Professional Documents
Culture Documents
Mushrooms and Mushroom Cultivation
Mushrooms and Mushroom Cultivation
Cultivation
Shu-Ting Chang, The Chinese University of Hong Kong, Hong Kong, China
Secondary article
Article Contents
. Introduction
. Biodiversity of Mushrooms
. Poisonous Mushrooms
A mushroom is a macrofungus with a distinctive fruiting body, which can be found either
above ground (epigeous) or below ground (hypogeous), large enough to be seen with the
naked eye and to be picked by hand. Cultivation of mushrooms starts from spawn, which is
used to inoculate substrate. Under optimum conditions, mushrooms can be harvested
from 10 days to 6 months after spawning depending upon the strains of species used.
Introduction
Mushrooms have been found in fossilized wood 300
million years old (Editorial, 1997). Almost certainly
prehistoric people used mushrooms as food. The great
early civilizations of the Greeks, Egyptians, Romans,
Chinese and Mexicans prized mushrooms as a delicacy,
appreciated their therapeutic nature and, in some cases,
used them in religious rites. The association of mushrooms
with thunderstorms was common in mythology, and it was
formerly believed that mushrooms were formed by lightning and thunderstorms. Although some wild poisonous
mushrooms became objects of fear and distrust, it is not
surprising that the intentional cultivation of edible mushrooms had a very early beginning. Literature references
indicate (Miles and Chang, 1997) that Auricularia auricula
was cultivated in China as early as ad 600 on wood logs.
Other wood-rotting mushrooms such as Flammulina
velutipes and Lentinula edodes were later grown in similar
manner, but the biggest advance in mushroom cultivation
came in France around 1600 when Agaricus bisporus was
cultivated upon a composted substrate. In the Western
world A. bisporus (champignon or button mushroom) has
remained the mushroom that is produced in the greatest
amounts, but now other species long popular in Asia (e.g.
Lentinula edodes, Pleurotus spp.), and produced there in
large numbers, are making inroads into Western markets.
Mushroom cultivation provides both nutritious protein
rich food and medicinal products. Cultivated mushrooms
have now become popular all over the world. In 1994, the
total world production of edible and medicinal mushrooms
was estimated to be over ve million tonnes, with a value of
over 14 billion US dollars. The bioconversion of lignocellulosic biomass to food and useful products by mushrooms has already had an impact at national and regional
levels, and the predictions are that this impact will continue
to increase. Being without adverse legal, ethical or safety
eects, this form of bioconversion technology has only
favourable socioeconomic and employment impacts.
Mushrooms, like all other fungi, lack chlorophyll. They
are unable to convert solar energy to organic matter like
. Edible Mushrooms
. Mushroom Hunting
. Cultivation of Mushrooms
green plants, but they can convert the huge agricultural and
forest waste materials into human food. The byproducts,
spent substrates, can be used as animal feed and crop
fertilizers. Therefore, sustainable development of mushroom cultivation can be called the non-green lignocellulosic revolution, because mushroom cultivation can
generate equitable economic growth and protect and
regenerate the environment.
Biodiversity of Mushrooms
Mushroom biology is the branch of mycology that deals
with mushrooms. Fungi have been placed in a kingdom of
their own called the Myceteae. The word mushroom may
mean dierent things to dierent people in dierent
countries. In this article, the term mushroom is dened
(Chang and Miles, 1992) in the broad sense as a
macrofungus with a distinctive fruiting body which can
be either epigeous or hypogeous and large enough to be
seen with the naked eye and to be picked by hand. Thus,
mushrooms need not be Basidiomycetes, nor aerial, nor
eshy, nor edible. Some mushrooms belong to the
Ascomycetes, grow underground and have a noneshy
texture. The most common type of mushrooms is umbrella
shaped with pileus (cap) and stipe (stem), e.g. Lentinula
edodes (Figure 1), and some species additionally have a
volva (cup), e.g. Volvariella volvacea (Figure 2), or annulus
(ring), e.g. Agaricus campestris (Figure 3), or both, e.g.
Amanita muscaria (Figure 4). Furthermore, some mushrooms are in the form of pliable cups while others are round
like golf balls. Some are in the shape of small clubs; some
resemble coral; others are yellow or orange jelly-like
globes; and some even resemble the human ear. In fact,
there is a countless variety of forms. The structure that we
call a mushroom is in reality only the fruiting body of the
fungus. The vegetative part of the fungus, called the
mycelium, comprises a system of branching threads and
cord-like strands that branch out through soil, compost,
wood log or other lignocellulosic material on which the
ENCYCLOPEDIA OF LIFE SCIENCES 2002, John Wiley & Sons, Ltd. www.els.net
Figure 4 Amanita muscaris, with an annulus near the top and with the
bulbous base adorned with several concentric zones of white scales
representing the volva (courtesy of Mrs Helena Pereira Lima Caruccio,
Brazil).
(Agaricaceae, Coprinaceae, Tricholomataceae, Hypholomataceae, Strophariaceae, Pluteaceae, Phallaceae, Hericiaceae, Auriculariaceae and Tremellaceae) placed in ve
orders (Agaricales, Phallales, Aphyllophorales, Auriculariales and Tremellales) and two subclasses (Holobasidiomycetidae and Phragmobasidiomycetidae).
One of the basic requirements for breeding better quality
mushrooms in higher yields is the wider availability of a
large reserve of phenotypic variations (traits) which can be
used for selection purposes both by researchers and by the
mushroom industry. Since the mushrooms themselves are
the only source of this genetic material, extinction of a
single strain or species would mean the potential loss of
many thousands of unique genes that could be used for
breeding desirable new strains. The genes contained in
existing mushroom strains and species represent the total
genetic resource, i.e. the entire pool of mushroom
germplasm, which can be preserved by in situ conservation
and ex situ preservation. The maintenance of mushrooms
in natural preserves as part of a strategy for protecting an
ecosystem constitutes in situ conservation. Mushroom
germplasm can also be preserved ex situ as fungal spores or
tissue in the form of a culture collection or gene bank. The
process of collection and classication of information
pertaining to the morphological, physiological, biochemical and genetical characteristics of individual mushroom
strains, and the storage of this information in computer
databases, would provide valuable and readily accessible
information for breeding programmes and academic
research.
Poisonous Mushrooms
Poisonous mushrooms are often called toadstools. However, like the term mushroom, toadstool also may mean
dierent things to dierent people. To some it means any
mushroom except the eld mushroom (Agaricus campestris) or cultivated mushroom (Agaricus bisporus), to others
it is any inedible or poisonous mushroom. Scientically the
term has no meaning at all. It is suggested that the term
toadstool be dropped altogether in order to avoid
confusion and the terms edible, medicinal and poisonous
mushrooms used instead.
It has been estimated that the number of poisonous
mushroom species is fairly large, i.e. about 1000 species, of
which some 30 species are known to be lethal. Since there is
no known eld test to determine if a mushroom is edible or
not, people should never eat any mushroom unless they are
absolutely certain of its identity (Figure 5). Amateurs should
aim to have the mushrooms checked by a competent
mycological authority.
The toxins contained in various species are very dierent
in chemical composition, and thus the eects of poisoning
dier considerably according to the species involved. In
These mushrooms
are not to be
consumed as food.
They are a recipe
for disaster.
A very healthy
and therapeutic
dish of the day!
Edible Mushrooms
Of the 10 000 or more species of macrofungi (mushrooms),
about 5000 are considered to possess varying degrees of
edibility (Miles and Chang, 1997) and more than 2000
species from about 31 genera (Table 1) are regarded as
prime edible mushrooms (Figure 6). Of these, only about
100 are experimentally grown, 50 economically cultivated,
and around 30 commercially cultivated (Table 2); and only
ve or six species are cultivated on an industrial scale.
The moisture content of fresh edible mushrooms varies
in the range 7095% depending upon the harvest time and
environmental conditions, whereas it is about 1013% in
dried mushrooms. The protein content of cultivated
species ranges from 1.75 to 5.9% of their fresh weight,
Hypholoma
Hypsizygus
Lactarius
Lentinula
Lepista
Lyophyllum
Marasmius
Pleurotus
Pholiota
Polyporus
Russula
Stropharia
Termitomyces
Tremella
Tricholoma
Volvariella
Ascomycetes
Morchella
Tuber
Pholiota nameko
Pleurotus citrinopileatus
Pleurotus cornucopiae
Pleurotus cystidiosus
Pleurotus djamor
Pleurotus eryngii
Pleurotus orida
Pleurotus ostreatusa
Pleurotus sajor-caju
Stropharia rugoso-annulata
Tremella aurantia
Tremella fuciformis
Volvariella diplasia
Volvariella esculenta
Volvariella volvaceaa
Mushroom Hunting
The fungi are found just about everywhere. Mushrooms
are rather more selective than other fungi in that the size of
the fruiting body requires the availability of more nutrients
than are required for the production of asexual spores by
microfungi. In damp places, such as tree-fern gullies and
areas of rain forest, plentiful moisture leads to mushroom
formation. They can be collected at most times of the year,
but in drier regions they occur only after seasonal rains. In
these areas there may be a particular ora of mushroom
species associated with the seasons of autumn, summer and
spring. Relatively few mushrooms are produced during the
cold winter months although there are perennial fruiting
bodies that persist during the winter. Formation of
mushroom fruiting bodies depends very much on the
pattern of rain and, in some years, there may be virtually a
complete lack of fruiting.
Mushroom hunters, in addition to bringing basic
equipment and eld guide reference, which will vary
depending on personal requirements and regional conditions, should record such items as date, time, location,
smell, substrate (host) colour, milk (if present), habitat and
anything at all unusual about the specimen. Some
important characteristics for identication disappear
rapidly as the mushroom matures; these characteristics
have to be recorded accurately at the time of collection.
Ecologically, mushrooms can be classied into three
groups: the saprophytes, the parasites and the mycorrhizae. There are only a few parasitic mushrooms. Most of
the cultivated gourmet mushrooms are saprophytic fungi.
Some are mycorrhizal mushrooms, e.g. Perigold black
Cultivation of Mushrooms
Cultivation of edible mushrooms had its beginning in
China around ad 600 when Auricularia auricula was rst
cultivated. It is interesting to look at the historical record of
the mushrooms rst cultivated (Table 3). Eight of these
mushrooms, including ve of the currently most popular
species, were cultivated prior to the twentieth century,
while the remaining 31 were rst successfully cultivated in
the twentieth century. The great increase in the number of
species brought into cultivation in the 1980s and 1990s
corresponds with the dramatic acceleration in total
worldwide cultivated mushroom production (e.g. 1975,
0.90 million tonnes (Mt); 1981, 1.26 Mt; 1986, 2.18 Mt;
1990, 3.76 Mt; and 1994, 4.91 Mt from the 10 most popular
species, Agaricus bisporus/bitorgues, Lentinula edodes,
Pleurotus spp., Auricularia spp., Volvariella volvacea,
Flammulina velutipes, Tremella fuciformis, Hypsizygus
marmoreus, Pholiota nameko and Grifola frondosa). In
recent years, several new species of edible mushrooms, e.g.
Hericium erinaceus, Dictyophora indusiata, Stropharia
rugoso-anmulata, Agrocybe aegerita, Coprinus comatus,
Agaricus blazei and Agrocybe cylindracea have also been
commercially cultivated. Although total production by
these new species has been minimal to date, it does reect
the great consumer interest in new varieties of edible
mushrooms. Mushroom scientists are making a great eort
to bring additional wild species under cultivation to satisfy
consumer demand.
5
Species
Species
600
800
1000
1232
1600
1621
1700
1800
1900
1950
1958
1958
1958
1960
1961
1962
1969
1973
1974
Auricularia auricula
Flammulina velutipes
Lentinula edodes
Poria cocos
Agaricus bisporus
Ganoderma spp.
Volvariella volvacea
Tremella fuciformis
Pleurotus ostreatus
Agrocybe cylindracea
Pleurotus orida
Pleurotus ferulae
Pholiota nameko
Hericium erinaceus
Agaricus bitorquis
Pleurotus abellatus
Pleurotus cystidiosus
Hypsizigus marmoreus
Pleurotus sajor-caju
1981
1982
1982
1982
1983
1983
1983
1984
1984
1984
1985
1985
1986
1987
1988
1989
1990
1991
1991
1997
Pleurotus citrinopileatus
Dictyophora duplicata
Oudemansiella radicata
Hohenbuehelia serotina
Armillaria mellea
Grifola frondosa
Pleurotus sapidus
Coprinus comatus
Amanita caesarea
Hericium coralloides
Tremella mesenterica
Sparassis crispa
Morchella spp.
Lyophyllum ulmarium
Lentinus tigrinus
Gloestereum incarnatum
Tricholoma lobayense
Tricholoma mongolicum
Tricholoma gambosum
Cantharellus cibarius
Reproduced with permission from Chang (1993) and Danell and Camacho (1997).
References
Chang ST (1993) Mushroom biology: the impact on mushroom
production and mushroom products. In: Chang ST, Buswell JA and
Chiu SW (eds) Mushroom Biology and Mushroom Products, pp. 320.
Hong Kong: Chinese University Press.
Chang ST and Buswell JA (1996) Mushroom nutriceuticals. World
Journal of Microbiology and Biotechnology 12: 473476.
Chang ST and Miles PG (1989) Edible Mushrooms and Their Cultivation,
p. 345. Boca Raton, FL: CRC Press.
Chang ST and Miles PG (1992) Mushroom biology a new discipline.
The Mycologist 6: 6465.
Chilton WS (1978) Chemistry and mode of action of mushroom toxins.
In: Rumack BH and Salzman E (eds) Mushroom Poisoning: Diagnosis
and Treatment, pp. 88117. West Palm Beach, FL: CRC Press.
Further Reading