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6 May 2026

22 Pages

Comparative Efficacy of × Hesperotropsis leylandii, Platycladus orientalis, and Juniperus communis Essential Oils Against Four Phytophthora Species Common in Nurseries and Forests

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Faculty of Forestry, University of Belgrade, Kneza Višeslava 1, 11030 Belgrade, Serbia
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Faculty of Forestry and Wood Technology, Mendel University in Brno, Zemědělská 3, 613 00 Brno, Czech Republic
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Institute for Medicinal Plant Research “Dr. Josif Pančić”, Tadeuša Košćuška 1, 11000 Belgrade, Serbia
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Authors to whom correspondence should be addressed.
Forests2026, 17(5), 566;https://doi.org/10.3390/f17050566
This article belongs to the Special Issue Integrative Phytochemistry and Structural Traits of Forest Trees
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Abstract

Phytophthora species are devastating oomycete pathogens affecting agriculture, horticulture, and natural ecosystems globally, primarily spread through the international trade of nursery stock. While synthetic fungicides remain a primary control method, there is an increasing demand for sustainable biocontrol agents. This study evaluated the inhibitory potential of essential oils from three Cupressaceae species, × Hesperotropsis leylandii (needles), Platycladus orientalis (needles), and Juniperus communis (ripe berries), against four common nursery and forest pathogens: P. cactorum, P. plurivora, P. pseudocitrophthora, and P. × cambivora. The essential oils were evaluated at concentrations of 0.1%, 0.25%, and 0.5% (v/v), while the commercial fungicide Infinito® (propamocarb + fluopicolide) served as a positive control. Results demonstrated significant dose-dependent inhibition across all treatments, with the 0.5% concentration yielding the highest efficacy. Notably, essential oil from J. communis achieved 90.2% inhibition against P. × cambivora, while Pl. orientalis essential oil reached 82.8% and 73.1% inhibition relative to the Infinito® effect against P. × cambivora and P. cactorum, respectively, underscoring the potential antimicrobial properties of these coniferous essential oils compared to the tested synthetic standard. Although the chemical analysis revealed that all investigated essential oils (× H. leylandii, Pl. orientalis, and J. communis) contained significant quantities of α-pinene (32.11, 16.01, and 32.29%, respectively), their chemical compositions differed. Namely, GC analysis revealed the presence of δ-3-carene in × H. leylandii and Pl. orientalis (18.51 and 37.98%, respectively), while cedran-8-ol was detected in significant quantity in Pl. orientalis (19.96%). Sabinene and myrcene (18.52 and 14.57%, respectively), besides α-pinene, were most abundant in J. communis essential oil. The observed differences in the evaluated activity might be due to the determined essential oils’ chemical composition.

1. Introduction

Phytophthora de Bary species are fungus-like organisms within the kingdom Chromista/Stramenopiles [1]. Alongside the downy mildews (DMs) within the family Peronosporaceae, they represent some of the major and widespread plant pathogens worldwide [2,3,4,5]. There are over 260 Phytophthora species currently known within this cohesive genus [3,4,5,6,7]. It is also worth noting that their other sister genera are Calycofera R. Bennet & Thines, Halophytophthora H.H. Ho & S.C. Jong, Nothophytophthora T. Jung et al., Synchrospora T. Jung et al., Phytopythium Abad et al., and 20 genera of DMs [8,9,10,11,12,13]. Phytophthora species are highly aggressive, having caused numerous epidemics in agriculture, ornamental plants, and natural and semi-natural ecosystems [5,7,14,15,16,17,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34] while manifesting various symptoms [14,18,35,36,37,38,39]. Also, Phytophthora species are demonstrated to be invasive [5,40,41,42,43,44,45] and easily spread particularly with plants for planting [39,46,47,48,49,50]. Specifically, it has been demonstrated that their ubiquitous global presence is due to the expansion of international trade in living plants [51,52,53] and their subsequent introduction into natural and semi-natural ecosystems via infected nursery stock pathways [39,46,47,48,50,54,55]. Furthermore, ornamental plants are highly vulnerable to Phytophthora infections both in nurseries [47,56,57,58,59,60,61,62,63] and in ornamental plantings [64,65,66,67,68,69,70,71,72].
Control of Phytophthora diseases is challenging; therefore, experience and timely detection relying on the wider adoption of advanced molecular techniques plays a crucial role [60,62,63,68,73,74,75,76,77,78,79]. Preventive management measures and integrated approaches [39,80,81,82,83], including water management [84,85,86], are needed to reduce the risks of infection in nurseries and, subsequently, in natural forests and other plantings. Phosphonate salts were found to be effective in controlling Phytophthora infections [87,88,89,90,91], and against other pathogens [92], but their use is rather limited nowadays [93]. Conventional fungicides have long been considered an effective option for the direct suppression of Phytophthora diseases [94,95,96]. However, their use poses several challenges, as resistance in target pathogens is easily achieved [97,98]. Furthermore, the widespread application of these fungicides has significant negative impact on human health and the environment. Consequently, there is a justified argument that fungicide use and chemical compounds in general should be significantly reduced in the future.
The introduction of various ecologically friendly, biologically based compounds and microbial control agents could be a viable strategy, although they are generally not widely used in forestry [99]. Their potential has been demonstrated in multiple studies [100,101,102,103,104,105,106]. Although their application may have challenges, these natural agents represent promising tools to replace or supplement conventional fungicides against Phytophthora species and other soil-borne pathogens, leading to their reduced usage. Additionally, progress has been made in studying mycovirus diversity in Phytophthora [107,108,109,110,111,112] and in their sister genus Halophytophthora [113]. While mycoviruses have been proven to be effective against other aggressive pathogens [114,115], the field remains largely unexplored in this specific context.
Plant extracts and essential oils (EOs) show significant potential in controlling various pests and pathogens [116,117,118,119,120], including Phytophthora pathogens [121,122,123,124]. These compounds originate from diverse botanical sources [125] and possess strong antifungal properties due to the presence of phenolic components, which induce the disruption of the pathogens’ cell membranes [126,127,128]. Volatility and environmental instability of EOs during application have been the main limiting factors in their wider use. Hence, modern technologies, primarily nanotechnology, enabled the development of formulations with EOs for their more effective utilization [129,130,131,132], including some studies against Phytophthora infections [124,133]. The Cupressaceae family is recognized as a significant source of bioactive secondary metabolites, particularly their EOs [134,135,136], which are characterized by an abundance of monoterpenoids and sesquiterpenoids [137,138], which contribute to their diverse biological activities [137,138,139,140,141,142,143,144,145,146]. In particular, a literature survey revealed antimicrobial activity of plants belonging to the Cupressaceae family against Phytophthora pathogens [121,123,133,147,148,149]. Given their abundance in nurseries and various crops, as well as their ease of propagation and rapid growth, members of the Cupressaceae family represent significant resources for the further exploitation of plant-derived compounds as biopesticides. The aim of this study was to evaluate the inhibitory potential of EOs from three frequently grown and easily propagated Cupressaceae species: × Hesperotropsis leylandii (Jacks. Dallim.) Garl. and Moor. (needles), Platycladus orientalis (L.) Franco (needles), and Juniperus communis L. (berries). These were tested against four common nursery and forest pathogens: P. cactorum (Lebert and Cohn) Schröt., P. pseudocitrophthora Jung, Cacciola, Bakonyi and Horta Jung, P. plurivora Jung and Burgess, and P. × cambivora (Petri) Buisman.

2. Materials and Methods

2.1. Plant Material

Branches with needles of × H. leylandii and Pl. orientalis were collected from five-year-old plants grown in an ornamental nursery near Belgrade, Serbia. The needles from randomly selected trees (voucher specimens coded as CLEY01 and THOR02, respectively) were harvested and stored in bulk in polyethylene bags in a freezer at −20 °C. Voucher specimens of both species were deposited in the University of Belgrade Faculty of Forestry under the codes indicated above. In parallel, juniper berries were collected from J. communis plants and also stored in a freezer at −20 °C at the Institute for Medicinal Plant Research “Dr. Josif Pančić” (voucher specimens coded as JUNF26).

2.2. Isolation of EOs

Isolation of the EOs from branches with needles of × H. leylandii and Pl. orientalis and ripe berries of J. communis was performed by hydrodistillation in Clevenger-type apparatus (Medilab, Ambala Cantt, Haryana, India), as prescribed by [150]. In short, plant material was grounded and transferred into Clevenger apparatus, where hydrodistillation was performed for 2 h, after which the obtained EOs were extracted using diethyl ether. The samples were then filtered through anhydrous sodium sulfate to remove any remaining moisture, while the solvent was removed using the gentle stream of nitrogen. The EO content was expressed as a percentage of the dry mass of the plant material. The yields of obtained essential oils were 0.93, 0.69 and 2.90% for × H. leylandii, Pl. orientalis and ripe berries of J. communis, respectively. EO was stored at a temperature of 4 °C until analysis.

2.3. Chemical Analysis of the EOs

Chemical analysis of the obtained EOs was carried out using gas chromatography (GC) and gas chromatography coupled with mass spectrometry (GC–MS).

2.3.1. GC Analysis

GC analysis was performed using an apparatus HP-5890 Series II GC (Hewlett-Packard, Waldbronn, Germany) that was equipped with a split-splitless injector, an automatic liquid sampler and a flame ionization detector (FID). The employed column was HP-5 (25 m × 0.32 mm, 0.52 μm film thickness). The flow rate of H2, used as the carrier gas, was 1 mL/min, while split ratio was 1:30. Temperature of the injector was 250 °C, while detector temperature was 300 °C. Temperature of the column was programed to increase linearly from 40 °C to 260 °C with a rate of 4 °C/min and was then kept at a temperature of 260 °C for 10 min. Tested EOs were dissolved in alcohol, injected in the amount of 1 μL. Quantification was carried out using area percent reports, obtained as a result of standard processing of chromatograms. The composition of the tested EOs was expressed in percentages computed from GC peak areas with the response factor considered to be 1.

2.3.2. GC–MS Analysis

For GC–MS analysis 1800C Series II GCD system (Hewlett-Packard, Palo Alto, CA, USA) equipped with a column HP-5MS (30 m × 0.25 mm, 0.25 μm film thickness) was used, using helium as the carrier gas. The transfer line was heated at 260 °C. Mass spectra were acquired in EI mode (70 eV), in m/z range 40–450. An amount of 0.2 μL of EOs dissolved in alcohol was injected. The components of the EOs were identified using comparison of their mass spectra to the ones available in libraries Wiley 275 and NIST/NBS. Calibration was carried out by means of linear n-paraffins mixture (C6–C40) as a standard. The experimental values for retention indices were determined employing calibrated Automated Mass Spectral Deconvolution and Identification System Software (AMDIS ver. 2.1), compared to those from the available literature, and used as an additional tool to confirm the MS findings.

2.4. Antimicrobial Assays

The inhibitory activity was tested in vitro using the poisoned food method. Following the protocol of Milanović et al. [149], selected EOs were used to evaluate the mycelial growth of four Phytophthora species. These isolates, frequently found in both nurseries and planted forests [47], were obtained from the culture collection of the Faculty of Forestry at the University of Belgrade. Specifically, the following four Phytophthora species were used: P. cactorum (JX276090), originating from Persian walnut (Juglans regia L.) (Milenković, unpublished); P. pseudocitrophthora (OR518930), originating from sessile oak (Quercus petraea (Matt.) Liebl.) [5]; P. plurivora (JX276057), originating from pedunculate oak (Quercus robur L.) (Milenković, unpublished); and P. × cambivora (MG595173), originating from cherry laurel (Prunus laurocerasus L.) [66]. All isolates were cultivated on V8 agar (V8A) media [100 mL/L V8 juice (Biotta®, Tägerwilen, Switzerland), 2 g/L CaCO3, and 20 g/L agar (Torlak, Belgrade, Serbia)] at 22–25 °C in the dark until they reached approximately half the diameter of a 90 mm Petri dish. The EOs of × H. leylandii (EOHL), J. communis (EOJC), and Pl. orientalis (EOPO) were diluted in 0.5% (v/v) Tween 80. Before pouring the V8A media (at ca. 50 °C), the EOs were added to reach volume concentrations of 0.1%, 0.25%, and 0.5%. Subsequently, 12 mL of the media was poured into each 90 mm Petri dish. The control group received a mock dilution of 0.5% Tween 80 and sterile distilled water at the same concentrations. From the edges of actively growing colonies, 6 mm agar plugs were taken with a sterile metal cork borer and placed in the center of the V8A plates containing EOs. Six replicates per concentration, EO, and Phytophthora species were prepared, including the mock-diluted control. Two lines at a 90° angle were drawn on the bottom of each Petri dish. All dishes were incubated at 20 °C in the dark until one of the treatments reached the edge of the Petri dish. Mycelial growth was determined with a steel needle, and the edges of the colonies were marked. The mean growth values were calculated from the total diameter measured in two directions and expressed in mm/day. The percent inhibition of mycelial growth relative to the control was determined according to the methods previously described by Milanović et al. [149], using the following formula:
I = (C − T)/C × 100 [%],
where I is the inhibition rate (%); C is the growth diameter of the control; and T is the growth diameter of different treatments.

2.5. Statistical Analyses

Prior to analysis, the data were tested for normality and homogeneity of variances using the Shapiro–Wilk test and Levene’s test, respectively. Since the data about inhibition rate (I) did not meet the assumptions of normal distribution and equal variances, a non-parametric PERMANOVA was applied. The analysis was performed separately for each species to evaluate differences among treatments within species (PAST software, version 4.03). Figures were plotted in OriginPro 2023b (OriginLab Corporation, Northampton, MA, USA).

3. Results

3.1. Chemical Composition of the Tested EOs

The chemical analysis revealed that all investigated EOs (× H. leylandii, Pl. orientalis, and J. communis, EOHL, EOPO and EOJC, respectively) were the most abundant in monoterpenes (90.21, 65.05 and 81.46, respectively), containing significant quantities of α-pinene at levels of 32.11, 16.01, and 32.29%, respectively (Table 1). However, their chemical compositions differed. Namely, GC analysis revealed the presence of δ-3-carene in × H. leylandii and Pl. orientalis in quantities of 18.51 and 37.98%, respectively, whereas cedran-8-ol was detected in significant quantity in Pl. orientalis (19.96%) (Table 1). On the other hand, sabinene and myrcene at levels of 18.52 and 14.57%, respectively, besides α-pinene, were most abundant in ripe berries J. communis essential oil (Table 1).
Table 1. Comparative chemical profile of the essential oils obtained from needles of the Leyland cypress (× H. leylandii)—EOHL and Oriental thuja (Pl. orientalis)—EOPO and ripe berries of common juniper (J. communis)—EOJC.

3.2. Antimicrobial Assay

The inhibitory effect differed significantly among treatments across all four Phytophthora species (PERMANOVA, p < 0.001 in all cases; Figure 1). The commercial fungicide Infinito® consistently demonstrated the strongest inhibitory activity (Figure 2, Figure 3, Figure 4 and Figure 5). In contrast, the × H. leylandii EO (EOHL) induced the lowest inhibition relative to Infinito® across all species (Figure 1).
Figure 1. Inhibitory activity of three essential oils and the standard fungicide Infinito® against four Phytophthora species. Panels (A–D) represent different Phytophthora species: (A) P. × cambivora; (B) P. cactorum; (C) P. pseudocitrophthora; and (D) P. plurivora. HL, JC, and PO denote the essential oils of × Hesperotropsis leylandii, Juniperus communis, and Platycladus orientalis, respectively. These were applied at three concentrations (0.1%, 0.25%, and 0.5%), whereas INF denotes the standard fungicide Infinito®. Bars represent mean values ± SE. Different letters above bars indicate significant differences among treatments within each species.
Figure 2. Inhibitory activity of × Hesperotropsis leylandii (EOHL), Juniperus communis (EOJC), and Platycladus orientalis (EOPO) essential oils, and the fungicide Infinito® against Phytophthora cactorum after six days of incubation at 20 °C in the dark: (A) control; (B–D) EOHL at 0.5%, 0.25%, and 0.1%, respectively; (E) fungicide Infinito®; (F–H) EOJC at 0.5%, 0.25%, and 0.1%, respectively; (I–K) EOPO at 0.5%, 0.25%, and 0.1%, respectively.
Figure 3. Inhibitory activity of × Hesperotropsis leylandii (EOHL), Juniperus communis (EOJC), and Platycladus orientalis (EOPO) essential oils, and the fungicide Infinito® against Phytophthora plurivora after six days of incubation at 20 °C in the dark: (A) control; (B–D) EOHL at 0.5%, 0.25%, and 0.1%, respectively; (E) fungicide Infinito®; (F–H) EOJC at 0.5%, 0.25%, and 0.1%, respectively; (I–K) EOPO at 0.5%, 0.25%, and 0.1%, respectively.
Figure 4. Inhibitory activity of × Hesperotropsis leylandii (EOHL), Juniperus communis (EOJC), and Platycladus orientalis (EOPO) essential oils, and the fungicide Infinito® against Phytophthora pseudocitrophthora after six days of incubation at 20 °C in the dark: (A) control; (B–D) EOHL at 0.5%, 0.25%, and 0.1%, respectively; (E) fungicide Infinito®; (F–H) EOJC at 0.5%, 0.25%, and 0.1%, respectively; (I–K) EOPO at 0.5%, 0.25%, and 0.1%, respectively.
Figure 5. Inhibitory activity of × Hesperotropsis leylandii (EOHL), Juniperus communis (EOJC), and Platycladus orientalis (EOPO) essential oils, and the fungicide Infinito® against Phytophthora × cambivora after six days of incubation at 20 °C in the dark: (A) control; (B–D) EOHL at 0.5%, 0.25%, and 0.1%, respectively; (E) fungicide Infinito®; (F–H) EOJC at 0.5%, 0.25%, and 0.1%, respectively; (I–K) EOPO at 0.5%, 0.25%, and 0.1%, respectively.
At a volume concentration of 0.5%, EOHO achieved 56.8% of the Infinito® effect in P. × cambivora, 33.5% in P. cactorum, 25.2% in P. pseudocitrophthora, and 19.7% in P. plurivora; inhibition was markedly lower at reduced concentrations (Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5). Among the EO treatments, J. communis berry EO (EOJC) most closely resembled the efficacy of Infinito®. At a 0.5% concentration, EOJC exhibited 90.2% inhibition in P. × cambivora, 55.3% in P. plurivora, 52.6% in P. pseudocitrophthora, and 32.3% in P. cactorum. Notably, at a 0.25% concentration, EOJC still maintained 42.6% of the Infinito® effect in P. × cambivora.
The EO from Pl. orientalis needles (EOPO) also showed significant inhibition at a 0.5% concentration, reaching 82.8% of the Infinito® effect in P. × cambivora, 73.1% in P. cactorum, 42.5% in P. plurivora, and 31.3% in P. pseudocitrophthora (Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5). Interestingly, EOPO maintained considerable inhibition at lower concentrations in specific species, showing 56.5% and 35.8% of the Infinito® effect in P. × cambivora and P. cactorum at 0.25%, respectively, and 28.3% at 0.1% in P. × cambivora (Figure 1).
Conversely, the weakest inhibition was consistently recorded at the lowest concentration (0.1%), specifically for EOHL in P. plurivora (7.1%), P. cactorum (4.8%), and P. × cambivora (2.5%), as well as EOPO in P. pseudocitrophthora (0.1%). Thus, while inhibitory activity generally increased with concentration, the treatment ranking and degree of similarity to the standard fungicide were clearly species-specific (Figure 2, Figure 3, Figure 4 and Figure 5).

4. Discussion

Our findings demonstrated that, while the commercial fungicide Infinito® remained the most potent inhibitor, the EOs of J. communis berries and Pl. orientalis needles (EOJC and EOPO, respectively) at a concentration of 0.5% exhibited significant activity, what might be considered as promising biocontrol agents, particularly against P. × cambivora (Figure 1 and Figure 5). This is of particular importance given the high significance of P. × cambivora as a destructive pathogen. Specifically, this hybrid species, originating from Southeast Asia [151,152], is highly aggressive and represents a primary cause of ink disease in chestnut (Castanea sativa Mill.) [18,22,153,154]. Beyond chestnuts, this pathogen has been found to be aggressive on numerous other hosts [18,66,155], most notably European beech (Fagus sylvatica L.), causing a variety of deteriorating symptoms [17,18,36,38,156,157]. Promising results were also obtained in the case EOPO at 0.5% concentration against another widespread and notorious pathogen P. cactorum (Figure 1 and Figure 2). This species has a long history of causing various damages in different crops in agriculture, horticulture and forestry [23,110,124,158,159,160,161,162]. In the cases of P. plurivora and P. pseudocitrophthora, the highest inhibition was achieved by EOJC at 0.5% concentration, which reached 55.3% and 52.6% efficacy in comparison to positive control, respectively (Figure 1, Figure 3 and Figure 4). P. plurivora [163] is one of the most widespread species worldwide, damaging a broad range of hosts [5,18,39,156,157,158,163,164,165,166]. Beyond causing direct damage, this species exacerbates the impact of spongy moth (Lymantria dispar L.) infestations [167,168], further contributing to the decline of affected forest ecosystems. Additionally, this species accelerated the root deterioration of Fraxinus excelsior L., leading to a progressive decline in conjunction with the bark pathogen Hymenoscyphus fraxineus (T. Kowalski) Baral, Queloz & Hosoya [169]. Phytophthora pseudocitrophthora was recently described [5] following detailed analyses of its closely related species, P. citrophthora (R.E. Sm. & E.H. Sm.) Leonian [162]. The latter species has a long history and is one of the earliest identified members of the genus Phytophthora [5,6,162]. Although the isolate (OR518930) used in this study was obtained from sessile oak, this species has been recorded on various hosts worldwide [5,6,164]. The lower efficacy observed in P. plurivora and P. pseudocitrophthora, compared to noted activity against P. × cambivora, suggested that these species might possess inherent physiological mechanisms, such as distinct cell wall composition or specialized enzymatic detoxification pathways, that provide greater resilience against the bioactive compounds found in the investigated EOs. However, further research, including biochemical assays or gene expression studies, is required to elucidate these underlying mechanisms.
Phytophthora species are among the most damaging plant pathogens worldwide [3,4,5,6,18,170,171,172,173,174,175,176,177]. These species are highly invasive [39,40] and spread easily via ‘plants for planting,’ particularly through infested nursery stock [39,46,47,48,49,50,171].
Given that numerous studies have shown afforestation programs to be severely affected by Phytophthora pathogens [39,47,48,49], the implementation of European Union (EU) Regulation 2024/1991 [178] on the restoration of natural sites is of particular concern. As this regulation will likely necessitate a vast quantity of new planting material, the risk of widespread disease incidence and the subsequent introduction of pathogens into restored areas is significantly heightened [47,179]. Therefore, the adoption of strict preventative measures to reduce the presence of these pathogens [39,47,85] is essential. Furthermore, seeking sustainable, ecologically friendly methods for the suppression of Phytophthora species is now urgently required to ensure the success of large-scale restoration efforts. Our results demonstrated that EOs, specifically EOJC and EOPO, offered a viable path toward these sustainable suppression goals, providing high levels of inhibition against aggressive hybrids like P. × cambivora.
Control of these pathogens remains highly challenging. While conventional fungicides have traditionally been regarded as an effective option for the direct suppression of Phytophthora diseases [94,95,96,180,181,182], their continued use faces significant hurdles. Beyond the growing issue of pathogen resistance [97,98], numerous negative environmental consequences have been documented. Consequently, the use of synthetic chemical compounds is likely to be increasingly restricted, as seen with the current limitations on copper-based fungicides within the EU [183]. Phosphonate salts were considered the effective and ecologically acceptable tool in controlling Phytophthora infections [87,88,89,90,91]. However, their usage has been strictly limited or banned since (EU) Regulation 2019/1009 [93] entered into force in July 2022. Therefore, in addition to the detailed preventive measures indicated by Horta Jung et al. [39], various host resistance and breeding programs [184,185,186,187] or the application of biocontrol agents could provide suitable solutions for the management of Phytophthora diseases [100,101,102,103,104,105,106,188]. In this context, EOs obtained from the Cupressaceae family are promising additional tools; they are widespread in nurseries, easily propagated, fast-growing, and have been proven to be effective in antimicrobial assays against Phytophthora pathogens [121,122,123,133,147,148,149].
Our study revealed that chemical compositions differed among the investigated EOs of × H. leylandii, Pl. orientalis, and J. communis (Table 1). All tested EOs contained significant quantities of α-pinene at 32.11%, 16.01%, and 32.29% (Table 1). Additionally, δ-3-carene was recorded in × H. leylandii and Pl. orientalis at 18.51% and 37.98%, respectively, whereas cedran-8-ol was detected in significant amounts in Pl. orientalis (19.96%). The chemical composition of the × H. leylandii EO corresponded well with previous findings [189,190]. Although × H. leylandii EO was highly efficient in suppressing arthropod larvae in wheat [190], in our study, it showed the lowest inhibition at the tested concentrations against all Phytophthora species used (Table 1; Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5).
Chemical analyses of Pl. orientalis EO were congruent with previous studies [134,136], as well as with species from the sister genus Thuja, such as Thuja plicata Donn ex D.Don [123,133,134,135,137,143,145]. In contrast, a slightly different composition with a lower α-pinene content was recorded for T. occidentalis L. EO [146]. Platycladus orientalis EO showed promising antimicrobial activity against the tested Phytophthora species, particularly P. plurivora, P. pseudocitrophthora, and P. × cambivora (Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5); this aligns with earlier testing of the antimicrobial activity of this species [139]. Furthermore, its related species T. plicata has shown high potential in suppressing Phytophthora pathogens [123,133]. Due to its antimicrobial activity, vigorous growth, easy propagation, and ubiquitous presence [191], Pl. orientalis EO could be considered as showing great promise for further exploitation as an eco-friendly biopesticide.
α-pinene was the most abundant compound in J. communis berries EO (32.29%), followed by sabinene and myrcene at concentrations of 18.52% and 14.57%, respectively. These findings are consistent with previous studies [142,192], although significantly higher amounts of α-pinene have been recorded in some cases [193]. The EO of this species has demonstrated significant antimicrobial and inhibitory properties in several studies [142,192,193,194,195]. In the present study, this oil was highly efficient and showed strong inhibitory properties against aggressive Phytophthora pathogens, suggesting it could be considered for further in vivo experiments and potential use as a biopesticide. This is particularly relevant in the context of EOs nanoencapsulation [124,132,133,196] for slow-release purposes to enhance their properties.
While the present study clearly demonstrates the potent antifungal activity of the × H. leylandii, Pl. orientalis, and J. communis EOs through in vitro assays, the specific underlying mechanisms of action remain to be fully elucidated. The fundamental mechanisms of action by which essential oils and their constituent phenolic compounds disrupt fungal cell walls and membranes are well-documented in the existing literature (e.g., [126,127,128]). Future investigations employing microscopic techniques, such as Scanning Electron Microscopy (SEM) or fluorescence microscopy, would be beneficial to visualize the precise morphological changes induced in the fungal cells and to confirm these mechanistic pathways. Future research should also focus on the synergistic effects of the primary constituents within the tested EOs, specifically α-pinene, δ-3-carene, sabinene, and cedran-8-ol, as these compounds play a decisive role in antimicrobial activity. Understanding these interactions is essential for optimizing bio-fungicides as sustainable alternatives to synthetic chemicals in nursery and forest management.

5. Conclusions

As the global demand for sustainable plant production grows, the identification of bio-based alternatives to synthetic fungicides is crucial. This study demonstrates that essential oils from three Cupressaceae species possess remarkable inhibitory potential against four devastating Phytophthora pathogens. At a concentration of 0.5%, these natural extracts achieved mycelial growth inhibition levels comparable to the commercial synthetic fungicide. The high efficacy of Juniperus communis and Platycladus orientalis essential oils suggests that their specific chemical profiles, rich in α-pinene, δ-3-carene, sabinene, and cedran-8-ol, play a decisive role in their antimicrobial activity. Future research should focus on the synergistic effects of these primary constituents to optimize these bio-fungicides as sustainable alternatives to synthetic chemical treatments in nursery and forest management.

Author Contributions

Conceptualization, I.M., S.M. and V.T.; methodology, I.M., A.Ž., S.M., J.D., M.M. and V.T.; validation, S.M., A.Ž. and J.D.; formal analysis, I.M., A.Ž., S.M., J.D., M.M. and V.T.; investigation, I.M., A.Ž. and V.T.; data curation, S.M. and V.T.; writing—original draft preparation, I.M., S.M. and V.T.; writing—review and editing, A.Ž., J.D. and M.M.; visualization, S.M.; supervision, V.T.; funding acquisition, I.M., S.M. and V.T. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Ministry of Science, Technological Development and Innovation of the Republic of Serbia, grant numbers 451-03-34/2026-03/200169 and 451-03-33/2026-03/200003, and the European Regional Development Fund, project ‘Phytophthora Research Centre’, grant number CZ.02.1.01/0.0/0.0/15_003/0000453.

Data Availability Statement

Data is available from the corresponding author on a reasonable request.

Acknowledgments

We are grateful to Jovan Sremčević, and the ‘Omorika’ nursery in Belgrade for their support in providing plants for EO extraction. We also thank Petar Vujičić, for his excellent assistance with the laboratory work.

Conflicts of Interest

The authors declare no conflicts of interest.

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