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ABSTRACT
Tropical regions have the potential to be one of the richest sources of cultivatable
fungal species. Even though there has been considerable research on the taxonomy and
phylogeny of these mushrooms, there has been far less research on their domestication.
The purpose of this paper is to review and detail the methods we have used for the
discovery and domestication of wild tropical mushrooms. As it is difficult to cultivate
mycorrhizal species we have mainly concentrated on saprobic species. Methods include
collection, isolation, spawn production and fruiting body production testing in sawdust
and compost media. We also discuss a semi-industrial approach of inoculating wild edible
mushroom spawn into the natural environment to produce seasonal mushrooms. We
have collected and isolated numerous strains of species of wild mushrooms and present
initial results on domestication attempts. It is hoped to be able to introduce these to the
mushroom growing industry in the future.
noted, more than 2,000 years ago [3]. developed by Fritsche [19]. Commercial
Edible mushrooms were gathered from production of mushrooms began in the
forests in ancient Greek and Roman times United States in the 1880s [8].
and were highly valued, though more by Agaricus is the leading mushroom crop
high-ranking people than by peasants [4]. worldwide and accounted for 99% of the
The British colonial records in Africa United States’ mushroom production in
contain little information about the local 1997 [7, 20]. The oyster mushroom
use of wild edible mushrooms, despite the (Pleurotus spp.) has been domesticated
fact that people throughout southern more recently and now ranks second in
Africa have eaten them for centuries [5, 6]. world production [21]. The Shiitake
While commercial harvesting of wild mushroom, which is very popular in Asian
mushrooms continues today, most of the cultures, ranks third in world production
world’s supply comes from commercial [22]. Many other edible mushrooms, such
mushroom growers [1, 7, 8]. The Chinese as the straw mushroom (Volvariella
first cultivated Shiitake (Lentinula edodes) volvacea) and wood ear mushrooms
mushrooms around 1,100 AD, with (Auricularia auricula), are gaining in
domestication efforts beginning century’s popularity [7]. It is estimated between 650-
earlier [7, 8]. The white button mushroom 700 mushroom species are edible [23], but
(Agaricus bisporus), which are most familiar only approximately 130 have been
to Americans and Europeans, was first domesticated (Table 1). The button, oyster,
domesticated in France in 1650 [8, 9]. The and shiitake mushrooms make up about
first truffle plantation took place in Italy 70% of the world’s production [24]. During
and France in 1970s [10]. For approximately the past 30 years, mushroom production
160 years, A. bisporus was grown in open worldwide increased twenty-fold, with
fields. At some point, it was realized that much of that increase occurring in the
mycelium, or what is referred to as the 1980s and 1990s [1, 8]. Increased demand
spawn of the mushroom, was what gave for specialty mushrooms has been
rise to the fruiting bodies and could be particularly strong. Asian countries
utilized much like the seed of plants to continue to dominate world production
grow mushrooms [11-13]. The first and consumption; however, consumption
cultivation in a cave (i.e. production year in the United States has increased sharply
round) began in Paris around 1800 and in recent years, providing potential
during the 19 th century production of opportunities for mushroom growers [1,
spawn by industrial spawn producers began 7, 8].
[14]. The first experimental culture of During the period between 1990-1994,
mycelium from spores was carried out by world mushroom production increased by
Costantin and Matruchot [15] in Paris at 30.5%, reaching about 4,909 thousand tons
the Pasteur Institute [14, 16], whereas the in 1994 [25-27]. The global economic value,
first strain isolation from cultures from although difficult to evaluate, has been
mushroom tissue was by Boyer [17]. The estimated to be more than 9.8 billion
first white cultivar derived from a white dollars per annum [27]. In Thailand
capped button mushroom was from a mushroom growing has also increased
culture in the USA in 1926 [18]; and the steadily [28, 29]. The saprobic mushrooms
first commercial hybrid strains were species that to our knowledge are cultivated
Chiang Mai J. Sci. 2014; 41(4) 733
in Thailand are listed in Table 2. The Thai lion’s mane (Hericium erinaceus) have been
government is trying to enhance people’s introduced to Thailand [29, 51]. A new
life by encouraging mushroom growing, hybrid strain from Thai and French strains
and an increase of mushroom production of Agaricus subrufescens were developed
at the community level is expected [29]. successfully between INRA, France and
Even though mushrooms have been Mae Fah Luang University, which fructifies
domesticated for cultivation since early in tropical climates [52].
times, the most commonly grown strains Mushrooms are not only used in
are temperate species [30-32]. Tropical traditional medicines but are known to
mushrooms are, however, numerous, and contain various bioactive components
recent studies on various genera have which can be used in cosmetics [29, 59], and
shown them to be specious. Numerous new medicine [54, 60, 61, 62]. People in most
species have been introduced to science parts of the world enjoy eating mushrooms
with some being edible and other and therefore there is enormous potential
having medicinal use [33-48]. Therefore for introducing new tropical mushrooms
domesticating tropical mushrooms provides to the global market. Even though there
an enormous opportunity for tropical and has been considerable research on the
subtropical countries [1, 49-52]. Most taxonomy and phylogeny of these
tropical mushrooms grow rapidly and mushrooms, there has been far less research
o
produce fruiting bodies at 25 C or higher on their domestication. The purpose of this
and thus can be produced more quickly than paper is to review and detail the methods
temperate species [51]. The tropical mushrooms we have used for the discovery and
can also be produced on readily available domestication of wild tropical cultivatable
and cheap waste products such as saw dust, mushrooms.
corn cobs, rice straw, sugarcane bagasse,
and other forest and agricultural wastes 1.1 Conservation of Unknown Genetic
[53]. Therefore growing new tropical Diversity During the Domestication
mushrooms will help recycle agricultural Process
and forest waste products, provide income The success of a newly cultivated strain
to various entrepreneurs and local industries, depends on both economical and biological
provide nutritional and medicinal foods factors. There are two groups of biological
and prevent pollution through less factors: the first group relates to mushroom
dumping and burning of agricultural waste production and the second to mushroom
[41, 51, 54]. consumption (edibility/toxicity, the
Several new wild edible mushrooms nutritional and medicinal aspects, and
have been successfully domesticated over appetency). Biological factors affecting the
the last few years, especially in tropical areas success of production depend on the
[42, 48, 51, 52, 54-58]. Klomklung et al. environment and any methods to control
[51], has shown that it is possible to it, and equally on genetic factors. These
domesticate local strains of Pleurotus genetic factors depend on the intraspecific
giganteus that can grow at temperatures variability, of the genetic structure of the
consistent with Thailand farm production. population, of epigenetic factors and
More recently, medicinal mushrooms genomic stability [52]. At each step of the
such as reishi (Ganoderma lucidum) and domestication process we have to make a
734 Chiang Mai J. Sci. 2014; 41(4)
choice and thus we make a genetic or even spore print from the wild specimen (= the
epigenetic selection: which field specimens parent) can save time when performing
to choose; from which part of the specimen hybridization.
to isolate; which isolate to choose, and 7) For any isolate that successfully
later which subculture. There is some passes the fruiting test and which is
experimental knowledge guiding these maintained on non-synthetic medium (such
decisions, however these are also species as compost extract medium for Agaricus),
dependent. make in parallel a mycelium growth rate
To search for new species that can be test on a synthetic or semi synthetic
cultivated we make some general recom- reproducible medium (for example 14
mendations that the senior author P. Callac days-test on malt agar medium). The
has realized from decades of research in growth rate is characteristic of the strain
mushroom growing: that will be further improved.
1) If several sporocarps are found from 8) Each retained strain should be
the same site make isolates from at least two. conserved through at least two
2) If you have a good specimen make independent clonal lineages.
at least two isolates, one from the cap and Although the above recommendations
one from the stipe; all isolates must have are general, one should be careful not to
different identification code numbers even generalize results obtained from a single
if they come from the same piece of cap of field collection, as these may not be
the same sporocarp. representative of the entire species. For
3) Any isolate growing very slowly or example, isolates from some wild
developing sectors with different aspects or specimens of A. subrufescens never produce
growth rates can be discarded immediately. fruiting bodies when domesticated; in A.
4) Although the optimal conditions for bisporus isolates from field specimens
each species is unknown, where possible generally fructify, but some with a very
strain isolation must be made on a culture low yield and some have yields as great as
medium similar the one that will be used the best cultivars (but not their level of
to maintain the strain over many years. quality).
5) Avoid taking too much precaution
against potential contamination; for 2. RECOMMENDATIONS FOR DISCOVERING
example in most cases it is not necessary to CULTIVABLE MUSHROOMS
use antibiotics in the culture medium to 2.1 Edible Cultivatable Mushrooms
isolate a strain and it is not necessary to To discover new industrial mushrooms,
use very thin slices of mushroom tissue. one must 1) make certain if it is edible and
Thus the strain may be less fastidious in it 2) establish if it can readily be cultivated.
sterile requirements when producing Information on edibility can be gained from
fruiting bodies. local traditional or scientific knowledge.
6) Obtaining a good isolate is more Local traditional knowledge is the best way
important than obtaining a spore print, to find out whether a species is edible or
since the isolate will allow you to get a medicinal. Leaning from indigenous people
spore print from a future fruit body, while including local names, seasonality, harvest
the spore print will never allow you to get techniques and habitat management is a
the parental strain; however a quickly made short-cut for mushroom identification.
Chiang Mai J. Sci. 2014; 41(4) 735
Key informants include local mushroom safely consume the delicious collection
collectors, housewives, and herbal doctors. (MRC unpub. data). Pleurotus giganteus is
A scientific name can also be used to find considered as a non-edible mushroom by
out whether a species is edible or poisonous, locals in Thailand, but is known as a
or if it has medicinal or other useful delicious edible mushroom in Sri Lanka
properties [1, 63, 64]. Local traditional [51, 67].
knowledge, taxonomic and hybrid Once it is established that the
knowledge (scientific and local) can also collection is edible we must then establish
establish whether species are edible. In the if it can be cultivated as not all mushrooms
genus Agaricus all the species are edible are cultivatable. Hints to their cultivat-
except in the section Xanthodermatei ability can be gleaned from existing
[42, 65], and excellent edible frequently knowledge. If other species in the genus or
eaten species belong to sections Bivelares, section of the genus are already cultivated,
Arvenses, Sanguinolentes, and Agaricus it is more likely to have success with novel
[40, 41]. Species of Minores are also cultivatable species (INRA unpub. data).
probably good, but generally they are too For an example Agaricus subrufescens is a
small [66]. Species of section Chitonioides well known edible, medicinal, and cultivated
are also sometimes eaten, although they can mushroom in the section Arvenses. We have
have a bad smell before cooking. Edibility collected and described the new species,
of other sections is not well known. In Agaricus flocculosipes, from northern
Lentinus, almost all the species are edible Thailand, in the same section, and obtained
unless they are too hard to eat [40-42]. successful cultivation results [42, INRA
Volvariella species are also edible. Local unpub. data]. On the other hand the new
people did not realize that a giant species Agaricus flavicentrum, which is a
Volvariella growing at the Mushroom delicious member of section Agaricus is
Research Centre in northern Thailand was unlikely to be cultivatable as others species
edible (they said no with horror). It was in this section cannot be produced
identified using morphology and named as commercially (MFU unpub. data).
a Volvariella species and we were able to
Table 1. Continued
Scientific names Common names References
Antrodia cinnamomea Stout camphor fungus, Zhan ku, [83]
Ni zh ng zh , Kusunoki shiba,
Niu-chang-ku,
Cinnamomum kanehir,
Grib kamphorniy
Antrodia camphorata Stout camphor fungus, Zhan ku, [83]
Ni zh ng zh , Kusunoki shiba,
Niu-chang-ku,
Cinnamomum kanehir,
Grib kamphorniy
Armillaria mellea Chiodini, honey [80, 84-86]
Armillaria tabescens [86]
Auricularia auricula Wood ear [25, 73, 80, 86, 87]
Auricularia
Ear fungus [1, 73, 86]
fuscosuccinea
Auricularia nigricans Black chinese mushroom, [30, 80, 86, 88]
(=Auricularia Wood ear fungus, Wood fungus,
polytricha) Ear fungus, or Tree ear fungus
Auricularia cornea Hairy jew’s ear, [86]
Wood ear, Cloud ear.
Auricularia peltata - [86]
Auricularia
- [86]
mesenterica
Calvatia gigantea Giant puffball [1, 89]
Coprinus comatus Lawyer’s wig, [30, 71, 73, 86, 89]
shaggy-mane mushroom
Coprinus
Inky cap [29]
atramentarius
Cordyceps sinensis Cordyceps Mushroom, [83]
Caterpillar fungus
Cordyceps militaris Caterpillar fungus, Orange club [83]
Cordyceps sobolifera The vegetable fly, [83]
Cordyceps cicadae, Insect flower,
Crown cicada
Collybia reinakeana - [89]
Creolophus pergamenus Bear’s head [30, 78]
Daedalea quercina Oak mazegill, Maze-gill fungus [1, 90]
Dictyophora duplicata Netted stinkhorn, Wood witch [1, 86, 91]
Dictyophora indusiata Stinkhorn [80, 91]
Dictyophora
Stinkhorn [86]
rubrovolvata
Chiang Mai J. Sci. 2014; 41(4) 737
Table 1. Continued
Scientific names Common names References
Flammulina velutipes Enokitake, Winter mushroom, [25, 29, 30, 70, 71, 73, 86]
Velvet stem
Fomes fomentarius Tinder fungus, Hoof fungus, [1, 91]
Tinder conk, Tinder polypore,
Ice man fungus
Ganoderma Artist’s bracket, Artist’s conk, [92-95]
applanatum or Flacher lackporling
Ganoderma australe Southern bracket, Reishi australe [83, 92, 94]
Ganoderma curtisii Southern bracket, Reishi australe [83, 94]
Ganoderma lucidum Reishi [29, 30, 70, 73, 86, 89,
92-94, 96]
Ganoderma oregonense Western varnish shelf, [83, 93]
Oregon polypore,
American reishi
Ganoderma resinaceum Lacquered bracket, L ngzh [83, 92, 93]
resinaceum de, Reishi resinaceum,
Ganoderme r sineux
Ganoderma sinense Black reishi [1, 83, 86, 93]
Ganoderma tenus [1, 83, 93]
Ganoderma tsugae Hemlock varnish shelf [83, 93]
Grifola frondosa Maitake [25, 30, 70, 97]
Grifola albicans Choreimaitake [86, 98]
Hericium coralloides Icicle tooth fungi, Comb tooth, [1, 99]
Conifer coral, Coral spine,
Coral tooth
Hericium erinaceus Lion’s mane mushroom [29, 30, 70, 73, 86, 99, 100]
Hypholoma capnoides - [1, 30, 101]
Hypholoma
Brick cap [1, 30, 102]
sublateritium
Hypsizygus marmoreus Bunashimeji [25, 86, 100, 103]
Hypsizygus tessulatus Shimeji [30, 70, 100, 104]
Hypsizygus ulmarius Shirotamogitake [30]
Inonotus obliquus Chaga, Hu sh g , [83, 105]
Naname inonotus, Polypore
cendr , Polypore oblique
Inonotus levis NA [106]
Kuehneromyces
Sheathed woodtuft [1, 107]
mutabilis
Laetiporus sulphureus Chicken-of-the-woods [1, 108]
Laricifomes officinalis
Agaric, Agarikon, Quinine conk [109]
(Fomitopsis officinalis)
738 Chiang Mai J. Sci. 2014; 41(4)
Table 1. Continued
Scientific names Common names References
Lentinula edodes Shii-take [29, 30, 70, 71, 73, 110]
Lentinus connatus - [111, 112]
Lentinus cladopus - [111, 112]
Lentinus polychrous - [111]
Lentinus sabnudus - [111]
Lentinus squarrulosus Tropical white rot fungus [89, 111, 112]
Lentinus strigosus
Ruddy panus [1, 113]
(=Panus rudis)
Lentinus tigrinus - [1, 89]
Lentinus tuber-regium King tuber mushroom [1, 86]
Lepista nuda Blewit [71, 73, 114]
Lepista sordid Flesh-brown blewit [115]
Lignosus rhinoceros Tiger milk mushroom, [116]
Tiger’s milk ling-zhi
Lyophyllum decastes Fried chicken, Hatakeshimaji [80]
Lyophyllum fumosum - [1, 117]
Lyophyllum ulmarium
(=Hypsizygus Ulmenr sling, Elm leech [1]
ulmarium)
Macrocybe gigantea
(=Tricholoma Giant mushroom [1, 118]
giganteum)
Macrolepiota gracilenta Parasol mushroom [29, 119]
Macrolepiota dolichaula Parasol mushroom MFLU unpub. data
Macrolepiota procera Parasol mushroom [1, 120, 121 ]
Marasmius oreades The scotch bonnet [1]
*Morchella angusticeps Black morel [1, 122]
*Morchella esculenta Common morel, Morel, [1, 123 ]
Yellow morel, True morel,
Morel mushroom, Sponge morel
Naematoloma
Bricktop, Chestnut, Kuritake [80]
sublateritivum
Neolentinus lepideus
Scaly Lentinus, Train wrecker [1, 124]
(=Lentinus lepidus)
Oligoporus spp. - [1]
Oudemansiella radicata Rooted Collybia, Rooted agaric [1, 86]
Oudemansiella
submucida Porcelain fungus [1, 125]
Oxyporus nobilissimus Giant polypore fungus [1]
Panaeolus cyanescens Pan cyan [30, 71]
Panaeolus subbalteatus The belted cap Panaeolus [71]
Chiang Mai J. Sci. 2014; 41(4) 739
Table 1. Continued
Scientific names Common names References
Panaeolus tropicalis Magic mushroom [1]
Panellus serotinus Green oyster, Last fall oyster, [80]
Hiratake
Phallus impudicus Common stinkhorn [1]
Phellinus linteus Black hoof fungus, Fire sponge [83]
Pholiota nameko Nameko, Viscid mushroom [25, 30, 86, 100]
Pholiota adipose Fat Pholiota [80]
Piptoporus betulinus Birch polypore,Birch bracket, [1, 126]
Razor strop
Piptoporus indigenous - [1]
Pleurocybella porrigens Angel’s wings [1]
Pleurotus abalones Abalone [80, 86]
Pleurotus citrinopileatus Golden oyster mushrooms [1, 29, 30, 86, 73]
Pleurotus cornucopiae Golden oyster [80, 127, 128]
Pleurotus cystidiosus Oyster mushrooms, Ohritake [29, 30, 80, 127, 129]
Pleurotus djamor Rose, Pink oyster [30, 80]
Pleurotus eryngii King oyster mushrooms [29, 30, 73, 86, 127, 128, 130]
Pleurotus euosmus Oyster mushrooms [1, 30]
Pleurotus florida Oyster mushrooms [127]
Pleurotus ostreatus Oyster, white mushrooms [29, 30, 70, 71, 131]
Pleurotus pulmonarius Phoenix-tail [30, 80]
Pleurotus rhodophyllus - [86]
Pleurotus sajor-caju Oyster mushrooms, [86, 89, 100, 112, 127,
Himarayahiratake, 132, 133]
Angel mushroom
Pleurotus sapidus - [69, 73]
Pleurotus giganteus Seri pagi in Malaysia, [73]
Urupaha in Sri Lanka
Pluteus cervinus Deer, Fawn mushroom [1]
Polyporus indigenous - [1]
Polyporus
- [89]
grammocephalus
Polyporus saporema - [1]
Polyporus umbellatus Lumpy bracket, [1, 30]
(= Dendropolyporus Umbrella polypore
umbellatus)
Poria cocos Hoelen, poria, Tuckahoe, [69, 134]
(Wolfporia extensa) China root, Fu ling, and
Matsuhodo.
Psilocybe cubensis San Lsidro, Cubensis [71]
Psilocybe cyanescens Cyan, Grandote [71]
740 Chiang Mai J. Sci. 2014; 41(4)
Table 1. Continued
Scientific names Common names References
Psilocybe mexicana Mushroom of the gods, Pajaritos [71]
Psilocybe tampanensis The tempa Psilocybe [71]
Schizophyllum
Split gill 1, 89, 135]
commune
Sparassis crispa Cauliflower mushroom [1, 136]
Stropharia rugoso- The wine red Stropharia, [30, 69, 71, 73]
annulata The giant Stropharia
Trametes cinnabarina - [1]
Trametes versicolor Turkey tail [1, 69]
Tremella fuciformis Snow-fungus, Jelly fungus [25, 29, 69, 70, 73]
Tremella aurantia - [69]
Tremella cinnabarina - [73]
*Tricholoma giganteum Nioushimeji [69, 73, 100, 137]
*Tricholoma crassum Matsutake crassum [29]
*Tuber aestivum Summer truffle [80, 138-140]
*Tuber indicum Chinese truffle [138-141]
*Tuber magnatum Piedmont white truffle [80, 140]
*Tuber melanosporum Perigord black truffle [80, 138-140]
Volvariella bombycina Silky sheath, Silky rosegill, [1, 30]
Silver-silk straw mushroom,
Tree mushroom
Volvariella diplasia Banana, Straw [80]
Volvariella volvacea Padi straw mushroom, [29, 30, 69-71, 73, 89]
The Chinese mushroom
V. volvacea var. Padi straw mushroom, [1, 30]
gloiocephala The Chinese mushroom
* Semi-industrial mushrooms
samples are placed in a plastic container and peculiarities of the type of fungi/
with small compartments and returned to mushrooms that are routinely grown [161].
the laboratory. Published references are Media normally affects colony morphology
used for correct identification, i.e. and colour, whether particular structures
Heinemann’s taxonomic system [147] for are formed or not, and may affect whether
Agaricus species which utilizes morphology the fungus/mushroom will even grow in
[146] are recommend publications to use, culture [162]. The widely used media for
as is Kerrigan’s work on Agaricus and mushroom culture is Potato Dextrose Agar
molecular approaches [65, 148-154] and for (PDA) [163]. The mushroom mycelia grow
other mushroom genera [155-159]. The well on PDA. Other media: Potato Sucrose
method for obtaining cultures is well- agar (PSA), Potato Dextrose Yeast Agar
established and routinely used by the (PDYA), Malt Extract Agar (MEA), Yeast
INRA laboratory and our group. These Malt Agar (YMA) and, Mulasaki and Skoog
methods of obtaining cultures from (MS) are also used to culture mushrooms
Agaricus blazei, A. bisporus, Coriolus [164, 165].
versicolor, Lentinula edodes, Pleurotus Maintenance of strains and genetic
ostreatus, P. tuber-regium, and Tremella characteristics of axenic cultures are
fuciformis and their cultivation are detailed important for mushroom cultivation.
in Stamets [30, 71]. In addition single spore There are various methods to maintain
isolates of the strains can be obtained short term and long term viability of
following the protocols of Raper [160], strains such as storage in liquid nitrogen,
prior to sub-culture for breeding. mineral oil, distilled water, or sawdust-
freezing method [166-170].
3.2 Media and Long Term Storage
Isolates of mushrooms are grown in a 3.3 Testing Cultivability
wide range of culture media and used for Approximately 30,000 mushroom
inoculation spawn and maintaining strains. species have been described [70] however,
Most mycologists develop preferences for less than 150 species are cultivated (Table 1)
certain types of media based on experience [1, 30, 71] and in Thailand only about 22
species are produced industrially (Table 2). Over the previous five years, we have
The number being cultivated is steadily isolated several wild mushrooms strains
increasing and information concerning collected from Northern Thailand (e.g.
their production is readily available [30]. species of Agaricus, Lentinus, Lepista, Lepiota,
o
extract, and peptone, ammonium hydrogen and autoclaved twice at 121 C, 15 psi for
phosphate ((NH 4) 2 HPO 4), ammonium 15 minutes. After cooling, the mycelia
nitrate (NH 4NO 3) and calcium nitrate from the pure mushroom cultures with a
(Ca(NO3)2) can be used as N sources [176, portion of PDA are inoculated to sorghum
180]. Media should be supplemented with grain medium [183]. The grains invaded by
each C and N source and the volume mycelia are used as spawn after the
depends on the media and mushroom. mycelium has invaded all grain medium
Subculture of the mushroom mycelia in inside the bottles [183].
different media and incubation in the same
conditions can be used to establish optimum 3.4.6 Preparing Sawdust Bags
growth conditions by measuring the Wood inhabiting mushrooms can be
diameter of the mushroom colony daily cultivated in any type of lignocellulosic
[183]. material such as straw, sawdust, and rice
hull [186]. It has been shown that oyster
3.4.5 Spawn Production mushroom cultivation (P. ostreatus) on
Spawn is the media for transfer of the various sawdust types, gives different
mushroom mycelium to the growing fruiting yields [187]. Presently sawdust is
substrate by the cultivator [175]. Depending commonly used and is the preferred
on the substrate to be inoculated, the medium at the commercial scale [188].
vehicle can be grain, sawdust, wood chips, Furthermore, it has been shown that P.
dowels, or rope. There are many types of ostreatus can give maximum biological
spawn, including virgin spawn (spawn efficiency on rubber tree sawdust [187].
collect from the pastures & meadows), Softwood sawdust such as coconut, cashew,
flake spawn (breaking of beds through mango, and rubber are known to be more
which mushroom mycelium has run), mill suitable than hardwood sawdust [189]. The
track spawn (bricks dried and made from availability of raw materials (sawdust, rice
mixture of horse dung, cow dung and lawn straw, sugarcane wastes) are key factors for
soil) and manure spawn (on sterilized choosing agricultural wastes for growing
horse manure or manure compost) [16]. mushrooms in any region [187]. The most
Technological developments have meant commonly and easily cultivated mushrooms
that there is good equipment available to in Thailand and in South East Asian
produce high quality spawn. For example countries are oyster mushrooms, ear
in INRA we use rye grain spawn made by mushrooms, and straw mushrooms.
Euromycel, France [184, 185]. In Thailand Lentinula edodes, Lentinus species (e.g.
we mostly use sorghum as the spawn Lentinus squarrosulus), Ganoderma species
medium, because it is readily available and (e.g. Ganoderma lucidum), Macrolepiota sp.
inexpensive. The grains of sorghum are (Macrolepiota gracilenta), Agrocybe sp. (e.g.
cleaned manually to remove inert matter, Agrocybe cylindracea,) can also be cultivated
stubble and debris. The cleaned grains are successfully [29]. It is recommended that a
thoroughly washed and soaked in tap water newcomer to mushroom cultivation starts
for 12 hours or overnight. Thereafter, the with easy to grow and commercially viable
soaked grains are drained until the excess mushrooms such as shitake, straw and
water is removed. The cleaned dry grains oyster mushrooms.
(50 g) are filled in to narrow neck bottles In tropical areas different types of
Chiang Mai J. Sci. 2014; 41(4) 745
sawdust are used depending on the area and in the compost. It is then cooled for 24 hrs
o
the trees available, with Hevea brasiliensis to 25 C, when the compost is generally
being most popular [29, 51, 190], followed used and inoculated with spawn.
by Acacia auriculiformis [191], Mangifera
indica [180] and Tamarindus indica. For 1 3.4.8 Trying Out Strains
kg saw dust bag, 10 g of calcium carbonate, Fruiting tests of wild strains are
50 g of rice bran, 10 g of pumice, 10 ml of important to introduce new mushrooms
molasses, 10 g of flour, 10 g of brewers to market. Depending on the type of the
waste are added. The components are mushroom we use compost or saw dust
mixed with water to make moisture around media. As a rule of thumb, for wood-
65-70% and then 800 g of the substrate are inhabiting mushrooms we use sawdust
tightly packed in 25 × 8 cm poly propylene media in bags and for soil-inhabiting
bags and capped with plastic ring or bottle mushrooms (e.g. Agaricus, Macrolepiota)
neck leaving a space to later inoculate the we used rice straw compost.
mycelium [29-30, 51, 100]. The sawdust bags For wood-inhabiting mushrooms
are sealed with cotton wool plugs and protocols adapted from Klomklung et al.
covered with newspaper, and tied with a [51] and Namphung Klomklung (pers.
rubber band. The sawdust bags are comm.) are followed. Spawn (2% weight
o
sterilized at 121 C for 15 minutes or at 90– of bags) is inoculated onto the surface of
o
100 C for 3 h. After the temperature drops sawdust growing bags. The bags are kept
o o
down to 25 C, spawn of 10% of weight of in a dark incubation room at 25 ± 1 C with
the sawdust bag is inoculated to the sawdust 70%-80% relative humidity and opened
bags. Sawdust bags are kept at room when the mycelium had completely
o
temperature (25 C) with humidity around colonized the substrate. The upper portions
70-80% in order to produce fruiting bodies of the bags are then opened and surface of
[51]. the substrate is scraped slightly with a sterile
teaspoon to remove the thin whitish
3.4.7 Preparing Compost mycelia. The substrate bags are then placed
The compost process for mushroom on the shelf and covered with black cloth
cultivation in Thailand is prepared from to give appropriate ventilation. To
rice straw as the main substrate. The maintain 80-85% relative humidity in the
chopped rice straw is supplemented with growing house, water is sprinkled inside
rice bran, urea and ammonium phosphate the growing house. Water spraying is
to increase the N source in compost. carried out daily until pin heads and
Gypsum and calcium carbonate are added eventually fruiting bodies fully develop.
to compost for buffering the pH to 7.5–8 The fruiting bodies are manually harvested,
(Royal Project method, unpub. data). The counted and weighed [51].
compost stack is homogeneously mixed and For soil-inhabiting mushrooms
moistened to 70% [water added]. The protocols adapted from Klomklung et al.
outdoor process of compost lasts 20 to 25 [51] and Naritsada Thongklang (pers.
days with eight turn cycles. The compost comm.) are followed. Spawn (2% weight
o
is heated to 55–60 C for 3–6 hrs with hot of compost) is inoculated into compost
o
air and steam for pasteurization, which media and incubated at 25 C. After the
neutralizes all harmful microbes and insects mycelia fully covers and grows throughout
746 Chiang Mai J. Sci. 2014; 41(4)
the compost a casing layer is applied [192]. mushrooms. There have however, been a few
A casing layer can be made from soil and studies on inoculating edible mushrooms
sand (1:1) and mixed with 2% calcium in natural environments so that they can be
carbonate at a pH 7.5–8 and a thickness of naturally harvested [1, 194, 199]. Research
o
1–1.5 inch is added and incubated at 25 C on planting mycelia in soils, could lead to
for Agaricus, Lepista, and Macrolepiota [184], the establishment of successful and
Royal Project method, unpub. data. An sustainable production technologies [200].
ideal environment, e.g. 90–95% of humidity In this paper we term this semi-industrial
o
at 25–27 C and low CO2 concentration is mushroom cultivation.
maintained. After casing, watering begins
usually once a day. The fruiting test is 4.2 Choosing Mushrooms and Location
carried out with four replicates trays. The for Semi-Industrial Mushroom Cultiva-
fruiting bodies, including those with open tion
and closed caps, are manually harvested, and Saprobic mushrooms colonize and
counted and weighed [193]. derive their nutrition from rotting wood and
organic matter found in soils [1, 201, 202]
4. SEMI-INDUSTRIAL APPROACH OF and there are numerous species that could
INOCLATING WILD EDIBLE MUSHROOM SPAWN be utilized in semi-industrial mushroom
INTO THE NATURAL ENVIRONMENT TO production. Mycorrhizal mushrooms on
PRODUCE SEASONAL MUSHROOMS the other hand form symbiotic relationships
4.1 Introducing Edible Mushrooms to with their host trees and plants and have
Grasslands, Orchards or Forests fundamental roles in determining plant
Uncertainty of harvests of wild health and in the functioning of forest
mushrooms from one year to the next ecosystems [1, 23, 203]. Tree species can
makes commercial exploitation difficult form mycorrhizal associations with more
and some attempts have been made to than one fungus, and a fungus may be
overcome this by cultivating high demand associated with more than one tree [1, 204-
mycorrhizal species such as Tricholoma 211]. Both types of fungus can be used in
matsutake, as then these mushrooms semi-industrial mushroom production, but
become available year round [23, 194, 195]. research is need for the best methodologies
Trees have successfully been infected with for each potential species. To semi-
truffles [194, 196-198] and managed under industrialize mycorrhizal mushrooms,
controlled conditions in Italy and elsewhere their spawn is inoculated directly in the
to naturally produce these expensive soils of specific hosts in forests. As the trees
mushrooms [1, 138, 139, 141, 196, 197]. grow, besides timber, a secondary crop of
Although industrial cultivation is the mushrooms is produced. Normally
only way to provide a year round supply inoculation has to be done twice (spring
of edible mushrooms, this can be supple- and autumn) in the first year and it will
mented by semi-industrial methods where fruit in the following 20 years [138, 139,
the mushrooms are inoculated directly into 141, 195-197]. Tree seedlings in pots can
natural environments and sporulate during also be inoculated with mycorrhiza, so that
ideal weather. This section deals with the they already have mycorrhizal associations
semi-industrial methodologies that are with edible mushrooms when they are
useful for saprobic as well as mycorrhizal
Chiang Mai J. Sci. 2014; 41(4) 747
o o o
ostreatus (Figure 3) is 25 C, while primordia production is 20–25 C and 25–30 C
formation and fruiting body production respectively [127]. Both mushrooms are
o o
were 10–1 C and 10–17 C, respectively from the same genus; illustrating how
[127]. In the case of P. cystidiosus the suitable knowledge of optimum conditions for media
o
temperature for spawn running is 25-35 C, and spawn running and fructification are
and primordia formation and fruiting body important for growing mushrooms.
The genus Agaricus is particularly yields were obtained. The yield of Thai A.
interesting for study as most species are subrufescens strain was lower than the
edible and others (e.g. A. bisporus, A. French control strain, but had a better
subrufescens) have medicinal properties yield than the Brazilian commercial strain
[221, 222]. In Thailand we have potentially (INRA unpub. data), while A. flocculosipes
discovered more than 70 new Agaricus strains produced mushrooms after 32 days.
species and have isolated several strains This is the first report of cultivation of a
(e.g. A. subrufescens, A. flocculosipes) for wild strain of the A. subrufescens and
fruiting testing. The mycelium was isolated A. flocculosipes collected in Asia (Thailand),
and subcultured on compost agar media and fruiting test as a tool and concluded
and inoculated onto spawn grain media. that it is possible to grow the wild strains
The spawns were inoculated into standard [185, 193].
compost media based on wheat straw mixed Our effort to grow Pleurotus giganteus
with horse manure and various additives has been partially successful [51]. P. giganteus
like rice bran, gypsum and calcium has been shown to have medicinal properties
carbonate. Comparative studies of the [223], and the dry weight protein content
cultivation of the Thai (A. subrufescens and of P. giganteus is 37.8%, which is high in
A. flocculosipes) and French strains of A. comparison to most other cultivated
subrufescens, as a control, were carried out. mushrooms [51, 67]. Our study has also
The mycelium fully covered the compost shown that it is possible to domesticate
within 15 days and a casing layer was local strains of P. giganteus that can grow
applied. The first primordia of the French at temperatures consistent with Thailand
A. subrufescens, and Thai A. subrufescens farm production [51]. However, the yields
strains appeared after 12 days, and 24 days were relatively low and methodology or
after casing, respectively. Significant strain selection needs improving to make
Chiang Mai J. Sci. 2014; 41(4) 749
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