Food Chemistry 113 (2009) 578584

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Food Chemistry
journal homepage: www.elsevier.com/locate/foodchem

Flavour components and antioxidant properties of several cultivated mushrooms

Shu-Yao Tsai a, Shih-Jeng Huang b, Sheng-Hua Lo c, Tsai-Ping Wu c, Pei-Ying Lian c, Jeng-Leun Mau c,*
a
b
c

Department of Health and Nutrition Biotechnology, Asia University, 500 Liufeng Road, Wufeng, Taichung 41354, Taiwan, ROC
Department of Nutrition and Health Science, Chungchou Institute of Technology, Yuanlin, Changhua 51003, Taiwan, ROC
Department of Food Science and Biotechnology, National Chung-Hsing University, 250 Kuokuang Road, Taichung 40227, Taiwan, ROC

a r t i c l e

i n f o

Article history:
Received 7 March 2008
Received in revised form 2 June 2008
Accepted 12 August 2008

Keywords:
Mushrooms
Volatile components
Taste components
Antioxidant properties

a b s t r a c t
Three mushrooms, Clitocybe maxima, Pleurotus ferulae and Pleurotus ostreatus grey strain were used to
study their avour components and antioxidant properties. The volatile avour components found comprised of six eight-membered carbon compounds and two aromatic compounds. The content total of soluble sugars and polyols was 125270 mg/g. The content of monosodium glutamate-like components was
1.768.89 mg/g. The contents of avour 50 -nucleotides ranged from 1.89 to 7.59 mg/g. Based on the
results obtained, three mushrooms possessed highly intense umami taste. Ethanolic extracts were more
effective in the inhibition of conjugated diene and scavenging ability on 1,1-diphenyl-2-picrylhydrazyl
radicals, whereas hot water extracts were more effective in the scavenging ability of hydroxyl radicals.
EC50 values were less than 14 and 30 mg/ml for ethanolic and hot water extracts, respectively, indicating
that the three mushrooms were relatively effective as they exhibited antioxidant properties, despite having scavenging abilities for hydroxyl radicals. Phenols were the major antioxidant components and the
total contents were 5.1011.1 mg gallic acid equivalents/g.
2008 Elsevier Ltd. All rights reserved.

1. Introduction
Edible mushrooms are commonly used as foods and food-avouring substances and also traditional Chinese medicines. Some
edible mushrooms have also been reported as therapeutic foods
which are useful in preventing diseases such as hypertension,
hypercholesterolaemia and cancer. These functional characteristics
are mainly due to chemical composition (Manzi, Aguzzi, & Pizzoferrato, 2001). Mushrooms have become attractive as functional
foods and a source of physiologically benecial substances.
Recently, fresh fruit bodies of mushrooms from three species
have become available in the market, including giant clitocybe (Clitocybe maxima Gaertn. et G. Mey.: Fr., big cup mushroom), ferulae
mushroom [Pleurotus ferulae (DC.: Fr) Quel.] and Japanese oyster
mushroom [Pleurotus ostreatus (Jacquin: Fries) Kummer (grey
strain), grey tree oyster mushroom]. However, the volatile components, non-volatile taste components and antioxidant properties of
C. maxima, P. ferulae and P. ostreatus are unknown. Therefore, our
objective was to examine the volatile and taste components
including proximate compositions, soluble sugars, free amino acids
and 50 -nucleotides in these three mushrooms. The antioxidant
properties of the ethanolic and hot water extracts assayed included
inhibition of conjugated diene, reducing capability, scavenging
abilities of radicals and chelating abilities to metal ions. The con* Corresponding author. Tel.: +886 4 2285 4313; fax: +886 4 2287 6211.
E-mail address: jlmau@dragon.nchu.edu.tw (J.-L. Mau).
0308-8146/$ - see front matter 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.foodchem.2008.08.034

tents of potential antioxidant components in these extracts were

also determined.
2. Materials and methods
2.1. Mushrooms
Three species of fresh mushrooms (C. maxima, P. ferulae and P.
ostreatus) were obtained from Tatsuen Farm, Pushin, Chunghua
County, Taiwan. Fruit bodies from each species were randomly divided into six trays without cover, each containing about 200 g.
Immediately after sampling, three trays of fresh samples from each
species were used for the analysis of volatile components. The
other three trays of C. maxima were divided into three trays of caps
and three trays of stipes based on its separate and distinct morphological appearance. All trays of mushrooms were then freeze dried
and ground using a mill (Retsch ultracentrifugal mill and sieving
machine Haan, Germany) to obtain a coarse powder (24 opening/
cm).
2.2. Extraction of volatile compounds
A tray of mushrooms (200 g, about 100 g for C. maxima caps or
stipes) was cut into small cubes and blended with 600 ml of 0.1 M
sodium phosphate buffer (pH 6.5, Wako Pure Chemical Co., Osaka,
Japan) containing 0.15% Tween 80 (Wako) and 1 ml of methanol
containing 1000 lg of 1-nonanol (Sigma Chemical Co., St. Louis,

S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

MO, USA) used as an internal standard. After 1 min of blending, the

homogenate was mixed with 50 ml n-hexane (Mallinckrodt Baker,
Phillipsburg, NJ, USA), and centrifuged at 10,000g for 10 min at
4 C. The n-hexane layer was preconcentrated with a distillation
apparatus at 40 C and carefully reconcentrated to approximately
50 ll using a 0.2 mm i.d.  10 cm Vigreux column (Tung Kawn
Glass Co., Hsinchu, Taiwan) at 40 C.
2.3. Determination of volatile compounds using GCMS
A HP 5890A Series II GC coupled to an HP 5972A MSD mass
spectrometer was used. A fused silica column (0.32 mm  30 m,
J&W, Folsom, CA, USA) coated with DB-Wax (1.0 lm thickness)
was used. The oven temperature was programed from 50 to
200 C at 2 C/min. The operating conditions were as follows: injector temperature, 250 C; GCMS interface temperature, 265 C; helium carrier ow rate, 1.0 ml/min; electron multiplier voltage,
1500 V; electron ionisation energy, 70 eV. Volatile components
were identied by comparing the mass spectral data with those
spectra available from the Wiley MS Chemstation Libraries (6th
edition, G1034, Rev. C.00.00, HewlettPackard) and GC retention
times of the components with those of authentic compounds,
which were commercially available (Aldrich Chemicals Co., Milwaukee, WI, USA; Penta Manufacturing Co., Livingston, NJ, USA).
Kovats retention indices of the volatile components were calculated with n-parafn (C8C25, Alltech Associates, Deereld, IL,
USA) as references. The amount of each component was determined using an internal standard method and calculated from
the peak area of gas chromatogram.
2.4. Soluble sugar, polyol, free amino acid and 50 -nucleotide assays
Soluble sugars and polyols were extracted and analysed as described by Ajlouni, Beelman, Thompson, and Mau (1995). Dried
powder (600 mg) was extracted with 50 ml of 80% aqueous ethanol
(95% pure, Taiwan Tobacco & Wine Monopoly Bureau, Taipei). This
suspension was shaken for 45 min at room temperature and ltered through Whatman No. 4 lter paper. The residue was washed
ve times with additional 25 ml portions of 80% ethanol. The combined ltrate was then concentrated under reduced pressure at
40 C and redissolved in deionised water to a nal volume of
10 ml. The aqueous extract was then passed through a lter unit
(13 mm, Lida, Corp., Kenosha, WI), and ltered using a 0.45 lm
CA non-sterile lter (Lida) prior to HPLC analysis. The HPLC system
consisted of a Hitachi L-6000 pump, a Rheodyne 7161 injector, a
20 ll sample loop, a Hitachi D-2500 chromato-integrator, a Shimadzu RID-10A detector and a Phase Sep-NH2 column (4.6  250 mm,
5 lm, Phase Separation Inc., Norwalk, CT). The mobile phase was
acetonitrile (LC grade, Tedia Co., Faireld, OH)/deionised water
85:15 (v/v) at a ow rate of 1 ml/min.
Free amino acids were extracted and analysed as described by
Mau, Chyau, Li, and Tseng (1997). Dried powder (500 mg) was
shaken with 50 ml of 0.1 N HCl (Union Chemical Co., Hsinchu,
Taiwan) for 45 min at ambient temperature and ltered through
Whatman No. 4 lter paper. The ltrate was then passed through
a lter unit (13 mm, Lida), and ltered using a 0.45 lm CA nonsterile lter (Lida). This ltrate was mixed with o-phthalaldehyde
reagent (Sigma) in an eppendorf tube, shaken to facilitate derivatisation and then immediately injected onto the HPLC. The HPLC
system was the same as for sugar analysis but included a Hitachi
L-7485 uorescence detector with uorescence excitation at
340 nm and emission at 450 nm and a LiChrospher 100 RP-18 column (4.6  250 mm, 5 lm, Phenomenex Inc., Torrance, CA). The
mobile phases were A, 50 mM sodium acetate (pH 5.7) containing
0.5% tetrahydrofuran; B, deionised water; and C, methanol. The
gradient was A:B:C 80:0:2033:0:67 for 038 min, 0:33:67 for

579

3840 min, and 0:100:0 for 4043 min. The ow rate was
1.2 ml/min.
50 -Nucleotides were extracted and analysed as described by
Taylor, Hershey, Levine, Coy, and Olivelle (1981). Dried powder
(500 mg) was extracted with 25 ml of deionised water. This suspension was heated to boiling for 1 min, cooled and then centrifuged at 22,200g for 15 min. The extraction was repeated once
with 20 ml of deionised water. The combined ltrate was then
evaporated, and ltered prior to HPLC injection in the same manner as in the soluble sugar and polyol assays. The HPLC system
was the same as for sugar and polyol assay except for a Shimadzu
UV detector and a LiChrospher 100 RP-18 column (4.6  250 mm,
5 lm, Merck). The mobile phase was 0.5 M KH2PO4/H3PO4 (pH
4.3, Wako Pure Chemical Co., Osaka, Japan) at a ow rate of 1 ml/
min and UV detection at 254 nm. Each sugar or polyol, amino acid
and 50 -nucleotide was identied using an authentic sample (Sigma) and quantied by its respective calibration curve.
2.5. Extraction of antioxidant components
For ethanolic extractions, a subsample (10 g) was extracted by
stirring with 100 ml of 95% ethanol at 25 C at 20g for 24 h and
ltration through Whatman No. 1 lter paper. The residue was
then extracted with two additional 100 ml portions of ethanol as
described above. The combined ethanolic extracts were then concentrated under reduced pressure at 40 C to dryness. For hot
water extractions, a subsample (10 g) was extracted by stirring
with 100 ml of boiling water at 100 C at 20g for 10 min, centrifugation at 5000g for 15 min and ltration through Whatman No.
1 lter paper. The residue was then extracted with two additional
100 ml portions of boiling water as described above. The combined
hot water extracts were then freeze dried. The dried extract was
used directly for analyses of antioxidant components or redissolved in water or ethanol to a concentration of 50 mg/ml and
stored at 4 C for further use.
2.6. Evaluation of antioxidant properties
The inhibition of conjugated diene was determined according to
the method of Lingnert, Vallentin, and Eriksson (1979). Each extract (0.520 mg/ml) in deionised water or ethanol (100 ll) was
mixed with 2 ml of 10 mM linoleic acid (Sigma) emulsion in
0.2 M sodium phosphate buffer (pH 6.6, Wako) in test tubes and
placed in the dark at 37 C to accelerate oxidation. After incubation
for 15 h, 6 ml of 60% methanol (Mallinckrodt Baker) in deionised
water was added, and the absorbance of the mixture was measured
at 234 nm against a blank in a Hitachi U-2001 spectrophotometer.
The inhibition of conjugated diene was calculated as follows:
inhibitory ability (%) = [(DA234 of control  DA234 of sample)/
DA234 of control]  100%.
The reducing capability was determined according to the method of Oyaizu (1986). Each extract (0.520 mg/ml) in deionised
water or ethanol (2.5 ml) was mixed with 2.5 ml of 200 mM sodium phosphate buffer (pH 6.6, Wako) and 2.5 ml of 1% potassium
ferricyanide (Sigma), and the mixture was incubated at 50 C for
20 min. After 2.5 ml of 10% trichloroacetic acid (Wako) was added,
the mixture was centrifuged at 200g for 10 min. The upper layer
(5 ml) was mixed with 5 ml of deionised water and 1 ml of 0.1%
ferric chloride (Wako), and the absorbance was measured at
700 nm against a blank.
Scavenging ability on 1,1-diphenyl-2-picrylhydrazyl (DPPH)
radicals was determined according to the method of Shimada,
Fujikawa, Yahara, and Nakamura (1992). Each extract (0.5
20 mg/ml) in deionised water or ethanol (4 ml) was mixed with
1 ml of methanolic solution containing DPPH (Sigma) radicals,
resulting in a nal concentration of 0.2 mM DPPH. The mixture

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S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

was shaken vigorously and left to stand for 30 min in the dark and
the absorbance was then measured at 517 nm against a blank. The
scavenging ability was calculated as follows: Scavenging ability
(%) = [(DA517 of control  DA517 of sample)/DA517 of control]  100.
The scavenging ability on hydroxyl radicals was determined
according to the method of Shi, Dalal, and Jain (1991). The hydroxyl radicals reacted with the nitrone spin trap 5,5-dimethyl pyrroline-N-oxide (DMPO, Sigma) and the resultant DMPOOH adducts
were detected with an electron paramagnetic resonance (EPR)
spectrometer. The EPR spectrum was recorded 2.5 min after mixing each extract (520 mg/ml) in deionised water or ethanol
(200 ll) with 200 ll of 10 mM H2O2 (Merck, Darmstadt, Germany),
200 ll of 10 mM Fe2+ (Sigma) and 200 ll of 10 mM DMPO using a
Bruker EMX-10 EPR spectrometer at the following settings: 0.348 T
magnetic eld, 1.0  104 T modulation amplitude, 0.5 s time constant and 200 s scan period. The scavenging ability was calculated
by subtracting the relative EPR signal intensity from 100. The relative EPR signal intensity was calculated as follows: Relative EPR
signal intensity (%) = [hDH2 (sample)/hDH2 (control)]  100; h is
the width of the peak, DH is the length of the peak.
The chelating ability was determined according to the method
of Dinis, Madeira, and Almeida (1994). Each extract (0.520 mg/
ml) in water or ethanol (1 ml) was mixed with 3.7 ml of methanol
and 0.1 ml of 2 mM ferrous chloride (Merck). The reaction was initiated by the addition of 0.2 ml of 5 mM ferrozine (Sigma). After
10 min at room temperature, the absorbance of the mixture was
determined at 562 nm against a blank. EC50 value (mg extract/
ml) is the effective concentration at which the inhibition of conjugated diene was inhibited by 50%; the absorbance was 0.5 for
reducing capability; DPPH or hydroxyl radicals were scavenged
by 50%; and ferrous ions were chelated by 50%, respectively. EC50
value was obtained by interpolation from linear regression
analysis.
2.7. Determination of antioxidant components
b-Carotene was extracted and analysed as described by Rundhaug, Pung, Read, and Bertram (1988). Each dried extract (100 mg)
was extracted with 1 ml of ethanol, 2 ml of n-hexane containing
BHA (25 lg/ml) and 1 ml of deionised water at 20g for 45 min
at room temperature and then centrifuged at 400g for 10 min.
After the removal of the n-hexane layer by N2 gas, the volume
was adjusted to 1 ml using n-hexane and ltered through a syringe-driven lter unit (13 mm, Millipore, Billerica, MA) using a
0.45 lm PVDF non-sterile lter paper. Immediately after ltration,
the ltrate was injected into a HPLC detector. The HPLC system was
the same as for the 50 -nucleotide assay. The mobile phase was
75 ml methanol/25 ml toluene at a ow rate of 1.5 ml/min and
UV detection was at 450 nm.
Tocopherols were extracted and analysed according to the
method of Carpenter (1979). Each extract (50 mg) was suspended
in 6 ml of pyrogallol (6% in ethanol) and 4 ml of 60% aqueous
potassium hydroxide solution. The resulting mixture was saponied at 70 C for 20 min. Deionised water (15 ml) was added and
the mixture was extracted with 15 ml of n-hexane. The organic
layer was washed with deionised water until neutral, then dried
over anhydrous sodium sulfate, and concentrated under reduced
pressure to dryness (<40 C). The residue was redissolved in 5 ml
of n-hexane and ltered prior to HPLC injection in the same manner as in the b-carotene assay. The HPLC system was the same as
for the 50 -nucleotide assay. The mobile phase was acetonitrile/
methanol (85:15, v/v) at a ow rate of 1.0 ml/min and UV detection
was at 295 nm.
The total phenol content was determined according to the
method of Taga, Miller, and Pratt (1984). Each extract (20 mg)
was dissolved in a solution of 5 ml of 3% HCl (Merck) in 60% meth-

anol and the resulting mixture (100 ll) was added to 2 ml of 2%

aqueous sodium carbonate (Wako) solution. After 3 min, 100 ll
of 50% Folin-Ciocalteau phenol reagent (Sigma) were added to
the mixture. After 30 min standing, the absorbance was measured
at 750 nm against a blank. The content of antioxidant components
was calculated on the basis of the calibration curve of the corresponding authentic compounds and gallic acid for total phenols
(Sigma).
2.8. Statistical analysis
For each mushroom species, three samples were used for the
determination of every quality attribute. For each of ethanolic
and hot water extractions, three samples were prepared for assays
of every antioxidant attribute and component. The experimental
data were subjected to an analysis of variance for a completely random design to determine the least signicant difference amongst
means at the level of 0.05.
3. Results and discussion
3.1. Volatile components
Volatile avour components found in the three mushrooms
were six eight-membered carbon compounds and two aromatic
compounds (Table 1). Typical mushroom aroma was attributed to
eight-membered carbon compounds, especially 1-octen-3-ol
(Mau & Hwang, 1997). However, P. ostreatus showed much higher
concentrations of eight-membered carbon compounds (>1100 mg/
kg). The major compound in C. maxima and P. ferulae was 1-octen3-ol whereas for P. ostreatus was 1-octen-3-one. It seems that aromatic compounds were insignicant in Pleurotus mushrooms, 6%
and 1% the major compound of aromatics was obtained for P. ferulae and P. ostreatus, respectively. However, in addition to the typical mushroom aroma, aromatic compounds in C. maxima might
give rise to an almond-like and fruity aroma (Arctander, 1969).
Volatile contents in Volvariella volvacea, Pleurotus eryngii and Agaricus bisporus were 8.3519.7 mg/kg (Mau et al., 1997), 17.8
29.1 mg/kg (Mau, Lin, Chen, Wu, & Peng, 1998) and 53.0 mg/kg

Table 1
Content of volatile components in Clitocybe maxima, Pleurotus ferulae and Pleurotus
ostreatus
Peak no.

Compound

RIa

Content (mg/kg fresh weight)

[% total volatiles]b

1
2
3
4
5
6
7
8

3-Octanone
1-Octen-3-one
3-Octanol
1-Octen-3-ol
Benzaldehyde
1-Octanol
2-Octen-1-ol
Benzyl alcohol

1270
1316
1406
1472
1534
1579
1637
1893

2.21 0.33
13.8 3.79
13.6 3.50
65.4 0.15
29.0 0.84
30.8 2.20
4.76 0.98
30.7 6.63

C. maxima

Total
Eight carbon
compounds
Aromatic
compounds

P. ferulae
Bc
B
B
C
A
A
B
A

3.49 0.08
1.13 0.55
0.94 0.26
98.8 5.25
0.85 0.09
5.28 0.64
0.52 0.05
6.28 0.40

P. ostreatus
B
B
B
B
C
B
C
C

138 43.9 A
506 34.9 A
181 19.4 A
254 1.17 A
2.72 0.23 B
32.5 2.06 A
6.22 0.30 A
12.9 0.23 B

190 9.39 B
131 [69]

117 4.57 B
110 [94]

1130 145 A
1120 [99]

59.6 [31]

7.13 [6]

15.6 [1]

a
Linear retention index was determined on a DB-Wax column using n-parafns
(C8C25) as reference standards.
b
% total volatiles = (the content of eight-membered carbon or aromatic compounds/total volatiles contents in each mushroom)  100%.
c
Each value is expressed as mean SD (n = 3). Means with different letters
within a row are signicantly different (P < 0.05).

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S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

Table 2
Content of soluble sugars and polyols in Clitocybe maxima, Pleurotus ferulae and
Pleurotus ostreatus
Sugar or polyol

Arabitol
Fructose
Glucose
Mannitol
Myo-inositol
Ribose
Sucrose
Trehalose
Total

Content (mg/g dry weight)

C. maxima cap

C. maxima stipe

P. ferulae

P. ostreatus

3.60 0.25 Ca
nd
1.25 0.05 C
4.45 0.45 D
nd
4.77 0.42
18.2 0.72
156 3.89 B
188 4.58 B

ndb
4.55 0.46
1.29 0.38 C
12.9 0.54 C
20.7 0.77 C
nd
nd
231 9.01 A
270 3.43 A

24.1 1.48 A
nd
13.2 0.68 A
20.6 1.22 B
34.0 0.31 B
nd
nd
33.3 1.15 C
125 3.02 D

17.1 1.35 B
nd
11.5 0.26 B
31.6 2.57 A
45.8 2.58 A
nd
nd
32.8 2.43 C
139 1.57 C

a
Each value is expressed as mean SD (n = 3). Means with different letters
within a row are signicantly different (P < 0.05).
b
Not detected.

(Mau & Hwang, 1997), respectively. It seems that these three

mushrooms showed much stronger aroma than other mushrooms
mentioned above.
3.2. Soluble sugars and polyols
The contents of total soluble sugars and polyols ranged from
125 to 270 mg/g and in the descending order of C. maxima stipe > C.
maxima cap > P. ostreatus > P. ferulae (Table 2). Stipes of C. maxima
contained much more sugars and polyols than caps and the other
two mushrooms. Obviously, C. maxima stipe was the energy reservoir for the cap. Mannitol and trehalose, which were the major
mushroom polyol and sugar, respectively (Mau et al., 1997), were
found in these three mushrooms. High amounts of trehalose were
found in C. maxima stipe and cap (156 and 230 mg/g, respectively)
whereas those in P. ferulae and P. ostreatus were relatively low
(33.3 and 32.8 mg/g, respectively). The proles of Pleurotus mushrooms were similar and consisted of arabitol, glucose, mannitol,
myo-inositol and trehalose.

As compared to other mushrooms, the total content of soluble

sugars or polyols were found to be 349457 mg/g in V. volvacea
(Mau et al., 1997), 205320 mg/g in A. bisporus (Tseng & Mau,
1999), 98.7316 mg/g in Auricularia spp. and Tremella fuciformis
(Mau & Tseng, 1998), 150225 mg/g in Agaricus blazei, A. cylindracea and Boletus edulis (Tsai, Tsai, & Mau, 2007) and 6.96
20.8 mg/g in P. eryngii (Mau et al., 1998). It seems that the content of sugars and polyols in these three mushrooms were in
the middle range. Soluble sugars contained in the mushroom contributed a sweet taste (Litcheld, 1967). Therefore, the high content of sugars and polyols would give rise to a moderately sweet
taste perception.
3.3. Free amino acids and taste components
The contents of total free amino acids in the three mushrooms
ranged from 10.5 to 37.6 mg/g and were in the descending order
of C. maxima cap > C. maxima stipe > P. ostreatus > P. ferulae (Table
3). Interestingly, the content of total free amino acids in C. maxima
cap was 2-fold higher than that found in stipe and 3-fold higher
than that found in P. ostreatus and P. ferulae. In addition, the free
amino acid proles of the three mushrooms were found to be considerably different. Surprisingly, c-aminobutyric acid (GABA), a
hypotensive agent (Kohama et al., 1987), was found in these three
mushrooms. The GABA content ranged from 0.25 to 0.45 mg/g and
was in descending order of C. maxima cap  C. maxima stipe > P.
ferulae > P. ostreatus. Furthermore, Tsai et al. (2007) found that
the content of GABA in A. blazei, A. cylindracea and B. edulis was
0.36, 0.21 and 0.11 mg/g, respectively. Since GABA is a biologically
active compound, the presence of GABA in these three mushrooms
would be benecial to humans in addition to their palatable taste
and other therapeutic effects.
In Table 3, free amino acids were divided into several classes on
the basis of their taste characteristics, as described by Komata
(1969). Aspartic and glutamic acids were the monosodium glutamate-like (MSG-like) components, which gave the most typical
mushroom taste, the umami taste or palatable taste that was the

Table 3
Content of free amino acids and taste components in Clitocybe maxima, Pleurotus ferulae and Pleurotus ostreatus
Amino acid

Content (mg/g dry weight)

C. maxima cap

C. maxima stipe

P. ferulae

L-Alanine
L-Aspartic acid
GABAa
L-Glutamic acid

5.00 0.10 Ac
2.48 0.11 A
1.48 0.07 A
0.45 0.03 A
7.41 0.43 A

2.46 0.01
0.35 0.04
0.65 0.01
0.42 0.01
4.55 0.03

B
D
B
A
B

1.30 0.07
1.30 0.06
0.36 0.02
0.31 0.02
1.40 0.09

Glycine
+ L-Threonine
L-Histidine
L-Isoleucine
L-Leucine
L-Lysine
L-Methionine
L-Phenylalanine
L-Serine
L-Tyrosine
L-Valine

0.68 0.03 B
11.78 0.35 A
0.72 0.07 B
0.06 0.01 B
1.92 0.09 A
0.23 0.02 B
0.28 0.03 B
2.82 0.12 A
0.89 0.04 A
1.44 0.11 A

0.50 0.01
3.84 0.03
0.27 0.05
1.01 0.33
0.73 0.02
0.14 0.01
0.46 0.14
1.26 0.02
0.32 0.01
0.64 0.08

C
B
C
A
C
BC
A
B
D
B

Taste componentb
Bitter
MSG-like
Sweet
Tasteless
Total

17.0 0.04
8.89 0.08
8.50 0.37
3.26 0.20
37.6 1.05

6.71 0.35
5.20 0.18
4.22 0.59
1.47 0.19
17.6 1.76

B
B
B
C
B

L-Arginine

a
b
c

A
A
A
A
A

P. ostreatus
C
B
C
B
C

1.14 0.03
1.13 0.08
0.58 0.01
0.25 0.02
1.56 0.02

D
C
B
C
C

0.80 0.01 B
0.39 0.04 D
0.72 0.06 B
0.32 0.01 B
0.75 0.02 C
1.04 0.11 A
0.42 0.09 AB
0.59 0.07 C
0.73 0.03 B
0.05 0.01 D

2.27 0.13
1.27 0.07
1.09 0.03
0.83 0.03
0.86 0.03
0.11 0.01
0.44 0.02
0.12 0.01
0.51 0.08
0.36 0.01

A
C
A
A
B
C
AB
D
C
C

4.24 0.15
1.76 0.09
2.69 0.11
1.79 0.01
10.5 0.09

5.23 0.06
2.14 0.03
3.53 0.09
1.62 0.15
12.5 0.16

C
C
C
BC
C

D
D
D
B
D

GABA, c-aminobutyric acid.

Bitter, Arg + His + Ile + Leu + Met + Phe + Val; MSG-like, monosodium glutamate-like, Asp + Glu; sweet, Ala + Gly + Ser + Thr; tasteless, Lys + Tyr + GABA.
Each value is expressed as mean SD (n = 3). Means with different letters within a row are signicantly different (P < 0.05).

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S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

characteristic taste of MSG and 50 -nucleotides (Yamaguchi, Yoshikawa, Ikeda, & Ninomiya, 1971). The contents of MSG-like components ranged from 1.76 to 8.89 mg/g and their order was the
same as that of the content of total free amino acid. For three
mushrooms, the content of bitter components was higher than
the contents of MSG-like components.
Chen (1986) conducted a series of sensory evaluations 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 was
found to be taste-active. The bitterness from the bitter components
in three mushrooms could probably be masked by the sweetness
from sweet components and mainly the high amount of soluble
sugars and polyols. Therefore, MSG-like and sweet components
would be responsible for the natural taste of these three
mushrooms.
With regard to the contents of MSG-like components in A. bisporus (22.747.1 mg/g; Tseng & Mau, 1999), these three mushrooms
possessed comparable content of MSG-like components. Furthermore, Tsai et al. (2007) found that contents of MSG-like components in A. blazei, A. cylindracea and B. edulis were 8.9714.9 mg/
g. Yang, Lin, and Mau (2001) found that contents of MSG-like components in Lentinula edodes, Flammulina velutipes strain white,
Pleurotus cystidiosus and P. ostreatus), ranged from 0.84 to
1.93 mg/g. Mau, Lin, Ma, and Song (2001a) found that contents of
MSG-like components in Dictyophora indusiata, Grifola frondosa,
Hericium erinaceus and Tricholoma giganteum ranged from 0.68 to
1.09 mg/g. Yang et al. (2001) also found that that in the yellow
strain of F. velutipes was 7.06 mg/g of MSG-like components. However, Mau, Lin, and Chen (2001b) found that contents of MSG-like
components in Ganoderma lucidum, Ganoderma tsugae and Coriolus
versicolor were in the range of 0.170.50 mg/g. Yang et al. (2001)
divided contents of MSG-like components into three ranges: low
(<5 mg/g), middle (520 mg/g) and high (>20 mg/g). Based on the
previous results, contents of MSG-like components in C. maxima
cap and stipe were in the middle range, but those in P. ostreatus
and P. ostreatus were in the low range.
3.4. 50 -Nucleotides
The contents of total 50 -nucleotides in C. maxima stipe, P. ferulae
and P. ostreatus (5.647.59 mg/g) were higher than that in C.
maxima cap (Table 4). Flavour 50 -nucleotides, which also gave the
umami or palatable taste, were found to be 50 -guanosine monophosphate (50 -GMP), 50 -inosine monophosphate (50 -IMP) and
50 -xanthosine monophosphate (50 -XMP) (Chen, 1986). Contents
of avour 50 -nucleotides in three mushrooms ranged from 1.89
to 7.59 mg/g and in the descending order of C. maxima stipe > P.
ferulae > P. ostreatus > C. maxima cap.
The contents of avour 50 -nucleotides were found to be 4.19
6.13 mg/g in A. bisporus (Tseng & Mau, 1999) and 1.634.89 mg/g
in P. eryngii (Mau et al., 1998). In addition, Yang et al. (2001) found
that contents of avour 50 -nucleotides in F. velutipes white strain, P.
cystidiosus and P. ostreatus, ranged from 5.52 to 8.60 mg/g. Mau
et al. (2001b) found that contents of avour 50 -nucleotides in D.
indusiata, G. frondosa, H. erinaceus and T. giganteum were 9.04,
0.64, 0.62 and 13.6 mg/g, respectively. The contents of avour
50 -nucleotides in G. lucidum, G. tsugae and C. versicolor were 1.18
5.65 mg/g (Mau et al., 2001a). The contents of avour 50 -nucleotides in A. blazei, A. cylindracea and B. edulis were 2.015.15 mg/g
(Tsai et al., 2007). Yang et al. (2001) reported that the contents of
avour 50 -nucleotides could be divided into three ranges: low
(<1 mg/g), middle (15 mg/g) and high (>5 mg/g). Therefore, the
content of avour 50 -nucleotides in C. maxima stipe, P. ferulae and

Table 4
Content of 50 -nucleotides in Clitocybe maxima, Pleurotus ferulae and Pleurotus ostreatus
50 -Nucleotidea

Content (mg/g dry weight)

C. maxima
cap

C. maxima
stipe

P. ferulae

P. ostreatus

0.32 0.01 C

50 -CMP

0.17 0.02
Dc
0.90 0.10 B

2.96 0.23 A

50 -GMP

0.03 0.01 D

0.11 0.01 C

50 -IMP

0.22 0.02 C

0.20 0.02 C

5 -UMP

0.28 0.04 D

0.77 0.07 C

50 -XMP

0.29 0.04 D

3.23 0.24 A

Flavour 50 nucleotideb
Total

0.54 0.08 D

3.54 0.02 A

1.89 0.61 D

7.59 0.13 A

1.48 0.02
A
0.32 0.01
C
0.58 0.02
B
0.69 0.02
B
1.94 0.01
A
1.78 0.04
B
3.05 0.01
B
6.79 0.11
B

1.39 0.02
B
0.72 0.06
B
0.61 0.01
A
0.74 0.03
A
1.05 0.01
B
1.13 0.02
C
2.48 0.01
C
5.64 0.14
C

50 -AMP

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
Flavour 50 -nucleotide, 50 -GMP + 50 -IMP + 50 -XMP.
c
Each value is expressed as mean SD (n = 3). Means with different letters
within a row are signicantly different (P < 0.05).

P. ostreatus were in the high range whereas those in C. maxima

cap were in the middle range.
50 -GMP gave the meaty avour, and is a much stronger avour
enhancer than MSG (Litcheld, 1967). The synergistic effect of avour 50 -nucleotides with MSG-like components might greatly increase the umami taste of mushrooms (Yamaguchi et al., 1971).
Using the equation derived from sensory evaluation (Yamaguchi
et al., 1971), the equivalent umami concentrations (EUC) were
46.3, 139, 56.8 and 60.4 g MSG per 100 g dry weight for C. maxima
cap and stipe, P. ferulae and P. ostreatus, respectively. Mau (2005)
reported that EUC values could be grouped into four levels: rst level of >1000 g per 100 g dry matter, second level of 1001000 g per
100 g, third level of 10100 g per 100 g, and fourth level of <10 g
per 100 g. Therefore, EUC value of C. maxima stipe was at the second level, and C. maxima cap, P. ferulae and P. ostreatus were at the
third level. In other words, the umami intensity of 1 g of C. maxima
cap and stipe, P. ferulae and P. ostreatus was equivalent to the umami intensity given by 0.46, 1.39, 0.57 and 0.60 g of MSG.
Based on the results obtained, C. maxima cap and stipe, P. ferulae
and P. ostreatus possessed a highly intense umami taste in addition
to its pharmacological properties. The sensory EUC values of three
mushrooms might be benecial for its use as foods or food-avouring materials or in the formulation of nutraceuticals and functional
foods with a palatable umami taste.
3.5. Antioxidant properties
The yields extracted by ethanol were in the descending order
of P. ferulae (18.9%) > P. ostreatus (15.3%) > C. maxima cap
(11.2%) > C. maxima stipe (5.89%). The yields extracted by hot
water were in the descending order of P. ferulae (57.2%) > P. ostreatus (44.5%) > C. maxima stipe (39.2%)  C. maxima cap (38.3%).
The yields of the hot water extracts were higher than those of
the ethanolic extracts. The use of hot water to extract soluble
components from three mushrooms was to simulate the making
of Chinese medicine and the brewing of herbal tea. Therefore,
as compared to other solvent extraction, the information obtained
by use of hot water would be more valuable for these extracts
used in human diets.

583

S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

The antioxidant properties assayed herein were summarised in

Table 5 and the results were normalised and expressed as EC50 values (mg dry weight of various extracts per ml) for comparison.
Effectiveness of antioxidant properties inversely correlated with
their EC50 values. Generally, ethanolic extracts were more effective
than hot water extracts in the inhibition of conjugated diene and
scavenging ability on DPPH radicals; whereas hot water extracts
were more effective in the scavenging ability on hydroxyl radials
as evidenced by their lower EC50 values. EC50 values were less than
14 and 30 mg/ml for ethanolic and hot water extracts, respectively,
indicating that the three mushrooms were relatively effective in
antioxidant properties, except for scavenging ability on hydroxyl
radicals.
Amongst the ethanolic extracts, P. ferulae and P. ostreatus were
more effective than C. maxima cap and stipe in antioxidant activities and chelating ability on ferrous ions. In contrast, C. maxima cap
and stipe were more effective in scavenging ability on DPPH radicals. However, none of the ethanolic extracts were effective in their
scavenging ability on hydroxyl radicals. The ethanolic extract from
P. ostreatus was more effective in the inhibition of conjugated diene
and chelating ability on ferrous ions whereas C. maxima cap was
more effective in reducing capability and scavenging ability on
DPPH radicals.
Like ethanolic extracts, hot water extracts from P. ferulae and P.
ostreatus were more effective than C. maxima cap and stipe in antioxidant activities. However, the effectiveness in reducing capability was in the descending order of C. maxima cap > C. maxima
stipe > P. ferulae > P. ostreatus. Only hot water extracts from P. ferulae and P. ostreatus showed scavenging ability on DPPH radicals.
The hot water extract from C. maxima cap was more effective in
its chelating ability on ferrous ions compared to the water extract
from P. ferulae, but lower those from C. maxima stipe and P. ostreatus. In scavenging ability on hydroxyl radicals, the effectiveness
was in the descending order of P. ferulae > P. ostreatus > C. maxima
cap whereas C. maxima stipe was not effective. However, amongst
antioxidant properties assayed, the hot water extract from P. ostreatus was slightly more effective. Overall, for both extracts, P. ostreatus was more effective amongst antioxidant properties assayed
than C. maxima and P. ferulae.
Although BHA and a-tocopherol showed good inhibitory ability
on lipid oxidation, reducing capability and scavenging ability on
DPPH radicals and EDTA was excellent for chelating ferrous ions,

they are additives and used or present in milligram levels in foods.

However, C. maxima cap and stipe, P. ferulae and P. ostreatus could
be used in gramme levels as food or a food ingredient. Therefore,
these three mushrooms might serve as possible protective agents
in human diets to help human reduce oxidative damage.
3.6. Antioxidant components
Naturally occurring antioxidant components, including b-carotene, tocopherols and total phenols, were found in two extracts
of three mushrooms (Table 6). However, ascorbic acid was not detected in two extracts and b-carotene was not detected in hot
water extracts. Total phenols were the major naturally occurring
antioxidant components found in the range of 5.519.66 mg gallic
acid equivalents/g and 5.1011.1 mg gallic acid equivalents/g for
ethanolic and hot water extracts, respectively. Contents of total
phenols were similar for both extracts of C. maxima cap and stipe
whereas for P. ferulae and P. ostreatus, contents of total phenols
were higher in hot water extracts than in ethanolic extracts.

Table 6
Contents of b-carotene, tocopherols and total phenols in ethanolic and hot water
extracts from Clitocybe maxima, Pleurotus ferulae and Pleurotus ostreatus
Contenta (mg/g dry weight)
C. maxima cap

C. maxima stipe

P. ferulae

P. ostreatus

Ethanolic
b-Carotene
a-Tocopherol
c-Tocopherol
d-Tocopherol
Total phenols

0.05 0.01 A
b 0.01 0.00 D
a 0.09 0.02 B
a 0.01 0.00 C
a 9.66 0.03 A

0.04 0.01 A
a 0.04 0.02 C
0.18 0.01 A
0.39 0.02 A
a 5.51 0.02 D

ndb
a 0.31 0.03 B
0.19 0.01 A
0.08 0.01 B
b 6.71 0.29 C

nd
0.50 0.02 A
0.17 0.03 A
0.03 0.01 C
b 7.11 0.24 B

Hot-water
b-Carotene
a-Tocopherol
c-Tocopherol
d-Tocopherol
Total phenols

nd
a 0.04 0.01 A
a 0.06 0.00
b 0.02 0.00
a 9.71 0.05 B

nd
a 0.01 0.00 B
nd
nd
b 5.10 0.01 D

nd
b 0.01 0.00 B
nd
nd
a 7.73 0.23 C

nd
nd
nd
nd
a 11.1 0.25 A

a
Each value is expressed as mean SD (n = 3). Means with different capital letters within a row are signicantly different (P < 0.05). Means with different small
letters within a column are signicantly different (P < 0.05).
b
Not detected.

Table 5
EC50 values of ethanolic and hot water extracts from Clitocybe maxima, Pleurotus ferulae and Pleurotus ostreatus in antioxidant properties
EC50 valuea (mg extract/ml)
C. maxima cap

C. maxima stipe

P. ferulae

P. ostreatus

Ethanolic
Inhibition of conjugated diene
Reducing capability
Scavenging ability on DPPH radicals
Scavenging ability on OH radicals
Chelating ability on ferrous ions

b 12.9 0.19 Ab
b 0.59 0.02 D
0.81 0.02 D
c
a 5.73 0.03 A

b 13.2 0.42 A
b 2.92 0.19 C
1.68 0.08 C
c
a 5.03 0.02 B

b 0.74 0.04 B
a 8.40 0.17 A
b 4.55 0.04 B
c
b 4.85 0.15 C

b 1.16 0.29 B
a 6.76 0.34 B
b 5.58 0.24 A
c
b 3.42 0.08 D

Hot-water
Inhibition of conjugated diene
Reducing capability
Scavenging ability on DPPH radicals
Scavenging ability on OH radicals
Chelating ability on ferrous ions

a 17.6 1.06 B
a 1.99 0.06 C
c
55.1 0.78 Ad
a 6.40 0.79 B

a 29.5 0.19 Ad
a 4.43 0.54 A
c
c
b 3.22 0.08 C

a 15.2 0.27 C
b 2.86 0.09 B
a 22.9 0.03 Ad
11.7 0.55 C
a 10.9 0.84 A

a 9.86 0.25 D
b 4.01 0.03 A
a 14.8 0.49 B
21.3 0.51 Bd
a 4.22 0.13 C

a
EC50 value: the effective concentration at which the inhibition of conjugated diene was inhibited by 50%; the absorbance was 0.5 for reducing capability; 1,1-diphenyl-2picrylhydrazyl (DPPH) or hydroxyl (OH) radicals were scavenged by 50%; and ferrous ions were chelated by 50%, respectively. EC50 value was obtained by interpolation from
linear regression analysis.
b
Each value is expressed as mean SD (n = 3). Means with different capital letters within a row at a specic antioxidant attribute signicantly different values (P < 0.05).
Means with different small letters within a column are signicantly different (P < 0.05).
c
No effect.
d
Obtained by extrapolation from linear regression analysis.

584

S.-Y. Tsai et al. / Food Chemistry 113 (2009) 578584

Phenols such as BHT and gallate were known to be effective

antioxidants (Madhavi, Singhal, & Kulkarni, 1996). Due to their
scavenging abilities on free radicals and chelating abilities on ferrous ions, phenols might possess good antioxidant, antimutagenic
and anticancer properties (Ahmad & Mukhtar, 1999). Tsai et al.
(2007) found that the contents of total antioxidant components
were moderately to highly associated (r = 0.636  0.907) with antioxidant properties. Furthermore, Dubost, Ou, and Beelman (2007)
found a good correlation between oxygen radical absorbance
capacity and polyphenols (R2 = 0.86). Therefore, the total phenols
in extracts were responsible for their effective antioxidant properties. Due to wide variations found in effectiveness of antioxidant
properties, the correlation between the total phenols and each
antioxidant attribute was not established.

4. Conclusion
Volatile avour components found were six eight-membered
carbon compounds and two aromatic compounds and the volatile
content of P. ostreatus was >1100 mg/kg. The major compound in C.
maxima and P. ferulae was 1-octen-3-ol whereas for P. ostreatus the
major compound was 1-octen-3-one. Contents of total soluble sugars and polyols ranged from 125 to 270 mg/g. High amounts of trehalose were found in C. maxima stipe and cap. However, the
proles of Pleurotus mushrooms were similar. Contents of MSGlike components ranged from 1.76 to 8.89 mg/g. The contents of total 50 -nucleotides in C. maxima stipe, P. ferulae and P. ostreatus
(5.647.59 mg/g) were higher than that in C. maxima cap. The contents of avour 50 -nucleotides ranged from 1.897.59 mg/g. Based
on the results obtained, C. maxima cap and stipe, P. ferulae and P.
ostreatus possessed highly intense umami taste.
Generally, ethanolic extracts were more effective in inhibition
of conjugated diene and scavenging ability on DPPH radicals
whereas hot water extracts were more effective in scavenging ability on hydroxyl radials. EC50 values were less than 14 and 30 mg/
ml for ethanolic and hot water extracts, respectively, indicating
that three mushrooms were effective in antioxidant properties, except for scavenging ability on hydroxyl radicals. Overall, for both
extracts, P. ostreatus was more effective amongst the antioxidant
properties assayed. The total phenols were the major naturally
occurring antioxidant components found in the range of 5.51
9.66 mg gallic acid equivalents/g and 5.1011.1 mg gallic acid
equivalents/g for ethanolic and hot water extracts, respectively.
The information obtained from this research would be helpful for
promotion of the cultivation and consumption of these three
mushrooms.
Acknowledgements
The study was supported by Council of Agriculture, ROC, Project
No. 94-AS-1.3.2-FD-Z2(14). We thank Mr. Shih-Wen Fang, Tatsuen
Farm for providing the mushrooms.
References
Ahmad, N., & Mukhtar, H. (1999). Green tea polyphenols and cancer: Biologic
mechanisms and practical implications. Nutrition Review, 57, 7883.

Ajlouni, S. O., Beelman, R. B., Thompson, D. B., & Mau, J.-L. (1995). Changes in soluble
sugars in various tissues of cultivated mushrooms, Agaricus bisporus, during
postharvest storage. In G. Charalambous (Ed.), Food avours (pp. 18651880).
Amsterdam, Netherlands: Elsevier Sci. Publ.
Arctander, S. (1969). Perfume and avour chemicals (aroma chemicals). Montclair, NJ:
Steffen Arctander.
Carpenter, A. P. (1979). Determination of tocopherols in vegetable oils. Journal of the
American Oil Chemists Society, 56, 668671.
Chen, H.-K. (1986). Studies on the characteristics of taste-active components in
mushroom concentrate and its powderization. Masters thesis, National ChungHsing University, Taichung, Taiwan.
Dinis, T. C. P., Madeira, V. M. C., & Almeida, L. M. (1994). Action of phenolic
derivatives (acetaminophen, salicylate, and 5-aminosalicylate) as inhibitors of
membrane lipid peroxidation and as peroxyl radical scavengers. Archives of
Biochemistry and Biophysics, 315, 161169.
Dubost, N. J., Ou, B., & Beelman, R. B. (2007). Quantication of polyphenols and
ergothioneine in cultivated mushrooms and correlation to total antioxidant
capacity. Food Chemistry, 105, 727735.
Kohama, Y., Matsumoto, S., Mimura, T., Tanabe, N., Inada, A., & Nakanishi, T. (1987).
Isolation and identication of hypotensive principles in red-mold rice. Chemical
and Pharmaceutical Bulletin, 35, 24842489.
Komata, Y. (1969). The taste and constituents of foods. Nippon Shokuhin Kogyo
Gakkaishi, 3, 26.
Lingnert, H., Vallentin, K., & Eriksson, C. E. (1979). Measurement of antioxidative
effect in model system. Journal of Food Processing and Preservation, 3, 87104.
Litcheld, J. H. (1967). Morel mushroom mycelium as a food avouring material.
Biotechnology and Bioengineering, 9, 289304.
Madhavi, D. L., Singhal, R. S., & Kulkarni, P. R. (1996). Technological aspects of food
antioxidants. In D. L. Madhavi, S. S. Deshpande, & D. K. Salunkhe (Eds.), Food
antioxidants: Technological, toxicological, and health prespectives (pp. 159265).
New York: Marcel Dekker.
Manzi, P., Aguzzi, A., & Pizzoferrato, L. (2001). Nutritional value of mushrooms
widely consumed in Italy. Food Chemistry, 73, 321325.
Mau, J.-L. (2005). The umami taste of edible and medicinal mushrooms.
International Journal of Medicinal Mushrooms, 7, 113119.
Mau, J.-L., Chyau, C.-C., Li, J.-Y., & Tseng, Y.-H. (1997). Flavour compounds in straw
mushrooms Volvariella volvacea harvested at different stages of maturity.
Journal of Agricultural and Food Chemistry, 45, 47264729.
Mau, J.-L., & Hwang, S.-J. (1997). Effect of gamma-irradiation on avour compounds
of fresh mushrooms. Journal of Agricultural and Food Chemistry, 45, 18491852.
Mau, J.-L., Lin, H.-C., & Chen, C.-C. (2001b). Non-volatile components of several
medicinal mushrooms. Food Research International, 34, 521526.
Mau, J.-L., Lin, Y.-P., Chen, P.-T., Wu, Y.-H., & Peng, J.-T. (1998). Flavour compounds
in king osyter mushrooms Pleurotus eryngii. Journal of Agricultural and Food
Chemistry, 46, 45874591.
Mau, J.-L., Lin, H.-C., Ma, J.-T., & Song, S.-F. (2001a). Non-volatile taste components
of several speciality mushrooms. Food Chemistry, 73, 461466.
Mau, J.-L., & Tseng, Y.-H. (1998). Non-volatile taste components of three strains of
Agrocybe cylindracea. Journal of Agricultural and Food Chemistry, 46, 20712074.
Oyaizu, M. (1986). Studies on products of browning reactions: Antioxidative
activities of products of browning reaction prepared from glucosamine.
Japanese Journal of Nutrition, 44, 307315.
Rundhaug, J. E., Pung, A., Read, C. M., & Bertram, J. S. (1988). Uptake and metabolism
of b-carotene and retinal by C3H/10T1/2 cells. Carcinogenesis, 9, 15411545.
Shi, X., Dalal, N. S., & Jain, A. C. (1991). Antioxidant behaviour of caffeine: Efcient
scavenging of hydroxyl radicals. Food Chemistry and Toxicology, 29, 16.
Shimada, K., Fujikawa, K., Yahara, K., & Nakamura, T. (1992). Antioxidative
properties of xanthan on the autoxidation of soybean oil in cyclodextrin
emulsion. Journal of Agricultural and Food Chemistry, 40, 945948.
Taga, M. S., Miller, E. E., & Pratt, D. E. (1984). Chia seeds as a source of natural lipid
antioxidants. Journal of the American Oil Chemists Society, 61, 928931.
Taylor, M. W., Hershey, R. A., Levine, R. A., Coy, K., & Olivelle, S. (1981). Improved
method of resolving nucleotides by reverse-phase high performance liquid
chromatography. Journal of Chromatography A, 219, 133139.
Tsai, S.-Y., Tsai, H.-L., & Mau, J.-L. (2007). Antioxidant properties of Agaricus blazei,
Agrocybe cylindracea and Boletus edulis. LWT Food Science and Technology, 40,
13921402.
Tseng, Y.-H., & Mau, J.-L. (1999). Contents of sugars, free amino acids and free 50 nucleotides in mushrooms, Agaricus bisporus, during postharvest storage.
Journal of the Science of Food and Agriculture, 79, 15191523.
Yamaguchi, S., Yoshikawa, T., Ikeda, S., & Ninomiya, T. (1971). Measurement of the
relative taste intensity of some a-amino acid and 50 -nucleotides. Journal of Food
Science, 36, 846849.
Yang, J.-H., Lin, H.-C., & Mau, J.-L. (2001). Non-volatile taste components of several
commercial mushrooms. Food Chemistry, 72, 465471.