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Growing Gourmet and Medicinal Mushrooms; A Companion Guide to the Mushroom Cultivator (3rd Ed.) - Ten Speed Press-Crown Publishing-Random House

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This book should help advance the cause of mycology and mushroom
biology worldwide. It will be an important reference for those who
are interested in research as well as in the cultivation of mushrooms.
Growing Gourmet and Medicinal Mushrooms is unique not only in its
treatment of the technical aspects of growing gourmet and
medicinal mushrooms, but also in its emphasis on the environmental
importance of mushrooms in terms of world biological diversity.
—S. T. Chang, Ph.D., The Chinese University of Hong Kong
Growing Gourmet and Medicinal Mushrooms is a visionary quest—
and Paul Stamets is your best possible guide—not just for informing
you about growing mushrooms, but for transforming you into a
myco-warrior, an active participant in a heroic, Gaian process of
planetary health through mushroom cultivation.
—Gary Lincoff, author of The Audubon Field Guide to Mushrooms
Stamets draws on the collective experience of centuries of mushroom
cultivation, creating a revolutionary model for the use of higher
fungi. Not only does he cover every aspect of cultivation, he also
addresses the issues of environmentalism, health, and business.
—Alan Bessette, Ph.D., Utica College of Syracuse University
Growing Gourmet and Medicinal Mushrooms is the most
comprehensive treatment of the subject I have seen in my thirty
years as a mycologist and mushroom specialist.
—S. C. Jong, Ph.D., The American Type Culture Collection
Pick up this book and prepare to be swept away into the world of
mushroom cultivation on the tide of Paul’s contagious enthusiasm.
Doers and dreamers, students and teachers will all find something to
enjoy in this book.
—Nancy Smith Weber, Ph.D., Forest Sciences Department, Oregon State
This book, a true labor of love, makes a major contribution to our

knowledge of the practical production of gourmet and medicinal
—Dan Royse, Ph.D., Penn State College of Agricultural Sciences

​C opyright © 1993, 2000 by Paul Stamets
All rights reserved. Published in the United States by Ten Speed P; ress, an imprint of
the Crown Publishing Group, a division of Random House, Inc., New York.
Ten Speed Press and the Ten Speed Press colophon are registered trademarks of Random House, Inc.
Photograph of Chinese rank badge (Wild Goose, 4th Rank) on this page courtesy of Beverley Jackson.
Cover design by Andrew Lenzer and Jeff Brandenburg
Interior design by Jeff Brandenburg
Library of Congress Cataloging-in-Publication Data
Stamets, Paul.
Growing gourmet and medicinal mushrooms = [Shokuyo oyobi yakuyo kinoko no saibai] : a companion guide
to The Mushroom Cultivator / by Paul Stamets. — 3rd ed.
p. cm.
Includes bibliographical references (p.).
eISBN: 978-1-60774-138-1
1. Mushroom culture. I. Title: Growing gourmet and medicinal mushrooms. II. Title: Shokuyo oyobi yakuyo
kinoko no saibai.
SB353.S73 2000


An environment wherein ecological equilibrium is enhanced through
the judicious use of fungi for the betterment of all lifeforms.

To my family

the Warriors of Hwa Rang Do


C HAPTER 1. Mushrooms, Civilization, and History
C HAPTER 2. The Role of Mushrooms in Nature
The Mycorrhizal Gourmet Mushrooms: Matsutake, Boletus, Chanterelles, and Truffles
Parasitic Mushrooms: Blights of the Forest
Saprophytic Mushrooms: The Decomposers
The Global Environmental Shift and the Loss of Species Diversity
Catastrophia: Nature as a Substrate Supplier
Mushrooms and Toxic Wastes
Mushroom Mycelium and Mycofiltration
C HAPTER 3. Selecting a Candidate for Cultivation
C HAPTER 4. Natural Culture: Creating Mycological Landscapes
Methods of Mushroom Culture
Spore-Mass Inoculation
Transplantation: Mining Mycelium from Wild Patches
Inoculating Outdoor Substrates with Pure Cultured Spawn
When to Inoculate an Outdoor Mushroom Patch
Site Location of a Mushroom Patch
Stumps as Platforms for Growing Mushrooms
Log Culture
C HAPTER 5. Permaculture with a Mycological Twist: The Stametsian Model for a
Synergistic Mycosphere
C HAPTER 6. Materials for Formulating a Fruiting Substrate
Raw Materials
Suitable Wood Types: Candidate Tree Species
Cereal Straws
Paper Products: Newspaper, Cardboard, Books
Corncobs and Cornstalks
Coffee and Banana Plants
Sugarcane Bagasse
Seed Hulls
Soybean Roughage (Okara)

Structure of the Habitat
C HAPTER 7. Biological Efficiency: An Expression of Yield
C HAPTER 8. Homemade vs. Commercial Spawn
C HAPTER 9. The Mushroom Life Cycle
C HAPTER 10. The Six Vectors of Contamination
C HAPTER 11. Mind and Methods for Mushroom Culture
Overview of Techniques for Cultivating Mushrooms
C HAPTER 12. Culturing Mushroom Mycelium on Agar Media
Preparing Nutrified Agar Media
Pouring Agar Media
Starting a Mushroom Strain by Cloning
Cloning Wild vs. Cultivated Mushrooms
How to Collect Spores
Germinating Spores
Purifying a Culture
C HAPTER 13. The Stock Culture Library: A Genetic Bank of Mushroom Strains
Preserving the Culture Library
The Stamets “P” Value System
Iconic Types of Mushroom Mycelium
The Event of Volunteer Primordia on Nutrified Agar Media
C HAPTER 14. Evaluating a Mushroom Strain
Features for Evaluating and Selecting a Mushroom Strain
C HAPTER 15. Generating Grain Spawn
Formulas for Creating Grain Spawn
First-Generation Grain Spawn Masters
Second-and Third-Generation Grain Spawn
Autoclavable Spawn Bags
Liquid-inoculation Techniques
Spore-Mass Inoculation
Liquid-inoculation Techniques: Mycelial Fragmentation and Fermentation
Pelletized (Granular) Spawn
Matching the Spawn with the Substrate: Critical Choices on the Mycelial Path
Spawn Storage
C HAPTER 16. Creating Sawdust Spawn
C HAPTER 17. Growing Mushrooms on Enriched Sawdust
The Supplemented Sawdust “Fruiting” Formula: Creating the Production Block

Testing for Moisture Content
Choosing a Sterilizer, aka Retort or Autoclave
Sterilization of Supplemented Substrates
Unloading the Autoclave
Atmospheric Steam Sterilization of Sawdust Substrates
Inoculation of Supplemented Sawdust: Creating the Production Block
Incubation of the Production Blocks
Achieving Full Colonization on Supplemented Sawdust
Handling the Blocks Post-Full Colonization
C HAPTER 18. Cultivating Gourmet Mushrooms on Agricultural Waste Products
Alternative Fruiting Formulas
Heat-Treating the Bulk Substrate
The Hot Water Bath Method: Submerged Pasteurization
The “Phase II” Chamber: Steam Pasteurization
Alternative Methods for Rendering Straw and Other Bulk Materials for Cultivation
The Hydrated Lime Bath
The Bleach Bath
The Hydrogen Peroxide Technique
The High-Pressure Extrusion Method
The Detergent Bath
Yeast Fermentation
C HAPTER 19. Cropping Containers
Tray Culture
Vertical Wall Culture
Slanted Wall or A-Frame Culture
Bag Culture
Column Culture
Bottle Culture
C HAPTER 20. Casing: A Topsoil Promoting Mushroom Formation
C HAPTER 21. Growth Parameters for Gourmet and Medicinal Mushroom Species
Spawn Run: Colonizing the Substrate
Primordia Formation: The Initiation Strategy
Fruitbody (Mushroom) Development
The Gilled Mushrooms
The Himematsutake Mushroom Agaricus blazei
The Portobello Mushroom Agaricus brunnescens
The Black Poplar Mushroom Agrocybe aegerita
The Shaggy Mane Coprinus comatus
The Enoki Mushroom Flammulina velutipes
The Clustered Wood-lovers
The Brown-Gilled Woodlover Hypholoma capnoides
Kuritake (The Chestnut Mushroom) Hypholoma sublateritium

The Beech Mushrooms
Buna-Shimeji Hypsizygus tessulatus
Shirotamogitake Hypsizygus ulmarius
The Shiitake Mushroom Lentinula edodes
The Nameko Mushroom Pholiota nameko
The Oyster Mushrooms
Golden Oyster Mushroom Pleurotus citrinopileatus
The Abalone Mushroom Pleurotus cystidiosus
The Pink Oyster Mushroom Pleurotus djamor
The King Oyster Mushroom Pleurotus eryngii
The Tarragon Oyster Mushroom Pleurotus euosmus
The Tree Oyster Mushroom Pleurotus ostreatus
The Phoenix or Indian Oyster Mushroom Pleurotus pulmonarius, “P. sajor-caju”
The King Tuber Oyster Mushroom Pleurotus tuberregium
The Caramel Capped Psilocybes Psilocybe cyanescens complex
The King Stropharia Mushroom Stropharia rugosoannulata
The Paddy Straw Mushroom Volvariella volvacea
The Polypore Mushrooms
Reishi or Ling Chi Ganoderma lucidum
Maitake or Hen-of-the-Woods Grifola frondosa
Zhu Ling or the Umbrella Polypore Polyporus umbellatus
Turkey Tail or Yun Zhi Trametes versicolor
The Lion’s Mane Hericium erinaceus
The Wood Ears Auricularia polytricha
The Jelly Mushrooms
White Jelly Mushroom Tremella fuciformis
The Cauliflower Mushrooms
The Cauliflower Mushroom Sparassis crispa
The Morels (Land-Fish Mushrooms)
The Morel Life Cycle
The Development of Indoor Morel Cultivation
The Black Morels Morchella angusticeps and Allies
C HAPTER 22. Maximizing the Substrate’s Potential through Species Sequencing
C HAPTER 23. Harvesting, Storing, and Packaging Mushrooms for Market
Harvesting the Crop
Packaging and Storing the Crop for Market
Drying Mushrooms
Marketing the Product
C HAPTER 24. Mushroom Recipes: Enjoying the Fruits of Your Labors
C HAPTER 25. Cultivation Problems and Their Solutions: A Troubleshooting Guide
Agar Culture
Grain Culture
Straw Culture

Supplemented Sawdust Culture
Pre-harvest Period
Harvest Stage
APPENDIX 1. Description of Environments for a Mushroom Farm
The Laboratory Complex
The Growing Room Complex

APPENDIX 2. Designing and Building a Spawn Laboratory

Design Criteria for a Spawn Laboratory
Good Clean Room Habits: Helpful Suggestions for Minimizing Contamination in the

APPENDIX 3. The Growing Room: An Environment for Mushroom Formation and Growth
Design Criteria for the Growing Rooms
Managing the Growing Rooms: Good Habits for the Personnel

APPENDIX 4. Resource Directory

Recommended Mushroom Field Guides
Mushroom Book Suppliers

Annual Mushroom Festivals and Events
Mushroom Cultivation Seminars and Training Centers
Mushroom Study Tours and Adventures
International Mushroom Associations

North American Mushroom Societies and Associations
Mushroom Growers Associations
Sources for Mushroom Cultures
Sources for Mushroom Spawn

Sources for Marketing Information
Mushroom Newsletters and Journals

Mushroom Museums
Sources for Medicinal Mushroom Products
Mycological Resources on the Internet

APPENDIX 5. Analyses of Basic Materials Used in Substrate Preparation
APPENDIX 6. Data Conversion Tables

Photo and Illustration Credits

Mushrooms—fleshy fungi—are the premier recyclers on the planet.
Fungi are essential to recycling organic wastes and the efficient
return of nutrients back into the ecosystem. Not only are they
recognized for their importance within the environment, but also for
their effect on human evolution and health. Yet, to date, the
inherent biological power embodied within the mycelial network of
mushrooms largely remains a vast, untapped resource. As we begin
the twenty-first century, ecologists, foresters, bioremediators,
pharmacologists, and mushroom growers are converging at a new
frontier of knowledge, wherein enormous biodynamic forces are at
Only recently have we learned enough about the cultivation of
mushrooms to tap into their inherent biological power. Working
with mushroom mycelium en masse will empower every country,
farm, recycling center, and individual with direct economic,
ecological, and medical benefits. Through the genius of evolution,
the Earth has selected fungal networks as a governing force
managing ecosystems. This sentient network responds quickly to
catastrophia. I believe the mycelium is Earth’s natural Internet, a
neural network of communicating cells. All landmasses are crisscrossed with interspersing mosaics of mycelial colonies. With more
than a mile of cells in a cubic inch of soil, the fungi are moving
steadily, although silently all around us. This vast mass of cells, in
the hundreds of billions of tons, represents a collective intelligence,
like a computer honed to improve itself. Only now are scientists
discovering that it is the microbial community upon which all higher
life forms are dependent. And only now do we know how to join in
alliance with them to improve life. As we begin a new century,
myco-technology is a perfect example of the equation of good
environmentalism, good health, and good business.

This book strives to create new models for the future use of higher
fungi in the environment. As woodland habitats, especially old
growth forests, are lost to development, mushroom diversity also
declines. Wilderness habitats still offer vast genetic resources for
new strains. The temperate forests of North America, particularly
the mycologically rich Pacific Northwest, may well be viewed in the
twenty-first century as pharmaceutical companies viewed the
Amazon Basin earlier in the twentieth century. Hence, mushroom
cultivators should preserve this gene pool now for its incalculable,
future value. The importance of many mushroom species may not be
recognized for decades to come.
In many ways, this book is an offspring of the marriage of many
cultures arising from the worldwide use of mushrooms as food, as
religious sacraments in Mesoamerica, and as medicine in Asia. We
now benefit from the collective experience of lifetimes of mushroom
cultivation. As cultivators we must continue to share, explore, and
expand the horizons of the human/fungal relationship. In the future,
humans and mushrooms must bond in an evolutionary partnership.
By empowering legions of individuals with the skills of tissue culture
and mycelial management, future generations will be able to better
manage our resources and improve life on this planet.
Now that the medical community widely recognizes the healthstimulating properties of mushrooms, a combined market for
gourmet and medicinal foods is rapidly emerging. People with
compromised immune systems would be wise to create their own
medicinal mushroom gardens. I envision the establishment of a
community-based, resource-driven industry, utilizing recyclable
materials in a fashion that strengthens ecological equilibrium and
human health. As recycling centers flourish, their by-products
include streams of organic waste, which cultivators can divert into
mushroom production.
I foresee a network of environmentally sensitive and imaginative
individuals presiding over this new industry, which has previously
been controlled by a few mega-businesses. The decentralization
began with The Mushroom Cultivator in 1983, and continues with

Growing Gourmet and Medicinal Mushrooms. Join me in the next
phase of this continuing revolution.

I first acknowledge the Mushrooms who have been my greatest
teachers. They are the Body Intellect, the Neural Network of this
My family has been extremely patient and forgiving during this
multiyear project. Azureus and LaDena have tolerated my insistent
need for their modeling talents and have helped on many mushroom
projects. Dusty Wu Yao is credited for her research skills, support,
humor, and love.
My parents have taught me many things. My father championed
education and science and impressed upon me that a laboratory is a
natural asset to every home. My mother taught me patience and
kindness, and that precognition is a natural part of the human
experience. My brother John first piqued my interest for mushrooms
upon his return from adventures in Colombia and Mexico.
Additionally, his knowledge on the scientific method of photography
has greatly helped improve my own techniques. In some mysterious
way, their combined influences set the stage for my unfolding love
of fungi.
Other people warrant acknowledgement in their assistance in the
completion of this book. Andrew Weil played a critical role in
helping build the creative milieu, the wellspring of spiritual chi from
which this manuscript flowed. Gary Lincoff was extremely helpful in
uncovering some of the more obscure references and waged
intellectual combat with admirable skill. Brother Bill Stamets is
thanked for his critical editorial remarks. Satit Thaithatgoon, my
friend from Thailand, is appreciated for his insights about mushroom
culture and life. I must thank Kit and Harley Barnhart for their
advice on photographic technique. Michael Beug deserves
acknowledgment for his unwavering support through all these years.

Erik Remmen kept me healthy and strong through the many years of
rigorous training in the ancient and noble martial art of Hwarang
Joseph Ammirati, David Arora, Julie Bennett, Alan and Arleen
Bessette, David Brigham, Janet Butz, Jonathan Caldwell, Alice Chen,
Jeff Chilton, Ken Cochran, Don Coombs, Kim and Troy Donahue,
Robert Ellingham, Gaston Guzman, Paxton Hoag, John Holliday,
Rick Hoss, Lou Hsu, Eric Iseman, Loren Israelson, Omon
Isikhuemhen, Barbara King, Mike Knoke, Alexander Krenov, Gary
Leatham, Andrew Lenzer, Mike Maki, Andrew H. Miller, Orson and
Hope Miller, Scott Moore, Tomiro Motohashi, Peter Mohs, Yoshikazu
Murai, Takashi Mizuno, Takeshi Nakazawa, Louise North, George
Osgood, Christiane Pischl, David Price, Paul Przybylowicz, Warren
Rekow, Scott Redhead, Rusty Rodriguez, Maggie Rogers, Luiz Amaro
Pachoa de Silva, Bulmaro Solano, Lillian Stamets, David Sumerlin,
Ralph Tew, Harry Thiers, Tom O’Dell, James Trappe, Solomon
Wasser, Dusty Yao, and Rytas Vilgalys all helped in their own
special ways.
The late Jim Roberts, of Lambert Spawn, gained my respect and
admiration for his devotion to helping the gourmet mushroom
industry. And, I will never forget the generosity shown to me by the
late Alexander Smith and Daniel Stuntz who were instrumental in
encouraging me to continue in the field of mycology—in spite of
those who fervently opposed me.
Companies that unselfishly contributed photographic material to
this work, and to whom I am grateful, are The BOTS Group, The
Minnesota Forest Resource Center, The Growing Company, DXN
Company, Morel Mountain, Organotech, and Ostrom’s Mushroom
Farms. I would also like to thank The Mushroom Council and the
American Mushroom Institute. The Evergreen State College
generously supported my studies with Psilocybe mushrooms and in
scanning electron microscopy.
Finally, I wish to acknowledge all those bewildered and
bemushroomed researchers who have paved the way into the future.
For your help on this odyssey through life, I will forever be in your


Mushrooms have never ceased to amaze me. The more I study them,
the more I realize how little I have known, and how much more
there is to learn. For thousands of years, fungi have evoked a host of
responses from people—from fear and loathing to reverent
adulation. And I am no exception.
When I was a little boy, wild mushrooms were looked upon with
foreboding. It was not as if my parents were afraid of them, but our
Irish heritage lacked a tradition of teaching children anything nice
about mushrooms. In this peculiar climate of ignorance, rains fell
and mushrooms magically sprang forth, wilted in the sun, rotted,
and vanished without a trace. Given the scare stories told about
“experts” dying after eating wild mushrooms, my family gave me
the best advice they could: Stay away from all mushrooms, except
those bought in the store. Naturally rebellious, I took this
admonition as a challenge, a call to arms, firing up an already
overactive imagination in a boy hungry for excitement.
When we were seven, my twin brother and I made a startling
mycological discovery—Puffballs! We were told that they were not
poisonous but if the spores got into our eyes, we would be instantly
blinded! This information was quickly put to good use. We would
viciously assault each other with mature puffballs, which would
burst upon impact and emit a cloud of brown spores. The battle
would continue until all the puffballs in sight had been hurled. They
provided us with hours of delight over the years. Neither one of us
ever went blind—although we both suffer from very poor eyesight.
You must realize that to a seven-year-old these free, ready-made
missiles satisfied instincts for warfare on the most primal of levels.
This is my earliest memory of mushrooms, and to this day I consider
it to be a positive emotional experience. (Although I admit a
psychiatrist might like to explore these feelings in greater detail.)

Not until I became a teenager did my hunter-gatherer instincts
resurface, when a relative returned from extensive travels in South
America. With a twinkle in his eyes, he spoke of his experiences with
the sacred Psilocybe mushrooms. I immediately set out to find these
species, not in the jungles of Colombia, but in the fields and forests
of Washington State where they were rumored to grow. For the first
several years, my searches provided me with an abundance of
excellent edible species, but no Psilocybes. Nevertheless, I was
When hiking through the mountains, I encountered so many
mushrooms. Each was a mystery until I could match them with
descriptions in a field guide. I soon came to learn that a mushroom
was described as “edible,” “poisonous,” or my favorite, “unknown,”
based on the experiences of others like me, who boldly ingested
them. People are rarely neutral in their opinion about mushrooms—
either they love them or they hate them. I took delight in striking
fear into the hearts of the latter group, whose illogical distrust of
fungi provoked my overactive imagination.
When I enrolled in the Evergreen State College in 1975, my skills
at mushroom identification earned the support of a professor with
similar interests. My initial interest was taxonomy, and I soon
focused on fungal microscopy. The scanning electron microscope
revealed new worlds, dimensional landscapes I never dreamed
possible. As my interest grew, the need for fresh material year-round
became essential. Naturally, these needs were aptly met by learning
cultivation techniques, first in petri dishes, then on grain, and
eventually on a wide variety of materials. In the quest for fresh
specimens, I had embarked upon an irrevocable path that would
steer my life on its current odyssey.

Paul Stamets in the virgin rainforest of Washington State, in route to collect new
strains of wild mushrooms.

Humanity’s use of mushrooms extends back to Paleolithic times. Few
mushrooms have been in affecting the course of human evolution.
They have played pivotal roles in ancient Greece, India, and
Mesoamerica. True to their beguiling nature, fungi have always
elicited deep emotional responses: from adulation by those who
understand them to outright fear by those who do not.
The historical record reveals that mushrooms have been used for
less than benign purposes. Claudius II and Pope Clement VII were
both killed by enemies who poisoned them with deadly Amanitas.
Buddha died, according to legend, from a mushroom that grew
underground. Buddha was given the mushroom by a peasant who
believed it to be a delicacy. In ancient verse, that mushroom was
linked to the phrase “pig’s foot” but has never been identified.
(Although Truffles grow underground, and pigs are used to find
them, no deadly poisonous species are known.)
The oldest archaeological record of probable mushroom use is a
Tassili image from a cave dating back 5000 years B.C. (Lhote, 1987).
The artist’s intent is clear. Mushrooms with electrified auras are
depicted outlining a bee-masked dancing shaman. The spiritual
interpretation of this image transcends time and is obvious. No
wonder the word “bemushroomed” has evolved to reflect the devout
mushroom lover’s state of mind.
In the fall of 1991, hikers in the Italian Alps came across the wellpreserved remains of a man who died over 5,300 years ago,
approximately 1,700 years later than the Tassili cave artist. Dubbed
the “Iceman” or “Oetzi” by the news media, he was well-equipped
with a knapsack, flint axe, a string of dried Birch Polypores

(Piptoporus betulinus), a tinder fungus (Fomes fomentarius), and
another as-yet-unidentified mushroom that may have had magicospiritual significance (Peintner et al. 1998). Polypores can be used
as spunk for starting fires and medicine for treating wounds.
Further, a rich tea with immuno-enhancing and antibacterial
properties can be prepared by boiling these mushrooms. Equipped
for traversing the high alpine wilderness, this intrepid adventurer
had discovered the value of the noble polypores. Even today, this
knowledge can be life-saving for anyone astray in the wilderness.

2000+ year-old Mesoamerican mushroom stone.

Fear of mushroom poisoning pervades every culture, sometimes
reaching phobic extremes. The term mycophobic describes those

individuals and cultures who look upon fungi with fear and loathing.
The English and Irish epitomize mycophobic cultures. In contrast,
mycophilic societies can be found throughout Asia and Eastern
Europe, especially among Polish, Russian, and Italian peoples. These
societies have enjoyed a long history of mushroom use, with as
many as a hundred common names to describe the mushroom
varieties they love.
An investment banker named R. Gordon Wasson intensively
studied the use of mushrooms by diverse cultures. His studies
concentrated on the use of mushrooms by Mesoamerican, Russian,
English, and Indian cultures. With the French mycologist Dr. Roger
Heim, Wasson published research on Psilocybe mushrooms in
Mesoamerica, and on Amanita mushrooms in Eurasia/Siberia.
Wasson’s studies spanned a lifetime marked by a passionate love for
fungi. His publications include Mushrooms, Russia, and History; The
Wondrous Mushroom: Mycolatry in Mesoamerica; Maria Sabina and Her
Mazatec Mushroom Velada; and Persephone’s Quest: Entheogens and
the Origins of Religion. More than any individual of the twentieth
century, Wasson kindled interest in ethnomycology to its present
state of intense study. Wasson died on Christmas Day in 1986.
One of Wasson’s most provocative findings can be found in Soma:
Divine Mushroom of Immortality (1976) where he postulated that the
mysterious Soma in Vedic literature, a red fruit leading to
spontaneous enlightenment for those who ingested it, was actually a
mushroom. The Vedic symbolism carefully disguised its true identity:
Amanita muscaria, the hallucinogenic Fly Agaric. Many cultures
portray Amanita muscaria as the archetypal mushroom, invoking
both fear and admiration. Although some Vedic scholars disagree
with his interpretation, Wasson’s exhaustive research still stands
(Brough, 1971 and Wasson, 1972).

Meso-American mushroom stones, circa 300 B.C., from the Pacific slope of

Aristotle, Plato, Homer, and Sophocles all participated in religious
ceremonies at Eleusis where an unusual temple honored Demeter,
the Goddess of Earth. For over two millennia, thousands of pilgrims
journeyed fourteen miles from Athens to Eleusis, paying the

equivalent of a month’s wage for the privilege of attending the
annual ceremony. The pilgrims were ritually harassed on their
journey to the temple, apparently in good humor.
Upon arriving at the temple, they gathered in the initiation hall, a
great telestrion. Inside the temple, pilgrims sat in rows that
descended step-wise to a hidden, central chamber from which a
fungal concoction was served. An odd feature was an array of
columns, beyond any apparent structural need, whose designed
purpose escapes archaeologists. The pilgrims spent the night
together and reportedly came away forever changed. In this
pavilion crowded with pillars, ceremonies occurred, known by
historians as the Eleusinian Mysteries. No revelation of the
ceremony’s secrets could be mentioned under the punishment of
imprisonment or death. These ceremonies continued until repressed
in the early centuries of the Christian era.
In 1977, at a mushroom conference on the Olympic Peninsula, R.
Gordon Wasson, Albert Hofmann, and Carl Ruck first postulated that
the Eleusinian Mysteries centered on the use of psychoactive fungi.
Their papers were later published in a book entitled The Road to
Eleusis: Unveiling the Secret of the Mysteries (1978). That Aristotle and
other founders of Western philosophy undertook such intellectual
adventures, and that this secret ceremony persisted for nearly 2,000
years, underscores the profound impact that fungal rites have had
on the evolution of Western consciousness.

Pre-classic Mayan mushroom stone from Kaminaljuyu Highlands of Guatemala, circa
500 B.C.

Mushrooms can be classified into three basic ecological groups:
mycorrhizal, parasitic, and saprophytic. Although this book centers on
the cultivation of the gourmet and medicinal saprophytic species,
other mushrooms are also discussed.

The Mycorrhizal Gourmet Mushrooms:
Matsutake, Boletus, Chanterelles, and Truffles
Mycorrhizal mushrooms form a mutually dependent, beneficial
relationship with the roots of host plants, ranging from trees to
grasses. “Myco” means mushrooms, while “rhizal” means roots. The
collection of filament of cells that grow into the mushroom body is
called the mycelium. The mycelia of these mycorrhizal mushrooms
can form an exterior sheath covering the roots of plants and are
called ectomycorrhizal. When they invade the interior root cells of
host plants they are called endomycorrhizal. In either case, both
organisms benefit from this association. Plant growth is accelerated.
The resident mushroom mycelium increases the plant’s absorption of
nutrients, nitrogenous compounds, and essential elements
(phosphorus, copper, and zinc). By growing beyond the immediate
root zone, the mycelium channels and concentrates nutrients from
afar. Plants with mycorrhizal fungal partners can also resist diseases
far better than those without.
Most ecologists now recognize that a forest’s health is directly
related to the presence, abundance, and variety of mycorrhizal
associations. The mycelial component of topsoil within a typical

Douglas fir forest in the Pacific Northwest approaches 10% of the
total biomass. Even this estimate may be low, not taking into
account the mass of the endomycorrhizae and the many yeast-like
fungi that thrive in the topsoil.
The nuances of climate, soil chemistry, and predominant
microflora play determinate roles in the cultivation of mycorrhizal
mushrooms in natural settings. Species native to a region are likely
to adapt much more readily to designed habitats than exotic species.
I am much more inclined to spend time attempting the cultivation of
native mycorrhizal species than to import exotic candidates from
afar. Here is a relevant example.

A Truffle market in France.

Truffle orchards are well established in France, Spain, and Italy,
with the renowned Perigold Black Truffle, Tuber melanosporum,
fetching up to $500 per pound. Only in the past thirty years has
tissue culture of Truffle mycelia become widely practiced, allowing
the development of planted Truffle orchards. Landowners seeking

an economic return without resorting to cutting trees are naturally
attracted to this prospective investment. The idea is enticing. Think
of having an orchard of oaks or filberts, yielding pounds of Truffles
per year for decades at several hundred dollars a pound! Several
companies in this country have, in the past twenty years, marketed
Truffle-inoculated trees for commercial use. Calcareous soils (i.e.,
high in calcium) in Texas, Washington, and Oregon have been
suggested as ideal sites. Tens of thousands of dollars have been
exhausted in this endeavor. Only two would-be Truffle orchards have
had any success thus far, with only a small percentage of trees
producing. This discouraging state of affairs should be fair warning
to investors seeking profitable enterprises in the arena of Truffle
cultivation. Suffice it to say that the only ones to have made money
in the Truffle tree industry are those who have resold “inoculated”
seedlings to other would-be trufflateurs.
A group of Oregon trufflateurs has been growing the Oregon
White Truffle, Tuber gibbosum. Douglas fir seedlings are inoculated
with mycelium from this native species and planted in plots similar
to Christmas tree farms. Several years pass before the first harvests
begin. However, since Oregon White Truffles were naturally
occurring nearby, whether or not the inoculation process caused the
truffles to form is unclear.
In Sweden, Eric Danell (1994; 1997), who is the first to grow
Chantarelles (Cantharellus cibarius) with a potted pine tree in a
greenhouse, is continuing an ambitious project of cultivating
mycorrhizal mushrooms using a community of microorganisms as
allies. (See photo here.) At the Invermay Agricultural Center in New
Zealand, scientists have succeeded in inoculating pines with
Matsutake (Tricholoma magnivelare) mycelia in the hope that
mushrooms will appear a decade later. In New Zealand, mycorrhizal
inoculations are more successful because of the extremely limited
number of natural mycorrhizal candidates, in contrast to the
hundreds seen in the forestlands of North America. These pilot
projects hold great promise for replenishing the fungal genome of
threatened mycorrhizal mushrooms in endangered ecosystems.

Mycorrhizal mushrooms in Europe have suffered a radical decline
in years of late. The combined effects of acid rain and other
industrial pollutants, even the disaster at Chernobyl, have been
suggested to explain the sudden decline of both the quantity and
diversity of wild mycorrhizal mushrooms. Most mycologists believe
the sudden availability of deadwood is responsible for the
comparative increase in the numbers of saprophytic mushrooms. The
decline in Europe portends, in a worst case scenario, a total
ecological collapse of the mycorrhizal community, followed by a
widespread die-back of the forests. In the past ten years, the
diversity of mycorrhizal mushrooms in Europe has fallen by more
than 50%! Some species, such as the Chanterelle, have all but
disappeared from regions in the Netherlands where it was abundant
only twenty years ago (see Arnolds, 1992; Leck, 1991; Lizon 1993,
1995). Many biologists view these mushrooms as indicator species,
the first domino to fall in a series leading to the failure of the
forest’s life-support systems.
One method for inoculating mycorrhizae calls for the planting of
young seedlings near the root zones of proven Truffle trees. The new
seedlings acclimate and become “infected” with the mycorrhizae of a
neighboring, parent tree. In this fashion, a second generation of
trees carrying the mycorrhizal fungus is generated. After a few
years, the new trees are dug up and replanted into new
environments. This method has had the longest tradition of success
in Europe.

Scanning electron micrograph of an emerging root tip being mycorrhized by
mushroom mycelia.

Scanning electron micrograph of mycelia encasing the root of a tree, known as

Another approach, modestly successful, is to dip exposed roots of
seedlings into water enriched with the spore-mass of a mycorrhizal
candidate. First, mushrooms are gathered from the wild and soaked
in water. Thousands of spores are washed off the gills, resulting in
an enriched broth of inoculum. A spore-mass slurry coming from
several mature mushrooms and diluted into a 5-gallon bucket can
inoculate a hundred or more seedlings. The concept is wonderfully
simple. Unfortunately, success is not guaranteed.
Broadcasting spore-mass onto the root zones of likely candidates is
another venue that costs little in time and effort. Habitats should be
selected on the basis of their parallels in nature. For instance,
Chanterelles can be found in oak forests of the Midwest and in
Douglas fir forests of the West. Casting spore-mass of Chanterelles
into forests similar to those where Chanterelles proliferate is

obviously the best choice. Although the success rate is not high, the
rewards are well worth the minimum effort involved. Bear in mind
that tree roots confirmed to be mycorrhized with a gourmet
mushroom will not necessarily result in harvestable mushrooms.
Fungi and their host trees may have long associations without the
appearance of edible fruitbodies. (For more information, consult
Fox, 1983.)
On sterilized media, most mycorrhizal mushrooms grow slowly,
compared to the saprophytic mushrooms. Their long evolved
dependence on root by-products and complex soils makes media
preparation inherently more complicated. Some mycorrhizal species,
like Pisolithus tinctorius, a puffball favoring pines, grow quite readily
on sterilized media. A major industry has evolved providing foresters
with seedlings inoculated with this fungus. Mycorrhized seedlings
are healthier and grow faster than non-mycorrhized ones.
Unfortunately, the gourmet mycorrhizal mushroom species do not
fall into the readily cultured species category. The famous Matsutake
may take weeks before its mycelium fully colonizes the media on a
single petri dish! Unfortunately, this rate of growth is the rule rather
than the exception with the majority of gourmet mycorrhizal species.

The first authenticated success in the cultivation of the Chantarelle: Pinus sylvestris in
companionship with Cantharellus cibarius.

Chanterelles are one of the most popularly collected wild
mushrooms. In the Pacific Northwest of North America the
harvesting of Chanterelles has become a controversial, multi-million
dollar business. Like Matsutake, Chanterelles also form mycorrhizal
associations with trees. Additionally, they demonstrate a unique
interdependence on soil yeasts and pseudomonads. This type of
mycorrhizal relationship makes tissue culture most difficult. At least
three organisms must be cultured simultaneously: the host tree, the
mushroom, and soil microflora. A red soil yeast, Rhodotorula glutinis,
is crucial in stimulating spore germination. The Chanterelle life
cycle may have more dimensions of biological complexity.
Cultivators have yet to grow Chanterelles to the fruitbody stage
under laboratory conditions. Not only do other microorganisms play
essential roles, the timing of their introduction appears critical to

success in the mycorrhizal theater.
Senescence occurs with both saprophytic and mycorrhizal
mushroom species. Often the first sign of senescence is not the
inability of mycelia to grow vegetatively, but the loss of the
formation of the sexually reproducing organ: the mushroom.
Furthermore, the slowness from sowing the mycelium to the final
stages of harvest confounds the quick feedback all cultivators need
to refine their techniques. Thus, experiments trying to model how
Matsutakes grow may take twenty to forty years each, the age the
trees must be to support healthy, fruiting colonies of these prized
fungi. Faster methods are clearly desirable, but presently only the
natural model has shown any clue to success.
Given the huge hurdle of time for honing laboratory techniques, I
favor the “low-tech” approach of planting trees adjacent to known
producers of Chanterelles, Matsutake, Truffles, and Boletus. After
several years, the trees can be uprooted, inspected for mycorrhizae,
and replanted in new environments. The value of the contributing
forest can then be viewed, not in terms of board feet of lumber, but
in terms of its ability for creating satellite, mushroom/tree colonies.
When industrial or suburban development threatens entire forests,
and is unavoidable, future-oriented foresters may consider the
removal of the mycorrhizae as a last-ditch effort to salvage as many
mycological communities as possible by simple transplantation
techniques, although on a much grander scale.
Until laboratory techniques evolve to establish a proven track
record of successful marriages that result in harvestable crops, I
hesitate to recommend mycorrhizal mushroom cultivation as an
economic endeavor. Mycorrhizal cultivation pales in comparison to
the predictability of growing saprophytic mushrooms like Oyster and
Shiitake mushrooms. The industry simply needs the benefit of many
more years of mycological research to better decipher the complex
models of mycorrhizal mushroom cultivation.

Oyster and Honey mushrooms sharing a stump.

Parasitic Mushrooms: Blights of the Forest?
Parasitic fungi are the bane of foresters. They do immeasurable
damage to the health of resident tree species, but in the process
create new habitats for many other organisms. Although the
ecological damage caused by parasitic fungi is well understood, we
are only just learning of their importance in the forest ecosystem.
Comparatively few mushrooms are true parasites.
Parasites live off a host plant, endangering the host’s health as it
grows. Of all the parasitic mushrooms that are edible, the Honey
mushroom, Armillaria mellea, is the best known. One of these Honey
mushrooms, known as Armillaria gallica, made national headlines
when scientists reported finding in Michigan a single colony
covering 37 acres, weighing at least 220,000 pounds, with an
estimated age of 1,500 years! Washington State soon responded with
reports of a colony of Armillaria ostoyae covering 2,200 acres and at
least 2,400 years old. With the exception of the trembling Aspen
forests of Colorado, this fungus is the largest known living organism
on the planet. And, it is a marauding parasite!
On a well-traveled trail in the Snoqualmie Forest of Washington
State, hikers have been stepping upon the largest and perhaps oldest

polypore: Bridgeoporus (Oxyporus) nobilissimus, a conk that grows up
to several feet in diameter and can weigh hundreds of pounds!1 This
“parasitic” species grows primarily on old-growth Abies procera
(California red fir) or on their stumps. Less than a dozen specimens
have ever been collected. This mushroom is the first ever to be listed
on any list, private or public, as an endangered species. Known only
from the old-growth forests of the Pacific Northwest, the Noble
Polypore’s ability to produce a conk that lives for hundreds of years
distinguishes it from any other mushroom known to North America.
This fact—that it produces a fruiting body that survives for centuries
—suggests that the Noble Polypore has unique anti-rotting
properties, antibiotics, or other compounds that could be useful
medicinally. Located at the Kew Gardens in Scotland, another
ancient polypore, Rigidioporus ulmarius, might also be medically
significant, holding the Guinness Book of Records for the largest
mushroom in the world—with an estimated weight of more than 625
pounds (284 kilograms). These examples from the fungal kingdom
attract my attention in the search for candidates having potentially
new medicines. With the loss of old-growth forests, cultivator–
mycologists can play an all-important role in saving the fungal
genome, especially in old-growth forests, a potential treasure trove
of new medicines.

Intrepid amateur mycologist Richard Gaines points to parasitic fungus attacking a yew

In the past, a parasitic fungus has been looked upon as
biologically evil. This view is rapidly changing as science progresses.
Montana State University researchers have discovered a new
parasitic fungus attacking the yew tree. This new species is called
Taxomyces andreanae and is medically significant for one notable
feature: it produces minute quantities of the potent anticarcinogen
Taxol, a proven treatment for breast cancer (Stone, 1993). This new
fungus was studied and now a synthetic form of this potent drug is
available for cancer patients. Recently, a leaf fungus isolated in the
Congo has been discovered that duplicates the effect of insulin, but
is orally active. Even well known medicines from fungi harbor
surprises. A mycologist at Cornell University (Hodge et al. 1996)

recently discovered that the fungus responsible for the multibillion
dollar drug, cyclosporin, has a sexual stage in Cordyceps subsessilis, a
parasitic mushroom attacking scarab beetle larvae. Of the estimated
1,500,000 species of fungi, approximately 70,000 have been
identified (Hawksworth et al. 1995), and about 10,000 are
mushrooms. We are just beginning to discover the importance of
species hidden within this barely explored genome.
Many saprophytic fungi can be weakly parasitic in their behavior,
especially if a host tree is dying from other causes. These can be
called facultative parasites: saprophytic fungi activated by favorable
conditions to behave parasitically. Some parasitic fungi continue to
grow long after their host has died. Oyster mushrooms (Pleurotus
ostreatus) are classic saprophytes, although they are frequently
found on dying cottonwood, oak, poplar, birch, maple, and alder
trees. These appear to be operating parasitically when they are only
exploiting a rapidly evolving ecological niche.
Most of the parasitic fungi are microfungi and are barely visible to
the naked eye. In mass, they cause the formation of cankers and
shoot blights. Often their preeminence in a middle-aged forest is
symptomatic of other imbalances within the ecosystem. Acid rain,
groundwater pollution, insect damage, and loss of protective habitat
all are contributing factors unleashing parasitic fungi. After a tree
dies, from parasitic fungi or other causes, saprophytic fungi come
into play.

The cultivation of the Button mushroom in caves near Paris in 1868. Note candle used
for illumination. (Robinson, 1885)

Saprophytic Mushrooms: The Decomposers
Most of the gourmet mushrooms are saprophytic, wood-decomposing
fungi. Saprophytic fungi are the premier recyclers on the planet. The
filamentous mycelial network is designed to weave between and
through the cell walls of plants. The enzymes and acids they secrete
degrade large molecular complexes into simpler compounds. All
ecosystems depend upon fungi’s ability to decompose organic plant
matter soon after it is rendered available. The end result of their
activity is the return of carbon, hydrogen, nitrogen, and minerals
back into the ecosystem in forms usable to plants, insects, and other
organisms. As decomposers, they can be separated into three key
groups. Some mushroom species cross over from one category to
another depending upon prevailing conditions.
Primary Decomposers: These are the fungi first to capture a twig, a
blade of grass, a chip of wood, a log or stump. Primary decomposers
are typically fast-growing, sending out ropy strands of mycelium

that quickly attach to and decompose plant tissue. Most of the
decomposers degrade wood. Hence, the majority of these
saprophytes are woodland species, such as Oyster mushrooms
(Pleurotus species), Shiitake (Lentinula edodes), and King Stropharia
(Stropharia rugosoannulata). However, each species has developed
specific sets of enzymes to break down lignin-cellulose, the structural
components of most plant cells. Once the enzymes of one mushroom
species have broken down the lignin-cellulose to its fullest potential,
other saprophytes utilizing their own repertoire of enzymes can
reduce this material even further.
Secondary Decomposers: These mushrooms rely on the previous
activity of other fungi to partially break down a substrate to a state
wherein they can thrive. Secondary decomposers typically grow
from composted material. The actions of other fungi, actinomycetes,
bacteria, and yeasts all operate within compost. As plant residue is
degraded by these microorganisms, the mass, structure, and
composition of the compost is reduced, and proportionately
available nitrogen is increased. Heat, carbon dioxide, ammonia, and
other gases are emitted as by-products of the composting process.
Once these microorganisms (especially actinomycetes) have
completed their life cycles, the compost is susceptible to invasion by
a select secondary decomposer. A classic example of a secondary
decomposer is the Button Mushroom, Agaricus brunnescens, the most
commonly cultivated mushroom. Another example is Stropharia
ambigua, which invades outdoor mushroom beds after wood chips
have been first decomposed by a primary saprophyte.
Tertiary Decomposers: An amorphous group, the fungi represented
by this group are typically soil dwellers. They survive in habitats
that are years in the making from the activity of the primary and
secondary decomposers. Fungi existing in these reduced substrates
are remarkable in that the habitat appears inhospitable for most
other mushrooms. A classic example of a tertiary decomposer is
Aleuria aurantia, the Orange Peel Mushroom. This complex group of

fungi often poses unique problems to would-be cultivators. Panaeolus
subbalteatus is yet another example. Although one can grow it on
composted substrates, this mushroom has the reputation of growing
prolifically in the discarded compost from Button mushroom farms.
Other tertiary decomposers include species of Conocybe, Agrocybe,
Pluteus, and some Agaricus species.
The floor of a forest is constantly being replenished by new
organic matter. Primary, secondary, and tertiary decomposers can
all occupy the same location. In the complex environment of the
forest floor, a “habitat” can actually be described as the overlaying
of several, mixed into one. And, over time, as each habitat is being
transformed, successions of mushrooms occur. This model becomes
infinitely complex when taking into account the interrelationships of
not only the fungi to one another, but also the fungi to other
microorganisms (yeasts, bacteria, protozoa), plants, insects, and
Primary and secondary decomposers afford the most opportunities
for cultivation. To select the best species for cultivation, several
variables must be carefully matched.
Climate, available raw materials, and the mushroom strains all
must interplay for cultivation to result in success. Native species are
the best choices when you are designing outdoor mushroom
Temperature-tolerant varieties of mushrooms are more forgiving
and easier to grow than those that thrive within finite temperature
limits. In warmer climates, moisture is typically more rapidly lost,
narrowing the opportunity for mushroom growth. Obviously,
growing mushrooms outdoors in a desert climate is more difficult
than growing mushrooms in moist environments where they
naturally abound. Clearly, the site selection of the mushroom habitat
is crucial. The more exposed a habitat is to direct midday sun, the
more difficult it is for mushrooms to flourish.
Many mushrooms actually benefit from indirect sunlight,
especially in the northern latitudes. Pacific Northwest mushroom
hunters have long noted that mushrooms grow most prolifically, not

in the darkest depths of a woodlands, but in environments where
shade and “dappled” sunlight are combined. Sensitivity-to-light
studies have established that various species differ in their optimal
response to wavebands of sunlight. Nevertheless, few mushrooms
enjoy prolonged exposure to direct sunlight.

The Global Environmental Shift and the Loss
of Species Diversity
Studies in Europe show a frightening loss of species diversity in
forestlands, most evident with the mycorrhizal species. Many
mycologists fear many mushroom varieties, and even species, will
soon become extinct. As the mycorrhizal species decline in both
numbers and variety, the populations of saprophytic and parasitic
fungi initially rise as a direct result of the increased availability of
deadwood debris. However, as woodlots are burned and replanted,
the complex mosaic of the natural forest is replaced by a highly
uniform, mono-species landscape. Because the replanted trees are
nearly identical in age, the cycle of debris replenishing the forest
floor is interrupted. This new “ecosystem” cannot support the
myriad fungi, insects, small mammals, birds, mosses, and flora so
characteristic of ancestral forests. In pursuit of commercial forests,
the native ecology has been supplanted by a biologically anemic
woodlot. This woodlot landscape is barren in terms of species
With the loss of every ecological niche, the sphere of biodiversity
shrinks. At some presently unknown level, the diversity will fall
below the critical mass needed for sustaining a healthy forestland.
Once passed, the forest may not ever recover without direct and
drastic counteraction: the insertion of multiage trees of different
species, with varying canopies and undergrowth. Even with such
extraordinary action, the complexity of a replanted forest cannot
match that which has evolved for thousands of years. Little is

understood about prerequisite microflora—yeasts, bacteria, and
micro-fungi—upon which the ancient forests are dependent. As the
number of species declines, whole communities of organisms
disappear. New associations are likewise limited. If this trend
continues, I believe the future of new forests, indeed the planet, is
Apart from the impact of wood harvest, the health of biologically
diverse forests is in increasing jeopardy due to acid rain and other
airborne toxins. Eventually, the populations of all fungi—
saprophytic and mycorrhizal—suffer as the critical mass of dead
trees declines more rapidly than it is replenished. North Americans
have already experienced the results of habitat loss from the
European forests. Importation of wild picked mushrooms from
Mexico, the United States, and Canada to Europe has escalated
radically in the past twenty years. This increase in demand is not
due just to the growing popularity of eating wild mushrooms. It is a
direct reflection of the decreased availability of wild mushrooms
from regions of the world suffering from ecological shock. The
woodlands of North America are only a few decades behind the
forests of Europe and Asia.
With the loss of habitat of the mycorrhizal gourmet mushrooms,
market demands for gourmet mushrooms should shift to those that
can be cultivated. Thus, the pressure on this not-yet-renewable
resource would be alleviated. I believe the judicious use of
saprophytic fungi by homeowners as well as foresters may well
prevent widespread parasitic disease vectors. Selecting and
controlling the types of saprophytic fungi occupying these ecological
niches can benefit both forester and forestland.

Catastrophia: Nature as a Substrate Supplier
Many saprophytic fungi benefit from catastrophic events in the
forests. When hurricane-force winds rage across woodlands,
enormous masses of dead debris are generated. The older trees are

especially prone to fall. Once the higher canopy is gone, the growth
of a younger, lower canopy of trees is triggered by the suddenly
available sunlight. The continued survival of young trees is
dependent upon the quick recycling of nutrients by the saprophytic
fungi in decomposing deadwood.
Every time catastrophes occur—hurricanes, tornadoes, volcanoes,
floods, and even earthquakes—the resulting deadwood becomes a
stream of inexpensive substrate materials. In a sense, the cost of
mushroom production is underwritten by natural disasters.
Unfortunately, to date, few individuals and communities take
advantage of catastrophia as a fortuitous event for enhancing
mycelial growth. However, once the economic value of recycling
with gourmet and medicinal mushrooms is clearly understood, and
with the increasing popularity of backyard cultivation, catastrophia
can be viewed as a positive event, at least in terms of providing new
economic opportunities and positive environmental consequences
for those who are mycologically astute.

Mushrooms and Toxic Wastes
In heavily industrialized areas, the soils are typically contaminated
with a wide variety of pollutants, particularly petroleum-based
compounds, polychlorinated biphenols (PCBs), heavy metals,
pesticide-related compounds, and even radioactive wastes.
Mushrooms grown in polluted environments can absorb toxins
directly into their tissues, especially heavy metals (Bressa, 1988;
Stijve 1974, 1976, 1992). As a result, mushrooms grown in these
environments should not be eaten. Recently, a visitor to Ternobyl, a
city about 60 miles from Chernobyl, the site of the world’s worst
nuclear power plant accident, returned to the United States with a
jar of pickled mushrooms. The mushrooms were radioactive enough
to set off Geiger counter alarms as the baggage was being processed.
Unfortunately, most toxins are not so readily detected.

A number of fungi can, however, be used to detoxify contaminated
environments, in a process called “bioremediation.” The white rot
fungi (particularly Phanerochaete chrysosporium) and brown rot fungi
(notably Gloephyllum species) are the most widely used. Most of
these wood-rotters produce lignin peroxidases and cellulases, which
have unusually powerful degradative properties. These extracellular
enzymes have evolved to break down plant fiber, primarily lignincellulose, the structural component in woody plants, into simpler
forms. By happenstance, these same enzymes also reduce
recalcitrant hydrocarbons and other manufactured toxins. Given the
number of industrial pollutants that are hydrocarbon-based, fungi
are excellent candidates for toxic waste cleanup and are viewed by
scientists and government agencies with increasing interest. Current
and prospective future uses include the detoxification of PCB
pesticide/herbicide residues. They are even being explored for
ameliorating the impact of radioactive wastes by sequestering heavy
A far-reaching patent has been applied for using mycelial mats to
break down toxic wastes, particularly those that are hydrocarbon
based, including most petroleum products, pesticides, PCBs
(polychlorobiphenols), and PCPs (pentachlorophenols), and for
eliminating the flow of pathogenic bacteria into sensitive
watersheds. This revolutionary patent also describes methods for
effectively destroying nerve gas surrogates, including Sarin and VX,
as well as chemical and biological warfare components by training
the mushroom mycelium (Venter, A. J., 1999; Word et al. 1997;
Thomas et al. 1998).
Bioremediation of toxic waste sites is especially attractive because
the environment is treated in situ. The contaminated soils do not
have to be hauled away, eliminating the extraordinary expense of
handling, transportation, and storage. Since these fungi have the
ability to reduce complex hydrocarbons into elemental compounds,
these compounds pose no threat to the environment. Indeed, these
former pollutants could even be considered “fertilizer,” helping

rather than harming the nutritional base of soils.
Dozens of bioremediation companies have formed to solve the
problem of toxic waste. Most of these companies look to the
imperfect fungi. The higher fungi should not be disqualified for
bioremediation just because they produce an edible fruitbody.
Indeed, this group may hold answers to many of the toxic waste
problems. The most vigorous rotters described in this book are the
Ganoderma and Pleurotus mushrooms. Mushrooms grown from toxic
wastes are best not eaten, as residual heavy metal toxins may be
concentrated within the mushrooms. However, one experiment using
Oyster mushrooms to degrade petroleum residues on an oil-saturated
Department of Transportation lot near Bellingham, Washington, not
only largely decomposed the oil, but the mushrooms were free of
petroleum residues when analyzed (Stamets, 1999).

Scanning electron micrograph of the mycelial network.

Mushroom Mycelium and Mycofiltration
The mycelium is a fabric of interconnected, interwoven strands of
cells. One colony can range in size from a half-dollar to many acres.
A cubic inch of soil can host up to a mile of mycelium. This organism
can be physically separated, and yet behave as one.
The exquisite lattice-like structure of the mushroom mycelium,
often referred to as the mycelial network, is perfectly designed as a
filtration membrane. Each colony extends long, complex chains of
cells that fork repeatedly in matrix-like fashion, spreading to
geographically defined borders. The mushroom mycelium, being a
voracious forager for carbon and nitrogen, secretes extracellular
enzymes that unlock organic complexes. The newly freed nutrients
are then selectively absorbed directly through the cell walls into the
mycelial network.
In the rainy season, water carries nutritional particles through this
filtration membrane, including bacteria, which often become a food
source for the mushroom mycelium. The resulting downstream
effluent is cleansed of not only carbon/nitrogen-rich compounds but
also bacteria, in some cases nematodes, and legions of other
microorganisms. The voracious Oyster mushrooms been found to be
parasitic against nematodes (Thorn and Barron, 1984; Hibbett and
Thorn, 1994). Extracellular enzymes act like an anesthetic and stun
the nematodes, thus allowing the invasion of the mycelium directly
into their immobilized bodies.
The use of mycelium as a mycofilter is currently being studied by
this author in the removal of biological contaminants from surface
water passing directly into sensitive watersheds. By placing sawdust
implanted with mushroom mycelium in drainage basins downstream
from farms raising livestock, the mycelium acts as a sieve, which
traps fecal bacteria and ameliorates the impact of a farm’s nitrogenrich outflow into aquatic ecosystems. This concept is incorporated
into an integrated farm model and explored in greater detail in
Chapter 5: Permaculture with a Mycological Twist.

Oyster mushrooms fruiting on diesel-contaminated soil at a test bioremediation site
near Bellingham, Washington, effectively “de-contaminating” the soil to a level where
it could be used for highway landscaping.

1. Oxyporus nobilissimus has been placed in its own genus, Bridgeoporus (Burdsall et al. 1996).

Many mushroom hunters would love to have their favorite edible
mushroom growing in their backyard. Who would not want a patch
of Matsutake, Shaggy Manes, Giant Puffballs, or the stately Prince
gracing their property? As the different seasons roll along, gourmet
mushrooms would arise in concert. Practically speaking, however,
our knowledge of mushroom cultivation is currently limited to 100
species of the 10,000 thought to exist throughout the world. Through
this book and the works of others, the number of cultivatible species
will enlarge, especially if amateurs are encouraged to boldly
experiment. Techniques for cultivating one species may be applied
for cultivating another, often by substituting an ingredient,
changing a formula, or altering the fruiting environment. Ironically,
with species never before grown, the strategy of “benign neglect”
more often leads to success than active interference with the natural
progression of events. I have been particularly adept at this
nonstrategy. Many of my early mushroom projects only produced
when I left them alone.
A list of candidates, which can be grown using current methods,
follows. Currently we do not know how to grow those species
marked by an asterisk (*). However, I believe techniques for their
cultivation will soon be perfected, given a little experimentation.
This list is by no means exhaustive, and will be much amended in
the future. Many of these mushrooms are described as good edibles
in the field guides, as listed in the Resource Directory in this book.
(See Appendix 4.)

Woodland Mushrooms
The Wood Ears

Auricularia auricula
Auricularia polytricha
The Prince
Agaricus augustus
The Almond Agaricus
Agaricus subrufescens
The Sylvan Agaricus
Agaricus sylvicola
Agaricus lilaceps*
Black Poplar Agrocybe
Agrocybe aegerita
The Clustered Woodlovers
Hypholoma capnoides
Hypholoma sublateritium
Psilocybe cyanescens and allies
Oyster-like Mushrooms
Hypsizygus ulmarius
Hypsizygus tessulatus (= H. marmoreus)
Pleurotus citrinopileatus (= P. cornucopiae var. citrinopileatus)
Pleurotus cornucopiae
Pleurotus cystidiosus (= P. abalonus, P. smithii (?))
Pleurotus djamor (=P. flabellatus, P. salmoneo-stramineus)
Pleurotus dryinus*
Pleurotus eryngii
Pleurotus euosmus
Pleurotus ostreatus
Pleurotus pulmonarius (= “sajor-caju”)
Tricholoma giganteum
The Deer Mushroom
Pluteus cervinus
Shiitake Mushroom
Lentinula edodes
Lentinula spp.

Garden Giant or King Stropharia
Stropharia rugosoannulata
Most polypore mushrooms

Grassland Mushrooms

Meadow Mushrooms
Agaricus arvensis
Agaricus blazei
Agaricus campestris
Lepiota procera
Horse Mushroom
Agaricus arvensis
The Giant Puffball
Calvatia gigantea and allies*
Smooth Lepiota
Lepiota naucina*
The Parasol Mushroom
Lepiota procera
Fairy Ring Mushroom
Marasmius oreades

Dung Inhabiting Mushrooms

The Button Mushrooms
Agaricus brunnescens
Agaricus bitorquis (= rodmanii)
The Magic Mushrooms
Psilocybe cubensis
Panaeolus cyanescens (= Copelandia cyanescens)
Panaeolus subbalteatus
Panaeolus tropicalis (Copelandia tropicalis)

Compost/Litter/Disturbed Habitat Mushrooms
Shaggy Manes

Coprinus comatus
Scaly Lepiota
Lepiota rachodes*
The Termite Mushrooms
Termitomyces spp.*
The Blewit
Lepista nuda

Termitomyces robustus is one of the best of the edible mushrooms but defies human
attempts at cultivation. So far only ants know the secret to growing this delicacy.

Gardening with gourmet and medicinal mushrooms.

Natural culture is the cultivation of mushrooms outdoors. After
mycological landscapes are constructed and inoculated, the forces of
nature take control. For these mycological landscapes to be
sustainable, a continual flow of organic debris is essential. Although
the cultivator may choose to install desired species, respect towards
nature’s selection of preferred mushrooms is the only path to
successful cultivation. Wild species in the landscape are natural
allies. The responsibility of the cultivator is to design a habitat
incorporating both wild and cultivated mushrooms, and seeking the
right fits. Yet, the complex nature of creating species mosaics is still
being understood. Only through the cumulative experiences of
mycological landscapers can the knowledge base of this new model
I also call this laissez-faire cultivation. After the mushroom patch
has been inoculated, it is left alone, subject to the whims of nature,
except for some timely watering. The mushroom habitat is
specifically designed, paying particular attention to site location,
topography, sun exposure, and the use of native woods and/or
garden by-products. Once prepared, the cultivator launches the
selected mushroom species into a constructed habitat by spawning.
In general, native mushroom species do better than exotic ones.
However, even those obstacles to growing exotic species are easily
overcome with some forethought to design, and the helpful
suggestions of an experienced cultivator.
Every day, gardeners, landscapers, rhododendron growers,
arborists, and nurseries utilize the very components needed for

growing mushrooms. Every pile of debris, whether it is tree
trimmings, sawdust, wood chips, or a mixture of these materials, will
support mushrooms. Unless selectively inoculated, debris piles
become habitats of miscellaneous “weed” mushrooms, making the
likelihood of growing a desirable mushroom remote.
When inoculating an outdoor environment with mushroom spawn,
the cultivator relinquishes much control to natural forces. There are
obvious advantages and disadvantages to natural culture. First, the
mushroom patch is controlled by volatile weather patterns. This also
means that outdoor beds have the advantage of needing minimum
maintenance. The ratio of hours spent per pound of mushrooms
grown becomes quite efficient. The key to success is creating an
environment wherein the planted mycelium naturally and
vigorously expands. A major advantage of growing outdoors
compared to growing indoors is that competitors are not
concentrated in a tight space. When cultivating mushrooms outdoors
entropy is your ally.
The rate of growth, time to fruiting, and quality of the crop
depends upon the quality of the spawn, substrate materials, and
weather conditions. Generally, when mushrooms are fruiting in the
wild, the inoculated patches also produce. Mushrooms that fruit
primarily in the summer, such as the King Stropharia (Stropharia
rugosoannulata) require frequent watering. Shaggy Manes (Coprinus
comatus) prefer the cool fall rains, thus requiring little attention. In
comparison to indoor cultivation, the outdoor crops are not as
frequent. However, the crops can be just as intense, sometimes more
so, especially when paying modest attention to the needs of the
mushroom mycelium at critical junctions in its life cycle.
While the cultivator is competing with molds indoors, wild
mushrooms are the major competitors outdoors. You may plant one
species in an environment where another species is already firmly
established. This is especially likely if you use old sawdust, chips, or
base materials. Starting with fresh materials is the simplest way to
avoid this problem. Piles of aged wood chips commonly support four
or five species of mushrooms within just a few square feet. Unless,

the cultivator uses a high rate of inoculation (25% spawn/substrate) and
uniformly clean wood chips, the concurrence of diverse mushroom
species should be expected. If, for instance, the backyard cultivator
gets mixed wood chips in the early spring from a county road
maintenance crew, and uses a dilute 5–10% inoculation rate of
sawdust spawn into the chips, the mushroom patch is likely to have
more wild species emerging along with the desired mushrooms.
In the Pacific Northwest of North America, I find a 5–10%
inoculation rate usually results in some mushrooms showing late in
the first year, the most substantial crops occurring in the second and
third years, and a dramatic drop-off in the fourth year. As the patch
ages, it is normal to see more diverse mushroom varieties cooccurring with the planted mushroom species.
I am constantly fascinated by the way nature reestablishes a
polyculture environment at the earliest opportunity. Some
mycologists believe a predetermined sequence of mycorrhizal and
saprophytic species prevails, for instance, around a Douglas fir tree,
as it matures. In complex natural habitats, the interlacing of
mycelial networks is common. Underneath a single tree, twenty or
more species may thrive. I look forward to the time when
mycotopian foresters will design whole species mosaics upon whose
foundation vast ecosystems can flourish. This book will describe
simpler precursor models for mixing and sequencing species. I hope
imaginative and skilled cultivators will further develop these
In one of my outdoor wood-chip beds, I created a “polyculture”
mushroom patch about 50 by 100 feet in size. In the spring I
acquired mixed wood chips from the county utility company—mostly
alder and Douglas fir—and inoculated three species into it. One year
after inoculation, in late April through May, Morels showed. From
June to early September, King Stropharia erupted with force,
providing our family with several hundred pounds. In late
September through much of November, an assortment of Clustered
Woodlovers (Hypholoma-like) species popped up. With noncoincident
fruiting cycles, this Zen-like polyculture approach is limited only by

your imagination.
Species succession can be accomplished indoors. Here is one
example. After Shiitake stops producing on logs or sawdust, the
substrate can be broken apart, remoistened, resterilized, and
reinoculated with another gourmet mushroom; in this case, I
recommend Oyster mushrooms. Once the Oyster mushroom life cycle
is completed, the substrate can be again sterilized, and inoculated
with the next species. Shiitake, Oyster, King Stropharia, and finally
Shaggy Manes can all be grown on the same substrate, increasingly
reducing the substrate mass, without the addition of new materials.
The majority of the substrate mass that does not evolve into gases, is
regenerated into mushrooms. The conversion of substrate mass-tomushroom mass is mind boggling. These concepts are further
developed in Chapter 22.
The following is a list of decomposer mushrooms most frequently
occurring in wood chips in the northern temperate regions of North
America. In general, these natural competitors are easy to
distinguish from the gourmet mushroom species described in this
book. Those that are mildly poisonous are labeled with*; those that
are deadly have two**. This list is by no means comprehensive.
Many other species, especially the poisonous mycorrhizal Amanita,
Hebeloma, Inocybe, and Cortinarius species are not listed here.
Mushrooms from these genera can inhabit the same plot of ground
where a cultivator may lay down wood chips, even if the host tree is
far removed.
The mushrooms in the Galerina autumnalis and Pholiotina filaris
groups are deadly poisonous. Some species in the genus Psilocybe
contain psilocybin and psilocin, compounds that often cause
uncontrolled laughter, hallucinations, and sometimes spiritual
experiences. Outdoor cultivators must hone their skills at mushroom
identification to avert the accidental ingestion of a poisonous
mushroom. Recommended mushroom field guides and mushroom
identification courses are listed in the Resource Directory in this

Some Wild Mushrooms Naturally Found in Beds of Wood
Ground Lovers
Agrocybe spp. and Pholiota spp.
The Sweaters
Clitocybe spp.*
The Inky Caps
Coprinus atramentarius*
C. comatus
C. disseminatus
C. lagopus
C. micaceus and allies
The Vomited Scrambled Egg Fungus
Fuligo cristata
The Deadly Galerinas
Galerina autumnalis and allies**
Red-Staining Lepiotas
Lepiota spp.*
The Clustered Woodlover
Hypholoma capnoides
The Green-Gilled Clustered Woodlover
Hypholoma fasciculare *
The Chestnut Mushroom
Hypholoma sublateritium
The Deadly Ringed Cone Head
Pholiotina filaris and allies **
Pholiota terrestris and allies
The Deer Mushroom
Pluteus atrocapillus (= cervinus)
Black Spored Silky Stems
Psathyrella spp.
The Caramel Capped Psilocybes

Psilocybe cyanescens and allies

Methods of Mushroom Culture
Mushrooms can be cultivated through a variety of methods. Some
techniques are exquisitely simple, and demand little or no technical
expertise. Others—involving sterile tissue culture—are much more
technically demanding. The simpler methods take little time, but
also require more patience and forgiveness on the part of the
cultivator, lest the mushrooms do not appear on your timetable. As
one progresses to the more technically demanding methods, the
probability of success is substantially increased, with mushrooms
appearing exactly on the day scheduled.
The simpler methods for mushroom cultivation, demanding little
or no technical expertise, are outlined in this chapter. They are
spore-mass inoculation, transplantation, and inoculation with pure
cultured spawn.

Collecting the spores of the delicious Lepiota rachodes, a Parasol Mushroom, on two
panes of glass, which are then folded together, creating a spore booklet.

Spore-Mass Inoculation

By far the simplest way to grow mushrooms is to broadcast spores
onto prepared substrates outdoors. First, spores of the desired
species must be collected. Spore collection techniques vary,
according to the shape, size, and type of the mushroom candidate.
For gilled mushrooms, the cap can be severed from the stem, and
laid, gills down, on top of clean typing paper, glass, or similar
surface. A glass jar or bowl is placed over the mushroom to lessen
the loss of water. After 12 hours, most mushrooms will have released
thousands of spores, falling according to the radiating symmetry of
the gills, in a symmetrically attractive outline called a sporeprint.
This method is ideal for mushroom hunters “on the go” who might
not be able to make use of the spores immediately. After the spores
have fallen, the spore print can be sealed, stored, and saved for
future use. It can even be mailed without harm.
By collecting spores of many mushrooms, one creates a species
library. Spore collections can resemble stamp or coin collections, but
are potentially more valuable. A mushroom hunter may find a
species only once in a lifetime. Under these circumstances, the
existence of a spore print may be the only resource a cultivator has
for future propagation. I prefer taking spore prints on panes of
glass, using duct tape as binding along one edge. The glass panes
are folded together, and masking tape is used to seal the three
remaining edges. This spore booklet is then registered with written
notes affixed to its face as to the name of mushroom, the date of
collection, the county and locality of the find. Spores collected in
this fashion remain viable for years, although viability decreases
over time. They should be stored in a dark, cool location, low in
humidity and free from temperature fluctuation. Techniques for
creating cultures from spores are explained further on.
For those wishing to begin a mushroom patch using fresh
specimens, a more efficient method of spore collection is
recommended. This method calls for the immersion of the mushroom
in water to create a spore-mass slurry. Choose fairly mature
mushrooms and submerge them in a 5-gallon bucket of water. A
gram or two of table salt inhibits bacteria from growing while not

substantially affecting the viability of the spores. With the addition
of 50 milliliters of molasses, spores are stimulated into frenzied
germination. After 4 hours of soaking, remove the mushrooms from
the bucket. Most mushrooms will have released tens of thousands of
spores. Allow the broth to sit for 24 to 48 hours at a temperature
above 50°F (10°C) but under 80°F (26.7°C). In most cases, spores
begin to germinate in minutes to hours, aggressively in search of
new mates and nutrients. This slurry can be expanded by a factor of
10 in 48 hours. I have often dreamed, being the mad scientist that I
am, of using spore-mass slurries of Morels and other species to
aerially “bomb” large expanses of forestlands. This idea, as crazy as
it may initially sound, warrants serious investigation.
During this stage of frenzied spore germination, the mushroom
patch habitat should be designed and constructed. Each species has
unique requirements for substrate components for fruiting. However,
mycelia of most species will run through a variety of lignin-cellulosic
wastes. Only at the stage when fruitbody production is sought does
the precise formulation of the substrate become crucial.
Oyster (Pleurotus ostreatus; P. eryngii, and allies), King Stropharia
(Stropharia rugosoannulata), and Shaggy Mane (Coprinus comatus)
mushrooms thrive in a broad range of substrate formulations. Other
mushrooms such as Morels (Morchella angusticeps and esculenta) are
more restrictive in their requirements. Since there are several tracks
that one can pursue to create suitable habitats, refer to Chapter 21
for more information.

Transplantation: Mining Mycelium from Wild Patches
Transplantation is the moving of mycelium from natural patches to
new habitats. Most wild mushroom patches have a vast mycelial
network emanating from beneath each mushroom. Not only can one
harvest the mushroom, but portions of the mycelial network can be
gathered and transferred to a new location. This method ensures the
quick establishment of a new colony without having to germinate
spores or buy commercial spawn.

When transplanting mycelium, I use a paper sack or a cardboard
box. Once mycelium is disturbed, it quickly dries out unless measures
are taken to prevent dehydration. After it is removed from its
original habitat, the mycelium will remain viable for days or weeks,
as long as it is kept moist in a cool, dark place.
Gathering the wild mycelium of mycorrhizal mushrooms could
endanger the parent colony. Be sure you cover the divot with wood
debris and press tightly back into place. In my opinion, mycorrhizal
species should not be transplanted unless the parent colony is
imminently threatened with loss of habitat—such as logging,
construction, etc. Digging up mycelium from the root zone of a
healthy forest can jeopardize the symbiotic relationship between the
mushroom and its host tree. Exposed mycelium and roots become
vulnerable to disease, insect invasion, and dehydration.
Furthermore, transplantation of mycorrhizal species has a lower
success rate than the transplantation of saprophytic mushrooms.
If done properly, transplanting the mycelium of saprophytic
mushrooms is not threatening to naturally occurring mushroom
colonies. Some of the best sites for finding mycelium for
transplantation are sawdust piles. Mycelial networks running
through sawdust piles tend to be vast and relatively clean of
competing fungi. Fans of mycelium are more often found along the
periphery of sawdust piles than within their depths. When sawdust
piles are a foot deep or more, the microclimate is better suited for
molds and thermophilic fungi. These mold fungi benefit from the
high carbon dioxide and heat generated from natural composting. At
depths of 2–6 inches, mushroom mycelia run vigorously. It is from
these areas that mushroom mycelium should be collected for
transplantation to new locations. One, in effect, engages in a form
of mycelial mining by encouraging the growth and the harvesting of
mycelium from such environments. Ideal locations for finding such
colonies are sawmills, nurseries, composting sites, recycling centers,
rose and rhododendron gardens, and soil mixing companies.

Establishing an outdoor mushroom bed in a garden.

Sprinkling spawn on top of mulch layer.

Adding more moist mulch over the spawn layer.

Cross section of garden bed showing mycelium and mushroom growth.

Inoculating Outdoor Substrates with Pure Cultured
In the early history of mushroom cultivation, mycelium was
collected from the wild and transplanted into new substrates with
varying results. Soon compost spawn (for the Button Mushroom,
Agaricus brunnescens) evolved with greater success. In 1933, spawn
technology was revolutionized by Sinden’s discovery of grain as a
spawn carrier medium. Likewise, Stoller (1962) significantly
contributed to the technology of mushroom cultivation through a
series of practical advances in using plastic bags, collars, and

filters.1 The Mushroom Cultivator (Stamets and Chilton, 1983)
explained the process of producing tissue culture for spawn
generation, empowering far more cultivators than ever before.
Legions of creative individuals embarked on the path of exotic
mushroom production. Today, thousands of cultivators are
contributing to an ever-expanding body of knowledge, and setting
the stage for the cultivation of many gourmet and medicinal fungi of
the future.
The advantage of using commercial spawn is in acquiring
mycelium of higher purity than can be harvested from nature.
Commercial spawn can be bought in two forms: grain or wood
(sawdust or plugs). For the inoculation of outdoor, unpasteurized
substrates, wood-based spawn is far better than grain spawn. When
grain spawn is introduced to an outdoor bed, insects, birds, and
slugs quickly seek out the nutritious kernels for food. Sawdust spawn
has the added advantage of having more particles or inoculation
points per pound than does grain. With more points of inoculation,
colonization is accelerated. The distances between mycelial
fragments are lessened, making the time to contact less than that
with grain spawn. Thus the window of vulnerability is closed to
many of the diseases that eagerly await intrusion.
Before spawn is used, the receiving habitat is moistened to near
saturation. The spawn is then mixed thoroughly through the new
habitat with your fingers or a rake. Once inoculated, the new bed is
again watered. The bed can be covered with cardboard, shade cloth,
scrap wood, or a similar material to protect the mycelium from sun
exposure and dehydration. After inoculation, the bed is ignored,
save for an occasional inspection and watering once a week, and
then only when deemed necessary.
Certain limitations prevail in the expansion of mycelium and its
ability for colonizing new substrates. The intensity or rate of
inoculation is extremely important. If the spawn is too dispersed
into the substrate, the points of inoculation will not be close enough
to result in the rapid reestablishment of one large contiguous
mycelial mat. My own experiences show that success is seen with an

inoculation rate of 5–50%, with an ideal of 20%. In other words, if
you gather a 5-gallon bucket of naturally occurring mycelium, 20
gallons of prepared substrate can be inoculated with a high
probability of success. Although this inoculation rate may seem high,
rapid colonization is assured. More skilled cultivators, whose
methods have been refined through experience, often use a less
intensive inoculation rate of 10%. Inoculation rates of 5% or less
often result in “island” colonies of the implanted species
interspersed among naturally occurring wild colonies.
At a 20% inoculation rate, colonization can be complete in as
short as 1 week and as long as 8 weeks. After a new mycelial mat
has been fully established, the cultivator has the option of further
expanding the colony by a factor of 5, or triggering the patch into
fruiting. This usually means providing shade and frequent watering.
Should prevailing weather conditions not be conducive to fruiting
and yet are above freezing, then the patch can be further expanded.
Should the cultivator not expect that further expansion would result
in full colonization by the onset of winter, then no new raw material
should be added, and mushrooms should be encouraged to form. The
widely cast mycelial mat, more often than not, acts as a single
organism. At the time when mushrooms are forming, colonization of
new organic debris declines or abates entirely. The energy of the
mycelium is now channeled to fruitbody formation and

Healthy Stropharia rugosoannulata mycelium tenaciously gripping alder chips and
sawdust. Note rhizomorphs.

From the same patch, a year later, the wood chips have decomposed into a rich soilloam.

The mycelium of saprophytic mushrooms must move to remain
healthy. When the mycelium reaches the borders of a geographically
or nutritionally defined habitat, a resting period ensues. If not soon
triggered into fruiting, over-incubation is likely, with the danger of
“dieback.” Only very cold temperatures will keep the patch viable
for a prolonged period. Typically, dieback is seen as the drastic
decline in vigor of the mycelium. Once the window of opportunity
has passed for fruiting, the mushroom patch might be salvaged by
the reintroduction of more undecomposed organic matter, or by
violent disturbance. The mycelium soon becomes a site for
contamination with secondary decomposers (weed fungi) and
predators (insects) coming into play. It is far better is to keep the
mycelium running until fruitings can be triggered at the most
opportune time. Mushroom patches are, by definition, temporary

King Stropharia lasts 3 to 4 years on a hardwood chip base. After
the second year more material should be added. However, if the
health of the patch has declined and new material is mixed in, then
the mushroom patch may not recover to its original state of vigor.
Mycelium that is healthy tends to be tenacious, holding the substrate
particles together. This is especially true with Stropharia spp. and
Oyster mushrooms. (Hericium erinaceus and Morchella spp. are
exceptions.) Over-incubation results in a weakened mycelial
network whereby the mycelium is incapable of holding various
substrate particles together. As mycelial integrity declines, other
decomposers are activated. Often, when mixing in new material at
this stage, weed fungi proliferate to the decided disadvantage of the
selected gourmet species. To the eye, the colony no longer looks like
a continuous sheet of mycelium, but becomes spotty in its growth
pattern. Soon islands of mycelia become smaller and smaller as they
retreat, eventually disappearing altogether. The only recourse is to
begin anew, scraping away the now-darkened wood/soil, and
replacing it with a new layer of wood chips and/or other organic

When to Inoculate an Outdoor Mushroom Patch
Outdoor beds can be inoculated in early spring to early fall. The key
to a mushroom bed is that the mycelium has sufficient time to
establish a substantial mycelial mat before the onset of inclement
weather conditions. Springtime is generally the best time to
inoculate, especially for creating large mushroom patches. As fall
approaches, more modestly sized beds should be established, with a
correspondingly higher rate of inoculation for faster growth. For
most saprophytic species, at least 4 weeks are required to form the
mycelial network with the critical mass necessary to survive the
Most woodland species survive wintering temperatures.
Temperate woodland mushrooms have evolved protective
mechanisms within their cellular network that allow them to tolerate

cold temperature extremes. Surface frosts usually do not harm the
terrestrially bound mushroom mycelium. As the mycelium
decomposes organic matter, heat is released, which benefits
subsurface mycelium. Mycelial colonization essentially stops when
outdoor temperatures fall below freezing.

Site Location of a Mushroom Patch
A suitable site for a mushroom patch is easy to choose. The best clue
is to simply take note of where you have seen mushrooms growing
during the rainy season. Or just observe where water traverses after
a heavy rain. A gentle slope, bordered by shrubs and other shadegiving plants, is usually ideal. Since saprophytic mushrooms are
noncompetitive to neighboring plants, they pose no danger to them.
In fact, plants near a mushroom bed often thrive—the result of the
increased moisture retention and the release of nutrients into the
root zone.
An ideal location for growing mushrooms is in a vegetable,
flower, and/or rhododendron garden. Gardens are favored by
plentiful watering, and the shade provided by potato, zucchini, and
similar broadleaf vegetable plants tend to keep humidity high near
the ground. Many gardeners bring in sawdust and wood chips to
make pathways between the rows of vegetables. By increasing the
breadth of these pathways, or by creating small cul-de-sacs in the
midst of the garden, a mushroom bed can be ideally located and
Other suitable locations are exposed north sides of buildings, and
against rock, brick, or cement walls. Walls are usually heat sinks,
causing condensation, which provides moisture to the mushroom site
as temperature fluctuates from day to night. Protected from winds,
these locations have limited loss of water due to evaporation.
Mushrooms love moisture. By locating a mushroom bed where
moisture naturally collects, colonization is rapid and more complete,
and the need for additional water for fruiting is minimized. The

message here: Choose your locations with moisture foremost in
mind. Choose shady locations over sunny ones. Choose north-facing
slopes rather than south facing. Choose companion plants with
broad leaves or canopies that shade the midday sun but allow rain
to pass. The difference in results is the difference between a
bountiful success or dismal failure.

Giant Oyster mushrooms fruiting from a stump.

Stumps as Platforms for Growing Mushrooms
Stumps are especially suitable for growing gourmet mushrooms.
There are few better, or more massive platforms, than the stump.
Millions of stumps are all that remain of many forests of the world.
In most cases, stumps are seen as having little or no economic
potential. These lone tombstones of biodegradable wood fiber offer
a unique, new opportunity for the mycologically astute. With
selective logging being increasingly practiced, cultivating gourmet
and medicinal mushrooms on stumps will be the wave of the future.
The advantage of the stump is not only its sheer mass, but with
roots intact, water is continuously being drawn via capillary action.
Once mycelium has permeated through wood fiber, the stump’s
water-carrying capacity is increased, thus further supporting

mycelial growth. Candidates for stump culture must be carefully
selected and matched with the appropriate species. A stump
partially or fully shaded is obviously better than one in full sunlight.
Stumps in ravines are better candidates than those located in the
center of a clear-cut area. An uprooted stump is not as good a
candidate as a well-rooted one. The presence of mosses, lichens,
and/or ferns is a good indicator that the microclimate is conducive
to mushroom growth. However, the presence of competitor fungi
generally disqualifies a stump as a good candidate. These are some
of the many factors that determine the suitability of stumpage.
Cultivating mushrooms on stumps requires forethought. Stumps
should be inoculated before the first season of wild mushrooms. With
each mushroom season, the air becomes laden with spores, seeking
new habitats. The open face of a stump, essentially a wound, is
highly susceptible to colonization by wild mushrooms. With the
spore cast from wild competitors, the likelihood of introducing your
species of choice is greatly reduced. If stumps are not inoculated
within several months of being cut, the probability of success
decreases. Therefore, old stumps are poor candidates. Even so, years
may pass after inoculation before mushrooms form on a stump. But
once a colony begins, the same species may predominate for many
Large-diameter stumps can harbor many communities of
mushrooms. On old-growth or second-growth Douglas fir stumps
common to the forests of Washington State, finding several species
of mushrooms is not unusual. This natural example of
“polyculture”—the simultaneous concurrence of more than one
species in a single habitat—should encourage experimentally
inclined cultivators. Mushroom landscapes of great complexity could
be designed. However, the occurrence of poisonous mushrooms
should be expected. Two notable toxic mushrooms frequent stumps:
Hypholoma fasciculare (=Naematoloma fasciculare) which causes
gastrointestinal upset but usually not death, and Galerina autumnalis,
a mushroom that does kill. Because of the similarity in appearance
between Flammulina velutipes (Enoki) and Galerina autumnalis, I

hesitate to recommend the cultivation of Enoki mushrooms on
stumps unless the cultivator is adept at identification. (To learn how
to identify mushrooms, please refer to the recommended mushroom
field guides listed in Appendix 4.)
Several polypores are especially good candidates for stump
cultivation, particularly Grifola frondosa, Maitake, Ganoderma
lucidum, Reishi, and its close relatives. As the antiviral and
anticancer properties of these mushrooms become better understood,
new strategies for the cultivation of medicinal mushrooms will be
developed. I envision the establishment of Maitake and Reishi
mushroom tree farms wherein stumps are purposely created and
selectively inoculated for maximum mushroom growth, interspersed
among shade trees. Once these models are perfected, other species
can be incorporated in creating a multicanopy medicinal forest.

Drilling and inoculating a stump with plug spawn.

Small-diameter stumps rot faster and produce crops of mushrooms

sooner than bigger stumps. However, the smaller stump has a
shorter mushroom-producing life span than the older stump. Often
with large-diameter stumps, mushroom formation is triggered when
competitors are encountered and/or coupled with wet weather
conditions. The fastest I know of a stump producing mushrooms
from time of inoculation is 8 weeks. In this case, an oak stump was
inoculated with plug spawn of Chicken-of-the-Woods, Laetiporus
(Polyporus) sulphureus. Notably, the stump face was checkered, with
multiple fissures running vertically through the innermost regions of
the wood. These fissures trapped water from rainfall and promoted
fast mycelial growth. Another technique that improves success rates
is to girdle the stump with a chain saw, interrupting direct contact
between the root zone and the above-air portion. A local
mycological society found that all their inoculated stumps produced
which were girdled compared to a small fraction for those that were
not. As with the growing of any mushrooms, the speed of colonization
is a determining factor in the eventual success or failure of any
cultivation project.
For foresters and ecologists, actively inoculating and rotting
stumps has several obvious advantages. Rather than allowing a
stump to be randomly decomposed, species of economic or ecological
significance can be introduced. For instance, a number of Honey
mushrooms, belonging to the genus Armillaria, can operate as both
saprophytes or parasites. Should clear-cuts become colonized with
these deadly, root-rotting species, satellite colonies can be spread to
adjacent living trees. Now that burning is increasingly restricted
because of air pollution concerns, disease vectors coming from
stumpage could present a new as-yet-unmeasured threat to the forest

The Honey mushroom, Armillariella mellea, growing on a fir stump in a rhododendron
garden, an ideal locus for creating a mycological landscape.

The advantages of growing on stumps can be summarized as
Developing a new, environmentally friendly wood products
Recycling wood debris of little or no economic value.
Prevention of disease vectors from parasitic fungi.
The rapid return of wood debris back into a nutritious food for
the benefit of other citizens of the forest. Since the food chain is
accelerated and enriched, ecosystems are invigorated.
Few studies have been published on recycling stumps with
mushrooms. One notable work from Eastern Europe, published by
Pagony (1973), describes the cultivation of Oyster mushrooms
(Pleurotus ostreatus) on large-diameter poplars with a 100% success
rate. Inoculations in the spring resulted in fruitings appearing the
ensuing fall, and continued for several years hence. An average of 4
pounds of Oyster mushrooms were harvested over 4 years (i.e., 1

pound/year/stump). Hilber (1982) also reported on the utility of
using natural wood (logs and stumps) for growing Oyster
mushrooms, and that per cubic meter of elm wood, the yield from
one season averaged 17–22 kilograms. A study in France by Anselmi
and Deandrea (1979), where poplar and willow stumps were
inoculated with spawn of the Oyster mushroom, showed that this
mushroom favored wood from newly felled trees, in zones that
received speckled sunlight. This study confirmed that Pleurotus
ostreatus only attacked deadwood and never became parasitic. Their
study supports my opinion (Stamets, 1990) that the purposeful
inoculation of stumps can forestall the invasion by parasites like
Honey mushrooms of the Armillaria mellea complex. Mushrooms of
this group first kill their host and then continue to live
saprophytically. A stump with Honey mushrooms can later destroy
neighboring living trees. In Washington State, one colony of Honey
mushrooms is blamed for destroying thousands of acres of conifers.

Inoculating a stump with the wedge and the spawn disk technique.

Stumps can be inoculated by one of several simple procedures.
Plug spawn can be inserted into the open face of each stump. If the
stumps are checkered through with cracks, the plugs are best
inserted directly into the fissures. Another method is known as the

wedge or disk inoculation technique. With a chain saw, a wedge is
cut or a shallow disk is sliced from the open face of the stump. The
newly cut faces are packed with sawdust spawn. The cut disk is then
replaced. By hammering a few nails into the stump, you can assure
firm contact between the cut faces. Another method I have
developed is to add spores to biodegradable chainsaw oil so that as
trees are being cut, the stumps are inoculated. (Patent pending).

Plug spawn of Shiitake. Spirally grooved wooden dowels help the mycelium survive
from the concussion of inoculation.

The broadleaf hardwoods are easier to saprophytize with the
gourmet and medicinal mushroom species described in this book
than the softwood pines. And within the hardwood group, the
rapidly growing species such as the alders and poplars decompose
more rapidly—and hence give an earlier crop—than the denser
hardwoods such as the oaks. However, the denser and more massive
stumps sustain colonies of mushrooms for many more years than the
quick-to-rot, smaller-diameter tree species. In a colonial graveyard
in New York state, a four-foot-diameter oak has consistently
produced clusters of Maitake mushrooms, sometimes weighing up to
100 pounds apiece, for more than 20 years! See this image.
Inoculating stumps with strains cloned from native mushrooms is

favored over the use of exotic fungi. Spring inoculations give the
mycelium the longest possible growing season.
Stump cultivation has tremendous potential. These unexploited
resources can become production sites of gourmet and medicinal
mushrooms. Although more studies are needed to ascertain the
proper matching of species to the wood types, I encourage you to
experiment. Only a few minutes are required to inoculate a stump or
dead tree. The potential rewards could span a lifetime.

The “soak and strike” method for initiating Shiitake.

Figure 14. Natural culture of Shiitake in the mountains of Japan. (Photographs, both
from Mimura, Japan, circa 1915.)

Log Culture
Log culture was developed in Japan and China more than a
millennium ago. Even today, thousands of small-scale Shiitake
growers use log culture to provide the majority of mushrooms sold to
markets. In their backyards and along hillsides, inoculated logs are
stacked like cordwood or in fence-like rows. These growers supply
local markets, generating a secondary income for their families.
Attempts to reproduce this model of Shiitake cultivation in North
America and Europe has met with modest success.
The advantage of log culture is that it is a simple and natural
method. The disadvantage is that the process is labor-intensive, and
slow in comparison to growing mushrooms on sterilized sawdust.
Besides Shiitake, many other mushrooms can be grown on logs,
including Nameko (Pholiota nameko), all the true Oyster mushrooms
(Pleurotus and Hypsizygus spp.), Lion’s Mane (Hericium erinaceus),
Wood Ears (Auricularia auricula), Clustered Woodlovers (Hypholoma