In vitro antifungal and sporulation-inhibitory activity of selected essential oils against Aspergillus niger and GC-MS characterization of selected highly active oils

Introduction

Fungal diseases of crops are a major constraint to agricultural productivity and postharvest quality worldwide. Species of Aspergillus, including A. niger, are particularly important because they cause seedling blights, collar rot, and storage rots of oilseed and grain crops, and can reduce seed germination and marketability (Aiswarya et al., 2024) [2] . In groundnut, for example, A. niger is associated with collar rot and storage rot, leading to significant yield and quality losses in many production systems (Kishore et al., 2007) [15] . Conventional management of fungal diseases relies heavily on synthetic fungicides and, in some cases, antibiotics and copper compounds. While such chemicals are effective, their intensive and often indiscriminate use has raised environmental and human health concerns, including pesticide residues in food, non-target toxicity, and the evolution of fungicide-resistant pathogen populations (Raveau et al., 2020; Samal et al., 2024) [26] . Similar issues are well documented for antibiotic use in plant agriculture, where resistant bacterial strains and offtarget impacts on microbiomes have been observed (McManus et al., 2002) [20] . These challenges have driven a strong interest in developing eco-friendly alternatives and integrating them into disease management programs. Plant-derived products, especially essential oils (EOs), have emerged as promising candidates for green disease management. EOs are complex mixtures of volatile secondary metabolites, primarily terpenes, terpenoids, phenylpropanoids, and related compounds, produced by aromatic plants. They are generally recognized as biodegradable, often have relatively low mammalian toxicity, and can show broad-spectrum antimicrobial activity (Nazzaro et al., 2017) [21] . Over the last two decades, numerous studies and reviews have demonstrated the antifungal properties of EOs against postharvest and field fungi, including Aspergillus, Penicillium, Botrytis, Colletotrichum, and others, and have proposed their use as fumigants, vapours, coatings, or components of active packaging (Sivakumar & Bautista-Baños, 2014; Allagui et al., 2023) [33] . The antifungal activity of EOs is closely related to their chemical composition. Phenylpropanoids such as (E)-cinnamaldehyde and eugenol, and oxygenated monoterpenes such as citral, thymol, and carvacrol, are often identified as key bioactive constituents International Journal of Agriculture and Food Science https://www.agriculturaljournals.com (Powers et al., 2018) [23] . Cinnamon (Cinnamomum spp.) oils are typically rich in (E)-cinnamaldehyde and have shown strong inhibition of growth and spore germination in a range of fungi, including Aspergillus spp. (Zhang et al., 2025) [37] . Likewise, lemongrass (Cymbopogon citratus) oil, dominated by citral (neral + geranial), and bay laurel (Laurus nobilis) oil, containing eugenol and monoterpene hydrocarbons, have been reported to inhibit mycelial growth and reduce mycotoxin production by Aspergillus species (Belasli et al., 2020; Adebayo and Osuolale, 2024) [6] . Despite this growing body of work, there is still value in comparative screening of multiple commercial EOs against important storage fungi, especially when combined with chemical profiling that allows a preliminary structureactivity interpretation. Such data can help to prioritize candidate oils for formulation development (e.g., coatings, nanoemulsions, or EO-fungicide combinations) and to identify oils that reduce sporulation, which is especially critical for lowering inoculum pressure and airborne spread (Rabari et al., 2018) [25] .

In this study, we evaluated the in vitro antifungal activity of 14 commercial EOs against Aspergillus niger using a disc diffusion assay. We recorded both inhibition of mycelial growth (zone of inhibition) and the qualitative effect on sporulation. Two selected highly active oils, Cinnamomum cassia and Laurus nobilis, were further subjected to GC-MS analysis to determine their major chemical constituents and to relate composition to bioactivity. The overarching aim was to identify promising EO candidates for the development of plant-based fungicidal or fungistatic products against A. niger and similar storage pathogens.

Materials and Methods

Isolation of Fungi

Infected Arachis hypogaea roots were collected from a farm in Jamvadi village, Gondal, Rajkot, Gujarat. The infected material was washed under running tap water for 3 h and surface sterilized with 0.1% HgCl2 for 15 min. It was then washed three times with sterile distilled water. After surface sterilization, infected root pieces were cut using sterile forceps and a scalpel. The cut infected pieces were inoculated on PDA medium under aseptic conditions and incubated at 37 ºC. After 48 h of incubation, mycelial growth was observed around the leaf. Morphological identification was carried out by observing the growth pattern and colour of the hyphae.

Essential oils

Fourteen EOs were obtained from Vimal Research Society for Agro-Biotech and Cosmic Powers (VIRSACO). The oils evaluated is listed in Fig. 1 together with their antifungal activity data.

Disc diffusion assay for antifungal activity

Antifungal activity of the 14 EOs against A. niger was evaluated using a disc diffusion method adapted from Rabari et al. (2018) [25] . A. niger was grown on PDA plates for 96 h at 37 °C to obtain actively growing mycelium and conidia. After 96 h, fungal mycelium and conidia were suspended in 10 mL sterile distilled water (D/W). One hundred microlitres (100 µL) of fungal suspension was spread over PDA plates using a glass spreader to obtain uniform fungal growth on test plates. For preparing the test discs, 5 µL of EO was pipetted onto a 5-mm Whatman filter paper disc and placed carefully on the PDA medium. All Petri plates were sealed with sterile laboratory parafilm to avoid evaporation of the test samples. The plates were left for 30 min at room temperature to allow diffusion of the oil and then incubated at 37 °C for 96 h. After incubation, the zone of inhibition was measured for all 14 oils in millimetres. The experiment was performed in triplicate, and the mean value was calculated. The antifungal activity of the oils is summarized in Figs. 1 and2. In addition to mycelial growth inhibition, the effect of each EO on sporulation was qualitatively recorded as complete inhibition of growth (no visible mycelium), sporulation inhibition (mycelial growth present but with markedly reduced or absent conidiation around the disc), or no apparent effect on sporulation relative to the control.

GC-MS analysis of selected essential oils

The two selected highly active oils, C. cassia and L. nobilis, were analyzed using gas chromatography-mass spectrometry (GC-MS) to identify major constituents. Analyses were carried out on a Perkin Elmer AutoSystem XL gas chromatograph coupled to a Perkin Elmer TurboMass mass spectrometer. Separation was achieved on a DB-1 capillary column (30 m x 0.25 mm, 0.25 µm film thickness) with helium as carrier gas at a flow rate of 1 mL min-1. The injector and detector temperatures were set at 250 °C and 280 °C, respectively. The oven temperature was programmed from 60 to 250 °C at 5 °C/min. A 2 µL sample of each EO diluted in an appropriate solvent was injected in split mode. The transfer line temperature was maintained at 280 °C. Mass spectra were acquired over an appropriate mass range, and EO components were identified by comparing their retention times and mass spectra with those in the NIST library and, when possible, with published data. Relative percentages of components were calculated from peak area normalization (Table 1).

Data handling and R-based visualization

All data handling and visualization were carried out in R (version 4.5.0; R Core Team, Vienna, Austria). The dataset containing botanical name, family, common name, zone of inhibition, and qualitative sporulation response for each EO was entered into a comma-separated values (CSV) file and imported into R using the readr package. Data wrangling and variable derivation were performed using functions from dplyr and tidyr within the tidyverse ecosystem. For each EO, an efficacy class was assigned based on the measured zone of inhibition: very low (<10 mm), low (10-30 mm), moderate (30-60 mm), and high (>60 mm). These classes were defined a priori to facilitate visualization and are not intended as formal breakpoints. A numerical rank variable (1 = highest activity) was generated by ordering oils in descending order of zone diameter and resolving ties by assigning the same rank. These derived variables are included in Fig. 1. Graphical representation of the antifungal activity was produced using ggplot2. A Cleveland dot (lollipop) plot was constructed with zone of inhibition (mm) on the x-axis and EO name on the y-axis. Horizontal segments were drawn from zero to the observed zone value for each oil, with points at the terminal zone value. Panels were faceted according to the qualitative sporulation response category (no apparent effect, sporulation inhibited, complete inhibition of growth and sporulation), and points International Journal of Agriculture and Food Science https://www.agriculturaljournals.com were coloured by efficacy class to emphasize differences among oils.

Results and Discussion Antifungal and sporulation-inhibitory activity of essential oils

Thirteen of the [15] . Similarly, lemongrass oil, dominated by citral, has been documented to completely inhibit mycelial growth of several Aspergillus species and other fungi at relatively low concentrations (Matasyoh et al., 2011) [19] . Recent studies on L. nobilis EO also highlight its strong antifungal and antitoxigenic effects against Aspergillus and yeast pathogens, supporting the high activity observed in our work (Belasli et al., 2020) [6] .

At the same time, some oils with modest zones (e.g., M. alternifolia and L. officinalis) nonetheless markedly suppressed sporulation. This duality, moderate growth inhibition but strong impact on conidiation, suggests that different EO components or concentration thresholds may differentially affect hyphal extension and spore differentiation. Such effects have been noted in mechanistic studies showing that EOs can alter membrane integrity, interfere with ergosterol biosynthesis, and disrupt signaling pathways involved in sporulation and secondary metabolism (Leiva-Mora et al., 2025) [16] . For storage disease management, oils that primarily reduce sporulation could be combined with other agents (e.g., biological control organisms or reduced-dose fungicides) to achieve both growth suppression and inoculum reduction. [32] . For L. nobilis EO, eugenol (50.40%) was the predominant peak, followed by benzaldehyde (10.96%), caryophyllene (10.23%), beta-linalool (9.90%), beta-pinene (3.17%), limonene (3.71%), 1,8-cineole (2.63%), 1-terpinene (2.10%), and other minor constituents (7.99%) (Table 1). This eugenol-rich chemotype is consistent with earlier reports in which L. nobilis leaf and flower oils contain large proportions of eugenol, 1,8-cineole, and monoterpene hydrocarbons, and exhibit notable antifungal activity (Belasliet al., 2020) [6] .

GC-MS

The strong antifungal effects of C. cassia and L. nobilis against A. niger in our assay therefore likely stem from the high abundance of phenylpropanoids such as (E)cinnamaldehyde and eugenol. Both compounds have been shown to disrupt fungal membranes, increase cell permeability, and cause leakage of intracellular constituents, ultimately leading to growth inhibition and cell death (Powers et al., 2018) [23] . Aldehydes like cinnamaldehyde are particularly reactive toward nucleophiles in proteins and may interfere with key metabolic enzymes, while eugenol can insert into lipid bilayers and disturb membrane structure (Al-Harrasi et al., 2022) [3] .

Sporulation-targeted EO activity and formulation potential

A notable feature of this study is the differentiated response of A. niger in terms of hyphal growth versus sporulation. Oils such as C. citratus and L. nobilis completely inhibited both mycelial growth and sporulation, whereas M. alternifolia, G. macrorhizum, A. graveolens, and C. cassia strongly reduced sporulation despite moderate growth zones. This pattern suggests that some EO components may preferentially target developmental processes leading to conidiation rather than primary vegetative growth. From a disease management standpoint, this is an under-exploited angle: many fungicide screening assays focus only on radial growth, ignoring sporulation behaviour. Recent reviews on EO mechanisms emphasize the need to move beyond simplistic MIC values and to examine multiple endpoints, including membrane integrity, oxidative stress, sporulation, biofilm formation, and mycotoxin production (Mani-López et al., 2021) [18] . In the context of A. niger, EOs that primarily suppress sporulation could help maintain low inoculum loads on seeds and in storage environments even when some mycelial growth persists, thereby reducing spread and potentially lowering mycotoxin risk. Future work could quantify conidial counts microscopically and evaluate effects on spore viability and germination.

Another forward-looking avenue is the incorporation of these highly active oils into delivery systems such as nano emulsions, edible coatings, and EO-loaded films, which can stabilize volatile compounds and provide controlled release.

Recent studies have demonstrated the success of EO nano emulsions (including L. nobilis EO) and EO-incorporated biopolymer films in inhibiting Aspergillus spp. on food substrates while improving barrier properties (Singhet al., 2023) [30] . The strong activity of C. cassia, L. nobilis, C. citratus, and S. aromaticum in our in vitro assay suggests that these oils would be promising candidates for such formulations aimed at protecting stored seeds or grains.

Comparison with previous EO screenings against Aspergillus

Large screening studies of EOs have shown that a relatively small subset of oils display consistently strong antifungal activity across multiple fungi. For example, Rabari et al.

(2018) [25] screened 75 EOs and identified cinnamon, clove, lemongrass, and thyme among the most broadly active oils. Similar conclusions emerge from other surveys and metaanalyses of EOs against Aspergillus and Penicillium spp., where phenylpropanoid-rich and citral-rich oils typically rank high in potency. Our results align with these patterns and reinforce the idea that these chemotypes could serve as backbone components in botanical fungicide formulations (Powers et al., 2018) [23] . However, we also observed that some oils traditionally considered weaker fungicides, such as sandalwood and jasmine, still produced small but measurable inhibition zones. While these effects may not be sufficient on their own, they could potentially contribute to synergistic or additive interactions in multi-component blends, for instance, cinnamon-clove or cinnamon-lemongrass mixtures, which have shown synergistic effects against A. niger and other microbes in previous work (Purkait et al., 2020) [24] . Exploring such combinations could allow dose reduction and help address cost and sensory constraints (e.g., strong odour or flavour) in practical applications. . Follow-up studies using broth microdilution and vapor-phase assays, together with quantification of spore germination and mycelial growth rates at different concentrations, would allow a more precise determination of fungistatic and fungicidal thresholds for the most promising oils.

Limitations and future directions

Finally, the current experiments were performed under controlled in vitro conditions. For practical application in crop or storage protection, the most active oils, particularly C. cassia, L. nobilis, C. citratus, and S. aromaticum, should be evaluated in more realistic systems, such as EO-based coatings, nanoemulsions, or active packaging applied to seeds or stored commodities. Such studies could be extended to examine seed germination, phytotoxicity, residue behaviour, and safety for mammalian cells. Despite these natural limitations of a first screening, the present results provide a clear and useful baseline for prioritizing candidate oils and guiding the design of more detailed mechanistic and formulation studies.

Conclusion

The present study demonstrates that selected plant essential oils can strongly inhibit the growth and sporulation of Aspergillus niger under in vitro conditions. Among the 14 essential oils tested, Laurus nobilis, Cymbopogon citratus, Cinnamomum cassia, and Cinnamomum zeylanicum showed the highest antifungal activity, while several oils also reduced conidial production. This sporulation-inhibitory effect is important because suppression of spores can reduce inoculum load and may help limit the spread of A. niger in crop and storage environments. GC-MS analysis showed that the strong activity of C. cassia and L. nobilis was associated with major bioactive constituents such as (E)-cinnamaldehyde and eugenol, respectively. These compounds are known to affect fungal membrane integrity and may contribute substantially to the observed inhibition. Although the work is an initial laboratory screening, the results identify these essential oils as promising candidates for further development as botanical fungistatic or fungicidal agents. Future studies should evaluate dose-response relationships, vapor-phase activity, effects on spore germination and mycotoxin production, and performance in seed coating, nano emulsion, or storage-protection systems. Overall, this study supports the potential use of essential oils as eco-friendly, residue-conscious tools for integrated management of A. niger and related storage pathogens. [13] Inhibits Aspergillus flavus and other fungi; fungicidal against various yeasts and molds (Li et al., 2022) [17] (E)-Cinnamaldehyde 15.369 66.36 Phenylpropanoid aldehyde

Potent antioxidant, broad antimicrobial, anti-inflammatory, insecticidal, anti-aflatoxigenic, gut-protective, possible neuroprotective (Sharma et al., 2016) [29] Strong inhibition of A. flavus, A. niger, A. fumigatus growth and aflatoxin B₁ production; effective in vapor and contact assays (Kim et al., 2018) [14] Eugenol 15.956 7.74 Phenylpropanoid phenolic ether

Strong antioxidant, broadspectrum antimicrobial, analgesic, anti-inflammatory, antitumor; food-preservative potential (Ulanowska and Olas, 2021) [34] Active against Aspergillus spp., Candida and dermatophytes; inhibits cell envelope and virulence factors (Didehdar et al., 2022) [9] Cinnamyl acetate 17.110 3.97 Phenylpropanoid ester

Fragrance component; contributes to antioxidant and antimicrobial effects of cinnamon phenylpropanoids (Shinde et al., 2023) [30] Limited direct data; activity usually attributed to co-occurring cinnamaldehyde/eugenol; may support overall antifungal effect (Sharma et al., 2016) [29] Eugenyl acetate 17.862 0.52 Phenylpropanoid ester Biotransformation product of eugenol; shares aromatic backbone, contributes to aroma and mild antimicrobial/antioxidant effects (Shinde et al., 2023) [30] Specific anti-Aspergillus data scarce; usually discussed with eugenol-rich EOs (Pandey et al., 2017) [22] Benzyl benzoate 21.181 10.24 Aromatic ester Acaricidal and insecticidal; used clinically for scabies/lice; repellent activity; some antimicrobial contribution in EOs (Saltan et al., 2024) [27] Primarily insecticidal; EOs rich in benzyl benzoate show activity vs yeasts and Saccharomyces; direct anti-Aspergillus data limited (Farjam, 2012) [11] Benzaldehyde 8.916 0.15 Aromatic aldehyde Flavoring and preservative; antimicrobial, antioxidant, antiinflammatory, sometimes enhances Used as antibacterial /anti-fungal preservative; component of antimicrobial EOs, but usually as antibiotic activity (Aljaafari et al., 2022) [4] minor contributor (Xiao et al., 2024) [36] Benzyl alcohol 10.474 0.23 Aromatic alcohol

Widely used bacteriostatic/antifungal preservative in pharmaceuticals and cosmetics; local anesthetic properties (European Medicines Agency, 2017) [10] Provides preservative-level antimicrobial activity; usually supportive rather than primary antifungal agent in EOs (European Medicines Agency, 2017) [10] Others -1.63

Mixed minor constituents

Mixture of low-abundance terpenes/phenolics that may synergize with major compounds in antimicrobial and antioxidant activity (Pandey et al., 2017) [22] Often contribute to synergy but individual roles not defined 2 L. nobilis β-Pinene 6.300 3.17 Monoterpene hydrocarbon Antimicrobial, antioxidant, antiinflammatory; contributes to respiratory and bronchodilatory effects of coniferous EOs (Bhavaniramya et al., 2019) [7] EOs rich in α/β-pinene show antifungal activity vs plant pathogens and Aspergillus spp.

(Kim et al., 2018) [14] 1,8-Cineole [7] Contributes to antifungal activity of citrus and laurel EOs; inhibits storage molds at higher concentrations (Bhavaniramya et al., 2019) [7] β-Linalool 7.925 9.90 Monoterpene alcohol

As above for linalool: antimicrobial, antioxidant, antiinflammatory, CNS-active (sedative, anxiolytic, antidepressant) (Kamatou and Viljoen, 2008) [13] Inhibits A. flavus and other fungi; effective as biofumigant; antifungal vs Candida spp. (Li et al., 2022) [17] 1-Terpinene (terpinene-type monoterpene) 9.275 2.10 Monoterpene hydrocarbon/terpenoid Part of terpinene/terpineol family; antioxidant, antimicrobial, antiinflammatory; contributes to respiratory and skin-care effects (Bhavaniramya et al., 2019) [7] Terpinen-rich EOs show fungicidal activity vs Aspergillus and dermatophytes, usually synergistic with other monoterpenes (Pandey et al., 2017) [22] Benzaldehyde 9.467 10.96 Aromatic aldehyde See row above: antimicrobial, antioxidant, food preservative, sometimes enhances antibiotic effects (Aljaafari et al., 2022) [4] Antibacterial and antifungal preservative; part of EO antimicrobial spectrum, but rarely dominant antifungal driver (Xiao et al., 2024) [36] Eugenol 11.525 50.40 Phenylpropanoid phenolic ether

As above: strong antioxidant, antimicrobial, analgesic, antiinflammatory, potential neuroprotective; major contributor to clove/bay activity (Ulanowska and Olas, 2021) [34] Significant antifungal action against Aspergillus spp. and other pathogens; damages cell envelope and inhibits virulence factors (Didehdar et al., 2022) [9] Caryophyllene (βcaryophyllene) 12.358 10.23 Sesquiterpene hydrocarbon Analgesic, antioxidant, antimicrobial, potent antiinflammatory and immunomodulatory via CB2 agonism; neuroprotective and cardioprotective potential (Gushiken et al., 2022) [12] Contributes to antifungal and antibacterial effects of spice EOs; often synergistic with phenylpropanoids (Gushiken et al., 2022) [12] Others -7.99 Mixed minor constituents Monoterpenes, sesquiterpenes and phenylpropanoids that modulate aroma, volatility and contribute to antioxidant/antimicrobial synergy (Caputo et al., 2017) [8] Likely to enhance or modulate antifungal action in combination with major components

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