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Cultivated Mushrooms: Preservation and Processing
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22
Cultivated Mushrooms:
Preservation and Processing
Panagiota A. Diamantopoulou and Philippoussis N. Antonios
Contents
22.1 Introduction................................................................................................................................... 495
22.2 Mushroom Composition and Nutritional Aspects........................................................................ 496
22.3 Health Benefits 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
22.6.1.1 Cooling: Refrigeration...................................................................................... 504
22.6.1.2 Minimal Processing.......................................................................................... 506
22.6.1.3 Effect of Packaging on Mushroom Shelf-Life...................................................510
22.6.2 Processing for Long-Term Preservation............................................................................510
22.6.2.1 Freezing.............................................................................................................510
22.6.2.2 Canning..............................................................................................................511
22.6.2.3 Drying................................................................................................................511
22.7 Value–Added Mushroom Products and By-Products....................................................................512
22.7.1 Food, Beverage, and Beauty Mushroom Products............................................................512
22.7.1.1 Food Products and Food Additives....................................................................514
22.7.1.2 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
References................................................................................................................................................517
22.1 Introduction
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 specific aroma, texture, and taste. According to the data of FAOSTAT (2014), total
world production of mushrooms (including truffles) 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
495
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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 fiber, vitamins, and
minerals (Crisan and Sands 1978; Bano et al. 1988; Barros et al. 2008). Apart from valued compounds
affecting taste, mushrooms are nutritionally desirable because of their low energy value, fiber 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 et al. (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 et al. (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 deficient in sulfur-containing amino acids, including methionine and cysteine (Mattila et al. 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 fiber, comprising mainly water-insoluble fiber
in the form of chitin (polymer of N-acetyl-glucosamine) and nonstarch polysaccharides like β-glucans
(Sadler 2003). The water-soluble dietary fiber 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 et al. 2010; Reis et al. 2012). Among them, according to Diamantopoulou
et al. (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 significant quantities of oleic acid Δ9C18:1. Palmitic
acid is the main saturated fatty acid (Diamantopoulou et al. 2012b, 2014). Mushrooms are fairly good
source of vitamins, particularly thiamine, riboflavin, 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, detoxification, hepatoprotective, and antidiabetic activities,
thoroughly reviewed by many (Wasser and Weis 1999; Wasser 2002; Sadler 2003; Lindequist et al. 2005;
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Cultivated Mushrooms 497
Cheung 2008; Patel and Goyal 2012; Rathee et al. 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 classified 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 et al. 2003). For example, L. edodes has been reported to possess antitumor,
antihypertensive, hypocholesterolemic, and antibacterial activities (Israilides et al. 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 et al. 2006; Ramberg et al. 2010; Wasser 2010b). Mushrooms of Pleurotus species were
reported to have hypocholesterolemic, anti-inflammatory, and immunostimulatory activities (Alam et al.
2009, 2011; Patel et al. 2012). The potential therapeutic implications of mushrooms are enormous, but
detailed mechanisms of the various health benefits of mushrooms to humans still require intensive inves-
tigation. Isolation of their active ingredients, with mechanism-based potential therapeutic value, remains
a challenge (Rathee et al. 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 filling the room with the inoculated substrate, highlighting significant 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 influence the productivity and the quality of the final 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
et al. 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 et al. 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 et al. 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 fructification
and to favor the development of mushrooms (Van Griensven 1988; Sánchez 2004; Ahlawat 2011). The
first pin initials begin to appear about 2 weeks after casing. From one layer of compost, two to three
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crops (called flushes) are harvested. In commercial practice, mushrooms are harvested before the
stage of maturation and death and flushes develop almost at weekly intervals (Van Griensven 1988;
Moore and Chiu 2001; Straatsma et al. 2013).
Yields are influenced 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 et al. 2001a; Straatsma et al. 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 first of all determined by the quality and amount
of compost filled 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 et al. 2013).
As far as harvesting period is concerned, it commences at the first sign of buttons, often on a 7–10
day cycle and usually lasts for 1–1.5 months. Mushrooms may be picked at the button (small unopened
mushrooms), cup (the cap has begun to open), or flat (fully expanded caps exposing all of the gills) stage
depending on market requirements. Timing is important as mushrooms grow quickly, doubling their size
within 24 h. 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 first flush occur on
the first 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 flush most problems occur on the final
picking day (ripening) and with the third flush 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 et al. 1998; Diamantopoulou and Philippoussis
2001; Chikthimmah et al. 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 et al. 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
flush, 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. flabellatus) 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 et al. 2001b;
Royse 2004). The spawned substrate is filled into perforated polyethylene—PE blocks (bags and bottles
are also used) and at the end of the spawn run period, fructification of P. ostreatus is triggered by lower-
ing the air temperature and CO2 levels, while light essential for pinning (8–12 h 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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Table 22.1
Preharvest and Harvesting Methods and Techniques Used in Agaricus spp. Cultivation to Promote
Mushroom Productivity and Quality
Parameters
and Methods Description of the Techniques and Target Results
Preharvesting Compost • Compost quality and well-filling of beds (e.g., edges of the bed)
period • The filling weight is determinant for both the quality and quantity of the second
flush. Filling less than 90 kg of compost/m2 usually results in quick lose of activity
and in lighter mushrooms of the second flush that mature too quickly
Casing • A rougher textured (good moisture retention properties) and wetter casing soil gives
soil— better mycelium quality and good quality of the later flushes mushrooms
moisture • 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
bodies
Ruffling • Serves to increase mycelium quantity in the casing soil. This results to heavier
mushrooms and a better spread of the flush. Better planning for the first day of
picking, improved uniformity over the beds, and higher production are achieved.
• Lightly ruffled casing soil produces less mycelium with less pinheads and greater
percentage of large mushrooms (cap diameter >45–50 mm)
• Deep ruffling (up to 1 cm 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
Evaporation— • Control of the evaporation between spraying actions has better watering results and
Growing enhances mushroom yield (mushroom number and crop timing) and quality (by
room climate minimizing mushroom discoloration and internal moisture)
Pinhead • Determining the moment of pin formation, the number of pinheads, the amount, and
formation size of mushrooms (by air temperature and CO2) is crucial for the harvest
Harvesting Thinning of • The formation of clusters of mushrooms, between which a too high CO2 content is
period clusters 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 first picking day thinning has to be done by removing all mushrooms with a
diameter of 25–35 mm 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
Selective • The mushrooms are harvested up to the max diameter (heavier mushrooms) and by
picking— picking several times a day the yield (~2–3 kg/m2) and quality (~5%) is increased
organized • Provide the unpicked mushrooms with space and nutrition by higher picking
harvesting frequency. The result is higher productivity, better quality (up to 85%) and color, and
longer shelf-life
Moisture • Moisture management during the first flush is essential for good quality mycelium
management that transports the nutrients required, has positive effect on the second flush 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, AQ1
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. et al., Fungal Biol.,
117, 697, 2013.
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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 et al. 2001b; Royse 2002; Mandeel et al. 2005; Kirbag and Akyuz
2008; Onuoha et al. 2009; Philippoussis 2009; Sánchez 2009; Fanadzo et al. 2010). An average
biological efficiency (kg of fresh mushrooms per 100 kg 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 3–5 days (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 efficiencies (BE) are achieved with sunflower 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)
(Philippoussis 2009).
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, flavor, 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 flies 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 et al. (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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Table 22.2
K21711_C022.indd 501
Management of Parameters and Cultivation Procedure in Different Stages of Growth and Fruiting of Lentinula edodes Grown in Bags
Relative
Stages of Temperature Humidity
Cultivation (°C) (%) CO2 (ppm) Light (lux) Duration Cultivation Procedures and Remarks
Substrate 21–27 90–100 1,500–10,000 No or short 30–45 days • Substrate type and its nutrients as well as texture-porosity (aeration) affect
Cultivated Mushrooms
inoculation exposure incubation/preharvest period and yielda,b,c

and mycelium 50–100 • Spawn-run: strain-dependant duration, there are cold and warm temperature strainsa,c,d
growth (1–4 h/day) • Toward the later stages of the spawn run 1, a thick mycelial coat forms on the outer
(spawn-run 1) surface of the colonized substrate blocka,b,d
Mycelial 21–27 70–90 >1,500 No or short 45–60 days • Clumps of mycelia (bumps) and early primordia are produceda,d
maturation exposure • Great numbers of bumps (prone to abortion) must be eliminated by aeration (cutting
(bumping and 50–100 slits on bags)
browning; (1–4 h/day) • Bag removal for browning of the mycelial coat and bark formationa,b,d
spawn-run 2)
Primordia 10–21 90–100 <1,000 500–2000 5–7 days • Fruiting induction by aeration and cold-shock through: water soaking (4–24 h at
formation 80–90 (12 h/day) 12°C)a,b,e, or temp. reduction to 15°Ca,b,c,d
• Temperature flactuationa,d,e
Fruit body 16–18 98–100 <1,000 500–2000 5–7 days • Relative humidity (RH) fluctuation during fruit body developmente
development (12 h/day) • RH is lowered to 60% for 6–12 h before crop harvesting for better shelf-life of
mushroomsa,e
Harvest of first 16–18 60–70 <1,000 500–2000 5–10 days • Harvest when the edge of the mushroom cap is still in-rolled, or when it is only
flush (12 h/day) partly extendedd
• Trimming the end of the stalk and cutting off residual stalk stubs from the
substratea,d,e
Interval 18–21 50–60 >1,500 Short 15 days • Lower the humidity to 30%–50% RH at 21°C during rest-periode
between exposure • For the second flush: soaking the substrate block for up to 12 h and incubation for
flushes 1 week at higher temperaturesb,c,d,f
Harvest of 16–18 60–70 <1,000 500–2000 5–10 days • After cooling to 16°C for 1 week a second flush will be harvestedc,d,f
second flush (12 h/day)
a Chen (2001).
b Royse (2004).
c Philippoussis et al. (2003).
d Chen (2005).
e Stamets (2000).
f Philippoussis et al. (2007).
501
4/30/2015 12:32:19 AM
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, firmness, maturity
stage, clearness, blemish-free, flavor, nutritional value, and safety and are affected by preharvest treat-
ments (see Section 22.4) as much as postharvest processing and storage conditions (Burton et al. 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 et al. 2008). Throughout storage, mushroom quality is usually assessed by pileus
color, transpiration and respiration rate, weight loss, and disease incidence (Burton et al. 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 reflect 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 influenced by the rate of local market development. For example, a reflectance
(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 et al. 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-
specific 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 et al. 1999; Brennan et al. 2000; Mahajan et al. 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 et al. 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 et al. 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
firmness. 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 et al. 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
Table 22.3
Effect of Respiration on Postharvest Physiology and Storage of Mushrooms
Effect Results References
Degradation and depletion of Mushroom deterioration associated with: Hammond and Nichols
soluble compounds (mannitol, • loss of texture-softening (1975) and Tseng and
trehalose, glycogen, fructose, • brown coloration in Mau (1999)
and proteins) Agaricus sp.—mucilage in Pleurotus sp., pileus
opening in Volvariella sp.
Movement of dry matter and Loss of texture Hammond and Nickols
water from stipe to/and pileus (1975) and McGarry
and gills and Burton (1994)
Changes in the nutritional and • Decrease of protein content with accumulation of total free Hammond (1979) and
medicinal attributes of amino acids Tseng and Mau
mushrooms during storage • Decrease in free amino acids (after the activation of (1999)
and temperatures 18°C–20°C 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)
Condensation of vapor is Texture changes—softening of the flesh Nichols (1985) and
observed in the inner package Mahajan et al.
when nonperforated or even (2008b)
the conventional PVC
stretchable films are used
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bacteria were detected in cultivated mushrooms (Venturini et al. 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

22.6.1.1 Cooling: Refrigeration
Mushrooms at ambient temperatures (ca. 22°C) have a short shelf-life of 1–3 days (Burton and Twyning
1989), at 15°C their shelf-life is 2–3 days (Gormley 1981), whereas in the tropics they count only 24 h
(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 field 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 firm,
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 10 kg of mushrooms or with 15 cm 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 6 days. It is
widely accepted that the ideal conditions for retarding the metabolic activities of tissues and decreas-
ing microbial growth up to 9 days are storage temperature 0°C–2°C and relative humidity 85%–90%
(for A. bisporus and L. edodes mushrooms; Beelman et al. 1973; Nichols 1985; López-Briones et al.
1992; Minato et al. 1999). Gormley (1975) indicated that white mushrooms can be stored for 7–9 days
at 0°C–1°C. The shelf-life of fresh sliced white mushrooms is shorter, mainly because of the effects
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Table 22.4
Methods for Precooling Fresh Mushrooms
Methods Features References
Air-cooling • Air velocity at least 60 m/min Nichols (1985) and
• Mushrooms usually placed in boxes, various packages or punnets TNAU (2014)
and cooled within 24 h, in a rate of 0.5°C/h, but unevenly
• Air is usually dehumidified 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 films (e.g., PVC) in mushroom packaging is essential for
preventing great weight losses even if it hampers the quick cooling
Forced cooling • Higher air pressure is developed on one side of the packages than Wakchaure (2011) and
the other and air is directed through the produce cooling it rapidly TNAU (2014)
• 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
Ice bank cooling • Air used to cool mushrooms is almost saturated as it has previously Nichols (1985)
passed through ice-cold water, cooled by direct heat exchange with it
and filled with vapor, a system preventing mushrooms from water loss
Vacuum cooling • It is a rapid and uniform (temperature reduction from ambient to Gormley (1975), Burton
2°C in 15–25 min), yet expensive method et al. (1987a), McGarry
• Subjecting mushrooms at very low pressures water from their and Burton (1994), and
surface is evaporated resulting in lower temperatures TNAU (2014)
• 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 films is essential for maintaining the
appearance of vacuum cooled mushrooms
• Adding reselected amount of water before or during precooling can
prevent weight loss
Hydrocooling • Large butches of mushrooms are spayed or immersed with/or chilled Nichols (1985) and
or cold water (~0°C) before further packing TNAU (2014)
• It is a rapid cooling technique (the high heat-transfer rates remove
field heat at 20–30 min) 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
Evaporative cooling • Inexpensive method based on the cooling effect created by TNAU (2014)
evaporation of water when dry air is shown over the wet product.
Suitable with areas with low ambient humidity (<65% RH) air
of washing and cutting process (Brennan et al. 2000), for example, 7.5 and 4 days at 0°C and 5°C,
respectively (Oliveira et al. 2012). The shelf-life of the oyster mushroom can be 9 days at 2°C or 3 days
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 3 days (Ahlawat and Tewari 2007). Finally,
optimum in-package O2 is 6% to reduce pileus opening (Roy et al. 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).
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22.6.1.2 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 Modified Atmosphere Packaging (MAP), Controlled Atmosphere
Packaging (CAP), washing, use of chemicals, blanching, radiation, use of moisture absorbers (e.g., sor-
bitol), or coatings (Beelman et al. 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 films 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 sulfites 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
sulfite, 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 et al. 2009), or treating L. edodes mushrooms with chi-
tosan–glucose complex coating and then packaging in low-density polyethylene film (Jiang et al. 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-
flavors 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 filling operation (Wu et al. 1981). However, some negative features of blanching are
the extraction of useful nutritious components (Wu et al. 1981; Gormley 1984), the undesired changes
in aroma (Le Loch-Bonazzi and Wolff 1991; Mau et al. 1992), the lower taste compared to fresh cooked
mushrooms (Wu et al. 1981), the damages of pileus’ texture that cause remarkable toughness after thaw-
ing and cooking (Gormley and Walsh 1982; Reyes De Corcuera et al. 2004), the mushroom weight loss
during the blanching procedure, and the increase in mannitol concentration in mushroom mass and
blanching fluid (Biekman et al. 1996).
To avoid general biochemical effects, blanching time should be kept to the minimum (Matser et al.
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 15 min (Coşkuner and Özdemir 2000). Temperatures
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Table 22.5
Chemical Compounds Used in the Preliminary Processing of Mushrooms
Compounds Features References
Oxine (stabilized chlorine • Very effective antimicrobial agent (50 ppm with a 2 min or longer Beelman (1987),
dioxide) wash period at ~12°C), but with variable results on whiteness Brennan and
• Its activity is due primarily to oxidation not chlorination and Gormley (1998),
although it is up to five times more effective, it is noncorrosive and Wakchaure
(2011)
Hydrogen peroxide • Known (5%, v/v) for its antibacterial and whitening actions in Brennan and
(H2O2) mushrooms Gormley (1998)
• Whole fresh mushrooms soaked for 10 min in solutions of H2O2 and Brennan (1999)
(or citric acid) then sliced, packed, and stored at 4°C for up to
19 days. Both treatments reduced the number of pseudomonad
bacteria and improved their quality of when compared to control
(water soaked) slices
Varsenic (EDTA) and • Both acidic and metal chelators, having the ability to inhibit Brennan and
citric acid enzymatic browning and microbial growth (Pseudomonas sp.) on Gormley (1998)
mushrooms
• Compared to H2O2, they are easier to handle, not corrosive and do
not deteriorate with age
• Fresh sliced mushrooms were soaked for 10 min 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
the panel
• As citric acid is the most widely used in foods and current
legislation supports its use over H2O2 or EDTA, it is the treatment
most recommended
Oxine (50 ppm), sodium • Beneficial effect on color and number of bacteria Wakchaure (2011)
erythorbate (0.1%) and
calcium chloride (0.5%)
Ascorbic acid (AC) and • AC is an antioxidant and reducing pH agent with wide use as Beelman et al.
calcium chloride (CaCl2) pretreatment for mushroom extension of shelf-life and CaCl2 is a (1973) and
firming agent Niazmand et al.
(2009)
Sodium metabisulfite • Extensively used in mushroom industry as whitening and Brennan and
(SM) preservative agent (1 g/L for 1–2 min), recommended for frozen Gormley (1998),
and sterilized products Brennan (1999),
• It may result in potentially harmful sulfur dioxide residues Czapski and
• Fresh mushrooms washed in 0.05% SM not only improved their Szudyga (2000),
initial whiteness, but also kept it for longer during storage and Wakchaure
• Washing A. bisporus mushrooms with water containing 3 g/L SM (2011)
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 • Positive effect on the color of A. bisporus pileus and general Czapski (2001)
jasmonate ethyl alcohol appearance during storage at 13°C/5 days
(sole or in combination)
commonly used in industry for blanching (90°C–100°C) can cause undesirable tissue softening; how-
ever, the blanching with boiling water for 2 min caused increased hardening in the oyster mushroom
(Kotwaliwale et al. 2007) and better color when it was freeze-dried (Fang et al. 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 et al. 2014). According to Fan et al. (2005), mushrooms
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of the genus Lentinula before canning are usually blanched in water at 90°C for 5 min, 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 8–10 min (Vivar-Quintana et al. 1999), 1% NaCl and 0.1%
citric acid at 95°C–100°C for 5–6 min (Wakchaure 2011), or 1 g/L NaCl and various concentrations of
citric acid or EDTA at 95°C–100°C for 15 min (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 et al. 1999). Prior to freezing, B. edulis
mushrooms were blanched at 96°C–98°C (3 min the pilei and 1.5 min 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 1 h 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 8 months (as compared to
unblanched samples); nevertheless, both color and nutritional value were better in fresh than in mush-
AQ2 rooms pretreated and frozen (Jaworska et al. 2008). Apart from the enzyme inhibitors mentioned above
(NaCl, organic acids, and EDTA) blanching may also contain metabisulfite and ascorbic acid hydrogen
peroxide (Coşkuner and Özdemir 2000; Czapski 2002). Blanching reduced the attractiveness of the
dry P. ostreatus, whereas sodium metabisulfite improved it (Martínez-Soto et al. 2001). The blanching
process of P. ostreatus with hot water was optimized at 150 s and 70°C for mushrooms prior to freezing
(Vullioud et al. 2011), or at 1 min and 70°C–75°C for those immersed at 35% brine solution and then
preserved at tropical room temperature for 6 months (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 efficient 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 l-ascorbic acid (Beelman et al. 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 et al. 1986). Additionally, in order to avoid the undesirable texture changes but obtain a
white color, mushrooms prior to freezing were dipped in metabisulfite solutions (0.5% and 1.0%) for 5 min
after blanching (Fang et al. 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 et al. (2010) reported that combina-
tion of gamma radiation (1.0 kGy) and passive MAP extended the shelf-life of Lentinula up to 20 days.
Its use is limited by the cost and the perception of consumers to irradiation on food (Przybylowicz and
Donoghue 1988).
MAP and CAP include the use of plastic permeable film 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 et al. 2006). In order to devise an MA package, which provides optimum gas conditions, it is
necessary to select a film with suitable gas permeability properties, not only to design a pack with the
appropriate ratio of weight of product to surface area of film, 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 et al. 1995; Simón et al. 2005; Kim
et al. 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 modification, where the required CO2 is produced
by the mushrooms and O2 is consumed with respiration, the appropriate film permeability allows the
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entrance of required O2 and the exit of surplus CO2 until eventually equilibrium is established. In active
modification, the atmosphere inside the packages is achieved by, firstly, 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 et al. (2008) showed that the equilibrium atmosphere con-
ditions for both O2 and CO2 were reached for active modified atmosphere packages containing L. edodes
mushrooms after 10 days 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 et al. (1992) suggested that to optimize marketing
conditions for white mushrooms, the storage atmosphere should contain 5%–10% O2 and 2.5%–5% CO2.
The films 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 films were used in prepacks,
concentrations depended on the film 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 modified atmosphere cre-
ated in the packages affect bacterial growth and consequent color change (Burton et al. 1987b), film
perforation is strongly suggested along with the use of new technology films with chosen (in advance)
permeability (Falguera et al. 2011).
Regarding research on fresh sliced A. bisporus mushrooms MAP, using PVC and PP films 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 et al. 2005). Nevertheless, with storage temperature
increase more perforations are needed in order to obtain the optimum MAP conditions for sliced mush-
rooms (Oliveira et al. 2012). On the other hand, MAP did not reduce mushroom catabolism (Varoquaux
et al. 1999) and induced internal and external discoloration of carposomes (López Briones et al. 1992) AQ3
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 7 days at 4°C. Low O2 (2%) and high CO2
(30%) concentrations significantly 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 florida (Jafri et al. 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 et al. (2006) using three different films (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 finding of Parentelli et al. (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 12 days of storage, whereas when macroperforated films were used, the
weight loss was as high as 15% (Antmann et al. 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 et al. 2008).
Additionally, the unique gas barrier properties of hydrophilic films (wheat gluten-based material and
synthetic polymer) have been tested through the MAP of A. bisporus mushrooms packed under micro-
porous and hydrophilic films 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 film used, owing to their high selectivity to gas diffusion (Barron et al.
2002). Also, biobased packaging made from paper coated with a gluten solution improved the preser-
vation of fresh A. bisporus mushrooms (Guilaume et al. 2010) under MAP at 20°C (80% RH), but the
weight loss was quite high (4%). At last, it should be noted that fluctuation of storage temperature, even
if it occurs only once, can restrict the benefits from the MAP application (Sandhya 2010).
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22.6.1.3 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 films, usually PE and PVC of different permeability and stored under refrigeration temperature.
Macroperforated PP films 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 et al. 1987b). However, as most plastic films 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 films are used or in MAP and the water condensation develop-
ment (Gormley and MacCanna 1967; Nichols and Hammond 1973; Roy et al. 1996). Recommendation
for the use of perforated films 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 film although prevents mushroom water loss,
particular at ambient temperature (18°C), it also favors water condensation on the underside of the film
at 2°C (Nichols and Hammond 1973) and at 20°C (Guillaume et al. 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 films, has been studied in relation to different films 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 et al. 2008b) and for best color 87%–90% (Roy et al. 1996; Wakchaure 2011).
Modified Humidity Packaging is another improved packaging system; as an additional tool to plastic
films and for better control of the inside package relative humidity, moisture absorbers such as sorbi-
tol (Roy et al. 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 sufficiently. Therefore, Mahajan et al. (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 5–6 days 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 films with suitable moisture permeability, for example, hydrophilic (Barron
et al. 2002).
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 et al. 2001).
22.6.2.1 Freezing
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 flush 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-flavor development,
loss of firmness, 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 influ-
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).
22.6.2.2 Canning
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 et al. 2006; Ahlawat and Tewari 2007). Canned mush-
rooms are clearly defined in the “Definitions 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 et al. 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, filling, sterilization, cooling,
labeling, packing, and storage (Fan et al. 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 8 min (Rodrigo et al. 1999), 121°C–130°C for 15 min (Fan et al. 2005), or at 118°C–121°C for 20 min
(Vivar-Quintana et al. 1999). The mushrooms with a stem length of 1 cm 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 et al. 1999) and
methods such as soaking and blanching in water or in aquatic solution containing various compounds is
the first measure to limit the negative effects of canning process.
22.6.2.3 Drying
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 specific level (normally less
than 10%) so that it is microbiologically and physicochemically stable (Krokida et al. 2003). Pleurotus,
Lentinula, Volvariella, Agaricus, Auricularia, and most wild mushrooms are commercially dried with
satisfactory rehydration and flavor retention. Dehydrated mushrooms are also valuable ingredients in
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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/firmness, cohesive-
ness, springiness, and chewiness of mushrooms usually change during drying (Kotwaliwale et al. 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 et al. (2014). Color, texture, density, porosity, and sorption are the properties
characterized the dried products affected by temperature and duration of drying (Krokida et al. 1998).
Pretreatments of mushrooms before drying are often utilized for color stabilization, flavor enhance-
ment, and texture retention (Singh et al. 2001). These include washing with (chlorinated) water, dipping
in citric acid, sodium chloride, or potassium metabisulfite, blanching in hot water, blanching followed
with soaking in whey and curd fermentation (Martínez-Soto et al. 2001; Walde et al. 2006), or steam
blanching followed by sulfiting and citric acid (Pal and Chakraverty 1997; Kotwaliwale et al. 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 fluidized bed and micro-
wave-vacuum drying (see Table 22.6).
The final 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 et al. 1997). According to Walde et al.
(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 > fluidized bed dryer > microwave oven, with fluid-
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 significantly 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 et al. 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 30 min, after samples were dried with hot air at 40°C/RH 75% (Apati et al. 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 significant quality deterioration after
this time (Jaworska et al. 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 flavor 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 et al. 2011). Some examples of long preserved food and beverage mushroom prod-
ucts are given below.
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Table 22.6
Methods for Drying Fresh Mushrooms
Methods Description References
Hot air • Method comparatively cheap, frequently used, involving thermal and/or Pal and Chakraverty
drying chemical pretreatment and then drying at temperatures 50°C–70°C and air (1997), Krokida and
velocities 1.0–5.0 m/s for 3–24 h depending the mushroom species, the Marinos-Kouris
product volume, and the chamber size (2003), and Giri and
• Due to long drying time and surface overheating, darkening in color, cellular Prasad (2007, 2013)
rupture, loss in flavor, 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 sulfiting 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 flavor of the
treated were better.
Vacuum • Drying temperature and pressure are important and because there is a high Martínez-Soto et al.
drying drying rate and low drying temperature and oxygen-deficient drying (2001), Alibas (2007),
environment, the quality characteristics of the products (e.g., shape, color, Giri and Prasat (2007),
and aroma) are maintained and Artnaseaw et al.
• In fresh Lentinula mushroom, vacuum dehydration at 50°C–65°C and (2010)
0.1–0.4 bar decreased initial moisture from 92% to 13% (w.b.) in 6 h and
revealed that temperature and pressure significantly 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
operation cost
Freeze- • Pleurotus mushrooms were firstly washed, frozen at −80°C, and then placed Kompany and René
drying in the freeze-dryer viz. at 0°C (condenser temperature −55°C, vacuum at 7 (1995), Martínez-Soto
Pa) for about 24 h. Although flavor was not significantly different, et al. (2001), and Giri
mushrooms had superior quality as compared to hot air and vacuum-dried and Prasat (2007)
ones.
• A. bisporus the quantity of aromatic compound 1-octen 3-ol was
significantly reduced
operation cost
Fluidized • The quality of both Agaricus and Pleurotus mushrooms was good and the Gothandapani et al.
bed drying time short (1997), Walde et al.
• The quality of Pleurotus sp. dried in fluidized bed condition at 50°C for (2006), and Giri and
80–120 min with 0.5 potassium metabisulfite was superior and with reduced Prasat (2007)
microbial spoilage to sun, thin layer drying and fluidized bed drying with
blanching
operation cost
Microwave • Is rapid, more uniform, and energy-efficient compared to conventional hot Riva et al. (1991),
drying air drying, but although it combines the advantages of drying at low Walde et al. (2006),
temperatures, it has not very satisfactory results due to temperature Giri and Prasad (2007),
instability and tissue hardening and Lombraña et al.
• Temperature control and pressure will contribute to the continued (2010)
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
Microwave- • Resulted in 70%–90% decrease in the drying time to a moisture content of Giri and Prasad (2007,
vacuum 6% (db) and had better rehydration characteristics in sliced white 2013)
drying mushrooms than the hot air dried samples
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22.7.1.1 Food Products and Food Additives

Mushroom pickling, a procedure that involves fermentation, is an economical way for mushroom pres-
ervation and has a beneficial 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
10 days at 21°C with freshly shredded cabbage. In another process (Singh et al. 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 15 days 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 final salt concentration reached to 6.6% by weight. Products of this
type can be stored for 6 months at the ambient temperature (Singh and Bano 1977), the pasteurization of
mushrooms before storage being unnecessary. Fermented mushrooms can be also used as a semifinished
product in the preparation of marinades and sauces of good quality (Bernś et al. 2006). Pickling is also
an interesting option of saving loss material (about 10%) in the canning process of mushrooms (Rai and
Arumuganathan 2008).
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 et al. 2006). The method is
simple, economical, and mushrooms can be preserved for periods ranging from 3 to 6 months 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 filling in the sterilized bottles or jars (Rai and Arumuganathan 2008). For mushroom
chips production, mushrooms are sliced (2 mm), 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 finally fried in oil (Rai and
Arumuganathan 2008).
Mushroom powder is used as a direct food additive to increase content of dietary fibbers in vari-
ous foods or as a partial substitute for wheat flour in bakery products. It is obtained after pulveriza-
tion of dried mushroom slices and used to enhance flavor of a dish or to provide specific 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 flour, 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 finally 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).
22.7.1.2 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 flavor and aroma profiles 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 et al. 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 40 days 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 14 days (Okamura-Matsui et al. 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-d-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 et al. 1995). Thus, alcoholic beverages made using mushrooms, except of
characteristic flavor and aroma, seem to be a functional source which can be expected to have also health
benefits (Okamura-Matsui et al. 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 artificial 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 et al. (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
Ganoderma tsugae.
22.7.2 Dietary Supplements: Nutraceutical Products

Mushroom nutraceuticals are preparations (refined 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, “Sonifilan,” “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,
isoflavones, 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 et al. 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 et al. 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 et al. 2009). Due to their immuno-
modulatory properties, purified β-glucans have been used clinically as part of a combination therapy
for a variety of cancers (Thompson et al. 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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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 et al. 2005). Among mushrooms, L. edodes
(shiitake) has been studied most extensively (antitumor and immunostimulating properties revised by
Shen et al. (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 et al. 2005). Moreover, the Phellinus
linteus extract antitumor activity has been demonstrated by Collins et al. (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 et al. 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 findings indicate that P.
linteus has a synergistic effect with Dox to activate a beneficial 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 significant advantages in survival over chemotherapy
alone. Next to a direct influence 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 et al. 2013). However, this
fruit body waste, apart from the nutritional use as animal feed, is a good source of insoluble dietary fibers
and glucans that can be used for the preparation of biologically active polysaccharide complexes utiliz-
able as food supplements (Synytsya et al. 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 et al. 2009; Chou et al. 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 (1 kg of mushrooms usually generates 5 kg 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-filling). Actually, the mushroom industry faces challenges in storing
AQ4 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).
AQ5 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,
K21711_C022.indd 516 4/30/2015 12:32:20 AM

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 et al.
2009; Phan and Sabaratnam 2012; Philippoussis and Diamantopoulou 2012; Papadopoulou et al. 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 et al. 2006; Kim et al. 2010).
22.8 Conclusions
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 difficult 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 scientific and market research, development, and technical support.
Mushroom harvesting wastes and SMS need to be reclassified 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 beneficial 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-
room constituents, especially those with positive health effects.
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Cultivated Mushrooms: Preservation and Processing

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22.2 Mushroom Composition and Nutritional Aspects

22.3 Health Benefits of Mushrooms

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22.4 Cultivation and Harvesting Methods Affecting

22.4.1 Agaricus bisporus

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22.4.2 Pleurotus spp.

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22.4.3 Lentinula edodes

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inoculation exposure incubation/preharvest period and yielda,b,c

22.5 Postharvest Handling of Mushrooms

22.5.1 Mushroom Grading and Quality

22.5.2 Postharvest Physiology and Storage

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22.6 Processing Methods of Mushrooms

22.6.1 Processing for Short-Term Preservation

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22.6.1.2 Minimal Processing

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22.6.1.3 Effect of Packaging on Mushroom Shelf-Life

22.6.2 Processing for Long-Term Preservation

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22.7 Value–Added Mushroom Products and By-Products

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22.7.1.1 Food Products and Food Additives

22.7.1.2 Beverages and Beauty Products

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22.7.2 Dietary Supplements: Nutraceutical Products

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22.7.3 Mushroom Industry Spent By-Products

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22.4 Cultivation and Harvesting Methods Affecting