Preservation and Processing
Panagiota A. Diamantopoulou and Philippoussis N. Antonios
Mushroom cultivation is a rather complicated process that produces a highly nutritious food of excellent
taste from waste materials, contributing to the nutrition and economic welfare of the people. Fresh and
preserved fruit bodies of about 200 mushroom species are consumed throughout the world as a delicacy
particularly for their specic aroma, texture, and taste. According to the data of FAOSTAT (2014), total
world production of mushrooms (including trufes) was nearly 8 million metric tons in 2012, with China
being by far the leading producer (5.2 million tons). Among over 20 cultivated species, Agaricus bisporus
22.1 Introduction .................................................................................................................................. 495
22.2 Mushroom Composition and Nutritional Aspects ....................................................................... 496
22.3 Health Benets of Mushrooms .................................................................................................... 496
22.4 Cultivation and Harvesting Methods Affecting Productivity, Quality, and Shelf-Life ............... 497
22.4.1 Agaricus bisporus ............................................................................................................ 497
22.4.2 Pleurotus spp. .................................................................................................................. 498
22.4.3 Lentinula edodes .............................................................................................................500
22.5 Postharvest Handling of Mushrooms ........................................................................................... 502
22.5.1 Mushroom Grading and Quality ..................................................................................... 502
22.5.2 Postharvest Physiology and Storage ................................................................................ 502
22.6 Processing Methods of Mushrooms ............................................................................................. 504
22.6.1 Processing for Short-Term Preservation .......................................................................... 504
184.108.40.206 Cooling: Refrigeration ..................................................................................... 504
220.127.116.11 Minimal Processing ......................................................................................... 506
18.104.22.168 Effect of Packaging on Mushroom Shelf-Life ..................................................510
22.6.2 Processing for Long-Term Preservation ...........................................................................510
22.214.171.124 Canning .............................................................................................................511
126.96.36.199 Drying ...............................................................................................................511
22.7 Value–Added Mushroom Products and By-Products ...................................................................512
22.7.1 Food, Beverage, and Beauty Mushroom Products ...........................................................512
188.8.131.52 Food Products and Food Additives ...................................................................514
184.108.40.206 Beverages and Beauty Products ........................................................................514
22.7.2 Dietary Supplements: Nutraceutical Products .................................................................515
22.7.3 Mushroom Industry Spent By-Products ...........................................................................516
22.8 Conclusions ...................................................................................................................................517
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496 Handbook of Vegetable Preservation and Processing
(button mushroom, white mushroom, brown mushroom, or portobello) dominates worldwide, followed
by Lentinula edodes (shiitake) and Pleurotus spp. (particularly Pleurotus ostreatus, oyster mushroom).
In this chapter, dealing with cultivated mushrooms’ preservation and processing, their nutritional value
and health-promoting effects are initially presented as well as the cultivation and harvesting methods
affecting crop productivity, quality, and self-life. The next two sections focus on mushrooms postharvest
physiology and handling, storage, and processing methods for short- and long-term preservation. Finally,
different added-value mushroom-derived products as well as mushroom industry waste by-products,
exhibiting exploitation potentials, are presented.
22.2 Mushroom Composition and Nutritional Aspects
Mushrooms are a healthy food that is low in calories but rich in protein, dietary ber, vitamins, and
minerals (Crisan and Sands 1978; Bano etal. 1988; Barros etal. 2008). Apart from valued compounds
affecting taste, mushrooms are nutritionally desirable because of their low energy value, ber content,
and high antioxidant capacity (Kalač 2013). They furnish good quality proteins, but the crude protein
content of cultivated mushrooms varies greatly. According to data of Mattila etal. (2002) for A. bisporus,
P. ostreatus, and L. edodes the net protein content ranges from 1.6 to 2.1 g/100 g, while the respective
range mentioned by Reis etal. (2012) is 0.8–1.2 g/100 g. Mushroom proteins are relatively rich in the
amino acids glutamic acid (12.6%–24.0%), aspartic acid (9.10%–12.1%), and arginine (3.70%–13.9%)
but decient in sulfur-containing amino acids, including methionine and cysteine (Mattila etal. 2002;
Cheung 2008). The moisture content of mushrooms ranges from 85% to 95% of their fresh weight and
it is affected by the time of cropping, watering conditions as well as temperature and relative humidity
during cultivation and postharvest period (Cheung 2008). The total carbohydrate content of mushrooms
(35%–70%, including digestible and nondigestible carbohydrates), varies with species (Cheung 2010). In
general, mushroom fruit bodies are a good source of dietary ber, comprising mainly water-insoluble ber
in the form of chitin (polymer of N-acetyl-glucosamine) and nonstarch polysaccharides like β-glucans
(Sadler 2003). The water-soluble dietary ber is less than 10%, with glucose, mannitol, and glycogen
being the predominant digestible carbohydrates (Cheung 2010). The fat contain is generally low (usually
<10% dw), especially in some species like L. edodes and P. ostreatus (~2%–3%; Reguła and Siwulski
2007). Cultivated mushrooms are mainly a source of unsaturated fatty acids (FAs), accounting ~75%
(w/w) of the total FAs (André etal. 2010; Reis etal. 2012). Among them, according to Diamantopoulou
etal. (2012a) linoleic acid (Δ9,12C18:2) is the predominant, reaching ~74%–82% in L. edodes, ~73% in
P. ostreatus and Ganoderma lucidum, 70% in Auricularia auricula but less than 50% in Volvariella
volvacea and Morchella esculenta which contain signicant quantities of oleic acid Δ9C18:1. Palmitic
acid is the main saturated fatty acid (Diamantopoulou etal. 2012b, 2014). Mushrooms are fairly good
source of vitamins, particularly thiamine, riboavin, niacin, biotin, and pantothenic acid. Folic acid and
vitamin B12 which are absent in most vegetables are present in the mushrooms which also supply a range
of valuable minerals, containing macroelements such as calcium, magnesium, sodium, potassium, and
phosphorus and microelements such as copper, iron, manganese, and zinc (Cheung 2008). However, the
ability of some species to accumulate detrimental elements including radioisotopes has to be taken into
consideration (Kalač 2013).
22.3 Health Benefits of Mushrooms
In recent years, increased interest in human health, nutrition, and disease prevention has enlarged
consumer demand for functional foods. In fact, many mushrooms have become attractive as much
research focused on their health promoting effects attributed to their bioactive compounds that present
immunomodulating, antitumor, antioxidant, radical scavenging, cardiovascular, antibacterial, antiviral,
antihypertensive, antihyperholesterolemia, detoxication, hepatoprotective, and antidiabetic activities,
thoroughly reviewed by many (Wasser and Weis 1999; Wasser 2002; Sadler 2003; Lindequist etal. 2005;
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Cheung 2008; Patel and Goyal 2012; Rathee etal. 2012). The market value of dietary supplements from
mushrooms is quickly growing and estimated over U.S. $15 billion (Wasser 2012).
Some mushrooms, in addition to nutritional importance, have gained special consideration due to
their various medicinal values, as they contain compounds that have been classied as Host Defense
Potentiators (HDP) and can have immune system enhancement properties. These compounds include
polysaccharides (β-glucans), polysaccharide-peptides, nucleosides, triterpenoids, complex starches, and
other metabolites (Yassin etal. 2003). For example, L. edodes has been reported to possess antitumor,
antihypertensive, hypocholesterolemic, and antibacterial activities (Israilides etal. 2008, Wasser 2010a;
Rahman and Choudhury 2012). G. lucidum has been proved to have antimicrobial and anti-HIV effects,
while the β-glucan polysaccharide and the ganoderic acid of this mushroom have shown antitumorogenic
effects (Tang etal. 2006; Ramberg etal. 2010; Wasser 2010b). Mushrooms of Pleurotus species were
reported to have hypocholesterolemic, anti-inammatory, and immunostimulatory activities (Alam etal.
2009, 2011; Patel etal. 2012). The potential therapeutic implications of mushrooms are enormous, but
detailed mechanisms of the various health benets of mushrooms to humans still require intensive inves-
tigation. Isolation of their active ingredients, with mechanism-based potential therapeutic value, remains
a challenge (Rathee etal. 2012).
22.4 Cultivation and Harvesting Methods Affecting
Productivity, Quality, and Shelf-Life
Mushroom cultivation technology involves several different phases (i.e., development of spawn, prepa-
ration of substrate, spawn running, and mushroom development) that are well known and extensively
described (Van Griensven 1988; Chang and Miles 2004; Philippoussis 2009). Although for achieving
consistent high culture performance the very best substrate ingredients and the high yielding strains
have to been chosen (Ahlawat 2011; Philippoussis and Diamantopoulou 2012), this section deals with the
cultivation process after lling the room with the inoculated substrate, highlighting signicant cultiva-
tion parameters involved in the preharvesting and harvesting stages and contributing to high yield and
production of quality mushrooms.
In general, the aim of the grower is to manage good mushroom productivity and quality with per-
fect shelf-life properties. However, this is not an easy task as there are many factors, depending on the
cultivated species of mushrooms that inuence the productivity and the quality of the nal product.
Following inoculation, the mycelium, grows through the substrate, biodegrades its ingredients and sup-
ports the formation of fruiting bodies. Mycelial growth and fruiting phases of mushrooms life cycle
are regulated by temperature, gaseous environment, nutrient status, water activity, and in certain cases
by light, for example, Pleurotus spp. and L. edodes have an obligate requirement for light for fruiting
induction and Agaricus spp. have no light requirement (Stamets 2000; Chang and Miles 2004; Zadrazil
etal. 2004). The level of environment and cultural control used is determined by the type of production
technology. Depending on the fate of the harvested product as fresh or preserved material, the fruit bod-
ies are harvested at different developmental stages according to the grades used in marketing, either by
hand or mechanically and processed accordingly (Singh etal. 2011).
22.4.1 Agaricus bisporus
This most intensively cultivated mushroom, presents high-technology cultivation systems and grow-
ing particularities as the development of fruit bodies requires a nonnutritional layer of casing soil on
top of the nutritious compost (Straatsma etal. 2013). The mushroom mycelium grows into the casing
layer in similar conditions to those of compost colonization, and when it reaches the upper surface of
the casing layer the fruiting process starts through environmental manipulation comprising reduction
of the temperature and the concentration of carbon dioxide through aeration to trigger fructication
and to favor the development of mushrooms (Van Griensven 1988; Sánchez 2004; Ahlawat 2011). The
rst pin initials begin to appear about 2 weeks after casing. From one layer of compost, two to three
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498 Handbook of Vegetable Preservation and Processing
crops (called ushes) are harvested. In commercial practice, mushrooms are harvested before the
stage of maturation and death and ushes develop almost at weekly intervals (Van Griensven 1988;
Moore and Chiu 2001; Straatsma etal. 2013).
Yields are inuenced by compost depth and quality, length of cropping and grade of mushrooms
picked, spawn productivity, moisture and climatic conditions, and disease factors. It is well known that
the product quality during storage is mainly dependent on the at-harvest quality attributes, which can be
positively affected mainly by cultivation (Philippoussis etal. 2001a; Straatsma etal. 2013). Table 22.1
summarizes the major techniques applied in the cultivation process of A. bisporus for good productivity
of the crop as well as quality and self-life of mushrooms.
Regarding preharvesting processes, total crop yield is rst of all determined by the quality and amount
of compost lled in the cultivation area and by the texture and the humidity of the casing layer. On the
other hand, mushroom numbers and their distribution over the bed are strongly modulated by the airing
(blow down), for example, if airing is delayed, or if it is carried out with a slowly decreasing temperature,
relatively much mycelium becomes visible at the surface of the casing and relatively few fruit bodies
develop. Moreover, an increasing evaporation during pinheading is very important, while an even spread
of fruits is good for the quality. Moreover, a prerequisite for good crop is maintaining the proper hygiene
inside the growing room, along with uniformity of the temperature, RH, and the CO2 concentration (Van
Griensven 1988; Straatsma etal. 2013).
As far as harvesting period is concerned, it commences at the rst sign of buttons, often on a 7–10
day cycle and usually lasts for 1–1.5months. Mushrooms may be picked at the button (small unopened
mushrooms), cup (the cap has begun to open), or at (fully expanded caps exposing all of the gills) stage
depending on market requirements. Timing is important as mushrooms grow quickly, doubling their size
within 24h. The fruiting bodies are harvested by hand with a twisting motion, the stems are trimmed
and the mushrooms are usually graded straight into boxes for transport and sale (Philippoussis et al.
2001a; Ahlawat 2011). Whiteness and cleanliness of fresh mushrooms are the principal factors determin-
ing quality. Consumers prefer to purchase mushrooms that are bright white, free of casing material, and
free of brown spots (blotches). The greatest problems with mushroom quality in the rst ush occur on
the rst picking day (grey color and water blotches) and the last picking day (veils, long stalks, and grey
discoloration particularly at the edge of the cap). In the second ush most problems occur on the nal
picking day (ripening) and with the third ush most problems relate to mushroom color. Various irriga-
tion treatments, involving the addition of hydrogen peroxide and calcium chloride, applied throughout
the growth of a mushroom crop, have shown to reduce bacterial populations and to improve the initial
quality and postharvest shelf-life of mushrooms (Kukura etal. 1998; Diamantopoulou and Philippoussis
2001; Chikthimmah etal. 2005). Moreover, the picking management (Table 22.1) determines the qual-
ity and production of mushrooms from the very moment that harvesting starts (Van Griensven 1988;
Straatsma etal. 2013). The decision whether a mushroom has reached the required quality or not and
which mushrooms will be left on the beds until the following picking is very crucial, while the graze
picking system (several times a day) is modulator for better retail quality, color and shelf-life, longer
ush, and higher productivity.
22.4.2 Pleurotus spp.
The production of Pleurotus species (oyster mushrooms like P. ostreatus, P. sajor-caju, P. pulmonarius,
P. eryngii, P. cornucopiae, P. tuber-regium, P. citrinopileatus, and P. abellatus) is a sharp contrast with
the technology used for Agaricus production. Both pasteurized and sterilized substrate of a wide range of
residues can be used, no casing is required, while fruiting is light dependent (Philippoussis etal. 2001b;
Royse 2004). The spawned substrate is lled into perforated polyethylene—PE blocks (bags and bottles
are also used) and at the end of the spawn run period, fructication of P. ostreatus is triggered by lower-
ing the air temperature and CO2 levels, while light essential for pinning (8–12h cycle with light intensity
150–250 lux) is provided. The mushrooms begin to form around the edges of block perforations (Royse
2003; Sánchez 2010).
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Preharvest and Harvesting Methods and Techniques Used in Agaricus spp. Cultivation to Promote
Mushroom Productivity and Quality
and Methods Description of the Techniques and Target Results
Compost • Compost quality and well-lling of beds (e.g., edges of the bed)
• The lling weight is determinant for both the quality and quantity of the second
ush. Filling less than 90kg of compost/m2 usually results in quick lose of activity
and in lighter mushrooms of the second ush that mature too quickly
• A rougher textured (good moisture retention properties) and wetter casing soil gives
better mycelium quality and good quality of the later ushes mushrooms
• A good casing layer structure is vital for good CO2 and heat exchange and to spread
the mycelium throughout the entire layer for uniform recovery growth
• More compacted casing soil creates less gas exchange and less but heavier fruit
Rufing • Serves to increase mycelium quantity in the casing soil. This results to heavier
mushrooms and a better spread of the ush. Better planning for the rst day of
picking, improved uniformity over the beds, and higher production are achieved.
• Lightly rufed casing soil produces less mycelium with less pinheads and greater
percentage of large mushrooms (cap diameter >45–50mm)
• Deep rufing (up to 1cm above the compost) has positive effects on an improved
heat, moisture, and CO2 exchange, gives better quality mushrooms (less susceptible
to internal moisture), and also leads to higher productivity
• Control of the evaporation between spraying actions has better watering results and
enhances mushroom yield (mushroom number and crop timing) and quality (by
minimizing mushroom discoloration and internal moisture)
• Determining the moment of pin formation, the number of pinheads, the amount, and
size of mushrooms (by air temperature and CO2) is crucial for the harvest
• The formation of clusters of mushrooms, between which a too high CO2 content is
created (e.g., on the third day of picking) results in softer caps and wet stalks that
elongate fast. Space must be crated between the mushrooms for air circulation
• The rst picking day thinning has to be done by removing all mushrooms with a
diameter of 25–35mm in order to give the remaining fruits space to develop into
bigger and heavier mushrooms during the rest of the harvesting period, thus
enhancing also quality and shelf-life. Thinning prevents the compost temperature
rising too quickly and the CO2-content between the clusters becoming too high
• The mushrooms are harvested up to the max diameter (heavier mushrooms) and by
picking several times a day the yield (~2–3kg/m2) and quality (~5%) is increased
• Provide the unpicked mushrooms with space and nutrition by higher picking
frequency. The result is higher productivity, better quality (up to 85%) and color, and
• Moisture management during the rst ush is essential for good quality mycelium
that transports the nutrients required, has positive effect on the second ush quality
Watering • Watering on the mushrooms is necessary to maintain quality, but less spraying has as
result average better quality, with less discoloration and better keepability
• Irrigation treatments involving the addition of calcium salts, have shown to improve
the initial quality and postharvest shelf-life of mushrooms
Hygiene • Cleaning beds (broken stems, stumps, fallen mushrooms, etc.) after harvesting
Sources: Van Griensven, L.J.L.D., The Cultivation of Mushrooms, Darlington Mushroom Laboratories Ltd., Rustington,
U.K., p. 515, 1988; Chang, S.T. and Miles, P.G., in: Chang, S.T. and Miles, P.G. (eds.), Mushrooms: Cultivation,
Nutritional Value, Medicinal Effect and Environmental Impact, 2nd edn., CRC Press LLC, Boca Raton, FL,
p.453, 2004; Ahlawat, O.P., Crop management of white button mushroom (Agaricus bisporus), in: Singh, M.,
Vijay, Β., Kamal, S., and Wakchaure, G. (eds.), Mushrooms: Cultivation, Marketing and Consumption,
Directorate of Mushroom Research (ICAR), Solan, India, pp. 85–96, 2011; Straatsma, G. etal., Fungal Biol.,
117, 697, 2013.
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500 Handbook of Vegetable Preservation and Processing
The right shape for picking can be judged by the shape and size of the fruit body. The fruit bodies
should be harvested before the mushrooms show slightly curled edges and before spore release, by
twisting so that the stubs are not left on the block holes. It is advisable to pick all the mushrooms
at one time from a block and the next flush will appear at one time (Chang and Miles 2004). There
are numerous studies on different Pleurotus species cultivation that evaluate the use of different
strains, types of spawn, methods of cultivation, and different lignocellulosic substrates for their
crop productivity (Philippoussis etal. 2001b; Royse 2002; Mandeel etal. 2005; Kirbag and Akyuz
2008; Onuoha etal. 2009; Philippoussis 2009; Sánchez 2009; Fanadzo et al. 2010). An average
biological efficiency (kg of fresh mushrooms per 100kg of dry substrate) can range from 80% to
120% for P. ostreatus and 90% to 150% for P. pulmonarius (Philippoussis and Diamantopoulou
2011). However, the crop is prone to fungal diseases (like green mold, Trichoderma spp.), bacterial
spots (like brown blotch, Pseudomonas spp.), and is suspect to attacks from flies (sciarid and cecid)
and mites (Cha 2004). General control measures which are needed include: good pasteurization of
the substrate (spraying with fungicide like prochloraz–manganese complex before pasteurization),
use of healthy spawn, proper management of temperature and humidity during growing period,
sanitation and hygiene, and regular application of chlorinated water containing 100–150 ppm of
freely available chlorine at an interval of 3–5days (Cha 2004). The spore load generated within the
growing room is another disadvantage of the crop, as they can become a potential health hazard to
workers allergic to the spores. Sporeless strains are highly sought after by oyster mushroom grow-
ers (Sánchez 2010).
22.4.3 Lentinula edodes
The current trend of Lentinula growing is in plastic bags (synthetic logs) containing sawdust-based
or other lignocellulosic substrates supplemented with nitrogen sources. This method decreases the
production time and increases productivity (Philippoussis et al. 2003, 2007; Royse 2004; Chen
2005). Actually, high average biological efciencies (BE) are achieved with sunower seed hulls
(BE: 107.5%) (Curvetto et al. 2005), sugarcane bagasse (BE: 87.4%) (Salmones et al. 1999), and
corn cobs (BE: 80.6%) (Philippoussis and Diamantopoulou 2011). More or less similar results (BE
≈ 80%) have been obtained with hard-wood residues and cereal straws (mainly barley and wheat)
The management of parameters, in different stages of growth and fruiting of L. edodes, for good pro-
ductivity and quality of mushrooms is presented in Table 22.2.
Harvesting is performed when the edge of the mushroom cap is still in-rolled, or when the mush-
room cap is only partly extended (60%–70%). In general, Lentinula quality is determined by shape
(rounded with downward in-rolled edge before the cap is fully extended and central stalk), texture
(thick and tight context), size, color, avor, and aroma (Chen 2005). Freshness and freedom from
pests and impurities are also critical factors for high-quality mushrooms (Chen 2001). Actually,
during bag cultivation of shiitake many pests (like ies and mites) and diseases (like green mold,
Trichoderma spp.) can occur as they thrive in warm and humid conditions. If unnoticed, these pests
often lower productivity and quality and can sometimes cause total crop failure. The use of pesticide
chemicals, however, is not advisable, as these materials can affect the mycelial growth and reduce the
quality of the shiitake. As a result, for mushroom growers, energetic precautionary measures should
be taken to avoid contamination. According to Fan etal. (2005), some good sanitation and hygienic
practices for successful pest and disease management are as follows: selection of fresh, pathogen and
pest-free substrates and supplements, sterilization of substrates, clean and disinfected inoculation
room and box as well as hands during spawning, keeping the incubation room clean and well-aerated,
frequent and carefully inspection of the bags and elimination of contaminated bags immediately,
sterilization of the contaminated bags before their disposal, removal of fruit bodies stumps which
remain after harvest and might attract pests, disinfection of the spent substrate, and growing houses
on a regular basis.
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Management of Parameters and Cultivation Procedure in Different Stages of Growth and Fruiting of Lentinula edodes Grown in Bags
(%) CO2 (ppm) Light (lux) Duration Cultivation Procedures and Remarks
21–27 90–100 1,500–10,000 No or short
30–45days • Substrate type and its nutrients as well as texture-porosity (aeration) affect
incubation/preharvest period and yielda,b,c
• Spawn-run: strain-dependant duration, there are cold and warm temperature strainsa,c,d
• Toward the later stages of the spawn run 1, a thick mycelial coat forms on the outer
surface of the colonized substrate blocka,b,d
21–27 70–90 >1,500 No or short
45–60days • Clumps of mycelia (bumps) and early primordia are produceda,d
• Great numbers of bumps (prone to abortion) must be eliminated by aeration (cutting
slits on bags)
• Bag removal for browning of the mycelial coat and bark formationa,b,d
5–7days • Fruiting induction by aeration and cold-shock through: water soaking (4–24h at
12°C)a,b,e, or temp. reduction to 15°Ca,b,c,d
• Temperature actuationa,d,e
16–18 98–100 <1,000 500–2000
5–7days • Relative humidity (RH) uctuation during fruit body developmente
• RH is lowered to 60% for 6–12h before crop harvesting for better shelf-life of
Harvest of rst
16–18 60–70 <1,000 500–2000
5–10days • Harvest when the edge of the mushroom cap is still in-rolled, or when it is only
• Trimming the end of the stalk and cutting off residual stalk stubs from the
18–21 50–60 >1,500 Short
15days • Lower the humidity to 30%–50% RH at 21°C during rest-periode
• For the second ush: soaking the substrate block for up to 12h and incubation for
1week at higher temperaturesb,c,d,f
16–18 60–70 <1,000 500–2000
5–10days • After cooling to 16°C for 1 week a second ush will be harvestedc,d,f
a Chen (2001).
b Royse (2004).
c Philippoussis etal. (2003).
d Chen (2005).
e Stamets (2000).
f Philippoussis etal. (2007).
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502 Handbook of Vegetable Preservation and Processing
22.5 Postharvest Handling of Mushrooms
The white mushroom (A. bisporus, A. bitorquis, etc.) is still nowadays the most cultivated mushroom
worldwide (Giri and Prasad 2013). It seems therefore reasonable that most aspects and data concerning
mushrooms’ postharvest physiology, processing, and storage focus on this mushroom species. However,
other mushrooms such as the oyster (P. ostreatus, P. pulmonarius, etc.), shiitake (L. edodes), straw (V.
volvacea), and ear mushroom (Auricularia spp.), widely cultivated and processed, are also referred in
the following text.
22.5.1 Mushroom Grading and Quality
Quality characteristics of mushrooms include many parameters such as color, size, rmness, maturity
stage, clearness, blemish-free, avor, nutritional value, and safety and are affected by preharvest treat-
ments (see Section 22.4) as much as postharvest processing and storage conditions (Burton etal. 1989).
Actually, as mushrooms continue to develop during storage, their quality seems to be determined by the
stage of sporophores’ maturity at harvest (Hammond and Nichols 1975). This stage can differ between
species as for proper harvesting; mushrooms should be collected while their pilei are still closed and
part of the veil is intact (e.g., Agaricus spp.), their edges are uncurled and their gills well formed (e.g.,
Pleurotus spp.), before the pileus is fully expanded and with the edges rolled-in (e.g., Lentinula spp.)
or before volva breaks/immediately after rupture (e.g., Volvariella spp.). Nevertheless, variability as
expressed by mushroom heterogeneity resulting from the different batches of mushrooms, harvested at
different maturity stages, is one of the main problems in mushroom technology related with important
storage losses (Aguirre etal. 2008). Throughout storage, mushroom quality is usually assessed by pileus
color, transpiration and respiration rate, weight loss, and disease incidence (Burton etal. 1987b; Burton
1989), whereas consumers’ acceptability is mainly based on the external characteristics of the product
and to a less extent its taste, rendering quality an individual perception (Eastwood and Burton 2002).
Mushrooms during or after picking are usually sorted into several grades, according to standards set
basically by the market that reect their quality and determine their price. General criteria for mush-
rooms grading into quality categories are size, color, maturity, and range of damage, but they can differ
in each country as they are inuenced by the rate of local market development. For example, a reectance
(L-value) greater than 86 is considered prerequisite of whole white acceptance from the whiteness point
of view (Gormley 1975). Nevertheless, good agricultural practices for mushroom cultivation, handling,
processing, and marketing, for example, those described in AMI (2009) or by the application of HACCP
system (Pardo etal. 2013), are essential for attaining premium quality. There is no world standard for
grading fresh, processed, or canned mushrooms, although there are EU standards or U.S. standards for
Grades of Mushrooms (i.e., CODEX STAN 39 1981; CODEX STAN 297 2009) that apply to whole or
sliced fresh, dried, or canned edible mushroom species. Finally, as supermarkets increasingly compete
with each other on the grounds of food safety and quality, they have developed their own company-
specic standards or “supplier criteria.” For example, in the United Kingdom “Tesco” has developed
“Tesco’s Nature Choice” a quality assurance scheme for its own brand; similarly in France “Casino” has
developed “Terre et Saveur” and Carrefour “Filiere Qualite” (Bord Glas 2002).
22.5.2 Postharvest Physiology and Storage
Mushrooms represent one of the most perishable commodities, being so delicate by nature that they need
special postharvest treatment. As a number of physiological processes take place in freshly cut mush-
rooms and during storage (pileus and veil opening, stipe elongation, browning, etc.) resulting in matura-
tion and senesce, their commercial and nutritional value can be easily decreased. The rate of which these
processes occur in mushrooms during storage is affected by factors such as strain resistance and physi-
ological behavior, room temperature and relative humidity, and the presence of microorganisms (Burton
and Noble 1993; Varoquaux etal. 1999; Brennan etal. 2000; Mahajan etal. 2008a).
The most important characteristic of mushroom metabolism is the high respiration rate because of
which, a constant giving off water vapor is exiting from mushroom surface resulting in constant weight
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loss (Nichols 1985; Mahajan etal. 2008a,b). In products with such high water content (>85%) and with
no conventional cuticle as mushrooms, evaporation and consequently loss of weight usually have detri-
mental effect on quality and shelf-life; therefore, mushroom respiration rate is an index of their shelf-life.
Weight loss is greater in sliced mushrooms or whole with open veils/bruised, particular when tempera-
ture increases, humidity of the storage room is low, and partial pressure of oxygen is high. The optimum
temperature of mushroom respiration activity after harvesting is 15°C–20°C with CO2 production rate
280 mg CO2/kg/h at 18°C, whereas at 0°C only 28–44 mg CO2/kg/h are produced (Hammond and
Nichols 1975; Nichols 1985). Burton and Noble (1993) recorded that Agaricus mushrooms during stor-
age in open punnets lost 4% per day of their weight at 5°C (73% RH) and 6% (90% RH) per day at 18°C.
Nevertheless, a weight loss of 2% during storage is generally accepted. The effect of respiration activity
of mushrooms on their postharvest physiology and storage is given in Table 22.3.
Spoilage of mushrooms during storage, associated with the presence of microorganisms (mainly bac-
teria and fungi) as well as enzymes, is another aspect of their physiology strongly affecting their shelf-
life (Gormley 1975; Beelman etal. 1989). Bacteria may activate (and increase) even in cold-storage
conditions and in the high-moisture mushroom surface along with the enzymatic action occurred on
mushroom tissues can cause rapid deterioration of mushrooms when heated, such as tissue browning,
presence of brown or yellowish spots and slime in pileus or stipe (e.g., Pseudomonas sp.), and loss of
rmness. Enzymatic browning in many foods is caused by the polyphenol oxidase group of enzymes,
in which tyrosinase is comprised. Although tyrosinase is particularly abundant and active in mushroom
tissues, it is unable to react in intact cells. The brown color of aged or damaged mushrooms is a result of
a succession of biochemical and chemical reactions of (colorless) phenolic compounds, tyrosinase and
oxygen (Burton and Noble 1993). High storage temperatures are responsible for the increased browning
of mushrooms as a result of polyphenol oxidase increased activity (Ratcliffe etal. 1994). Nonenzymatic
browning is also inevitable as mushrooms contain carbohydrates, proteins, and amino acids that interact
and (particular at temperatures above 5°C) can result in tissue darkening. Browning reactions in fruits
and vegetables comprise as a serious problem in the Food Industry and is one of the most detrimental fac-
tors to mushroom’s quality, followed by the loss of texture and cap opening (Burton 1986) and they are
usually encountered by sterilization and blanching (Gormley and Walsh 1982). However, no food-borne
Effect of Respiration on Postharvest Physiology and Storage of Mushrooms
Effect Results References
Degradation and depletion of
soluble compounds (mannitol,
trehalose, glycogen, fructose,
Mushroom deterioration associated with:
• loss of texture-softening
• brown coloration in
Agaricus sp.—mucilage in Pleurotus sp., pileus
opening in Volvariella sp.
Hammond and Nichols
(1975) and Tseng and
Movement of dry matter and
water from stipe to/and pileus
Loss of texture Hammond and Nickols
(1975) and McGarry
and Burton (1994)
Changes in the nutritional and
medicinal attributes of
mushrooms during storage
and temperatures 18°C–20°C
• Decrease of protein content with accumulation of total free
• Decrease in free amino acids (after the activation of
enzyme protease) and cell wall glucans
• Decrease of total carbohydrates and phenol content
• Decrease of substances giving the pilei their taste
(5′-nucleotides and 1-octen-2-ol)
• Reduction in the content of polysaccharide lentinan
(Hammond 1979; Tseng and Mau 1999)
Hammond (1979) and
Tseng and Mau
Condensation of vapor is
observed in the inner package
when nonperforated or even
the conventional PVC
stretchable lms are used
Texture changes—softening of the esh Nichols (1985) and
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504 Handbook of Vegetable Preservation and Processing
bacteria were detected in cultivated mushrooms (Venturini etal. 2011), pathogenic Campylobacter bac-
teria are usually killed through compost pasteurization and do not grow below 28°C, and bacterium
Clostridium botulinum forms its toxin only in semiperforated packages (O2 < 2%) of mushrooms stored
in high temperatures (Sugiyama and Yang 1975). As for fungi grown on mushrooms during storage (e.g.,
Trichoderma sp.), spore germination and mycelial development are restricted due to low temperatures,
despite their resistance at cold and dry conditions.
22.6 Processing Methods of Mushrooms
Although the major part of cultivated mushrooms is consumed in the fresh condition, trading mushrooms
totally at fresh status seems unfeasible for every point of the chain and for all year around. Practices and
aspects for mushroom postharvest care include proper storage/packaging and/or minimal processing of
fresh mushrooms for their short-term maintenance, as well as various processing techniques for their
long-term preservation. However, traditional eating patterns have been changed in more selective and
individual preference eating behaviors, along with a corresponding abatement of formal eating times
and occasions. This has increased the demand for individual portion packs and convenience formats in
every day diet and has lead to reduction of the demand for whole processed frozen or canned mushrooms.
Also, increase of out-of-home consumption (hotels, restaurants, pubs, etc.) and institutional catering
(workplace canteens, universities, schools, etc.) has led in increased requirement for fresh, prepared, and
processed mushrooms from these sectors (Bord Glas 2002). Another aspect of modern mushroom pro-
cessing concerns techniques aiming at the value-addition of the product as well as environmental aspects
such as waste disposal and utilization of waste and off-grade mushrooms (Zivanovic 2006), presented
in Section 22.7.
22.6.1 Processing for Short-Term Preservation
220.127.116.11 Cooling: Refrigeration
Mushrooms at ambient temperatures (ca. 22°C) have a short shelf-life of 1–3days (Burton and Twyning
1989), at 15°C their shelf-life is 2–3days (Gormley 1981), whereas in the tropics they count only 24h
(Wakchaure 2011). Cooling is still the most effective technology available for retarding mushroom
deterioration process. As mushrooms’ respiration rate is three times higher at 10°C than at 0°C, their
immediate cooling after harvest (to 4°C–5°C) is needed for rapid removal of the eld heat, slow down
of metabolism, and deterioration prior to storage or transportation (Wakchaure 2011). In low tempera-
tures, apart from limiting weight loss and retaining freshness, mushroom pilei maintain closed and rm,
whereas browning and stipe elongation are decelerated. High relative humidity is also essential as it
prevents drying and retains pileus glossiness. Precooling at this stage is the key component in the pres-
ervation of quality characteristics of many perishable fresh produce, including mushrooms. This can be
achieved using (forced) air-, ice bank, vacuum, hydrocooling, and evaporative cooling, dependant on the
quantity of mushrooms to be handled (see Table 22.4).
After the stage of precooling, mushrooms should be placed in chambers with constant low tempera-
ture and air circulating in a uniform and freely way between mushroom punnets/packs/packages until
selling. However, packs with more than 10kg of mushrooms or with 15cm thick layers may cause
problems (Wakchaure 2011). The cooling systems can differ (simple refrigerator to large sophisticated
systems) depending on the quantity of mushrooms harvested daily. Relative humidity in any case
should be high (85%–95%) and this can be achieved by using large evaporators (Przybylowicz and
Donoghue 1988). In this manner, mushroom physiology remains almost unactivated up to 6days. It is
widely accepted that the ideal conditions for retarding the metabolic activities of tissues and decreas-
ing microbial growth up to 9days are storage temperature 0°C–2°C and relative humidity 85%–90%
(for A. bisporus and L. edodes mushrooms; Beelman etal. 1973; Nichols 1985; López-Briones etal.
1992; Minato etal. 1999). Gormley (1975) indicated that white mushrooms can be stored for 7–9days
at 0°C–1°C. The shelf-life of fresh sliced white mushrooms is shorter, mainly because of the effects
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of washing and cutting process (Brennan et al. 2000), for example, 7.5 and 4days at 0°C and 5°C,
respectively (Oliveira etal. 2012). The shelf-life of the oyster mushroom can be 9days at 2°C or 3days
at 18°C (Lukasse and Polderdijk 2003). Volvariella sp., a mushroom that undergoes autolysis at 4°C,
should be stored at 10°C–15°C and has a shelf-life of only 3days (Ahlawat and Tewari 2007). Finally,
optimum in-package O2 is 6% to reduce pileus opening (Roy etal. 1995). Usual hazards throughout
cold storage are incorrect determination of temperature, high weight loss (in dry environment), and
growth of microorganisms (in wet environment). As the inner temperature of mushrooms is more
important than that of the environment, an accurate monitoring of the pileus/stipe temperature is
essential (Nichols 1985).
Methods for Precooling Fresh Mushrooms
Methods Features References
Air-cooling • Air velocity at least 60 m/min
• Mushrooms usually placed in boxes, various packages or punnets
and cooled within 24h, in a rate of 0.5°C/h, but unevenly
• Air is usually dehumidied during refrigeration, so the vapor pressure
of the cooled air is low and dehydrates the mushroom surface
• There is about 3% weight loss in cooling from ambient temperature
to 2°C in unwrapped mushrooms that reach quicker the store
temperature particularly if they are placed on the top of the pile
• The use of lms (e.g., PVC) in mushroom packaging is essential for
preventing great weight losses even if it hampers the quick cooling
Nichols (1985) and
Forced cooling • Higher air pressure is developed on one side of the packages than
the other and air is directed through the produce cooling it rapidly
• It is accomplished by using three systems cold-wall, forced air
tunnel, and serpentine cooling
• A spray-moist chiller can be also used for cooling mushrooms
rapidly in an hour at 16°C–18°C, without moisture loss
Wakchaure (2011) and
Ice bank cooling • Air used to cool mushrooms is almost saturated as it has previously
passed through ice-cold water, cooled by direct heat exchange with it
and lled with vapor, a system preventing mushrooms from water loss
Vacuum cooling • It is a rapid and uniform (temperature reduction from ambient to
2°C in 15–25min), yet expensive method
• Subjecting mushrooms at very low pressures water from their
surface is evaporated resulting in lower temperatures
• The color of vacuum-cooled mushrooms is superior to conventional
cooled mushrooms, but the weight losses reported are 1% per 5°C of
drop, much greater than those of forced dry or wet air
• No differences in quality or hyphal structure were detected between
vacuum and conventionally cooled mushrooms
• Overwrapping with perforated lms is essential for maintaining the
appearance of vacuum cooled mushrooms
• Adding reselected amount of water before or during precooling can
prevent weight loss
Gormley (1975), Burton
etal. (1987a), McGarry
and Burton (1994), and
Hydrocooling • Large butches of mushrooms are spayed or immersed with/or chilled
or cold water (~0°C) before further packing
• It is a rapid cooling technique (the high heat-transfer rates remove
eld heat at 20–30min) that diminishes product water loss, yet with
no uniform effect
• The risk of spoilage due to microbes accumulated at recirculated
water can be prevented by the addition of chlorine solutions (100
ppm) or approved phenol compounds
Nichols (1985) and
Evaporative cooling • Inexpensive method based on the cooling effect created by
evaporation of water when dry air is shown over the wet product.
Suitable with areas with low ambient humidity (<65% RH) air
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506 Handbook of Vegetable Preservation and Processing
18.104.22.168 Minimal Processing
Minimal processing is a new approach for extending mushrooms’ shelf-life more easy and natural than
conventional processing techniques (e.g., canning and drying), with little effect on primary charac-
teristics of the produce. It includes Modied Atmosphere Packaging (MAP), Controlled Atmosphere
Packaging (CAP), washing, use of chemicals, blanching, radiation, use of moisture absorbers (e.g., sor-
bitol), or coatings (Beelman etal. 1973; Bernaś et al. 2006; Wakchaure 2011). Although treatments
involving chemicals are simpler and probably cheaper for processors to implement rather than those
involving packaging, attempts concerning the extension of mushrooms’ shelf-life by the use of chemi-
cal treatments have yet little success, basically because of the consumers’ preference in chemical free
products. New materials and lms in mushroom packaging that would replace any chemical pretreat-
ment and compound added in the packs are therefore welcomed by the consumers. Likewise, it would
be preferable for any new treatment solution to be applied in the same way as for sultes or stabilized
chlorine dioxide. The current practice of washing or dipping whole mushrooms in treatment solutions
prior to slicing is more effective, in terms of shelf-life, than dipping, spraying, or brushing solutions onto
sliced mushrooms (Brennan and Gormley 1998).
Chemicals may control microorganisms causing decay and diseases, eliminating or inhibiting the
adverse changes in the color and texture of pilei and can be applied by spraying, soaking, vacuum moist-
ening, and dipping in solutions/emulsions. Washing is occasionally used by mushroom growers for
removing soil, as mushrooms are not generally washed; this is because after washing, mushrooms have
a water-soak appearance and moisture penetration eventually leads to increased microbe populations.
Also, darkening and bronzing of the pilei often occur after washing as a result of enzyme o-phenolox-
idase release from damaged substrates (Burton and Noble 1993), not only in fresh but also in frozen
mushrooms (Czapski and Szudyga 2000). Therefore, addition in water of several reducing agents with
antibacterial effect can retard browning and pileus opening. Among these tested, the most popular are
Oxine (stabilized chlorine dioxide), sodium chloride, sodium hypochloride, calcium chloride, sodium
sulte, methyl jasmonate ethyl alcohol sole or in combination, and others are given in Table 22.5.
Addition of semipermeable edible coatings in freshly harvested mushrooms is another physical method
to reduce their water loss, browning, and microbial growth and to improve their texture during storage at
low temperature (4°C). Immersion of A. bisporus mushrooms in sodium alginate solution (Nussinovitch
and Kampf 1993), dipping freshly cut white mushrooms in solutions containing chitosan (Eissa 2007),
coating with various polysaccharides (Niazmand etal. 2009), or treating L. edodes mushrooms with chi-
tosan–glucose complex coating and then packaging in low-density polyethylene lm (Jiang etal. 2012),
all were effective in improving the shelf-life of mushrooms.
Blanching is an important treatment applied during preliminary processing of mushrooms (mainly
of the Agaricus species) after washing and prior to freezing and canning, with the use of hot water or
steam. Blanching inhibits tissue browning (inactivation of polyphenol oxidase) and production of off-
avors during mushroom frozen storage and defrosting. It also removes trapped air and decreases weight
losses during canning; the main purpose of blanching in canning however is to inactivate the enzymatic
browning by the thermal inactivation of the enzyme polyphenol oxidase and then to induce mushroom
shrinkage, so that that it will not occur during sterilization, making additionally the product more pli-
able to facilitate the lling operation (Wu etal. 1981). However, some negative features of blanching are
the extraction of useful nutritious components (Wu etal. 1981; Gormley 1984), the undesired changes
in aroma (Le Loch-Bonazzi and Wolff 1991; Mau etal. 1992), the lower taste compared to fresh cooked
mushrooms (Wu etal. 1981), the damages of pileus’ texture that cause remarkable toughness after thaw-
ing and cooking (Gormley and Walsh 1982; Reyes De Corcuera etal. 2004), the mushroom weight loss
during the blanching procedure, and the increase in mannitol concentration in mushroom mass and
blanching uid (Biekman etal. 1996).
To avoid general biochemical effects, blanching time should be kept to the minimum (Matser etal.
2000), determination of which is achieved by the peroxidase test (Reyes De Corcuera et al. 2004).
Mushroom grading under different sizes is also desirable so that each size is blanched separately to avoid
under- or overblanching. The length of the blanching treatment can vary, for example, 20 s (Wakchaure
2011), 7 min (Baldwin et al. 1986), or even 15min (Coşkuner and Özdemir 2000). Temperatures
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commonly used in industry for blanching (90°C–100°C) can cause undesirable tissue softening; how-
ever, the blanching with boiling water for 2min caused increased hardening in the oyster mushroom
(Kotwaliwale etal. 2007) and better color when it was freeze-dried (Fang etal. 1974). Water blanching
(98°C/90 s) of Boletus edulis prior to drying was found to have a negative effect on the level of antioxi-
dants, regardless of the drying method (Jaworska etal. 2014). According to Fan etal. (2005), mushrooms
Chemical Compounds Used in the Preliminary Processing of Mushrooms
Compounds Features References
Oxine (stabilized chlorine
• Very effective antimicrobial agent (50 ppm with a 2min or longer
wash period at ~12°C), but with variable results on whiteness
• Its activity is due primarily to oxidation not chlorination and
although it is up to ve times more effective, it is noncorrosive
• Known (5%, v/v) for its antibacterial and whitening actions in
• Whole fresh mushrooms soaked for 10min in solutions of H2O2
(or citric acid) then sliced, packed, and stored at 4°C for up to
19days. Both treatments reduced the number of pseudomonad
bacteria and improved their quality of when compared to control
(water soaked) slices
and Brennan (1999)
Varsenic (EDTA) and
• Both acidic and metal chelators, having the ability to inhibit
enzymatic browning and microbial growth (Pseudomonas sp.) on
• Compared to H2O2, they are easier to handle, not corrosive and do
not deteriorate with age
• Fresh sliced mushrooms were soaked for 10min in citric acid
solution (up to 40 g/L) and retained their shelf-life (whiteness)
for longer, whereas none acidic taste was detected in samples by
• As citric acid is the most widely used in foods and current
legislation supports its use over H2O2 or EDTA, it is the treatment
Oxine (50 ppm), sodium
erythorbate (0.1%) and
calcium chloride (0.5%)
• Benecial effect on color and number of bacteria Wakchaure (2011)
Ascorbic acid (AC) and
calcium chloride (CaCl2)
• AC is an antioxidant and reducing pH agent with wide use as
pretreatment for mushroom extension of shelf-life and CaCl2 is a
• Extensively used in mushroom industry as whitening and
preservative agent (1 g/L for 1–2min), recommended for frozen
and sterilized products
• It may result in potentially harmful sulfur dioxide residues
• Fresh mushrooms washed in 0.05% SM not only improved their
initial whiteness, but also kept it for longer during storage
• Washing A. bisporus mushrooms with water containing 3 g/L SM
affected positively their color of during 3-month (freezing)
storage. Immersion in water (20 s) of washed with SM
mushrooms affected negatively their color and texture, yet
decreased the SO2 content.
• In other studies, the use of SM has been proved as ineffective in
bleaching fresh Agaricus slices, reducing the number of
pseudomonad bacteria or improving their quality during cold-storage
Jasmonic acid and methyl
jasmonate ethyl alcohol
(sole or in combination)
• Positive effect on the color of A. bisporus pileus and general
appearance during storage at 13°C/5days
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508 Handbook of Vegetable Preservation and Processing
of the genus Lentinula before canning are usually blanched in water at 90°C for 5min, at a ratio of 1
(mass):1.5 (water volume), so that air is removed and bacteria population is reduced.
Blanching procedure involving just water reduces the initial mushroom whiteness; therefore, water
blanching has been performed in a variation of treatments: hot water containing 1%–2% NaCl and 0.5%–
1% citric acid at temperatures of 95°C for 8–10min (Vivar-Quintana etal. 1999), 1% NaCl and 0.1%
citric acid at 95°C–100°C for 5–6min (Wakchaure 2011), or 1 g/L NaCl and various concentrations of
citric acid or EDTA at 95°C–100°C for 15min (Coşkuner and Özdemir 2000). When glucono-δ-lactone
replaced citric acid during blanching, canned white mushrooms not only retained their color, taste, and
yield but also did not have pilei with strange acidic taste (Rodrigo etal. 1999). Prior to freezing, B. edulis
mushrooms were blanched at 96°C–98°C (3min the pilei and 1.5min the stipes) in a ratio by mushroom
mass to water of 1:5 (Jaworska and Bernaś 2009) resulting in good quality frozen product. However,
when various solutions containing citric (0.5%) and ascorbic acid (0.1%)/lactic (1.0%) and ascorbic acid
(0.1%) were used during 1h mushroom soaking before blanching, the quality of the Boletus mush-
rooms was not superior to those blanched with water. On the other hand, similar preliminary treatments
applied to A. bisporus mushrooms and extended their storage life from 4 to 8months (as compared to
unblanched samples); nevertheless, both color and nutritional value were better in fresh than in mush-
rooms pretreated and frozen (Jaworska etal. 2008). Apart from the enzyme inhibitors mentioned above
(NaCl, organic acids, and EDTA) blanching may also contain metabisulte and ascorbic acid hydrogen
peroxide (Coşkuner and Özdemir 2000; Czapski 2002). Blanching reduced the attractiveness of the
dry P. ostreatus, whereas sodium metabisulte improved it (Martínez-Soto etal. 2001). The blanching
process of P. ostreatus with hot water was optimized at 150 s and 70°C for mushrooms prior to freezing
(Vullioud etal. 2011), or at 1min and 70°C–75°C for those immersed at 35% brine solution and then
preserved at tropical room temperature for 6months (Victor and Obele 2013).
Steam blanching is applied to cut mushrooms, it requires less time than water blanching, and produces
mushrooms with darker color but better taste. Steam blanching during 90 s (Yan 2012) and microwave
blanching for 59 s at 570 W were applied to P. eryngii, both being efcient for sensory and nutritional
quality of the product. Two other treatments recommended before blanching for preserving the color and
reducing the weight loss associated with blanching are vacuum moistening and soaking in solutions con-
taining, for example, citric acid and -ascorbic acid (Beelman etal. 1973). Mushrooms that were vacuum
moistened prior to blanching had a water uptake during evacuation as much as 80% of the fresh mushroom
weight (Baldwin etal. 1986). Additionally, in order to avoid the undesirable texture changes but obtain a
white color, mushrooms prior to freezing were dipped in metabisulte solutions (0.5% and 1.0%) for 5min
after blanching (Fang etal. 1974). As soon as blanching is complete, mushrooms should be cooled quickly
with running water to thoroughly to stop the cooking process (Coşkuner and Özdemir 2000).
Radiation preservation of mushrooms by using gamma or x-rays at the dose of 100–150 Krad has been
found to restrict the postharvest growth and discoloration/deterioration of mushrooms, yet decrease with
increase the level of irradiation dose the level of octo-carbon aromatic compounds (1-octen-3-ol, 3-octa-
none, etc.) present in the mushrooms (Mau and Huang 1997). Jiang etal. (2010) reported that combina-
tion of gamma radiation (1.0 kGy) and passive MAP extended the shelf-life of Lentinula up to 20days.
Its use is limited by the cost and the perception of consumers to irradiation on food (Przybylowicz and
MAP and CAP include the use of plastic permeable lm system which overwrapping packed mush-
rooms and as a result of mushroom respiration, restricts the transfer of gases (CO2 and O2) and creates
an atmosphere poor in O2 and reach in CO2. It is the simplest, most economical, and effective method for
limiting tissue respiration rate, reducing microbial growth, and extending the shelf-life of mushrooms
(Kim etal. 2006). In order to devise an MA package, which provides optimum gas conditions, it is
necessary to select a lm with suitable gas permeability properties, not only to design a pack with the
appropriate ratio of weight of product to surface area of lm, but also to predict the respiration rate at
the desired O2 and CO2 gas concentrations (Cliffe-Byrnes and O’Beirne 2007). MAP has been mostly
studied at A. bisporus mushrooms (Nichols and Hammond 1975; Roy etal. 1995; Simón etal. 2005; Kim
etal. 2006); however, it can be applied in almost all species of mushrooms, packed whole or sliced. The
application of MA can be passive or active. In passive modication, where the required CO2 is produced
by the mushrooms and O2 is consumed with respiration, the appropriate lm permeability allows the
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entrance of required O2 and the exit of surplus CO2 until eventually equilibrium is established. In active
modication, the atmosphere inside the packages is achieved by, rstly, the creation of vacuum and, then,
by introducing of the appropriate gas mixture. In this case, steady state atmosphere is reached quickly
after packaging (Wakchaure 2011). Antmann etal. (2008) showed that the equilibrium atmosphere con-
ditions for both O2 and CO2 were reached for active modied atmosphere packages containing L. edodes
mushrooms after 10days of storage. Also, in MAP the gas composition is controlled once at the begin-
ning of storage, whereas respiration along with interaction between the product and the package modify
the atmosphere. On the other hand, in CAP the gas atmosphere is controlled constantly and accordingly
regulated (Sandhya 2010).
In MAP/CAP, normally the concentration of O2 is reduced from 21% to 2%–5% and that of CO2 is
increased from 0.03% to 16%–19%. López Briones etal. (1992) suggested that to optimize marketing
conditions for white mushrooms, the storage atmosphere should contain 5%–10% O2 and 2.5%–5% CO2.
The lms usually used are (perforated and no-perforated) LDPE (low-density polyethylene), PVC (poly-
vinyl chloride), and PP (polypropylene). A gas composition of 1%–2% O2 and 8%–15% CO2 at 18°C or
11%–17% O2 and 4%–10% CO2 at 2°C has been reported when PVC and PP lms were used in prepacks,
concentrations depended on the lm used (Nichols and Hammond 1975). However, O2 levels below 2%
are related to anaerobiosis and in combination with room temperatures (18°C–20°C) may result in the
growth of C. botulinum (Nichols and Hammond 1973). As the moisture and modied atmosphere cre-
ated in the packages affect bacterial growth and consequent color change (Burton et al. 1987b), lm
perforation is strongly suggested along with the use of new technology lms with chosen (in advance)
permeability (Falguera etal. 2011).
Regarding research on fresh sliced A. bisporus mushrooms MAP, using PVC and PP lms with atmo-
spheres of 10%–20% O2 and 2.5% CO2 at 5°C improved the appearance and reduced the microbial
counts when compared to the air atmosphere (Simón etal. 2005). Nevertheless, with storage temperature
increase more perforations are needed in order to obtain the optimum MAP conditions for sliced mush-
rooms (Oliveira etal. 2012). On the other hand, MAP did not reduce mushroom catabolism (Varoquaux
etal. 1999) and induced internal and external discoloration of carposomes (López Briones etal. 1992)
and that was the reason why MA-packed, whole mushrooms were be available on the European market.
Villaescusa and Gil (2003) proposed an MAP with a combination of 15% O2 and 5% CO2 for maintain-
ing the good quality characteristics of P. ostreatus stored for 7days at 4°C. Low O2 (2%) and high CO2
(30%) concentrations signicantly prolonged the shelf-life of P. eryngii stored at 20°C–25°C (90%–95%
RH) for 5 days (Li et al. 2013). Additionally, a combination of chemical treatment and MAP using
10% O2 and 5% CO2 provided better quality retention of Pleurotus orida (Jafri etal. 2013). The effect
of passive MA storage (5°C, 73%–77% RH) on the sensory characteristics and shelf-life of L. edodes
mushrooms was investigated by Ares etal. (2006) using three different lms (low-density PE, PP, and
PP macroperforated). The results showed that mushrooms stored under MA had higher deterioration rate
than those stored at PP macroperforated, indicating that CO2 concentrations above 9% accelerate dete-
rioration and that L . edodes mushrooms are more susceptible to high CO2 concentrations than Agaricus
ones. Similar were the nding of Parentelli etal. (2007) regarding the use of passive and active MAP on
the sensory quality and deterioration of the mentioned species. Application of active MAP on L. edodes
mushrooms packaged on bags of low-density PE under 15% and 25% O2 (80%–85% RH) resulted in the
development of off-odors after 12days of storage, whereas when macroperforated lms were used, the
weight loss was as high as 15% (Antmann etal. 2008). However, washing with citric or ascorbic acid had
positive effect on the L-value of L. edodes during 10-day storage at 7°C (Santana etal. 2008).
Additionally, the unique gas barrier properties of hydrophilic lms (wheat gluten-based material and
synthetic polymer) have been tested through the MAP of A. bisporus mushrooms packed under micro-
porous and hydrophilic lms and stored at 10°C and 20°C under high relative humidity (>92%). Unique
steady state atmospheres, poor in both oxygen and carbon dioxide, were observed, regardless of the
temperature and the hydrophilic lm used, owing to their high selectivity to gas diffusion (Barron etal.
2002). Also, biobased packaging made from paper coated with a gluten solution improved the preser-
vation of fresh A. bisporus mushrooms (Guilaume etal. 2010) under MAP at 20°C (80% RH), but the
weight loss was quite high (4%). At last, it should be noted that uctuation of storage temperature, even
if it occurs only once, can restrict the benets from the MAP application (Sandhya 2010).
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510 Handbook of Vegetable Preservation and Processing
22.214.171.124 Effect of Packaging on Mushroom Shelf-Life
Fresh mushrooms are packed in many ways according to wholesale, retail, and transport requirements
and their species characteristics. They are sold as either loose or in trays/punnets overwrapped with
plastic lms, usually PE and PVC of different permeability and stored under refrigeration temperature.
Macroperforated PP lms that limit dehydration without modifying the atmosphere within the package
are also used. However, overperforation may result in excessive mushroom water loss causing wrin-
kling and development of brown patches on the mushroom surfaces (Gormley and MacCanna 1967).
Proper packaging is very important, as it protects mushrooms from bruises throughout transportation
and extends shelf-life through reduction of desiccation and vapor condensation, but also attracts cus-
tomer interest on the product. Design, color, and labeling contribute to the overall impression of quality
(Burton etal. 1987b). However, as most plastic lms available commercially nowadays exhibit too low
gas permeability, there is the need for developing new, environmental friendly microperforated materials
to provide a larger range of O2 and CO2 permeability.
There are two main problems related with mushroom packaging; the creation of an in-package anaero-
bic atmosphere, particular if nonperforated lms are used or in MAP and the water condensation develop-
ment (Gormley and MacCanna 1967; Nichols and Hammond 1973; Roy etal. 1996). Recommendation
for the use of perforated lms has been given by FDA already by 1978 (Herr 1991) so that the growth
of anaerobic C. botulinum and condensation are prevented, fact that however excludes the possibility of
application MA or CA in the packages. Stretchable PVC lm although prevents mushroom water loss,
particular at ambient temperature (18°C), it also favors water condensation on the underside of the lm
at 2°C (Nichols and Hammond 1973) and at 20°C (Guillaume etal. 2010), which is detrimental on the
freshness and consequently the shelf-life of mushrooms. The undesirable condensation observed inside
the packages, created as a result of high humidity due to high transpiration rate of mushrooms and poor
water vapor permeability of the lms, has been studied in relation to different lms used and their effect
on microbial growth, color, and mushroom texture by many researchers (e.g., Gormley and MacCanna
1967; Cliffe-Byrnes and O’Beirne 2007). The optimum humidity level for mushroom packaging was sug-
gested to be 96% (Mahajan etal. 2008b) and for best color 87%–90% (Roy etal. 1996; Wakchaure 2011).
Modied Humidity Packaging is another improved packaging system; as an additional tool to plastic
lms and for better control of the inside package relative humidity, moisture absorbers such as sorbi-
tol (Roy etal. 1995, 1996) and sorbitol and silica gel (Villaescusa and Gil 2003) have been tested for
Agaricus and Pleurotus mushrooms. However, existing moisture absorbers either have low absorption
capacity either and/or absorb moisture quickly and increase mushroom weight loss, not improving the
quality parameters sufciently. Therefore, Mahajan etal. (2008b) suggested a mixed moisture absorber
containing bentonite, sorbitol, and CaCl2 that has high moisture capacity and stays in the powder form
for at least 5–6days of storage. In addition, montmorillonite clay and silica gel can be use to extend the
shelf-life of mushroom in packs (Wakchaure 2011). However, a more practical way to control in-package
humidity would be the use of lms with suitable moisture permeability, for example, hydrophilic (Barron
22.6.2 Processing for Long-Term Preservation
Mushrooms are mainly consumed in their fresh state, but there is also a worldwide trade in the process-
ing of mushrooms (through freezing, canning, pickling, drying, etc.) that alters their nature and extends
shelf-life, allowing the transportation of processed mushrooms to be traded internationally as a com-
modity (Manzi etal. 2001).
Freezing is an increasingly popular method of mushroom preservation that provides storage stability and
allows mushroom consumption year round. It also offers to consumers a product with high nutritional value
and quality attributes. Almost all mushroom species can be consumed as frozen products. The mushroom
freezing process includes preparation steps similar to those used for canning viz. mushroom cleaning,
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washing, cutting, grading/sorting, and blanching. Blast freezing is the most common method used in mush-
rooms, followed by the cryogenic method (Jaworska and Bernaś 2009). Blast freezers rapidly bring the
temperature of foods down, freezing them extremely quickly from −25°C to −30°C, creating small ice crys-
tals that damage less the mushroom cells. Freon, plate, and individual quick freezing are additional meth-
ods used in mushrooms (Coggins and Chamul 2004). Gormley (1972) showed that the white mushrooms
although were frozen more quickly in Freon than in air blast freezing, they had larger drip loss, result quite
surprising, indicating that drip loss of frozen mushroom is dependent on the ush they came from. Once
the mushrooms have been frozen, they can be moved to a more conventional freezer for storage, as long as
the freezer stays cold enough to keep them frozen. However, various factors are involved in the quality loss
during frozen storage of mushrooms, for example, weight loss, undesired color and off-avor development,
loss of rmness, decrease in nutritive value, and the most detrimental of all being the enzymatic browning
reaction (Coggins and Chamul 2004). Czapski and Szudyga (2000) showed that strain remarkably inu-
ences the whiteness of frozen mushrooms. In order to assure good quality frozen mushrooms, preliminary
processing (e.g., blanching or dipping) in solutions is required. Finally, packaging used for quick frozen
vegetables should be in accordance with the relevant provisions of the Code of Practice for the Processing
and Handling of Quick Frozen Foods (CODEX CAC/RCP 8 1976).
Although the amount of fresh mushrooms used for canning has dropped over the last years, about 38%
of them are canned nowadays, holding a major share in world trade (Ravi and Siddiq 2011). Through
canning (sterilization), mushrooms can be stored for a period up to 2 years with storage costs being
relatively low. The white mushroom has been traditionally used for canning, but other species such as P.
ostreatus, L. edodes, and V. volvacea and the wild Cantharellus cibarius, Boletus edulis, and Lactarius
deliciosus are also canned or bottled (Bernaś etal. 2006; Ahlawat and Tewari 2007). Canned mush-
rooms are clearly dened in the “Denitions and Standards of Identity” for canned vegetables in 21 CFR
51.990 (USDA-AMS 1962). The following styles (according to USDA standards) should also be included
as part of the name or in close proximity to the name: “Buttons,” “Sliced Buttons,” “Whole,” “Sliced” or
“Sliced Whole,” “Random Sliced,” “Quarters,” “Stems and Pieces (Cut),” and “Grilling,” as appropriate.
Also, guidelines on the quality of canned mushrooms of special types and containing special ingredients
are given in CODEX STAN 297 (2009). Desalted mushrooms can be also used in canning in air-tight
containers (Czapski 2003).
In order to produce good quality canned mushrooms, these should be processed as soon as possible
after harvest or stored at 4°C–5°C until processed. Nevertheless, storage at low temperature 1 day before
canning was suggested to reduce weight loss and enhance water retention (Beelman etal. 1973). Canning
(and bottling) is an established process of preserving mushrooms in brine, butter, oil, vinegar, etc. that
involves six basic operations viz. selection, trimming, cleaning, blanching, lling, sterilization, cooling,
labeling, packing, and storage (Fan etal. 2005; Ravi and Siddiq 2011). It is important that before sealing
the can, the air is removed and then sterilization (by immersion in water or with steam) follows at 126°C
for 8min (Rodrigo etal. 1999), 121°C–130°C for 15min (Fan etal. 2005), or at 118°C–121°C for 20min
(Vivar-Quintana etal. 1999). The mushrooms with a stem length of 1cm are preferred and are canned
whole, sliced, and stems-and-pieces as per demand (Beelman and Edwards 1989). Some of the qual-
ity characteristics of canned mushrooms are color, weight, and grade (Vivar-Quintana etal. 1999) and
methods such as soaking and blanching in water or in aquatic solution containing various compounds is
the rst measure to limit the negative effects of canning process.
Drying is the oldest and yet one of the most important preservation methods of a number of mushrooms.
It is based on the principle that the water activity of a product is reduced at a specic level (normally less
than 10%) so that it is microbiologically and physicochemically stable (Krokida etal. 2003). Pleurotus,
Lentinula, Volvariella, Agaricus, Auricularia, and most wild mushrooms are commercially dried with
satisfactory rehydration and avor retention. Dehydrated mushrooms are also valuable ingredients in
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512 Handbook of Vegetable Preservation and Processing
a variety of food—products such as snacks, instant soups, and sauces. Taking account the quality of
the drying mushrooms, much attention is paid nowadays in the drying methods utilized, as they affect
their physiology and hence their quality. Textural characteristics, namely, hardness/rmness, cohesive-
ness, springiness, and chewiness of mushrooms usually change during drying (Kotwaliwale etal. 2007).
Color also changes during drying; browning mostly occurs due to enzymatic or nonenzymatic reac-
tions between carbohydrate and amino acids in high temperatures. However, little or no change in the
proximate composition (expressed in dry weight) of B. edulis mushrooms that were air- and freeze-dried
was detected by Jaworska etal. (2014). Color, texture, density, porosity, and sorption are the properties
characterized the dried products affected by temperature and duration of drying (Krokida etal. 1998).
Pretreatments of mushrooms before drying are often utilized for color stabilization, avor enhance-
ment, and texture retention (Singh etal. 2001). These include washing with (chlorinated) water, dipping
in citric acid, sodium chloride, or potassium metabisulte, blanching in hot water, blanching followed
with soaking in whey and curd fermentation (Martínez-Soto etal. 2001; Walde etal. 2006), or steam
blanching followed by sulting and citric acid (Pal and Chakraverty 1997; Kotwaliwale etal. 2007).
Methods for drying mushrooms include the conventional hot air drying, thin layer drying, vacuum
drying, freeze-drying, microwave drying, and the more recently introduced uidized bed and micro-
wave-vacuum drying (see Table 22.6).
The nal temperature and the drying rate are the most important factors of the process. The drying
rate is dependent on several parameters, such as the use of pretreatments, temperature, mushroom thick-
ness, method of drying, and moisture diffusivity (Gothandapani etal. 1997). According to Walde etal.
(2006), the time taken for drying both white and oyster mushrooms from 7.5% (db) to 2.0% (db) was in
the order of vacuum dryer > cabinet moisture dryer > uidized bed dryer > microwave oven, with uid-
ized bed drying being a promising method as regard to time and quality to the faster microwave drying.
Drying kinetics and rehydration characteristics were mainly affected by the microwave power level, fol-
lowed by sample thickness, while system pressure had little effect on drying rate. As for rehydration rate,
it was signicantly affected by the system pressure.
The rehydration characteristics of a dried product also constitute a quality parameter that indicates
if physical or chemical changes occurred during drying, the pretreatments, and sample composition
(Funebo and Ohlsson 1998). Also, the rehydration of the mushrooms is temperature and pressure depen-
dant (Garćia-Segovia etal. 2011). It was suggested therefore that vacuum rehydration of air-dried shii-
take could replace the conventional process as lower immersion time was needed and a more desirable
texture was achieved. The best rehydration process for P. ostreatus occurred at room temperature in
water during 30min, after samples were dried with hot air at 40°C/RH 75% (Apati etal. 2010). Funebo
and Ohlsson (1998) found that A. bisporus mushrooms had greater rehydration capacity using hot air
dehydration without the use of microwaves. Regardless the method, after drying, dehydrated products
are packed in air-tight polyethylene containers and stored at low temperatures. Theoretically, these prod-
ucts have unlimited shelf-life, practically this is over a year, due to signicant quality deterioration after
this time (Jaworska etal. 2014). The great advantage of this method is the lower cost in transportation,
handling, and storage compared to other preservation methods.
22.7 Value–Added Mushroom Products and By-Products
22.7.1 Food, Beverage, and Beauty Mushroom Products
In view of their high perishable nature, the fresh mushrooms have to be processed to extend their shelf-
life for off-season use and also add value to the product. This can be achieved by adopting appropriate
postharvest technology to process surplus mushrooms into novel value-added products like food prepa-
rations (soup powder, pickles, chips, paste and ketchup, pâté, noodles and pasta, biscuits, and nuggets),
mushroom-based avor enhancers or as additives in beverages and beauty products (Zivanovic 2006; Rai
and Arumuganathan 2008). The value-added products are the current need for the mushroom growers
not only to reduce the losses, but also to enhance the income by value-addition and boost the mushroom
consumption (Mehta etal. 2011). Some examples of long preserved food and beverage mushroom prod-
ucts are given below.
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Methods for Drying Fresh Mushrooms
Methods Description References
• Method comparatively cheap, frequently used, involving thermal and/or
chemical pretreatment and then drying at temperatures 50°C–70°C and air
velocities 1.0–5.0 m/s for 3–24h depending the mushroom species, the
product volume, and the chamber size
• Due to long drying time and surface overheating, darkening in color, cellular
rupture, loss in avor, aroma, and dehydration ability may occur
• A combination of a drying air temperature of 50°C and an air velocity of 0.9
m/s for drying untreated and treated (blanching followed by sulting and
citric acid pretreatment) Pleurotus mushrooms was proposed. The texture
and appearance of the untreated mushrooms after rehydration were better
than those of the treated mushrooms, whereas the color and avor of the
treated were better.
Pal and Chakraverty
(1997), Krokida and
(2003), and Giri and
Prasad (2007, 2013)
• Drying temperature and pressure are important and because there is a high
drying rate and low drying temperature and oxygen-decient drying
environment, the quality characteristics of the products (e.g., shape, color,
and aroma) are maintained
• In fresh Lentinula mushroom, vacuum dehydration at 50°C–65°C and
0.1–0.4 bar decreased initial moisture from 92% to 13% (w.b.) in 6h and
revealed that temperature and pressure signicantly affected color
degradation. Rehydration capacity decreased with the increased in vacuum
pressure, yet it was not affected by temperature
• P. ostreatus maintained at 55°C and pressure at 1334 Pa for about 12 h
• Limited use in mushrooms because of their high installation and
(2001), Alibas (2007),
Giri and Prasat (2007),
and Artnaseaw etal.
• Pleurotus mushrooms were rstly washed, frozen at −80°C, and then placed
in the freeze-dryer viz. at 0°C (condenser temperature −55°C, vacuum at 7
Pa) for about 24h. Although avor was not signicantly different,
mushrooms had superior quality as compared to hot air and vacuum-dried
• A. bisporus the quantity of aromatic compound 1-octen 3-ol was
• Limited use in mushrooms because of their high installation and
Kompany and René
etal. (2001), and Giri
and Prasat (2007)
• The quality of both Agaricus and Pleurotus mushrooms was good and the
• The quality of Pleurotus sp. dried in uidized bed condition at 50°C for
80–120min with 0.5 potassium metabisulte was superior and with reduced
microbial spoilage to sun, thin layer drying and uidized bed drying with
• Limited use in mushrooms because of their high installation and
(1997), Walde etal.
(2006), and Giri and
• Is rapid, more uniform, and energy-efcient compared to conventional hot
air drying, but although it combines the advantages of drying at low
temperatures, it has not very satisfactory results due to temperature
instability and tissue hardening
• Temperature control and pressure will contribute to the continued
development of microwave technology applied to food dehydration
• Combination of air-drying and microwave treatment gave encouraging
results with acceptable drying rates provided heat level and heating intensity
are combined properly
Riva etal. (1991),
Walde etal. (2006),
Giri and Prasad (2007),
and Lombraña etal.
• Resulted in 70%–90% decrease in the drying time to a moisture content of
6% (db) and had better rehydration characteristics in sliced white
mushrooms than the hot air dried samples
Giri and Prasad (2007,
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514 Handbook of Vegetable Preservation and Processing
126.96.36.199 Food Products and Food Additives
Mushroom pickling, a procedure that involves fermentation, is an economical way for mushroom pres-
ervation and has a benecial effect on the human organism and impact a pleasant aroma and taste to
pickled food (Zivanovic 2006). Among other mushrooms, Volvariella andPleurotus species are suitable
for pickling. According to Kreb and Lelley (1991), good results are obtained by pickling P. ostreatus for
10days at 21°C with freshly shredded cabbage. In another process (Singh etal. 1995), cleaned mush-
rooms are blanched in hot water (80°C for 5 min), rapidly cooled and added to 60% brine to obtain
mushroom to brine ratio of 7:3 by volume. The mixture is maintained at 15°C–20°C for 15days for fer-
mentation and further kept at 0°C–4°C to obtain a pH of 3.9. Sugar is added to the preparation at the rate
of 3.3% by weight to the brine and nal salt concentration reached to 6.6% by weight. Products of this
type can be stored for 6months at the ambient temperature (Singh and Bano 1977), the pasteurization of
mushrooms before storage being unnecessary. Fermented mushrooms can be also used as a seminished
product in the preparation of marinades and sauces of good quality (Bernś etal. 2006). Pickling is also
an interesting option of saving loss material (about 10%) in the canning process of mushrooms (Rai and
The raw material used for mushroom steeping or marinade (usually A. bisporus,Pleurotus spp., L.
edodes, etc.),being fresh, salted, or pickled mushrooms, is processed with acetic acid or citric acid (2%–
5%). The steeping solution usually contains also salt and sugar (Bernaś etal. 2006). The method is
simple, economical, and mushrooms can be preserved for periods ranging from 3 to 6months by steep-
ing them in concentrated solutions of salts and or acids (Rai and Arumuganathan 2008).
Mushroom paste and ketchup is prepared after boiling the sliced mushrooms in water and grinding
them in a mixer. Then acetic acid, salt, sugar, onion, garlic, pepper, and other ingredients are mixed in
the paste before lling in the sterilized bottles or jars (Rai and Arumuganathan 2008). For mushroom
chips production, mushrooms are sliced (2mm), blanched in 2% brine solution, and dipped overnight
in a solution of 0.1% of citric acid + 1.5% of NaCl + 0.3% of red chilli powder. After draining off
the solution, the mushrooms are subjected to drying at 60°C for 8 h and nally fried in oil (Rai and
Mushroom powder is used as a direct food additive to increase content of dietary bbers in vari-
ous foods or as a partial substitute for wheat our in bakery products. It is obtained after pulveriza-
tion of dried mushroom slices and used to enhance avor of a dish or to provide specic mushroom
aroma for soups, biscuits, nuggets, and snacks preparation (Zivanovic 2006). According to Rai and
Arumuganathan (2008), soup is prepared by mixing the powder with milk power, corn our, and other
ingredients. Biscuits are prepared by mixing mushroom powder with ingredients like sugar, oil, baking
powder, ammonium bicarbonate, salt, vanilla, milk powder, and glucose and nally the required shape is
given to the dough before baking in the oven. Nuggets and snacks are also made (e.g., in India) after mix-
ing the powder with different vegetable (like soybean) powder, water, and spices. Other mushroom-based
food products (like bread, cake, roasted mushrooms in oil, mushroom pâté, etc.) have been presented by
Zivanovic (2006) and Ravi and Siddiq (2011).
188.8.131.52 Beverages and Beauty Products
Flavor and bioactive compounds can be extracted with water and/or alcohol and the extracts can be used
for preparation of mushroom beers, wines, spirits, or prophylactic drinks (Zivanovic 2006). Different
mushroom species, like A. bisporus, L. edodes, and Grifola frondosa, are used alone or in mixtures to
obtain the avor and aroma proles that work well with the alcoholic beverages. In order to utilize the
mushrooms in the process (brewing, wine making, etc.), their carposomes are dried and then ground into
a powder or mushroom mycelia are used. It is well known that Saccharomyces cerevisiae is the main
microorganism used in alcoholic beverage brewing, because this microbe has alcohol dehydrogenase
(ADH) activity. However, some genera of mushrooms produce ADH and wine, beer, and sake can be
made by using mushrooms in place of S. cerevisiae (Okamura-Matsui etal. 2003). According to them,
in wine making 2 g of mushroom mycelia were added to 30 mL of autoclaved grape juice, which was
incubated at 20°C for 40days while in beer brewing 2 g of mushroom mycelia were added to autoclaved
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hopped malt extract medium (pH 5.8) containing 10% malt extract and 0.1% hop extract and incubated
at 20°C for 14days (Okamura-Matsui etal. 2003). The highest alcohol concentrations in the wine, beer,
and sake were achieved with P. ostreatus (12.2%), Tricholoma matsutake (4.6%), and A. blazei (8.0%).
In the case of A. blazei, the produced wine contained about 0.7% β--glucan, which is known to have
preventive effects against cancer. The wine made using F. velutipes showed thrombosis-preventing activ-
ity. Alcoholic drinks based on extracts of G. lucidum and other mushrooms are produced by adding
fresh or dried mushrooms into low-quality distilled spirits with more than 35 proofs and mixing it with
sugar and molasses (Mizuno etal. 1995). Thus, alcoholic beverages made using mushrooms, except of
characteristic avor and aroma, seem to be a functional source which can be expected to have also health
benets (Okamura-Matsui etal. 2003).
Finally, some beauty products, like skin creams and lotions, contain mushroom extracts. For example
L. edodes preparations came in market containing the mushroom compound kojic acid, a natural alter-
native to hydroquinone that improves appearance of skin by bleaching it to fade scars and age spots
(Rahman and Choudhury 2012). Another example of mushroom application is articial skin and wound
cover products, based on the high content of β-glucan and chitin of the fungal cell walls, as these sub-
stances enhance wound healing. The basic of the production of the fungal-based skin patches is in
extensive extraction and washing of mycelia (or fruit bodies), so that only water insoluble biopolymers,
like β-glucan and chitin, remain (Zivanovic 2006). Sua etal. (1997) mention “Sacchachitin” as a product
of this kind, prepared as a woven skin substitute from the mycelium or waste residue of the fruit body of
22.7.2 Dietary Supplements: Nutraceutical Products
Mushroom nutraceuticals are preparations (rened or not) derived from fruit bodies or fungal mycelium
(produced in submerged culture) that possess nutritional and/or health-promoting properties and which
are consumed in the form of tonics, capsules, or tablets as a dietary supplement (Chang and Buswell
1996). Many commercial products from medicinal mushrooms are available worldwide. Some exam-
ples, mentioned by Zivanovic (2006), include “Concord Sunchih” and “Reishi Plus” from G. lucidum,
“Grifon” from G. frondosa, “Didanosine” from Cordyceps militaris, “Calvacin” from Calvatia gigan-
tea, “Lentinan,” “Lentinacin” or “Lentysine,” “KS-2,” “LEM,” and “LAP” from L. edodes, “PSK”
and “Krestin” from Trametes versicolor, “Befungin” from Inonotus obliquus, “Sonilan,” “SPG,” and
“Schizophyllan” from Schizophyllum commune, “ATOM” and “AB-FP” from Agaricus blazei, and
“Lovastatin” from P. ostreatus. These products include water or alcohol extracts, concentrates, and pow-
ders in bulk or tablet form (Zivanovic 2006).
Mushroom substances include low-molecular-weight compounds (LMW, e.g., quinones, cerebrosides,
isoavones, catechols, amines, triacylglycerols, sesquiterpenes, steroids, organic germanium, and sele-
nium) and high-molecular-weight compounds (HMW, e.g., homo- and heteropolysaccharides, glyco-
proteins, glycopeptides, proteins, and RNA–protein complexes) (Ferreira etal. 2010). The mushrooms
might be used directly in the diet to promote health, taking advantage of the additive and synergistic
effects of the mentioned bioactive compounds present in them. The majority of mushroom-based nutra-
ceutical products are not single bioactive compounds but combinations of several individual components
that together contribute to the overall bioactivity of the product (Reis etal. 2012). It is believed that
combinations of these bioactive components target the human immune system as well as aid in neuron
transmission, metabolism, hormonal balance, and the transport of nutrients and oxygen.
Mushroom immunomodulatory effects are mostly attributed to β-glucans. The β-glucans are of par-
ticular interest since they are naturally occurring polymers of glucose, are orally active when taken as
food supplements, and have a long track record of safe use (Chan etal. 2009). Due to their immuno-
modulatory properties, puried β-glucans have been used clinically as part of a combination therapy
for a variety of cancers (Thompson etal. 2010). In Japan, Russia, China, and the United States several
different polysaccharide antitumor agents have been developed from the fruiting body, mycelia, and
culture medium of various medicinal mushrooms (L. edodes, G. lucidum, S. commune, T. versicolor, I.
obliquus, Flammulina velutipes, etc.). Mushroom polysaccharides have recently attracted attention and it
is not surprising that a number of companies process medicinal mushrooms into extracts, which can then
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516 Handbook of Vegetable Preservation and Processing
be used in the production of health foods, drinks, and dietary supplements. As a result, functional food
market reached an approximately 50%–60% growth in value sales over a 5-year period, being estimated
in excess of €10 billion in Europe (Van Griensven 2013).
Regarding clinical studies of antitumor activity and immunomodulating action of biologically active
metabolites, mostly polysaccharides (β-glucans) from mushroom fruit bodies and/or cultured mycelium,
lentinan from L. edodes, schizophyllan from S. commune, MD-fraction from G. frondosa, and com-
pounds from T. versicolor (PSK and PSP) have been in clinical use, especially in Japan and China,
for the adjuvant tumor therapy (immunotherapy) in addition to the major cancer therapies like surgi-
cal operation, radiotherapy, and chemotherapy (Lindequist etal. 2005). Among mushrooms, L . edodes
(shiitake) has been studied most extensively (antitumor and immunostimulating properties revised by
Shen etal. (2011), while epidemiological evidence has shown a correlation between daily mushroom
consumption and a low rate of cancer mortality in Japan. Application of lentinan (parenteral) in addi-
tion to chemotherapy led to prolongation of survival time, restoration of immunological parameters,
and improvement of life quality in patients with stomach cancer, colon cancer, and other carcinomas in
comparison to patients who had chemotherapy alone (Lindequist etal. 2005). Moreover, the Phellinus
linteus extract antitumor activity has been demonstrated by Collins etal. (2006). The anticancer drug
doxorubicin (Dox), known to induce apoptosis (programmed cell death) in cancer cells, can damage
healthy cells in higher doses. The effect of P. linteus extract on Dox-induced apoptosis was investigated
in prostate cancer cells (Collins etal. 2006). The research showed that P. linteus or Dox, at relatively
low doses, could not kill these cells. However, combination treatment at low doses of P. linteus and Dox
results in a synergistic effect and brought about death in prostate cells. These ndings indicate that P.
linteus has a synergistic effect with Dox to activate a benecial decline in prostate cancer cells, indicat-
ing its therapeutic potential. Additionally, it has been shown that applying extracts of T. versicolor to
postoperative immunochemotherapy has led to signicant advantages in survival over chemotherapy
alone. Next to a direct inuence on survival, mushroom extracts could possibly affect the side effects of
radio- and chemotherapy (Van Griensven 2013).
22.7.3 Mushroom Industry Spent By-Products
Mushroom harvesting furnishes a great amount of waste consisting mainly of stipes (stems or stalks) and
mushrooms of irregular shape. Depending on the size of the mushroom farm, the amount of waste ranges
between 20% and 30% of production volume. This results in thousands of metric tons of waste material
per year with no suitable commercial use. A great part of mushroom wastes consist of the bases or stipes
that have tough texture and are usually considered to be a waste product when mushrooms are harvested.
These cut-off bases or stipes make up about 25%–33% of the weight of fresh mushrooms, and they are
normally used to make low-economic value animal feed and compost (Chou etal. 2013). However, this
fruit body waste, apart from the nutritional use as animal feed, is a good source of insoluble dietary bers
and glucans that can be used for the preparation of biologically active polysaccharide complexes utiliz-
able as food supplements (Synytsya etal. 2008). Also, stipe residue is a potential source of fungal chitin,
chitosan, and their derivatives that can be used as an antimicrobial, emulsifying, thickening, and stabiliz-
ing agent in food industry (Yen and Mau 2007). Additionally, the underutilized wastes, which are good
nitrogen sources, have been found to have a high content of polysaccharides, which exhibited prebiotic
activity and might become an important source of novel prebiotics (Synytsya etal. 2009; Chou etal. 2013).
Moreover, mushroom industries generate a big supply of an organic by-product called spent mushroom
substrate (SMS). This is the unutilized substrate (composted agro-residues) and the mushroom mycelium
left after harvesting of mushrooms (1kg of mushrooms usually generates 5kg of SMS). As the mush-
room industry is steadily growing, the volume of SMS generated annually is increasing along with the
concerns of its environmental impact (the usual disposal strategy of SMS is by burning, spreading on
land, burying, composting, or land-lling). Actually, the mushroom industry faces challenges in storing
and disposing the SMS, using environmentally friendly practices. These challenges include the relatively
high salt and water content of SMS, its bulkiness (resulting in high transportation costs).
However, SMS has desirable chemical, physical, and biological properties that could be explored for
new value-added uses and products, such as soil conditioner and organic fertilizer, land restoration,
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casing for mushroom production, plant disease control, bedding for livestock housing, feedstock for heat
and power production, in vermiculture as a growing medium, bioremediation using SMS as a source
of crude oxidizing enzymes like laccase (restoration of contaminated soils and water, e.g., from bio-
cides, fungicides, and phenolic compounds), production of lignocellulosic enzymes such as laccase, xyla-
nase, lignin peroxidase, cellulase, and hemicellulase (Rinker 2002; Suess and Curtis 2006; Rigas etal.
2009; Phan and Sabaratnam 2012; Philippoussis and Diamantopoulou 2012; Papadopoulou etal. 2013).
Furthermore, numerous studies have shown the feasibility of using SMS to produce animal feed (Rinker
2002), mainly the use of the protein-enriched residues of Pleurotus spp. production as forage source in
maintenance rations for ruminants (Bae etal. 2006; Kim etal. 2010).
This chapter contains an integrated presentation of mushrooms’ physiology and particular features and
how these affect their storage and processing, indicating that edible mushrooms are difcult materials
for storage and processing. However, there are numerous methods for mushroom processing, the most
popular of which being drying, canning, and freezing. Under this view and keeping in mind that mush-
rooms are important mainstream for production of food and nonfood items, further innovation of the
sector is needed along with constant scientic and market research, development, and technical support.
Mushroom harvesting wastes and SMS need to be reclassied as by-products of the mushroom growing
process and not as wastes as they can be used in many biotechnological and environmental applications.
Moreover, as there is increasing interest worldwide in the many bioactive compounds and metabolites
that are found in cultivated and wild fungi, a large number of mushrooms have been studied and used
as food additives, as a source of biologically active constituents (contained in fruiting bodies or cul-
tured mycelia) or health food supplements. However, additional work is needed toward exploitation of
the health benecial compounds in mushrooms from a pharmaceutical and functional-food perspective.
Future medicinal mushroom research should provide trustworthy results of clinical tests in humans as
well as information on the effects of various cultivation, storage, and preservation treatments on mush-
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[AQ1] Please provide chapter title for reference “Chang and Miles (2004).”
[AQ2] Please check if the sentence “Apart...” conveys the intended meaning.
[AQ3] Please check if the usage of the word “carposomes” is appropriate here.
[AQ4] Please check if the sentence “These...” conveys the intended meaning.
[AQ5] Please check if the sentence “However...” conveys the intended meaning.
[AQ6] Please provide chapter title for reference “Bord Glas (2002)” if appropriate.
[AQ7] Please provide editor(s) details for references “Cha (2004), Chen (2005), Curvetto etal. (2005),
Fan etal. (2005), Przybylowicz and Donoghue (1988), Royse (2004), and Van Griensven (2013).”
[AQ8] Please provide chapter title for reference “Chang and Miles (2004).”
[AQ9] Please provide title for reference “FAOSTAT (2014).”
[AQ10] Please check the inserted city in “Shen etal. (2011)” for correctness.
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