Received: 8 November 2020 Revised: 26 August 2021 Accepted: 28 August 2021

DOI: 10.1002/ptr.7281

REVIEW

Antifungal and mycotoxin detoxification ability of essential

oils: A review

Jing Cai1 | Rong Yan1 | Jichao Shi2 | Jia Chen1 | Miao Long1 |
3,4 4
Wenda Wu | Kamil Kuca

1
Key Laboratory of Zoonosis of Liaoning
Province, College of Animal Science & With increased popular awareness of food safety and environmental protection,
Veterinary Medicine, Shenyang Agricultural
plant essential oil has attracted interest due to the absence of residue, its high effi-
University, Shenyang, China
2
Liaoning Service Development Center, ciency, antioxidant, immune regulation, antibacterial, insecticidal, and other advan-
Shenyang, China tages. Their application in degradation and elimination of mycotoxin toxicity has
3
MOE Joint International Research Laboratory
attracted increasing attention. This paper reviews the structure, antibacterial activity,
of Animal Health and Food Safety, College of
Veterinary Medicine, Nanjing Agricultural antibacterial mechanism, and toxic effects of essential oils. The inhibitory effects of
University, Nanjing, China
various essential oils on different mycotoxins were studied. The research progress on
4
Department of Chemistry, Faculty of Science,
University of Hradec Kralove, Hradec Kralove,
the inhibitory effects of plant essential oils on fungi and mycotoxins in recent years
Czech Republic was summarized to provide reference for the application of plant essential oils.

Correspondence
KEYWORDS
Miao Long, Key Laboratory of Zoonosis of
Liaoning Province, College of Animal Science & essential oils, fungi, inhibition, mycotoxin
Veterinary Medicine, Shenyang Agricultural
University, Shenyang 110866, China.
Email: longmiao@syau.edu.cn (M. L.)

Wenda Wu, MOE Joint International Research

Laboratory of Animal Health and Food Safety,
College of Veterinary Medicine, Nanjing
Agricultural University, Nanjing 210095,
China.
Email: wuwenda@njau.edu.cn (W. W.)
Kamil Kuca, Department of Chemistry, Faculty
of Science, University of Hradec Kralove,
50003 Hradec Kralove, Czech Republic.
Email: kamil.kuca@uhk.cz (K. K.)

Funding information
China Postdoctoral Science Foundation, Grant/
Award Number: 2016T90477; National Key
R & D Program, Grant/Award Numbers:
2016YFD0501207, 2016YFD0501009;
National Natural Science Foundation of China,
Grant/Award Numbers: 31972746, 31772809,
31872538; UHK (University of Hradec
Kralove), Grant/Award Number: VT2019-2021

1 | INTRODUCTION
List of Abbreviations: ATP, adenosine triphosphate; AFB1, fumonisin B1; AFB2, fumonisin
B2; AFG1, aflatoxin G1; AFG2, aflatoxin G2; CO2, carbondioxide; DNA, deoxyribonucleic
acid; DON, deoxynivalenol; HGB, hemoglobin; LYM, ymphocytes; MAE, microwave-assisted Plant essential oil is a substance originally derived from plants, also
extraction; mRNA, messenger RNA; OTA, ochratoxin; PLE, macrocyclic dibenzyl; RNA,
known as aromatic oil or volatile oil. It is a secondary metabolite
ribonucleicacid; ROS, reactive oxygen species; UAE, ultrasonic-assisted extraction; WBC,
white blood cell. extracted from plant tissues such as buds, stems, leaves, flowers,

Phytotherapy Research. 2021;1–11. wileyonlinelibrary.com/journal/ptr © 2021 John Wiley & Sons Ltd. 1
2 CAI ET AL.
seeds, roots, or bark. It is extracted from different plants and different
plant essential oils are generally of different colors. In most cases, they
are oily. Generally, they are named according to the source of plant
essential oils. Volatile oils with strong aromaticity are usually present
in secretory tissue cells (Perczak et al., 2016). Some plant essential oils
can also precipitate solid components at room temperature. Most
essential oils are colorless or light yellow, contain hydrophobic groups,
are easily soluble in organic solvents, are volatile at room temperature,
impart a particular, strong flavor, have strong penetration of flavor
components, long duration, a relative boiling point between 70 C and
 3
300 C, and a relative density of between 0.850 and 1.065 g/cm .
Mycotoxins are metabolites produced by fungi growing in food or
feed. They are highly toxic, and exhibit carcinogenesis, mutagenesis,
teratogenesis, immune system inhibition, nephrotoxicity, and hepato-
toxicity. Their harm is far greater than pesticide residues and food
additives (Chen et al., 2020). A variety of mycotoxins can induce
tumors in different tissues and organs of experimental animals
(Wagacha & Muthomi, 2008). Given mycotoxins are produced by
fungi, it is essential to find some new antifungal substances to control
or detoxify mycotoxins; however, the traditional method of adding
antibiotics to feed is likely to lead to adverse consequences such as
drug resistance and drug residues in the body of livestock and poultry
and incurs certain production and food safety risks. Due to the poten-
tial risk of bacterial resistance to synthetic antimicrobial agents, more
attention has been paid to the use of natural antimicrobial agents,
especially those extracted from plants, such as essential oils. Com-
pared with traditional chemical fungicides, natural plant essential oils
exhibit the advantages of broad-spectrum antibacterial effect, low
toxicity, and a high rate of utilization (Baczek et al., 2017; Miguel
et al., 2018; Oliveira et al., 2019; Puškárová, Bučková, Kraková, Pan-
gallo, & Kozics, 2017). The combination of essential oils may benefit
gastrointestinal health and be applied as an alternative to antibiotics
(Xue et al., 2020). Therefore, analysis of the inhibitory effect of plant
essential oil on fungi and mycotoxins will help us to eliminate pollu-
tion arising from fungi and mycotoxins and provide better protection
for human and animal health. In this article, we review the antifungal
and detoxification mycotoxin ability of essential oils.
1.1 | Plant essential oils
Essential oils are a mixture of many complex compounds, and the
chemical composition and concentration of different plant essential
oils differ. For example, thymol and vanillin are the main components
of thyme essential oils. The two can account for 3–62% of the total
essential oils of thyme (Arras, 1978). Cinnamaldehyde, the main
component of cinnamon essential oils, accounts for 60–75% of the
total essential oil thereof (Austin & Therapies, 2003). The compo-
nents of plant essential oils are relatively complex, and mainly com-
posed of aliphatic compounds, aromatic compounds, nitrogen-
containing sulfur compounds, terpenes, and their derivatives. Among
them, terpenoids are the most common and the prevalent compo-
nents in plant essential oils, accounting for more than 70%, and some
even exceeding 85%, especially monoterpenes (such as nerol and cit-
ronellol). The bioactivity of plant essential oils is mainly determined
by sesquiterpenes (such as nerolidol and terpineol) and diterpenoids
(such as phytol, vitamin A, and alcohols) (Cui, Zhang, Zhou, Zhao, &
Lin, 2015).
1.2 | Extraction of plant essential oils
Although plant essential oils come from plants, not all subsces
extracted from plants are essential oils, because plants contain a vari-
ety of active substances, and essential oils have their own unique
physical and chemical properties. The common extraction methods of
plant essential oils include distillation, organic solvent extraction,
pressing, adsorption extraction, lipolysis, enzyme fermentation, super-
critical CO2 extraction, etc. (Moumni et al., 2020). The extraction effi-
ciency of essential oils by distillation and pressing was lower than that
achieved when using microwave-assisted extraction (Markovi
c
et al., 2018; Reyes-Jurado, Franco-Vega, Ramírez-Corona, Palou, &

pez-Malo, 2014). Table 1 lists the extraction methods used for sev-
Lo
eral essential oils.
Steam distillation uses steam to release aromatic compounds
from plants and transport them for cooling with resulting water vapor
(Zhang et al., 2012). The purpose of separation is to exploit the differ-
ence in density between oil and water (Pateiro et al., 2018). The disad-
vantage of this method is that the process is slow and the extraction
temperature is high, leading to the degradation of some components
(Gavahian, Hashemi, Khaneghah, & Tehrani, 2013).
Cold pressing is a method that destroys the cell structure of
plants and makes essential oils flow from the secretory tissue under
the influence of mechanical extrusion. The advantage of cold pressing
is that it can be undertaken at room temperature without damaging
the components of the essential oils. The disadvantage is that the
yield is low, and the purification steps in the later stage are compli-
cated, which renders it unsuitable for large-scale production
(Kapsaski-Kanelli et al., 2017).
The extraction method uses hydrophobic and non-polar charac-
teristics of essential oils to extract lipophilic substances from plants
with non-polar solvents such as ethanol (Toma, Vinatoru, Paniwnyk, &
Mason, 2001), however, this method is only suitable for laboratory
research, because this method can produce many non-volatile macro-
molecular substances which are difficult to separate, making subse-
quent purification difficult.
Supercritical CO2 extraction was used to change the density of
supercritical CO2 by changing the pressure or temperature according
to the relationship between solubility and density of supercritical
CO2. At the critical state, the components with different boiling points
and different molecular weights were selectively extracted. Ara et al.
studied the extraction of essential oils from Cornus officinalis by super-
critical CO2 extraction. It was proved that supercritical CO2 extraction
is a feasible method of extraction of essential oils from natural sub-
stances (Ara, Jowkarderis, & Raofie, 2015; Ismahane& Abdeslam-
Hassen, 2017).
CAI ET AL. 3

TABLE 1 Methods of extraction of essential oils

Method Advantages Disadvantages Application

Hydrodistillation Simple operation, low cost and
environmental protection
Cold pressing The process is simple and keeps the
original aroma of essential oil
Solvent extraction Low temperature to avoid damage of
heat sensitive components
Supercritical CO2 Nontoxic, pollution-free, low cost
extraction
Microwave assisted Fast and efficient, saving solvent
Biological enzymolysis The extraction speed is fast and the
technique efficiency is high
Ultrasound-assisted Save time and energy
The extraction efficiency is low
The purity of essential oil is low
There are many impurities in essential
oil
The operation temperature is high,
which is not suitable for the
extraction of heat sensitive
substances
Easy to degrade essential oil
Expensive, not easy to promote
There are many influencing factors
Rosemary essential oil (Sharifi-Rad
et al., 2020)
Citrus limon (Ferhat, Meklati, &
Chemat, 2010)
Rehmannia glutinosa Libosch oil;
essential oils of Origanum onites
(Ozel & Kaymaz, 2004; Xu, Liu, Hu,
& Cao, 2016; Zhao et al., 2016)
Algerian Argan essential oil (Haloui &
Meniai, 2017)
Cottonseed oil (Taghvaei &
Jafari, 2015)
Ginsenosides (Hsu, Lu, Chen, Wang, &
Hwang, 2013)
Lavandula angustifolia Caraway
essential oils, Polysaccharides from
Radix astragali (Li, Lu, & Zhao, 2013;
Wu, Ma, Kan, & Guo, 2014)

Ultrasonic-assisted extraction can break the plant cell wall by
ultrasonic wave energy to enhance their permeability to the solvent,
thus improving the extraction efficiency. This method has the advan-
tages of needing less solvent and being faster and is often used to
extract plant essential oils with poor thermal stability (Tavakolpour
et al., 2017).
Microwave-assisted extraction is a method to separate the
extracted substances from the system by heating selectively according
to the differences in microwave absorption ability of different sub-
stances (Razzaghi et al., 2019). The advantages of this method are that
it is less time-consuming, provides high resource utilization, and yields
a high aromatic content in the final plant essential oils (Lucchesi,
Smadja, Bradshaw, Louw, & Chemat, 2007).
1.3 | Biological activity of plant essential oils
Plant essential oils have multiple pharmacological effects, such as anti-
oxidant effects, action as analgesic, antipyretic, hypoglycemic,
antiulcer, anticonvulsant, anti-pathological aging, and anticancer
agents, and other potential therapeutic effects (Uma, Huang, &
Kumar, 2017). Plant essential oils play an important role in anti-
oxidation by acting directly on the diphenyl bitter acyl radical and
hydroxyl radical (Jayasena & Cheorun, 2014). Through olfactory path-
way, lavender and perilla essential oils can help to improve the immu-
nity of rats, and increase the number of white blood cell (WBC),
lymphocytes (LYM), and hemoglobin (HGB) in white rats, effectively
regulating the blood lipid metabolism of anxiety rats, balancing the
levels of serum total cholesterol, triglyceride, and glutamic acid, and
maintaining stable levels of high-density lipoprotein, and low-density
lipoprotein, cholesterols. Furthermore, lavender essential oil can
reduce systolic blood pressure, diastolic blood pressure, and mean
arterial pressure through the olfactory pathway (Hamzeh, Safari-
Faramani, & Khatony, 2020; Kazemi et al., 2020). Madder oil, garlic
essential oil, Atractylodes macrocephala oil, rosemary essential oil, and
volatile oil of phlegm fire have certain anticancer activity. Some plant
essential oils play the role of contact killing and repellent in the
growth and development of insects. Moreover, plant essential oils
show advantages of low molecular weight, simple composition, good
volatility, abundant sources, low environmental pollution load, and
low toxicity and are suitable for indoor pest control. For example, the
essential oils extracted from Cinnamomum osmophloeum of the family
Lauraceae exhibit good contact activity against three kinds of mosqui-
toes (Cheng et al., 2009). The essential oils extracted from Eucalyptus
globulus of Myrtaceae have contact killing activity against Aedes
albopictus larvae and repellent activity against adults of three kinds of
mosquitoes (Amer & Mehlhorn, 2006). Chegini was fumigated by thy-
mus oil on eggs, larvae, and adult-stage tomato spider mites. The mor-
tality of tomato spider mites increased with increasing concentration
of essential oils (Goudarzvande, Habib, Jaber, & Alireza, 2018). There
are also reports that the use of plant essential oils can inhibit the num-
ber of pests in beans: clove essential oil, cinnamon essential oil, and
Cinnamomum camphora essential oil can decrease the rate of growth
of cowpea nose worms (Viteri Jumbo et al., 2018). Agnus castus (Vitex
agnus castus) for premenstrual complaints, flaxseed (Linum
usitatissimum) for hypertension, feverfew (Tanacetum partenium) for
migraine prevention, ginger (Zingiber officinalis) for pregnancy-induced
nausea, ginseng (Panax ginseng) for improving fasting glucose levels as
well as phytoestrogens and St John's wort (Hypericum perforatum) for
the relief of some symptoms in menopause (Izzo, Hoon-Kim,
4 CAI ET AL.

Radhakrishnan, & Williamson, 2016) are studied. The main pharmaco-
logical activities of these plant essential oils are listed in Table 2.
Many essential oils and their volatile components have been
found to have strong antibacterial activity, which inhibit the growth of
some bacteria, fungi, and other microorganisms, and suppress the pro-
duction or accumulation of mycotoxins (Bluma, Amaiden, Daghero, &
Etcheverry, 2008; Moumni et al., 2020). The antibacterial activity of
essential oils is mainly caused by its complex active components
(mainly including alcohols, aldehydes, ketones, and phenols) (Li, Cai, &
Liu, 2019; Yang et al., 2011). Essential oils can use its own hydropho-
bicity to penetrate lipids, thus destroying the structure of the cell wall
and changing the permeability of the cell membrane: this leads to the
outflow of ions and matter within the cyst, causing cell death
(Cosentino et al., 1999; Dorman & Deans, 2010). Since Hoffmann and
Evans first proved that plant essential oils have antibacterial proper-
ties, many scholars have investigated the bactericidal effects of plant
essential oils. Through their research, essential oils are proven to have
certain inhibitory effects on fungi, bacteria, and viruses (Dwivedy,
Prakash, Chanotiya, Bisht, & Dubey, 2017). Carvacrol (35.2%),
followed by γ-terpinene (20.1%), was the main constituent of the
essential oils of oregano (Origanum vulgare), while cinnamaldehyde
(69.1%) represents the major component of the essential oils of cinna-
mon (Cinnamomum cassia). Active products extracted from the essen-
tial oils studied exhibited antibacterial activity against salmonella
strains. Essential oils from thyme, oregano, mint, cinnamon, sage, and
clove were found to have the strongest antibacterial properties (Chen
et al., 2020; Moumni et al., 2020). O. vulgare can inhibit the mycelial
growth of Enicillium verrucosum and Aspergillus westerdijkiae and the
production of ochratoxin A (Schlösser & Prange, 2019). Table 3 lists
the antibacterial properties of the main active components of several
plant essential oils and the microorganisms affected.
1.4 | Safety evaluation of plant essential oils
Plant essential oils are not only beneficial special commodity but can
also be toxic. The safety of plant essential oil is directly related to its
type, dosage, receptor, and action time. The acute toxicity of Artemisia
annua L. and Artemisia scoparia Waldst. on mouse, quail, bee,
trichogramma, ladybird, and red crucian carp by oral, percutaneous,
inhalation, and skin irritation is low. Different doses of Sabina essen-
tial oil can inhibit the rate of growth and food intake of animals. High-
dose Sabina essential oil has certain teratogenic toxicity and can result
in sperm aberration. Chronic poisoning symptoms may also occur
after long-term low-dose exposure to Sabina essential oil. Table 4 dis-
plays the safety evaluation of several plant essential oils and their
active components.
2 | THE MECHANISM OF ANTIFUNGAL
ACTION
The antibacterial activity of plant essential oils is an important indica-
tor of their biological activity. As a new type of antibacterial agent,
plant essential oil shows different antibacterial abilities because of its
complex composition and diverse mechanisms of action.
2.1 | Effects on fungal cell wall and cell membrane

TABLE 2 Main pharmacological activities of plant essential oils The cell wall and cell membrane play an essential role in maintaining
the normal life of cells. Plant essential oils can penetrate the cells of
Pharmacological
activity Essential oils Reference bacteria owing to their volatility and water solubility, thus affecting
the normal activities of cells (Dwivedy et al., 2017; Vijayasteltar, Nair,
Antibacterial Eucalyptus Cermelli, Fabio, Fabio, and
Quaglio (2008) Maliakel, Kuttan, & I, 2016). Phenolic compounds in plant essential

Antiviral Aloysia triphylla Silva, Figueiredo, Byler, and oils attack the cell wall or cell membrane of microorganisms, damaging
Royle Setzer (2020) the structure of cell wall and cell membrane, thus affecting the func-
Antiinflammatory Rosmarinus Borges, Ortiz, Pereira, Keita, tion thereof: this causes the contents of the cells to leak (carbohy-
officinalis and Carvalho (2019) drates, fatty acids, phospholipids, etc.), causing cell death. The
Regulating Oregano Rienth, Crovadore, Ghaffari, hydrophobic components in the essential oil can also change the per-
immunity and Lefort (2019) meability of the cell membrane and the outflow of cations such as H+
Antioxidant Lavandula Insawang et al. (2019) and K+, thus changing the pH value of cells and affecting the compo-
stoechas sition and activity of cells, causing the death of bacteria and inducing
Anticancer Arnica Montana Sugier, Sugier, Jakubowicz- bacteriostasis (Dwivedy et al., 2017). The main components of gera-
L. Achenes Gil, Winiarczyk, and
niol and citronella essential oils promote the movement of potassium
Kowalski (2019), Andrew
and Izzo (2017) ions from the cell to the outside of the cell and increase the fluidity of

Insect resistance Citrus Campolo et al. (2020) the fungal cell membrane (Bard, Albrecht, Gupta, Guynn, &
Stillwell, 1988); carvacrol can reduce the fluidity of the cell membrane
Anti-mutagenic Artemisia Taherkhani (2016)
ciniformis by changing the proportion and structure of the fatty acids in the cell

Anaesthesia Ocimum Khumpirapang, Pikulkaew, membrane, and then play an inhibitory role thereon (Di Pasqua,
basilicum Anuchapreeda, and Hoskins, Betts, & Mauriello, 2006). The antifungal macrocyclic
Okonogi (2018) dibenzyl (PLE) isolated from Herba Lysimachiae could inhibit chitin
CAI ET AL. 5

TABLE 3 Antibacterial properties of the main active components of several plant essential oils and the microorganisms affected

Active
Essential oils Scientific name ingredient Affected microorganisms References
Garlic essential oil Allium sativum L. Diallyl sulfide Enteropathogenic bacterial Plata-Rueda et al. (2017)
Capsicum essential Capsicum annuum L. Capsaicin Gram-positive and gram-negative Li et al. (2019)
oil bacteria
Cinnamon oil Cinnamomum cassia Presl Cinnamaldehyde Escherichia coli, Staphylococcus Lyu et al. (2018)
aureus, Listeria monocytogenes,
Salmonella enteritidis
Oregano essential Origanum vulgare L. Carvacrol, Gram-positive and gram-negative Lee, Kim, Beuchat, Kim, and
oil thymol Ryu (2020), Dogruyol, Mol, and
Cosansu (2020)
Rosemary essence Rosmarinus officinalis 1,8-cineole S. aureus, monocytogenes, Hussein, Lee, and Joo (2020),
Campylobacter jejuni Bertocchi et al. (2020)
Fennel essential oil Foeniculum vulgare Anethol Aeromonas hydrophila, Kwiatkowski et al. (2019)
Brevibacterium linens,
Brochothrix thermosphacta
Clove oil Syzygium aromaticum (L.) Eugenol E. coli, S. aureus, L. monocytogenes, Vurmaz and Gülten (2020)
Merr. Enteritidis, C. jejuni
Lemon essential oil Anethum grabeolens L. Limonene, S. Aureus, monocytogenes Le, P. (2020)
Carvacrol
Ginger oil Zingiber officinale Rosc. Zingiberene, Gram-positive and gram-negative Amalraj, Haponiuk, Thomas, and
Zingerone bacteria Gopi (2020)

TABLE 4 Evaluation on safety limit of PEOs and their components

1
Active ingredient Evaluation method Subjects LD50/(g kg ) Toxicity level Reference
Azadirachtin Oral acute Wistar rat 13.000 Practically non-toxic Rajashekar, Bakthavatsalam, and
Shivanandappa (2012)
Thymus vulgaris Oral acute Mice 17.500 Practically non-toxic El Euony, Elblehi, Abdel-Latif,
Abdel-Daim, and El-Sayed (2020)
Mentha haplocalyx Oral acute Mice 2.000 Low toxicity Zhang et al. (2015)
Eugenol Oral acute Mice 3.000 Low toxicity Zin et al. (2012), Cansian
et al. (2017), Abdelkhalek, Risha,
El-Adl, Salama, and
Dawood (2020), Vurmaz and
Gülten (2020)
Litsea cubeba Oral acute Wistar rat 4.000 Low toxicity Luo, Jiang, and Zou (2005)
Cinnamaldehyde Oral acute Wistar rat 2.220 Chahbi et al. (2020), Netopilova
et al. (2019), Kumar, Kumari, and
Mishra (2019)
Cymbopogon citratus Oral acute Mice 3.500 Low toxicity Costa et al. (2011)

synthesis of the Candida albicans cell wall both in vitro and in vivo.
The inhibitory effect of PLE on chitin synthesis in the cell wall of
C. albicans may be the main reason for its antifungal activity (Wu,
Cheng, Sun, & Lou, 2008; Singh et al., 2009).
2.2 | Effect on fungal lipid layer (nucleic acid and
protein)
Most of essential oils contain surfactants, which can change the per-
meability of cell membranes, allow extracellular substances to enter
the cell, and then destroy cell activity. Knobloch et al. (1986) found
that most of the active components of plant essential oils had inhibi-
tory effects on the aforementioned processes, verifying that plant
essential oils could have effects on the growth, development, and res-
piration of bacteria at a certain concentration. Bard et al. proved that
geraniol, an essential oil component, can decrease the temperature of
the lipid layer and affect the fluidity of the cell membrane (Bard
et al., 1988). Some plant essential oils can enter the cell and combine
with the bioactive substances therein, thus interfering with the normal
function of the cell and influencing the metabolism of the organism.
For example, cinnamaldehyde can damage the plasma membrane of
6 CAI ET AL.
cells, and then enter the cells to destroy the spatial structure of mac-
romolecular substances within, thus destroying the normal material
absorption, transport, and metabolism pathways (Yin et al., 2020). A
certain concentration of citric acid can change the permeability of the
cell wall, mitochondrial membrane, and plasma membrane of Aspergil-
lus flavus, and then inhibit the synthesis of DNA, RNA, lipids, and pro-
teins, resulting in the failure of metabolism, regulation, and other
functions necessary for normal life, that is, it causes irreparable and
lethal DNA damage to A. flavus. Luo et al. found that a concentration
of citral of 1.5 mg/L could cause lethal DNA damage to A. flavus. This
result revealed that citral first damages the cells by changing the
selective permeability of the cell wall, plasma membrane, and mito-
chondrial membrane of A. flavus. After entering the cell, the mycelium
is damaged. The inhibition of DNA, RNA, lipid, and protein biosynthe-
sis caused the related genes to become unable to be expresses, forc-
ing a loss of regulation and control of metabolism and normal self-
replication, ultimately leading to cell death (Luo et al., 2005).
2.3 | Effects on energy metabolism and reductase
of fungi
Mitochondria provide energy for various life activities in cells through
respiration. Each step in the process of respiration is inseparable from
the participation of enzymes. Plant essential oil can inhibit the growth
of mycelium and the production of toxins. It can be observed (using a
transmission electron microscope) that the essential oil of plant mainly
acts on the inner membrane system and affects mitochondria, thus
influencing respiration and causing cell death (Nogueira et al., 2010).
It has been reported that dill seed essential oil can inhibit the activity
of mitochondrial dehydrogenase, significantly reducing the synthesis
of ATP in cells, and interfering with the normal energy metabolism of
cells. In the hydrolysis reaction, ATPase pumps many hydrogen ions in
mitochondria into the gap of the mitochondrial membrane, which
leads to abnormal mitochondrial membrane potential and a large
amount of reactive oxygen species (ROS) accumulating in mitochon-
dria. ROS can cause A. flavus to die by oxidizing biological macromole-
cules or mediating apoptosis (Tian et al., 2012). Some essential oils,
such as lactate dehydrogenase, malate dehydrogenase, and succinate
dehydrogenase, affect the function of mitochondria by inhibiting the
activity of mitochondrial dehydrogenase. Echinacea can inhibit the
effects of the citric acid cycle and ATP synthesis in C. albicans mito-
chondria (Chen et al., 2013). Terpenoids, the main components of
essential oil, can significantly decrease the number of mitochondria in
cells, and then affect the amount of ROS and ATP (Haque, Irfan, &
Kamil, 2016).
2.4 | Effect on fungal gene expression
Gene determines protein, protein determines function. Mycotoxins
are produced by fungi, and the genes controlling mycotoxins are
closely related. Precocene II and piperitone decreased the mRNA
levels of Tri4, Tri5, Tri6, and Tri10 encoding proteins required for
deoxynivalenol biosynthesis. Methyl syringate is the main active com-
ponent of Betula platyphylla essential oil. Jermnak et al. found that the
rate of inhibition of afIR, pksA, and omtA gene transcription increased
with the concentration of methyl syringate. Yaguchi et al. found that
Precocene II and piperone could significantly inhibit the transcription
of tri4, tri5, tri6, and tri10 genes when the volume fraction of
piperone was 300 or 1,000 μl/ml (Dwivedy et al., 2017). Studies
showed that plant essential oils could change the conception of mem-
brane protein, destroy the integrity of cell membranes, increase the
fluidity and permeability thereof, and lead to the outflow of intracellu-
lar substances (Jin et al., 2019).
The antibacterial mechanism of plant essential oils is complex and
remains incomplete in terms of the understanding thereof. Further
research to provide a feasible method for degradation of mycotoxins
in food and feed, thus ensuring food safety, is warranted. This will
provide a scientific theoretical basis for the use of plant essential oils
as a natural antibacterial agent instead of chemical synthetic
antibacterial agents. It is believed that, with a scientific research foun-
dation and good production techniques, natural fungicides with plant
essential oils and active ingredients as raw materials will show great
development potential and broad application prospects. However, the
components of plant essential oil are highly variable. Therefore, it is
urgent to standardize the feed additives related to practical applica-
tion and product stability, and it is an ideal choice to synthesize or
extract specific active ingredients from natural materials. Therefore, in
the future, we should strengthen the research on the function and
coordination of specific active components of plant essential oil, clar-
ify its mechanism from the molecular and cellular point of view, and
provide theoretical basis for industrial application. The following table
shows the action mechanism of common plant essential oils (Table 5).
3 | DETOXIFICATION OF PLANT
E S S E N T I A L OI L S TO M Y C O T O X I N S
The research into the antibacterial activity of plant essential oils
mainly focused on the inhibition of bacteria and some mold growth
characteristics of plant essential oils and the accumulation of toxin-
producing mycotoxins. There are very few studies on the
TABLE 5 Action mechanism of several plant essential oils
Represents
Mechanism of action essential oil Reference
Destroy the integrity of Clove oil Xu et al. (2016)
cell wall and membrane
Affects bacteria and Cinnamon oil Yin et al. (2020)
protein synthesis
Interference with bacterial Dill seed Tian et al. (2012)
cell metabolism essential oil
Interference with bacterial Betula platyphylla Jermnak
cell metabolism essential oil et al. (2012)
CAI ET AL.
detoxification of mycotoxins by plant essential oils. Some plant essen-
tial oils contain specific components, which can degrade mycotoxins
produced by some fungi to a certain extent. The studies showed that
the essential oils of Aflamomum danielli could degrade ochratoxin OTA
when the concentration thereof was 1,000 or 2,000 mg/kg, and the
rates of degradation were 90% to 95%, and 100%, respectively
(Aroyeun, Adegoke, Varga, & Teren, 2009).
The abilities of essential oils and various extraction solvents (eth-
anol, 70% ethanol, and water extracts) of three medicinal plants (Thy-
mus daenensis, Satureja khozistanica, and Satureja macrosiphonia) were
investigated for their ability to reduce the content of Fumonisin B1
(AFB1). In general, ethanol and 70% ethanol extracts of all three
plants at 2000 mg/L and their essential oils at 125 mg/L inhibited
(51–87%) AFB1 production and had stronger anti-AFB1-biosyntesis
activity than aqueous extract (Tian et al., 2019). On the contrary,
essential oils, ethanol and 70% ethanol extracts of these plants could
not significantly degrade AFB1-contamination whereas their aqueous
extracts could significantly degrade AFB1. The aqueous extract of
Thymus daenensis at a concentration of 2000 mg/L resulted in reduc-
tion of 97% of the AFB1 present, while those extracted from
S. khozistanica and S. macrosiphonia reduced it by 6–15%. In general,
essential oils and ethanol extracts of both Thymus daenensis and
S. khozistanica largely inhibited A. flavus growth as well as
AFB1-biosynthesis while aqueous extracts of Thymus daenensis
showed strong AFB1-degradation activity and were deemed to be a
promising candidate for degradation of AFB1 (Gorran, Farzaneh, Shiv-
azad, Rezaeian, & Ghassempour, 2013). These results indicated that
the essential oils derived from the use of different extraction methods
had different mechanisms of degradation, so more methods should be
used to explore the best extraction methods with which to obtain the
essential oils necessary for the degradation of mycotoxins.
Sandosskumar et al. found that when the Allium cepa extract was
incubated with 70 ng/ml AFB1, the concentration of AFB1 decreased
(58.5%) after 5 days of incubation (Sandosskumar et al., 2007). The
seed extract of Ajowan (Trachyspermum ammi (L.) Sprague ex Turrill)
showed the maximum degradation of AFG1 at up to 65%. The dia-
lysed T. ammi extract was more effective than the crude extract, capa-
ble of degrading more than 90% of the toxin. Significant levels of
degradation of AFB1 (61%), AFB2 (54%), and AFG2 (46%) were
observed. A time-course study of AFG1 detoxification by dialysed
T. ammi extract showed that more than 78% of AFG1 was degraded
within 6 hr and 91% was degraded at 24 hr after incubation
(Velazhahan et al., 2010).
Perczak et al. assessed the ability of selected oils to degrade
deoxynivalenol (DON) under in vitro conditions, and found that
palmarosa and lemon oils provided effective reductions in the levels
of mycotoxins (Perczak, Ju
s, Gwiazdowska,
ska,
Marchwin
Wa
skiewicz, 2019). The higher reduction of DON content by these
oils reached 72% at pH 3 to 6 and a temperature of 20 C after inter-
action for 24 hr. A study showed that phenolic compounds from
plants could reduce FB1 production by Fusarium verticillioides, and
found that chlorophorin was the most effective compound in reducing
7
FB1 production (94% reduction), followed by caffeic acid, ferulic acid,
vanillic acid, and iroko, which reduced FB(1) levels by 90–91%
(Beekrum, Govinden, Padayachee, & Odhav, 2003). Cinnamon oil from
plants also could degrade FB1 and the rate of degradation of FB1 was
94.06% under optimal conditions (Xing et al., 2014). Further study
showed that cinnamon oil emulsion droplets could protect against
FB1 and/or AFB1 induced oxidative stress, cytotoxicity, and repro-
ductive toxicity in male rats (Abdel-Wahhab, El-Nekeety, Hassan,
Gibriel, & Abdel-Wahhab, 2018).
Plant essential oil can reduce mycotoxins, which provides a new
way of reducing their harm to animals and humans; however, some
problems remain. First, different extraction methods of essential oil
have different degradation effects on mycotoxins, so many more
experiments are needed to optimize the extraction process. In addi-
tion, the degradation mechanism of mycotoxins by plant essential oil
has not been studied, especially with regard to whether the degrada-
tion products remain toxic, or not.
4 | SUMMARY AND PROSPECTS
As a natural plant metabolite, essential oil mainly consists of terpenes,
oxygen-containing derivatives and aliphatic compounds, which have
antibacterial, antioxidant, anticancer, and other functions. With the
further study of plant essential oils, it will be developed in multiple
directions and be multi-disciplinary in the future. Plant essential oils
are a potentially useful natural product with low toxicity and an abun-
dance of sources. It will play a greater role in environmental protec-
tion, coatings, medicine, health products, feed, food, and other
aspects. It is important to prevent and treat the harm arising from
mycotoxins to human and animal health. With the application of most
chemical anti-corrosion and antibacterial agents becoming more
restricted, plant-derived fungicides with natural active antifungal
ingredients as raw materials will be favored by the market. Therefore,
the development and application of some plant essential oils in live-
stock and poultry production, show the antibacterial ability of plant
essential oils, the better to enhance the immunity of livestock and
poultry, improve the health of livestock and poultry, and improve the
production performance of livestock and poultry. At present, plant
essential oil has not been popularized for use in animal husbandry. In
the future, plant essential oil can be used to replace drugs in animal
husbandry, perhaps to improve animal growth performance, prevent
feed mildew, and improve animal intestinal health. The many applica-
tions of plant essential oils in animal production warrant further
investigation.
& AC KNOW LEDG EME NT S
This work was financially supported by the National Natural Science
Foundation of China (31972746, 31772809, 31872538), Liaoning
Provincial Natural Fund Guidance Program Project (2019-ZD-0708);
China Postdoctoral Science Foundation (2016T90477), PAPD, and
Project UHK VT2019-2021.
8 CAI ET AL.
CONF LICT OF IN TE RE ST
The authors declare no conflict of interest.
AUTHOR CONTRIBUTIONS
Jing Cai wrote the paper; Rong Yan, Jichao Shi and Jia Chen revised;
Miao Long, Wenda Wu and Kamil Kuca revised and supervised.
DATA AVAI LAB ILITY S TATEMENT
There are no data associated with this manuscript, as it is a review
article.
ORCID
Miao Long https://orcid.org/0000-0001-6683-1956
Kamil Kuca https://orcid.org/0000-0001-9664-1109
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How to cite this article: Cai, J., Yan, R., Shi, J., Chen, J., Long,
M., Wu, W., & Kuca, K. (2021). Antifungal and mycotoxin
detoxification ability of essential oils: A review. Phytotherapy
Research, 1–11. https://doi.org/10.1002/ptr.7281