REPRINT OF: Moore, D. & Chiu, S. W. (2001). Fungal products as food. Chapter 10 in Bio-Exploitation of
Filamentous Fungi (ed. S. B. Pointing & K. D. Hyde), pp. 223-251. Fungal Diversity Press: Hong Kong.

FUNGAL PRODUCTS AS FOOD

David Moore and Siu Wai Chiu

School of Biological Sciences, The University of Manchester, U.K. and Department of Biology, The Chinese
University of Hong Kong, Shatin, Hong Kong S. A. R., China

10.1 Summary
Fungi are ideal food because they have a fairly high content of protein (typically 20-30% dry matter as crude
protein) which contains all of the essential amino acids. Fungal biomass is also a source of dietary fibre, and is
virtually free of cholesterol. Mushrooms are cultivated around the world, global annual production being in the
region of 8 million metric tonnes. Agaricus spp. account for something close to 30% of the total. About 60% of
the world’s mushrooms are grown in China. The biggest change during the last quarter of the century has been
an increasing interest in a wider variety of mushrooms. Supplies of fresh mushrooms are now intercontinental
commodities. The only successful fermenter-grown fungal food on the market is the myco-protein Quorn, the
mycelium of a species of Fusarium. Marketing the material emphasises its ability to simulate the fibrous nature
of meat and it is sold as a healthy alternative to meat. Large scale collecting of mushrooms for food has become
an industry in many regions and the commercial picking industry is bound to continue to expand. It raises
several issues, including conservation, ownership, and sustainability of supplies. The key reaction seems to be
effective holistic management of the forest resource.

10.2 Introduction
Fungi are an ideal food because they have a fairly high content of protein (typically 20-30% crude protein as a
percentage of dry matter) which contains all of the amino acids which are essential to human and animal
nutrition. Fungal biomass is easily digested, the chitinous wall provides a source of dietary fibre, and although
filamentous fungi, in contrast to yeasts, have a relatively low vitamin content, they do contain B-vitamins, are
characteristically low in fat. Also, an extremely important attribute of all fungal food is that it is virtually free of
cholesterol. Consequently, fungal protein foods compete successfully with animal protein foods (i.e. meat) on
health grounds. Since, in principle, fungal foods can be produced readily using waste products as substrates,
fungal foods should also be able to compete successfully on grounds of primary cost. In this chapter we will
emphasise the technology, which is often hidden, on which traditional exploitation processes depend. We
emphasise ‘exploitation’ not just usage. Exploitation is the act of successfully applying industry to any object.
This chapter is not a guide for mushroom pickers with notes about tasty subjects and piquant recipes. This
chapter deals with fungi as food on an industrial scale and will indicate how both old and new industries could,
in some cases should, develop in the future. We include some mention of supplementary nutrients and health
products under the heading ‘food’, and will also deal with products which have important roles in processing or
as components of food. Our focus is on the filamentous fungi but we must acknowledge that the yeasts play a
dominant role where biotechnology is applied to the food industry, being essential in brewing and bread making,
and important sources of single-cell protein and dietary supplements. Although these fungal activities are crucial
to human existence (life without bread and wine would be poor life indeed!) and support massive industries
(annual global consumption of ethanol is currently 30 billion litres) they are excluded from this account because
the yeasts concerned are not filamentous fungi. On the other hand, traditional solid state fermentations for
producing mushrooms and other food products and in recent years the Quorn fermentation provide us with a
sufficient range of examples of filamentous fungi being almost equally crucial to human affairs (Table 1).
Judging from archaeological and similar finds, mushrooms, toadstools and bracket fungi have been
used since before recorded history for both food and medicinal purposes. Western recorded fungal history
includes ancient Egyptian murals and tomb ornaments depicting bread and wine making, but probably more
relevant to the present topic is that the Greek first century physician, Dioscorides, wrote that a type of bracket
fungus was effective against cuts and sores, fractured limbs and bruises from falls and was also valuable for
liver complaints, asthma, jaundice, dysentery, kidney diseases and cases of hysteria. Extravagant claims like this
are readily found also in Chinese and Japanese traditional medicine. For example lingzhi (= fruit bodies of the
fungus Ganoderma) is described as ‘the rarest and most precious Chinese medical herb’ which legend claims
even to have ‘... the miraculous power of raising the dead to life ...’.

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Table 1. The most important direct food uses of fungi 

Application  Species 
Cultivated edible macrofungi   

  Champignon, Button mushroom  Agaricus bisporus (= A.

brunnescens), A. bitorquis 

  Shiitake or shiang-gu,  Lentinula edodes 

  Chinese or straw mushroom  Volvariella volvacea 

  Winter mushroom, enoki  Flammulina velutipes 

  Oyster mushroom  Pleurotus sp. 

  Truffle  Tuber melanosporum 

Cheeses   

  Roquefort, ‘blue’ Stilton   Penicillium roquefortii 

  Camembert, Brie, soft ripened

cheeses  Penicillium camembertii 
Oriental food fermentations   

  Ang-kak  Monascus purpurea 

  Hamanatto  Aspergillus oryzae 

  Miso  Aspergillus oryzae, A. sojae 

  Ontjom  Neurospora intermedia 

  Shoyu (soy sauce)  Aspergillus oryzae, A. sojae 

  Tempeh  Rhizopus oligosporus 

Adapted from Wainwright (1992) 

Less extravagant claims are for lingzhi ‘... preventing and mitigating a variety of clinical conditions: chronic
bronchitis, asthma, neurasthenia, insomnia, amnesia, hypertension and hypotension, coronary heart disease,
arrhythmia, stroke, hyperlipidemia, thombosis, female endocrine disorder, female physiological disease,
menstrual disorder, chronic hepatitis, gastric diseases and duodenal ulcer, allergic and chronic rhinitis, dysuria,
arthritis, rheumatism, allergic dermatosis, cancer.’ Now, that quotation does not come from some ancient
medical text, but from a leaflet picked up in a departmental store in Hong Kong in April of 1999! So the
material is being sold now to sophisticated and highly educated people - to the engineer who designed and built
your lap-top computer, or the pilot of your next trans-Pacific flight, for example. And it’s being sold on the basis
of a written medical tradition which goes back more than 5000 years.
That a similarly ancestral, but sadly largely unwritten, tradition occurred in Europe is indicated by
material carried by the Alpine traveller who has become known as ‘The Iceman’ (Peinter, Pöder & Pümpel,
1998). About 3200 BC a Neolithic traveller set out across the Alps. He didn’t make it. Somehow he was caught
in the ice and snow, and died, to be entombed and preserved in the glacier. Eventually, as a result of the
glacier’s slow descent of the mountains, his corpse was exposed at the edge of the ice sheet in 1991 close to the
present Austrian/Italian border. A well-preserved 5000 year old corpse with all of its clothes and equipment is a
remarkable find by any measure. But possibly most remarkable is that there were three separate fungal products
among the Iceman’s equipment. One of these is easy to account for. It was a mass of fibrous material in a leather

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pouch together with flints and a bone tool like an awl. This fungus has been identified as one with a long history
of use as a tinder, so clearly it was part of the Iceman’s fire-making kit. The other two are more problematical.
Both are pieces of a bracket fungus (Piptoporus betulinus) and both are threaded onto leather thongs. One piece
is essentially conical, about 5 cm in its longest dimension, and is on a simple leather thong. The other is
spheroidal, about 5 cm diameter, and is on a thong which has a lobed tassel at one end. These objects were
clearly carefully made and must have been important to the owner to be included as part of the kit he chose to
take with him in his trek across the mountains. Piptoporus is known to produce (and accumulate in its fruit
bodies) antiseptics and pharmacologically active substances which are claimed to reduce fatigue and sooth the
mind. With due ceremony and additional magic, these objects may well have been seen as essential to a traveller
in the mountains. The conical one might be a sort of styptic pencil to be applied to scratches and grazes. Perhaps
the flattened, spheroidal one was chewed or sucked when the going got tough and the tough needed just a little
help to keep going.
Our distant European and Asian ancestors held fungal products in such high esteem that they were
necessary accessories for daily life, including the most hazardous of journeys. Today, alcohol and citric acid are
the world’s most important fungal metabolites in terms of production volume, although, penicillin can still lay
claim to be the most 'important' in social and medical terms. Since the introduction of penicillin, many millions
of chemicals and metabolites have been screened for antimicrobial and other pharmaceutical activities. The
lesson has been well learned and chemicals screening is a major activity of the pharmaceutical and agrochemical
industries around the world. Antibiotics obtained from fungi which are presently of clinical use as antibacterial
agents include the still-important penicillin, cephalosporin and fusidic acid (both of the latter are useful against
penicillin-resistant bacteria), and the antifungal griseofulvin (used to control fungal infections of the skin, nails
and hair). Obviously, antibiotics are the products which come to mind first when thinking of medically-useful
fungal products. More recently, though, several of the mushrooms cultivated in Asia, especially shiitake or
shiang-gu (Lentinula) have been shown to produce materials with antitumour, anticancer, antiviral,
antihypertensive and anticholesterol effects. Indeed, it may be that the consumption of wood ear (Auricularia) is
a contributing factor in the low incidence of atherosclerosis amongst Asians.
Cultivation of mushrooms for food is on the increase, but wild mushrooms have declined disturbingly
in some parts of Europe, possibly as a result of air pollution. Such losses would be of obvious concern to
gourmets, but there are much deeper implications. The species most affected seem to be those that form
mycorrhizas with the roots of forest trees. Dying fungi might mean dying forests. These fungi are more
important than meets the eye!

10.3 Cultivated mushrooms and other fruit bodies

Agaricus bisporus (replaced by A. bitorquis in regions with Mediterranean climates) is by far the most
commonly cultivated mushroom around the world (Table 2; cultivation procedures are described in the next
section). In the mid 1970s the Agaricus crop accounted for over 70% of total global mushroom production.
Today, it accounts for something closer to 30% even though production tonnage has more than doubled in the
intervening years. The biggest change during the last quarter of the twentieth century has been the increasing
interest shown in a wider variety of mushrooms. Even in the most conservative of markets (like the United
Kingdom) so-called ‘exotic’ mushrooms have now penetrated the market and supplies of fresh Lentinula and
Pleurotus are routinely to be found alongside Agaricus in local supermarkets (Fig, 10). Most of these
mushrooms are cultivated fairly close to the point of sale. Otherwise, preserved mushrooms are imported as
canned or dried products, sometimes at a lower retail price. In some markets the demand for fresh mushrooms is
so great that it easily exceeds the ability of the local farming community to satisfy it and the current efficient
transport system for chilled products enables the import of good quality fresh mushrooms. For example, most
UK imported mushrooms originate in the Netherlands or Ireland. In Hong Kong, nearly all the mushrooms are
imported across the border from ‘mainland’ China, from Taiwan and from Japan. The industry is truly
intercontinental, however, and a small supermarket local to DM’s home in south Manchester regularly displays
punnets of fresh enoki (Flammulina) which are grown in Chile. This indicates (a) that intercontinental air
transport makes the approx. 12000 km distance irrelevant and (b) that the production costs are sufficiently low
to enable reasonable pricing in such a distant market. Another reason for the remarkable increases seen in
production of certain mushrooms has been the use of substrates which are waste products from other industries.
For example, oyster mushroom species (Pleurotus ostreatus, P. cystidiosus, P. sajor-caju) are all easily grown
on cotton wastes. Similarly, although the straw mushroom (Volvariella volvacea) is traditionally grown in
South-East Asia on rice straw, it too can be grown on cotton waste. Cotton waste gives higher yields and is also
more widely available than is rice straw so it is a far cheaper substrate (the higher cost of rice straw does not
derive from any intrinsic value but in the cost of transporting it to a non-rice-growing region). Cotton waste
substrates are generated by the textile and garment industries (major industries in the past, and still strong, in
places like Hong Kong and the UK) and are produced in bulk by recycling schemes around the world.

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Table 2. The mushroom cultivation industry over the final quarter of the twentieth century 

  Proportion of total
World production production
(metric tonnes × 1000)  (1991figures)  Production increase
  1976  1991  %  1976-1991 
Agaricus bisporus
and A. bitorquis  675  1590a  37.2  2.4 
Lentinula edodes  130  526b  12.3  4.1 
Volvariella volvacea  49  253  5.9  5.2 
Pleurotus spp.  15  917  21.5  61.1 
Flammulina velutipes  38  187  4.4  4.9 
Auricularia spp. and
Tremella spp.  7  605  14.2  81.8 
Pholiota nameko  15  40  0.9  2.7 
Hericium erinaceus  ?  66  1.5  n/a 
Others incl. Tuber  <1  89  2.1  29.7 

TOTALS  930  4273c  100  4.6 

a
Agaricus spp. global production in 1996 reached 2 million tonnes.
b
Lentinula edodes global production in 1995 was 840,000 tonnes fresh weight of which more than 80% was
produced in China.
c
Total world production of mushrooms in 1996 was estimated to be around 5.8 million tonnes (China
contributing 3.5 million tonnes). 

Disposal of an abundant bulky solid waste coupled with currency earning by sale of a mushroom crop is a good
example of an organic farming system integrated with a waste treatment system. The concept of using
mushroom cultivation as a waste remediation has become a popular model in recent years. All agricultural
production generates enormous waste because so little of each crop is actually used; 95% of the total biomass
produced in palm and coconut oil plantations is discarded as waste, 98% of the sisal plant is waste, 83% of sugar
cane biomass is waste, and in 1995 the global production of cereal straw was estimated at 3.2 × 109 metric
tonnes (Chang, 1998). Pleurotus spp. in particular grow readily on so many lignocellulose agricultural wastes
that it becomes an attractive notion to use the fungus to digest the waste and by so doing produce a cash crop of
mushrooms. Even more attractive is that after the mushrooms have been harvested the ‘spent compost’ can be a
useful animal feed (the mushroom mycelium boosts its protein content), soil conditioner (it is a compost still
rich in nutrients and with polymeric components that enhance soil structure) and even used to digest pollutants
(like polychlorinated phenols) on land-fill waste sites because it contains populations of microorganisms able to
digest the natural phenolic components of lignin (Chiu et al., 1998a). Some care must be exercised in applying
this attractive concept, though, because Pleurotus is also able to accumulate certain metal ions in the fruit body.
If the waste material which is to be used as substrate comes from an industrial source which is likely to be
contaminated by ‘heavy metals’ then the mushroom crop may be unsuitable for consumption. For example,
Chiu et al. (1998b) showed that cadmium could be accumulated in Pleurotus fruit bodies to such high levels that
a single modest serving of mushrooms could cause the consumer to exceed the tolerable food limit
recommended for a full week of intake of this metal.
Truffle cultivation is rather different from that of the other fungi listed in Table 2 which can be grown
intensively in mushroom farms. The truffle, however, is the underground fruit body of an ascomycete that is
mycorrhizal on oak (Quercus), so it is dependent on its host tree. The truffle develops a few centimetres below
the soil surface as an irregularly shaped, 2 - 8 cm diameter fruit body which has an outer rind of tissue
protecting an inner mass of ascospores. There are almost seventy different species of truffle, and since they are
mentioned in ancient Greek texts so they must have been collected for over 2000 years. The most highly prized -
the ‘diamond of French cuisine’ - is the black truffle of the Périgord (actually more common in Provence),
Tuber melanosporum. Only 200 tons are harvested every year in France; they sell for nearly 3,000 francs per
pound on average. Traditionally, truffles are found using pigs or dogs trained to detect the volatile metabolites
produced by the fruit body. Truffle ‘cultivation’ was first achieved early in the nineteenth century when it was
found that when seedlings adjacent to truffle-producing trees were transplanted, they too began producing

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truffles in their new location. ‘Truffières’ or truffle groves have been established throughout France in the past
hundred years and the value of the crop is such that the practice is now extending around the world. Truffières
are started by planting oak seedlings in areas known to be infested with truffle fungi. The truffles begin to
appear under such trees 7 to 15 years after planting and cropping will continue for twenty to thirty years. Most
plants infected with Tuber melanosporum are now raised in greenhouses although pure cultures of this species
cannot be used yet to inoculate the roots of oak seedlings. Recently, methods have been developed to colonise
plant roots with the related T. magnatum (a white truffle), encouraging the hope that the same might be done
with other truffle species.
A cultivated mushroom which does not appear in Table 2 is Ganoderma lucidum, even though the
global production in 1997 was about 4,300 metric tonnes (about 3,000 tonnes of which were grown in China).
Ganoderma, however, is unique in being consumed for its pharmaceutical value (real or imagined) rather than
as a food. Under the names lingzhi or reishi, several Ganoderma spp. of the G. lucidum complex provide
various commercial brands of nutriceuticals, in the form of health drinks, powders, tablets, capsules and diet
supplements. Ganoderma is highly regarded as a traditional herbal medicine (some of the claims made for it
have been outlined above), and its popularity in China has spread to other Asian countries, and also to the wider
world. Current research is focussed on purification and characterization of the bioactive components and
determination of clinical value, especially putative anti-tumour and anti-aging properties. As a nutriceutical,
though, it is strictly outside the scope of this chapter. It is cultivated by being inoculated into short segments of
wooden logs which are then covered in soil in an enclosure (often a plastic-covered ‘tunnel’) which can be kept
moist and warm. The fruit bodies then emerge in large number quite close together and the conditions encourage
the fungus to form the desirable long stemmed fruit body.

10.4 The mycelium as food

The emphasis given in the 1950s to the 1970s to the production of single-cell protein (SCP) is now almost
forgotten. The hope that microbes could provide a means of solving the world’s food shortage by industrial
production of cheap protein alternatives to meat protein were dashed by the realisation that resolution of
problems on that scale has more to do with politics and economics than with biotechnology. Today, the
emphasis has moved towards the use of wastes as substrates for microbial biomass production for animal feed
and towards production of novel ‘health’ foods for sale in advanced economies. Fungal SCP must compete with
established animal feeds like soya meal, the price of which can be kept artificially low for long periods. Any
novel microbial protein that requires costly research and development and/or unusual and expensive production
facilities will not to be able to compete. The two most successful applications for fungal protein in the food
industry illustrate the different ways of coping with this economic fact of life. One example is the conversion of
brewery wastes, using low-technology processes into flavourings, diet supplements and products like Marmite
and Vegemite. Of course, these are yeast products but the example illustrates one successful model - start with a
cheap (ideally, a waste-material) feedstock and use conventional production processes to make a product that
sells at a relatively high retail price. The oriental example is the production of Monascus products which can be
obtained from submerged fermentation of food processing wastes and were valued at over US$2 million in the
Japanese market in 1992. The alternative successful model with a fungal product currently on the market is the
myco-protein Quorn. This is the mycelium of a species of the filamentous soil-fungus Fusarium. Production
details are given below but the mycelium is grown in a very large air-lift fermenter in a continuous-culture
mode. It is consequently a high-technology product. The market virtues of the material centre around its
filamentous structure which enable it to simulate the fibrous nature of meat (Fig. 12). Coupled with the inherent
nutritional value of fungal biomass, this permits the product to be sold as a low-fat, low-calorie, cholesterol-free
health food to consumers who can afford to choose Quorn as a meat substitute. At a price. The retail prices of
Quorn and some other foods in the south Manchester supermarket to which we have already referred are shown
in Table 3. This little survey illustrates the difference between market value and market price. Quorn is more
expensive than most meat products (and most mushrooms), but it is sold as a ‘meat alternative’. Evidently, it is
not positioned in the market to be sold to consumers who cannot afford meat, but rather to those who can afford
to pay a premium price for a health food. The retail price of Quorn reflects the perceived value created by the
marketing, not the intrinsic value of the product. On the other hand, the retail price of mushrooms is their
intrinsic value because severe competition in this sector of the market ensures that retail price very closely
reflects actual costs of production, distribution and sale.
With a view to future product developments there is a good deal of research under way on growing
fungi such as Trichoderma sp. on cheap cellulosic materials, mainly agricultural wastes. Fungi can also be used
to improve the nutritional quality of cereals like barley to compensate for the latter’s deficiency in the amino
acid lysine. Inoculating soaked barley with Aspergillus oryzae or Rhizopus arrhizus increases the protein
content as the fungus grows. The product is used as pig food.

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Table 3. Retail price comparisons of fungal food products and other equivalents 

Product  pack size pack price price per comments 

(g)  (£)  100 g (£) 
Quorn mince  350  2.99  0.86   

Quorn pieces  350  2.99  0.86   

Unwashed ‘value’ pack, grown

750  1.49  0.20 
locally in UK 
Agaricus white mushrooms  500  1.29  0.26  closed-cups, grown locally in UK 
‘Baby-buttons’, grown locally in
150  0.99  0.66 
UK 
Agaricus brown cap sold loose, imported from Ireland 
1000  3.50  0.35 
mushrooms 
Oyster mushrooms grown locally in UK 
125  0.99  0.79 
(Pleurotus sp.) 
Shiitake mushrooms sold fresh, grown locally in UK 
100  1.59  1.59 
(Lentinula edodes) 
sold fresh, imported from Italy,
Girolle mushrooms
125  1.79  1.43  presumably collected, not
(Cantharellus cibarius) 
cultivated 
Beef, mince  500  1.03  0.21 
Beef, diced  750  4.99  0.67 
Pork, ‘casserole’ style  500  1.99  0.40 
Pork, ‘stir-fry’ style  340  2.49  0.73 
Lamb, diced  400  3.69  0.92  Fresh meats, all UK produce 

Lamb, diced (‘organic’)  400  4.59  1.15 

Chicken, diced skinless  400  2.99  0.75 
Turkey, diced  300  0.99  0.33 
Turkey, ‘stir-fry’  600  3.99  0.67 
Sold frozen and bread-crumbed,
Fish, ‘Cod nuggets’  600  1.49  0.25  North Atlantic origin but probably
landed in UK 
Fat-reduced, cheddar-style
Cheese ‘medium fat’  554  3.54  0.64 
‘industrial’ cheese 
Prices shown in £-sterling. Survey done 5 September 1999 at Tesco, East Didsbury, Manchester. 

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10.5 Fermented foods
As well as being used directly as food, fungi are also used in the processing of various food products. In these
applications the fungus is primarily responsible for the production of some characteristic odour, flavour, or
texture and may or may not become part of the final edible product. Growing filamentous fungi on water-soaked
seeds of plants is the basis for production of several human food products in Asia, including soy sauce and
various other fermented foods. In soy sauce production soybeans are soaked, cooked, mashed and fermented
with Aspergillus oryzae and A. sojae. Depending on the size of the factory, the soybeans may be fermented in
fist-sized balls (the traditional method) or on trays. When the substrate has become overgrown with the fungus
the material is mixed with salt and water and the fermentation is completed in the brine. The biggest industrial
units today use a continuous process in which defatted soybean flakes, moistened and autoclaved are mixed with
ground, roasted wheat. The mixture is turned mechanically to ensure even growth of Aspergillus oryzae and A.
sojae for two to three days; then it is transferred to brine and inoculated with Pediococcus halophilus and 30
days later with Saccharomyces rouxii. The brine fermentation takes six to nine months to complete, after which
the soy sauce is pressure-filtered, pasteurised and bottled.
Indonesian tempeh is a white cake produced by fermentation of partially cooked soybean cotyledons
with Rhizopus oligosporus. The fungus binds the soybean mass into a protein-rich cake that can be used as a
meat substitute which is being increasingly widely sold into the vegetarian market. There are a variety of other
fermented products of this sort. Ang-kak is a rice product popular in China and the Philippines which is
fermented using Monascus species. Monascus purpureus produces the characteristic pigments and ethanol
which are used for red rice wine and food colouring. The pigments are a mixture of red, yellow and purple
polyketides and about ten times more pigment is obtained from solid state fermentation than from submerged
liquid fermentation.
Cheese could be considered the occidental equivalent of the fermented soya products which are popular
in Asia. Cheese is a solid or semisolid protein food product manufactured from milk. Before the advent of
modern methods of food processing, like refrigeration, pasteurisation and canning, cheese manufacture was the
only method of preserving milk. Although basic cheese making is a bacterial fermentation, there are two
important processes to which filamentous fungi contribute; these are the provision of enzymes for coagulation
and mould-ripening. Cheese production relies on the action of enzymes which coagulate the proteins in milk,
forming solid curds (from which the cheese is made) and liquid whey. Traditional cheese-making uses animal
enzymes, specifically chymosin and pepsin, extracted from the stomach membranes of unweaned ruminants.
Rapid expansion of the cheese-making industry caused attention to shift to alternative sources of such enzymes
and moulds like Aspergillus spp. and Mucor miehei have supplied these to the extent that around 80% of
cheesemaking now uses non-animal coagulants. Very recently, animal enzymes produced by genetically-
modified microbes have entered the market, but for the moment most industrial cheese production still depends
on enzymes from filamentous fungi for the coagulation step. Mould ripening is another matter, being a
traditional method of flavouring cheeses which has been in use for at least two thousand years. Blue cheeses,
like Roquefort, Gorgonzola, Stilton, Danish Blue, Blue Cheshire, use Penicillium roquefortii which is
inoculated into the cheese prior to storage at controlled temperature and humidity. The fungus grows throughout
the cheese, producing methyl ketones, particularly 2-heptanone, as the major flavour and odour compounds.
Camembert and Brie are ripened by Penicillium camembertii, which changes the texture of the cheese rather
than its flavour. This fungus grows on the surface of the cheese producing extracellular proteases which digest
the cheese to a softer consistency from the outside towards the centre.

10.6 Industrial cultivation methods

On all criteria, the biggest (non-yeast) biotechnology industry in the world is mushroom cultivation. Most
definitions, however, take the much narrower view that industrial microbial cultivations, the very stuff of
biotechnology, are submerged cultivations done in stirred tank reactors (Fig. 8). These are popularly thought of
as the 'real' fermentations, where the activities of the microorganisms generate a vigorous ferment within the
medium. However, if brewing activities are put to one side (on the grounds that they exploit a yeast rather than a
filamentous fungus) these aspects of biotechnology are only a very small fraction of the whole - whether
calculated in terms of value of product, mass of product, number of people involved in the industry, or
geographical area over which the industry is practised. By far the greatest level of exploitation of filamentous
fungi uses cultivation in the absence of free-flowing water - the so-called solid state fermentations.

10.6.1 Solid state fermentations

The mushroom industries of the world all depend on some form of solid state fermentation. In the European
tradition (Fig. 9) this has come to mean cultivation of a mushroom crop on composted plant litter. Similar
approaches were developed for Oyster and Paddy straw mushrooms in the Orient, though in the Chinese
tradition (Fig. 10) the typical approach is to cultivate the crop of choice (Lentinula) on wood logs.

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10.6.1.1 Commercial mushroom production in the European tradition

The European mushroom industry is said to have originated in the caves beneath Paris at the end of the
nineteenth century. It probably emerged from the food provisioning functions of the kitchen gardens on the
estates of the European aristocracy. Some of the surviving records of such estates refer to manured and
composted plots set aside for mushroom production. The compost used, and its preparation, would very
definitely be familiar to the very competent gardeners of the day. The current industry depends on a compost
which is very selective for the crop species Agaricus bisporus. Although widely distributed in nature, this
fungus is rarely noticed because established mycelia produce relatively few mushrooms, and them only
infrequently. The industry we know today seems, therefore, to be the result of a remarkable 'joint evolution'
during which an otherwise ordinary horticultural compost was developed that achieves high cropping densities
with an otherwise unremarkable and not very abundant mushroom. And all without a genetic engineer in sight!

10.6.1.1.1 Composting
Good compost is the essential prerequisite for successful mushroom farming. For Agaricus farming, compost
preparation is a smelly process because a nitrogen-rich animal waste is included. Even the most modern
installations have a severe impact on their neighbours! The human nose can sense 10 ppm ammonia but the
ammonia emitted from a fermenting compost can reach 600 to 1000 ppm. To make mushroom composting
environmental friendly, the air from compost producers must be collected and treated chemically and
biologically to remove the generated pollutants (ammonia and sulfur-containing compounds). The basic raw
material for mushroom compost in Europe is wheat straw, although straws of other cereals are sometimes used.
Ideally, the straw is obtained after it has been used as stable bedding and is already mixed with horse manure.
On commercial scale this is not possible and other animal wastes, like chicken manure, are mixed with the
straw, together with gypsum and large quantities of water. The use of gypsum (calcium sulphate) in the
preparation of compost was generally introduced in the late 1930s. Excess calcium reacts with, and precipitates,
the mucous and slimy components of manure and so prevents water logging of the compost and generally
improves aeration and, most importantly, the mechanical properties which aid thorough mixing. Its introduction
resulted in large crops being grown reliably for the first time because uniformly digested compost could be
produced reliably for the first time.
Composting proceeds in two phases. In phase 1 the straw, manure and other components are mixed into
large heaps. After the water is added the heaps are thoroughly mixed by mechanical compost turning machines.
This ‘pre-wetting’ treatment continues for a few days and then the machines arrange the compost into long
stacks about 2 m wide, 2 m high and many metres long. Within a few days the bacterial activity heats the stack
to around 70 ΕC in the centre, though it is considerably cooler at the surface. Higher temperatures, which would
kill the microorganisms, are avoided by regular ‘inside-out’ turning of the compost heap. As well as heat, the
bacterial degradation process releases large amounts of ammonia. An important aim in phase 1 of composting is
to achieve uniformity by thorough mixing (so that all of the compost spends some time within the hotter core of
the stack). A week after the stack was first laid it is mixed, or 'turned', by large, self-propelled ‘turning’
machines (Fig. 1). It is left for a further week, then turned again. Three weeks after the process was started, the
compost is ready for phase 2.
This, also known as peak-heat, pasteurisation or sweat-out is a continuation of the composting process
but without further mixing and under more controlled conditions. The compost may be treated in bulk or loaded
into the eventual growing containers. In either case the process is done in a building which allows air to be
circulated around the growing containers or through the bulk of the compost. To begin with, air and compost
temperatures are raised to about 60 ΕC for several hours. This pasteurisation stage is usually completed in a day
and then the amount of ventilation is increased and compost temperature is kept at about 50 ΕC for 4 - 6 days.
The beds are then allowed to cool to around 25 ΕC and are ready for use. Natural drop in temperature and
absence of free ammonia are signs that the composting process has been completed.

10.6.1.1.2 Spawning
Spawning is the process that introduces the mushroom mycelium into the compost. This is generally done with
some form carrier that can be easily-mixed into the compost, fungus-coated cereal grains (often barley) being
the most usual. About 5 kg spawn per tonne of compost (= 0.5% by weight) is used. From these inoculation
centres the mycelium grows out to invade the compost (= 'spawn running'), filling the compost bed after 10-14
days at a compost temperature of 25 ΕC. Slow-release nutrients might be added at the casing step or separately.

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10.6.1.1.3 Casing
To encourage fruiting of Agaricus, the spawn run compost must be covered with a 'casing layer' - originally of
soil but now most usually a mixture of moist peat and chalk, the chalk being used to adjust the otherwise acid
pH to a neutral one. Casing is needed only by Agaricus, the procedure is not necessary when cultivating other
species such as Volvariella spp., Pleurotus spp., Auricularia spp. and Lentinula edodes. The optimum depth of
the casing is 3 - 5 cm and it should be an even layer applied to a level compost surface. The mushroom
mycelium grows into the casing layer in similar conditions to those for spawn running. But the mycelium
reaches the upper surface of the casing layer as strands - a necessary start to the fruiting process. To encourage
completion of fruiting the growing room is ventilated to lower the concentration of carbon dioxide (usually to
<0.1%) and to help reduce the temperature to 16 - 18 ΕC. Throughout these steps the casing layer must be kept
moist by mist-spraying with water at intervals. Moisture, temperature and atmospheric gases all have to be
stringently controlled to match the requirements of the particular mushroom strain being cultivated. After
allowing 7 to 9 days for the Agaricus mycelium to grow into the casing layer, a machine with rotating tines is
run across the mushroom bed to mix the casing layer thoroughly. This is called ‘ruffling’ and it serves to break
up the mycelial strands and encourage the mushroom mycelia to grow and colonize the surface of the casing
layer.

10.6.1.1.4 Cropping
A few days after ruffling, the sudden change in microclimate sensed by the mushroom mycelium on the surface
of the casing soil triggers the formation of mushroom primordia in Agaricus. These beginnings of mushroom
fruit bodies, called 'pins' or 'pinheads' which are more or less spherical and have a smooth surface, will be seen
about 7 to 10 days after casing (Fig. 2). It will be 18 to 21 days after casing before marketable mushrooms can
be harvested (Figs 3, 11). Successions of mushrooms then develop in a series of flushes about 8 days apart, and
each taking about 5 days to clear from the beds. During the cropping period, the casing needs to be kept moist
and the air temperature must be maintained in the 16 to 18 ΕC range. Ventilation must also be maintained to
keep carbon dioxide levels low. Accurate balance is required here: humidification is essential to minimise
dessication, but too high a level of humidity encourages disease.
Growers expect to harvest between three and five flushes from each spawning cycle, with a total yield
of around 25 kg m-2 of growing tray. After the final pick (seven to ten weeks after spawning) the compost is
spent, and the cropping room is emptied, cleaned, sterilised and filled with the next crop. On most large
commercial farms a new crop is filled every one or two weeks throughout the year. So a mushroom farmer is
likely to see more crops in one year than a cereal farmer will see in a lifetime!

10.6.1.1.5 Cropping and Service Systems

Although the basics of mushroom production are the same however the crop is produced, growing containers
differ and the process can be separated into specialised stages. Mushrooms may be produced in large wooden
trays, in beds on shelving and in plastic bags. Trays, made of wood in sizes varying from 0.9 × 1.2 m to 1.2 ×
2.4 m and 15 to 23 cm deep, are arranged in tiers 3 or more high, separated by wooden legs. Fork lift trucks are
needed to move the trays and some sort of mechanised tray handling line is necessary. Shelving is usually made
of metal and arranged to give four to six layers of fixed shelves in a cropping room with centre and peripheral
access gangways. Each shelf is about 1.4 m wide and extends almost the whole length of the room. Special
machinery for compost filling, emptying, spawning, casing and other cultivation operations is necessary.
Growing bags of about 25 kg are usually supplied to the farm already spawned and may be arranged on the floor
of the cropping house or on tiered shelving. Each arrangement makes its own demands on techniques and
equipment. It is also often the case that the outdoor stages of composting are completed by a specialist compost
producer and the ready-to-use compost delivered in bulk to a mushroom farm. Indeed, the compost may even be
delivered ready spawned so that the farm process starts with spawn running. Nowadays, the mushroom
production industry comprise spawn makers, phase I compost, phase II compost and phase III compost
suppliers. Phase III compost is completely colonized by the mushroom mycelia which if placed in a suitable
environment in a mushroom farm will produce the fruit body crop readily. Cardboard boxes of phase III
compost are now widely available as ‘home grown mushroom kits’. For a commercial mushroom farmer, the
use of phase I compost gives the most flexibility to optimize farm conditions for cultivation of any mushroom
strain. Purchase of phase II compost enables a farmer to choose which mushroom strain to spawn. The use of
phase III compost, though it is obviously more costly, guarantees the production of a crop in a short time and
requires the least prior investment in facilities for mushroom production.

The production of Pleurotus (oyster) mushrooms is a sharp contrast with the approach just described
for Agaricus because the needs of the organism are much less stringent. This goes some way to explaining the
rapid boom in oyster mushroom farming from the late 1980s to the present day. Both composted/pasturized and

  9
  
sterilized, but uncomposted, substrate of a wide range (sawdust, wheat straw, etc.) can be used. No casing is
required. The crop can be adapted to different countries depending on their climates by growing different
species of oyster mushrooms, e.g. Pleurotus pulmonarius (misnamed as P. sajor-caju) in India; P. ostreatus
(commercially called P. florida - another inaccurate name) in Europe.

10.6.1.2 Commercial mushroom production in the Asian tradition

Lentinula edodes is traditionally grown on evergreen hardwood logs (oak, chestnut, hornbeam) and is still very
widely grown like this in the central highlands in China. To put this statement into perspective, the traditional
log-pile approach is still the most frequently used method in China over a growing region which covers an area
about equal to the entire land area of the European Union. The logs considered suitable for shiitake production
are over 10 cm diameter and 1.5 to 2 m long. The logs are normally cut in spring or autumn of each year to
minimise pre-infestation by wild fungi or insects. Holes drilled in the logs (or saw- or axe-cuts) are packed with
spawn, and the spawn-filled hole then sealed with wax or other sealant to protect the spawn from weather. The
logs are stacked in laying yards on the open hillside in arrangements which permit good air circulation and easy
drainage and provide temperatures between 24 Εand 28 ΕC (Fig. 4). The logs remain here for the five to eight
months it takes for the fungus to grow completely through the log. Finally, the logs are transferred to the raising
yard to promote fruit body formation. This is usually done in winter to ensure the lower temperature (12 - 20
ΕC) and increased moisture which are required for fruit body initiation. The first crops of mushrooms appear in
the first spring after being moved to the raising yard. Each log will produce 0.5 to 3 kg of mushrooms, each
spring and autumn, for 5 to 7 years.
This traditional approach to shiitake production is expensive and demanding in its use of both land and
trees. Some commentators estimate that there are 10 million mushroom farmers in China; if true, the traditional
use of locally-cut logs is likely to devastate the hill forests. For this and other reasons more industrial
approaches are being applied to shiitake growing. Hardwood chips and sawdust packed into polythene bags as
'artificial logs' provide a highly productive alternative to the traditional technique (Fig. 5), and the cultivation
can be done in houses (which may only be plastic-covered enclosures) in which climate control allows year-
round production.
The straw mushroom (Volvariella volvacea) is grown mainly on rice straw (Fig. 6), although several
other agricultural wastes make suitable substrates. Preparation of the substrate is limited to tying the straw into
bundles which are soaked in water for 24 to 48 hours. The soaked straw is piled into heaps about 1 m high
which are inoculated with spent straw from a previous crop. In less than one month, a synchronized flush of
egg-like fruit bodies appear. These immature fruit bodies (in which the universal veil is intact and completely
encloses the immature fruit body, Fig. 7) are sold for consumption just like the young fruits (‘baby buttons’, Fig.
11) of Agaricus (though this is not the case with oyster and shiitake mushrooms which are sold mature).
Comparatively low yields of V. volvacea are generated from the substrate, and it is difficult to maintain a good
quality in post-harvest storage. Within 2-3 days the crop turns brown and autolyses even in cold storage. These
factors restrict production of the crop.

10.7 Quorn production

Myco-protein is the term coined by the UK Foods Standards Committee to serve as the generic name for a food
product resulting from the continuous fermentation of a selected strain of Fusarium venenatum (originally called
F. graminearum). The material is produced by Marlow Foods Limited, a joint venture set up in 1984 between
RHM (Ranks Hovis McDougall) and ICI. The product was originally intended to be dried and powdered for sale
as a high-protein SCP flour but the organoleptic qualities of the hyphal mass were recognised and it has been
developed as a meat substitute under the brand name Quorn.
The fungus is grown on food-grade glucose as the carbon source, usually derived from wheat or maize
starch, and a 45 m tall airlift fermenter is used in continuous culture mode (i.e. medium is added continuously
and mycelium + spent medium is harvested at a rate equal to the production of new hyphae) (Trinci, 1992).
Fast-growing fungal cells contain 8 to 9% RNA (w/w) but humans are intolerant to high levels of nucleic acids
because they are metabolised to the sparingly-soluble uric acid which can accumulate in the joints (in other
vertebrates the much more soluble allantoin is the end-product of nucleic acid catabolism). SCP intended for
human consumption must therefore be processed so that the recommended maximum daily intake of nucleic
acid (4 g) is not easily exceeded. Immediately after harvesting the temperature of the biomass is raised (in less
than 10 sec) to 64ΕC and held for 20 to 30 min. This stops growth, disrupts ribosomes and activates endogenous
RNAases which digest cellular RNA to nucleotides which then diffuse out of the tissue. Loss of protein and
fibrous structure is minimised though about 30% of the dry weight is lost in the treatment. The RNA content is
reduced to about 1%, well within the 2% upper limit recommended by the World Health Organisation. After
RNA reduction, the mycelium is harvested on a continuous belt filter to give a raw product containing about
30% solids. Myco-protein typically contains 44% protein, 18% dietary fibre and only 13% fat (the values for

  10
  
beef are 68%, 0% and 30% respectively)(and see Table 4).

Table 4. Nutritional comparison of Quorn products with their meat equivalents 

    Skinless Quorn Breaded Low Fat Low Fat

Quorn Quorn Beef Quorn
    Chicken Southern Chicken Beef Pork
Pieces  Burgers Burgers Sausages
Per 100g  Units  Pieces  Burgers Burgers Burgers  Sausages

energy  (kJ) 355.0  621.0 746.0 1128.0 490.0 1192.0 777.0  491.0  728.0
energy  (kcals) 85.0  148.0 178.0 270.0 117.0 287.0 186.0  115.0  174.0
Protein  (g) 12.3  24.8 10.7 12.9 12.8 15.0 17.6  13.5  13.0
Carbohydrate  (g) 1.8  0.0 12.3 17.1 5.8 3.5 3.7  5.2  9.6
of which: sugars    0.8  0.0 1.5 0.5 2.5 0.7 0.7  1.0  0.4
oil/Fat  (g) 3.2  5.4 9.6 16.7 4.6 23.8 11.3  4.7  9.3
of which: saturates    0.6  1.6 1.2 6.5 2.3 10.0 4.8  2.8  3.5
Fibre  (g) 4.8  0.0 3.1 0.6 4.1 0.4 trace  3.4  1.6
Sodium  (g) 0.2  0.1 0.6 0.5 0.5 0.5 0.5  0.6  0.9

The table shows typical values for the same amount of six meal types. The data were obtained from the Marlow Foods
Limited web site (http://www.quorn.com/) which states that the information comes from official sources. Quorn pieces
are fully textured but only mildly flavoured so they are close to representing the basic myco-protein. Note how the
composition depends more on the product recipe than on the myco-protein. 

The sole use for myco-protein at the moment is as the primary ingredient of the Quorn range of ‘meat-
alternative’ products. The Fusarium biomass is mechanically processed to align the hyphae and this alignment is
‘set’ by mixing with small amounts of a binder, such as egg albumin, which is coagulated by heating to stabilise
the filaments. This mimics the muscle fibre and connective tissue structure of meat. Quorn is marketed as ‘The
tasty, healthy, alternative to meat ...’ and the makers claim ‘...It is mushroom in origin and provides the taste and
texture of a full range of meat products and ready meals but is entirely meat free...’ (see Marlow Foods Limited
web site at http://www.quorn.com/). The web site glossary describes myco-protein, more accurately, as ‘The
harvested filamentous cells of a distant relative of the mushroom family (Fusarium species (Schwabe)
ATCC20334) grown by a continuous fermentation process.’
The simplest forms of myco-protein which are marketed are Quorn mince and Quorn pieces intended
for use in home cooking recipes in much the same way as meat products. A widening variety of products is
emerging, which now include: Quorn burgers, sausages and fillets; Quorn ready meals; and the ‘Quorn Deli’
range of cold cuts - sliced substitute meats for salads, sandwiches or snacks (Fig. 12). The main thrust of the
marketing for all of these products is ‘healthy eating’ - the headlined ‘Quorn product facts’ being: low in fat,
good source of protein, good source of dietary fibre, low in calories, cholesterol free. The absolute content of the
various components in the retail product depends more on the process recipe than the myco-protein (Table 4).
Given that the sparsely-branched filamentous structure of Fusarium is crucial to the creation of the
meat-like texture of Quorn, it could be argued that the Quorn fermentation is the only true example we have in
which it is the filamentous character of the fungus that is exploited. Although this is the only liquid
fermentation which yields fungal biomass as the sole food product, there are several others which are used to
produce fungal metabolites. Monascus pigments (Fig. 8) and ethanol, and enzyme production by Aspergillus
and Mucor (in relation to cheese production) have been mentioned earlier. Many other enzymes produced by
filamentous fungi in fermentation cultures are used for processing foods. For example, amylases ( from A. niger,
A. foetidus, Rhizopus foetidus) are used to convert starchy substrates to sugars prior to alcoholic fermentation,
and also to make chocolate syrups from cocoa, and invertase (from Aspergillus oryzae, A. niger) is used for
sucrose conversion in confectionery. Some other industrial uses for fungal enzymes include proteinases and
lipases from Aspergillus oryzae used in detergents and proteinases used in hide processing in the tanning

  11
  
industry. The global value of enzymes from filamentous fermentations is in the region of US$ one billion. The
single most important fermentation metabolite is citric acid, 300,000 tonnes of which are produced (mostly by
Aspergillus niger and A. wentii) each year to be used mainly in effervescent soft drinks. The standard production
method is a fermentation of sugar beet or cane molasses or a glucose syrup. Important though these liquid
fermentations are, in all cases the enzyme or metabolite product is purified from the fermentation liquor (in
some cases the enzyme is extracted from the biomass), so the fungus itself is not a component of the final
product.

10.8 Wild harvests

Collecting mushrooms for food is an age-old tradition which is on a par with collection of berries and other
forest fruits. It becomes an exploitation when the collection becomes an industry which is pursued in order to
supply a commodity.

10.8.1 Commercial mushroom picking

Several wild mushrooms have reached the ‘exploitation’ state. Masutake, chanterelles, morels and truffles
probably represent the best expression of this in the popular imagination because of the history and mystique
associated with the industries in their home countries; in Europe, particularly in France and Italy. These histories
include festivals and markets which associate folk events and heighten interest in and appreciation of the
qualities of the products themselves. Those qualities contribute to the mystique, prompting discussion of how
tasty and flavourful competing collections might be. The truffle probably has more mystique than most because
this subterranean delicacy is still hard to find and harvest so there is the added mystery of detecting the presence
of a truffle ten or twenty cm underground with the help of forest lore, flies, or trained pigs and dogs. There are
about seventy different truffle species. The most highly prized in French cuisine is the black truffle of the
Périgord, Tuber melanosporum, but in Italy it is the white truffle of Alba which is considered the true delicacy.
Deciding these matters creates lots of fun and innocent enjoyment. And makes lots of money. The world market
for chanterelles (collected, not cultivated) was estimated recently at more than 1.5 × 109 US$ (Watling, 1997).
Add the value of tourism and peripheral matters like cookery programmes on TV and recipe books and
magazine articles and the collection and appreciation of these fungi becomes a very big industry indeed.
Unfortunately, mycorrhizal species like these have shown a significant decline in fruit body formation in recent
years (Arnolds, 1991, 1995). The widened and increasing market and heightened commercial demands for forest
fungi are appearing at a time when the woods and forests are under pressure from many other dangers like
urbanization, global warming, deforestation and industrial pollution.
Although mushrooms have been important sources of food and medicine for a great many years, the
demand for wild mushrooms has grown sharply since about the early 1980s. Speciality mushrooms have always
been harvested and shipped to distant markets. There is a story that as early as 1872 New Zealand had a fungus-
based industry earning, eventually, hundreds of thousands of pounds annually by collecting the wood ear fungus
(Auricularia polytricha) for sale in China. This fungus trade was initiated by a New Plymouth merchant, Chew
Chong and provided much-needed income for New Zealand’s settlers.
The then Colonial Secretary of Hong Kong, in answer to an inquiry from the New Zealand Colonial
Secretary in 1871, reported that the fungus was used as a medicine “...much prized by the Chinese
community...” It was also used as a food, 'mu-erh', eaten with noodles and bean curd, as well as in soups. Chong
paid colonial farmers four pence per pound weight of sun-dried fungus and is said to have purchased an average
of £65-worth on each New Plymouth market day (equivalent to 1.8 metric tonnes!). Retail prices in the Hong
Kong market were four to ten times that. This trade in New Zealand wood ear was maintained until the 1950s,
eventually being effectively killed off when commercial cultivation of Auricularia made collection from natural
sources uncompetitive. This story emphasises the rural ideal of collection of a cash crop by local residents for
sale elsewhere. Generally small and scattered enterprises where pickers sell and ship most of their own harvest.
The trade may be seasonal, and the volume of mushrooms picked may be relatively small.
All of this changed during the late 1980s, mainly in the amount and manner in which mushrooms were
harvested, sold, and handled. The commercial picking industry has now expanded to a system of harvesters,
buyers, processors, and brokers. Harvesters locate and pick the mushrooms. Buyers, typically associated with a
specific processor, set up buying stations near wooded areas known to produce mushrooms and advertise their
willingness to buy. Processors grade, clean, pack, and ship the product while providing the cash directly to the
field workers. Brokers market the mushrooms around the world. This is a model which has become common in
Europe and the United States. One of the things that makes it viable is the easy access to rapid trans- and
intercontinental transport. As transport and communications continue to improve, the commercial picking
industry is bound to continue to expand.

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10.8.2 Regulate or be damned
In most of the areas in which commercial mushroom pickers operate the operation is seen as a 'problem', though
the exact perception of what the problem consists of depends very much on the standpoint of the observer. In
any one region there may be thousands of pickers harvesting fungi for commercial purposes from both private
and public lands. The knowledge we have about the ecology of wild edible mushrooms is incomplete and this
ignorance is at the centre of the conflicts which are arising between commercial pickers and conservationists
and local residents. The three parties do not always align as might be expected. A successful commercial
picking job can see a region of woodland completely denuded of marketable mushrooms in just a few hours.
Local residents see this as destruction of a natural resource which 'belongs to the people' and expect the support
of conservation-minded mycologists in the campaigns which result. Unfortunately, ownership of the resource is
not always entirely transparent. 'The people' may be allowed to enjoy a woodland for what it is by a generous
landowner who subsequently is enlightened to the cash value of its mushroom crop. Similarly, it is not at all
clear that picking mushrooms does any damage that a conservationist should be concerned about. In the UK the
activities of commercial pickers have been likened to the activities of factory fishing boats which take fish of all
ages and consequently damage the breeding stock of the fishery to the point where the fish population declines
catastrophically. This is an emotive comparison for the UK, which has experienced a catastrophic decline in its
own fishing industry, but it has almost no biological relevance to commercial mushroom picking. Mushrooms
are not individuals, but simply the fruiting structures arising on underground mycelia. Removing one generation
of fruit bodies will probably encourage a new generation to emerge. Certainly, continued productivity of
mushroom farms beyond the first flush of cultivated mushrooms is enhanced by regular harvesting.
What must be safe-guarded, of course, is the health of the mycelium and so there probably is a need for
regulation aimed at avoiding unnecessary trampling and disturbance. But this is not the sole need for regulation.
The USDA has studied commercial mushroom picking in forests in the North American continent, highlighting
the issues in several reports (Molena et al., 1993; Pilz & Molina, 1996; Hosford et al., 1997). These reports
make it clear that mushroom harvesters range from the curious recreational collector, via temporarily
unemployed workers needing additional income to highly skilled and almost professional commercial pickers.
The ranks of the latter often include individuals who work in forestry supplementing income during periods of
slack employment by collecting and selling mushrooms. Mushroom harvest dollars are often part of the 'black
economy' in which much of the cash value is not reported. Even if unreported, harvest dollars benefit local
communities.
Even remote areas may experience commercial picking of especially valuable species. Apparently,
helicopters are routinely used to transport matsutake collected in roadless areas of interior British Columbia
(Hosford et al, 1997)! Relatively inaccessible areas are unlikely to experience significant commercial picking
when retail prices are low. There is significant international competition and international markets and prices
can fluctuate wildly from year to year, and even within a season, as global weather patterns produce good or
poor crops in various locations. Competition and price fixing affect prices paid to pickers. When prices are high,
large numbers of pickers may congregate in small areas '... sometimes to the consternation of land managers and
the local communities' (Amaranthus & Pilz, 1996).
The regulatory issues created by commercial mushroom picking across the Pacific Northwest include:
overharvesting, undesirable harvest methods, competition between harvesters for 'picking rights' (sometimes
armed conflict! See Hosford et al, 1997, p. 45; McRae, 1993), wildlife harassment, traffic safety and road
repairs, campground crowding and maintenance, littering, trespass, firearm safety, vandalism, and research site
security. The report concludes: '...Regulations for protecting the mushroom resource may take the form of
implementing harvest rules and permit systems, limiting permit numbers, allocating or rotating collection areas,
or providing contracts for exclusive harvest rights. Prevention of inappropriate harvest methods requires
communication through meetings, videos, posters, handouts, press releases, presentations, and other public
education efforts. Road closures may be necessary to control traffic, road deterioration, and wildlife harassment
or to protect research and monitoring areas. Law enforcement is needed to deter unauthorized or illegal activities
in the forest. Large numbers of harvesters and avid competition for a valuable commodity can increase the
potential for crime, and because buyers often handle large amounts of cash, they are tempting targets for
robbery' (Amaranthus & Pilz, 1996).

10.8.3 Managing the ecosystem towards sustainable development

The growth in commercial mushroom picking of recent years is unlikely to have generated so much concern had
it not coincided with the growing debate about conservation and the damage done to natural biodiversity by
commercial pressures. Revelations of the extent of commercial picking, or indeed the arrival of commercial
pickers in a woodland, fuel concern that both timber and mushroom harvesting adversely affect the
sustainability of wild mushroom populations. You don't have to be an expert to recognise that the woods and
forests are being changed by commercial pressures of all sorts; from pressure to use the land for other purposes

  13
  
to pressure to grow a more profitable tree crop. Couple this with the known adverse effects of atmospheric
pollution (especially severe in northern Europe) and one can readily appreciate how commercial mushroom
picking amplifies existing concern among mycologists, forest managers, recreational pickers, commercial
harvesters, buyers, and processors alike.
The decline in fungal populations in Europe and its correlation with increases in various types of
pollution are well documented. Acid rain, caused by the use of high-sulfur coal for power generation, has been
linked to decline in European temperate forests and mycorrhizal fungi. Decline in Cantharellus cibarius has
been correlated with acid rain deposition patterns (and specifically not with mushroom harvesting). Pollutants
from heavy industry harm trees directly and thereby damage the soil and mycorrhizal fungal associates. These
sorts of pollutants are at much lower levels in the Northwest US forests, but it is not known whether nitrous
oxides and ozone values typical of the Pacific Northwest could influence production of wild edible mushrooms.
As many of the commercially valuable fungi are mycorrhizal, genuine fears have been raised that mushroom
harvesting at current rates could adversely affect forest health and productivity or food webs for wildlife species.
Unfortunately, we again encounter the problem that so little research is done on Kingdom Fungi that we just do
not know whether the catastrophe is happening, is about to happen, or has already happened!
Some US agencies are restricting mushroom harvest in particular forest areas because of these
uncertainties and additional regulation and legislation will no doubt be called for. Data from Europe, however,
has not attributed decline in populations of mycorrhizal fungi in the last thirty years to collection of mushrooms
by commercial pickers. Rather, alterations in forest habitats by agriculture and urban development has led to
changes in fungus composition. Importantly, though, decline in mushroom populations over these years has
outpaced the loss or alteration of habitat, changes in forest age, change in tree composition, and change in forest
structure. Again, we find our own ignorance at the end of our search for reasons.
The key reaction to all this seems to be to call for effective management of the forest resource.
Coupled, of course, with the recognition that the mushroom harvest is part of that resource and its demands must
be anticipated. Land managers play key roles here because their tree management decisions influence the
abundance and distribution of wild mushrooms. Managers must appreciate that wild mushrooms are an
important forest product, both economically and ecologically. Mycologists must contribute, too. Historically,
mycologists have not been used to thinking in terms of conservation, and, for that matter, conservationists have
not paid much attention to fungi. Even though the decline of biodiversity has been expressed in Red Data Books
of threatened plants and animals for several years, fungi are only just making an appearance. We need to
encourage 'ecosystem management', which is an holistic approach to managing land, based on understanding
and maintaining the components, interactions, and processes of ecological systems, while simultaneously
producing products and services for society. The USDA principles are worthy of wider adoption: '... to be
effective, ecosystem management must be "adaptive;" that is: (i) the best information currently available should
be used for immediate decisions; (ii) monitoring and research activities should be started to address critical areas
of ignorance; (iii) management strategies should be adapted (modified) as improved information becomes
available...' (Pilz et al., 1996).
An ecosystem is an interactive network of the organisms and their environment. Every organism plays
a part. Thus changing a natural ecosystem into a monoculture (say, a plantation with just one type of tree) in the
hope of creating a mushroom farm in forest area, is unlikely to be successful. Rather, by preserving the natural
habitat as a whole and implementing a proper, holistic, management, a consistent mushroom yield can be
guaranteed with secure replenishment from natural resources. This should be a way to maintain a sustainable
resource which the public can enjoy while the mushroom harvester profits from it.

10.9 Concluding remarks

As a readily manipulated group of saprotrophic and mycorrhizal organisms, fungi will continue be used for
commercial exploitation. Future use will not be limited solely to food production. Opportunities for exploitation
are already expanding into the fields of nutriceuticals (nutrients which have some additional medical value) and
food supplementation with a special flavour, taste, texture and/or colour. Advances in cultivation technology
guarantee control of product quality and consistency and permit modification as required to meet market
demands.
Both developed and developing countries grow and sell mushrooms, and the cultivation industry is
currently structured in a way that allows considerable benefit to be returned to the mushroom farmers of the
world. If there are ten million mushroom farmers in China, and there are certainly over a million decentralized
mushroom farms, the cash return on the mushroom crop is making a considerable contribution to rural income.
This remains true even though farm-gate prices are much lower than retail prices. Exploitation, however, is an
economic concept; it has nothing to do with altruism and little to do with science. Indeed, the view from the
academic’s office may be very different from the view from the accountant’s. Current biotechnology babble is
likely to include the phrases ‘market pull’ (the extent to which product success is determined by the demands of

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the market and positive response of the consumer) and ‘science push’ (the introduction of innovative products to
the market as a result of scientific innovation). We hope it is not too cynical to suggest that the really critical
contribution is made by what we would call ‘accountant shove’ and define this as the tendency to take decisions
that seek to maximise return on past investments as well as future profits solely for the benefit of the balance
sheet. Future developments in the exploitation of filamentous fungi must give due weight to the community
impact of the exploitation. In developing cultivation methods we think this means applying the latest research in
adapting traditional methods, and integrating mushroom cultivation with waste disposal and remediation. In
developing commercial picking we think it means combining conservation with picking so that the industry is
sustainable. In the fermentation industries we think this means ensuring that the real value of the product far
outweighs any adverse environmental impact of the industrial process itself or its wastes. Above all, there is a
need for wider and deeper research into the biology of fungi. Too often, it seems, literature surveys reveal our
ignorance more explicitly than our knowledge. Exploit the fungi, not the consumers, communities or cultivators.

References
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Ecosystems to Conserve Fungus Diversity and Sustain Wild Mushroom Harvests, General Technical Report PNW-
GTR-371 (eds: D. Pilz and R. Molina). USDA Forest Service, Pacific Northwest Research Station, Portland,
Oregon, USA , pp. 42-61.
Arnolds, E. (1991). Decline of ectomycorrhizal fungi in Europe. Agriculture, Ecosystems, and Environment 35: 209-244.
Arnolds, E. (1995). Conservation and management of natural populations of edible fungi. Canadian Journal of Botany 73
(supple. 1): 987-998.
Chang, S. T. (1998). A global strategy for mushroom cultivation - a challenge of a ‘non-green revolution’. In Proceedings of
the ‘98 Nanjing International Symposium, Science and Cultivation of Mushrooms (eds: M. Lu, K. Gao, H. -F. Si
and M. -J. Chen). JSTC-ISMS, Nanjing, China, pp. 5-15.
Chiu, S. W., Ching, M. L., Fong, K. L. and Moore, D. (1998a). Spent oyster mushroom substrate performs better than many
mushroom mycelia in removing the biocide pentachlorophenol. Mycological Research 102: 1553-1562.
Chiu, S. W. Chan, Y. H., Law, S. C., Cheung, K. T. and Moore, D. (1998b). Cadmium and manganese in contrast to calcium
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Hosford, D., Pilz, D., Molina, R. and Amaranthus, M. (1997). Ecology and Management of the Commercially Harvested
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McRae, M. (1993). Mushrooms, guns and money. Outside 18: 64-69 and 151-154.
Molina, R., O’Dell, T, Luoma, D., Amaranthus, M., Castellano, M and Russell, K. (1993). Biology, Ecology and Social
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Peinter, U., Pöder, R. and Pümpel, T. (1998). The iceman's fungi. Mycological Research 102: 1153-1162.
Pilz, D. and Molina, R (eds) (1996). Managing Forest Ecosystems to Conserve Fungus Diversity and Sustain Wild
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Pilz, D., Molina, R., Amaranthus, M., Castellano, M.and Weber, N. S. (1996). Forest fungi and ecosystem management. In:
Managing Forest Ecosystems to Conserve Fungus Diversity and Sustain Wild Mushroom Harvests, General
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Trinci, A. P. J. (1992). Myco-protein: a twenty-year overnight success story. Mycological Research 96: 1-13.
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Figs 1 - 7. Mushroom production. Fig. 1, turning the hot, steaming Agaricus compost. Fig. 2, “... never mind the
pins, feel the compost ...” it is considered very important for the farmer to feel the texture and moisture level of
the compost. This sort of subjective judgement of the state of the compost is highly valued even though most of
the environmental conditions (light, relative humidity, air temperature, compost temperature) for Agaricus
cultivation are likely o be computer-controlled. Fig. 3, the middle tray in a three-tier Agaricus growing house in
Holland. Fig. 4, an outdoor shiitake farm using natural wood logs in Hubei Province, China. Fig. 5, Lentinula
mushrooms fruited on an artificial log in the Department of Biology of the Chinese University of Hong Kong.
Fig. 6, Volvariella volvacea growing on rice straw. Fig. 7, immature Volvariella ‘eggs’ grown on cotton waste
compost (mixed with lime and wheat bran) in the Department of Biology of the Chinese University of Hong
Kong.

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Fig. 8. Fermenter growth of fungi. An experimental fermenter growing Monascus purpurea in the laboratory in
the Biology Department, Chinese University of Hong Kong.

Figs 9 - 12. Point of sale. Fig. 9, supermarket mushrooms in Didsbury, Manchester. Fig. 10, selling mushrooms
in Nathan Road, Hong Kong. Figs 11, 12. Fungi as food for sale in Manchester. Fig. 11, fresh Agaricus bisporus
button mushrooms in their natural state. Fig. 12, three packs of Fusarium venenatum (previously called
Fusarium graminearum) pretending to be meat.

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