Fore wore

It is an honor and a pleasure for me to write the Foreword for "The

Biology and Cultivation of Edible Mushrooms."

Increasing interest in the cultivation of mushrooms- has been amply

demonstrated by the succession of international congresses on the science

and cultivation of edible mushrooms, that have attracted more and more

participants. Since 1950 these congresses have been organized every three

years. All but one were held in Western Europe; the 1962 congress was

held in the United States.

At these congresses most attention has focused on the continuing re￾search on the growth and cultivation of Agaricus bisporus ("cultivated

mushroom," "champignon de couche," "champignon de Paris"), cultiva￾tion (on beds or couches of composted horse manure) of which began

in the vicinity of Paris about 1650.

The early organizers of the congresses were almost exclusively involved

with this species. However, it was gradually discovered that Agaricus

was not the only edible higher fungus being grown commercially. At the

International Mushroom Congress held in Amsterdam in 1965, Dr. K.

Mori of Japan showed his film on the cultivation of several species of

edible fungi in eastern Asia, which "opened the eyes" of the western

world. The cultivation of Pleurotus ostreatus and of Stropharia rugoso￾annulata started independently in Eastern Europe shortly after this

congress.

Matsutake (Tricholoma matsutake = T. caligatum) is the most popular

fungus in Japan; it has challenged Japanese researchers and growers for

many years. A symposium was held in 1964 in Kyoto to discuss modes

of cultivating this fungus. However, even at present it still has to be col￾lected in the woods and has not been cultivated artificially.

The famous "truffle du Pιrigord" (Tuber mehnosporum) of France

and other truffle species from the countries around the Mediterranean

are also highly valued fungi. International congresses on the cultivation

xvii

xviii Foreword

of truffles were organized in 1968 in Italy and in 1971 in France, and

successful cultivation of these mycorrhizal fungi has been initiated.

It was fortunate that the Ninth International Congress (1974) could

be held in eastern Asia (partly in Japan and partly in the Republic of

China). Field excursions enabled the participants to observe commercial

cultivation of various fungi. During the East Asian gathering the name

of the international congresses was changed to "International Congress

on the Cultivation of Edible Fungi."

Mutual discussions among the participants of this 9th Congress re￾sulted in Dr. S. T. Changes securing the cooperation of prominent re￾searchers and practical workers on mushrooms and other edible fungi to

write about their special fields of interest. Their contributions comprise

this comprehensive volume.

Books in the Japanese language by Tottori and Sakari Hisamune on

the cultivation of Lentinus, Flammulina, Volvariella, Pholiota, Pleurotus,

Agaricus, and Ganoderma are basically manuals for cultivating, their

language making them accessible to only a limited circle of readers. This

volume, edited by Dr. Chang and Dr. W. A. Hayes, should reach a much

wider audience with a much greater variety of interests.

The fungi dealt with in this book, which are presently under cultivation

or may be cultivated in the future, can tentatively be divided into the

following groups, according to their occurrence in nature.

1. Fungi that grow on fresh or almost fresh plant residues: Lentinus,

Pleurotus, Flammulina, Auricularia, Pholiota, Tremella, Agrocybe,

Ganoderma, Coprinus

2. Fungi that grow on only little composted material: Volvariella,

Stropharia, Coprinus

3. Fungi that grow on well or very well composted material: Agaricus

4. Fungi that grow on soil and humus: Lepiota, Lepista, Morchella,

Gyromitra

5. Mycorrhizal fungi: Boletes, Cantharellus, Amanita, Tuber, Matsu￾take, Morchella, Lactarius

The coverage in each chapter varies in depth; for example, more ex￾tensive research has been done on Agaricus bisporus and Lentinus edodes

than on various other fungi. Gradually more and more will be known

about the biology of other fungi. It is my sincere hope that this book will

lead to new ideas and stimulate further research.

P. J. Bels

President of Society International Commission

on Mushroom Science

January, 1976 Horst, Limburg, Holland

Preface

The word mushroom may mean different things in diflFerent books

and countries. In this book edible mushrooms refer to both epigeous

and hypogeous fruiting bodies of macroscopic fungi that are already

commercially cultivated or grown in half-culture processes or potentially

implanted under controlled conditions. Mushrooms are occasionally de￾scribed as all kinds of fungi (Thallophyta) that lack chlorophyll; but in

this book we deal only with those mushrooms that are edible, i.e., usable

as food and in some manner grown commercially.

Two motivating forces have been responsible for the compilation of

this book. The first is academic, which we believe is common to any

attempt of this kind. It is our zealous desire to combine the knowledge

of the biological nature of edible mushrooms with established methods

of practical application (cultivation). In so doing, the information on

the subject of edible mushrooms made available by many biologists and

researchers during recent years has been organized and summarized in

a manner readily understandable, and hence accessible, to any scientist

or layman. This work is the product of the collaborative eflForts of a

group of the most knowledgeable men in their respective fields, consisting

of 33 fungal specialists representing scholars from 11 diflFerent countries.

The second motivating force, more commanding than the one just

considered, stems from the fact that edible mushrooms are rich in pro￾teins, and thus constitute a valuable source of supplementary food (ac￾cording to current statistics, worldwide mushroom production annually

is of the order of 700,000 to 900,000 metric tons). In fact, mushrooms

have played an important role in the diet of many people for thousands

of years.

That mushrooms can serve as food is not, itself, of major importance.

However, the way in which edible mushrooms can be grown greatly

enhances their importance. As this book shows, a great variety of cheap

(some almost worthless) materials, e.g., agricultural wastes, industrial

XX Preface

wastes, and family wastes, can be successfully used as media for growing

the various edible mushrooms. It is reahzed also that these cultivation

methods, which have already proved practical, may be just a beginning.

More efficient and far cheaper means are certainly within the realm of

realization as more and more people become interested in mushroom

research. With this rapidly expanding human population (at the current

rate of 2.1%, representing a rise of about 75 million people per year, or

200,000 daily), the possibility of using edible mushrooms to assist in

alleviating the worldwide food shortage should obviously be vigorously

explored.

This book emphasizes both the academic (biology) and the applied

(cultivation) aspects of edible mushrooms. The reason for this is simple.

In practically every science there are always these two aspects to con￾sider—it is like the two faces of a coin, inseparable and yet distinct.

Knowledge of one is bound to enhance that of the other. The two are

invariably complementary to one another, and only by combining the

existing knowledge of both in the same treatise, as is done in this book,

can we hope to integrate the science of edible mushrooms to the fullest

extent, hopefully for the mutual benefit of the researchers, on the one

hand, and the growers, on the other. This book, therefore, aims to bring

these two groups into closer touch with one another than ever before.

It is an indisputable fact that a gap still exists beween the academic

fungal researchers and the commercial mushroom growers. It is our hope

to integrate the knowledge of researchers and the advanced techniques

of the growers so that both will be mutually benefited. We are convinced

that researchers should know the problems faced by growers and that

the growers, in turn, should understand some of the basic biological facts

about the mushrooms which they cultivate. In this connection, one might

add a significant point. Growers should bear in mind that climatic con￾ditions as well as economic situations vary from one locale to another in

various parts of the world, thus creating differences in availability of

materials for media, methods for producing these media, and possibly

the use of manual labor/machines, etc. These variables are always present

in contrast to the "biology*' of any given species of mushroom, which is

more or less the same all over the world.

This book is intended for anyone who is interested in edible mush￾rooms: the experienced mushroom specialist, seasoned commercial grow￾ers, biology students, or simply the layman.

Our principal hope for this book is that it will encourage and stimulate

further research on all aspects of edible mushrooms, with special attention

directed toward discovering new edible species and improving both the

quality and yield of existing ones. This should eventually lead to their

Preface xxi

mass production accompanied by reduction in cost. A number of require￾ments are needed to realize these hopes. First, more extensive chemical

analysis should be carried out, subjecting the various components to well￾planned tests in an attempt to assay their nutritional values. These data

are important since they can provide us with a basis for objectively eval￾uating any edible mushroom. Second, there is a great need for experi￾ments aimed at producing mutations which will improve the productive

capacity and efficiency of growing methods as well as possibly increasing

food values. These are important steps which should lead to a substantial

boost in the uses of edible mushrooms and their ultimate contribution

to mankind.

Last, but not least, we look forward to the day when basic and applied

research on edible mushrooms commands financial support of similar

magnitude to that devoted to other crops. The position we are taking is,

indeed, not difficult to justify. At present, edible mushrooms have made

a positive contribution to the diet of millions of people. Faced with the

prospect of a deteriorating food supply, should we not endeavor to launch

a program to increase the production of edible mushrooms? If this book

serves as a stimulus in hastening that day, then the efforts of all those

who contributed to it will not have been spent in vain.

We have been greatly encouraged by the financial support from the

Asia Foundation for the preparation of this book. We would like to ex￾press our appreciation to P. J. Bels for writing the Foreword and to the

many contributing authors. Special thanks are due to R. J. Bandoni, Uni￾versity of British Columbia; K. M. Graham, University of Malaya; J. W.

Kimbrough, University of Florida; B. C. Lu, University of Guelph; L. B.

Thrower, Chinese University of Hong Kong; and L. C. Wu, University

of Wisconsin for reviewing some of the manuscripts. We are also grateful

to Academic Press for publishing the book.

S. T. Chang

W. A. Hayes

EDIBLE MUSHROOMS

1

Morphology and Classification

ALEXANDER H. SMITH

I. Introduction 3

II. The Ascomycetes 5

III. The Basidiomycetes 6

A. The Agaricales 8

1. The Basidiocarp 9

2. Hyphal Systems and Hyphal Characters 18

3. The Spores 23

IV. Edibihty and Poisonous Properties 25

V. Family Agaricaceae Fries 26

A. Type Genus: Agaricus Fries 26

Agaricus Fries , 26

B. Type Species: Agaricus campestris Fries 26

VL Family Strophariaceae Singer & Smith 28

A. Type Genus: Stropharia (Fr.) Quelet 28

VIL Family Pluteaceae Kotl. & Pouzar 29

A. Type Genus: Volvariella Speg 30

Volvariella Speg 30

B. Type Species: Volvarielh argentina Speg 30

VIIL Family Tricholomataceae Roze 30

References 33

I. INTRODUCTION

To understand the present grouping of diverse species in a single

volume such as this, one must keep in mind that the higher fungi are a

large and heterogeneous group, numbering in the thousands of species,

and that people all over the world, at one time or another have tried them

as a source of food. Some of the experimentation on edibility, undoubtedly,

3

Copyright © 1978 by Academic Press, Inc.

All rights of reproduction in anv form reserved.

ISBN 0-12-168050-9

4 Alexander Η. Smith

were disastrous and some highly successful. It is logical to assume that in

different regions different species were tried, and the technique of obtain￾ing the desired ones was in a large measure the same as it is now: in spite

of our present degree of development of mushroom growing as an in￾dustry, most of the species eaten are collected in the wild. We assume

that the first attempts at mushroom growing were made by some who

observed that a desirable species was usually found around a straw stack

or a heap of cow manure, and they decided to enhance the habitat in one

way or another to obtain larger yields. Also, they no doubt transplanted

samples from the original pile to new piles of substrate by way of in￾creasing total production. From such beginnings, as new knowledge was

uncovered relative to the nature of the mushroom plant, it took only a step

to transfer this primitive industry into the laboratory and to concentrate on

the species which could be made to produce mushrooms on a predictable

regime. At present these studies are being continued relative to bringing

more species into the commercial trade and to obtaining optimum yields

at lowest possible cost. In short, the cultivation of mushrooms on an inter￾national basis does not appear to have a single point of origin nor was it

limited to a single species. Consequently, in such a work as this, one finds

an assemblage of species selected purely on the basis that at some time in

the past it was found that their fruit bodies (the mushrooms) were edible

and that their production could be programmed in one way or another.

In order to establish a background for the discussion of each species

treated here, it is proper to fit them into the system of classification cur￾rently in use by mycologists, and which, by and large, is thought to indi￾cate relationships based on descent from a common, remote ancestor (or

ancestors). Such a scheme is termed a natural or phylogenetic arrange￾ment. In it species are considered closely related if they have in common

a large number of features found to be relatively stable in the population.

Since the sporocarp is composed of threads (hyphae) of microscopic

size, many of the features can be determined only with the aid of a micro￾scope. Such features are termed microscopic characters. Those which can

be observed with the naked eye or with the help of a hand lens are termed

macroscopic or gross characters. Both sets must be considered in postulat￾ing the degree of relationship between any two populations. In systematic

studies any one of the various recognized units (as categories in a hierar￾chy of categories) is termed a taxon (pi. taxa). An order, a family, a

species, and each subdivision of any of the above, is a taxon. Since our

current classification of the higher fungi is based on both micro- and

macroscopic details, one must be familiar with the structural details of

the fruit bodies and spores to understand it. Currently, "chemicar char￾acters are also used. These involve both empirical tests for color changes

1. Morphology and Classification 5

II. THE ASCOMYCETES

The edible fungi under cultivation for the production of sporocarps all

fall into two major groups: Ascomycetes and Basidiomycetes. Considering

the Ascomycetes first, much attention has been given to culturing (or try￾ing to culture) members of the Discomycetes and the related order Tuber￾ales (truffles). In the Discomycetes (Cup Fungi), attention has been

focused on the genus Morchella of the family Morchellaceae. The asco￾carps in this family are more or less club-shaped and the upper part of

the club is pitted. The club consists of a thin shell of tissue around a

hollow interior. Thus, in spite of the size of the ascocarp, there is relatively

little substance to it. The ascospores are produced in the asci which in

turn are arranged in an hymenium (Fig. 1), and are forcibly discharged

from the asci at maturity. The discharge of many asci together often

produces a visible cloud of spore dust and this is accompanied by a slight

hissing sound. In technical descriptions the stalk is referred to as a stipe.

The fertile upper portion is the head even though the shape is not as a

rule globose. The taxonomy of the species of Morchella is still in an un￾settled state. One finds some authors recognizing many species and some

only about a half dozen. The species most used in culture attempts is

Morchella esculenta Fries. Uncertainty as to the concept of this species

still exists as is evident if one compares the illustrations which have ap￾peared in fairly recent times. Although morels are easy to culture in the

laboratory from tissue or spores, so far attempts to fruit them on a com￾mercial regime have been unsuccessful. Thus, in a sense, the group is

peripheral to the present work. It is the group, however, receiving the

most attention relative to introducing another type of edible mushroom

to the world market. It constitutes a great challenge to the mushroom￾growing industry.

The second group of Ascomycetes, the truffles, belong to the Tuberales,

as previously stated. The ascocarp of the truffle plant may be characterized

as a cup fungus in which the cup remains more or less closed, and the

caused by application of chemicals and the chemical analysis of the

basidiocarp.

In keeping with the concept that our classification is based on relation￾ship by descent, we consider the characters of the reproductive structures

to be of paramount importance, and consequently those cells or tissues in

which nuclei fuse or undergo meiosis or are associated directly with this

process are the starting points in systematics, not only for fungi but for

living organisms generally.

Alexander Η. Smith

III. THE BASIDIOMYCETES

Among the Basidiomycetes which are used commercially for food are

some species of Auricularia and one of Tremella. These will be considered

first, ahead of the main group, the Agaricales, because the fruiting bodies

(basidiocarps) are more or less shapeless to saucerlike and are carti￾laginous to jellylike in consistency. These genera are now included in the

Phragmobasidiomycetidae (Ainsworth et ah, 1973), a group encompassing

three orders: Tremellales, Auriculariales and Septobasidiales. The various

types of basidia featured by this group are shown in Figures 2, 3, 7, and

interior becomes filled with folds and veins growing inward, more or less,

from the shell (peridium). The fruiting bodies are usually referred to as

"tubers,'' no doubt because of the superficial resemblance to a small

potato. The term tuber in mycology is not used in the strict sense of the

vascular plant taxonomist. The ascocarps develop in the soil and rarely

become exposed over the upper surface. This pattern of fruiting, of course,

makes them very difficult to find, and has long furnished a source of

humor to collectors. The spores, when mature, cannot be discharged into

the air as in a morel, and it is assumed that other methods of dispersal

have been evolved, such as dispersal by insects. Evidence for the fact

that the Tuberales are closely related to the Discomycetes, however, is

found in the fact that in some species previously classed as Tuberales,

the asci still discharge spores forcibly even though the ascocarp never

becomes open as in a cup. This can be demonstrated by breaking open an

ascocarp at just the right stage of spore development. A small puff of

spores can be obtained, and an examination of the ascus under the micro￾scope will show that the spores have been discharged in the usual man￾ner of the operculate Discomycetes.

There are a number of species of truffles which are commonly used as

food: Singer (1961) lists Tuber aestivum Vitt., Τ. uncinatum Chatin, Γ.

mesentericum Vitt., Τ. montanum Chatin, and Τ. brummale Vitt. A num￾ber of species of Tuber occur in North America, but the tuber flora of this

continent involves species mostly different from those harvested in France

and Italy. Gilkey (1939) recognized 17 species in North America, but

none, to my knowledge, are currently of commercial importance. The

method of cultivation in France and Italy is based on the environmental

approach. Since the mycelium of the tuber lives in the soil and forms

mycorrhiza with certain species of oak, the technique is to plant the

proper species of oak in the right kind of soil in an area where truffles

have been found, and let nature do the rest.