Food Chemistry 124 (2011) 1076–1082

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Food Chemistry
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Chemical composition and non-volatile components of Croatian

wild edible mushrooms
S. Beluhan a,*, A. Ranogajec b
a
Department of Biochemical Engineering, Faculty of Food Technology and Biotechnology, University of Zagreb, Pierottijeva 6, 10000 Zagreb, Croatia
b
Laboratory for Liquid Chromatography, Institute of Public Health Dr. Andrija Stampar, Mirogojska 16, 10000 Zagreb, Croatia

a r t i c l e i n f o a b s t r a c t

Article history: The chemical composition (dry matter, crude protein, crude fat, total carbohydrates, and ash) and non-
Received 6 April 2010 volatile components content (soluble sugars, free amino acids, and 50 -nucleotides) of 10 popular Croatian
Received in revised form 10 June 2010 wild edible mushroom species (Agaricus campestris, Boletus edulis, Calocybe gambosa, Cantharellus cibarius,
Accepted 26 July 2010
Craterellus cornucopioides, Entoloma clypeatum, Flammulina velutipes, Macroleptiota procera, Morchella
elata, and Pleurotus ostreatus) were determined. All investigated mushrooms were found to be good
sources of proteins and total carbohydrates, with contents varying in the ranges of 27.95–38.89, and
Keywords:
42.62–66.78 g/100 g, respectively. In addition, the fat contents were very low 1.34–6.45 g/100 g. B. edulis
Wild edible mushrooms
Chemical composition
(19.87 mg/g) showed the highest concentration of essential amino acids and M. elata (14.25 mg/g) the
Free amino acids lowest concentration. Monosodium glutamate-like components and total flavour 50 -nucleotides were
50 -Nucleotides the highest in C. cornucopioides (45.85 and 13.88 mg/g, respectively), and lowest in F. velutipes (7.63
Taste components and 1.05 mg/g, respectively). Equivalent umami concentration values in 10 Croatian wild edible mush-
Equivalent umami concentration rooms ranged from 73.78 to 1186.45 g MSG/100 g dry weight, and overall, all these mushrooms pos-
sessed highly umami taste.
Ó 2010 Elsevier Ltd. All rights reserved.

1. Introduction searched group of living organisms. It is assumed that in Croatia

Wild mushrooms are becoming more and more important in
our diet for their nutritional (Barros et al., 2007; Manzi, Gambelli,
Marconi, Vivanti, & Pizzoferrato, 1999), organoleptic (Maga, 1981)
and medicinal (Lee, Jian, & Mau, 2009) characteristics. Numerous
species of wild growing mushrooms are widely consumed as a del-
icacy in Central and Eastern Europe. In Croatia, mushroom picking
is a hobby growing in popularity in which all can enjoy. A large
part of people believe that the autumn is the best time for picking
mushrooms, but mushrooms are one of the few wild fruits that are
available throughout the year.
Nine wild edible mushroom species: field mushroom (Agaricus
campestris), porcini mushroom (Boletus edulis), golden chantarelle
(Chantarella cibarius), black trumpet (Craterellus cornucopioides),
winter mushroom (Flamumulina velutipes), hedgehok mushroom
(Hydnum repardum), red pine mushroom (Lactarius deliciosus),
black morel (Morchella elata), and oyster mushroom (Pleurotus
ostreatus) are very popular and the most harvested wild edible
mushrooms in Croatia.
During the development of the National Strategy and Action
Plan for the Conservation of biological and landscape diversity in
Croatia, obtained results showed that the fungi are the least re-
* Corresponding author. Tel.: +385 1 4604 058; fax: +385 1 4836 424.
E-mail address: sunbel@pbf.hr (S. Beluhan).
are about 20,000 living biological species, and to date it has found
and recorded (including lichens) only about 3800 mushroom spe-
cies (both edible and poisonous) which makes it a little more than
20% of the estimated number of biological species (Biodiversity of
Croatia, 2006).
Besides all the nutritional properties already available in the lit-
erature, there are no reports dealing about the chemical composi-
tion and nutritional value of Croatian wild edible mushrooms.
Number of studies have been carried out on the chemical compo-
sition and nutrition value of European edible mushrooms from dif-
ferent countries (Kalač, 2009), particularly on Finnish (Mattila,
Väänänen, Könkö, Aro, & Jalava, 2002), Polish (Jaworska & Bernaś,
2009), Spanish (Diéz & Alvarez, 2001), Portuguese (Barros, Cruz,
Baptista, Estevinho, & Ferreira, 2008a; Barros, Venturini, Baptista,
Estevihno, & Ferreira, 2008b; Barros et al., 2007; Heleno, Barros,
Sousa, Martins, & Ferreira, 2009), Italian (Manzi, Aguzzi, & Pizzofe-
rato, 2001; Manzi, Marconi, Aguzzi, & Pizzoferato, 2004), Macedo-
nian (Bauer-Petrovska, 2001), Greek (Ouzouni, Petridis, Koller, &
Riganakos, 2009; Ouzouni & Riganakos, 2007), and Turkish (Colak,
Faiz, & Sesli, 2009) species.
The fungal kingdom possesses certain natural advantages in
terms of dietary supremacy over the rest of the vegetarian platter.
According to Ghorai et al. (2009) these are: (a) a good protein con-
tent (20–30% of dry matter) having all the essential amino acids
(especially enriched in lysine) thus capable of substituting meat,

0308-8146/$ - see front matter Ó 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.foodchem.2010.07.081
 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082
(b) chitinous wall to act as a source of dietary fibre, (c) high vita-
min B content, (d) low in fat, and (e) virtually free of cholesterol.
They are easily cultivable and are consumed either in fresh or pro-
cessed form.
In general, the fruiting bodies of mushrooms, on dry weight
basis, contain about 57% carbohydrate, 25% protein, 5.7% fats, and
12.5% ash. Dry matter of mushrooms is very low, usually in the
range of 60–140 g/kg. Such high water content and water activity
(aw) affect the texture and participate in the short shelf life of fruit-
ing bodies (Kalač, 2009). Low dry matter and lipid contents result
in the low energy value of mushrooms. The values of 87–155 kJ/
100 g of fresh mushrooms was reported by Barros et al. (2007).
Thus, mushrooms are a food item of low energy value.
Flavour represents one of the most important quality attributes
contributing to the widespread consumption of both cultivated
and wild edible mushrooms. The typical flavour of mushrooms is
due to both volatile compounds (Maga, 1981) and non-volatile
compounds (Litchfield, 1967). The content of volatile compounds,
especially 1-octen-3-ol, decreased significantly during storage
time, cooking, and processing (Lee et al., 2009). The non-volatile
compounds, such as free amino acids, 50 -nucleotides, soluble sug-
ars, and polyols (Mau, Beelman, Ziegler, & Royse, 1991; Mau, Lin,
& Chen, 2001; Mau, Lin, Chen, Wu, & Peng, 1998; Tseng, Lee, &
Mau, 2005; Tseng & Mau, 1999; Yang, Lin, & Mau, 2001) are
responsible for the taste of mushrooms. However, until now, the
profile of non-volatile compounds of most Croatian wild edible
mushrooms was not available.
Umami taste, also called the palatable taste or the perception of
satisfaction, is a good taste commonly induced or enhanced by
monosodium glutamate (MSG) and 50 -nucleotides. Besides the four
basic tastes (sour, sweet, bitter, and salty), the umami taste is the
fifth taste in mouth perception (Yamaguchi, 1979).
Generally, mushrooms posses four functionalities, including
nutritional values, tasty properties, physiological effects, and cul-
tural characteristics (Mau, 2005). Mushroom has long been used
as food or food-flavouring material due to their unique and subtle
flavour including aroma and taste components, which add func-
tionality to foods. Mau (2005) calculated the equivalent umami
concentrations (EUC) of mushrooms, based on their contents of
non-volatile components. EUC is the concentration of MSG equiva-
lent to the umami intensity given by the mixture of MSG and the
50 -nucleotides. However, the profile of taste components of Cro-
atian wild edible mushrooms was not available.
Accordingly, this research was designed to examine the proxi-
mate composition and non-volatile taste components in 10 wild
species of popular Croatian edible mushrooms, particularly soluble
sugars and polyol, free amino acids including taste amino acids,
and 50 -nucleotides. Also, equivalent umami concentration (EUC)
values for studied mushrooms and the difference between species
were also evaluated and compared.
2. Materials and methods
2.1. Mushroom samples
Ten wild edible species (A. campestris, B. edulis, Calocybe gamb-
osa, Cantharelluscibarius, C. cornucopioides, E. clypeatum, F. velutipes,
Macroleptiota procera, M. elata, P. ostreatus), belonging to six differ-
ent families, were collected from forests in Croatian regions of
Istria (northwest) and Slavonia (northeast) from September 2008
to May 2009. The mushroom samples were cleaned from forest
debris (without washing) with a plastic knife, transported to the
laboratory within 12 h of collection and freeze-dried. Dried sam-
ples (50 g each) were ground to obtain fine powder. For all the
mushroom species three samples were analysed.
1077
2.2. Standards and reagents
Adenosine 50 -monophosphate sodium salt (AMP), cytidine 50 -
monophosphate disodium salt (CMP), guanosine 50 -monophos-
phate disodium salt (GMP), inosine 50 -monophosphate disodium
salt (IMP), uridine 50 -monophosphate disodium salt (UMP), and
xanthosine 50 -monophosphate (XMP) were purchased from Sigma
Chemical Co. (St. Louis, MO, USA). The amino acid standards
(aspartic acid, glutamic acid, serine, histidine, glycine, threonine,
alanine, arginine, cystine, valine, methionine, phenylalanine, iso-
leucine, leucine, lysine, tyrosine, and tryptophan) were obtained
from Fluka (Buchs, Switzerland). Boric acid, Na2HPO4, and derivat-
isation reagents (3-MPA, OPA, and FMOC-Cl) were from Sigma–Al-
drich (Steinheim, Germany). Potassium dihydrogen phosphate,
dipotassium hydrogen phosphate, HCl, and acetic acid were sup-
plied by Merck (Darmstadt, Germany). Orthophosphoric acid, for-
mic acid, ammonium acetate, sodium hydroxide, potassium
hydroxide, and NaOH were supplied by Kemika (Zagreb, Croatia).
Methanol and acetonitrile (HPLC gradient grade) were from J. T. Ba-
ker (Deventer, Holland).
2.3. Stock standard solutions
Stock standard solutions of individual 50 -nucleotides were pre-
pared by weighing of approximately 10 mg of each compound into
volumetric flask (10 ml). Standards were redissolved in deionised
water apart from GMP which needed a few drops of 10% KOH for
dissolution. Stock standards were stored at 4 °C in dark for up to
one month. Working standard solution mix of nucleotides was pre-
pared daily by dilution with water. For method optimisation on
column Luna Hilic working standard solution mix was prepared
in acetonitrile/water (9:1) (Ranogajec, Beluhan, & Šmit, 2010).
The stock solutions of individual amino acids were prepared in
0.1 M HCl except of tryptophan which was prepared in water. A
mix of all amino acids standards was prepared from stock solutions
of individual amino acids with a concentration of approximately
8 lg/ml of individual amino acid.
2.4. Equipment
The HPLC system Agilent 1100 Series consisted of a binary
pump G1312A, thermostated autosampler G1329A, vacuum degas-
ser G1379A, column owen G1330B, DAD detector G1315C, and FLD
detector G1321A. Column ZORBAX Eclipse-AAA (3.0  150 mm,
3.5 lm), thermostated at 40 °C was used for analysis. The mobile
phases were A: 40 mM Na2HPO4 pH 7.8, pH adjusted 7.8 with
NaOH solution (10 M) and B: ACN:MeOH:water (45:45:10), HPLC
gradient grade, at flow rate of 0.8 ml/min. Gradient was 0–57% B
for 1.9–18.1 min, 57–100% B for 18.1–18.6 min, 100% B for 18.6–
22.3 min, 100–0% B for 22.3–23.2 min. Run time was 26 min.
Detection DAD: 338 nm (for OPA-amino acids) and 262 nm (for
FMOC-amino acids) FLD was time programmed: 0–15.6 min ? ex
340, em 450, 15.6–26.0 min ? ex 266, em 350, PMT gain 10.
2.5. Chemical composition
The chemical composition of 10 wild edible Croatian mush-
rooms, including moisture, ash, total carbohydrates, crude fat,
and crude protein, were determined according to AOAC methods
(1995). To obtain moisture contents, samples of the mushrooms
were dried in an oven at 105 °C overnight until constant weight.
The ash content was determined by incineration at 550 °C for
24 h. The crude protein content of the samples was estimated by
the micro-Kjeldhal method, in which the sample was digested with
a known quantity of concentrated sulphuric acid in the Kjeltec
digestion apparatus (1007 Digestion Unit, Tecator, Sweden). For
1078 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082
the calculation of crude protein in mushrooms, the nitrogen con-
tent was multiplied by a factor of 4.38 (Crisan & Sands, 1978).
The crude fat content of the samples was determined by extracting
a known weight of powdered mushroom sample with petroleum
ether as a solvent, using a Soxhlet apparatus. The amount of total
sugars was calculated by difference: Total sugars = 100  (g mois-
ture  g protein  g crude fat  g ash). Total energy was calculated
according to the following equation: Energy (kJ) = 17  (g pro-
tein + g carbohydrate) + 37  (g lipid) (Manzi et al., 2004).
2.6. Soluble sugar assay
Soluble sugars were extracted and analysed as described by
Ajlouni, Beelman, Thompson, and Mau (1995). Freeze-dried pow-
der (300 mg) was extracted with 50 ml of 80 ml/l aqueous ethanol
and xylose (50 mg) was added as an internal standard. This sus-
pension was shaken for 45 min at ambient temperature and fil-
tered through Whatman No. 4 filter paper. The residue was
washed five times using 5 ml of aqueous ethanol each time. The
combined filtrate was rotary-evaporated and redissolved in deion-
ised water to a final volume of 10 ml. The aqueous extract was fil-
tered through 0.45 lm Acrodisc-CR filter prior to injection 10 ll of
the sample into the HPLC.
2.7. Free amino acid assay
Amino acids were derivatised on-line automatically with 3-
mercaptopropionic acid, 3-MPA/OPA. For derivatisation of amino
acids, the following reagents were used: 0.1 M borate buffer in
water with pH 10.2; 3-MPA/OPA reagent: 10 mg/ml of OPA was
dissolved in 0.02 M borate buffer with 1.0% of 3-MPA. Reagents
were stored at 4 °C. Prior derivatisation 20 ll of 3-MPA/OPA re-
agent was diluted with 140 ll of 0.1 M borate buffer. Derivatisa-
tion was performed using the automatic injector: successive
sampling of 2.5 ll of borate buffer and 0.5 ll of sample, then mixed
two times with a wait time of 0.5 min. Subsequently, 0.5 ll of 3-
MPA/OPA reagent was added. After mixing six times, 32 ll of water
was added, mixed two times and finally 18 ll of the mixture was
injected. Column Zorbax Eclipse-AAA (3.0  150 mm, 3.5 lm),
thermostated at 40 °C was used for analysis. The mobile phases
were A: 40 mM Na2HPO4 pH 7.8, pH adjusted 7.8 with NaOH solu-
tion (10 M) and B: ACN:MeOH:water (45:45:10), HPLC gradient
grade, at a flow rate of 0.8 ml/min, fluorescence detection (ex
340 nm, em 450 nm). Gradient was 0–57% B for 1.9–18.1 min,
57–100% B for 18.1–18.6 min, 100% B for 18.6–22.3 min, 100–0%
B for 22.3–23.2 min. Run time was 26 min.
2.8. Free 50 -nucleotide assay
50 -Nucleotides were extracted from homogenised sample
(500 mg) with 50 ml of deionised water. The suspension was
heated in a water bath (100 °C) for 1 min, vortex mixed for 30 s,
cooled to room temperature and filtered prior to HPLC injection
through a 0.2 lm nylon filter (Acrodisc, Pall Life Science). Determi-
nation was performed on column Synergy Hydro thermostated at
20 °C. The mobile phases were A: 50 mM phosphate buffer (pH
5.8) and B: methanol at a flow rate of 0.4 ml min1 and detection
at 254 nm. Gradient was 0–50% B for 3–12 min, run time was
13.5 min, post run time was 5 min. Multilevel calibration (n = 6)
by linear least-squares regression was established, with quantifica-
tion by the external standard technique. At the end of each sample
sequence, the column was rinsed with 300 column volumes of
water and stored in methanol:water (95:5). Peaks were identified
by comparison of retention time and similarity of chromatographic
peak spectrum with authentic standards, as estimated by a similar-
ity index of 0.95.
2.9. Equivalent umami concentration
The equivalent umami concentration (EUC, g monosodium glu-
tamate (MSG) per 100 g) is the concentration of MSG equivalent to
the umami intensity given by the mixture of MSG and the 50 -nucle-
otide and is represented by the following addition equation (Yam-
aguchi, Yoshikawa, Ikeda, & Ninomiya, 1971):
X X  X 
Y¼ ai bi þ 1218 ai bi aj bj ð1Þ
where Y is the EUC of the mixture in terms of g MSG/100 g; ai is the
concentration (mg/100 g) of each umami amino acid [aspartic acid
(Asp) or glutamic acid (Glu)]; aj is the concentration (g/100 g) of
each umami 50 -nucleotide [50 -inosine monophosphate (50 -IMP), 50 -
guanosine monophosphate (50 -GMP), 50 -xanthosine monophos-
phate (50 -XMP) or 50 -adenosine monophosphate (50 -AMP)]; bi is
the relative umami concentration (RUC) for each umami amino acid
to MSG (Glu, 1 and Asp, 0.077); bj is the RUC for each umami
50 -nucleotide to 50 -IMP (50 -IMP, 1; 50 -GMP, 2.3; 50 -XMP, 0.61 and
50 -AMP, 0.18); and 1.218 is a synergistic constant based on the con-
centration (g/100 g) used.
2.10. Statistical analysis
For each of the 10 mushroom species three samples were used
for the determination of every quality attribute, and also all of the
assays were carried out in triplicate. The results are expressed as
mean values and standard deviation (SD). The experimental data
were subjected to one-way analysis of variance (ANOVA OneWay)
for a completely random design to determine the least significant
difference amongst means at the level of 0.05. This test was carried
out using the OriginPro 8 program.
3. Results and discussion
We studied the chemical composition and non-volatile compo-
nents content of the 10 wild edible mushroom species, which are
most popular and regularly consumed by the Croats. The non-vol-
atile components content, especially flavour-enhancing 50 -nucleo-
tides and amino acid contents of most of these mushrooms were
not reported previously and none of the studied mushroom species
were commercially grown. Some of the mushroom species, such as
C. gambosa, E. clypeatum, and M. procera were collected by special-
ist collectors and were procured by theirs courtesy.
The results of the chemical composition and calculated energy
values (expressed on dry weight basis) for investigated mushroom
species are shown in Table 1. While examining the nutritional
composition of mushroom samples, the maturation stage of them
was not considered. When the nutrition value of mushrooms is
evaluated, the most important factor is their dry weight content.
It is known that the dry weight content of fresh mushrooms are
generally 5–15% and the nutritional profiles of mushrooms are di-
rectly affected with their moisture content (Bano & Rajarathnam,
1988; Crisan & Sands, 1978; Manzi et al., 1999; Mattila et al.,
2002). In addition, it is also known than moisture content of mush-
rooms depends on the harvesting time, maturation period and
environmental conditions such as humidity and temperature in
growing period (Crisan & Sands, 1978). The dry weight content of
all studied mushroom species ranged from 9.97% to 14.89%. In
the literature, dry weight content of A. campestris, B. edulis, C. ciba-
rius, M. procera, and P. ostreatus were 8.62%, 14.27%, 9.72%, 10.89%,
and 8.01%, respectively (Bauer-Petrovska, 2001). When these
results were compared with the values obtained in this study for
the same mushroom species (Table 1), it can be easily seen that
with exception of B. edulis (12.23%), the values from our study
for A. campestris (14.89%), C. cibarius (14.24%), M. procera
 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082 1079

Table 1
Proximate composition of Croatian wild edible mushroom species in a dry weight basis.

Mushroom species Dry matter (%) Proteins (g/100 g) Fat (g/100 g) Total sugars (g/100 g) Ash (g/100 g) Energy (kJ)
A
Agaricus campestris 14.89 ± 0.07b 38.89 ± 0.67a 2.7 ± 0.56c 47.56 ± 0.89a 3.5 ± 0.66c 1569.55 ± 0.24b
Boletus edulis 12.23 ± 0.41c 36.91 ± 0.02b 2.92 ± 0.41c 64.27 ± 0.21a 5.3 ± 0.87c 1488.10 ± 0.18c
Calocybe gambosa 13.89 ± 0.39b 36.65 ± 0.34a 1.34 ± 0.12c 42.65 ± 0.97a 7.98 ± 0.79b 1397.68 ± 0.29c
Cantharellus cibarius 14.24 ± 0.21c 30.91 ± 0.28b 1.9 ± 0.61c 52.5 ± 0.24a 8.8 ± 0.05c 1488.27 ± 0.28c
C. cornucopioides 10.06 ± 0.01b 47.21 ± 0.34a 4.87 ± 0.21c 44.54 ± 0.69a 10.08 ± 0.83b 1729.94 ± 0.14a
Entoloma clypeatum 13.99 ± 0.22c 27.98 ± 0.21b 6.21 ± 0.44c 50.75 ± 0.42a 9.21 ± 0.91c 1568.18 ± 0.19b
Flammulina velutipes 11.95 ± 0.02b 27.95 ± 0.34a 6.45 ± 0.43c 42.6 ± 0.10a 7.39 ± 0.56b 1438.00 ± 0.16c
Macroleptiota procera 13.23 ± 0.78c 24.22 ± 0.67b 2.23 ± 0.22c 66.78 ± 0.12a 5.37 ± 0.04c 1629.51 ± 0.14c
Morchella elata 9.97 ± 0.45b 35.75 ± 0.99a 3.91 ± 0.77c 43.98 ± 0.01a 9.01 ± 0.05b 1500.08 ± 0.44b
Pleurotus ostreatus 11.67 ± 0.23c 24.90 ± 0.89b 2.08 ± 0.06c 61.9 ± 0.13a 7.62 ± 0.23c 1552.26 ± 0.16b
A
Each value is expressed as mean ± SD (n = 3). Means with different letters within a row are significantly different (P < 0.05).

(13.23%), and P. ostreatus (11.67%) were higher. But, dry weight
content of each studied mushroom species was generally similar
to each other. Based on dry weight, contents of other estimated
components were in the order: total carbohydrates (42.6–66.78
g/100 g) > crude protein (24.22–47.21 g/100 g) > crude ash (3.5–
10.08 g/100 g) > crude fat (1.9–6.45 g/100 g). Obviously, mush-
rooms were high in carbohydrate and protein contents but low
in ash and fat. Although, it is difficult to compare all obtained
results with results reported in literature because of lack of infor-
mation about some of the species studied in this work, particularly
C. gambosa and E. clypeatum, some of the results are consistent
with the literature related to M. procera, A. campestris, B. edulis (Tsai,
Tsai, & Mau, 2008), P. ostreatus, C. cibarius (Ouzouni et al., 2009), C.
cornucopioides, F. velutipes (Yang et al.,2001), and M. elata (Tsai,
Weng, Huang, Chen, & Mau, 2006). The major compounds of mush-
rooms are proteins and sugars. It was reported that the protein
contents of mushrooms are affected by a number of factors, namely
the type of mushroom, the stage of development, the part sampled,
level of nitrogen available and the harvest location (Colak et al.,
2009). In this study, the highest protein content was found for C.
cornucopioides (47.21 g/100 g) which is in agreement with Colak
et al., 2009) who found 50.10 g/100 g proteins, but Barros et al.
(2008a) reported a higher protein content (69.45 g/100 g) for the
same mushroom species. The lowest protein content was found
for M. procera (24.22 g/100 g), and the obtained result was similar
to 23.9 g/100 g as reported Ouzouni and Riganakos (2007). The dis-
tribution of proteins within a fruiting body and changes in protein
content during the development of a fruiting body remain mostly
unclear (Kalač, 2009). Vetter and Rimóczi (1993) reported the
highest crude protein content in cultivated P. ostreatus at a cap
diameter of 5–8 cm. At this stage of the development, crude
protein was 36.4 g/100 g and 11.8 g/100 g in cap and stipe, respec-
tively. In our work, protein content in wild edible P. octreatus caps
diameter between 4 and 4.7 cm were approximately 24.90 g/100 g
dry weight. Lipid compounds such as free fatty acids, tri-, di- and
monoglycerides, phospholipids, sterols, and derivatives can be ex-
tracted from mushrooms as crude fat (Crisan & Sands, 1978).
Mushrooms are consumed for low-calorie diet because of their
low crude fat content. The fat contents widely ranged from
1.34 g/100 g (C. gambosa) to 6.45 g/100 g (F. velutipes) in dry mush-
rooms, and the proximate descending order were F. velutipes, C.
cornucopioides, M. elata, B. edulis, C. gambosa, respectively. Gener-
ally, carbohydrate contents of mushrooms fruit bodies were in
the range of 44.0–74.3 g/100 g (Crisan & Sands, 1978). The carbo-
hydrate contents of some wild edible mushrooms from nitrogen
free extracts were found to be between 41 and 65 g/100 g (Sanmee,
Dell, Lumyong, Izumori, & Lumyong, 2003). In this study, the total
carbohydrate content of M. procera was the highest (66.78 g/100 g)
and it was similar with that reported by Ouzouni and Riganakos
(2007), 68.4 g/100 g. The lowest carbohydrate content was assayed
in F. velutipes and C. gambosa (42.6 and 42.65 g/100 g, respectively).
Ash content of mushroom is usually between 5 and 12% of dry
weight (Barros et al., 2008b; Ouzouni & Riganakos, 2007). In our
work, ash contents varied among species, so C. cornucopioides, E.
clipeatum, M. elata, and C. cibarius (10.08, 9.21, 9.01, and 8.8 g/
100 g, respectively) were higher in ash content than C. gambosa,
P. ostreatus, and F. velutipes (7.98, 7.62, and 7.39 g/100 g, respec-
tively). The lowest ash contents were found in M. procera, B. edulis
and A. campestris (5.37, 5.3, and 3.5 g/100 g, respectively). On the
basis of proximate analysis, it can be calculated that the edible por-
tion of 100 g dry weight of these mushrooms provides, on average,
1537 kJ (363 kcal). The highest value (1731 kJ) is guaranteed by C.
cornucopioides, while C. gambosa, give the lowest (1398 kJ) energy
contribution.
The sugar compositions of the wild edible mushrooms studied
in this work are shown in Table 2. Mannitol and trehalose, which
were major mushroom polyol and sugar, respectively, according
to Bano and Rajarathnam (1988) and Mau, Chyau, Li, and Tseng
(1997), were found in all 10 mushrooms, as well as glucose,
whereas mannose was not detected in C. cornucopioides and E.

Table 2
Content of soluble sugars and polyol (g/100 g) of Croatian wild edible mushrooms.

Mushroom species Mannitol Trehalose Mannose Glucose Total

Agaricus campestris 16.34 ± 0.78bA 0.72 ± 0.07c 65.61 ± 0.01a 15.21 ± 0.34b 97.88 ± 0.45a
Boletus edulis 3.72 ± 0.21c 9.92 ± 0.04b 36.23 ± 0.12a 6.28 ± 0.15b 59.89 ± 0.13b
Calocybe gambosa 0.34 ± 0.09c 7.69 ± 0.24b 17.21 ± 0.35a 11.15 ± 0.02b 36.39 ± 0.18b
Cantharellus cibarius 8.56 ± 0.02b 6.68 ± 0.09b 28.56 ± 0.02a 7.98 ± 0.13b 48.78 ± 0.12b
C. cornucopioides 11.21 ± 0.12a 0.09 ± 0.05c ndB 2.78 ± 0.54b 14.08 ± 0.32c
Entoloma clypeatum 6.67 ± 0.07b 3.05 ± 0.42b nd 11.78 ± 0.12a 21.50 ± 0.37c
Flammulina velutipes 7.90 ± 0.15b 2.97 ± 0.67c 7.23 ± 0.72b 12.01 ± 0.01a 30.10 ± 0.59b
Macroleptiota procera 2.42 ± 0.18b 0.11 ± 0.02c 11.09 ± 0.25a 10.78 ± 0.02a 24.40 ± 0.09c
Morchella elata 5.43 ± 0.21b 1.09 ± 0.02c 43.07 ± 0.12a 9.54 ± 0.23b 59.13 ± 0.12b
Pleurotus ostreatus 9.82 ± 0.55b 1.79 ± 0.12c 11.13 ± 0.29a 15.01 ± 0.35a 37.75 ± 0.39b
A
Each value is expressed as mean ± SD (n = 3). Means with different letters within a row are significantly different (P < 0.05).
B
Not detected.
1080 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082

clypeatum. Contents of total soluble sugars and polyol ranged from
14.08 g/100 g (C. cornucopioides) to 97.88 g/100 g (A. campestris).
Also, the highest amount of mannitol, mannose and glucose was
found in A. campestris (16.34, 65.61, and 15.21 g/100 g, respec-
tively). Soluble sugars contained in the mushroom contributed to
the sweet taste (Litchfield, 1967). Therefore, the high content of
sugars and polyols would give rise to a moderate sweet taste
perception.
Several authors referred to mushrooms as a good source of
essential amino acids such as: leucine, valine, threonine, lysine,
methionine, and tryptophan. Threonine and lysine were the major
essential free amino acids found in all mushrooms. Wild mush-
room proteins also contain considerable amounts of non-essential
amino acids such as: alanine, arginine, glycine, glutamic acid,
aspartic acid, and serine (Heleno et al., 2009). The contents of free
essential and non-essential amino acids are present in Table 3. Sev-
enteen free amino acids were detected at almost all studied mush-
room species and we found that threonine and lysine represented
the largest amount in C. cibarius (8.98 and 5.74 mg/g dry weight,
respectively). The total amino acid contents ranged from
54.20 mg/g (F. velutipes) to 72.04 mg/g (B. edulis). Several amino
acids were not detected, particularly, isoleucine an essential amino
acid in most of mushrooms, as well as tyrosine and aspartic acid as
non-essential amino acids. Contents of essential amino acids varied
among these 10 species and were in the same order as the total
amino acid contents for these mushrooms.
According to the classification described by Mau et al. (2001)
and Yang et al. (2001), amino acids in edible mushrooms were di-
vided into four groups on the basis of their taste characteristics
(Table 4). Group one was monosodium glutamate-like (MSG-like)
or palatable taste amino acids, including aspartic and glutamic
acids. Group two presented sweet taste amino acids, including ala-
nine, glycine, serine and threonine. The third class belonged to bit-
ter amino acids, including arginine, histidine, isoleucine, leucine,
methionine, phenylalanine, and valine. Last of all, lysine and tyro-
sine were contributed to tasteless mushrooms. Therefore, MSG-like
and sweet components could be responsible for the pleasant taste
of mushrooms. However, it is a vital that contents of MSG-like,

Table 3
Content of free amino acids of Croatian wild edible mushrooms.

Amino Mushroom species/Content (mg/g dry weight)

acidA
A. campestris B. edulis C. gambosa C. cibarius C. E. clypeatum F. velutipes M. procera M. elata P. ostreatus
cornucopioides
Ala 8.19 ± 0.27b 8.68 ± 0.04b 5.65 ± 0.05b 4.98 ± 0.25b 2.34 ± 0.16b 2.99 ± 0.13b 1.95 ± 0.29b 2.21 ± 0.04b 2.38 ± 0.11b 6.45 ± 0.13b
Arg 0.92 ± 0.39c 0.59 ± 0.17c 0.68 ± 0.18c 0.44 ± 0.14c 0.46 ± 0.09c 0.23 ± 0.34c 0.49 ± 0.11c 0.29 ± 0.33c 0.18 ± 0.21c 0.11 ± 0.12d
Asp 0.21 ± 0.04d 0.33 ± 0.18c 0.19 ± 0.02d 0.06 ± 0.17d ndC nd 2.59 ± 0.06b 0.12 ± 0.02d nd 0.17 ± 0.05c
Cys 1.74 ± 0.11b 2.17 ± 0.21b 1.56 ± 0.05b 1.99 ± 0.16b 1.87 ± 0.01b 1.21 ± 0.45b 1.39 ± 0.43b 1.45 ± 0.04b 1.37 ± 0.25b 1.81 ± 0.12b
Glu 34.78 ± 0.32a 39.09 ± 0.02a 25.67 ± 0.01a 29.99 ± 0.13a 45.85 ± 0.24a 23.89 ± 0.43a 29.98 ± 0.01a 33.65 ± 0.19a 38.29 ± 0.08a 41.09 ± 0.13a
Gly 0.12 ± 0.12d 0.27 ± 0.16c 0.22 ± 0.11d 0.13 ± 0.07c 0.17 ± 0.04d 0.24 ± 0.12c 0.15 ± 0.19d 0.09 ± 0.17d 0.31 ± 0.21c 0.34 ± 0.55c
HisB 2.02 ± 0.05c 2.34 ± 0.17c 3.91 ± 0.25a 3.15 ± 0.19b 3.61 ± 0.29a 2.98 ± 0.11b 2.54 ± 0.34b 3.29 ± 0.15a 3.61 ± 0.07a 4.42 ± 0.08a
IleB 0.41 ± 0.02d 0.12 ± 0.16e nd nd nd nd 0.44 ± 0.00c 0.19 ± 0.02d nd nd
LeuB 0.24 ± 0.09d 0.47 ± 0.27d 0.45 ± 0.12c 0.21 ± 0.04c 1.05 ± 0.07c 0.11 ± 0.34d 0.73 ± 0.01c 0.38 ± 0.22c 0.85 ± 0.06c 0.29 ± 0.02c
LysB 5.27 ± 0.12b 5.46 ± 0.13b 4.19 ± 0.11a 5.74 ± 0.27a 4.69 ± 0.02a 3.45 ± 0.49b 5.68 ± 0.11a 4.11 ± 0.17a 2.97 ± 0.01b 4.65 ± 0.65a
MetB 0.71 ± 0.42c 0.74 ± 0.01d 1.01 ± 0.21b 0.41 ± 0.03c 0.16 ± 0.14d 0.56 ± 0.45c 0.03 ± 0.01d 0.69 ± 0.11c 0.68 ± 0.02c 0.12 ± 0.03d
PheB 0.28 ± 0.09d 0.19 ± 0.89e 0.08 ± 0.11d 0.06 ± 0.08d 0.85 ± 0.01c 0.77 ± 0.02c 1.23 ± 0.06b 0.45 ± 0.30c 0.12 ± 0.00d 0.09 ± 0.17d
Ser 0.69 ± 0.19c 1.01 ± 0.89b 0.09 ± 0.11d 0.18 ± 0.08c 0.01 ± 0.01d 0.03 ± 0.32d 0.21 ± 0.06c 0.11 ± 0.30d 0.09 ± 0.10d 0.03 ± 0.27d
ThrB 7.89 ± 0.01a 9.14 ± 0.21a 5.71 ± 0.05a 8.98 ± 0.16a 4.56 ± 0.01a 6.23 ± 0.10a 5.21 ± 0.43a 5.78 ± 0.04a 4.21 ± 0.25a 6.99 ± 0.12a
Trp 0.08 ± 0.15d 0.03 ± 0.17d 0.09 ± 0.25d 0.02 ± 0.19d 0.12 ± 0.29d 0.03 ± 0.01d nd 0.09 ± 0.15d 0.05 ± 0.07d 0.01 ± 0.08d
Tyr nd nd 0.01 ± 0.42d nd 0.02 ± 0.03d 0.11 ± 0.02d 0.04 ± 011d 0.02 ± 0.21d nd nd
ValB 1.19 ± 0.07c 1.41 ± 0.01c 1.27 ± 0.03b 1.34 ± 0.36b 1.72 ± 0.07b 1.09 ± 0.29c 1.54 ± 0.23b 1.39 ± 0.11b 1.81 ± 0.13b 1.21 ± 0.43b
Total 64.74 ± 0.33b 72.04 ± 0.13a 50.78 ± 0.21c 57.68 ± 0.11c 67.48 ± 0.25b 43.92 ± 0.17d 54.20 ± 0.01c 54.31 ± 0.07c 56.92 ± 0.05c 67.78 ± 0.09b
A
Ala, L-Alanine; Arg, L-Arginine; Asp, L-Aspartic acid; Cys, L-Cystine; Glu, L-Glutamic acid; Gly, Glycine; His, L-Histidine; Ile, L-Isoleucine; Leu, L-Leucine; Lys, L-Lysine; Met,
L-Methionine; Phe, L-Phenylalanine; Ser, L-Serine; Thr, L-Threonine; Trp, L-Tryptophan; Tyr, L-Tyrosine; Val, L-Valine.
B
Essential amino acids.
C
Not detected.

Table 4
Taste characteristics of Croatian wild edible mushrooms.

Mushroom species Content (mg/g dry weight)

MSG-likeB SweetC BitterD TastelessE Total
A
Agaricus campestris 34.99 ± 0.36a 16.89 ± 0.09b 5.77 ± 0.03c 5.27 ± 0.12c 62.92 ± 0.20a
Boletus edulis 39.42 ± 0.20a 19.10 ± 0.12b 5.86 ± 0.29c 5.46 ± 0.13c 69.84 ± 0.22a
Calocybe gambosa 25.86 ± 0.03a 11.67 ± 0.27b 7.41 ± 0.11c 4.20 ± 0.53d 49.14 ± 0.30b
Cantharellus cibarius 30.05 ± 0.30a 14.27 ± 0.01b 10.15 ± 0.13c 5.74 ± 0.27d 60.21 ± 0.22a
C. cornucopioides 45.85 ± 0.24a 7.08 ± 0.18b 7.85 ± 0.08b 4.71 ± 0.05c 65.49 ± 0.17a
Entoloma clypeatum 23.89 ± 0.43a 9.49 ± 0.51b 5.74 ± 0.06c 3.56 ± 0.51c 42.68 ± 0.46b
Flammulina velutipes 7.63 ± 0.21a 7.52 ± 0.27b 7.00 ± 0.55a 5.72 ± 0.22b 27.87 ± 0.38c
Macroleptiota procera 33.77 ± 0.21a 8.19 ± 0.81b 6.68 ± 0.32b 4.13 ± 0.19c 52.77 ± 0.48b
Morchella elata 38.29 ± 0.08a 7.00 ± 0.06b 7.25 ± 0.19b 2.97 ± 0.07c 55.51 ± 0.12a
Pleurotus ostreatus 41.26 ± 0.18a 13.81 ± 0.03b 6.24 ± 0.14c 4.65 ± 0.08c 65.96 ± 0.13a
A
Means with different letters within a row are significantly different (P < 0.05).
B
Monosodium glutamate-like; Asp + Glu.
C
Sweet: Ala + Gly + Ser + Thr.
D
Bitter: Arg + His + Ile + Leu + Met + Phe + Val.
E
Tasteless: Lys + Tyr.
 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082 1081

sweet components, and total soluble sugars were considerably
high in edible mushrooms which made them distinguished from
medicinal mushrooms. On the basis of recommended classifica-
tion, investigated MSG-like amino acids in these 10 mushrooms
ranged from 7.63 mg/g (F. velutipes) to 45.85 mg/g (C. cornucopio-
ides). Contents of MSG-like components were found to be 22.67–
47.12 mg/g dry weight in common mushrooms (A. bisporus) (Tseng
& Mau, 1999) and 1.01–1.77 mg/g in king oyster mushrooms (P.
eryngii) (Mau et al., 1998). In addition, Yang et al. (2001) found that
contents of MSG-like components in several commercial mush-
rooms, including shiitake, winter (F. velutipes), and oyster mush-
rooms (P. ostreatus), ranged from 0.84 to 1.93 mg/g. In this work,
the sweet contents were moderate and ranged from 7.08 mg/g
(C. cornucopioides) to 19.10 mg/g (B. edulis). Due to their low solu-
ble sugar contents, the sweet taste of these mushrooms mainly de-
pends on the sweet amino acid contents. The bitter components
were predominantly present in all these 10 wild edible mush-
rooms, in which C. cibarius contained the highest amount
(10.15 mg/g). Chen (1986) conducted a series of sensory evalua-
tions on synthetic mushroom extracts prepared by omitting and
adding soluble components and found that alanine, glycine, and
threonine (sweet), and aspartic and glutamic acids (MSG-like)
were taste-active amino acids in A. bisporus, whereas none of the
bitter components were found to be taste-active in the overall
taste perception. The bitterness from the bitter components in
mushrooms could probably be masked by the sweetness from
the sweet components and additionally the high amount of manni-
tol. Therefore, MSG-like and sweet components would be respon-
sible for the delightful taste of mushrooms (Mau et al., 2001;
Tsai et al., 2009; Tseng & Mau, 1999).
Flavour 50 -nucleotides were found to be 50 -guanosine mono-
phosphate (50 -GMP), 50 -inosine monophosphate (50 -IMP) and 50 -
xanthosine monophosphate (50 -XMP) (Chen, 1986). 50 -GMP gave
the meaty flavour, and is a flavour enhancer much stronger than
MSG (Litchfield, 1967). Contents of flavour 50 -nucleotides were
found to be 4.19–6.30 mg/g dry weight in common mushrooms
(Tseng & Mau, 1999), 4.42–9.00 mg/g in paddy straw mushrooms
(Mau et al., 1997), and 1.63–4.89 mg/g in king oyster mushrooms
(Mau et al. 1998). Yang et al. (2001) reported that contents of fla-
vour 50 -nucleotides could be divided into three ranges: low
(<1 mg/g), middle (1–5 mg/g) and high ranges (>5 mg/g). Between
10 Croatian wild edible mushrooms, the ranges of total and flavour
50 -nucleotides, presented in Table 5, were highest in C. cornucopio-
ides, 35.36 and 13.88 mg/g, respectively. Also, high contents of fla-
vour 50 -nucleotides were found in M. elata (5.32 mg/g). P. ostreatus
have middle flavour 50 -nucleotides range of 3.43 mg/g. Other seven
wild edible mushrooms were comparably low in flavour 50 -nucle-
otides content, ranging from 1.01 to 1.63 mg/g.
The synergistic effects of flavour 50 -nucleotides with MSG-like
components might greatly increase the umami taste of mushrooms
(Yamaguchi, et al., 1971). Mau (2005) grouped EUC values from
mushroom taste components using the equation of Yamaguchi
et al. (1971) into four levels: first level of >1000 g/100 g dry weight
(>10 g MSG/g dry matter), second level of 100–1000 g/100 g (1–
10 g MSG/100 g), third level of 10–100 g/100 g (0.1–1 g MSG/g),
and fourth level of <10 g/100 g (<0.1 g MSG/g). So, the EUC value
of 100% represents that the umami intensity per 1 g dry weight
is equivalent to the umami intensity given by 1 g of MSG or, in
other words, 1 g MSG/g dry weight. With regard to EUC, values
of mushrooms investigated in our work were in the range of
73.78 g MSG/100 g in F. velutipes to 1186.45 g MSG/100 g dry
weight B. edulis (Table 6). The highest umami intensities of 1 g of
C. gambosa, A. campestris and B. edulis were equivalent to the uma-
mi intensities given by 6.4, 9.91, and 11.86 g of MSG, respectively.
Furthermore, C. cibarius, E. clypeatum and M. procera showed satis-
factory umami intensities by 2.48, 1.98, and 3.18 g of MSG, respec-
tively. Surprisingly, the EUC values of F. velutipes, C. cornucopioides,
and especially P. ostreatus and M. elata were lower than that is ex-
pected, 0.73, 1.21, 1.5, and 0.79 g of MSG, respectively. However,
Table 6
EUC of Croatian wild edible mushrooms.
EUCA (g MSG/100 g dry weight)
Agaricus campestris 991.61 ± 52.4aB
Boletus edulis 1186.45 ± 87.9a
Calocybe gambosa 640.63 ± 49.3b
Cantharellus cibarius 248.58 ± 41.1c
C. cornucopioides 120.91 ± 14.9d
Entoloma clypeatum 198.22 ± 29.9c
Flammulina velutipes 73.78 ± 9.1d
Macroleptiota procera 318.14 ± 57.6c
Morchella elata 79.52 ± 12.4d
Pleurotus ostreatus 150.55 ± 21.9d
A P P P 
Calculated based on the equation: Y ¼ ai bi þ 1:218 ð ai bi Þ aj bj (Yam-
aguchi et al., 1971), where Y is the EUC of the mixture in terms of g MSG/100 g; ai is
the concentration (mg/100 g) of each umami amino acid [aspartic acid (Asp) or
glutamic acid (Glu)]; aj is the concentration (g/100 g) of each umami 50 -nucleotide
(50 -IMP, 50 -GMP, 50 -XMP or 50 -AMP); bi is the relative umami concentration (RUC)
for each umami amino acid to MSG (Glu, 1 and Asp, 0.077); bj is the RUC for each
umami 50 -nucleotide to 50 -IMP (50 -IMP, 1; 50 -GMP, 2.3; 50 -XMP, 0.61 and 50 -AMP,
0.18); and 1.218 is a synergistic constant based on the concentration (g/100 g) used.
B
Each value is expressed as mean ± SD (n = 3). Means with different letters
within a column are significantly different (P < 0.05).

Table 5
Content of free 50 -nucleotides of Croatian wild edible mushrooms.

Mushroom species ContentA (mg/g dry weight)

50 -AMP 50 -CMP 50 -GMP 50 -IMP 50 -UMP 50 -XMP FlavourC Total
D
Agaricus campestris 0.73 ± 0.17b 0.78 ± 0.43b 0.61 ± 0.03c 0.12 ± 0.22d 1.11 ± 0.08a 0.56 ± 0.06c 1.29 ± 0.10a 3.91 ± 0.15c
Boletus edulis 1.65 ± 0.14a 0.16 ± 0.45c 0.64 ± 0.13b 0.28 ± 0.08c 1.21 ± 0.34a 0.71 ± 0.09b 1.63 ± 0.03a 4.65 ± 0.15c
Calocybe gambosa 2.21 ± 0.23b 0.28 ± 0.03d 0.60 ± 0.43c 0.03 ± 0.10d 3.57 ± 0.09a 0.38 ± 0.04c 1.01 ± 0.21b 7.07 ± 0.14b
Cantharellus cibarius 0.41 ± 0.21b 0.09 ± 0.02d 0.21 ± 0.07c 0.03 ± 0.12d 0.75 ± 0.23a 0.14 ± 0.07d 0.38 ± 0.03b 2.01 ± 0.08c
C. cornucopioides 0.35 ± 0.11c 5.13 ± 0.08b 2.88 ± 0.06c 3.97 ± 0.02b 2.12 ± 0.09d 7.03 ± 0.05a 13.88 ± 0.02a 35.36 ± 0.03a
Entoloma clypeatum 2.23 ± 0.34a 0.78 ± 0.12b 0.67 ± 0.17b ndB 2.71 ± 0.14a 0.57 ± 0.28c 1.24 ± 0.10a 6.95 ± 0.11b
Flammulina velutipes 1.48 ± 0.07b 4.01 ± 0.28a 0.45 ± 0.05c 0.28 ± 0.01d 1.42 ± 0.25b 0.32 ± 0.09d 1.05 ± 0.04b 7.96 ± 0.11b
Macroleptiota procera 1.03 ± 0.01b 1.39 ± 0.15a 0.12 ± 0.03c 0.16 ± 0.19c 1.61 ± 0.08a 0.24 ± 0.13c 0.52 ± 0.08c 4.55 ± 0.07c
Morchella elata 6.57 ± 0.45a 4.28 ± 0.33b 1.19 ± 0.01d 1.77 ± 0.04c 4.19 ± 0.09b 2.36 ± 0.03c 5.32 ± 0.01b 20.36 ± 0.18a
Pleurotus ostreatus 1.21 ± 0.03b 0.81 ± 0.29c 0.59 ± 0.03c 0.21 ± 0.09d 0.59 ± 0.06c 2.63 ± 0.21a 3.43 ± 0.09a 6.04 ± 0.11b
A
50 -AMP, 50 -adenosine monophosphate; 50 -CMP, 50 -cytosine monophosphate; 50 -GMP, 50 -guanosine monophosphate; 50 -IMP, 50 -inosine monophosphate; 50 -UMP, 50 -
uridine monophosphate; 50 -XMP, 50 -xanthosine monophosphate.
B
Non detected.
C
Flavour 50 -nucleotides: 50 -IMP + 50 -GMP + 50 -XMP.
D
Means with different letters within a row are significantly different (P < 0.05).
1082 S. Beluhan, A. Ranogajec / Food Chemistry 124 (2011) 1076–1082
sensory EUC values were not conducted in this work, but calcu-
lated EUC values of 10 wild edible mushrooms might be beneficial
for these species to be used as food-flavouring components or in
the formulation of nutraceutical and functional foods with a palat-
able umami taste.
In conclusion, the studied species of mushrooms picked and
consumed in Croatia have always been harvested wild, with the
exception of P. ostreatus, and no effort has been made to cultivate
these species on a commercial scale. Despite growing urbanisation
and changing food habits that accompany it, the tradition of pick-
ing and consuming wild edible mushrooms remains rooted in the
culture of the Croats. The high nutritional quality and unique fla-
vour and aroma of these mushrooms are likely to be poorly known,
so therefore it is imperative that the nutritional database of these
mushrooms is set up to retain and improve the information of
these unique species. Based on the results obtained, 10 mushroom
species possessed highly intense umami taste in addition to poten-
tial pharmacology properties for some of them. However, further
studies are also required on the content of possible anti-nutritional
or even toxic factors in Croatian wild edible mushrooms to estab-
lish their overall nutritional supremacy.
Acknowledgements
The authors are grateful for financial support from the Ministry
of Science and Technology of the Republic of Croatia, by a Grant
No. 058-190-2004. Also, the authors thank to Mr. Josip Marino Si-
mic from Mushroom Gatherer Association ‘‘Buletus” for generously
providing the mushrooms.
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