Impaginato


487

Adv. Hort. Sci., 2018 32(4): 487-493 Doi: 10.13128/ahs-22569

In vitro activity of some essential oils

against Penicillium digitatum

F. Khorram, A. Ramezanian (*), M.J. Saharkhiz

Department of Horticultural Science, School of Agriculture, Shiraz

University, Shiraz, Iran.

Key words: cinnamon, citrus, decay, postharvest, savory, summer.

Abstract: Natural plant essential oils (EOs) can be used instead of synthetic

fungicides because of human health concerns and environmental protection. In

this study, the in vitro activity of some plants EOs against Penicillium digitatum,

the cause of citrus green mold was evaluated during 8 days of incubation at

25°C. The EOs extracted from sweet orange (Citrus sinensis), lemon (Citrus

limon), lime (Citrus aurantifolia), and sour orange (Citrus aurantium) fruit peel

(500, 1000 and 2000 µl l-1 concentrations), cinnamon (Cinnamomum cassia) bark

and summer savory (Satureja hortensis) aerial parts (400, 500 and 600 µl l-1 con-

centrations) were used on Penicillium digitatum mycelium. None of the EOs

extracted from tested citrus in this study could inhibit mycelial growth com-

pletely even at concentration of 2000 µl l-1. The best results were obtained with

cinnamon and summer savory EOs at concentration of 500 and 600 µl l-1. So,

based on the results, cinnamon and summer savory EOs can be ideal candidates

to replace the synthetic fungicides to control postharvest green mold of citrus

fruit. GC-MS analysis showed that the most abundant of all constituents in EO

extracts were carvacrol and γ-terpinene in summer savory and (E)-cinnamalde-

hyde in cinnamon.

1. Introduction

Citrus spp. are the most important produced fruits in the world
(Sharma and Saxena, 2004), due to their good taste, useful nutrients, and
widespread availability (Liu et al., 2012). nevertheless, the high water
content and nutrient composition make them also very susceptible to
decay by pathogens after harvest (Tripathi and Dubey, 2004). one of the
most common diseases that infects citrus fruit is green mold caused by
Penicillium digitatum (zheng et al., 2005). The yield losses and the wors-
ening of the quality caused by the fungus are economically important.
This pathogen infects the fruit through wounds on the peel inflicted dur-
ing harvest, transportation, handling or commercialization. Penicillium
digitatum is one of the most important pathogen in citrus industry,
because one generation of green mold complete during 7-10 days in rot-
ten fruit at 20-25°C, and the large amounts of spores are disseminated
easily by air currents (Palou, 2014). 

Currently, the use of synthetic fungicides is the primary and most sim-

(*) Corresponding author: 

ramezanian@shirazu.ac.ir

Citation: 

kHorrAM f., rAMezAniAn A., SAHArkHiz M.J.,
2018 - In vitro activity of some essential oils
against Penicillium digitatum. - Adv. Hort. Sci.,
32(4): 487-493

Copyright:

© 2018 khorram f., ramezanian A., Saharkhiz
M.J. This is an open access, peer reviewed article
published by firenze University Press
(http://www.fupress.net/index.php/ahs/) and
distributed under the terms of the Creative
Commons Attribution License, which permits
unrestricted use, distribution, and reproduction
in any medium, provided the original author and
source are credited.

Data Availability Statement:

All relevant data are within the paper and its
Supporting information files.

Competing Interests:

The authors declare no competing interests.

received for publication 25 January 2018
Accepted for publication 21 May 2018

AHS
Advances in Horticultural Science



Adv. Hort. Sci., 2018 32(4): 487-493

488

ple method for the control of postharvest diseases of
citrus fruit (Palou et al., 2008). However, fungicides
consumption is strongly becoming restricted because
of residual toxicity, carcinogenicity, long degradation
p e r i o d  a n d  i n c r e a s i n g  h u m a n  h e a l t h  c o n c e r n s
(Tripathi and Dubey, 2004; Palou et al., 2008).

recently, researchers have been interested in
development of alternative methods to manage
postharvest decay. The essential oils (eos) are one of
non-chemical and useful control options for the man-
agement of fungal postharvest diseases (Sassi et al.,
2008). essential oils are complex compounds that are
natural and environmentally friendly, having antioxi-
dant, antimicrobial and medicinal properties (Bakkali
et al., 2008). So, they can be ideal candidates to
replace synthetic antimicrobials for maintenance of
harvested horticultural crops (Tripathi and Dubey,
2004).

Many studies reported the beneficial effects of eo
treatments for the control of postharvest decay
caused by P. digitatum, such as Thymus vulgaris at
concentration of 1000 ppm (fatemi et al., 2012),
Mentha spicata and Lippia scaberrima at concentra-
tions of 1000 and 3000 μl l-1, respectively (Du Plooy
et al., 2009), Bubonium imbricatum at concentration
of 1000 ppm (Alilou et al., 2008), Citrus spp. at con-
c e n t r a t i o n  o f  1 0 %  ( B a d a w y  e t  a l . ,  2 0 1 1 ) ,  a n d
Cinnamomum zeylanicum at concentration of 0.5%
(kouassi et al., 2012), thereby enhancing shelf life of
fruits and vegetables.

The purpose of this study was to investigate the in
vitro activity of eos obtained from sweet orange
(Citrus sinensis), lemon (Citrus limon), lime (Citrus
aurantifolia), and sour orange (Citrus aurantium) fruit
peel, cinnamon (Cinnamomum cassia) bark and sum-
mer savory (Satureja hortensis) aerial parts for the
control of green mold caused by P. digitatum as a
preliminary study to find a suitable and effective eo
as alternative to synthetic fungicides to control green
mold in citrus postharvest management. 

2. Materials and Methods

Extraction of essential oils

Plant materials used in this study are shown in
Table 1. The air-dried plants material (300 gr) were
cut into pieces, grounded into powder by blender,
then the eos extracted through hydro-distillation for
3-4 hours using a clevenger apparatus (Miquel et al.,
1976). Then the eos were dehydrated with anhy-
drous sodium sulfate and stored in dark bottles at
-20°C before using for antifungal study.

Isolation of fungus 

The fungus used throughout this study was P. digi-
tatum, the cause of citrus green mold. for isolation
of fungus colony, P. digitatum spores were isolated
from a decayed orange and cultured on potato dex-
trose agar (PDA) by the single spore procedure 
at 25°C. The isolates were maintained on PDA until
needed.

In vitro antifungal assay

The antifungal assay was performed on PDA
plates amended with three concentrations (500,
1000 and 2000 µl l-1) of sweet orange, lemon, lime
and sour orange eos and three concentrations (400,
500 and 600 µl l-1) of cinnamon and summer savory
eos. Tween 80 (Merck-kGaA, Germany) as an emulsi-
fier was mixed with 80 ml of sterilized and molten
PDA media, cooled to about 45°C, and then enriched
with eos. There were four 80 mm plates/replicates
per treatment. After one day, the mycelia of P. digi-
tatum from 4-days-old cultures were put in the cen-
ter of amended PDA petri plates with a cork borer. All
of the plates were sealed with parafilm. inoculated
plates were kept at 25°C for 8 days. Colony diameter
was determined daily by measuring the average radi-
al growth (obagwu and korsten, 2003). in order to
evaluate its effect on fungal growth, tween 80 (emul-
sifier) was also considered as a treatment in the
experiment.

Table 1 - Plant materials used for eos extraction

name family Used part origin

Sweet orange (Citrus sinensis cv. Thomson navel) rutaceae fruit rind tissue (flavedo and albedo) fars-iran

Lemon (Citrus limon cv. Lisbon) rutaceae fruit rind tissue (flavedo and albedo) fars-iran

Lime (Citrus aurantifolia cv. Mexican lime) rutaceae fruit rind tissue (flavedo and albedo) fars-iran

Sour orange (Citrus aurantium cv. amara) rutaceae fruit rind tissue (flavedo and albedo) fars-iran

Cinnamon (Cinnamomum cassia) Lauraceae Tree bark China

Summer savory (Satureja hortensis) Lamiaceae Aerial parts fars-iran



Khorram et al. - Essential oils against Penicillium digitatum

489

inhibition percentage (iP) of fungal growth was
calculated as the radial growth of treated fungus (T)
relative to the growth in control (C) treatment (plates
without eo and Tween 80) according to the following
formula:

Essential oils analysis

At the end of the study, the main components of
the most effective eos on P. digitatum were analyzed
by gas chromatography (GC) and gas chromatogra-
phy-mass spectrometry (GC-MS). The GC analysis was
c a r r i e d  o u t  b y  t h e  u s e  o f  A g i l e n t  G C  ( 7 8 9 0 - A ,
Perkinelmer, USA) and a flame ionization detector. it
was done on fused silica capillary HP-5 column. The
temperatures of injector and detector were held at
250°C and 280°C, respectively. nitrogen was selected
as carrier gas; oven temperature was 60-210°C at a
rate of 4°C/min, which was then increased to 240°C
at a rate of 20°C/min, and finally, kept for 8.5 min.

The GC-MS analysis was performed using an
Agilent GC series 7890-A (Perkinelmer, USA) with a
fused silica capillary HP-5MS column and 5975-C
mass spectrometer (UniCo, USA). Carrier gas was
helium. ion source and interface temperatures were
set at 230°C and 280°C, respectively. Mass range was
programmed from 45 to 550 amu. oven temperature

was 60-210°C at a rate of 4°C/min. n-alkanes was
used as a standard to determine the retention
indices for all constituents. The constituents were
recognized by comparing their retention indices with
literature reports, and their mass spectra comparison
with the Wiley, Adams and Mass finder 2.1 Library
data (Adams, 1997).

Statistical analysis

The experiment was distributed according to a
split plot in time design. The analysis of variance
(AnoVA) was performed. Mean comparisons were
conducted by LSD (least significant difference) at P≤
0.01. Data were analyzed by SAS software (v. 9.1).

3. Results and Discussion

I n h i b i t o r y  e f f e c t s  o f  d i f f e r e n t  t r e a t m e n t s  o n

Penicillium digitatum growth

The in vitro activity of the tested eos on colony
diameter of P. digitatum during 8 days of incubation
is summarized in Table 2.

our results indicated that colony radial growth of
P. digitatum was inhibited completely (100%) under
in vitro condition by both cinnamon and summer
savory eos at 500 and 600 µl l-1 concentrations dur-
ing 8 days of incubation. Also, the mycelial growth

Table 2 - inhibition percentage (%) of plant essential oils on in vitro radial growth of Penicillium digitatum

* for each column, similar letters (lower case, superscript) are not significantly different according to LSD (P≤0.01) test.
^ Means followed by similar letters (subscript), are not significantly different according to LSD (P≤0.01) test.

Treatment
eo

Concentration
(µl l-1)

Time (day)

1 2 3 4 5 6 7 8

Control - 0.00 g * S^ 0.00 
g

S
0.00 h

S
0.00 g

S
0.00 e

S
0.00 g

S
0.00 e

S
0.00 e

S

Tween 80 - 0.80 g Q-S 0.43 
g

rS
0.38 h

rS
1.29 g

P-S
1.24 e

P-S
5.09 fg

n-S
7.08 e

L-S
8.13 d e

k-S

Sweet orange 500 22.42 f W-i 5.81 
fg

L-S
10.34 gh

i-S
12.73 fg

D-Q
6.59 e

L-S
9.37 fg

J-S
6.55 e

L-S
0.00 e

S

Sweet orange 1000 23.81 f W-g 5.75 
fg

M-S
18.02 e-g

B-k
23.07 ef

W-h
12.56 e

D-r
16.26 e-g

C-n
11.55 e

G-S
0.00 e

S

Sweet orange 2000 75.80 b B 54.74 
b

e-i
48.24 c

i-o
50.72 c

G-k
32.09 cd

r-b
38.42 cd

L-S
41.76 c

J-S
36.56 bc

o-V

Lemon 500 32.81 ef r-a 17.41 
d-g

C-M
20.79 e-g 

z-j
23.18 ef

W-h
13.03 e

D-Q
12.79 fg

D-Q
12.19 e

f-S 0.00 
e S

Lemon 1000 46.91 c-e L-P 24.46 
c-f

V-e
38.18 cd

M-S
41.15 cd

J-S
17.42 c-e

C-M
22.27 d-f

X-i
20.14 de

B-k
12.50 c-e

e-r

Lemon 2000 61.38 b-d D-H 23.54 
c-f

W-H
45.84 c 

i-Q
50.87 c

G-k
34.39 c

Q-X
44.26 c

i-r
34.66 cd

P-W
25.31 b-e

T-c

Lime 500 36.98 ef M-U 20.71 
d-f

A-i
21.95 e-g

Y-i
17.98 f

C-M
15.74 de

C-n
16.98 e-g

C-n
17.25 de

C-n
0.00 e

S

Lime 1000 54.54 b-e e-i 30.49 
c-e

S-b
44.03 c

i-r
48.19 c

i-o
30.72 cd

S-B
33.27 c-e

r-Y
33.22 cd

r-Y
30.85 b-d

S-b

Lime 2000 70.43 bc B-D 42.48 
bc

i-S
49.22 c

H-M
50.62 c

G-L
34.11 c

Q-Y
41.72 c

J-S
40.39 c

k-S
32.50 b-d

r-a

Sour orange 500 48.92 c-e i-n 36.82 
b-d

n-U
14.64 fg

C-o
12.61 fg

D-r
11.36 e

H-S
16.16 e-g

c-n
17.89 de

C-M
0.00 e

S

Sour orange 1000 50.94 c-e G-k 24.00 
c-f

W-f
30.77 de

S-b
33.67 de

Q-Y
13.36 e

C-P
15.97 e-g

C-n
14.85 de

C-o
9.37 de

J-S

Sour orange 2000 61.44 b-d D-H 33.01 
c-e

r-z
44.12 c

i-r
53.39 c

f-J
33.12 c

r-Y
37.28 cd

M-T
41.08 c

J-S
40.94 b

k-S

Cinnamon 400 40.52 d-f k-S 12.63 
e-g

D-r
24.77 ef

U-d
22.22 ef

X-i
3.34 e

o-S
6.48 fg

L-S
9.40 e

J-S
3.44 e

o-S

Cinnamon 500 100.00 a A 100.00 
a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A

Cinnamon 600 100.00 a A 100.00 
a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A

Savory 400 100.00 a A 89.61 
a

A
74.01 b

BC
70.91 b

B-D
66.01 b

B-e
65.25 b

B-f
61.83 b

C-G
51.25 b

G-k

Savory 500 100.00 a A 100.00 
a

A
100.00 a

A
100.00 a

A
100.0 0 a

A
100.00 a

A
100.00 a

A
100.00 a

A

Savory 600 100.00 a A 100.00 
a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A
100.00 a

A



Adv. Hort. Sci., 2018 32(4): 487-493

490

was decreased by cinnamon and summer savory eos
at concentrations lower than 500 µl l-1, but it was not
suppressed completely. 

The main activity of the eos in the postharvest
fruit are derived from their ability to inhibit pathogen
growth (Periago et al., 2004). Cinnamon eo has the
potential to be employed as a natural antifungal agent
for fruit disinfectation, as cinnamaldehyde is its main
constituent (Xing et al., 2010). furthermore, it has been
reported that summer savory contains some substances
with antibacterial properties (Deans and Svoboda,
1989). in this research, cinnamon and summer savory
eos have the strongest effect on P. digitatum growth
(Table 2). it has been reported that eucalyptus and cin-
namon (Cinnamomum zeylanicum, Blume) oil vapour
(500 ppm) reduced decay almost by 50% in tomatoes
after 10 days of storage (Tzortzakis, 2007). Moreover,
Win et al. (2007) presented that eos from cinnamon at
the concentration of 5.0 g l-1 completely inhibited coni-
dial germination and mycelial growth of all fungi on
banana (Colletotrichum musae, Fusarium spp. and
Lasiodiplodia theobromae). in addition, Lopez-reyes et
al. (2010) showed that summer savory, oregano and
thyme eos at 10% showed significant inhibitory effect
(similar to chemical control) against P. expansum and
Botrytis cinerea on four cultivars of apples.

The mechanism by which eos suppress the micro-
bial growth is not fully understood, but a number of
possible explanations have been postulated. essential
oils are lipophilic and this property enables them to
preferentially move from an aqueous phase into fungi
membrane. This action leads to membrane expan-
sion, increasing in membrane fluidity and permeabili-
ty, membrane proteins disorder, respiration rate con-
trol, change of ion transportation in fungi and induced
cellular contents leakage (Burt, 2004; oonmetta-Aree
et al., 2006; khan et al., 2010; fadli et al., 2012).

in the present study, the lowest inhibition was
observed in control plates that contained only PDA
(0%); however, this was not significantly different
from plates containing PDA and the tween 80 with-
out the eos during 8 days. So, results indicated that
the tween 80 used as an emulsifier had no effect on
the mycelial growth (Table 2).

We observed an increase of antifungal effects of
the tested citrus fruits peel eos such as sweet
orange, lemon, lime and sour orange as the eos con-
centration increased, but the fungi growth was not
inhibited completely even at concentration of 2000
µl l-1. So, as the results showed, none of the tested
concentrations of citrus fruits eos in this study could
inhibit radial growth completely (Table 2).

essential oils are present in great quantities in the
flavedo of citrus fruit (Caccioni et al., 1998). The cit-
rus fruits eo consists a mixture of components such
as terpenes, hydrocarbons, ketones, aldehydes, alco-
hols, acids, and esters. The amount of them depends
on the citrus cultivar, the extraction and separation
techniques (fisher and Phillips, 2008).

The positive effect of the volatile components of
citrus fruit essential oils on P. digitatum and italicum
growth has been reported (Caccioni et al., 1998). The
s p o r e  g e r m i n a t i o n  a n d  m y c e l i u m  g r o w t h  o f  P .
italicum and digitatum were stimulated by the essen-
tial oil of Citrus reticulata Blanco at concentration of
more than 2.5 μl ml-1 (Wang et al., 2012). Moreover,
Badawy et al. (2011) reported that Citrus aurantifolia
eos had the antifungal effects against P. digitatum
pathogens at concentration of 10% (v/v). However, in
our study the application of Citrus spp. could not pro-
vide acceptable control of green mold disease.

Analysis of the summer savory and cinnamon EOs

The analysis of the volatile profiles in summer
savory and cinnamon eos are listed in Table 3 and 4,

Table 3 - Chemical composition of the summer savory essential
oil

* retention index

number Component ri* (%)

1 α- Thujene 924 1.15

2 α-Pinene 932 0.64

3 Camphene 946 0.06

4 Hepten-1-ol 958 0.05

5 Sabinene 969 0.01

6 β-Pinene 974 0.21

7 3- Myrcene 988 1.15

8 Phellandrene 1002 0.23

9 α-Terpinene 1014 3.75

10 p-Cymene 1020 2.19

11 Sylvestrene 1025 0.37

12 e-β- ocimene 1044 0.07

13 γ-Terpinene 1054 31.98

14 Terpinolene 1086 0.05

15 trans-α Sabinene hydrate 1098 0.07

16 isoborneol 1155 0.06

17 Terpinene-4-ol 1174 0.2

18 α-Terpineol 1186 0.1

19 carvacrol methyl ether 1241 0.09

20 Thymol 1289 0.8

21 Carvacrol 1298 55.66

22 Thymol acetate 1349 0.03

23 Carvacrol acetate 1370 0.07

24 Caryophyllene 1417 0.36

25 Aromadendrene 1439 0.08

26 α–Humulene 1454 0.01

27 Bicyclogermacrene 1500 0.18

28 Bisabolene 1505 0.21

29 Unknown - 0.01

30 Spathulenol 1577 0.04



Khorram et al. - Essential oils against Penicillium digitatum

491

respectively. A total of 30 different components of
summer savory, and 37 components of cinnamon
were identified and isolated by GC and GC-MS from
the eos. The principal components of the summer
savory eo were carvacrol (55.66%), γ-terpinene
(31.98%), α-terpinene (3.75%), p-cymene (2.19%), 3-
myrcene (1.15%), and α-thujene (1.15%). The major
components of the cinnamon eo were (e)-cin-
namaldehyde (70.04%), α-copaene (10.82%), δ-
cadinene (5.35%), α-muurolene (4.23%), and γ-
muurolene (1.25%). other constituents which were
less than 1% have been shown in Table 3 and 4. 

As shown in Table 2, both summer savory and cin-
namon eos were equally effective in inhibiting the
growth of P. digitatum. This is in accord with the
reported in vitro inhibitory effect of carvacrol against

pathogens (Periago et al., 2004). in fact, the main
c o m p o n e n t  o f  s u m m e r  s a v o r y e o  i s  a  p h e n o l
(Sacchetti et al., 2005), and its most important mech-
anism of antimicrobial activity is connected with the
phenolic ring in its chemical structure (Ultee et al.,
2002). furthermore, it has been reported that the
toxicity rate of the phenol ring is due to the site (s)
and number of hydroxyl groups (Cowan, 1999).
Concerning cinnamon eos, its major volatile com-
pound is cinnamaldehyde. Moreover, it has been
reported that cinnamon eo had potent anti-bacterial
and anti-fungal activities due to cinnamaldehyde (ooi
et al., 2006), because it acts as membrane irritants
(nabavi et al., 2015).

4. Conclusions

in this study, the in vitro activity of plants eos
against P. digitatum were tested at different concen-
trations during 8 days of incubation at 25°C. As
showed by the results, the stronger inhibitions were
obtained by cinnamon and summer savory eos at
concentration of 500 and 600 µl l-1. none of the citrus
eos could inhibit fungus radial growth completely
compared with cinnamon and savory eos. GC-MS
analysis showed that the most abundant of all con-
stituents in eo extracts were carvacrol and γ-ter-
pinene in summer savory and (e)-cinnamaldehyde in
cinnamon. 

Acknowledgements

We thank Shiraz University research Council for
financial supports, fars Agricultural and natural
resources research and education Center, Shiraz, iran
for technical assistance.

References

ADAMS r.P., 1997 - Identification of essential oil compo-
nents by gas chromatography/mass spectroscopy. - J.
Am. Soc. Mass Spect., 6(8): 671-672.

ALiLoU H., AkSSirAr M., HASSAni L.M.i., CHeBLi B., eL
HAkMoUi A., MeLLoUki f., roUHi r., BoirA H.,
BLÁzQUez M.A., 2008 - Chemical composition and anti-
fungal activity of Bubonium imbricatum volatile oil. -
Phytopathol. Mediterr., 47(1): 3-10.

BADAWY f.i., SALLAM M.n., iBrAHiM A., ASrAn, M., 2011
- Efficacy of some essential oils on controlling green
mold of orange and their effects on postharvest quality

Table 4 - Chemical composition of the cinnamon essential oil

* retention index

number Component ri* (%)

1 α-Pinene 932 0.57

2 Camphene 946 0.34

3 Benzaldehyde 958 0.49

4 β -Pinene 975 0.16

5 α-Phellandrene 1004 0.02

6 p-Cymene 1023 0.07

7 Limonene 1026 0.13

8 1,8-Cineole 1029 0.07

9 γ-Terpinene 1056 0.04

10 Benzenepropanal 1160 0.26

11 Borneol 1163 0.19

12 α-Terpineol 1189 0.03

13 (z)-Cinnamaldehyde 1217 0.73

14 (e)-Cinnamaldehyde 1271 70.04

15 Unknown 1333 0.06

16 Cyclosativene 1367 0.59

17 α-Copaene 1373 10.82

18 Unknown 1387 0.18

19 β-elemene 1389 0.14

20 Sativene 1393 0.44

21 (e)-Caryophyllene 1416 0.20

22 β-Gurjunene 1426 0.06

23 α -Humulene 1450 0.20

24 γ-Muurolene 1474 1.25

25 ar-Curcumene 1480 0.15

26 Viridiflorene 1492 0.28

27 α -Muurolene 1497 4.23

28 β-Bisabolene 1506 0.15

29 γ-Cadinene 1511 0.26

30 δ-Cadinene 1521 5.35

31 (e)-ortho-Methoxy cinnamaldehyde 1528 0.27

32 trans-Cadina-1(2),4-diene 1529 0.96

33 α -Calacorene 1540 0.50

34 epi-a-Muurolol 1639 0.40

35 α -Muurolol 1643 0.20

36 α -Cadinol 1651 0.03

37 Cadalene 1671 0.10



492

Adv. Hort. Sci., 2018 32(4): 487-493

parameters. - Plant Pathol. J., 10(4): 168-174.
BAkkALi f., AVerBeCk S., AVerBeCk D., iDAoMAr M.,

2008 - Biological effects of essential oils - A review. -
food Chem. Toxicol., 46(2): 446-475.

BUrT S., 2004 - Essential oils: their antibacterial properties
and potential applications in foods-a review. - int. J.
food Microbiol., 94(3): 223-253.

CACCioni D.r., GUizzArDi M., BionDi D.M., renDA A.,
rUBerTo G., 1998 - Relationship between volatile com-
ponents of citrus fruit essential oils and antimicrobial

a c t i o n  o n  P e n i c i l l i u m  d i g i t a t u m a n d  P e n i c i l l i u m
italicum. - int. J. food Microbiol., 43(1): 73-79.

CoWAn M.M., 1999 - Plant products as antimicrobial
agents. - Clin. Microbiol. rev., 12(4): 564-582.

DeAnS G., SVoBoDA k.P., 1989. Antibacterial activity of
summer savory (Satureja hortensis L) essential oil and
its constituents. - J. Hortic Sci., 64(2): 205-210.

DU PLooY W., reGnier T., CoMBrinCk S., 2009 - Essential
oil amended coatings as alternatives to synthetic fungi-

cides in citrus postharvest management. - Postharvest
Biol. Technol., 53(3): 117-122.

fADLi M., SAAD A., SAYADi S., CHeVALier J., MezrioUi
n.e., PAGÈS J.M., HASSAni L., 2012 - Antibacterial
activity of Thymus maroccanus and Thymus brous-
sonetii essential oils against nosocomial infection-bac-
teria and their synergistic potential with antibiotics. -
Phytomedicine, 19(5): 464-471.

fATeMi S., JAfArPoUr M., eGHBALSAieD S., 2012 - Study
of the effect of Thymus vulgaris and hot water treat-
ment on storage life of orange (Citrus sinensis CV.
Valencia). - J. Med. Plants res., 6(6): 968-971.

fiSHer k., PHiLLiPS C., 2008 - Potential antimicrobial uses
of essential oils in food: is citrus the answer?. - Trends
food Sci. Technol., 19(3): 156-164.

kHAn A., AHMAD A., AkHTAr f., YoUSUf S., XeSS i., kHAn
L.A., MAnzoor n., 2010 - ocimum sanctum essential
oil and its active principles exert their antifungal activi-

ty by disrupting ergosterol biosynthesis and membrane

integrity. - res. Microbiol., 161(10): 816-823.
koUASSi k.H.S., BAJJi M., JiJAkLi H., 2012 - The control of

postharvest blue and green molds of citrus in relation
with essential oil–wax formulations, adherence and vis-

cosity. - Postharvest Biol. Technol., 73: 122-128.
LiU Y., HeYinG e., TAnUMiHArDJo S., 2012 - History, glob-

al distribution, and nutritional importance of citrus

fruits. - Compr. rev. food. Sci. food. Saf., 11: 530-545.
LoPez-reYeS J.G., SPADAro D., GULLino M.L., GAriBALDi

A., 2010 - Efficacy of plant essential oils on postharvest
control of rot caused by fungi on four cultivars of

apples in vivo. - flavour fragr. J., 25(3): 171-177.
MiQUeL J., riCHArD H., SAnDreT f., 1976 - Volatile con-

stituents of Moroccan thyme oil. - J. Agric. food.
Chem., 24(4): 833-835.

nABAVi S.f., Di Lorenzo A., izADi M., SoBArzo-SÁnCHez
e., DAGLiA M., nABAVi S.M., 2015 - Antibacterial
effects of cinnamon: From farm to food, cosmetic and

pharmaceutical  industries. - nutrients, 7(9): 7729-

7748.
oBAGWU J., korSTen L., 2003 - Control of citrus green and

blue molds with garlic extracts. - eur. J. Plant Pathol.,
109(3): 221-225.

ooi L.S., Li Y., kAM S.L., WAnG H., WonG e.Y., ooi V.e.,
2006 - Antimicrobial activities of cinnamon oil and cin-
n a m a l d e h y d e  f r o m  t h e  C h i n e s e  m e d i c i n a l  h e r b

Cinnamomum cassia Blume. - Am. J. Chin. Med.,
34(03): 511-522.

oonMeTTA-Aree J., SUzUki T., GASALUCk P., eUMkeB G.,
2006 - Antimicrobial properties and action of galangal
(Alpinia galanga Linn.) on Staphylococcus aureus. -
LWT-food Sci. Technol., 39(10): 1214-1220.

PALoU L., 2014 - Penicillium digitatum, Penicillium italicum
(Green mold, Blue mold), Postharvest decay. Control

Strategies. - Academic Press, elsevier inc., London, Uk,
pp. 102.

PALoU L., SMiLAniCk J.L., DroBY S., 2008 - Alternatives to
conventional fungicides for the control of citrus

p o s t h a r v e s t  g r e e n  a n d  b l u e  m o u l d s . -  S t e w a r t
Postharvest rev., 2(2): 1-16.

PeriAGo P.M., DeLGADo B., fernÁnDez P.S., PALoP A.,
2004 - Use of carvacrol and cymene to control growth
and viability of Listeria monocytogenes cells and pre-
dictions of survivors using frequency distribution func-

tions. - J. food Prot., 67(7): 1408-1416.
SACCHeTTi G., MAieTTi S., MUzzoLi M., SCAGLiAnTi M.,

M A n f r e D i n i  S . ,  r A D i C e  M . ,  B r U n i  r . ,  2 0 0 5  -
Comparative evaluation of 11 essential oils of different

origin as functional antioxidants, antiradicals and

antimicrobials in foods. - food Chem., 91(4): 621-632.
SASSi A.B., HArzALLAH-SkHiri f., CHrAief i., BoUr-

GoUGnon n., HAMMAMi M., AoUni M., 2008 -
Chemical composition and antimicrobial activities of

the essential oil of (Tunisian) Chrysanthemum trifurca-
tum (Desf.) Batt. and Trab. flowerheads. - Comptes
rendus Chimie, 11(3): 324-330.

SHArMA r., SAXenA S., 2004 - Rootstocks influence granu-
lation in Kinnow mandarin (Citrus nobilis × C. deliciosa).
- Sci. Hortic., 101(3): 235-242.

TriPATHi P., DUBeY n., 2004 - Exploitation of natural prod-
ucts as an alternative strategy to control postharvest

fungal rotting of fruit and vegetables. - Postharvest
Biol. Technol., 32(3): 235-245.

TzorTzAkiS n.G., 2007 - Maintaining postharvest quality
of fresh produce with volatile compounds. - innov. food
Sci. emerg. Technol., 8(1): 111-116.

ULTee A., Bennik M., MoezeLAAr r. 2002 - The phenolic
hydroxyl group of carvacrol is essential for action

against the food-borne pathogen Bacillus cereus. -
Appl. environ. Microbiol., 68(4): 1561-1568.

WAnG H., TAo n., HUAnG S., LiU Y., 2012 - Effect of
Shatangju (Citrus reticulata Blanco) essential oil on
spore germination and mycelium growth of Penicillium
digitatum and P. italicum. - J. essent. oil Bear., 15(5):
715-723.

Win n.k.k., JiTAreerAT P., kAnLAYAnArAT S., SAnG-



Khorram et al. - Essential oils against Penicillium digitatum

493

CHoTe S., 2007 - Effects of cinnamon extract, chitosan
coating, hot water treatment and their combinations

on crown rot disease and quality of banana fruit. -
Postharvest Biol. Technol., 45: 333-340.

XinG Y., Li X., XU Q., YUn J., LU Y., 2010 - Antifungal activities
of cinnamon oil against rhizopus nigricans, Aspergillus

flavus and Penicillium expansum in vitro and in vivo fruit
test. - int. J. food Sci. Technol., 45(9): 1837-1842.

zHenG X.D., zHAnG H.Y., SUn P., 2005 - Biological control
o f  p o s t h a r v e s t  g r e e n  m o l d  d e c a y  o f  o r a n g e s  b y

rhodotorula glutinis. - eur. food res. Technol., 220(3-
4): 353-357.