Review

Biocontrol Activity of Aromatic and Medicinal Plants and Their

Bioactive Components against Soil-Borne Pathogens
Babett Greff 1, András Sáhó 2,3, Erika Lakatos 1 and László Varga 1,*

1 Department of Food Science, Albert Casimir Faculty at Mosonmagyarovar, Szechenyi Istvan University,
15-17 Lucsony Street, 9200 Mosonmagyarovar, Hungary
2 Wittmann Antal Multidisciplinary Doctoral School in Plant, Animal, and Food Sciences, Szechenyi Istvan

University, 2 Var Square, 9200 Mosonmagyarovar, Hungary

3 Kisalfoldi Agricultural Ltd., 1 Fo Street, 9072 Nagyszentjanos, Hungary

* Correspondence: varga.laszlo@sze.hu

Abstract: Soil-borne phytopathogens can have detrimental effects on both cereal and horticultural
crops resulting in serious losses worldwide. Due to their high efficiency and easy applicability,
synthetic pesticides are still the primary choice in modern plant disease control systems, but
stringent regulations and increasing environmental concerns make the search for sustainable
alternatives more pressing than ever. In addition to the incorporation of botanicals into agricultural
practices, the diversification of cropping systems with aromatic and medicinal plants is also an
effective tool to control plant diseases through providing nutrients and shaping soil microbial
communities. However, these techniques are not universally accepted and may negatively affect
soil fertility if their application is not thoroughly controlled. Because the biocontrol potential of
aromatic and medicinal plants has been extensively examined over the past decades, the present
study aims to overview the recent literature concerning the biopesticide effect of secondary
metabolites derived from aromatic and medicinal plants on important soil-borne plant pathogens
including bacteria, fungi, and nematodes. Most of the investigated herbs belong to the family of
Lamiaceae (e.g., Origanum spp., Salvia spp., Thymus spp., Mentha spp., etc.) and have been associated
with potent antimicrobial activity, primarily due to their chemical constituents. The most frequently
tested organisms include fungi, such as Rhizoctonia spp., Fusarium spp., and Phytophthora spp.,
Citation: Greff, B.; Sáhó, A.; Lakatos, which may be highly persistent in soil. Despite the intense research efforts dedicated to the
E.; Varga, L. Biocontrol Activity of development of plant-based pesticides, only a few species of aromatic herbs are utilized for the
Aromatic and Medicinal Plants and
production of commercial formulations due to inconsistent efficiency, lack of field verification,
Their Bioactive Components against
costs, and prolonged authorization requirements. However, recycling the wastes from aromatic and
Soil-Borne Pathogens. Plants 2023,
medicinal plant-utilizing industries may offer an economically feasible way to improve soil health
12, 706. https://doi.org/10.3390/
and reduce environmental burdens at the same time. Overall, this review provides comprehensive
plants12040706
knowledge on the efficiency of aromatic herb-based plant protection techniques, and it also
Academic Editors: Despina Vokou highlights the importance of exploiting the residues generated by aromatic plant-utilizing sectors
Received: 31 December 2022 as part of agro-industrial processes.
Revised: 30 January 2023
Accepted: 2 February 2023 Keywords: biopesticide; aromatic plant; herb; soil-borne pathogen; essential oil
Published: 5 February 2023

Copyright: © 2023 by the authors. 1. Introduction

Licensee MDPI, Basel, Switzerland. Continuously increasing human population, ever-growing demand for food, and
This article is an open access article agricultural intensification have been placing a tremendous pressure on soil health,
distributed under the terms and
leading to degradation and exhaustion of agricultural lands and declining yields and
conditions of the Creative Commons
productivity [1–4]. The evaluation of soil health and quality is a critically important step
Attribution (CC BY) license
in maintaining long-term agricultural sustainability [5,6]. Healthy soil is characterized by
(https://creativecommons.org/license
stability, continuous nutrient cycling, stress resilience, richness, evenness, abundance,
s/by/4.0/).
and diversity of microbial communities [7,8]. The interactions between plants and soil-

Plants 2023, 12, 706. https://doi.org/10.3390/plants12040706 www.mdpi.com/journal/plants

Plants 2023, 12, 706 2 of 27

dwelling microorganisms are important drivers of the agroecosystem, as these indigenous

microbes can control the development, defense, tolerance, and nutrition of plants [9–11].
However, the composition and abundance of microbial communities are highly
influenced by various parameters such as the type of cultivated plants, environmental
fluctuations, and conducted agricultural activities [7,10,12].
Reduced plant productivity and phytopathogen-related disease outbreaks can be
considered as indicators of soil instability, poor ecosystem health, and a lack of microbial
diversity [7,8]. Soil-borne pathogens including viruses, bacteria, fungi, protozoa, and
nematodes are well known for their socio-economic and ecological impacts on the
ecosystem by causing serious losses in the agro-industry [13–16]. Plant roots and their
exudates provide substrates and space for these phytopathogens leading to colonization
and infection. Generally, the eradication of these organisms is especially hard in
continuous-cropping systems, due to the intensive tillage and cultivation processes [7,17].
Both agricultural practices and directed agronomical measures can affect the
diversity of soil microbial communities [18]. Traditionally, crop rotation techniques had
been used by farmers to manage the productivity of soil and reduce the possibility of crop
diseases [19]; however, after World War II, shortened rotation periods, intensive tillage,
monocropping, and application of synthetic pesticides and fertilizers came to the fore [20].
Although numerous cultural, physical, biological, and genetic options are available
nowadays, the main strategies for controlling soil-borne pathogens are still highly
dependent on the use of agrochemicals [15,21] due to their high efficiency and easy
applicability [22]. Besides the positive effects, the indiscriminate use of these chemical
agents may induce undesirable changes to the beneficial soil microorganisms and human
and animal health [22–24]. The commonly applied chemical pesticides are usually
effective against their target organisms, but they are not specific for them [25]. According
to Ashraf and Zuhaib [26], only an estimated 0.1% of the used synthetic chemicals reach
the targeted pathogens, whereas the remaining 99.9% may contaminate the environment
and endanger nontarget organisms.
The application of many chemical agents has been restricted in certain parts of the
world [27]. Therefore, finding sustainable and safer solutions with minimum or no side
effects is essential to develop effective soil health management strategies and lower the
possible environmental pollution [23,28–30].
In recent years, there has been an increasing interest in the application of
allelochemicals isolated from higher plants in plant protection because they are safer than
their synthetic counterparts, effective at reduced doses, and biodegradable without
leaving harmful residues [31–34]. The use of pathogen-suppressing secondary metabolites
(e.g., alkaloids, terpenoids, phenols) is especially important in organic farming systems
[33,35], but conventional cultivation methods and agricultural practices utilizing aromatic
plants and their residues (i.e., solid biomass and hydrosols) may also be effective in
controlling the spread and severity of plant diseases (Figure 1).
This paper summarizes and reviews (a) the most important soil-borne pathogens
including bacteria, fungi, nematodes, and viruses; (b) the current status of botanical
pesticides; and (c) the beneficial effects of aromatic and medicinal plant-based cultivation
techniques on soil and plant health. Moreover, the reutilization of by-products generated
by aromatic plant-based industries as biocontrol agents is also discussed.
Plants 2023, 12, 706 3 of 27

Figure 1. Aromatic and medicinal plants as part of non-chemical plant protections systems: effects
and methods of application. The icons in circles were made by Blackwoods, Freepik, Kerismaker,
and Surang (www.flaticon.com). The plant illustration on the left is based on the work of Mariya
Kazakova (purchased from www.shutterstock.com).

2. Phytopathogens in Soil
Soil biology is in direct connection with agricultural sustainability [36], as healthy
soil can recycle nutrients, decompose organic matter, support biological activities,
suppress the growth of pathogens, and inactivate toxic compounds [37]. It is the habitat
of a myriad of organisms (e.g., invertebrate animals, plants, protozoa, fungi, and bacteria)
that play an important role in maintaining agro-ecosystem functions and productivity
[38,39]. As for microbial diversity, only a few grams of soil can contain millions of species
[40]. In addition to beneficial microbes, however, major and minor pathogens may also be
present [15,41]. Even though the occurrence of microbial plant diseases is a natural part
of crop production, the excessive accumulation and epidemic spread of phytopathogens
can induce serious crop damages and yield losses [12,36].
Soil-borne pathogenic fungi, bacteria, protozoa, viruses, and plant parasitic
nematodes can cause negative plant–soil feedback individually or collectively [42,43].
They usually affect the root system or the stem base of plants, but vascular diseases caused
by these organisms have also been reported [44]. Overall, approximately 15% of total crop
production is estimated to be lost to different biological threats annually [45], but microbe-
induced diseases may even lead to total crop failure [46].

2.1. Fungi
As far as fungal pathogens are concerned, infections can be transmitted through soil-
damaging agricultural and horticultural products [47]. Fusarium, Verticillium, Rhizoctonia,
Sclerotinia, Phytophthora, and Pythium species may survive in the soil and in plant residues
for longer periods by forming resistant structures such as sclerotia, microsclerotia,
oospores, chlamydospores, or hyphae [22,48]. Several of them can invade the host plant
through roots and stems or spread rapidly among the seedlings [49] causing root necrosis
Plants 2023, 12, 706 4 of 27

and vascular diseases, as well as rot, gall, and proliferation of roots and tubers [44]. Recent
studies have highlighted that the causal agents of root rot (Fusarium solani and Fusarium
oxysporum f. sp. dahliae-lycopersici) and wilting (Verticillium dahliae) are able to infect a
considerable range of crop plants, making the controlling process extremely difficult
without high-cost fumigants [50]. For instance, Verticillium dahliae can survive in the soil
for up to 14 years by forming microsclerotia and may affect more than 160 plant species
[51]. Furthermore, fungi belonging to the genera Pythium and Phytophthora may cause sim-
ilar symptoms in the crown, stem, and root tissues of certain crops leading to scars; girdled
stem; stunt; stem lesions; foliar blight; browning; wilting; decay; damp-off; loss of root
density; and rot of crown, roots, and fruits [52,53].

2.2. Bacteria
Along with fungal strains, important bacterial pathogens can be found in the soil
including Agrobacterium, Pectobacterium, Pseudomonas, Ralstonia, and Xanthomonas species
[36,54]; however, bacterial pathogens are less likely to cause plant diseases, as they need
a wound or a natural opening to invade the host plant [16]. Ralstonia solanacearum is a
causal agent of bacterial wilt disease in various crops, including potato, banana, tomato,
and peanut. The bacteria enter the plant through wounds, cracks, or root tips; trigger mor-
phological alterations in the root system of the infected host; invade the xylem vessels;
and cause wilting symptoms and plant death [54,55]. Pseudomonas syringae and Pectobac-
terium carotovorum are also important soil-borne bacteria inducing necrotic lesions on to-
mato and carrot soft root, respectively [56]. Pectobacterium atrosepticum is mainly respon-
sible for various field symptoms of potato, including reduced emergence, wilting, chloro-
sis, stem and tuber rot, haulm desiccation, blackleg, and plant death [57]. Xanthomonas
campestris pv. musacearum may infect banana plants through injured roots and stems. Since
no remedy is available against this pathogen, the infected plant may be cut down, and
fallowing or prolonged crop rotation technique may be introduced [58].

2.3. Nematodes
So far, over 4100 species of plant-parasitic nematodes have been identified. They
cause serious economic losses globally (approximately USD 173 billion a year [59]) due to
their wide variety of interactions and host range. These small roundworms can be catego-
rized as ectoparasitic and endoparasitic species [59–66]. Among them, the sedentary en-
doparasitic nematodes [66] are the most economically important species. The temperate
Meloidogyne hapla and the tropical Meloidogyne incognita, Meloidogyne arenaria, and Meloido-
gyne javanica belong to the most destructive root-knot nematodes that can infect almost all
vascular plant species [61]. The parasitism of these biotrophic organisms includes the es-
tablishment of permanent feeding sites in the root cortex, vascular parenchyma, endo-
dermis, and pericycle of host plants. The intensified metabolic activity in the multinucle-
ate giant cells mobilizes photosynthates and, consequently, decreases plant quality and
quantity with typical symptoms such as suppressed plant growth, wilting, leaf discolora-
tion, and root deformation [61,67]. Southern knot-nematodes may also interact with soil-
borne fungi resulting in disease complexes [62]. Regarding cyst nematodes, potato, cereal,
and soybean cyst nematodes have been identified as the most significant obligate bio-
trophs that may be highly persistent in the soil and can survive for longer periods as eggs
inside a cyst without a host making their eradication almost impossible [61,68]. In addi-
tion, ectoparasitic nematodes are an integral part of the soil fauna [69] moving freely be-
tween root-feeding sites. However, they are more likely exposed to various environmental
conditions and predators. Similarly to endoparasites, they cause localized damage weak-
ening the host plant defense against bacterial and fungal infections [70], whereas certain
species, including Xiphinema index, may act as a vector of viruses [63]. Currently, various
methods (e.g., use of chemical agents, bio-fumigants, sanitation, resistant species, solari-
zation, steaming, tilling, fallowing, escape cropping, cover cropping with trap crops, crop
Plants 2023, 12, 706 5 of 27

rotation with poor or non-hosts, soil amendments, biological control by natural antago-
nists, etc.) are employed to reduce nematode infestation in the soil [71]. Although chemical
nematicides are efficient for the control of root-knot nematodes, their use has been grad-
ually decreased due to their toxicity, limited availability in developing countries, high
costs, and impaired efficacy after repeated applications [72,73].

2.4. Viruses
Soil may contain enormous amounts of soil-borne plant viruses with an estimated
viral abundance of 1031 [74]. The annual crop losses caused by these tiny organisms are
estimated at USD 60 billion worldwide. Currently, more than 2000 species have been iden-
tified that affect economically important crops including barley, groundnut, wheat, sugar
beet, potato, and fruit crops [42,75]. The first report was released in 1925 in connection
with a soil-borne virus, the soil-borne wheat mosaic virus, and further 53 plant pathogens
belonging to 12 genera have been recognized as soil-borne viruses in the last decades [27].
They are generally transmitted via soil, fungi, nematodes, plasmodiophora, insects and
other arthropods, sap transmission, seed and pollen transmission, mechanical friction,
and vegetative propagation materials [42,74]. In general, they are extremely persistent [42]
just as with beet soil-borne mosaic virus that is spread by a soil-borne plasmodiophoro-
mycete, Polymyxa betae, which generates resting spores allowing the virus to survive in
the soil for decades [76]. Although different viruses have different host ranges, collec-
tively, they can infect almost all kinds of cultivated and noncultivated plants, and the in-
fections can be transmitted from the affected plants to the healthy ones [75]. As an exam-
ple, the tobacco mosaic virus is a common viral disease that affects over 1000 species in 85
plant families [75,77]. The remains of infected plants in the soil are the principal reservoirs
of the virus. Transmission mostly occurs when leaves are rubbed in the presence of virus-
containing soil or when injured root hairs come into contact with infected residues or free
virus [78]. Furthermore, tomato spotted wilt virus is capable of infecting more than 1000
plant species (e.g., ornamental plants, lettuce, pepper, potato, etc.) [79]. Since the limita-
tion of viral infections and the eradication of viruses from infected soil are almost impos-
sible, control methods have mostly relied on various agrochemicals that can reduce the
vector population, resistant plant varieties, and other preventive measures [27,42,80].

3. Non-Chemical Plant Protection Methods

The lack of effective pest control is still a major global obstacle to improved plant
health and productivity [81]. Synthetic agrochemicals have been widely used to control
soil-borne pathogens [21], but the regulations on the application of these products have
become more stringent because of their adverse effects [82]. After the implementation of
the Montreal Protocol, EU regulations EC 2037/2000 and EC 3093/1994 completely prohib-
ited the use of methyl bromide, the most popular fumigant against soil-borne bacteria,
fungi, and nematodes. Furthermore, European Directive 2009/128/EC on the sustainable
use of pesticides was issued to reduce the risk of pesticides to the environment and human
health [22,28,83–85], and European Regulation no. 1107/2009 has been controlling the
placement of plant protection products on the market [86]. This led to the prohibition of
several traditional pesticides [87] and fueled the search for other suitable candidates, in-
cluding existing and new agrochemicals and more sustainable, economically feasible al-
ternatives that are less dangerous to the environment [88,89]. However, these methods
must meet various standards regarding pest specificity, toxicity, pesticide resistance, cost,
and availability [90].
Organic farming is a rapidly developing more sustainable method that improves soil
quality [38,91,92], thereby lowering the risk of negative environmental impacts such as
ecosystem degradation and global warming [93]. This form of agricultural production
aims to utilize non-chemical disease prevention tools rather than other treatments [92].
For instance, ecofriendly farming systems rely on the use of different physical, cultural,
Plants 2023, 12, 706 6 of 27

and biological management techniques to suppress the spread of pathogenic microorgan-

isms [94]. These methods, with only a few exceptions, do not pollute the environment [95].
Nevertheless, the activity spectrum of non-chemical farming strategies is extremely vari-
able, and they are often less effective than their chemical analogues [96,97]. Therefore,
successful disease control may require the implementation of multiple methods [98]. Yoon
et al. [79] used soil-dwelling predatory mites and an essential oil mixture to control tomato
spotted wilt virus and thrips. Likewise, Schmitt and Seddon [99] reported that the simul-
taneous use of microbiological control agents and plant extracts may have an additive or
even a synergistic effect. Meanwhile, Baysal-Gurel et al. [48] showed that cover crops in
combination with solarization significantly reduced Rhizoctonia root rot severity.
As another option, Integrated Pest Management (IPM) is a redesigned form of inten-
sive agricultural systems, in which non-chemical methods, botanicals, or other curative
treatments are combined with synthetic chemicals [35]. Just as with organic production,
the goal is not to fully eliminate disease-causing organisms but to prevent them from be-
coming damaging or dominant in the cropping system. As a result, an integration of dif-
ferent preventive tactics (i.e., chemical, biological, physical, and genetic means with crop-
ping practices) may be more effective than a single method [100,101]. A survey conducted
by Kabir and Rainis [102] showed that vegetable farmers in Bangladesh used additional
methods (e.g., biological control, soil amendments, pheromone traps, manual cleaning,
soil solarization, and botanicals), while pure pesticides were only applied when there
were no other options, or the pest infestation was high. In a study by Fielding et al. [103]
eight medicinal plant extracts were applied successfully in combination with kresoxim-
methyl for the inhibition of gray mold on apples. Shlevin et al. [104] analyzed independent
studies to evaluate the efficacy of soil solarization with fumigants in controlling various
soil-borne pathogens (e.g., Fusarium species; root-knot nematodes; and the group of Scle-
rotium cepivorum, Verticillium spp., Pyrenochaeta spp., Rhizoctonia spp., and Pythium spp).
Overall, their results showed that the combined technique improved management effi-
cacy compared to soil solarization only. In contrast, Deguine et al. [101] reported that these
integrated methods were rarely adopted by farmers, and, as a result, pesticides remained
the cornerstone of crop protection around the world.

4. Aromatic Plants as Biocontrol Agents

There are approximately 17,500 aromatic plant species throughout the world, either
cultivated or collected in the wild, that are greatly used for cosmetic, preservation, flavor-
ing, therapeutic, and pharmacologic purposes [105–108]. These herbs are responsible for
the synthesis of a multitude of secondary metabolites (e.g., flavonoids, phenol com-
pounds, terpenes, and nitrogen-containing chemicals), which are formed from primary
compounds [47,109,110]. Some of these metabolites are produced and stored during the
normal growth of plants, whereas others accumulate only in response to stress conditions
[111].
Due to their strong bioactivity, aromatic plants and their constituents may be applied
as part of environmentally friendly management programs [64,112] to improve soil qual-
ity [113] and protect plants from pathogens and insects [114]. In a cropping system, aro-
matic plants and their bioactive metabolites can affect phytopathogens directly or indi-
rectly by inducing systemic resistance and defense responses, repelling viral vectors, or
inhibiting the growth of phytopathogens by changing the structure of soil microbial com-
munity [115–119]. Therefore, introduction of these noncrop biocontrol plants can affect
crop productivity [120], with the postextraction residues and other plant debris being fur-
ther sources of bioactive components that can be recycled as natural pesticides, organic
amendments or transformed into value-added products providing nutrients, organic mat-
ter, and beneficial microorganisms [64,121,122]. For these reasons, the following sections
provide background information on the applicability of aromatic and medicinal plants,
botanicals, postextraction residues, and composts derived from aromatic plant waste to
control the spread of major soil-borne pathogens.
Plants 2023, 12, 706 7 of 27

4.1. Use of Pure Extracts as Biopesticides

Microorganisms are considered to be the most important biopesticides in the fight
against plant diseases, whereas botanicals still represent a relatively small part of the mar-
ket. Prior to World War II, only four major components were commonly used, i.e., rote-
none and rotenoids, pyrethrum and pyrethrins, nicotine and alkaloids, and some vegeta-
ble oils [123], while other allelochemicals, such as derris and citronella oil, were applied
as insecticides [90]. Although their use declined following the commercialization of syn-
thetic agrochemicals due to their instability (pyrethrum) and toxicity to nontarget species
(rotenone and nicotine) [123,124], the multitudinal backlash against synthetic compounds
eventually led to the prosperity of biopesticides of plant origin [125].
The group of botanicals includes various plant volatiles, extracts, and natural oils that
are characterized by improved biocompatibility, little or no cross-resistance to commercial
chemical agents, and structural diversity [126,127]. Generally, these biologically active
substances are extracted or distilled from fresh or dried plant materials with classical
methods using water, alcohol, and other solvents [127]. To decrease the environmental
impact, green processes requiring no or low amounts of solvents and less energy and time
have also been developed including subcritical water extraction, pressurized ultrasound
and microwave extraction, and supercritical fluid extraction [128–130].

4.1.1. Essential Oils

Essential oils (EOs), biosynthesized, stored, and secreted in various parts of aromatic
herbs, e.g., buds, bark, leaves, zest, twigs, wood, stems, seeds, fruits, rhizome, roots, and
flowers [128,131–133], are complex mixtures of terpenoids and phenolic compounds that
play a critical role in plant defense mechanisms against abiotic and biotic stress factors
[128,132,134]. These volatile organic compounds can diffuse through water- and gas-filled
pores and serve as food sources, infochemicals, chemo-attractants, or antimicrobials [135].
Monoterpenes, as frequent constituents of EOs, are involved in chemical and ecological
interactions, and their production may be induced by the attack of pathogens and herbi-
vores [136,137]. Due to their bioactive potential, EOs and their constituents can be utilized
as components of ecological products with nematicidal, antimicrobial, antiviral, insecti-
cidal, repellent, antifeedant, and molluscicidal activity [23,118,138]. They can also support
the growth of other soil microbes indirectly by killing organisms and providing easily
decomposable carbon and energy sources [139]. This was confirmed by [140], who found
that Mentha × piperita EO had an instant antimicrobial effect on soil; however, a fraction of
soil microbes survived the treatment and used the killed microorganisms as fresh organic
carbon. As a result, an increased microbial biomass and respiration rate were recorded.
As for the efficiency of active substances derived from aromatic and medicinal plants,
EOs tend to possess stronger antimicrobial properties than other extracts [51]. Although
several studies investigated the biocidal effect of EOs and their constituents on phytopath-
ogens [46,141,142], the mechanism of their action is complex and not fully understood
[143,144]. Volatile oils containing oxygenated monoterpenes, primarily phenols and alde-
hydes, tend to be more active against pathogens [145,146]. This was confirmed by
Wogiatzi et al. [147], who found that the thymol-rich EO of Origanum vulgare collected
from the mountain Olympus in Greece had the strongest antifungal effect on Verticillium
dahliae, Pythium spp., and Sclerotinia sclerotiorum. Conversely, the hydrocarbonic fraction
of Salvia pomifera L. ssp. calycina (Sm.) Hayek was less effective against Sclerotinia scleroti-
orum and Rhizoctonia solani than was the oxygenated fraction [148].
The application of EOs can provoke chemical, physical, and biochemical changes in
the cell [149] including disruption of the permeability barrier and proton pump function,
damage of cytoplasmic membrane, increased permeability and leakage of cell contents,
decreased ATP synthesis and augmented ATP hydrolysis, cytoplasm coagulation, re-
duced membrane potential, and inhibition of the production of toxic microbial metabo-
lites [133,150]. As regards fungi, EOs can inhibit ergosterol biosynthesis leading to the
Plants 2023, 12, 706 8 of 27

depletion of sterol content in the cell membrane, disruption of cell membrane integrity
and permeability, loss of ions, and, eventually, inhibition of fungal growth [151]. The
study of Upadhyay et al. [151] has shown that a decreased level of methylglyoxal may
also be observed that was reported to be in correlation with aflatoxin B1 production in
Aspergillus flavus. In addition, Vokou et al. [139] claimed that volatile oils extracted from
Satureja thymbra could reduce the spore germination and mycelial growth of Penicillium
citrinum and Mucor hiemalis, respectively. Another study conducted by Sempere-Ferre et
al. [152] showed that EO constituents, either alone or in combination, inhibited the myce-
lial growth of Botryotinia fuckeliana and Rhizoctonia solani, although eugenol exerted a fun-
gistatic effect only. Similarly, testing the antifungal efficacy of various EO constituents,
Marei et al. [153] found that 1,8-cineole, (R)-camphor, (R)-carvone, camphene, cuminalde-
hyde, (R)-linalool, geraniol, (1R,2S,5R)-menthol, (S)-fenchone, myrcene, thymol, and (S)-
limonene were promising substances against Fusarium oxysporum, Penicillium digitatum,
Aspergillus niger, and Rhizoctonia solani. Thymol and (S)-limonene possessed strong inhib-
itory effect on cellulase and pectin methyl esterase activities. Lee et al. [154] stated that
Origanum vulgare EO had a broad antifungal spectrum against both postharvest (Botrytis
cinerea, Colletotrichum gloeosporoides) and soil-borne pathogenic fungi (Fusarium oxysporum,
Rhizoctonia solani, and Pythium ultimum). Bi et al. [155] reported that a 21-day-long soil
treatment with EOs from palmarosa (Cymbopogon martini), oregano (Origanum syriacum),
and red thyme (Thymus vulgaris) at concentrations of ≥0.1 μg/mL decreased the population
density of Phytophthora capsici below the limit of detection.
As nematicides, EOs can reduce gall formation, hinder the hatching of nematode
eggs, immobilize juveniles, and, as a result, completely suppress nematode infestation
[156]. This is consistent with the findings of Ntalli et al. [65], who reported the EOs of
Origanum vulgare, Origanum dictammus, Mentha pulegium, and Melissa officinalis to be effec-
tive against Meloidogyne incognita by irreversibly paralyzing the larvae (Table 1). In fact,
the oxygenated EO constituents were more active than hydrocarbons, including β-caryo-
phyllene, p-cymene, and limonene.
Overall, the studies presented thus far provide evidence that the biological activity
of EOs is usually strongly related to their chemical composition; the synergistic effect of
different bioactive components [65,148]; specific seasonal, geographic, and climatic con-
ditions [64]; genetic factors; and harvest and postharvest processes [157].

Table 1. Properties of essential oils and/or essential oil constituents, dealt with in this review paper,
against soil-borne organisms and related references.

Refer-
Essential Oil (EO) Target Organism Effect
ence
Thymus pallescens, Artemisia herba-alba Inhibition of mycelial
Fusarium oxysporum f. sp. ciceris
Asso, Laurus nobilis L., Cymbopogon growth, sporulation, and [46]
Padwick
citratus (de Candolle ex Nees) Stapf. spore germination
Mentha piperita L., Thymus vulgaris L., Inhibition of mycelial
Verticillium dahliae Kleb. isolates [51]
Lavandula angustifolia Mill. growth
High nematicidal activ-
Origanum dictamnus, Mentha pulegium, ity (irreversible motility
Meloidogyne incognita [65]
Origanum vulgare, Melissa officinalis loss of nematode juve-
niles)
Suppression of spore
Satureja thymbra, Rosmarinus officinalis Penicillium citrinum [139]
germination
Complete inhibition of
Satureja thymbra Mucor hiemalis [139]
mycelial growth
Inhibition of mycelial
Mentha spp. Penicillium expansum [141]
growth
Plants 2023, 12, 706 9 of 27

Alternaria alternata, Aspergillus spp.,

Strong antifungal activ-
Penicillium spp., Trichoderma viride,
Origanum onites L., Salvia thymbra L. ity against the microor- [142]
Cladosporium cladosporioides,
ganisms tested
Phomopsis helianthi
Verticillium dahliae, Fusarium ox-
Inhibition of mycelial
Origanum spp. ysporum f. sp. lycopersici, Sclerotinia [147]
growth
sclerotiorum, Pythium spp.
Fusarium oxysporum f. sp. dianthi,
Fusarium solani f. sp. cucurbitae,
Salvia pomifera L. ssp. calycina (Sm.) Inhibition of mycelial
Fusarium proliferatum, Verticillium [148]
Hayek growth
dahliae, Sclerotinia sclerotiorum,
Rhizoctonia solani
EO constituents (carvacrol, cinnamal- Inhibition of mycelial
Rhizoctonia solani [152]
dehyde, eugenol, thymol) growth
EO constituents [camphene, (R)-
camphor, (R)-carvone, 1,8-cineole,
Inhibition of mycelial
cuminaldehyde, (S)-fenchone, geraniol,
growth
(R)-linalool, (1R,2S,5R)-menthol, Aspergillus niger, Fusarium
myrcene] oxysporum, Penicillium digitatum, [153]
Rhizoctonia solani Inhibition of mycelial
growth, inhibitory effect
EO constituents [thymol, (S)-limonene]
on pectin methyl ester-
ase and cellulase
Cuminum cyminum, Eucalyptus Fusarium oxysporum, Pythium Inhibition of mycelial
[154]
citriodora, Origanum vulgare ultimum, Rhizoctonia solani growth
Strong antifungal activ-
ity,
control of production
and germination of spo-
rangia and zoospores,
Origanum syraicum, Cymbopogon mar- oospore production, and
Phytophthora capsici isolates [155]
tini, Thymus vulgaris mycelial growth
reduction of Phy-
tophthora capsici popula-
tion in soil,
suppression of infection
on zucchini fruits

4.1.2. Phenolic Compounds

Aromatic plants are rich sources of phenolic compounds (e.g., phenols, coumarins,
tannins, flavonoids, phenolic acids) derived from the phenylpropanoid metabolism and
shikimate pathway [158,159]. So far, around 8000 phenolics with highly diverse structures
have been reported in the literature [160,161]. They contain one or more aromatic rings
(C6) bearing at least one hydroxyl group [162,163]. These compounds are toxic to fungi,
bacteria, insects, nematodes, and weeds [164], but, unlike EOs, they are water-soluble be-
cause they mostly exist in glycosidic form in plants [165].
Although polyphenols were isolated from various plant parts [166], they are found
at higher levels in the outer layers of tissues [167] as the production of these metabolites
can be induced by both biotic and abiotic factors such as reactive oxygen and nitrogen
species, insufficient amounts of nutrients, low temperatures, UV light, predators, para-
sites, pathogens, wounding, and other stress conditions [163,168,169]. Nevertheless, the
Plants 2023, 12, 706 10 of 27

antimicrobial potential of polyphenols has not been fully deciphered [170] due to the com-
plexity of their natural mixtures in plants [171] and their inconsistent activity [172].
Among them, flavonoids are associated with the stronger biocidal effects, whereas non-
flavonoid compounds usually show a weak antagonistic behavior against pathogens
[173]. Their activity, however, depends not only on their purity, structure, and concentra-
tion but also on the tested microbial strains, solvents, and experimental conditions [172].
Natural phenolics combine seemingly antagonistic effects, including an antioxidant
activity, with variable degrees of cytotoxicity [174]. The mode of action of phenolics in-
volves modification of cell membrane permeability and changes in intracellular functions,
such as [170] inhibition of efflux pump and enzymatic (i.e., DNA gyrase, beta-ketoacyl
acyl carrier protein synthase II and III, FabG, FabZ, FabI, d-alanine–d-alanine ligase, ure-
ase, sortase A, dihydrofolate reductase, etc.) activities, leakage of intracellular constitu-
ents, disturbance of cell wall biosynthesis, and inhibition of bacterial biofilm formation
[175,176]. Important phenolics can repress or induce the expression of genes playing im-
portant roles in the pathogenesis of soft-rot-causing pathogenic bacteria or induce genes
encoding efflux pumps in soft-rot enterobacteria [174]. As Summers and Felton [177]
stated, phenolics could result in reduced protein digestibility, impaired enzymatic func-
tions, and reduced bioavailability of amino acids by forming complexes with proteins. In
addition, Simmonds [178] mentioned that the ingestion of phenolic compounds can re-
duce the nutritive value of the food consumed by nonadapted insects. This effect may be
due to the induction of oxidative stress to the digestive system [177]. Widmer and Laurent
[179] reported that lavender (Lavandula angustifolia), lavender hybrid (Lavandula angustifo-
lia × Lavandula spicata), and rosemary (Rosmarinus officinalis) leaf extracts reduced the ger-
mination of Phytophthora capsici, Phytophthora megakarya, and Phytophthora palmivora zoo-
spores. The aqueous extract of Acacia saligna H. L. Wendl. containing benzoic acid, caf-
feine, o-coumaric acid, naringenin, quercetin, and kaempferol effectively inhibited the
mycelial growth of Fusarium culmorum, Penicillium chrysogenum, and Rhizoctonia solani, es-
pecially at high concentrations (Table 2). At the same time, the minimal inhibitory con-
centration of the extract was 200, 300, 300, and 100 μg/mL against Agrobacterium tumefa-
ciens, Enterobacter cloacae, Erwinia amylovora, and Pectobacterium carotovorum subsp. caroto-
vorum, respectively [180]. The filter-sterilized water extracts of Origanum vulgare L.,
Melissa officinalis L., and Salvia officinalis L. shoots (0.5 and/or 2.0% w/v) suppressed the
propagation of Fusarium oxysporum f. sp. asparagi [181]. Similar findings were reported by
Ahmad and Matsubara [182], who demonstrated that the aqueous extract of Thymus vul-
garis L. (0.5 and 2.0%) had the maximum suppression effect on Fusarium oxysporum f. sp.
cyclaminis. Caffeic acid and rosmarinic acid, as the main constituents, also exhibited a
stronger antifungal activity. To induce the defense responses of soybean and sorghum,
Colpas et al. [112] used the aqueous extracts of Ocimum gratissimum leaves, which caused
an instant increase in phytoalexin production. López et al. [183] demonstrated that the
dichlormethanolic extracts of Melissa officinalis L. subsp. officinalis, Mentha longifolia (L.)
Hudson, Origanum vulgare L. subsp. virens Bonnier & Layens, and Salvia pratensis L., as
well as the ethyl acetate extract of Mentha × piperita L., Salvia pratensis L., and Thymus prae-
cox Opiz subsp. Polytrichus inhibited the growth of Rhizopus stolonifer over 20% under in
vitro conditions. The nematicidal activity of aromatic plant extracts was demonstrated by
Ntalli et al. [64]: Origanum vulgare L. and Thymus citriodorus (Schreb) extracts were effec-
tively utilized against Meloidogyne javanica (Treub) and Meloidogyne incognita (Kofoid and
White). However, elevated concentrations of the active constituents may have an inhibi-
tory effect on physiological processes of crop plants by increasing electrolyte leakage from
the cell membrane of seedlings and negatively affecting seed germination, chlorophyll
synthesis, and the functioning of photosystem II [184].

Table 2. Properties of extracts from aromatic and medicinal plants, dealt with in this review paper,
against soil-borne organisms and related references.
Plants 2023, 12, 706 11 of 27

Type of the Refer-

Plant Extract Target Organism Effect
Extract ence
Mentha piperita L., Thymus
Verticillium dahliae Kleb. iso- Inhibition of mycelial
vulgaris L., Lavandula angustifolia - [51]
lates growth
Mill.
Hydrosol
Orginum vulgare from EO Meloidogyne javanica
Nematostatic/nemati-
distillation
cidal activity (paraly-
Water ex- [64]
sis of second-stage ju-
Orginum vulgare, Thymus citriodo- tract of Meloidogyne incognita,
veniles)
rus plant pow- Meloidogyne javanica
ders
Rosmarinus officinalis, Lavandula
Hot water
angustifolia, Lavandula angustifolia Control of zoospore
extract of Phytophthora spp. isolates [179]
× Lavandula spicata, Salvia germination
leaves
officinalis
Water ex- Fusarium culmorum,
Inhibition of mycelial
Acacia saligna tract of Rhizoctonia solani, Penicillium [180]
growth
flowers chrysogenum
Water ex- Agrobacterium tumefaciens,
Inhibition of bacterial
Acacia saligna tract of Pectobacterium carotovorum [180]
growth
flowers subsp. carotovorum
Water ex-
Origanum vulgare L., Salvia Fusarium oxysporum f. sp. as- Suppression of fungal
tract of [181]
officinalis L., Melissa officinalis L. paragi propagation
shoots
Suppression of fungal
Water ex- pathogen,
Fusarium oxysporum f. sp.
Salvia officinalis L. tract of reduction of disease [182]
cyclaminis
shoots indices in cyclamen
roots and shoots

4.1.3. Applicability of Pure Metabolites in Plant Protection

Plant-derived biopesticides are inexpensive and readily available materials, which
do not contaminate the environment due to their biodegradable nature [47,127]. Despite
the extensive body of scientific literature and the tremendous amount of experience that
support the bioactivity of botanicals on plant pathogens, only a few botanical pesticides
are available and utilized in agriculture due to the lack of field verification [127,185]. In
Hungary, for instance, the following plant-based pesticides and plant conditioners have
been approved: Green NanoherbTM and Biomit® with a blend of EOs, DU-OLTM with lav-
ender and lemon oil, ORGANICTM with orange oil, and HerbalTM containing herbal ex-
tracts [186,187].
Although several plant-based protectants, e.g., spearmint oil and clove oil, have even
been included in the pesticide database of the European Union and approved by the Eu-
ropean Commission [149,188], the commercialization of botanical extracts has been lim-
ited by regulatory policies, costs, and prolonged procedures [47]. These bioactive sub-
stances can hinder the growth of soil-borne pathogens with a relatively mild effect on the
nontarget species [35]. However, they are prone to decompose rapidly, whereas moisture,
air, or sunlight may affect their stability, efficiency, and shelf-life. In addition, there are
further drawbacks that can complicate the applicability of botanical extracts in the field,
such as their nonselective nature, varying performance, and unavailability during certain
parts of the growing season, as well as the absence of legislation; established residue tol-
erance; scientific validation; and sufficient data on stability, quality, and effect on human
Plants 2023, 12, 706 12 of 27

health [127]. The applicability of volatiles or other extracts may be also compromised by
their dose-dependent properties [53,184].
Overall, plant-based bioactive substances may be integrated into common practices
to reduce the use of conventional chemical treatments [127]. Formulation with emulsify-
ing agents, polymers, surfactants, stabilizers, defoamers, solvents, and other substances
can be used to ensure their stability, improve their efficiency, and control the release of
biologically active components under field conditions [114].

4.2. Aromatic Plants as Part of Agricultural Practices: Cultivating Aromatic Plants and
Recycling the Waste as Green Manure and Compost
The aerial parts and roots of aromatic plants and their waste can affect the composi-
tion of microbial populations in soil by releasing complex mixtures of chemicals (Figure
2). These secondary metabolites, excreted directly into the soil from the living and decom-
posing tissues [109,189], may inhibit the growth of phytopathogens [190].

(a) (b)
Figure 2. (a) Post-extraction lavender (Lavandula angustifolia) waste; (b) Antifungal effect of lavender
waste extracts on phytopathogenic Sclerotinia sclerotiorum. Photographs taken by Babett Greff.

Over the last decades, various forms of these herbs including cultivated plants, green
manure, compost, and wastes from EO extraction (plant residues and hydrosol) have been
tested as potential non-chemical treatments against soil-borne plant pathogens. In the next
subsections, the potential effects of aromatic plants, their postextraction wastes, and com-
post on phytopathogens will be discussed.
Plants 2023, 12, 706 13 of 27

4.2.1. Diversifying Cropping Systems with Aromatic and Medicinal Plants

As continuous cropping systems have been constrained by various biotic and abiotic
factors (i.e., reduced soil quality, autotoxicity of the crop, shifts in microbial communities,
reduced microbial diversity and abundance, prosperity of plant pathogens, etc.) [19], ag-
ricultural diversity has become a critical part of sustainable agroecosystems [191]. Diver-
sification requires the combination of environmental and economic sustainability through
increasing crop species diversity (intercropping, crop rotation) or planting noncrops
(cover-cropping) [19,191–193]. These strategies may reduce production costs and improve
resource use and soil quality [192], while the highly diverse plant communities thus cre-
ated are less affected by phytopathogens [193]. However, the reduced activity of plant
pathogens in such communities depends both on host density and the effects of neighbor-
ing species on disease-causing organisms [43].
Under optimal conditions, the phytomanagement of soils through the cultivation of
aromatic and medicinal plant species can be a win-win approach that provides additional
profits and marketable products [194–196]. As a vegetation cover, herbs can increase the
fertility and the organic matter content of soils resulting in improved properties (e.g., wa-
ter retention, soil porosity, permeability, and bulk density) [105].
It is also believed that the microbiota around medicinal and aromatic plants is highly
plant-specific [196]. The rhizospheric zone in the vicinity of the plant roots is an attractive
environment for a large number of organisms because plants may release up to 20% of the
photosynthetically fixed carbon through their roots [119]. In addition to the typical root
exudates, such as sugars, organic acids, amino acids, and sterols, aromatic crops can re-
lease novel chemicals in the soil that may have a long-lasting effect on the processes arbi-
trated by the surrounding microorganisms. Although the production of plant secondary
metabolites is relatively low [110,116,197], volatiles can form a stable concentration gradi-
ent in the soil due to the lack of turbulence [109], and the build-up of pathogenic microor-
ganisms and other predators can thus be inhibited [191]. However, several rhizospheric
microorganisms have the potential to adapt and conduct endophytic or epiphytic life.
These microbes and the host plant will influence each other through biological and phys-
icochemical interactions [198]. For instance, the microorganisms associated with the roots
of aromatic plants take part in plant growth and nutrition, regulate disease interactions
[196], and may induce the production of certain bioactive compounds [117].
Over the past decades, several studies have confirmed the beneficial effects of aro-
matic herbs on soil microbial composition. Karthikeyan et al. [199] showed that the bacte-
rial, fungal, and actinomycete populations in the rhizosphere of Ocimum sanctum L. were
2.3 × 107, 1.9 × 105, and 1.2 × 106 CFU/g, respectively. These values were higher than the
corresponding ones observed in the rhizosphere of Coleus forskholii Briq. Similar results
were reported for diazotrophic bacterial populations (i.e., Azospirillum spp., Azotobacter
spp., and Pseudomonas spp.). Examining soil microbial populations, Adamović et al. [200]
observed an increase in the total number of microorganisms, cellulolytic microorganisms,
azotobacters, fungi, and free nitrogen-fixing microorganisms when mint (Mentha × piperita
L.) was planted. The cultivation of basil (Ocimum basilicum L.) had the same effect on the
total numbers of actinomycetes. Zhang et al. [113] showed that intercropping pear trees
with basil or summer savory (Satureja hortensis L.) could increase the organic matter con-
tent of soil and the activities of enzymes including invertase, urease, and catalase due to
the influence of root exudates on soil microbial diversity. Meanwhile, soil organic matter
was negatively correlated with pathotrophic groups of fungi, and, thus, intercropping sig-
nificantly reduced the relative abundance of pathotrophic and saprotrophic groups. Bais
et al. [111] reported that the fungal cell wall elicitors from Phytophthora drechsleri and Phy-
tophthora cinnamomi improved root growth of Ocimum basilicum L. and facilitated rosma-
rinic acid production in hairy root cultures. Former studies have demonstrated that both
EOs and aromatic water may possess antiphytoviral activity [80], but the physicochemical
properties of soil and root exudates can also influence the migration and location of these
organisms [74]. Due to their phytoremediation potential, several aromatic and medicinal
Plants 2023, 12, 706 14 of 27

plants are also used for the remediation of heavily contaminated soil by accumulating and
removing organic and inorganic contaminants [201].
It should be noted that consecutive monocultures of herbs may severely alter the bac-
terial and fungal populations in soil [202]. According to the research conducted by Tang
et al. [19], the continuous cropping of Salvia miltiorrhiza Bunge affected the microbial com-
position (e.g., fungal and actinomycete communities) of soil both in terms of structure
diversity and abundance. Overall, the productivity of these plants tends to decrease over
time because of the accumulation of their natural enemies [43] and the relatively low di-
versity of functional microbes [19].
Crop rotation is one of the best options to keep the levels of soil-borne pathogens and
weeds low, but, from an economical perspective, it is not always acceptable [44,203].
Cover cropping and intercropping are used more and more frequently in conventional
habitat management systems [138]. Aromatic herbs are excellent intercropping options for
different cultures due to their characteristic traits and resistance to adverse environmental
conditions [204]. Verma et al. [192] found that intercropping Pelargonium graveolens L. with
companion crops may improve the postharvest soil total Kjeldahl nitrogen; organic car-
bon; carbon-to-nitrogen ratio; and available N, P, and K contents. The cultivation of wheat,
oat, and barley also enhanced the biomass-specific respiration of soil that might be in as-
sociation with higher microbial activity. Khan et al. [194] reported that cropping treat-
ments with Ocimum basilicum L. cv. CIM-Saumya and Mentha arvensis L. cv. Kosi along
with crop residue retention showed reduced CO2-C emission and increased soil organic
carbon content compared to fallow soil. Chand et al. [205] showed that the cultivation of
Pelargonium graveolens, Rosmarinus officinalis, and Mentha piperita had a conservation func-
tion, and these herbs reduced soil erosion and slowed down the runoff. However, the
relative soil and water conservation efficiency of Thymus vulgaris was almost zero due to
its poor canopy coverage. The authors also suggested that geranium and rosemary were
good candidates for vegetative barriers, intercrops, and cover crops.
The plant material of herbs can be left on the top of the soil as mulch or incorporated
into green manure [206]. In this form, they can interfere with phytopathogens directly
through the release of toxic compounds or indirectly [207] by favoring the natural enemies
of soil-borne pathogens through habitat manipulation [35]. Furthermore, the supplemen-
tation of plant growth-promoting microorganisms associated with medicinal plants can
improve plant health by increasing their immunity to phytopathogens [208], improving
stress tolerance, and influencing nutrient availability and uptake, as well as the produc-
tion of growth-promoting metabolites and hormones, such as cytokinins and auxins [209].
Castronovo et al. [210] reported that microorganisms isolated from the bulk soil of Origa-
num vulgare L. exhibited antibacterial activity against various human pathogens. The Ag-
robacterium, Agromyces, Bacillus, and Chryseobacterium isolates inhibited the growth of sev-
eral Bacillus strains of environmental origin. Tiwari et al. [211] isolated potential plant
growth-promoting bacterial agents (Bacillus and Pseudomonas spp.) from the root vicinity
of medicinal aromatic plants such as Ocimum spp. Under greenhouse conditions, the iso-
lated bioinoculants alone or in combination with Trichoderma harzianum reduced the re-
production factor of Meloidogyne incognita by 46.4–72.3% in sterile and natural soils. Chow-
dhary and Kaushik [212] reported that approximately 12% of endophytic fungal isolates
from Mentha x piperita plants exhibited antifungal activity against three or more of the
tested phytopathogens.
In addition to the aforementioned beneficial effects on soil microbial composition,
aromatic plant-based cropping systems can provide significantly increased profits and
crop production rates [213] because herbs (e.g., Ocimum basilicum L.) are less prone to
compete with cultivated crops for nutrients [214,215]. Carvalho et al. [216] studied how
intercropping with aromatic plants, such as Ruta graveolens, can affect tomato yield. This
cultivation technique was found to increase the total yield of tomato fruits by 26%. In a
similar companion planting experiment, Ahmad et al. [214] observed that certain Lami-
aceae species (e.g., Ocimum basilicum L., Mentha piperita L., Hyssopus officinalis L.) promoted
Plants 2023, 12, 706 15 of 27

the growth of tomato plants and influenced the production and accumulation of certain
metabolites in the dominant crop. Peppermint and hyssop enhanced shikimic acid and
apigenin contents in all parts of tomato plants providing a product with improved quality
and yield. Likewise, cropping with basil resulted in higher free amino acid contents with
the leaves being rich in alanine, phenylalanine, iso-leucine, lysine, valine, and γ-amino-
butyric acid (GABA), whereas the stems contained elevated levels of valine, serine, ala-
nine, proline, GABA, and glutamine compared to the control treatment. In addition, the
1:1 tomato–aromatic plant companion setups boosted the growth of tomato plants with-
out the accumulation of competition pressure. Under certain circumstances, however,
temporal phytotoxic effects could be experienced due to the incorporation of aromatic
herb biomass. Nevertheless, other techniques (e.g., relay-planting) are available that can
optimize productivity [217].

4.2.2. Recycling the By-Products of Aromatic Plant-Utilizing Sectors

The demand for aromatic and medicinal plant-based products has increased contin-
uously, at the rate of 15–25%, for the last few years [218]. As the ratio between the pro-
cessed EO and the utilized plant material is low [219], large quantities of solid waste are
generated and remain unutilized [220]. From a sustainability perspective, the aromatic
biomass cannot be considered as waste [221], and it may be recycled as animal feed, bio-
sorbents, biogas, biofuel, chelating agents, biopesticides, or soil amendments [121,219].
Just as with the fresh plant material, the distilled solid biomass can contain volatile
components and phenolic compounds, albeit in different ratios [219,222]. According to
Kadoglidou et al. [33], the application of Mentha spicata L. and Origanum vulgare L. ssp.
hirtum plant material as soil amendments (4%) decreased the average degree of infection
(Fusarium oxysporun f. sp. lycopersici and Verticillium dahliae) of tomatoes. In addition to
plant disease reduction, the growth and physiology indices as well as soluble solids con-
tents and fruit yield were also affected. Moreover, the amendment of Mentha spicata L.
tissue at different concentrations may have a positive impact on tomato seedling emer-
gence and dry weight, the size of the most robust leaf, shoot length, physiological param-
eters (e.g., stomatal conductance, photosynthetic rate, and yield), and the size of fungal
and bacterial populations [223]. Klein et al. [224] used herbal biomass in combination with
soil solarization to control the spread of Fusarium oxysporum f. sp. radicis-lycopersici and
Meloidogyne javanica under farm conditions. The results showed that solarization with
rosemary (Rosmarinus officinalis L.), sage (Salvia officinalis L.), tarragon (Artemisia dracuncu-
lus L.), and thyme (Thymus vulgaris L.) residues was highly effective against the tested
fungi reducing their viability by 99–100%. As for the root-knot nematode, galling was re-
duced to zero in tomato roots in the solarized soil amended with sage and thyme residues.
Despite the positive effect of untreated herbal residues on soil productivity and disease
severity, their direct use may be compromised due to their unknown composition [157].
Furthermore, the introduction of novel plant metabolites may also affect indigenous mi-
crobial communities [116], critically the decomposition process [225], and endanger crop
growth and yield [203]. As an example of the negative impact, Gravanis et al. [226] re-
ported that the incorporation of dried Origanum vulgare biomass significantly decreased
the number of bacterial colonies in soil samples. Similar results were reported by Chouli-
aras et al. [227]. In contrast, Ainalidou et al. [228] showed that soil enrichment with the
green parts of Mentha spicata, Mentha piperita, and Rosmarinus officinalis increased the total
microbial biomass. However, the use of rosemary had a negative effect on the growth
indices of tomato seedlings, including weight, shoot length, and root length of fresh seed-
lings, as also reported by Argyropoulou et al. [229].
The meaningful utilization of aromatic biomass is still a great challenge. Composting
and vermitechnology are commonly applied to treat these agricultural wastes prior to use.
The mature and stable end product is a suitable soil amendment that enhances important
soil processes and features improving the diversity and activity of microbial communities
in soil [223,230]. Moreover, composts provide a new environmental source of microbes
Plants 2023, 12, 706 16 of 27

with antimicrobial activity [231]. Although their efficiency is variable, the microorganisms
involved in the decomposition of organic matter can suppress the growth of soil-borne
pathogenic microbes through antagonistic interactions, inducing systemic resistance in
the host plant or forming humic molecules and biostimulants [232]. Under glasshouse
conditions, the nematicide activity of vermicomposts containing Artemisia annua, Chrysan-
themum cinerariaefolium, Plantago ovata, Mentha arvensis, Pelargonium graveolens, and Tagetes
minuta was examined by Pandey and Kalra [233]. The maximum reduction in root galling
was recorded in tomato plants treated with menthol mint vermicompost [Root-Knot Index
(RKI): 1.33] followed by marigold (RKI: 1.66), isabgol (RKI: 1.66), and qinghao (RKI: 1.81).
Compared to one of the most common chemical nematicides (carbofuran), increased fresh
and dry root and shoot weights and fruit yields were achieved with certain herb-based
vermicompost treatments. Singh et al. [234] prepared vermicompost from distillation
waste of aromatic oil crops (e.g., Cymbopogon flexuosus and Cymbopogon winterianus) with
Eisenia fetida to control root-rot of Coleus forskohlii caused by Fusarium chlamydosporum and
Ralstonia solanacearum. Zhou et al. [235] co-composted food waste and sawdust with Chi-
nese medicinal herb residues (1:1:1). The acetone extract of the end products showed an-
tagonistic activity against Fusarium oxysporum and Alternaria solani, but the antimicrobial
properties of the mature compost were as strong as those of the Chinese herbal residues
only. It was concluded that the growth inhibition observed was partly due to the natural
microbiota of the compost. In eleven composts, Zaccardelli et al. [122] found 104 spore-
forming bacterial isolates that exerted antagonistic effects on Sclerotinia minor and Rhi-
zoctonia solani. Among the seven most promising isolates, the in vivo antipathogenic ac-
tivity of two Bacillus subtilis strains and a Bacillus amyloliquefaciens isolate was confirmed
against Sclerotinia minor on Diplotaxis tenuifolia L.
In addition to the solid by-products, the hydrolates remaining after the extraction
process may also possess biocidal properties [236] as a result of leaching of certain bioac-
tive compounds (i.e., trace amounts of EOs and other water-soluble components) during
hydrodistillation [219,237]. The hydrosol extract of Thymus capitatus L. rich in carvacrol
effectively inhibited the growth of important fungal pathogens of Citrus sinensis L. includ-
ing Aspergillus niger, Aspergillus oryzae, and Fusarium solani [238]. Gaspar-Pintiliescu et al.
[239] reported antibacterial activity for the aromatic water of Rosmarinus officinalis. Under
in vitro conditions, strong nematicidal activity was associated with the use of hydrolates
derived from Lavandula × intermedia Emeric ex Loisel. var. super and Lavandula luisieri
(Rozeira) Rivas-Martínez [240]. A similar effect of Thymus citriodorus (Schreb) was demon-
strated by Ntalli et al. [241] against Meloidogyne incognita and Meloidogyne javanica. Sainz
et al. [242] reported a high mortality rate for second-stage juveniles of Meloidogyne javanica
upon use of a hydrolate from Artemisia pedemontana subsp. assoana. However, hydrolate-
based natural pesticides may cause acute toxicity at higher doses for non-target organ-
isms, reduce the metabolism of the natural soil microbiota, and slightly influence the
physiological diversity of microbial communities [243].

5. Conclusions
Modern agriculture still relies heavily on the application of synthetic chemicals, even
though controversies have arisen regarding their safety. Due to the public’s concerns, the
strict regulations, and the pursuit of agricultural sustainability, low-risk and environmen-
tally compatible biocontrol alternatives have emerged as suitable substitutes for these ag-
rochemicals. Botanicals, including EOs and phenolic compounds, have been used for dec-
ades as non-chemical treatments thanks to their biodegradable nature and high selectivity.
Their market is continuously growing; however, they only make up a small percentage of
the global market, and just a few of these aromatic plant-based biopesticides are commer-
cially available because of their varying performance and the absence of field studies re-
garding their efficiency. The inclusion of aromatic and medicinal herbs into agricultural
practices is a possible way to increase crop diversity. In addition, the use of their industrial
Plants 2023, 12, 706 17 of 27

by-products as organic fertilizers may suppress the growth of major soil-borne phytopath-
ogens. The cultivation of aromatic herbs can also improve crop production rates and the
chemical composition of other plants. If used as part of integrated pest management pro-
grams, they are capable of reducing the environmental impacts of chemical pesticides. In
the last few decades, disposal of by-products (e.g., solid biomass, aromatic water, etc.)
generated by the aromatic and medicinal plant-utilizing sector has become a common is-
sue throughout the world. Since these wastes may be rich sources of bioactive compounds
and beneficial microorganisms, their use as natural biopesticides or organic amendments
has also gained popularity. Composting and vermicomposting technologies can offer a
meaningful way to produce a mature and stable end product that can improve important
soil processes and provide nutrients for the treated crops.
All in all, botanicals, and especially EOs, are the most studied and commonly used
treatments against soil-borne plant pathogens. There is substantial evidence that several
of them possess strong antifungal properties, thereby reducing mycelial growth and/or
spore germination of various fungal taxa such as Fusarium, Aspergillus, Penicillium, Rhi-
zoctonia solani, Verticillium dahliae, and Sclerotinia sclerotiorum. Moreover, they show pro-
nounced nematicidal activities against phytoparasitic, root-knot nematodes (Meloidogyne
spp.). Nevertheless, the application of these methods is far behind their true potential due
to some shortcomings, e.g., lack of in vitro experiments, varying selectivity, diversity of
bioactive compounds and complex mechanisms of action, rigorous approval procedures
and registration processes, etc. Therefore, more insight is required to overcome these ob-
stacles and improve the overall commercialization process of plant-based biocontrol
agents.

Author Contributions: Conceptualization, B.G., A.S., E.L. and L.V.; methodology, B.G. and L.V.;
formal analysis, L.V.; investigation, B.G. and A.S.; resources, E.L.; writing—original draft prepara-
tion, B.G. and A.S.; writing—review and editing, L.V.; supervision, E.L.; project administration, E.L.;
funding acquisition, E.L. All authors have read and agreed to the published version of the manu-
script.
Funding: This work was supported by the ÚNKP-22-4-II-SZE-24 New National Excellence Program
of the Ministry for Culture and Innovation (Hungary) from the source of the National Research,
Development, and Innovation Fund.
Data Availability Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.

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