Journal of Natural Pesticide Research 6 (2023) 100052

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Journal of Natural Pesticide Research

journal homepage: www.journals.elsevier.com/journal-of-natural-pesticide-research

Application of essential oils to control some important fungi and bacteria

pathogenic on cereals
Parissa Taheri *, Marjan Soweizy, Saeed Tarighi
Department of Plant Protection, Faculty of Agriculture, Ferdowsi University of Mashhad, Mashhad, Iran

A R T I C L E I N F O A B S T R A C T

Keywords: Among the poaceae family, cereals are known as the most important products because they are the main source
Antibacterial of energy and food. Different plant pathogens, including oomycetes, fungi, bacteria, nematodes and viruses, can
Antifungal infect cereal products. Fungal diseases are the most important group of cereal pathogens, causing significant crop
Cereal diseases
losses worldwide. Several techniques, including chemical control, the use of resistant cultivars, biological con­
Medicinal plants
Essential Oils
trol, and application of antimicrobial plant metabolites can be used to manage cereal diseases. The use of
Phytopathogen chemical products is limited due to undesirable environmental effects and the emergence of resistant pathogens
to fungicides. Due to the fact that medicinal plants produce a variety of secondary metabolites with antimicrobial
effects, they are considered as an alternative to chemical substances. In this study, we have discussed the
sensitivity of most important fungal and bacterial pathogens of cereal crops against different essential oils and
their main components, together with their modes of action in controlling cereal diseases.

1. Introduction
Plants are a large group of multicellular eukaryotes and are generally
attacked by various pathogens. Plant diseases have become a severe risk
that significantly reduce quality and quantity of agricultural products
(Pascale et al., 2020). In recent years, agricultural product safety has
become an important global issue, in which product problems caused by
pathogenic microorganisms have been mainly considered (Hou et al.,
2022; Benali et al., 2020). However, most of the chemical compounds
used in agriculture are still of interest due to their low price and effec­
tiveness against phytopathogens. Using chemical compounds for a long
time may lead to microbial resistance; on the other hand, it is a serious
threat to human health (Chanprapai and Chavasiri, 2017; Matusinsky
et al., 2015). Therefore, alternatives to chemical compounds have
become common due to the increasing demand for sustainable agricul­
ture and public concern about environmental pollution and human
health (Chidi et al., 2020). Nowadays, serious efforts are needed to find
new natural antimicrpbial compounds as alternative, safe, eco-friendly,
cheap and easily degradable compounds (Yang et al., 2010).
A range of medicinal plants contain active compounds, which can
inhibit microorganism’s growth (Shrivastava et al., 2011). The use of
natural antimicrobial compounds from plant origins, which are rich in
secondary metabolites, is of great interest because these products do not
pose any health or pollution risks. In addition, these natural products are
economically viable and have minimal side effects on the host (Sales
et al., 2016). Many plants drived essential oils and extracts show anti­
microbial effects against various plant pathogens.
The purpose of this article is to review the antimicrobial effects of
plant-derived essential oils, which are effective against the most
important bacterial and fungal diseases affecting cereals production,
worldwide.
2. Essential oils components and properties
The term "Essential Oil" (EO) was first introduced in sixteenth cen­
tury by Swiss medical reformer Paracelsus von Hohenheim. Plant based
oils are often complex combinations of non - polar and polar organic
products (Macwan et al., 2016, Guenther and Althausen, 1948). The EOs
are secondary metabolites that are found in a variety of plant parts,
including the roots, fruits, buds, flowers, twigs, seeds, leaves, woods,
and bark of aromatic medicinal plants. They are also produced in plas­
tids and cytoplasm of plant cells and stored in cavities, canals, epidermal
cells, secretory cells (Sumonrat et al., 2008; Bozin et al., 2006). The EOs
are mainly obtained from plants that belong to several families, such as
Rutaceae, Zingiberaceae Lauraceae, Myrtaceae, Lamiaceae and Aster­
aceae (Rasooli et al., 2005). The EOs are multi-component natural

* Corresponding author.
E-mail address: p-taheri@um.ac.ir (P. Taheri).

https://doi.org/10.1016/j.napere.2023.100052
Received 15 March 2023; Received in revised form 6 July 2023; Accepted 23 July 2023
Available online 24 July 2023
2773-0786/© 2023 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
P. Taheri et al. Journal of Natural Pesticide Research 6 (2023) 100052

Fig. 1. Different components of the essential oils obtained from various plant species.

compounds (Fig. 1), containing approximately 20–60 constituents of
different chemical structures and concentrations (Hammer et al., 1999).
The main components in the EOs are classified into different groups,
which are often responsible for their biological properties (Pichersky
et al., 2006). One of these groups includes terpene compounds that can
be tricyclic, bicyclic or monocyclic. The general formula of these com­
pounds is (C5H8)n, which is composed of isoprene units (Turek and
Stintzing, 2012). Other main groups include aldehydes, phenols,
alcohols and esters. Oxygenated compounds in the EOs are less abun­
dant than terpenes (Bakkali et al., 2008).
The synthesis of terpene hydrocarbons takes place in the cytoplasm
of plant cells. The most common terpenes include sesquiterpenes
(C15H24) and monoterpenes (C10H16), but hydrocarbons such as tri­
terpenes (C30H40) and diterpenes (C20H32), etc., which are terpenes
with longer chains, are also present in plant cells. The majority of ter­
penes lack strong inherent antimicrobial action (Caballero et al., 2003).

Fig. 2. Possible effects of essential oils on fungal cell components.

2
P. Taheri et al.
Fig. 3. Possible effects of essential oils on bacterial cell components.
The EO components may vary in quantity, quality and type of com­
pounds according to weather conditions, plant organ, age, vegetative
stage, soil composition, and plant interaction with various microor­
ganisms (Masotti et al., 2003). One of the most important properties of
the EOs that enables them to interact with lipids in the cell membrane of
microbes is their hydrophobicity. As a result, it makes them more porous
due to disruption of the cell structure. Due to the loss of ions and
essential chemicals from fungal and bacterial cell, this trait eventually
causes the microbial cell to die (Devi et al., 2010). The EO allows a plant
to control or regulate its environment (ecological role), thus acting as a
chemical signal to attract pollinators, inhibit seed germination,
communicate between plants, or repel predators (chemically signaling
the presence of herbivores). In addition, the EOs are outstanding anti­
microbials and may be present in all parts of aromatic plants (Sinkar
et al., 2021).
3. The EOs modes of action against bacteria and fungi
There are currently very few articles, which in detail demonstrated
the EOs modes of action against phytopathogens. The majority of EOs
interact with bacterial cell membrane processes such as ion gradients,
electron transport, protein translocation, phosphorylation, and other
enzyme-dependent events to exert their antimicrobial effects (Benchaar
et al., 2008). Multiple modes of action may be involved in the antibac­
terial activity of various EOs. By entering the cell wall and cytoplasmic
membrane, the EOs alter the structure of phospholipid layers, poly­
saccharides, and fatty acids (Akthar et al., 2014). Another function of
the EOs is to increase membrane permeability and cell membrane
leakage, also inhibit secretion of toxins (Dreger and Wielgus, 2013).
Gram-negative bacteria are less sensitive to EOs than Gram-positive
bacteria due to having an outer membrane rich in lipopolysaccharides
(Fig. 2) because the outer membrane limits direct contact between the
cytoplasmic membrane and the EOs (Raveau et al., 2020).
In the studies conducted by several researchers on the mechanisms of
action of EOs and plant extracts obtained using an organic solvent, six
different mechanisms have been mentioned about antifungal properties
of plant drived compounds (Fig. 3), as listed below (Tariq et al., 2019;
Akthar et al., 2014).
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Journal of Natural Pesticide Research 6 (2023) 100052
3.1. Affecting the fungal mitochondria
Some EOs disrupt the function of mitochondrial dehydrogenases and
electron transport chain, thus affecting mitochondrial activity in fungal
cells.
3.2. Inhibiting efflux pumps
H+ -ATPase controls fungal pathogenicity, cell proliferation, and
intracellular pH. Intracellular acidification and cell death result from
H+ -ATPase inhibition via application of the EOs.
3.3. Inhibiting cell wall formation in fungi
The shape and integrity of the fungal cells are destroyed by inhibiting
chitin and glucan biosynthesis of, and as a result, homeostasis is
disturbed, leading to cell death.
3.4. Inhibiting cell division
The EOs inhibit cell division and development of the mitotic spindle
by preventing microtubule polymerization.
3.5. Inhibition of RNA/DNA or protein synthesis
Inhibition of RNA and DNA synthesis followed by inhibition of
protein synthesis is known as an antifungal mechanism of various EOs.
3.6. Cell membrane disruption
Disturbance in ergosterol synthesis causes membrane destruction
and cell leakage, which destroys membrane functions such as protein
and enzyme activity, electron transfer, or absorption and transport of
nutrients (Fig. 3).
Almost all the EOs tested to date have shown some antimicrobial
activities. This activity varies from one plant to another and from one
pathogen to another. One of the EOs characteristics is hydrophobicity,
which is associated with antimicrobial effects of the EOs. It’s crucial to
P. Taheri et al.
understand that a damaged cell structure may have a cascading effect on
other cellular structures, increasing cell permeability and causing the
leakage of cell components (Faleiro, 2011). The kind and quantity of the
EO components affect its antibacterial action. The stage of plant devel­
opment, the plant part used for obtaining the EO, growth circumstances
(temperature, soil, fertilizers, interaction with microorganisms, etc.),
and climatic factors, as well as distillation and storage conditions, are
among the many other factors impacting composition of the EOs (Naz­
zaro et al., 2013; Faghih-Imani et al., 2020).
4. Effect of the EOs on mycotoxin production by fungi
Cereals are rich in nutrients that are used as human food and animal
feed all over the world. Cereal crops are infected in the field by a wide
range of fungal species due to the production of toxic secondary me­
tabolites (Wan et al., 2019). Generally, cereal pathogenic fungi are
divided into two main groups that attack seed crops. The first category is
referred to as "field fungi," and for their development, the substrate must
have a higher moisture content and be at a lower temperature (e.g.,
Fusarium spp, Alternaria spp. Cladosporium), and the second group
includes“storage” fungi (e.g., Penicillium; Aspergillus spp.), which
multiply during storage (Ismaiel and Papenbrock, 2015). Mycotoxins
are the toxins produced by fungi that cause disease or death in humans,
animals and reduced performance when ingested, inhaled or absorbed
through the skin. These are substances with a small molecular weight
(Stroka, 2011).
It is challenging to eliminate mycotoxins production in food because
they are resistant to heat and different food processing techniques (Pitt
and Hocking, 2009). Mycotoxins interact with nucleic acids at the
cellular level and prevent biosynthesis of the vital macromolecules DNA
and RNA, affect the structure and operation of biological membranes, or
interfere with metabolism of energy. Mycotoxins typically weaken the
immunity system in human and animals by preventing protein synthesis
and cell growth (Luo et al., 2018). Therefore, the quality and quantity of
agricultural products are significantly reduced due to contamination
with fungal pathogens and toxins that they produce, and it will result in
economic losses before and after harvest. To minimize the losses caused
by fungi in agricultural products, various management/ control strate­
gies are used (Shuping and Eloff, 2017). Phytochemicals are the most
promising natural compounds for production of novel and effective
eco-friendly fungicides (Anjorin et al., 2013) to be used in plant disease
management instead of hazardous synthetic agrochemicals.
Effectiveness of the EOs in preventing fungal contamination of ce­
reals has been examined and demonstrated in several studies. Soliman
and Badeaa (2002) found that toxigenic fungi are sensitive to several
EOs tested, and particularly sensitive to cinnamon and thyme EOs. The
results also showed that the EOs of anise, cinnamon, spearmint and
thyme have more effect indecreasing fungal growth and mycotoxins
production in wheat grains. At 5% (v/v) concentration, sweet basil EO
totally inhibited mycelial growth and prevented aflatoxin production,
showing that the oil’s minimum inhibitory concentration (MIC) is less
than or equivalent to 5% (Atanda et al., 2007). Numerous studies have
revealed that Aspergillus growth and mycotoxin production were
inhibited by many plant EOs (Jayashree and Subramanyam, 1999:
Paranagama et al., 2003: Rasooli et al., 2005; Shukla et al., 2009; Helal
et al., 2007). Effectiveness of the EOs in inhibiting synthesis of aflatoxin
B1 may be attributed to their capability to prevent A. flavus from
catabolizing carbohydrates by interfering with some important enzymes
(Dwivedy et al., 2016).
According to Perczak et al. (2016), under specific environmental
circumstances, the synthesis of zearalenone (ZEA) by Fusarium spp.
could be inhibited by the EOs of lemon, grapefruit, eucalyptus, and
palmarosa. Another study demonstrated that the use of Curcuma longa
EO could totally stop the growth of Fusarium graminearum and ZEA
production (K.N. Kumar et al., 2016; P. Kumar et al., 2016). Antifungal
and antimycotoxigenic properties of Fusarium spp. decreased by using
4
Journal of Natural Pesticide Research 6 (2023) 100052
the EOs of Cinnamomum zeylanicum, Citrus paradisi, Citrus limonum,
Eucalyptus radiata, Citrus grandis, and Cymbopogon martini (Perczak et al.,
2019). Also, several studies have shown that the EOs stop mycotoxin
production by strong fungicidal activity via affecting metabolic genes
and have (Divband, Shokri, and Khosravi, 2017; Kalagatur et al., 2015).
Plant products can be promising alternatives for degradation or inhibi­
tion of mycotoxins biosynthesis. Therefore, plant products as natural
compounds may be used in agriculture to control mycotoxins contami­
nation in food and feed products.
5. Effect of the EOs on quorum sensing
Antibiotics are often used to treat bacterial diseases. Organisms have
developed resistance as a result of misuse and overuse of antibiotics
(Kowalczyk et al., 2020; Zhuang et al., 2020). A worldwide hazard to
agricultural products has been created by the pathogenic bacteria’s
unparalleled rise in antibiotic resistance. With increasing resistance
against the available antibiotics, researchers are looking for alternative
methods to treat and prevent bacterial diseases (Zhuang et al., 2020;
Tiwary et al., 2017). Quorum sensing (QS) is among these new objec­
tives. Bacteria use the QS phenomenon, which is essential for many
pathogenic bacteria to produce virulence genes like Pyocyanin, Algi­
nate, Rhamnolipid, Pyoverdin, and the ability to form biofilms. The QS
has emerged as a crucial therapeutic target (Kalia and Purohit 2011),
which could potentially aid in reducing bacterial resistance and
combating infections (Mokhetho et al., 2018; Ahmad et al., 2015;
Elgaml and Miyoshi, 2017). A vast array of behaviors are controlled by
the communication and gene-regulatory mechanism known as QS such
as swarming, sporulation, the development of biofilms, conjugation of
antibiotic synthesis, and pathogenicity (Tariq et al., 2019).
In filamentous fungi, the QS mechanisms have only lately been
described. Aspergillus and Penicillium species appear to use signal
molecules to coordinate operations. Secondary metabolism in these
fungl is induced when the population density rises. Other microbes have
previously defined this occurrence as a competing tactic, attempting to
obtain nutrients and occupy niches (Padder et al., 2018). The QS in fungi
is mediated by accumulation of tiny diffusible molecules in the extra­
cellular environment. There is a great variety of signaling molecules
identified in fungi, and these compounds are typically not strain-specific
(Barriuso, 2015). A major development in QS phenomenon of eukary­
otes was identification of the quorum sensing molecule (QSM) farnesol
in the pathogenic fungus Candida albicans. The existence of fungal QS
systems has indicated that acetaldehydes, tyrosol, farnesol, tryptophol,
and several volatile substances, in addition to lipids (oxylipins), peptides
(pheromones), alcohols (tyrosol, farnesol, and 1-phenylethanol), are
actively involved in fungal QS (Albuquerque and Casadevall, 2012:
Padder et al., 2018). It is now well accepted that this type of microbial
communication is not limited to isolates of the same species or even
taxon. Multiple types of interaction between bacteria, bacteria and
fungi, and even bacteria or fungi with their plant (Geiger et al., 2004) or
mammalian (Cámara et al., 2002) hosts have been demonstrated to be
mediated via involvement of QSMs (Geiger et al., 2004: Cámara et al.,
2002: Padder et al., 2018). Therefore, the QS is a communication system
by using small diffusible signaling molecules known as autoinducers or
QSMs. One of the most suitable alternative methods for controlling
bacteril and fungal pathogens, which can be effective on the QSMs, in­
cludes application of the EOs obtained from various plant species (Ta
and Arnason, 2015).
Examples include Zingiber officinale (Kim et al., 2015), Curcuma longa
(turmeric) (Virk et al., 2019), Allium sativum (garlic) (Vadekeetil et al.,
2016) and Terminalia chebula (Mihalik et al., 2008) which have been
extensively studied as potent inhibitors of QS and QS-derived virulence
factors in P. aeruginosa. Yap et al. (2015) reported varying level of QS
inhibition in Escherichia coli by Trans-cinnamaldehyde, benzyl alcohol
and eugenol, which are the major components of the EOs of Cinnamo­
mum verum. Several studies have reported the anti-QS activities of pure
P. Taheri et al.
EO compounds (Cáceres et al., 2020; Bhargava, and Zhang et al., 2015,
2018). Consequently, the development of innovative anti-infective
techniques that do not rely on the use of antibiotics has found QS to
be an appealing target.
6. Review of controlling ten fungal and fungus- like pathogens
on cereals by the EOs
In this review, we discussed the antimicrobial effects of plant EOs
against some of the most harmful diseases, which affect cereal crops.
The top ten cereal fungal and fungus-like pathogens list includes, (1)
Fusarium graminearum (2) Magnaporthe oryzae (Rice blast), (3) Rhizoc­
tonia solani (4) Gaumannomyces graminis (Wheat Take-all), (5) Bipolaris
sorokiniana (6) Pythium spp, (7) Puccinia spp., (8) Fusarium verticillioides
and storage fungi which include species of (9) Aspergillus spp. and (10)
Penicillium spp. (Newitt et al., 2019; Sharma et al., 2021). Effect of
various EOs on these destructive phytopathogens are discussed below.
6.1. Fusarium graminearum
Fusarium graminearum is the most destructive pathogen of cereal
crops, which causes Fusarium head blight (FHB), Fusarium seedling
blight (FSB), root and crow rot diseases (Taheri, 2018). Antifungal ef­
fects of the EOs on F. graminearum have been reported by several re­
searchers. In this regard, lemongrass, cinnamon, oregano, clove and
palmarosa and their major components showed complete growth inhi­
bition of F. graminearum and the EOs had a statistically significant effect
on mycotoxin production by F. graminearum as reported by Velluti et al.
(2004). According to Ferreira et al. (2018), the EO of Zingiber officinale
reduced biosynthesis of the mycotoxin dioxynivalenol (DON) and
ergosterol at concentrations of 500 and 1000 g/ml, respectively. Ac­
cording to K.N. Kumar et al. (2016); P. Kumar et al. (2016), the EO of
Curcuma longa totally suppressed formation of zearalenone (ZEA) and
reduced fungal biomass at concentrations of 3500 and 3000 mg/ml,
respectively. Another important biological activity of C. longa EO is the
accumulation of free radicals in C. longa treated mycelium, which may
be responsible for the apoptotic cell death and altered toxin metabolic
pathway gene regulation, leading to decrease in mycotoxin production.
The mechanism of C. longa action in vivo was also directly correlated
with the presence of phenolic and other chemical compounds, which are
involved in the stress induction, leading to the altered metabolic
pathway gene regulation, decreased toxin production and fungal
growth.
The main component of the EO of Anacyclus valentinus was observed
to be − 3-carene (31%), followed by − 3-carene (31%), spathulenol
(14.2%), decanoic acid (5.5%), -cadinene (4.4%), anethole (3.4%), and
aromadendrene (3.3%). According to Houicher et al. (2018), these
compounds were responsible for antifungal activity of the EO with the
MIC values of 1.25–2.5 µl/ml against Aspergillus parasiticus, Aspergillus
flavus, Penicillium expansum, Penicillium citrinum, Aspergillus ochraceus,
Fusarium moniliforme, Fusarium graminearum. In the studies by Davari
and Ezazi (2017), the EO of Eucalyptus sp. revealed concentration of
44.08% of 1, 8-cineole. The authors found an antifungal effect of this
compound against Botrytis cinerea, Fusarium asiaticum, Fusarium oxy­
sporum, Aspergillus flavus, Fusarium verticillioides, F. gramineraum at a
concentration of 1000 µl/L with inhibition of 33–75%. Numerous plant
EOs and their major components have demonstrated antifungal efficacy
against several plant pathogenic fungus. Isman and Machial (2006) re­
ported that although their action mechanisms are unclear, they can be
connected to their ability to disintegrate or damage the integrity of the
fungal cell walls and membranes.
Kalagatur et al. (2015) reported that the EO of Ocimum sanctum could
inhibit F. graminearum growth and ZEA production with minimum
inhibitory concentration and minimum fungicidal concentration of 1250
and 1800 μg/ml, respectively. This study showed that O. sanctum is
effective via multiple ways in preventing F. graminearum from producing
5
Journal of Natural Pesticide Research 6 (2023) 100052
ZEA. With increased O. sanctum concentration, the mycelial biomass,
gene expression (PKS4 and PKS13), and ZEA production all showed a
sharp decline.
Harčárová et al. (2021) reported that the EO of Origanum vulgare
exhibited strong antifungal activity against F. graminearum at 100 µg/ml
concentration. Another study found that the EOs of Satureja hortensis and
Zataria multiflora were effective in reducing the synthesis of DON as well
as suppressing growth of F. graminearum (Lahooji et al., 2010). Anti­
fungal activity was observed using the EOs of a number of plant species,
including Eriocephallus panculatus, Artemesia afra, Conyza scabrida,
Mentha peripeta and Pelargonium graveolons against Fusarium spp. isolates
with 100% growth inhibition. Among various fungi tested, F. grami­
nearum was the most susceptible to the EOs tested in a liquid media with
the lowest MIC (0.24 mg/ml). This low MIC was obtained with the EOs
of A. afra, P. graveolens and E. punculatu (Amidou and Tshifhiwa, 2012).
More in vitro and in vivo studies seems to be required to further describe
the active ingredients and determine exact modes of action of these EOs
and their major ingredients against various fungal pathogens.
6.2. Fusarium verticillioides
Fusarium verticillioides (teleomorph: Gibberella moniliformis) is a fila­
mentous fungus and is one of the most important fungal pathogens of
cereals. The species F. verticillioides is one of the main producers of
mycotoxins resulting in low yield and grain quality as a consequence of
Fusarium ear rot (FER) disease, which damages the plant in all stages of
growth. Studies have been conducted to investigate antimicrobial effect
of various EOs against F. verticillioides (Achar and Sreenivasa, 2021; Tran
et al., 2021). Yamamoto et al. (2013) reported the antifungal activity of
ginger EO (GEO; Zingiber officinale Roscoe) and the minimum inhibitory
concentration (MIC) against F. verticillioides by micro-broth dilution.
Different concentrations (500–5000 µg/ml) of the EO were used. It was
found that antifungal activity increased by increasing the concentration
or volume of the EO. The GEO exhibited antifungal activity, with the
MIC of 2500 μg/ml against F. verticillioids.
Hydrophobic property of the EOs, which results in the breakdown of
membrane integrity and release of cellular substances, is responsible for
this activity (Yamamoto et al. 2013). At 4000 and 5000 g/ml, the GEO
prevented F. verticillioides from producing ergosterol. A sterol called
ergosterol influences the hormonal and structural functions of yeast
cells. It controls the movement and activation of several membrane
bound enzymes (Bendaha et al., 2011). Some antifungal substances
prevent cell development by preventing the manufacture of ergosterol,
which happens when antifungals bind to ergosterol on cellular mem­
branes. This results in osmotic disturbances, the development and
multiplication of fungal cells, and an alteration in the integrity and
functionality of several membrane bound proteins (Yamamoto et al.
2013; Bendaha et al., 2011).
Starovic et al. (2016) demonstrated antifungal activity of the EOs
obtained from medicinal plants including myrtle (Myrtus communis L.),
rosemary (Rosmarinus officinalis L.), sage (Salvia fruticosa L.), bitter
fennel (Foeniculum vulgare spp. piperituum L.), anise (Pimpinella anisum
L.), and mint (Mentha spicata L.) against some important pathogenic
fungi, such as, Fusarium subglutinans, Bipolaris/ Drechslera sorociniana, F.
tricinctum, F. vertricilioides, F. sporotrichioides, F. oxysporum, F. incarna­
tum, F. equiseti, Macrophomina phaseolina and F. proliferatum based on the
MIC value. The EOs of all tested plants showed considerable antifungal
activity against all pathogens investigated. The myrtle EO exhibited
maximum antifungal activity with the MIC values of
0.0003–3.25 mg/ml, then mint at 0.0003–7.75 mg/ml and sage at
0.0003–10 mg/ml.
Da Silva Bomfim et al. (2015) reported the GC-MS analysis of Ros­
marinus officinalis EO isolated from aerial plant parts by hydro­
distillation. The main components of this EO were α -pinene (12.4%),
camphor (15.2%) and 1.8 cineole (52.2%). This EO inhibited the growth
of F. verticillioides with the MIC and MFC values of 150 LG/ml. The
P. Taheri et al.
results of this study demonstrated that R. officinalis EO has potent
antifungal activity against F. verticillioides. This was demonstrated by
changes that R. officinalis EO caused in the weight of the mycelium, the
size of fungal structures, the morphology of microconidia, and its
inhibitory effects on the production of ergosterol and fumonisin. R.
officinalis EO may have fungitoxic effects on F. verticillioides due to cell
wall rupture and loss of cellular components. As a result, this EO can be
utilized as a substitute natural fungicide that poses fewer hazards to both
human and the environment (da Silva Bomfim et al., 2015).
In another study, Xing et al. (2014), tested the antifungal activity of
eight EOs against F. verticillioides and cinnamon EO showed the best
inhibitory effect. The main compounds in this EO included synthetic
cinnamaldehyde (99%), natural cinnamaldehyde (95%), and cinna­
maldehyde (85%) with minimal inhibitory concentrations (MICs) of 45,
60 and 50 μL/L against F. verticillioides. Additionally, it has been noted
that the crucial enzymes in yeasts and molds that produce cell walls, β -
(1, 3) - glucan synthase and chitin synthase 1, are inhibited by cinna­
maldehyde. The EO lipophilic qualities, which make it permeable to the
cell wall and cytoplasmic membrane and aid in the formation of poly­
saccharides under water stress circumstances, may be responsible for
these changes in the cell wall. As a result, the chance for cinnamon oil to
interact with the cell wall increases, the enzymatic processes that con­
trol creation of cell wall are hindered, and the cell integrity is compro­
mised (Xing et al., 2014).
Avanço et al. (2017) reported the antifungal property of Curcuma
longa EO along with its three major compounds, α-turmerone (42.6%),
β-turmerone (16.0%) and arturmerone (12.9%) against F. verticillioides.
It was found that C. longa EO inhibited the mycelial growth with MIC
and MFC of 73.7 μg/ml. On F. verticillioides, C. longa EO was found to
have antioxidant and antimycotoxigenic properties. The oil may reduce
processes such as lipid peroxidation and others caused by generation of
free radicals (Avanço et al., 2017).
The effects of fourteen EOs were investigated for antifungal activity
by measuring the IC50 (half maximal inhibitory concentration) against
F. verticillioides. The results revealed that the EOs of Origanum x appli,
Origanum vulgare., Origanum vulgare, Chenopodium ambrosoides and
Tagetes riojana were the most active EOs with the IC50 values of
66.79uL/L, 101.71uL/L and 108.27uL/L, respectively, followed by the
EOs of C. ambrosioides and T. riojana, with the IC50 values of 243.12 μL/
L and 761.75 μL/L, respectively (Pizzolitto et al., 2020). Numerous
studies have linked the key chemical constituents of the EOs phenols
> alcohols > aldehydes > ketones > esters > hydrocarbons to their
antifungal effects, and they have hypothesized that there is a correlation
between antifungal activity and these constituents chemical structures
(Andrade et al., 2019). The majority of the active EOs studied (Origanum
spp.) indicated a high amount of phenolic compounds, whereas the less
bioactive EOs revealed a high hydrocarbon monoterpene content. In
connection with this, phenolic chemicals’ antifungal activity has been
linked to a change in cell membrane permeability, which causes the loss
of cellular components and suppression of cell metabolisms (Andrade
et al., 2019; Pizzolitto et al., 2020).
Twenty five samples of the EOs from medicinal plants were studied
for their antifungal activity against various fungi, such as,
F. verticillioides, F oxysporum, Penicillium brevicompactum, Aspergillus
flavus, Penicillium expansum, and Aspergillus fumigates by Zabka et al.
(2009). Almost all 25 EOs tested had a positive effect on the pathogen
growth. The EO of Pimenta dioica was determined as the most effective
EO against F. verticillioides. Growth of F. verticillioides was strongly
inhibited at the MIC value of 0.6 µlml− 1.
Davari and Ezazi (2017) reported significant inhibition of
F. asiaticum, F. verticillioides, F. graminearum, F. oxysporum f.sp. lentis,
F. redo-lens f.sp. dianthus, A. flavus, Aspergillus tubingensis, Sclerotinia
sclerotiorum, Cladosporium cladosporioides, and Botrytis cinerea by the EO
obtained from Zhumeria majdae. Camphor and linalool are two mono­
terpenes that are abundant in Z. majdae EO. Effectiveness of the EOs
depends on a number of variables, including chemical structure, active
6
Journal of Natural Pesticide Research 6 (2023) 100052
biological ingredient, concentration of the EOs, and type of microor­
ganisms (Davari and Ezazi, 2017).
6.3. Magnaporthe oryzae
Magnaporthe oryzae (anamorph Pyricularia oryzae), which causes rice
blast, is the most economically important fungal pathogen on rice, world
wide. Application of natural antifungal compounds can be proposed as
an environmentally friendly method and alternative to chemical fungi­
cides for management of rice blast disease (Kumar and Ashraf, 2019).
Wang et al. (2010) reported antifungal activity of the EO obtained from
root bark of Periploca sepium against on M. oryzae with the MIC value of
300 µg/ml. The MIC value has been defined as the lowest EO concen­
tration inhibiting spore germination and mycelium growth. Aguiar et al.
(2014) found the compound citronellal in the composition of the EO of
Corymbia citriodora and Cymbopogon nardus with MIC values of 61.78%
and 36.6%, respectively and verified its antifungal activity. The EOs of
both C. nardus and C. citriodora had considerable impact on decreasing
the development of P. grisea, with MIC values of 100 ppm and 200 ppm,
respectively.
Scientists have found that P. grisea’s growth, spore formation, and
germination were all significantly inhibited by the EOs of C. citriodora
and C. nardus. Sesquiterpene hydrocarbons, phenolic chemicals, and
oxygenated monoterpenes may all contribute to the antifungal action of
the EOs. Therefore, it is anticipated that the EO treatment that sup­
presses spore formation in vitro, might significantly reduce spore pro­
duction in the storage period, thereby helping to prevent spread of the
infection. In general, morphological alterations in the cell wall and
interference in enzymatic activities of wall production, which impair
fungal growth and morphogenesis, are responsible for the antifungal
activity of C. nardus, C. citriodora, and citronellal against P. grisea.
Therefore, the fungal enzyme systems are compromised or ion perme­
ability increases, allowing important intracellular components to seep
out. Huang and associates, (2010) reported that the EO of fresh fruit of
Illicium verum was effective against rice blast disease and reduced dis­
ease severity. The IC50 value of the oil against spore germination of P.
oryzae was 0.32 mg/ml. High concentration of trans-anethole in some
EOs, which has been identified as the primary active ingredient among
the oil volatile components, is what’s responsible for its antifungal
properties. Sukanya (2011) reported that Coriander sativum, Curcuma
domestica and Piper nigrum EOs exhibited strong antifungal activity
against P. oryzae. Application of Neem (Azadirachta indica) oil and neem
oil plus neem leaf extract reduced blast disease on the Pusa Basmati 1121
rice variety by about 25%, according to a study by Kumar et al. (2017).
Olufolaji et al. (2015) discovered that the leaves of Ocimum gratissimum,
Chromolaena odorata and Cymbopogon citratus, the nuts of Garcinia kola,
the seeds of Eugenia aromatica and Piper guineense had antifungal prop­
erties against P. oryzae. Recently, Patil et al. (2020) tested several EOs of
different plants against P. oryzae and found that Nirgundi oil signifi­
cantly reduced the disease severity compared to the control. Therefore,
these EOs or their main antifungal ingredients might be used as natural
fungicides and plant protectants to control rice blast as a highly
destructive disease, worldwide.
6.4. Rhizoctonia solani
In many economically important crops, the fungus Rhizoctonia solani
(teleomorph: Thanatephorus cucumeris) causes root rot, crown rot,
damping off, and leaf blight. To find new and safe type of control agents
against R. solani, antifungal activities of many plant EOs have been
investigated by numerous researchers, so far (AYDIN and Filiz, 2021).
Nafiseh et al. (2011) demonstrated the antifungal activity of Eucalyptus
EO against several plant pathogenic fungi, such as, Aspergillus flavus,
Penicillium digitatum, Pythium ultimum, Colletotrichum gloeosporioides,
Bipolaris sorokiniana and R. solani. Eucalyptus EO was effective against
all tested fungi and completely inhibited mycelial growth of the two
P. Taheri et al.
most important soilborne pathogens, P. ultimum and R. solani. However,
further research is needed to assess the EOs potential for use on plants
and the quality of fruits and vegetables that have been treated in order to
develop them as synthetic fungicide alternatives. Marei et al. (2012)
investigated antifungal effects of several oxygenated monoterpenes of
the EOs against four plant pathogenic fungi, including R. solani. These
compounds included camphor, carvone, 1, 8-cineole, camphene, fen­
chone, cuminaldehyde, geraniol, limonene, menthol, linalool, myrcene,
and thymol. They discovered that a few monoterpenes had strong
inhibitory effects on the majority of the tested fungi. Three mono­
terpenes, including 1, 8-cineole, (S)-limonene and thymol are potent
antifungal compounds that can be used to control plant pathogenic
fungi. Uncertainty surrounds the processes through which mono­
terpenes function as antifungals. But as lipophilic agents, they carry out
their activity at the level of membrane and the enzymes that are
embedded in the membrane, according to various studies. Therefore,
these substances exert their antifungal effects by altering the fatty acid
content of fungal cell membran (Marei et al., 2012).
Additionally, Cakir et al. (2004) discovered that the growth of fungi,
such as R. solani and Fusarium spp., were completely inhibited at
1 g/mlconcentrations of terpineol and caryophyllene oxide as the main
components of the EOs obtained from Hypericum species. Seema and
Devaki (2010) reported that the EOs of cinnamon (Cinnamomum zeyla­
nicum), clove (Syzygium aromaticum), coriander (Coriandrum sativum),
fennel (Foeniculum vulgare) and nutmeg (Myristica fragrans) have strong
inhibitory effects against R. solani with MIC values of 500, 1000, 1000,
2000 and 2000 ppm, respectively. The effectiveness of the EOs utilized
in this investigation could result from the interaction of several bio­
molecules that are found in them. Five EOs have demonstrated potential
efficacy against R. solani. The outcomes demonstrated the antifungal
effects of these five EOs on both mycelial development and sclerotia
formation (Seema et al., 2010).
Antifungal activities of Myrtus communis and Vitex agnus-castus EOs
have been determined against R. solani (Yilar et al., 2016). Mycelial
growth of R. solani was suppressed by 10 ml of Vitex agnus-castus (VAC)
EO in 65.35% and by M. communis EO in 100%. Sixteen components
were identified in the EO of M. communis, and among the main ones,
Eucalyptol (49.15%), Myrtenol (19.49%) and α-Pinene (8.38%) stood
out. The EO of M. communis was tested for its antifungal activity against
R. solani, reaching MIC and LC50 values of 10 and 1.072 μL/ Petri,
respectively.
Nasseri et al. (2015) demonstrated antifungal effect of the EO ob­
tained from Zatania multiflora against R. solani, with the MIC of
200 ppm. This result affirms the effectiveness of the EO as an inhibitor of
the phytopathogenic fungi. Udomsilp et al. (2009) reported antifungal
activities of the EO obtained from two plant species, including, Boswellia
carteri and Acacia farnesiana. The inhibition rate of A. farnesiana EO
against rice pathogenic fungi, such as R. solani was 100% at 2% (v/v)
concentration. All types of pathogenic fungi were susceptible to B. carteri
oil in terms of mycelial growth and spore germination inhibition. In
2017, Siddiqui et al. reported that phytochemicals (b-caryophyllene,
D-cadinene, a-cubebene, 1, 2-benzen carboxylic acid and caryophyllene
oxide) of Mikania scandens EO could be used instead of systemic chem­
ical compounds against R. solani. Their results showed that b-car­
yophyllene, D-cadinene, a-cubebene, 1, 2-benzen carboxylic acid and
caryophyllene oxide exhibit suitable antifungal activities against Pytium
graminicola, Fusarium oxysporum, R. solani and Tricoderma harzianum
with MIC values of 125, 125, 250 and 250 µg/ml, respectively. The
antifungal action of bioactive components, such as eugenol, carvacrol,
cinnamaldehyde and thymol which are the main components of several
EOs, were analyzed and it has been demonstrated that carvacrol and
thymol had inhibitory action on R. solani at 100 mg/ml concentration.
Therefore, these natural compounds might be used for plant protection
against destructive fungal phytopathogens. Carvacrol and thymol were
categorized by the fungicide resistance action committee (FRAC) to their
mode of action (transport or synthesis of lipids / function or membrane
7
Journal of Natural Pesticide Research 6 (2023) 100052
integrity). This group covers the active compounds that act on the
permeability of cell membranes and impact glycolipid formation to
change production of lipids and the integrity of cell membranes.
Therefore, they interfere with fungal cell membrane function (Semper­
e-Ferre et al., 2021).
6.5. Gaeumannomyces graminis
Take-all, caused by the fungal pathogen Gaeumannomyces graminis
var. tritici, is another essential wheat disease, worldwide. The typical
symptom is black necrosis on the roots and destruction of root hairs.
Control of this disease in the seedling stage is done using chemical
fungicides. Therefore, natural agents are the best alternatives for long-
term control of this disease (M. Wang et al., 2018; Y. Wang et al.,
2018; Yang et al., 2017). In the search for natural fungicides for pro­
tecting cereal crops, effect of the EO obtained from Drimys winteri was
investigated by a number of researchers against the soil borne fungus
G. graminis. The plant genus Drimys is native to Argentina and chile
(Muñoz et al., 2011). This evergreen tree contains drimane- type ses­
quiterpenoids. The EOs of this plant and their major constituents are
toxic against a wide range of pathogens including, fungi and bacteria
(Paz Robles et al., 2014; Arias et al., 2018). Monsálvez et al. (2010)
suggested that oxygenated monoterpenes, such as α-pinene, β-pinene,
limonene and β-myrcene could be responsible for the antifungal efficacy
of D. winteri EO at 250 mg kg− 1 concentration. In vivo experiment with
n-hexane fraction from the D. winteri bark in growth substrate inoculated
with G. graminis allowed a significant increase in plant height, accu­
mulated biomass, chlorophyll content, and stomatal conductance of the
wheat plants as well as a significant decrease in the severity of the
damage caused by take-all disease. According to the findings, this
fraction protects wheat seedling development at concentrations between
250 and 500 mg kg− 1. D. winteri showed strong inhibition against G.
graminis. The most active responses were obtained from the n-hexane
fraction, followed by the acetone fraction, with IC50s of 198 and
234 mg/L, respectively (Zapata et al., 2011). According to findings of
this investigation, it is important to further explore substances with
antimicrobial activity and to ascertain their efficacy both in vivo and in
the field. Also, further research must be done on potential formulation
and application techniques.
Paz et al. (2020) showed that the EO of D. winteri could inhibit the
growth of G. graminis var. tritici. These researchers showed that the main
constituents of the D. winteri oil were valdiviolide (12 μg ml− 1), iso­
drimeninol (10 μg ml− 1), polygodial (7 μg ml− 1), drimenol
(150 μg ml− 1), drimenin (300 μg ml− 1) and drimendiol (60 μg ml− 1).
They found that polygodial has strong inhibitory effect against the
fungus and operating as a nonionic surface-active agent that disrupts the
lipid-protein interface. More research using omics assessment is
required to clarify the processes by which drimane sesquiterpenoids
exert their antimicrobial activity. The EO of Carpesium macrocephalum is
a source of sesquiterpenes with antimicrbial activity. Application of
Carabrone (bicyclic sesquiterpene) obtained from C. macrocephalum
fruits oil leads to mitochondrial destruction and vacuolation in G. tritici
(Wang et al., 2020).
6.6. Bipolaris sorokiniana
Bipolaris sorokiniana, with teleomorph Cochliobolus sativus, causes
spot blotch as an important fungal disease on wheat and barley
throughout South Asia (Ye et al., 2019; Al-Sadi, 2021; McDonald et al.,
2018). For management of spot blotch disease, antifungal effects of
natural substances such as the EOs obtained from various plant species
have been studied because of having antimicrobial ingredients in their
composition. The EO of Origanum vulgare subsp. viride have been shown
by Mehdizadeh et al. (2018) to have powerful fungistat effect against a
variety of fungal infections, including B. sorokiniana. The fungal in­
fections were successfully defeated by the EOs of O. vulgare subsp. viride
P. Taheri et al.
from two of the thymol-richest locations. Monoterpenes damage the
cytoplasmic membranes of microorganisms, causing the membrane to
be highly impermeable to protons and other big ions. It is difficult to
relate the action of a complex combination to a single or specific
component. Antifungal activity of the EO may be enhanced by major or
minor ingredients. Consideration should be given to the synergistic ef­
fects of the EOs constituents. As a result, the amount and characteristics
of the EOs active ingredients determine how susceptible microbes are to
it (Mehdizadeh et al., 2018). Numerous studies have demonstrated the
bactericide and fungicide effects of the EOs. Their light toxicity confirms
the validity of the ideas emphasizing the need to more focus on research
about the EOs effects on pathogenic fungi (Mehdizadeh et al., 2018).
Salgado et al. (2003) reported that the EO of Eucalyptus spp. inhibited
the mycelial growth of B. sorokiniana at 500 ppm concentration, which
suggested its antifungal activity. Zangoie et al. (2013) demonstrated that
asafoetida seed EO completely inhibited the mycelial growth of B. sor­
okiniana. Metoui et al. (2015) studied the antifungal activity of bitter
orange oil against Fusarium solani, Fusarium oxysporum, B. sorokiniana,
Fusarium avenaceum and Botrytis cinerea. They showed a remarkable
antifungal activity of C. aurantium EO, against B. sorokiniana with 81.75
(%) inhibitory effect at 4 ml/ml concentration. Bitter orange oil contains
linalool (39.30%), lianolyl acetate (32.10%), a-terpineol (10.75%), and
with only a small fraction of limonene (5.28%) all with antimicrobial
effects against various pathogens. The growth of fungi decreased with
increasing concentration of this EO. Chahal et al. (2016) reported the
antifungal potential of camphor and limonene isolated from dill seed EO
against Alternaria triticina and B. sorokiniana using spore germination
inhibition assay. Camphor was most effective against A. triticina whereas
limonene was most effective against B. sorokiniana.
Hamrouni et al. (2015) investigated the effect of Pinus halepensis EO
against various fungi, including Fusarium oxysporum, F. culmorum,
F. nygamai, F. subglutinans, Rhizoctonia sp., Microdochium nivale var
nivale, Alternaria sp, Fusarium verticillioides and B. sorokiniana. Among
the tested fungi, 50% growth inhibition was observed for F. avenaceum
and B. sorokiniana, F. subglutinans and F. nygamai with 52%, while the
inhibition percentage of other fungi was equal or less than 46%.
Although the herbicidal mechanism and probable phytotoxic effects of
the oils was not examined in detail, monoterpene components of the EOs
are known to have phytotoxicity that may cause anatomical and phys­
iological changes in plant seedlings, resulting in accumulation of lipid
globules in the cytoplasm and a reduction in some organelles, like
mitochondria, possibly because DNA synthesis is inhibited or mem­
branes encasing mitochondria and nuclei are disrupted (Hamrouni et al.,
2015). Bahadar et al. (2016) also reported similar findings when
working with: B. sorokiniana fungi of pathogenic on wheat. Thus, cau­
tions should be taken in applying low and effective concentrations of the
EOs using the best way of plant treatment for controlling
phytopathogens.
6.7. Pythium spp
Pythium spp., as one of the main soil-borne fungus-like organisms
belonging to Oomycota, is a widespread phytopathogen throughout the
world. To control Pythium diseases, efforts have been made to develop
natural agents, including plant EOs (Syed et al., 2020; Arora et al.,
2021). Bruni et al. (2004) studied antifungal and antioxidant activity of
the EO obtained from Ishpingo (Ocotea quixos, Lauraceae) fruit. The
main components detected in this EO were b-selinene (2.1%) benzal­
dehyde (3.6%), 1, 8-cineole (8.0%), methylcinnamate (21.6%) and
transcinnamaldehyde (27.9%), which had antimicrobial effect.
Furthermore, this study demonstrated that Ishpingo oil was able to
control the growth of P. ultimum, with MIC of 500 μg/ml. The oil was
found to be a free radical scavenger and an oxidation inhibitor in tests of
in vitro antioxidant activity. The oil has a considerable ability to func­
tion as a general donor of hydrogen atoms or electrons. As a result, these
findings demonstrated that the oil behaved differently as an oxidation
8
Journal of Natural Pesticide Research 6 (2023) 100052
inhibitor and a free radical scavenger. This discrepancy may be caused
by hydro/lipophilicity of the antioxidants and the various processes
engaged in various phases of oxidation.
Amini et al. (2012) studied effectiveness of the EO obtained from
Thymus kotschyanus, Thymus vulgaris and Zataria multiflora plants on the
mycelial growth of four pathogenic fungi, i.e. Fusarium graminearum,
Rhizoctonia solani (AG4), Sclerotinia sclerotiorum and Pythium aphani­
dermatum. The EOs extracted showed 100% growth inhibition on four
pathogenic species at 200 µl/l concentration. The mode of action of
thymus species EOs can be the loss of cytoplasm, folding of the nuclear
membrane, and thickening of fungal cell walls. These abilities of
Thymus species to penetrate the plasma membrane and have lipophilic
characteristics may explain their performance. Such disruptions of
thyme EOs can result from EO components interfering with enzymatic
wall synthesis activities, which alter the shape of fungus and prevent
their proliferation (Amini et al., 2012). Similar results were obtained by
Nafiseh et al. (2011), which demonstrated the inhibitory effect of
eucalyptus EO against mycelial growth of P. ultimum.
6.8. Puccinia spp
Puccinia species are a group of rust fungi, belonging to Basidiomycota
which cause devastating diseases on wheat and other cereal crops (Kaur,
2019; Mehmood et al., 2020). Several investigations have been done to
date on the antimicrobial activities of plant EOs against Puccinia spp.
Tohamey and El-Sharkawy (2014) studied effect the EOs obtained from
chamomile, thyme, cumin, basil, eucalyptus and garlic oils at different
concentrations on the growth of Puccinia triticina under greenhouse and
field conditions. They reported that the latent period (LP) and incuba­
tion period (IP) of the rust disease increased in all tested concentrations
and the numer of pustules decreased in comparison with the control.
Effectiveness of some EOs in controlling wheat stem rust disease in
the field conditions was also investigated. Among the EOs tested, garlic
oil showed the most significant effect and it was found that severity of
the disease has an inverse relationship with increasing the EO concen­
tration (Abdel-Kader et al., 2021). According to the report of Uwineza
et al. (2018), pennyroyal (Mentha pulegium), Atlas cedar (Cedrus atlan­
tica) and clove (Eugenia aromatica), are potentially effective plant
sources against yellow rust caused by P. striiformis on bread wheat. It
was observed that these EOs had no significant effect on the seed weight,
while the fungicide treatment affected the seed weight. Creation of
wheat cultivars, which are resistant to the pathogen has so far been a key
alternative disease control strategy for this disease. However, and
globally, the current status of this wheat constraint would suggest that
resistance breeding alone is ineffective at controlling stripe rust epi­
demics due to the macrocyclic lifecycle of P. striiformis. However,
application of the EOs together with resistant cultivars may alleviate this
constraint by preventing the emergence of new races like those recently
observed in Europe (Wan et al., 2017). The EOs main components,
monocyclic oxygenated monoterpenes, may be responsible for their
antimicrobial properties. Terpenes exact mode of action is unknown,
although it is most likely that these lipophilic substances cause fungi’s
membrane to become less stable (Hmiri et al., 2011).
6.9. Aspergillus flavus
The genus Aspergillus comprises more than 300 species of sapro­
phytic filamentous fungi belonging to the phylum Ascomycota, with a
typical yellow green appearance. In soil and organic debris, Aspergillus
flavus persists as conidia or sclerotia. Conidiophores are specialized
asexual structures that carry conidia. The B1 and B2 aflatoxins are
frequently produced by A. flavus. These poisons are primarily linked to
food products, like cereals, oilseeds, and nuts. The species A. flavus is one
of the leading causes of food spoilage due to the mycotoxins it produces;
that’s why many studies on the antifungal activity of various EOs have
been focused on Aspergillus (Bhatnagar-Mathur et al., 2015; Uka et al.,
P. Taheri et al.
2020; Cho et al., 2022).
The antifungal activity of Cymbopogon citratus oil was investigated
against Aspergillus ochraceus, A. niger and A. flavus by Sonker et al.
(2014) and the MIC value of the EO was 0.29 μL/ml for all fungi tested
and its MFC value was 0.57 µl/ml for A. ochraceus and 0.64 µl/ml for
A. niger and A. flavus. The EO of C. citratus caused 100% mycelial growth
inhibition against all mentioned fungi (Sonker et al., 2014). At 0.8 L/ml
in A. flavus, A. niger, and A. ochraceus, C. citratus oil significantly
inhibited the generation of ochratoxin production. Various aspects of
plant EOs antitoxigenic effects against fungal species have been
demonstrated, including cell membrane disruption, decreased fungal
growth, and suppression of carbohydrate catabolism (Burt, 2004; Sonker
et al., 2014)). Application of C. citratus oil at 200 and 300 L concen­
trations during an in vivo study led to superior control of grape related
fungus. In grapes treated with this EO, fungal decays might be mini­
mized along with weight loss and fruit firmness (Sonker et al., 2014).
Recently, Nerilo et al. (2016) reported that ginger EO (GEO)
exhibited strong antifungal activity against A. flavus having the MIC and
MFC values of 150 µg/ml. Additionally, it was claimed that mycelium
development slowed down by increasing concentrations of the GEO.
Aflatoxin formation was suppressed by the GEO compounds due to their
antioxidant effects, but because they also exhibited antifungal activity,
the reduction in biomass alone also reduced aflatoxin production.
Aflatoxin formation is prevented by a decrease in oxidative stress and
fungal biomass, possibly as a result of an additive action. Genetic
research revealed that when the fungus comes into contact with anti­
oxidants, the genes responsible for the transcription of proteins involved
in the aflatoxin production are suppressed (Nerilo et al., 2016). It should
come as no surprise that ginger includes a number of antioxidants,
including sesquiterpenes, monoterpenes, and flavonoids (Nerilo et al.,
2016), which are involved in decreasing mycotoxin production by
mycotoxigenic fungi.
In another similar work done by Tian et al. (2012) in vitro, antifungal
activity of Cinnamomum jensenianum EO was evaluated against A. flavus
at different concentrations (1, 2, 4, 6, and 8 µl/ml) and this EO showed
100% inhibition at 6 µl/ml concentration and the MIC of 8 µl/ml. These
findings demonstrated that A. flavus ability to produce aflatoxin B1 and
dry mycelium weight can be effectively inhibited by C. jensenianum.
Reduction in the ability of A. flavus to produce aflatoxin B1 can be linked
to the inhibition of carbohydrate catabolism in A. flavus by acting on
some important enzymes (Tian et al., 2012).
In another study by Tavares et al. (2008), inhibition of mycelial
growth was observed to test the fungicidal potential of Daucus carota
subsp. halophilus EO against A. flavus, A. fumigatus and A. niger. It was
found that the EO of D. carota subsp. halophilus possessed significant
antifungal activity against all the tested strains with the MIC values
ranging from 0.16 to 0.32 µl/ml.
In addition, Pinto et al. (2017) reported that the EO of Thapsia villosa
exhibited strong antifungal activity against A. flavus with the MIC value
of 1.25 µl/ml and the MFC of more than 5 µl/ml. Limonene was the main
component identified in this EO. The cyclic monoterpene hydrocarbon
family includes limonene, which is widely utilized in many different
industries and works well in dissolving cholesterol. The hydrophobicity
of limonene appears to make it easier for the lipids that have built up in
the microbial plasma membrane to dissolve, which leads to a breakdown
in the membrane integrity (Espina et al., 2013; Pinto et al., 2017). A
crucial target for the limonene component appears to be the microbial
cell envelope (Pinto et al., 2017). According to the study by Vale-Silva
et al. (2010), the EO of Thymus x viciosoi aerial parts was rich in oxygen
monoterpenes, especially γ-terpinene (7.5%), thymol (18.0%), p-cym­
ene (19.0%) and carvacrol (30.0%), which have considerable antifungal
effects against A. flavus.
6.10. Penicillium spp
Penicillium is a genus of anamorphic fungi. This genus belongs to
9
Journal of Natural Pesticide Research 6 (2023) 100052
Ascomycota and is phylogenetically represented by 429 globally valid
species, 2 subgenera and 26 sections under, according to the Interna­
tional Commission of Penicillium and Aspergillus (ICPA) (Ashtekar et al.,
2021). Penicillium species are widely studied in the agricultural, indus­
trial and medical fields because of their capability in producing wide
range of secondary metabolites (Toghueo and Boyom, 2020). Several
studies have been developed for testing the effect of medicinal plants
against Penicillium spp., causing storage molds on cereals and numerous
other agricultural crops.
The EO obtained from Gallesia integrifolia demonstrated antimicro­
bial potential in studies conducted by Raimundo et al. in 2018 against a
variety of microorganisms, including Penicillium ochrochloron, P. funi­
culosum and P. verrucosum var. cyclopium. Fungicidal activity of the EO
ranged from 0.02 to 0.18 mg/ml. The fungistatic activity (MIC) of this
EO ranged from 0.01 to 0.09 mg/ml. The antifungal activity of
G. integrifolia EO has been attributed to its main components. Organo­
sulfur was the dominant component of the EO of this plant. Its other
major compounds were 2, 8-dithianonane (52.63%), lenthionine
(14.69%) and dimethyl trisulfide (15.49%). The organosulfur can be
produced in vegetal tissues from sulfur containing amino acids like
cysteine and methionine. The existence of disulfide linkages in these
molecules is also related to the antimicrobial properties of organosulfur.
Therefore, the inclusion of sulfur compounds in the EOs that have pol­
ysulfide bridges (-Sn-) can improve their polarity and broaden their
chemical affinity with the chitin and ergosterol rich cellular membrane
and wall structures of microorganisms. As a result of this contact with
the membranes, membrane rupture and microbial cell imbalance may be
promoted (Levinson, 2016; Raimundo et al., 2018).
Tagetes patula EO is another EO with a broad range of antifungal
properties. Due to its potent medical qualities, this Asteraceae-family
plant is one of the most widely used medicinal plants in the world.
Romagnoli et al. (2005) reported that the EO of T. patula exhibits anti­
fungal action against B. cinerea and P. digitatum. At 1.25 l/ml, the
mycelial growth of P. digitatum was entirely inhibited. Against B. cinerea,
on the other hand, it displayed a dose-dependent growth suppression
with a MIC of 10 l/ml.
Using the microdilution technique, Džamić et al. (2013) assessed
Hyssopus officinalis subsp. pilifer oil’s effectiveness as an antifungal
agent. Strong antifungal action was observed for different components
of this EO, including transpinocamphone, isopinocamphone, pinene,
and 1.8-cineole, with the MIC values of 52.20 and 26.10 mg/ml,
respectively. Also, H. officinalis subsp. pilifer EO showed promising
antimicrobial activity against Penicillium funiculosum and Penicilium
ochrochloron.
In another study by Atrash et al. (2018) the fungicidal potential of
Savory EO (SEO) was evaluated against P. digitatum at concentrations
ranged between 200 and 1200 µl/L. The SEO was found to be the most
effective at concentrations of 1000 and 1200 µl/L with 100% inhibition.
In an in vivo investigation, the use of SEO at 800 or 1000 ml/L pre­
vented fruit rot for up to 15 days after storage. Plasma membrane of the
fungus may be damaged by SEO high carvacrol concentration, which
could limit fungal infection (Atrash et al., 2018).
Similarly, in another work by Cid-Pérez et al. (2016), Poliomintha
longiflora EO was reported against P. expansum, B. cinerea, A. flavus, and
Colletotrichum gloeosporioides, by using agar ilution method with MIC of
0.8–1.4 g/L. This activity was attributed to thymol as the main ingre­
dient of the EO. Therefore, monoterpene esters may interact with cell
wall enzymes, such as the α- and β glucanases, as well as with chitin
synthase/chitinase of pathogens. The EO obtained from Origanum vul­
gare spp., whose main constituent was thymol, showed inhibitory effect
against A. flavus, P. oxalicum, and P. minioluteum in a study by Camiletti
et al. (2014) and the MIC values of 350–650 ppm were reported.
7. Effect of different EOs on major bacterial pathogens of cereals
The top five cereal bacterial pathogens list include Pseudomonas,
P. Taheri et al.
Xanthomonas, Rathayibacter Pantoea and Bacillus cereus, which cause
destructive diseases worldwide (Mansfield et al., 2012: Abbas, 2022). In
this section, in vitro and in vivo antibacterial effects of various EOs are
reviewed against these major bacterial phytopathogens.
7.1. Pseudomonas syringae
Pseudomonas is a large and diverse genus of bacteria that is widely
distributed in various natural environments. The species of this genus
are found in soil, water, on plant organs and around the roots of plants.
This genus is placed in the Pseudomonadaceae family. Species of this
genus may be saprophytic, plant or animal pathogens, and some of them
produce ice nuclei (Saati-Santamaría et al., 2021). The ice damages the
plant cell wall (frost injury) and allows bacteria to get inside (Hirano and
Upper, 2000; Lukas et al., 2022). Some bacterial secretion systems have
been explained by studying the species of this genus. Pseudomonas spe­
cies are aerobic, rod-shaped, Gram-negative, bacteria measuring
0.5–0.8, μm by 1.5–3.0 µm.
Various species of the genus Pseudomonas cause disease in dicotyle­
donous and monocotyledonous plants, including cereals. For instance,
one of the most significant wheat diseases in the world, bacterial blight,
is caused by Pseudomonas syringae pv syringae. A study was conducted to
examine the impact of Thymus vulgaris and Origanum vulgare EOs on
production of virulence factors in plant pathogenic P. syringae species,
including anti-toxin and anti-biofilm activities. Coronatine, tabtoxin,
and syringomycin were also evaluated phenotypically and genotypi­
cally. The studies revealed that both EOs had potent inhibitory effects on
P. syringae, with the MIC ranges of 1.43–11.5 mg/ml for thyme and
5.8–11.6 mg/ml for oregano. Most strains of P. syringae were inhibited
from biofilms and cronatine, syringomycin, and tabetoxin production
when exposed to these EOs (Carezzano et al., 2017). Thyme and oregano
EOs were discovered to have good inhibitory action against P. syringae
(Carezzano et al., 2017).
Also, in a study by Shabani et al. (2019) to investigate the control
effect of three different EOs from the aerial parts of rosemary (Rosmar­
inus officinalis), oregano (Mentha spicata) and thyme (Thymus daenensis
Celak. Subsp. daenesis) on wheat bacterial blight, it was found that these
EOs are effective in preventing growth of bacterial colonies and thus
reducing the disease. Carvacrol was the main component identified in
Thymus vulgaris and Origanum vulgaris. Carvacrol has the ability to target
bacteria’s cytoplasmic membrane and damage the phospholipid bilayer
there. Carvacrol is an intrinsically hydrophobic monoterpene that
quickly penetrates bacterial cell membranes, leading to a loss of their
integrity and the release of bacterial cell contents, which ultimately
leads to cell death (Mechmechani et al., 2022).
In another study by Aghajanighara et al. (2020) using the EOs of 9
different medicinal plants, including Juncus sp., it was found that the EO
of this plant has controlling effect against various bacteria, including
Pseudomonas syringae pv syringae. Normal bacterial growth and meta­
bolism are significantly influenced by the integrity of the cell membrane.
Nucleic acids are considered as vital macromolecules in the living cells
and their leakage could cause cell death. The EO damaged and ruined
cell membrane integrity, causing macromolecules like nucleic acids to
flow out of the cells (M. Wang et al., 2018; Y. Wang et al., 2018). Ac­
cording to reports on the mechanism of the EOs action, they may
contribute to cell leakage, cytoplasmic membrane damage, and cell wall
breakdown (Goldbeck et al., 2014).
7.2. Xanthomonas spp
Xanthomonas is a short rod-shaped aerobic, Gram-negative bacte­
rium belonging to the family Xanthomonadaceae. The species of this
genus include a wide range of plant pathogens that cause numerous and
diverse diseases in all kinds of plant genera (da Silva et al., 2017; Afzaal
et al., 2022). The species Xanthomonas translucens is responsible for
significant yield losses in cereals. Various pathovars of this pathogen are
10
Journal of Natural Pesticide Research 6 (2023) 100052
defined according to their host range, molecular and biochemical
characteristics and are divided into two major groups: translucense and
graminis. The translucense group includes pathovars that cause bacte­
rial leaf streak (BLS) on cereals. The BLS is distributed as an important
disease in many wheat and barley production regions, worldwide.
Xanthomonas translucens pv translucens is seed-borne in wheat, barley,
rye and triticale as well as some grasses (Sapkota et al., 2020). Beyki and
Alizadeh (2006) investigated effect of the EOs obtained from several
plants on two pathovars (cerealis and hordei) of X. translucens. Ac­
cording to their report, the antibacterial effect of plant extracts and EOs
on the two investigated pathovars was almost similar, and in fact, the
two pathovars did not differ in terms of sensitivity to these compounds.
The EOs of Mentha aquatica, Echinophora sibthorpiana, Hyssopus offici­
nalis and Ziziphora persica had the highest antibacterial effect against the
pathogen. Fazeli-Nasab et al. (2022) investigated the antibacteial effect
of the EOs obtained from 10 medicinal plants against Rathayibacter tritici
and Xanthomonas translucens bacteria. Thyme EO had the lowest MIC,
which was 6.25 ppm and inhibited R. tritici colony growth.
Hypericum perforatum EO had the lowest MIC, which was 6.25 ppm
and inhibited X. translucens growth. The lowest concentration of the EO
of yew and fennel was 6.25 ppm, which inhibited the growth of both
bacteria. Xanthomonas oryzae pv oryzae, another plant pathogenic bac­
terium in this genus, is the causal agent of bacterial blight in rice. This
pathogen is damaging in tropical Asian countries and has been reported
from Australia, Africa and Latin America, as well as in North America
(Shen and Ronald, 2002). Depending on the stage of rice growth,
geographical location and seasonal conditions, this disease can cause a
severe yield reduction of up to 50% (Shen and Ronald, 2002). In the
studies of Singh et al. (2017), screening of natural EOs from different
parts of plants such as, leaves, seeds, roots, rhizomes was performed
against X. oryzae pv. oryzae. The results showed reduction in biofilm
formation in 9 out of 15 oils at 1000 ppm concentration without sig­
nificant reduction in total colony forming units. Investigations deter­
mined that movement characteristics (such as swimming) in the case of
Acorus calamus and Melaleuca leucadendron EOs and virulence traits such
as protease and exopolysaccharide, which are the pathogenic factors of
this bacteria, were reduced via using the EO obtained from Acorus cal­
amous and Nardostachys jatamansi. The highest decrease in lipase ac­
tivity of the pathogen was also observed by application of Vitex agnus -
castus root EO. These studies indicated that some plant EOs may be used
in the future to develop potential bactericides against X. oryzae pv.
oryzae. Also, in a study by Abdullahi et al. (2020), it was found that the
EO of ginger plant (Zingiber officinale) can be useful in controlling some
fungal and bacterial plant pathogens, including X. oryzae pv. oryzae.
7.3. Rathayibacter spp
Rathayibacter is one of the plant pathogenic gram-positive genera in
the Microbacteriaceae family. Species of this bacterial genus, such as
Rathayibacter tritici and Rathayibacter iranicus, cause tundu and gum­
ming disease in wheat, respectively, and Rathayibacter toxicus causes
annual ryegrass toxicity and bacterial head blight of grasses. Rathayi­
bacter tritici species is anaerobic, non-motile and rod-shaped bacterium
that causes spike blight in wheat and barley (Evtushenko and Dorofeeva,
2015; Park et al., 2017).The species R. toxicus is distinguished from the
other species by producing a type of toxin called corynetoxin in plants.
Toxin production is associated with infection by a bacteriophage
(Murray et al., 2017). The studies of Ghahari et al. (2015) on EO and
extract of the Seeds of Glycine max showed that the EO can control
R. toxicus at concentration of 25 µg/ml. In a research by Altundag et al.
(2010), the antibacterial effects of Satureja cuneifolia EO belonging to the
Labiatae family were tested on wheat pathogenic bacteria including
R. iranicus, R. tritici, Xanthomonas translucens pv. translucens, Pseudo­
monas syringae pv. atrofaciens and R. rathayi. The antibacterial effect was
tested based on the micro-well diffusion method at different concen­
trations, and the diameter of the inhibition zone caused by the EOs
P. Taheri et al.
ranged from 8 to 32 mm, for P. syringae pv. atrofaciens and R. rathayi
respectively. The data obtained by Fazeli-Nasab et al. (2022) revealed
that 6.25 ppm concentration of thyme EO inhibited the growth of
R. tritici. Due to the high hydrophobicity of the EOs, one of their primary
modes of action is the destruction of bacterial structures and the loss of
cell membrane-specific barrier function. This has an impact on a number
of processes, including solute transport, metabolic regulation, energy
state maintenance, and membrane-coupled energy transfer processes.
However, the EOs are also responsible for other side effects as membrane
protein destruction, cytoplasmic coagulation, and cell wall breakdown.
High amounts of the EO may denature enzymes, although sub-lethal
concentrations may inhibit them. Since thymol, a component of the
EOs, has already been demonstrated to have the ability to disrupt the
cytoplasmic membrane, its mechanism of action is well understood. This
is most likely because thymol can destabilize the phospholipid bilayer,
which results in an increase in permeability (Fazeli-Nasab et al., 2022).
The antibacterial effects of oregano and thyme EOs on pathogens in vivo
conditions remain to be determined, considering the fact that they
showed significant antibacterial activity in vitro.
7.4. Pantoea stewartii
Pantoea stewartii is a bacterium responsible for serious losses of corn
crop in the world and especially in America, where it is native (Duong
et al., 2017). This bacterium belongs to the family Enterobacteriaceae.
Pantoea stewartii subsp. stewartii is a gram-negative bacterium that
mainly causes disease in sweet corn (Walterson and Stavrinides, 2015;
Roper, 2011). Moghaddam et al. (2014) investigated the inhibitory ef­
fects of Ocimum ciliatum EO on 10 bacterial strains, including Ralstonia
solanacearum, Rhodococcus fascians, X. oryzae pv. oryzae, P. syringae pv.
syringae, Brenneria nigrifluens, Xanthomonas citri, Agrobacterium vitis,
P. stewartii subsp. indologenes, P. syringae pv. Lachrymans, and P. tolaasii.
The EO GC and GC-MS results showed that the EO of this plant is mainly
contains methyl chavicol (87.6%), methyl eugenol (2.6%) and 1,
8-cineole (1.7%). The results of antibacterial test showed that the oil has
antibacterial activity against all tested bacteria, including P. stewartii
subsp. indologenes.
7.5. Bacillus cereus
Bacillus cereus is a ubiquitous facultative anaerobic, gram-positive,
endospore forming bacterium belonging to the family Bacillaceae. B.
cereus is compatible with different environments due to the production
of different biofilms (Baldwin, 2020; Ehling-Schulz et al., 2019). Bos­
kovic et al. (2015) tested the antibacterial activity of Origanum vulgare
(oregano) EO against four pathogenic bacteria (Salmonella enteritidis,
Bacillus cereus, Salmonella typhimurium and Staphylococcus aureus) using
broth dilution assay with the MIC values of 160–640 µg/ml against all
strains. Carvacrol alters the membrane potential, depletes the intracel­
lular ATP pool, and makes the cytoplasmic membrane more permeable
to potassium ions in B. cereus (Boskovic et al., 2015).
Clemente et al. (2016) used broth dilution method to demonstrate
the antibacterial efficacy of Cinnamomum zeylanicum EO against
gram-positive and gram-negative bacteria. The antibacterial action of C.
zeylanicum EO was demonstrated by the MIC ranges of 100–400 g/ml
against all strains tested. Agar diffusion method was used by Ksouda
et al. (2019) to examine antibacterial effect of the EO obtained from the
seeds of Pimpinella saxifrage against Pseudomonas aeruginosa, Salmonella
Typhimurium, Escherichia coli, Micrococcus luteus, Listeria monocytogenes,
and B. cereus. This EO showed antibacterial activity, with the MIC values
of 0.78–3.12 mg/ml and IC50 of 6.81 g/ml against all strains tested.
Hence, the presence of anethole could be the cause of P. saxifrage EOs
antibacterial properties. P. saxifrage EO, which is primarily made of
anethole, displayed strong antibacterial and antioxidant properties. It is
generally challenging to pinpoint a single mechanism for the antibac­
terial action of EOs due to the abundance of their constituent parts. Yet,
11
Journal of Natural Pesticide Research 6 (2023) 100052
it has been shown in numerous studies that one of the primary anti­
bacterial action mechanisms is connected to the dissolving of EO com­
ponents in the lipids of the bacterial cell membrane and mitochondria
because of their lipophilic nature. This disrupts the cell membrane and
causes ions to flow out, which causes bacterial cell death (Ksouda et al.,
2019).
Purkait et al. (2020) tested the antibacterial activity of the EOs ob­
tained from of Syzygium aromaticum, Piger nigerium and Cinnamomum
zeylancium against B. cereus. All EOs tested showed significant antimi­
crobial activity with different degrees of inhibition. According to
Mohammed and Al-Bayati (2009), the EOs of Thymus kotchyanus aerial
parts had the best inhibitory effect against B. cereus, Klebsiella pneumo­
niae, and Escherichia coli, while Escherichia coli, Proteus mirabilis, and
Staphylococcus aureus were inhibited with higher concentration of the
EO. In general, phenolic compounds including carvacrol, eugenol, and
thymol are highly present in EOs with potent antibacterial effects
against microorganisms. It makes sense that their mode of action would
be similar to that of other phenolics, which is generally thought to
involve disruption of the cytoplasmic membrane, disruption of the
proton motive force (PMF), disruption of electron flow, disruption of
active transport, and disruption of the coagulation of cell contents (Burt
et al., 2004). Thymol can also break down the outer membrane of
gram-negative bacteria, releasing lipopolysaccharides (LPS), and
improving the cytoplasmic membrane’s permeability to ATP (Moham­
med and Al-Bayati, 2009).
In another study using Pinus halepensis EO, significant antimicrobial
activity of this EO was observed against Staphylococcus aureus, Bacillus
subtilis, Proteus mirabilis, Listeria monocytogenes, E. coli and P. aeruginosa
(Bouyahya et al., 2019). The outcomes demonstrated that all of the
examined microorganisms were significantly inhibited by the EO of P.
halepensis. This effect would be related to their main ingredients, espe­
cially caryophyllene, which is recognized for its antibacterial properties
yet. It can be challenging to pinpoint a single or specific constituent as
the active ingredient in a complicated mixture. In fact, some researches
suggest that minor ingredients are crucial to antibacterial activity,
maybe by creating a synergistic interaction with other ingredients.
Various antibacterial effects of the EOs may result from their varied
chemical composition (Dahham et al., 2015; Bouyahya et al., 2019). The
majority of studies have been focused on plant EOs modes of action.
However, research on the antibacterial properties of the EOs and their
constituents is still ongoing.
According to several reports, antibacterial properties of the EOs
might be related to their hydrophobicity, which causes a rise in cell
permeability and the subsequent leakage of cell components. The
disturbance of cell walls and membranes, modification of ATP genera­
tion and protein synthesis, pH disturbance, intracytoplasmic alterations,
DNA damage, and inhibition of quorum sensing are the most well-
known targets of EOs in bacteria (Cox et al., 2000). However, more
research is needed to purify and identify chemical structure of bioactive
components and clarify their antibacterial actions.
8. Conclusion
In this study, the antimicrobial activities of different EOs, as natural
plant protectant agents against most important fungal and bacterial
pathogens of cereal crops, were reviewed according to the latest reports.
It is demonstrated that the EOs of different plant species have strong
antimicrobial activities on numerous pathogens of cereals. The use of
EOs can be considered as an eco-friendly and safe alternative method to
be replaced with application of hazardous agrochemical which are
harmful for both consumers and the environment. The EOs are capable
of inhibiting growth and reproduction of various pathogens in the host
plant tissues and disrupting cell membrane of phytopathogens. Although
many EOs and their active compounds have been approved to be natural
control agents for numerous phytopathogens in vitro and in greenhouse
conditions, whether all EOs can be used in controling plant pathogens in
P. Taheri et al. Journal of Natural Pesticide Research 6 (2023) 100052

Table 1
Essential oils (EOs) active against pathogenic fungi in cereals and their modes of action.
Fungi EOs Assay MIC Mode of action of the References
EOs
Plant Part Major compounds

Fusarium graminearum Zingiber officinale Rhizomes A-zingiberene In - 1000 µg/ml Inhibited ergosterol Ferreira et al.
vitro and DON production (2018)
F. graminearum Curcuma longa Rhizomes Ar-turmerone In In 2450 mg/ml Inhibited ZON and K.N. Kumar et al.
vitro vivo fungal biomass (2016); P. Kumar
et al. (2016)
F. graminearum Anacyclus valentinus Aerial 3-carene In - 1.25–25 µl/ml Antifungal activity Houicher et al.
vitro (2018)
F. graminearum Eucalyptus sp Leaves 1,8 cineole In - 1000 µl/l Antifungal activity Davari and Ezazi
vitro (2017)
F. graminearum Ocimum sanctum Aerial Eugenol In - 1250 µg/ml Effect on spore Kalagatur et al.
vitro germination and (2015)
mycelia structure
F. graminearum Zataria multiflora - Thymol carvacrol In - 16 µl Inhibited DON Lahooji et al.
Satureja hortensis vitro 31.5 µl (2010)
F. graminearum Artemesia afra Leaves - In - 0.24 mg/ml Effect on mycelia Amidou et al.
Pelargonium vitro structure (2012)
graveolens
Eriocephallus
punculatus
Magnaporthe oryzae Periploca sepium Root 2-Hydroxy-4- In - 300 µg/ml Antifungal activity Wang et al. (2010)
methoxybenzaldehyde vitro
M. oryzae Corymbia citriodora Leaves Citronellal In - 200 ppm Germination Aguiar et al. (2014)
Cymbopogon nardus vitro 100 ppm inhibition
M. oryzae Illicium verum Fruit Trans- anethole In - Ic50:0.32 mg/ml Germination Huang et al. (2010)
vitro inhibition
M. oryzae Azadirachta indica Leaves - In In - - Kumar et al. (2017)
vitro vivo
M.oryzae Eugenia aromatica seed - In - 100% Antifungal activity Olufolaji et al.
vitro (2015)
Rhizoctonia solani Eucalyptus Leaves - In - - Antifungal activity Nafiseh et al.
camaldulensis vitro (2011)
R. solani Hypericum Aerial Caryophyllene In - 1gr/ml Antifungal activity Cakir et al. (2004)
hyssopifolium parts Terpineol vitro
H. heterophyllum
R. solani Cinnamomum - - In - 500 ppm Effect on mycelia Seema and Devaki
zeylanicum vitro structure and sclerotia (2010)
formation
R. solani Syzygium - - In - 1000 ppm Effect on mycelia Seema and Devaki
aromaticum vitro structure and sclerotia (2010)
formation
R. solani Coriandrum sativum - - In - 1000 ppm Effect on mycelia Seema and Devaki
vitro structure and sclerotia (2010)
formation
R. solani Foeniculum vulgare - - In - 2000 ppm Effect on mycelia Seema and Devaki
vitro structure and sclerotia (2010)
formation
R. solani Myrtus communis Total plant Ecalyptol In - 10 µl/Petri Antifungal activity Yilar et al. (2016)
vitro
R. solani Zaturia multiflora Total plant Thymol In - 200 ppm Effect on mycelia Nasseri et al. (2015)
vitro structure and sclerotia
formation
R. solani Acacia farnesiana - - In - 2 v/v Effect on mycelia Udomsilp et al.
vitro growth and (2009)
Germination
inhibition
R. solani Mikania scandens Leaves В- caryophyllene In - 250 µg/ml Antifungal activity Siddiqui et al.
vitro (2017)
− 1
Gaeumanomyces Drimys wintri Bark α - pinene In In 250 mg kg Effect on mycelia Monsálvez et al.
graminis vitro vivo growth (2010)
G. graminis Drimys wintri Bark N-hexane In - IC50: 198 mg/L Effect on mycelia Zapata et al. (2011)
vitro growth
G. graminis Drimys wintri Bark Polygodial In - LC50:7 µg/ml Fungal membrane Paz et al. (2020)
vitro damage increase of
lipid peroxidationn
Bipolaris sorokiniana Origanum vulgare Flower Thymol In - 2.1 g/ml Antifungal activity Mehdizadeh et al.
subsp. viride vitro (2018)
B. sorokiniana Eucalyptus Leaves Globulol In - 500 ppm Antifungal activity Salgado et al.
vitro (2003)
B. sorokiniana Asafoetida Seed - In - - Antifungal activity Zangoie et al.
vitro (2013)
B. sorokiniana Citrus aurantiom Leaves 9 Linalool In - 4 ml/ml Antifungal activity Metoui et al. (2015)
vitro
(continued on next page)

12
P. Taheri et al. Journal of Natural Pesticide Research 6 (2023) 100052

Table 1 (continued )
Fungi EOs Assay MIC Mode of action of the References
EOs
Plant Part Major compounds

B.sorokiniana Dill Seed Limonene In - - Antifungal activity Chahal et al. (2016)

vitro
B. sorokiniana Pinus halepensis Fresh plant Z- caryophyllene In - - Effect on seed Hamrouni et al.
vitro germination and (2015)
seeding grow
inhibition
Phythium spp Ocotea quixos Fruit Trancinnamalde In - 500 µg/ml Antifungal activity Bruni et al. (2004)
vitro and antioxidant
activity
P. aohanidermatum Thymus kotschyanus Aerial Carvacrol In - 200 µl/l Antifungal activity Amini et al. (2012)
Thymol vitro
P. aphanidermatum Thymus vulgaris Aerial Carvacrol In - 200 µl/l Antifungal activity Amini et al. (2012)
Thymol vitro
P. aphanidermatum Zataria multiflora Aerial Carvacrol In - 200 µl/l Antifungal activity Amini et al. (2012)
Thymol vitro
P. ultimum Ecalyptus Leaves - In - 100% Antifungal activity Nafiseh et al.
camadulensis vitro (2011)
Fusarium verticillioides Zingiber officinale Rhizomes α – Zingiberene and In - 2500 µg/ml Inhibited ergosterol Yammamoto et al.
Citral vitro And fumonisin (2013)
production
F. verticillioides Myrtus communis Total plant Linalool and 1,8 cineol In - 0.0003–3.25 mg/ Antifungal activity Starovic et al.
vitro ml (2016)
F.verticillioides Rosmarinus Leaves 1,8 Cineole In - 150LG/ml Inhibited ergosterol Da silva Bomfim
officinalis vitro And fumonisin et al. (2015)
production
F. verticillioides Cinnamon - Cinnamaldehyde In - 45 µl/L Antifungal activity Xing et al. (2014)
vitro
F.verticillioides Curcuma longa Rhizomes α - Turmerone and In - 73.7 mg/ml Inhibited ergosterol Avanço et al.
β-Turmerone vitro (2017)
F. verticilioides Origanum x appli - Thymol and cis sabinene In - Ic50: 66.79UL/L Antifungal activity Pizzolitto et al.
vitro (2020)
F. verticillioides Pimenta dioica - Eugenol In - 0.6 µl/ml Antifungal activity Zabka et al. (2009)
vitro
Fusarium Zhumeria majdae Aerial Linalool, Camphor In - 2.33 × /10–9 µl/ Antifungal activity Davari and Ezazi
verticillioides vitro 103spor (2017)
Fusarium Zhumeria majdae Aerial Linalool, Camphor In - 2.33 ×/10–9 µl/ Antifungal activity Davari and Ezazi
verticillioides vitro 103spor (2017)
Fusarium Mentha arvensis Seed Menthol and Menthone In - 1.53 × 10–4 µl/ Antifungal activity K.N. Kumar et al.
verticillioides vitro 103 (2016); P. Kumar
(KT855217) et al. (2016)
F.verticillioides Mentha arvensis Seed Menthol and Menthone In - 1.53 × 10–4 µl/10 Antifungal activity K.N. Kumar et al.
(KT855216) vitro (2016); P. Kumar
et al. (2016)
Aspergillus flavus Cymbopogon Leaves E- Citral In In 0.29 µl/ml Inhibited mycotoxin Sonker et al. (2014)
citratus vitro vivo
A.flavus Zingiber officinale Rhizome α-Zingiberene In - 150 µg/ml Effect on the conidial Nerilo et al. (2016)
geranial vitro germination and
antimycotoxin
A.flavus Cinnamomum bark Eucalyptol In - 8 µl/ml Effect on the conidial Tian et al. (2012)
jensenianum vitro germination
A.flavus Daucuscarota Flowering - In - 2.5 v/v Antifungal activity Tavares et al.
subsp. halophilus Umbel and vitro 20 v/v (2008)
seed
A.flavus Thapsia villosa Aerial and Limonene In - 1.25 µl/ml Antifungal activity Pinto et al. (2017)
fruit vitro
A.flavus Thymus villosa Aerial Carvacrol In - 0.32 µl/ml Antifungal activity Vale-silva et al.
vitro (2010)
Penicillium spp Gallesia integrifolia Fruit 2,8- dithianonane In - 0.01–0.09 mg/ml Antifungal activity Raimundo et al.
vitro (2018)
P. digitatum Tagetes patula Flower Piperiton In - 1.25 l/ml Antifungal activity Romagnoli et al.
vitro (2005)
P. funiculosum Hyssopus officinalis Aerial 1,8- cineole In - 52.20 mg/ml Antifungal activity Džamić et al.
L. subsp. pilifer vitro Antioxidant activity (2013)
P.digitatum Satureja hortensis Fruit Carvacrol and g- In In 1000 µl/L Antifungal activity Atrash et al. (2018)
terpinene vitro vivo Antioxidant activity
P.expansum Poliomintha Leaves Thymol and carvacrol In - 1.2 g/L Antifungal activity Cid-Pérez et al.
longiflora vitro (2016)
P. oxalicum Origanum vulgare Leaves Thymol In - 350–650 PPM Antifungal activity Camiletti et al.
P.minioluteum spp vitro (2014)

13
P. Taheri et al. Journal of Natural Pesticide Research 6 (2023) 100052

Table 2
Essential oils (EOs) active against pathogenic bacterial in cereals and their modes of action.
Bacteria EOs Assay MIC Modes of action of the EOs References

Plant Part Major compounds

1.43 − 11.5 mg/ml
Pseudomonas syringae Thymus vulgaris Leaves Carvacrol P- In - Antitoxin Antibiofilm Carezzano et al.
Cimene vitro (2017)
P. syringae Origanum vulgare Leaves Carvacrol P- In - 5.8–11.6 mg/ Antitoxin Carezzano et al.
cymene vitro ml Antibiofilm (2017)
Cis- sabinene
hydrate
Xanthomonas Hypericum Leaves - In In 6.25 ppm Antibacterial activity Fazeli-Nasab et al.
translucens perforatum vitro vivo (2022)
X. translucens Yew and fenneel Leaves - In - 6.25 ppm Antibacterial activity Fazeli-Nasab et al.
vitro (2022)
Rathayibacter tritici Thyme Leaves Thymol In - 6.25 ppm Antibacterial activity Fazeli-Nasab et al.
vitro (2022)
R. tritici Glycin max Seed - In - 25 µg/ml Antibacterial activity Ghahari et al. (2015)
vitro
R. iranicus subsp. Origanum Aerial Carvacrol In - 125 µg/ml Antibacterial activity Dagh et al. (2011)
iranicus minutiflorum vitro
Pantoea stewarti subsp. Ocimum ciliatum Aerial Methyl In - 4 mg/ml Antibacterial activity Moghaddam et al.
indologenes Carvacrol vitro (2014)
Bacillus cereus Origanum vulgar Aerial Carvacrol In - 160 µg/ml Antibacterial activity Boskovic et al. (2015)
vitro
B. cereus Cinnamomum Bark Carvacrol In - 200 µg/ml Antibacterial activity, cellular Clemente et al.
zeylanicum vitro damage, cell cycle arrest (2016)
B. cereus Pimpinella Seed Anethole In - 1.56 mg/ml Antibacterial activity, antioxidant Ksouda et al. (2019)
saxifrages Pseudoisoeugenol vitro activity and damage of DNA
B. cereus Syzygium Bud Eugenol In In 78.73 Antibacterial activity Purkait et al. (2020)
aromaticum vitro vivo ± 3.48 µg/ml
B. cereus Cinnamomum Bark Cinnamyl acetate In - 82.69 Antibacterial activity Purkait et al. (2020)
zeylancium vitro ± 3.21 µg/ml
B. cereus Thymus Aerial Thymol In - 15.6 mg/ml Antioxidant activity and Mohammed and
kotchyanus vitro antioxidant activity Al-Bayati (2009)
B. cereus Pinus halepensis Aerial β-caryophyllene In - 1000 v/v Antibacterial activity and Bouyahya et al.
vitro antioxidant activity (2019)

the fiels conditions, at which concentration and via which treatment
methods needs to be carefully evaluated. In general, the investigation
and collection of information about antimicrobial activity of various EOs
in vitro and in vivo conditions might be helpful for identifying natural
agents suitable as an alternative, safe, cheap and easily degradable
compounds with plant origin, which could be formulated and used in
disease management strategies to decrease the use of hazardous agro­
chemicals, such as synthetic fungicides and bactericides, to produce
high quality organic crops based on increasing demand for these healthy
agricultural products worldwide Tables 1 and 2.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data Availability
No data was used for the research described in the article.
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Further reading
Ghahari, S., Alinezhad, H., Nematzadeh, G.A., Tajbakhsh, M., Baharfar, R., 2017.
Chemical composition, antioxidant and biological activities of the essential oil and
extract of the seeds of glycine max (soybean) from North Iran. Curr. Microbiol. 74 (4),
522–531. https://doi.org/10.1007/s00284-016-1188-4.
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growth and aflatoxin production. Phytochemistry 66 (24), 2851–2856. https://doi.
org/10.1016/j.phytochem.2005.09.029.
Salmond, G.P., Reeves, P.J., 1993. Membrance traffic wardens and protein secretion in
Gram-negative bacteria. Trends Biochem. Sci. 18 (1), 7–12. https://doi.org/
10.1016/0968-0004(93)90080-7.