Silicon dioxide (SiO2) nanoparticles affect the morphology, sporulation, mycotoxin production, and pathogenicity of Fusarium brachygibbosum infecting olive trees

Introduction: This study investigates the In Vitro antifungal activity of silicon dioxide nanoparticles (SiO₂ NPs) against mycotoxigenic Fusarium brachygibbosum species, a fungus posing a significant threat to olive trees in Tunisia.

Methods: Two different doses of SiO₂ NPs (100 and 200 mg kg ^-1 ) were used to evaluate its effect on fungal growth, mycotoxin production, and virulence capability of tested F. brachygibbosum strain.

Results and Discussion: While mycelial growth was not influenced by SiO₂ NPs, a notable increase in macroconidia sporulation was observed at the highest dose tested. Scanning electron microscopy revealed structural alterations in fungal hyphae treated with SiO₂ NPs, including hyphal disorganization after the adherence of nanoparticles. Furthermore, SiO₂ NPs influenced oxidative stress in Fusarium, with varying effects on hydrogen peroxide levels, total antioxidant activity, and total phenolic compounds, modulating the capability of the fungus to produce mycotoxins. Indeed, fusaric acid and 15-acetyldeoxynivalenol amounts decreased in presence of SiO₂, while an increasing level of neosolaniol and diacetoxyscirpenol was observed. Pathogenicity tests on olive and sorghum leaves revealed a reduction of disease severity in SiO₂ treated samples compared to untreated controls, showcasing the potential of SiO₂ NPs as a sustainable alternative for managing Fusarium infections. These findings underline the potential use of SiO₂ NPs as environmentally friendly and effective tool in integrated pest management strategies against F. brachygibbosum as well as other Fusarium species occurring on olive trees. Further research is warranted to optimize their application and understand their interactions with both the pathogen and the host plant.

Highlights

• Fusarium brachygibbosum, an emerging olive tree pathogen in Tunisia, poses a significant threat to agriculture and requires innovative management solutions.

• Silicon dioxide nanoparticles (SiO₂ NPs) were tested for their effects on Fusarium, marking the first such study in Tunisia.

• While SiO₂ NPs did not inhibit mycelial growth, they induced pigmentation and significantly increased macroconidia sporulation at the highest concentrations (200 ppm).

• SiO₂ NPs induced an oxidative stress and a loss of membrane integrity in F. brachygibbosum.

• SiO₂ NPs influenced mycotoxin production in F. brachygibbosum reducing trichothecenes production

• SiO₂ NPs reduced Fusarium pathogenic effects on olive and sorghum leaves, showing promise as an alternative to chemical fungicides.

1 Introduction

Fusarium species are among the most significant phytopathogenic and mycotoxigenic fungi worldwide, causing severe diseases on various economically important crops (Munkvold et al., 2021). Although traditionally associated with major losses in cereals and vegetables, Fusarium infections are increasingly reported in olive trees showing symptoms of dieback in several countries worldwide (Nicoletti et al., 2020), including Morocco (Chliyeh et al., 2017), Spain (Hernández et al., 1998), Greece (Markakis et al., 2021), and Tunisia (Gharbi et al., 2020; Trabelsi et al., 2017).

This rise can be attributed to changes in olive cultivation methods, the expansion of new plantations into Fusarium-infected areas, and the use of infected planting material. Although Fusarium wilt has not yet reached the severity observed with other pathogens such as Verticillium species, Xylella fastidiosa, Pseudocercospora, and Venturia oleaginea, studies suggest that it has the potential to become more aggressive, similar to its impact on cereals and vegetables (Verma et al., 2024; Al-Hashimi et al., 2025).

Numerous agronomic strategies, including resistant varieties, chemical treatments, and crop rotation have been proposed to control Fusarium. Among these strategies, treatment with synthetic fungicides is the most effective tool against phytopathogenic Fusarium species. However, the repeated use of chemicals with the same mode of action could lead to a strong selection of resistant strains (Liu et al., 2019a; Yin et al., 2009; Yuan and Zhou, 2005), making the management of fungal diseases more difficult. Moreover, integrated pest management guidelines and environmental sustainability policies encourage researchers to develop newer eco-friendly strategies to control both fungal species and their mycotoxin accumulation, to reduce chemical treatments and, consequently, their negative environmental impact.

Recently, the new technologies and new knowledge on nanoparticles have encouraged researchers to develop effective nanoparticle-based bio-stimulants and pesticide more eco-friendly than chemical compounds (Duhan et al., 2017; Bhagat et al., 2023; Kumari et al., 2023). In addition, the nanoparticle based products allow the precise release of necessary and sufficient amounts of active ingredients and thus minimal residual presence, and release in the environment (Santosh P. and Yojana S., 2022). Notably, a reduced graphene oxide–copper oxide nanocomposite demonstrated superior antifungal activity at a significantly lower concentration (1 mg/L) compared to a conventional fungicide (2.5 g/L) in controlling Fusarium diseases in tomato and pepper plants (El-Abeid et al., 2020), highlighting the potential of nanotechnology in plant protection.

Within this framework, silicon, known as second most abundant element on earth after oxygen (Shahzad et al., 2022), was stated to have an important role in mitigating different stresses occurred on plants (Coskun et al., 2016; Elsheery et al., 2020). Especially its nanosized form which has shown promising results due to its good physical properties than bulk material (Siddiqui et al., 2014). Used as a pesticide, silica nanoparticles have shown potential in controlling various pests such as insects (Ziaee and Ganji, 2016). Moreover, studies who focused on the impact of silicon NPs on the interaction plant-pathogen have demonstrated the capability of silica nanoparticles in stimulating plant’s defense mechanisms and controlling phytopathogenic microorganisms (Baazaoui et al., 2021; Bathoova et al., 2021; Hassan et al., 2022; Parveen and Siddiqui, 2022). For instance, Yobo et al. (2019) reported that granular and foliar potassium silicate had a positive effect in controlling Fusarium Head Blight disease of wheat. Likewise, Sakr, (2021a) and Sakr & Shoaib, (2021) demonstrated that treatment of 1.50 g kg^-1 of soil and 1.7 mM soluble silicon reduced Fusarium Head Blight disease of wheat and barley. Previous studies suggest that SiO₂ NPs may affect fungal cells by disrupting membrane integrity through their accumulation on the cell wall. This accumulation can interfere with the transmembrane energy cycle, disrupt the electron transport chain, and compromise the overall structural integrity of the fungal cells (Derbalah et al., 2018; Abdelrhim et al., 2021). While SiO₂ NPs are reported to have antifungal properties, the underlying molecular mechanisms remain poorly understood.

This study aims to evaluate the in vitro antifungal potential activity of silicon dioxide nanoparticles (SiO₂ NPs) against F. brachygibbosum, an emerging pathogen threatening Tunisian olive trees and recognized as a potential Union quarantine pest (Bragard et al., 2021). This research pioneers the use of SiO₂ NPs as a novel strategy to control F. brachygibbosum, offering a promising tool to managing this emerging olive tree pathogen.

2 Material and methods

2.1 Nanoparticle synthesis

The synthesis of SiO₂ nanoparticles was carried out using the sol-gel process combined with supercritical ethanol drying. The preparation began with a solution containing tetraethyl orthosilicate (TEOS), ethanol, and distilled water, as described by Martínez et al. (2006). This mixture was stirred for 30 min to form a transparent sol. Subsequently, hydrofluoric acid (HF) was added gradually to the sol. The resulting mixture was subjected to supercritical drying to produce an aerogel. This step involved placing the sol in an ethanol-filled autoclave, which was heated to 243 °C and pressurized above ethanol’s critical point (Tc = 243 °C, Pc = 63 bar). Gelation occurred after maintaining the sol at this temperature for 1 h. The interstitial solvent was removed through a controlled depressurization process over 1 h using nitrogen gas. To prevent thermal strain and cracking, the autoclave was kept sealed for 24 h, allowing thermal equilibrium before it was gradually opened. The characteristic of the obtained SiO₂ aerogel was thoroughly detailed by Sghaier-Hammami et al. (2024).

2.2 Fungal isolation and identification

An orchard in the Kairouan region, located in southwestern Tunisia (35,67°N; 10,10°E), a major area for olive cultivation, was inspected for the presence of olive trees showing dieback symptoms on the aerial parts. A representative sample of olive roots was collected and carried to the laboratory for further analysis. For fungal isolation, root portions were thoroughly rinsed under running tap water for 15 min to remove soil and debris, dried on filter paper, and surface-sterilized using 2% (v/v) sodium hypochlorite for 2 min. They were then rinsed three times for 1 min with sterile distilled water and dried again on sterile filter paper. Small pieces from the margin of symptomatic tissues were transferred on Potato Dextrose Agar (PDA) medium amended with 100 mg L^-1 streptomycin sulfate and 50 mg L^-1 neomycin. Petri dishes were incubated at 25 °C for 7 days under an alternating light/darkness cycle of 12 h photoperiod.

After incubation, colonies developed from root pieces were transferred on new PDA plates and then purified by using the single spore isolation technique. All monosporic strains identified as Fusarium were transferred on PDA and Spezieller Nahrstoffarmer agar (SNA) media and morphologically identified according to Leslie & Summerell (2006). Morphological identification of each strain was also confirmed by sequencing the translation elongation factor 1-α gene by using the primer pair EF1/EF2 (O’Donnell et al., 1998), according to the procedure reported by Abi Saad et al. (2022).

2.3 Treatment of Fusarium brachygibbosum strain with nanoparticles

In order to test the effect of the SiO₂ NPs on fungal growth of F. brachygibbosum species, two different doses were used (100 and 200 ppm). In detail, SiO₂ NPs was added to PDA cooled down to 45°C–50°C after autoclaving and stirred for 10 min on a magnetic stirrer to ensure its homogeneous dispersion in the medium.

To evaluate the effect on fungal growth, F. brachygibbosum strain, MB1 stored in ITEM collection (Agro-Food Microbial Culture Collection, ISPA-CNR Bari, Italy, http://server.ispa.cnr.it/ITEM/Collection/) was used for mycelial growth assay. Briefly, mycelium disks (4 mm in diameter) from actively growing margins of 3–5-day old colonies cultured on PDA, were used to inoculate Petri dishes (90 mm in diameter) containing PDA or PDA amended with SiO₂ NPs. The inhibition activity of the SiO₂ NPs on colony growth was determined by measuring the diameter (in mm) of developing colonies after 3, 5, 7, and 10 days of incubation at 25°C, under darkness. The number and size of macroconidia and chlamydospores were also assessed after 7 and 14 days of incubation, respectively. The experiment was conducted with five replicates per treatment, and the trial was repeated twice.

2.4 In vitro pathogenicity test on detached olive leaves and stems, and shoots sorghum

The In vitro pathogenicity test on detached olive leaves was performed as described by Liu et al. (2007). To test the virulence of MB1 F. brachygibbosum strain treated with SiO₂, olive leaves and stems from the Chemlali cultivar, and leaves of sorghum were collected and injected with conidial suspension. The conidial suspension was prepared by carefully scraping the fungal culture from the amended medium, ensuring that no sterile distilled water was added to the surface to prevent any potential influence of SiO₂ NPs on the leaves.

Samples were collected at the same foliar stage and had approximately the same dimensions. For each treatment, six samples were placed in rectangular Petri dishes on soaked paper moistened with sterilized distilled water. For the control treatment, distilled water was injected into the samples, using the same number of replicates as for the other treatments. An aliquot of 10 µL of conidial suspension (10⁶ spore mL^-1) or distilled water was injected into the center of each leaf to ensure uniform conditions for all replicates.

2.5 SEM observations

For the morphological and chemical analysis of the fusarium samples, a Thermo Scientific™ Quanta™ 250 FEI scanning electron microscope (SEM), equipped with an Energy-Dispersive X-ray Spectroscopy (EDX) system, was used. This microscope is manufactured by Thermo Fisher Scientific Inc., located in Waltham, Massachusetts, USA.

The SEM provides high-resolution imaging of the sample surfaces at scales ranging from micrometers to nanometers. It uses an electron beam that interacts with the material, generating signals that are converted into images representing the surface topography. The integrated EDX system enables detailed chemical analysis by detecting and quantifying the elements present in the samples based on the X-ray emissions generated during electron beam interaction. This technique allowed for the identification of chemical elements and the mapping of their distribution within the samples.

Analyses were performed under low vacuum conditions to preserve the integrity of biological and non-conductive samples. Images were obtained at various magnifications to examine surface structures and elemental characteristics. Measurements were carried out in low-voltage mode to reduce the risk of damage to sensitive samples and to enhance electron penetration depth in the materials. The spatial resolution provided by this instrument allowed for the examination of fine surface details, and EDX analysis enabled the determination of the chemical composition at micro- and nanoscale levels.

2.6 Image analysis

Infected zone rate was calculated using the software Image Plus (Motic, version 2.0, 2008) using the formula:

$\text{Infected\hspace{0.17em}zone\hspace{0.17em}rate}\left(\%\right)=\frac{\text{Infected}\text{leaf}\text{surface}}{\text{Leaf}\text{surface}}\times 100$

For each leaf, the surface area of visible symptoms (necrotic or chlorotic spots) was measured and compared to the total leaf surface

Additionally, hyphal deformation severity was assessed by measuring the mycelium diameter in SEM images using ImageJ software (National Institutes of Health, Bethesda, MD, United States).

2.7 Cell membrane integrity

The integrity of the cell membrane of F. brachygibbosum was assessed by UV-VIS spectrophotometry following the method described by Ribeiro et al. (2024), with modifications. Specifically, a sporal suspension of the three treatments, adjusted to approximately 10⁶ spore mL^-1, was prepared in sterilized distilled water and incubated at 37°C for 2 h. After incubation, the samples were centrifuged, and the supernatant was collected for analysis. The absorbance of the supernatant was measured at 260 nm and 280 nm using a UV-VIS spectrophotometer (UV Spectrophotometer OPTIZEN 3220UV, Daejeon, South Korea).

2.8 Non-enzymatic antioxidant parameters measurements

The total antioxidant activity was measured following the method described by Koleva et al. (2002). Results were expressed in milligrams of gallic acid equivalents per Gram of dry weight (mg GAE/g DW), as outlined in the same study. The calibration curve was constructed within a concentration range of 0–500 μg mL^-1.

The total phenolic content of different species was assessed using a modified version of the Folin-Ciocalteu method as described by Pereira et al. (2018). Phenolic content was expressed as milligrams of gallic acid equivalents per Gram of dry residue (mg GAE/g DR), calculated from a calibration curve prepared with gallic acid standards in the range from 0.001 to 0.01 mg L^-1.

Hydrogen peroxide (H₂O₂) content in fungi was determined using a modified method based on Velikova et al. (2000). Fungi were rapidly frozen in liquid nitrogen and ground in 5 mL of 0.1% (w/v) trichloroacetic acid. The homogenate was centrifuged at 15,000 × g for 15 min at 4 °C to separate the supernatant. An aliquot of 0.5 mL of the supernatant was combined with 1 mL of 1 M KI and 0.5 mL of 10 mM K₂HPO₄ buffer (pH 7.0). The reaction mixture was kept in darkness for 60 min before measuring absorbance at 390 nm. The H₂O₂ concentration was calculated using a standard curve prepared beforehand.

2.9 Mycotoxins analysis

The effect of SiO₂ NPs, at two different concentrations 100 and 200 mg kg^-1, on mycotoxin production was evaluated culturing F. brachygibbosum MB1 strain on PDA and PDA amended with the compound. After 14 days of incubation at 25 °C in darkness, mycelial plugs (2 g) were collected in 50 mL tubes containing 10 mL acetonitrile/water/acetic acid (79:20:1, v/v/v). Mycotoxin extraction was obtained by 3 min of slurried. Extracts were diluted in water (ratio 1:10) and directly injected into the LC- HRMS instrument. Liquid chromatography–HMRS analysis was performed on Orbitrap ExplorisTM 240 E, equipped with a heated electrospray ion source (HESI II) coupled to a Vanquish UHPLC system (all from Thermo Fisher Scientific, San Jose, CA, United States). Chromatographic separation was performed at 40 °C on a Gemini^® C₁₈-column, 150 × 2 mm i. d., 5 μm particle size, equipped with a C₁₈ 4 × 3 mm i. d. security guard cartridge (Phenomenex, Torrance, CA, United States). The flow rate of the mobile phase was 350 μL/min, whereas the injection volume was 10 µL. Eluent A was water, and eluent B was methanol, both containing 0.5% acetic acid and 1 mM ammonium acetate. The HESI II ion source was operated in positive ion mode and in negative mode, with the following settings: sheath gas: 30 arbitrary units, auxiliary gas: five arbitrary units, sweep gas 1 arbitrary units, spray voltage: 3.5 kV for positive ion and 2,5 KV for negative ion, RF lens: 70 percentage (arbitrary units), ion transfer tube: 325°C, vaporizer temperature: 300°C. High-resolution mass spectrometry chromatograms were acquired in positive and negative ionization mod in the same run. Resolution was 120,000 and the scan range (m/z): 70-850. Mycotoxin identification was carried out by comparing retention times, fragmentation patterns, and collision cross-sections with reference standards compiled in our in-house UNIFI library. This library was built by analyzing a mixture of mycotoxin standards under the same analytical conditions.

2.10 Statistical analysis

All data are presented as means. Tukey’s HSD test was performed using one-way analysis of variance (ANOVA) by IBM SPSS software (Version 27.0. Armonk, NY: IBM Corp, United States) with a value of P < 0.05 being considered significant. Principal component and Pearson’s correlation analysis was executed utilizing the FactoMineR packages within the R software, version 4.2.3, with data normalized by applying Z-score standardization prior to PCA.

3 Results

3.1 Host symptoms and morphological characterization of Fusarium strains infecting olive roots

Leaves, roots, and stems were sampled from symptomatic olive trees cultivated in the semi-arid Kairouan region. The olive samples showed typical Fusarium infection symptoms, including leaf wilting, chlorosis, and necrosis along the midrib and edges. Stem cross-sections reveal vascular discoloration with brown streaks in the xylem, while roots appear dry, brittle, and necrotic with reduced branching (Figure 1).

Figure 1

Figure 1. Symptoms of Fusarium infection on olive trees: (a) wilted olive leaves with discoloration, (b) internal vascular browning in infected branches, (c) root damage and decay caused by the pathogen.

Only Fusarium strains with white to pink colonies were selected, as the focus is on studying F. brachygibbosum species, which are considered emerging pathogens. These strains have recently gained attention due to their characteristic abundant aerial mycelium, aligning with their increasing significance as a plant health threat highlighted by (Bragard et al., 2021).

Colonies grown on PDA reached a diameter of 90 mm at 28°C after 14 days in darkness, characterized by abundant aerial white mycelium, pink colonies (Figure 2a) and rare production of microconidia. Chlamydospores were commonly observed, ranging from 6 to 24 µm in diameter, with a spherical to globose shape, smooth or slightly roughened surfaces, and appearing terminally or intercalarily in chains of two or three (Figure 2b).

Figure 2

Figure 2. Cultural and morphological features of Fusarium brachygibbosum (a) microscopic view of Fusarium brachyggibosum conidia structures observed under compound microscope (b) (40X).

Molecular identification at species level was conducted using the internal transcribed spacer (ITS) region of rDNA and the translation elongation factor 1α (TEF-1α). Sequence homology confirmed the identification of the species as F. brachygibbosum.

3.2 Inhibitory effects of SiO₂ nanoparticles on fusarium growth

The effect of SiO₂ NPs on the fungal growth and structural development of F. brachygibbosum was evaluated in vitro conditions by using PDA and PDA amended with the compound. Two concentrations of SiO₂ NPs, 100 ppm and 200 ppm, were tested. Observations were conducted after 3, 5, 7, and 10 days of incubation, examining both the reverse and front views of the culture plates. In the control group, Fusarium growth exhibited consistent colony expansion and notable red pigmentation, which became more pronounced by day 10. In contrast, Fusarium cultured with SiO₂ NPs displayed slower growth and reduced pigmentation, with the inhibitory effect being more evident at the higher concentration of 200 ppm. By day 10, colonies grown with 200 ppm SiO₂ NPs showed smaller diameters and lighter pigmentation compared to both the control and the 100 ppm treatment. These findings suggest a dose-dependent inhibitory effect of SiO₂ nanoparticles on Fusarium growth and pigment production, with higher concentrations exerting a greater impact (Figure 3).

Figure 3

Figure 3. Impact of SiO₂ nanoparticles on Fusarium brachygibbosum mycelial growth over time on PDA medium: reverse and front views at 100 and 200 ppm compared to the control (NTF).

To quantify this effect, inhibition rates were measured over time. At 3 dpi, a slight mycelial growth inhibition was observed (3.45% and 1.53% on PDA amended with 100 ppm and 200 ppm, respectively). However, after 3 days, no significant difference was observed (p < 0.05) between control and treatments (Table 1). These results indicate that SiO₂ NP could influence fungal colony growth only in the early stages, but their effect diminishes over time. Interestingly, these findings imply that SiO₂ nanoparticles may primarily affect Fusarium structural components rather than directly inhibiting mycelium expansion.

Table 1

Table 1. Inhibition rate of growth of Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group.

Further structural analysis of F. brachygibbosum grown under different conditions revealed significant variations (Figure 4). In the control only the fungal hyphae and sparse structures were observed, with no visible macroconidia (Figure 4a). At 100 ppm SiO₂, hyphae and fungal fragments were present, but macroconidia were still absent (Figure 4b). However, treatment with 200 ppm SiO₂ led to the formation of macroconidia, clearly marked by their elongated, canoe-shaped morphology. These observations suggest that while lower concentrations of SiO₂ nanoparticles primarily affect fungal growth, higher concentrations promote macroconidia formation in F. brachygibbosum. This stimulatory effect on macroconidia formation at 200 ppm provides insight into the structural and developmental impacts of SiO₂ nanoparticles on fungal physiology.

Figure 4

Figure 4. Microscopic view of F. brachyggibosum conidia structures at 10x (a) and 40X (b) for.

The impact of SiO₂ nanoparticles (SiO₂ NPs) on F. brachygibbosum spore formation and morphology was assessed by counting the different structures of the fungus. Spore counts revealed clear differences between treatments. Chlamydospore production was highest in the untreated control, while macroconidia were absent in this condition. Treatment with 200 ppm SiO₂ NPs significantly increased the production of macroconidia, whereas the 100 ppm treatment resulted in minimal macroconidia and an intermediate level of chlamydospore production, suggesting a dose-dependent stimulation of macroconidia formation (Figure 5a). The assessment of spore area followed a similar trend. Macroconidia observed in the 200 ppm treatment had the largest area, significantly exceeding those in other treatments. In contrast, the area of chlamydospores remained consistent across the untreated and 100 ppm conditions, indicating a limited effect of lower SiO₂ concentrations on this structural feature (Figure 5b). Furthermore, the perimeter of macroconidia in the 200 ppm treatment was the largest, aligning with the elongated, canoe-shaped morphology noted in previous microscopic observations (Figure 4). Chlamydospores in the untreated and 100 ppm treatments exhibited significantly smaller perimeters, further highlighting the distinct morphological impact of the higher SiO₂ NP concentration (Figure 5c).

Figure 5

Figure 5. Assessement of spore number (a) Area (b) Perimeter (c) of Fusarium brachygibbosum spores grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group. Results represent the mean ± standard error (SE). Bars with a different letter indicate significant differences according to Tukey’s HSD test at P < 0.05.

These findings align with earlier results, showing that SiO₂ nanoparticles exert a limited effect on mycelial growth but significantly influence Fusarium structural development.

3.3 SEM analysis

The analysis of SEM images revealed significant differences in the morphology of Fusarium hyphae depending on the concentration of SiO₂ nanoparticles and the incubation time. At 3 days post-incubation (dpi), untreated Fusarium hyphae exhibited an organized filamentous structure (Figure 6a,d), while the addition of 100 ppm or 200 ppm SiO₂ induced progressive hyphal disorganization, with a more pronounced effect at 200 ppm (Figures 6b,e). At 8 dpi, untreated Fusarium showed dense hyphal proliferation, whereas SiO₂ treatments, particularly at 200 ppm, resulted in hyphal fragmentation and an apparent reduction in density.

Figure 6

Figure 6. Scanning electron micrographs of Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control. (a,d) Control fungi grown for 3 and 10 days post-incubation (dpi), respectively. (b,e): Fungi grown on PDA medium with 100 ppm SiO₂ for 3 and 10 dpi, respectively. (c,f) Fungi grown on PDA medium with 200 ppm SiO₂ for 3 and 10 dpi, respectively.

As a consequence, the assessment of hyphal deformation severity, indicated by mycelium diameter measurement, showed a significant reduction with increasing SiO₂ concentration. The control group exhibited the largest mycelium diameter (7.102 ± 0.463 µm), which was not significantly different from the group treated with 100 ppm SiO₂ (6.151 ± 0.404 µm). However, treatment with 200 ppm SiO₂ resulted in a significantly lower mycelium diameter (3.175 ± 0.158 µm), highlighting the inhibitory effect of higher SiO₂ concentrations on fungal growth (Table 2).

Table 2

Table 2. Mycelium diameter of Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group.

In the SEM micrographs of the treatments with 100 ppm and 200 ppm SiO₂, nanoparticles appeared to be visible on the surface of the hyphae. These particles manifested as small granular structures adhering to the hyphal walls, especially at 200 ppm (Figures 6c,f), where their density seems higher. This suggests direct interaction between the NPs and the hyphae, potentially contributing to the observed inhibitory effect. Such interactions could induce mechanical alterations, modify cell wall permeability, or disrupt normal biological functions of the hyphae.

3.4 Membrane integrity

The mean optical density values obtained at 260 and 280 nm were used to detect absorbing materials, reflecting the potential loss of macromolecules such as nucleic acids and proteins. This loss is associated with plasma membrane disruption and is evident in F. brachygibbosum treated with SiO₂ nanoparticles (Table 3). The control group exhibited low optical density values at both wavelengths, indicating intact fungal cells under the given culture conditions. Conversely, treatment with 100 ppm SiO₂ nanoparticles led to a significant rise in the optical density of fungal suspensions, suggesting increased membrane permeability. Surprisingly, the group treated with 200 ppm SiO₂ nanoparticles displayed the lowest absorbance values, a finding that will be further examined in the discussion.

Table 3

Table 3. Detection of absorbing material at 260 and 280 nm in suspensions of Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group.

3.5 Impact of SiO₂ on oxidative stress in Fusarium brachygibbosum

The impact of SiO₂ supplementation on H₂O₂ content and total antioxidant activity in Fusarium spp. was assessed in untreated Fusarium (NTF) and cultures grown on PDA supplemented with 100 ppm and 200 ppm SiO₂. The highest H₂O₂ content was recorded in the untreated control, with a significant reduction observed at 100 ppm SiO₂. However, at 200 ppm SiO₂, H₂O₂ levels increased compared to the 100 ppm treatment, though they remained slightly lower than those in the untreated control (Figure 7a). A similar trend was observed for total phenolic compound levels, illustrating the pattern of H₂O₂ variation (Figure 7b).

Figure 7

Figure 7. Assessment of the peroxyde hydrogen concentration (a) total polyphenols concentration (b) and total antioxidant activity (c) in Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group. Results represent the mean ± standard error (SE). Bars with a different letter indicate significant differences according to Tukey’s HSD test at P < 0.05.

The total antioxidant activity exhibited a dose-dependent response, with the lowest activity in the untreated control, a moderate increase at 100 ppm SiO₂, and the highest antioxidant activity observed at 200 ppm SiO₂ (Figure 7c).

These results indicate that 100 ppm SiO₂ reduces oxidative stress in F. brachygibbosum as evidenced by decreased H₂O₂ levels, while 200 ppm SiO₂ enhances antioxidant activity, potentially as a response to elevated H₂O₂ content.

3.6 Mycotoxin analyses of Fusarium brachygibbosum under SiO₂ NPS treatments

The mycotoxin analysis of F. brachygibbosum grown on PDA medium amended with SiO₂ nanoparticles (NPs) at concentrations of 100 and 200 ppm, compared to the non-treated fungal (NTF) control was able to highlight the change in the quantified mycotoxins including Fusaric Acid (FA), neosolaniol (NEO), 15-acetyldeoxynivalenol (3-15 AC DON), and 4,15-diacetoxyscirpenol (DAS) as represented in Table 4. In the non-treated Fusarium, FA was detected at 203.1 mg kg^-1, while 3-15 AC DON was present at 57.5 mg kg^-1, respectively. NEO and DAS were not detected under these conditions. Upon exposure to 100 ppm SiO₂ NPs, FA remained undetectable, whereas NEO was measured at 1308.4 mg kg^-1 3-15 AC DON was produced at a lower concentration (32.4 mg kg^-1), while DAS increased to 430.4 mg kg^-1. At the higher concentration of 200 ppm SiO₂ NPs, FA remained undetectable, while NEO levels further increased to 2197.3 mg kg^-1 3-15 AC DON was not detected in this condition, whereas DAS production rose to 734.2 mg kg^-1.

Table 4

Table 4. Trichothecenes production of Fusarium brachygibbosum grown on PDA medium amended with SiO₂ at concentrations of 100 and 200 ppm, compared to the control group.

These results indicate that SiO₂ NPs significantly influenced trichothecene biosynthesis in F. brachygibbosum, particularly enhancing neosolaniol and DAS production while suppressing FA synthesis. Additionally, 3-15 AC DON levels decreased with increasing SiO₂ NP concentrations, becoming undetectable at 200 ppm.

3.7 In vitro assessment of SiO₂ treatment on fusarium in olive and sorghum

The effect of SiO₂ NPs at two concentrations (100 and 200 ppm) was assessed on olive tree leaves and stems, and on sorghum shoots. Severe symptoms, including necrosis and browning on leaves and dark streaks or lesions on stems, indicating significant tissue damage, were observed on olive leaves inoculated with untreated F. brachygibbosum conidial suspension (Figure 8). Treatment with SiO₂ at 100 ppm reduced these symptoms, with leaves exhibiting milder necrosis and stems showing less browning and lesion formation. At 200 ppm, symptoms were further mitigated, with olive leaves displaying minimal necrosis and stems showing near-normal appearance with reduced streaking.

Figure 8

Figure 8. Binocular magnifier observations showing symptoms on olive tree leaves and shoots, and sorghum leaves after inoculation with sporal suspensions prepared under different treatments: Water, NTF (untreated), FSiO₂ at 100 ppm, and FSiO₂ at 200 ppm.

The measurement of the infected zone rate (%) validates the observed results in Figure 9. The untreated Fusarium showed the highest infection severity, with an infected zone rate exceeding 40%, which aligned with the severe necrosis, browning, and tissue damage observed in olive leaves and stems. In contrast, treatments with SiO₂ at 100 ppm and 200 ppm significantly reduce the infected zone rate to approximately 10%. These findings confirm the visual observations, where SiO₂ treatment, particularly at higher concentrations, mitigated symptoms such as necrosis, discoloration, and structural degradation, highlighting its effectiveness in reducing Fusarium-induced damage.

Figure 9

Figure 9. Infected surface of olive leaves treated with NTF, FSiO₂ 100 ppm and FSiO₂ 200 ppm compared to total leaf surface. Results represent the mean ± standard error (SE). Bars with a different letter indicate significant differences according to Tukey’s HSD test at P < 0.05.

For sorghum shoots, the NTF group exhibited extensive tissue degradation and mycelium development. Treatment with SiO₂ at 100 ppm led to moderate recovery, with a reduction in Fusarium mycelium development, while at 200 ppm, significant improvement was observed, with shoots showing minimal structural damage and reduced mycelium presence (Figure 8). These results support the observations made on olive trees and confirm the loss of F. brachygibbosum pathogenicity under SiO₂ treatment in both olive leaves and shoots and sorghum shoots.

3.8 Analysis of principal components and the relationships among various measured parameters

Based on the principal component analysis (PCA), the first two components, Dim1 (PC1) and Dim2 (PC2), explain 71.8% and 28.2% of the total variance, respectively, providing a comprehensive representation of the dataset’s variability (Figure 10). PC1 primarily separates FSiO₂ 100 ppm from FSiO₂ 200 ppm and NTF, driven by parameters such as H₂O₂ levels, infected zone rate, and fungal growth inhibition. In contrast, PC2 distinctly discriminates FSiO₂ 200 ppm from the other samples, with strong contributions from macroconidia-related traits (number, area, and percentage), chlamydospore parameters, and biochemical markers such as polyphenols and AAT activity. The PCA highlights a clear dose-dependent effect of FSiO₂ nanoparticles on Fusarium physiology and biochemistry, with higher concentrations (200 ppm) inducing more pronounced alterations. The clustering patterns and variable correlations suggest that FSiO₂ nanoparticles at higher concentrations effectively disrupt fungal growth and metabolism, with macroconidia and polyphenol-related traits emerging as key indicators of antifungal activity and stress responses.

Figure 10

Figure 10. Principal component analysis of (a) variables influencing the effect of SiO2 nanoparticles on Fusarium brachygibbosum and (b) treatments: non-treated fusarium, and fusarium treated with 100 ppm and 200 ppm SiO₂ nanoparticles.

4 Discussion

Fusarium species have frequently been isolated from olive tissues that display wilt symptoms in Tunisia (Trabelsi et al., 2017) probably due to the co-cultivation of olive trees with other crops susceptible to Fusarium infection. Several pathogenic Fusarium species were isolated from olive trees showing dieback and wilting symptoms, representing an increasing concern for olive producers. Indeed, some Fusarium species, including F. brachygibbosum, caused important damages in nurseries and newly established olive orchards (Trabelsi et al., 2017). Among the pest management strategies, treatment with synthetic fungicides is the most effective tool against phytopathogenic Fusarium species. However, despite the efficacy of chemical pesticides in reducing Fusarium growth, their heavy use can lead to the occurrence of resistant strains in the fungal populations (Lucas et al., 2015), as well as adverse environmental impact (Siamak and Zheng, 2018). For that reason, it is crucial to develop alternative eco-friendly tools for controlling Fusarium species (Balakumaran et al., 2015). In this study the activity of SiO₂ on F. brachygybbosum isolated from olive tree has been evaluated.

4.1 Impact of SiO₂ on mycelium growth and structure

The SiO₂ nanoparticles used in this study exhibit a spherical shape with a uniform and narrow size distribution, as mentioned in our previous work (Baazaoui et al., 2023). Their mesoporous structure and an average particle size of approximately 6 nm contribute to their distinctive physicochemical and morphological properties, which may facilitate effective contact with fungal cells. However, unlike previous studies that reported antifungal activity against Rhizoctonia solani (Abdelrhim et al., 2021), Alternaria (Derbalah et al., 2018), and Fusarium oxysporum (Parveen and Siddiqui, 2022), their presence did not inhibit mycelial growth. However, SEM analysis revealed significant structural damage to fungal hyphae treated with SiO₂ nanoparticles, including hyphal disorganization, irregular shrinkage, and nanoparticle adherence, particularly at higher concentrations. Similar structural damage was reported for Fusarium treated with ZnO or ZnO-CuO nanoparticles, where TEM analysis showed cell wall disintegration, plasma membrane damage, and destruction of intracellular contents (Gaber et al., 2024; Subba et al., 2024). The observed adherence of SiO₂ nanoparticles to the mycelium suggests a direct interaction with the fungal cell wall, potentially affecting its integrity and function. AgNPs have been shown to induce similar effects, forming pits and cavities on the Fusarium cell wall, leading to compromised structural integrity and increased permeability (ElSharawy et al., 2023; Todorova et al., 2024). These findings indicate that nanoparticles may disrupt fungal cell wall function, ultimately contributing to oxidative stress and cellular damage.

The harmful effects of nanoparticles on fungal mycelium are further supported by evidence that ions released from nanoparticles penetrate fungal cells, disrupting essential processes such as protein synthesis and DNA replication (Todorova et al., 2024). Our study confirmed these effects, showing DNA damage and loss of plasma membrane integrity when SiO₂ nanoparticles were added at 100 ppm to the Fusarium culture medium. However, the absence of effect at 200 ppm could be due to the observed adherence of nanoparticles to the mycelium surface at this concentration. The adherence of nanoparticles to microbial cell membranes, whether through direct or electrostatic interactions, influences membrane permeability (Zhang et al., 2007). This interaction may physically target the membrane, preventing further loss of integrity.

In response to this stress, F. brachygibbosum exhibited increased macroconidia sporulation, while chlamydospore production remained unaffected. Asexual spores play a crucial role in fungal survival, enabling colonization of new resource patches, particularly in suboptimal and disturbed environments (Wyatt et al., 2013; Boddy and Hiscox, 2016). Interestingly, stress-induced sporulation has been reported in fungi exposed to resource limitations, where increased investment in spore production allows fungal populations to persist under adverse conditions (Camenzind et al., 2022). Although asexual reproduction is a key factor in the rapid spread of fungal disease outbreaks (Ashu and Xu, 2015), the structural damage induced by SiO₂ NPs may influence the viability or pathogenic potential of the resulting spores, potentially affecting fungal dissemination and host infection dynamics. Given these findings, the increased sporulation observed in F. brachygibbosum following SiO₂ treatment may not indicate enhanced fungal proliferation but rather a stress-induced survival mechanism aimed at ensuring persistence under unfavorable conditions. This disruption in fungal cell integrity and function may also contribute to oxidative stress, further exacerbating cellular damage.

4.2 Impact of SiO₂ on fungal oxidative systems and mycotoxin production

SiO₂ nanoparticles demonstrated a concentration-dependent impact on oxidative stress and antioxidant activity in F. brachygibbosum. Silicon nanoparticles (SiNPs) have been shown to interact with cellular membranes and induce oxidative stress in various organisms. Specifically, SiO₂ NPs can damage phospholipid membrane integrity and fluidity, potentially leading to cytotoxicity (Wei et al., 2015). At 100 ppm, SiO₂ significantly reduced oxidative stress by lowering H₂O₂ levels and total polyphenol content, indicating that this concentration effectively mitigates oxidative damage, possibly by enhancing cellular defense mechanisms.

In contrast, at 200 ppm SiO₂, H₂O₂ levels increased compared to the 100 ppm treatment, although they remained lower than those in the untreated group. This suggests that at higher concentrations, SiO₂ exposure may disrupt redox balance, potentially through direct oxidative damage to cellular components. Furthermore, mesoporous silica nanoparticles have been found to induce intracellular peroxidation damage in Phytophthora infestans by selectively triggering excess production of reactive oxygen species (ROS), offering potential as a green fungicide for controlling late blight in potatoes (Chen et al., 2023). A similar mechanism may be involved in F. brachygibbosum, where the observed increase in total antioxidant activity at 200 ppm suggests an adaptive response to counteract ROS accumulation.

At the proteomic level, previous studies on AgNPs have shown that brief exposure of E. coli leads to changes in the expression of envelope proteins (OmpA, OmpC, OmpF, OppA, MetQ) and heat shock proteins (IbpA, IbpB, and ribosomal subunit 30S S6), indicating that the cell membrane is a primary target of nanoparticle-induced stress (Rodrigues et al., 2024). Similarly, SiO₂ NPs may interact with fungal membrane-associated proteins, potentially affecting cell wall integrity and cellular homeostasis. Proteomic analyses of bacteria treated with AgNPs have also revealed elevated levels of antioxidant enzymes such as superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD), along with hydroperoxide resistance proteins, suggesting that oxidative stress plays a central role in nanoparticle toxicity (Liao et al., 2019). Given that SiO₂ NPs induce oxidative stress in F. brachygibbosum, a similar proteomic response involving antioxidant defenses may occur in fungi.

The antimicrobial activity of AgNPs has been attributed to a synergistic effect of silver ion release and nanoparticle-specific interactions (Yan et al., 2018). While SiO₂ NPs do not release metal ions in the same manner, their ability to generate ROS and interact with cellular membranes suggests that they may share common cytotoxic mechanisms with AgNPs. However, the precise mode of action of SiO₂ NPs remains unclear. Further studies are needed to elucidate the proteomic changes in fungal cells exposed to SiO₂ NPs, particularly in membrane-associated proteins and oxidative stress response pathways, to better understand their antifungal potential.

Since oxidative stress is known to influence fungal secondary metabolism, these changes in redox balance may also impact the biosynthesis of mycotoxins (Reverberi et al., 2010). Given the observed oxidative stress modulation by SiO₂ nanoparticles, we investigated whether these changes correlated with alterations in trichothecene production by F. brachygibbosum.

Our analysis of mycotoxin profiles revealed a distinct pattern dependent on SiO₂ NP concentration. Fusaric acid production was completely inhibited in presence of SiO₂. Previous studies have demonstrated the virulence of this toxin, showing that deletion of key regulatory genes such as FUB1 and FUB4 resulted in reduced FA synthesis and decreased virulence of Fusarium oxysporum in both plant and mammalian hosts (Ding et al., 2018; López-Díaz et al., 2018). Additionally, FA has been shown to increase host susceptibility to fungal invasion (Liu et al., 2019b), suggesting that its inhibition by SiO₂ NPs could directly impact the pathogenicity of F. brachygibbosum. Supporting this, Singh et al. (2017) demonstrated that FA alone can induce Fusarium wilt symptoms in tomato plants, triggering cell death, oxidative stress, and disruptions in photosynthesis and protein expression upon injection of purified FA into tomato leaves.

In contrast to FA, SiO₂ NPs significantly increased NEO production. Previous studies demonstrated that 100 ppm silver nanoparticles reduced NEO production by 9% in F. chlamydosporum (Bakri et al., 2020). In our study, the inhibition of FA production under SiO₂ treatments suggests a shift in metabolic flux toward alternative pathways, including NEO biosynthesis (Wei et al., 2022). The absence of FA production could have allowed more precursors to be diverted toward the NEO biosynthesis pathway. Alternatively, oxidative stress induced by nanoparticles may have influenced mycotoxin biosynthesis in a different way. Ferrigo et al. (2021) reported that H₂O₂ treatment at a lower concentration (1 mM) caused an almost total disappearance of MON and a strong reduction of FBs content in two Fusarium species. This suggests that oxidative stress could either suppress specific toxin biosynthesis pathways or trigger compensatory mechanisms leading to increased production of other secondary metabolites. A similar trend was observed with the trichothecene toxin DAS which increased concentration with the increasing dose of nanoparticles. However, 15-Acetyldeoxynivalenol production progressively declined with increasing nanoparticle doses, reaching undetectable levels at highest tested dose.

4.3 Impact of SiO₂ on the reduction of fungal pathogenicity

Patogenicity test revealed a reduced pathogenic effect of F. brachygibbosum conidia treated with SiO₂ NPs. Similarly, reduction in H₂O₂ and increasing of total antioxidant activity were observed in F. brachygibbosum after treatment. This change in H₂O₂ levels played a significant role in impairing the pathogen’s ability to form infection structures necessary for effective host colonization (Scott and Eaton, 2008). ROS, including H₂O₂, are known to act as signaling molecules crucial for fungal differentiation, particularly in the development of structures such as appressoria, which facilitate plant penetration. NADPH oxidases (Nox) are key enzymes responsible for the localized generation of ROS during such processes, and their activity is often essential for fungal virulence (Zhang et al., 2020). Previous studies have demonstrated that fungal mutants with diminished ROS production due to impaired Nox activity exhibit delayed or defective development of primary lesions and a reduction in the functional efficiency of appressoria (Heller and Tudzynski, 2011). This aligns with our observations of reduced necrotic symptoms in leaves inoculated with treated Fusarium, suggesting that the treatment may have disrupted the ROS-mediated signaling pathways required for the differentiation of pathogenic structures. This phenomenon is similar to the mode of action of Succinate Dehydrogenase Inhibitor (SDHI) fungicides which impair spore germination by inhibiting succinate dehydrogenase (SDH) (Xu et al., 2019). This disruption prevents the development of infection structures necessary for host colonization, ultimately limiting fungal pathogenicity. Given this similarity, our findings suggest that silicon nanoparticles may have the potential to replace certain conventional pesticides by targeting fungal pathogenicity through comparable mechanisms.

To our knowledge, most studies on the role of SiO₂ NPs in plant-pathogen interactions focused on their ability to enhance plant defense mechanisms (Suriyaprabha et al., 2014; Albalawi et al., 2022; Goswami et al., 2022; Parveen and Siddiqui, 2022). Few in vitro studies have been conducted, based only on macroscopic observations rather than molecular analyses (Qin and Tian, 2005). Sakr (2016) demonstrated that the inhibition of Fusarium mycelial growth observed in silicon-amended PDA was due to the alkaline pH rather than silicon itself, suggesting that silicon does not have intrinsic antifungal properties. However, the same author hypothesized that the SiO₂ NPs ability to reduce fungal infection severity was due to host-mediated resistance (Sakr, 2021b).

While SiO₂ NPs show potential as nanofertilizers and biopesticides (Sghaier-Hammami et al., 2024), further research is needed to fully understand their long-term impacts on ecosystems and optimize their use in sustainable agriculture (Ahmad et al., 2023). Although SiNPs can potentially enhance soil microbiome and fertility (Rajput et al., 2021; Sghaier-Hammami et al., 2024), there are concerns regarding the accumulation of nanoparticles in soil, which may adversely affect soil microbiota, fertility, and plant growth, as reported by several researchers (Grün et al., 2018; Javed et al., 2019; Baazaoui et al., 2021). An important yet unexplored question is whether fungi can develop resistance to SiO₂ NPs, similar to what has been observed with silver nanoparticles. Studies on silver have shown that fungi can adapt through genetic mechanisms, such as mutations in the RLM1 gene, which strengthens cell wall integrity, or the activation of copper-transporting ATPases, which help detoxify metal stress (Antsotegi-Uskola et al., 2017; Terzioğlu et al., 2020). Investigating whether similar resistance mechanisms exist for SiO₂ NPs is crucial for assessing their long-term antifungal efficacy and environmental impact. Further research is imperative to better understand the metabolic activity of F. brachygibbosum in the presence of SiO₂, not only to explore its fungicidal potential but also to assess its environmental implications. Such inquiries hold the potential to open novel pathways in pesticide research, where silicon could serve not only as a bio-stimulant but also as a fungicide against Fusarium infections in olive trees.

5 Conclusion

This study sheds light on the potential of SiO₂ NPs as a sustainable alternative to conventional fungicides for managing F. brachygibbosum, a pathogen threatening olive tree cultivation in Tunisia. Despite SiO₂ NPs not exhibiting direct inhibitory effects on mycelial growth, significant alterations in fungal physiology and mycotoxin production were observed. These findings suggest that SiO₂ NPs interact with the fungus in ways that modify its developmental processes rather than directly inhibiting its growth. Pathogenicity tests further demonstrated that Fusarium treated with SiO₂ NPs caused less severe disease symptoms on olive leaves. This reduction in disease severity aligns with the observed changes in fungal sporulation and highlights the ability of SiO₂ NPs to mitigate the pathogen’s virulence. The study also revealed that SiO₂ NPs influenced oxidative stress within the fungus, as evidenced by altered H₂O₂ levels and increased antioxidant activity and total polyphenol compounds. The findings underscore the importance of exploring alternative and environmentally friendly strategies to address the challenges posed by Fusarium species in agriculture. While this study provides a foundation for using SiO₂ NPs as a tool for fungal management, further research is essential to elucidate their precise mechanisms of action and optimize their application. In particular, comparing their effects with non-nanostructured SiO₂ would help determine whether the observed pathogenicity reductions are specifically due to nanoscale properties. Although the application of nanoparticles in the field, outside of laboratory conditions is still restricted due to the lack of information on the toxicity of these compounds and their impact on the environment, the preliminary results of this study should encourage research to delve in depth in this aspect to minimize the use of conventional fungicides and limit their deleterious impact.

Data availability statement

The original contributions presented in the study are included in the article, further inquiries can be directed to the corresponding author.

Author contributions

MB: Conceptualization, Writing – original draft, Writing – review and editing, Investigation, Methodology, Validation. BS-H: Conceptualization, Methodology, Project administration, Supervision, Validation, Visualization, Writing – original draft, Writing – review and editing. MM: Investigation, Methodology, Validation, Writing – review and editing. HN: Methodology, Writing – review and editing. SS: Formal Analysis, Methodology, Validation, Writing – review and editing. LG: Methodology, Writing – review and editing. RS: Methodology, Writing – review and editing. MM: Methodology, Writing – review and editing. SL: Investigation, Methodology, Validation, Writing – review and editing. AM: Investigation, Supervision, Writing – review and editing. SBMH: Investigation, Supervision, Validation, Writing – review and editing.

Funding

The author(s) declare that financial support was received for the research and/or publication of this article. This work was supported by the Tunisian Ministry of Higher Education, Scientific Research and Technology under grant number PEJC2023-D4P03.

Acknowledgments

The authors extend their appreciation to the laboratory members for their technical help.

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.

Generative AI statement

The author(s) declare that no Generative AI was used in the creation of this manuscript.

Publisher’s note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

References

Abdelrhim, A. S., Mazrou, Y. S. A., Nehela, Y., Atallah, O. O., El-Ashmony, R. M., Dawood, M. F. A., et al. (2021). Silicon dioxide nanoparticles induce innate immune responses and activate antioxidant machinery in wheat against Rhizoctonia solani. Plants 10, 2758. doi:10.3390/plants10122758

PubMed Abstract | CrossRef Full Text | Google Scholar

Abi Saad, C., Masiello, M., Habib, W., Gerges, E., Sanzani, S. M., Logrieco, A. F., et al. (2022). Diversity of Fusarium species isolated from symptomatic plants belonging to a wide range of agri-food and ornamental crops in Lebanon. J. Fungi 8, 897. doi:10.3390/jof8090897

PubMed Abstract | CrossRef Full Text | Google Scholar

Ahmad, F., Jabeen, K., Iqbal, S., Umar, A., Ameen, F., Gancarz, M., et al. (2023). Influence of silicon nano-particles on Avena sativa L. to alleviate the biotic stress of Rhizoctonia solani. Sci. Rep. 13, 15191. doi:10.1038/s41598-023-41699-w

PubMed Abstract | CrossRef Full Text | Google Scholar

Albalawi, M. A., Abdelaziz, A. M., Attia, M. S., Saied, E., Elganzory, H. H., and Hashem, A. H. (2022). Mycosynthesis of silica nanoparticles using Aspergillus Niger: control of Alternaria solani causing early blight disease, induction of innate immunity and reducing of oxidative stress in eggplant. Antioxidants 11, 2323. doi:10.3390/antiox11122323

PubMed Abstract | CrossRef Full Text | Google Scholar

Al-Hashimi, A., Aina, O., Daniel, A. I., Du Plessis, M., Keyster, M., and Klein, A. (2025). Critical review on characterization, management, and challenges of fusarium head blight disease in wheat. Physiological Mol. Plant Pathology 136, 102557. doi:10.1016/j.pmpp.2024.102557

CrossRef Full Text | Google Scholar

Antsotegi-Uskola, M., Markina-Iñarrairaegui, A., and Ugalde, U. (2017). Copper resistance in Aspergillus nidulans relies on the PI-type ATPase CrpA, regulated by the transcription factor AceA. Front. Microbiol. 8, 912. doi:10.3389/fmicb.2017.00912

PubMed Abstract | CrossRef Full Text | Google Scholar

Ashu, E. E., and Xu, J. (2015). The roles of sexual and asexual reproduction in the origin and dissemination of strains causing fungal infectious disease outbreaks. Infect. Genet. Evol. 36, 199–209. doi:10.1016/j.meegid.2015.09.019

PubMed Abstract | CrossRef Full Text | Google Scholar

Baazaoui, N., Bellili, K., Messaoud, M., Elleuch, L., Elleuch, R., Labidi, S., et al. (2023). Bio-nano-remediation of olive oil mill wastewater using silicon dioxide nanoparticles for its potential use as biofertilizer for young olive plants. Silicon 15, 7395–7411. doi:10.1007/s12633-023-02585-2

CrossRef Full Text | Google Scholar

Baazaoui, N., Sghaier-Hammami, B., Hammami, S. B. M., Khefacha, R., Chaari, S., Elleuch, L., et al. (2021). A handbook guide to better use of nanoparticles in plants. Commun. Soil Sci. Plant Analysis 52, 287–321. doi:10.1080/00103624.2020.1836198

CrossRef Full Text | Google Scholar

Bakri, M. M., El-Naggar, M. A., Helmy, E. A., Ashoor, M. S., and Abdel Ghany, T. M. (2020). Efficacy of juniperus procera constituents with silver nanoparticles against Aspergillus fumigatus and Fusarium chlamydosporum. BioNanoSci. 10, 62–72. doi:10.1007/s12668-019-00716-x

CrossRef Full Text | Google Scholar

Balakumaran, M. D., Ramachandran, R., and Kalaichelvan, P. T. (2015). Exploitation of endophytic fungus, Guignardia mangiferae for extracellular synthesis of silver nanoparticles and their in vitro biological activities. Microbiol. Res. 178, 9–17. doi:10.1016/j.micres.2015.05.009

PubMed Abstract | CrossRef Full Text | Google Scholar

Bathoova, M., Švubová, R., Bokor, B., Neděla, V., Tihlaříková, E., and Martinka, M. (2021). Silicon triggers sorghum root enzyme activities and inhibits the root cell colonization by Alternaria alternata. Planta 253, 29. doi:10.1007/s00425-020-03560-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Bhagat, R., Ingle, A. P., and Chen, H. (2023). “5 - nanosensors and nanobiosensors for sustainable agriculture,” in Nanotechnology in agriculture and agroecosystems. Editor A. P. Ingle (Elsevier), 93–112. doi:10.1016/B978-0-323-99446-0.00014-3

CrossRef Full Text | Google Scholar

Boddy, L., and Hiscox, J. (2016). Fungal ecology: principles and mechanisms of colonization and competition by saprotrophic fungi. Microbiol. Spectr. 4, doi:10.1128/microbiolspec.funk-0019-2016

PubMed Abstract | CrossRef Full Text | Google Scholar

Bragard, C., Di Serio, F., Gonthier, P., Jaques Miret, J. A., and Justesen, A. F. (2021). Pest categorisation of Fusarium brachygibbosum. EFSA J. 19, e06887. doi:10.2903/j.efsa.2021.6887

PubMed Abstract | CrossRef Full Text | Google Scholar

Camenzind, T., Weimershaus, P., Lehmann, A., Aguilar-Trigueros, C., and Rillig, M. C. (2022). Soil fungi invest into asexual sporulation under resource scarcity, but trait spaces of individual isolates are unique. Environ. Microbiol. 24, 2962–2978. doi:10.1111/1462-2920.16012

PubMed Abstract | CrossRef Full Text | Google Scholar

Chen, S., Guo, X., Zhang, B., Nie, D., Rao, W., Zhang, D., et al. (2023). Mesoporous silica nanoparticles induce intracellular peroxidation damage of Phytophthora infestans: a new type of green fungicide for late blight control. Environ. Sci. Technol. 57, 3980–3989. doi:10.1021/acs.est.2c07182

PubMed Abstract | CrossRef Full Text | Google Scholar

Chliyeh, M., Msairi, S., Touhami, A. O., Benkirane, R., and Douira, A. (2017). Detection of Fusarium solani as a pathogen causing root rot and wilt diseases of young olive trees in Morocco. Annu. Res. and Rev. Biol. 13, 1–7. doi:10.9734/ARRB/2017/33744

CrossRef Full Text | Google Scholar

Coskun, D., Britto, D. T., Huynh, W. Q., and Kronzucker, H. J. (2016). The role of silicon in higher plants under salinity and drought stress. Front. Plant Sci. 7, 1072. doi:10.3389/fpls.2016.01072

PubMed Abstract | CrossRef Full Text | Google Scholar

Derbalah, A., Shenashen, M., Hamza, A., Mohamed, A., and El Safty, S. (2018). Antifungal activity of fabricated mesoporous silica nanoparticles against early blight of tomato. Egypt. J. Basic Appl. Sci. 5, 145–150. doi:10.1016/j.ejbas.2018.05.002

CrossRef Full Text | Google Scholar

Ding, Z., Yang, L., Wang, G., Guo, L., Liu, L., Wang, J., et al. (2018). Fusaric acid is a virulence factor of Fusarium oxysporum f. sp. cubense on banana plantlets. Trop. plant Pathol. 43, 297–305. doi:10.1007/s40858-018-0230-4

CrossRef Full Text | Google Scholar

Duhan, J. S., Kumar, R., Kumar, N., Kaur, P., Nehra, K., and Duhan, S. (2017). Nanotechnology: the new perspective in precision agriculture. Biotechnol. Rep. 15, 11–23. doi:10.1016/j.btre.2017.03.002

PubMed Abstract | CrossRef Full Text | Google Scholar

El-Abeid, S. E., Ahmed, Y., Daròs, J.-A., and Mohamed, M. A. (2020). Reduced graphene oxide nanosheet-decorated copper oxide nanoparticles: a potent antifungal nanocomposite against Fusarium root rot and wilt diseases of tomato and pepper plants. Nanomaterials 10, 1001. doi:10.3390/nano10051001

PubMed Abstract | CrossRef Full Text | Google Scholar

ElSharawy, A. A., Ibrahim, M. S. S., and Mossa, M. I. (2023). Effect of green biosynthesized silver nanoparticles using Cleome amblyocarpa on controlling chickpea wilt. Egypt. J. Phytopathology 51, 1–16. doi:10.21608/ejp.2023.186891.1081

CrossRef Full Text | Google Scholar

Elsheery, N. I., Helaly, M. N., El-Hoseiny, H. M., and Alam-Eldein, S. M. (2020). Zinc oxide and silicone nanoparticles to improve the resistance mechanism and annual productivity of salt-stressed mango trees. Agronomy 10, 558. doi:10.3390/agronomy10040558

CrossRef Full Text | Google Scholar

Ferrigo, D., Scarpino, V., Vanara, F., Causin, R., Raiola, A., and Blandino, M. (2021). Influence of H2O2-induced oxidative stress on in vitro growth and moniliformin and fumonisins accumulation by Fusarium proliferatum and Fusarium subglutinans. Toxins 13, 653. doi:10.3390/toxins13090653

PubMed Abstract | CrossRef Full Text | Google Scholar

Gaber, S. E., Hashem, A. H., El-Sayyad, G. S., and Attia, M. S. (2024). Antifungal activity of myco-synthesized bimetallic ZnO-CuO nanoparticles against fungal plant pathogen Fusarium oxysporum. Biomass Conv. bioref. 14, 25395–25409. doi:10.1007/s13399-023-04550-w

CrossRef Full Text | Google Scholar

Gharbi, Y., Bouazizi, E., Cheffi, M., Ben Amar, F., and Triki, M. A. (2020). Investigation of soil-borne fungi, causal agents of olive trees wilt and dieback in Tunisia. Archives Phytopathology Plant Prot. 53, 828–843. doi:10.1080/03235408.2020.1800559

CrossRef Full Text | Google Scholar

Goswami, P., Sharma, M., Srivastava, N., and Mathur, J. (2022). Assessment of the fungicidal efficacy of biogenic SiO2 NPs in Eruca sativa against fusarium wilt. J. Nat. Pesticide Res. 2, 100011. doi:10.1016/j.napere.2022.100011

CrossRef Full Text | Google Scholar

Grün, A.-L., Straskraba, S., Schulz, S., Schloter, M., and Emmerling, C. (2018). Long-term effects of environmentally relevant concentrations of silver nanoparticles on microbial biomass, enzyme activity, and functional genes involved in the nitrogen cycle of loamy soil. J. Environ. Sci. 69, 12–22. doi:10.1016/j.jes.2018.04.013

PubMed Abstract | CrossRef Full Text | Google Scholar

Hassan, I. F., Ajaj, R., Gaballah, M. S., Ogbaga, C. C., Kalaji, H. M., Hatterman-Valenti, H. M., et al. (2022). Foliar application of nano-silicon improves the physiological and biochemical characteristics of ‘kalamata’ olive subjected to deficit irrigation in a semi-arid climate. Plants 11, 1561. doi:10.3390/plants11121561

PubMed Abstract | CrossRef Full Text | Google Scholar

Heller, J., and Tudzynski, P. (2011). Reactive oxygen species in phytopathogenic fungi: signaling, development, and disease. Annu. Rev. Phytopathology 49, 369–390. doi:10.1146/annurev-phyto-072910-095355

PubMed Abstract | CrossRef Full Text | Google Scholar

Hernández, M. E. S., Dávila, A. R., de Algaba, A. P., López, M. A. B., and Casas, A. T. (1998). Occurrence and etiology of death of young olive trees in southern Spain. Eur. J. Plant Pathology 104, 347–357. doi:10.1023/A:1008624929989

CrossRef Full Text | Google Scholar

Javed, Z., Dashora, K., Mishra, M., D Fasake, V., and Srivastva, A. (2019). Effect of accumulation of nanoparticles in soil health-a concern on future. Front. Nanosci. Nanotechnol. 5, 1–9. doi:10.15761/FNN.1000181

CrossRef Full Text | Google Scholar

Koleva, I. I., Beek, T. A. V., Linssen, J. P. H., Groot, A. D., and Evstatieva, L. N. (2002). Screening of plant extracts for antioxidant activity: a comparative study on three testing methods. Phytochem. Anal. 13, 8–17. doi:10.1002/pca.611

PubMed Abstract | CrossRef Full Text | Google Scholar

Kumari, A., Rana, V., Yadav, S. K., and Kumar, V. (2023). Nanotechnology as a powerful tool in plant sciences: recent developments, challenges and perspectives. Plant Nano Biol. 5, 100046. doi:10.1016/j.plana.2023.100046

CrossRef Full Text | Google Scholar

Leslie, J. F., and Summerell, B. A. (2006). The Fusarium Laboratory Manual. Hoboken: Blackwell Publishing. 1–2. doi:10.1002/9780470278376

CrossRef Full Text | Google Scholar

Liao, S., Zhang, Y., Pan, X., Zhu, F., Jiang, C., Liu, Q., et al. (2019). Antibacterial activity and mechanism of silver nanoparticles against multidrug-resistant Pseudomonas aeruginosa. Int. J. Nanomedicine 14, 1469–1487. doi:10.2147/IJN.S191340

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, G., Kennedy, R., Greenshields, D. L., Peng, G., Forseille, L., Selvaraj, G., et al. (2007). Detached and attached arabidopsis leaf assays reveal distinctive defense responses against hemibiotrophic colletotrichum spp. MPMI 20, 1308–1319. doi:10.1094/MPMI-20-10-1308

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, S., Fu, L., Wang, S., Chen, J., Jiang, J., Che, Z., et al. (2019a). Carbendazim resistance of Fusarium graminearum from henan wheat. Plant Dis. 103, 2536–2540. doi:10.1094/PDIS-02-19-0391-RE

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, S., Li, J., Zhang, Y., Liu, N., Viljoen, A., Mostert, D., et al. (2019b). Fusaric acid instigates the invasion of banana by Fusarium oxysporum f. sp. cubense TR4. New Phytol. 225, 913–929. doi:10.1111/nph.16193

PubMed Abstract | CrossRef Full Text | Google Scholar

López-Díaz, C., Rahjoo, V., Sulyok, M., Ghionna, V., Martín-Vicente, A., Capilla, J., et al. (2018). Fusaric acid contributes to virulence of Fusarium oxysporum on plant and mammalian hosts. Mol. Plant Pathol. 19, 440–453. doi:10.1111/mpp.12536

PubMed Abstract | CrossRef Full Text | Google Scholar

Lucas, J. A., Hawkins, N. J., and Fraaije, B. A. (2015). “Chapter two - the evolution of fungicide resistance,” in Advances in applied microbiology. Editors S. Sariaslani, and G. M. Gadd (Academic Press), 29–92. doi:10.1016/bs.aambs.2014.09.001

CrossRef Full Text | Google Scholar

Markakis, E. A., Roditakis, E. N., Kalantzakis, G. S., Chatzaki, A., Soultatos, S. K., Stavrakaki, M., et al. (2021). Characterization of fungi associated with olive fruit rot and olive oil degradation in crete, southern Greece. Plant Dis. 105, 3623–3635. doi:10.1094/PDIS-10-20-2227-RE

PubMed Abstract | CrossRef Full Text | Google Scholar

Martínez, J. R., Palomares-Sánchez, S., Ortega-Zarzosa, G., Ruiz, F., and Chumakov, Y. (2006). Rietveld refinement of amorphous SiO2 prepared via sol–gel method. Mater. Lett. 60, 3526–3529. doi:10.1016/j.matlet.2006.03.044

CrossRef Full Text | Google Scholar

Munkvold, G. P., Proctor, R. H., and Moretti, A. (2021). Mycotoxin production in Fusarium according to contemporary species concepts. Annu. Rev. Phytopathology 59, 373–402. doi:10.1146/annurev-phyto-020620-102825

PubMed Abstract | CrossRef Full Text | Google Scholar

Nicoletti, R., Di Vaio, C., and Cirillo, C. (2020). Endophytic fungi of olive tree. Microorganisms 8, 1321. doi:10.3390/microorganisms8091321

PubMed Abstract | CrossRef Full Text | Google Scholar

O’Donnell, K., Kistler, H. C., Cigelnik, E., and Ploetz, R. C. (1998). Multiple evolutionary origins of the fungus causing Panama disease of banana: concordant evidence from nuclear and mitochondrial gene genealogies. Proc. Natl. Acad. Sci. U. S. A. 95, 2044–2049. doi:10.1073/pnas.95.5.2044

PubMed Abstract | CrossRef Full Text | Google Scholar

Parveen, A., and Siddiqui, Z. A. (2022). Impact of silicon dioxide nanoparticles on growth, photosynthetic pigments, proline, activities of defense enzymes and some bacterial and fungal pathogens of tomato. Vegetos 35, 83–93. doi:10.1007/s42535-021-00280-4

CrossRef Full Text | Google Scholar

Pereira, G. A., Arruda, H. S., and Pastore, G. M. (2018). Modification and validation of Folin-Ciocalteu assay for faster and safer analysis of total phenolic content in food samples. Braz. J. Food Res. 125, 125. doi:10.3895/rebrapa.v9n1.6062

CrossRef Full Text | Google Scholar

Qin, G. Z., and Tian, S. P. (2005). Enhancement of biocontrol activity of cryptococcus laurentii by silicon and the possible mechanisms involved. Phytopathology 95, 69–75. doi:10.1094/PHYTO-95-0069

PubMed Abstract | CrossRef Full Text | Google Scholar

Rajput, V. D., Minkina, T., Feizi, M., Kumari, A., Khan, M., Mandzhieva, S., et al. (2021). Effects of silicon and silicon-based nanoparticles on rhizosphere microbiome, plant stress and growth. Biology 10, 791. doi:10.3390/biology10080791

PubMed Abstract | CrossRef Full Text | Google Scholar

Reverberi, M., Ricelli, A., Zjalic, S., Fabbri, A. A., and Fanelli, C. (2010). Natural functions of mycotoxins and control of their biosynthesis in fungi. Appl. Microbiol. Biotechnol. 87, 899–911. doi:10.1007/s00253-010-2657-5

PubMed Abstract | CrossRef Full Text | Google Scholar

Ribeiro, R., Rentes, B. R., Honorato, L. A., and Kuhnen, S. (2024). Effect of nanoemulsion loaded with macela (Achyrocline satureioides) on the ultrastructure of Staphylococcus aureus and the modulating activity of antibiotics. Front. Nanotechnol. 6, 1466988. doi:10.3389/fnano.2024.1466988

CrossRef Full Text | Google Scholar

Rodrigues, A. S., Batista, J. G. S., Rodrigues, M. Á. V., Thipe, V. C., Minarini, L. A. R., Lopes, P. S., et al. (2024). Advances in silver nanoparticles: a comprehensive review on their potential as antimicrobial agents and their mechanisms of action elucidated by proteomics. Front. Microbiol. 15, 1440065. doi:10.3389/fmicb.2024.1440065

PubMed Abstract | CrossRef Full Text | Google Scholar

Sakr, N. (2016). The role of silicon (Si) in increasing plant resistance against fungal diseases. Hellenic Plant Prot. J. 9, 1–15. doi:10.1515/hppj-2016-0001

CrossRef Full Text | Google Scholar

Sakr, N. (2021a). pH variation inhibits the mycelial growth of fusarium spp. and cochliobolus sativus. Pak. J. Phytopathology 33, 265–273. doi:10.33866/phytopathol.033.02.0637

CrossRef Full Text | Google Scholar

Sakr, N. (2021b). Soluble silicon controls Fusarium head blight in bread and durum wheat plants. Gesunde Pflanz. 73, 479–493. doi:10.1007/s10343-021-00568-0

CrossRef Full Text | Google Scholar

Sakr, N., and Shoaib, A. (2021). Pathogenic and molecular variation of Fusarium species causing head blight on barley landraces. Acta Phytopathologica Entomologica Hung. 56, 5–23. doi:10.1556/038.2021.00006

CrossRef Full Text | Google Scholar

Santosh, P. S., and Yojana, S. M. (2022). Sustainable management of pesticides. IJZI 08, 827–836. doi:10.33745/ijzi.2022.v08i02.100

CrossRef Full Text | Google Scholar

Scott, B., and Eaton, C. J. (2008). Role of reactive oxygen species in fungal cellular differentiations. Curr. Opin. Microbiol. 11, 488–493. doi:10.1016/j.mib.2008.10.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Sghaier-Hammami, B., Mansour, R. B., Messaoud, M., Baazaoui, N., Ettlili, S., Elleuch, R., et al. (2024). Silicon dioxide (SiO2) nanoparticles as a Alternaria alternata fungi mitigator on biomass, photosynthetic machinery, nutriome and antioxidant capacity of barley (hordeum vulgare L.). Silicon 16, 4929–4944. doi:10.1007/s12633-024-03031-7

CrossRef Full Text | Google Scholar

Shahzad, S., Ali, S., Ahmad, R., Ercisli, S., and Anjum, M. A. (2022). Foliar application of silicon enhances growth, flower yield, quality and postharvest Life of tuberose (polianthes tuberosa L.) under saline conditions by improving antioxidant defense mechanism. Silicon 14, 1511–1518. doi:10.1007/s12633-021-00974-z

CrossRef Full Text | Google Scholar

Siamak, S. B., and Zheng, S. (2018). Banana Fusarium wilt (Fusarium oxysporum f. sp. cubense) control and resistance, in the context of developing wilt-resistant bananas within sustainable production systems. Hortic. Plant J. 4, 208–218. doi:10.1016/j.hpj.2018.08.001

CrossRef Full Text | Google Scholar

Siddiqui, M. H., Al-Whaibi, M. H., Faisal, M., and Al Sahli, A. A. (2014). Nano-silicon dioxide mitigates the adverse effects of salt stress on Cucurbita pepo L. Environ. Toxicol. Chem. 33, 2429–2437. doi:10.1002/etc.2697

PubMed Abstract | CrossRef Full Text | Google Scholar

Singh, V. K., Singh, H. B., and Upadhyay, R. S. (2017). Role of fusaric acid in the development of ‘Fusarium wilt’ symptoms in tomato: physiological, biochemical and proteomic perspectives. Plant Physiology Biochem. 118, 320–332. doi:10.1016/j.plaphy.2017.06.028

PubMed Abstract | CrossRef Full Text | Google Scholar

Subba, B., Rai, G. B., Bhandary, R., Parajuli, P., Thapa, N., Kandel, D. R., et al. (2024). Antifungal activity of zinc oxide nanoparticles (ZnO NPs) on Fusarium equiseti phytopathogen isolated from tomato plant in Nepal. Heliyon 10, e40198. doi:10.1016/j.heliyon.2024.e40198

PubMed Abstract | CrossRef Full Text | Google Scholar

Suriyaprabha, R., Karunakaran, G., Kavitha, K., Yuvakkumar, R., Rajendran, V., and Kannan, N. (2014). Application of silica nanoparticles in maize to enhance fungal resistance. IET Nanobiotechnology 8, 133–137. doi:10.1049/iet-nbt.2013.0004

PubMed Abstract | CrossRef Full Text | Google Scholar

Terzioğlu, E., Alkım, C., Arslan, M., Balaban, B. G., Holyavkin, C., Kısakesen, H. İ., et al. (2020). Genomic, transcriptomic and physiological analyses of silver-resistant Saccharomyces cerevisiae obtained by evolutionary engineering. Yeast 37, 413–426. doi:10.1002/yea.3514

PubMed Abstract | CrossRef Full Text | Google Scholar

Todorova, D., Yavorov, N., and Vrabič-Brodnjak, U. (2024). Impact of silver nanoparticle treatment and chitosan on packaging paper’s barrier effectiveness. Polymers 16, 2127. doi:10.3390/polym16152127

PubMed Abstract | CrossRef Full Text | Google Scholar

Trabelsi, R., Sellami, H., Gharbi, Y., Krid, S., Cheffi, M., Kammoun, S., et al. (2017). Morphological and molecular characterization of Fusarium spp. associated with olive trees dieback in Tunisia. 3 Biotech. 7, 28. doi:10.1007/s13205-016-0587-3

PubMed Abstract | CrossRef Full Text | Google Scholar

Velikova, V., Yordanov, I., and Edreva, A. (2000). Oxidative stress and some antioxidant systems in acid rain-treated bean plants: protective role of exogenous polyamines. Plant Sci. 151, 59–66. doi:10.1016/S0168-9452(99)00197-1

CrossRef Full Text | Google Scholar

Verma, R., Kushwaha, K. P. S., Kumar, S., Rawat, S., Pandey, R., Pandey, D., et al. (2024). Emerging threat to Indian agriculture: Fusarium incarnatum-equiseti species complex as a novel pathogen imperiling bajra, cowpea, finger millet, green gram, moth bean, and soybean crops. Crop Prot. 182, 106741. doi:10.1016/j.cropro.2024.106741

CrossRef Full Text | Google Scholar

Wei, X., Jiang, W., Yu, J., Ding, L., Hu, J., and Jiang, G. (2015). Effects of SiO2 nanoparticles on phospholipid membrane integrity and fluidity. J. Hazard. Mater. 287, 217–224. doi:10.1016/j.jhazmat.2015.01.063

PubMed Abstract | CrossRef Full Text | Google Scholar

Wei, X., Wang, W.-G., and Matsuda, Y. (2022). Branching and converging pathways in fungal natural product biosynthesis. Fungal Biol. Biotechnol. 9, 6. doi:10.1186/s40694-022-00135-w

PubMed Abstract | CrossRef Full Text | Google Scholar

Wyatt, T. T., Wösten, H. A. B., and Dijksterhuis, J. (2013). “Chapter two - fungal spores for dispersion in space and time,” in Advances in applied microbiology. Editors S. Sariaslani, and G. M. Gadd (Academic Press), 43–91. doi:10.1016/B978-0-12-407672-3.00002-2

CrossRef Full Text | Google Scholar

Xu, C., Li, M., Zhou, Z., Li, J., Chen, D., Duan, Y., et al. (2019). Impact of five succinate dehydrogenase inhibitors on DON biosynthesis of Fusarium asiaticum, causing Fusarium head blight in wheat. Toxins 11, 272. doi:10.3390/toxins11050272

PubMed Abstract | CrossRef Full Text | Google Scholar

Yan, X., He, B., Liu, L., Qu, G., Shi, J., Hu, L., et al. (2018). Antibacterial mechanism of silver nanoparticles in Pseudomonas aeruginosa: proteomics approach. Metallomics 10, 557–564. doi:10.1039/c7mt00328e

PubMed Abstract | CrossRef Full Text | Google Scholar

Yin, Y., Liu, X., Li, B., and Ma, Z. (2009). Characterization of sterol demethylation inhibitor-resistant isolates of Fusarium asiaticum and F. Graminearum collected from wheat in China. Phytopathology® 99, 487–497. doi:10.1094/PHYTO-99-5-0487

PubMed Abstract | CrossRef Full Text | Google Scholar

Yobo, K. S., Mngadi, Z. N. C., and Laing, M. D. (2019). Efficacy of two potassium silicate formulations and two trichoderma strains on Fusarium head blight of wheat. Proc. Natl. Acad. Sci. India, Sect. B Biol. Sci. 89, 185–190. doi:10.1007/s40011-017-0935-z

CrossRef Full Text | Google Scholar

Yuan, S., and Zhou, M. (2005). A major gene for resistance to carbendazim, in field isolates of Gibberella zeae. Can. J. Plant Pathology 27, 58–63. doi:10.1080/07060660509507194

CrossRef Full Text | Google Scholar

Zhang, L., Jiang, Y., Ding, Y., Povey, M., and York, D. (2007). Investigation into the antibacterial behaviour of suspensions of ZnO nanoparticles (ZnO nanofluids). J. Nanopart Res. 9, 479–489. doi:10.1007/s11051-006-9150-1

CrossRef Full Text | Google Scholar

Zhang, Z., Chen, Y., Li, B., Chen, T., and Tian, S. (2020). Reactive oxygen species: a generalist in regulating development and pathogenicity of phytopathogenic fungi. Comput. Struct. Biotechnol. J. 18, 3344–3349. doi:10.1016/j.csbj.2020.10.024

PubMed Abstract | CrossRef Full Text | Google Scholar

Ziaee, M., and Ganji, Z. (2016). Insecticidal efficacy of silica nanoparticles against Rhyzopertha Dominica F. and Tribolium confusum Jacquelin du Val. J. Plant Prot. Res. 56, 250–256. doi:10.1515/jppr-2016-0037

CrossRef Full Text | Google Scholar