Research Article
Bioactive Metabolites and Anti-Phytopathogenic Potential of Erica verticillata: A Targeted Leaf Extract Study
Abdallah Khalil1, Karrar A. Hamzah2, Shimaa Bashir3, Said Behiry4 and Ahmed Abdelkhalek3,5*
1Plant Protection Department, Faculty of Agriculture, Omar Al-Mukhtar University, Elbyda, Libya; 2Department of Horticulture and Landscape Planning, College of Agriculture, Al-Qasim Green University, Al-Qasim District 964, Babylon, 51013, Iraq; 3Plant Protection and Biomolecular Diagnosis Department, Arid Lands Cultivation Research Institute, City of Scientific Research and Technological Applications, Alexandria 21934, Egypt; 4Agricultural Botany Department, Faculty of Agriculture (Saba Basha), Alexandria University, Alexandria 21531, Egypt; 5Plant Protection Department, The National Institute of Horticultural Research, Konstytucji 3 Maja 1/3, Skierniewice, 96-100, Poland.
Abstract | Plant pathogenic microorganisms pose significant threats to agriculture, necessitating the exploration of natural alternatives to conventional pesticides. This study aimed to investigate the antimicrobial potential of methanolic extracts from Erica verticillata against various plant pathogenic bacteria and fungi. The antibacterial assays revealed substantial activity of the extract, indicating that Pectobacterium atrosepticum and Streptomyces scabiei exhibited sensitivity at a concentration of 300 µg/ mL. At the same time, Ralstonia solanacearum and Pectobacterium carotovorum displayed sensitivity at a concentration of 500 µg/ mL. Antifungal evaluations revealed potent inhibition against Fusarium oxysporum, Botrytis cinerea, and Rhizoctonia solani. The extract strongly inhibited B. cinerea (78.2 % at 5000 µg/ mL) outperforming copper hydroxide (58.7 %). Significant in vitro inhibition was also observed against R. solani and F. oxysporum, often exceeding the positive control. Phytochemical analysis using HPLC revealed that the predominant phenolic components were gallic acid, caffeic acid, and syringic acid, with concentrations of 2620.3, 606.5, and 580.9 µg/ g, respectively. The principal flavonoid was quercetin at a concentration of 169.4 µg/ g. Hexadecanoic acid and phorbol were the most prevalent secondary metabolites identified in the GC-MS analysis, comprising 26.4 % and 16.4 %, respectively. These findings highlight the rich phytochemical profile of E. verticillata and underscore its potential as a source of natural antimicrobial agents for managing plant diseases, including those caused by antibiotic-resistant pathogens.
Received | May 25, 2025; Revised | June 24 2025; Accepted | July 10, 2025; Published | July 17, 2025
*Correspondence | Ahmed Abdelkhalek, Plant Protection and Biomolecular Diagnosis Department, Arid Lands Cultivation Research Institute, City of Scientific Research and Technological Applications, Alexandria 21934, Egypt; Email: [email protected]
Citation | Khalil, A., K.A. Hamzah, S. Bashir, S. Behiry and A. Abdelkhalek. 2025. Bioactive metabolites and anti-phytopathogenic potential of Erica verticillata: A targeted leaf extract study. Novel Research in Microbiology Journal, 9(4): 273-288.
DOI | https://dx.doi.org/10.17582/journal.NRMJ/9.4.273.288
Keywords | Erica verticillata, Antimicrobial, Sustainable agriculture, HPLC, GC-MS
Copyright: 2025 by the authors. Licensee ResearchersLinks Ltd, England, UK.
This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/).
Introduction
Plant pathogenic microorganisms, encompassing bacterial and fungal diseases, represent a substantial risk to agricultural production and global food security (Aamer et al., 2024). Bacterial and fungal pathogens, including Pectobacterium carotovorum, Pectobacterium atrosepticum, Ralstonia solanacearum, Streptomyces scabiei, Fusarium oxysporum, Botrytis cinerea, and Rhizoctonia solani, are implicated in numerous plant diseases that impact crop quality and yield (Erbs and Newman, 2024; Ali et al., 2025). The increasing resistance of such pathogens to conventional antimicrobial agents necessitates the development of alternative control strategies. The emergence of antimicrobial resistance has become a global health concern, prompting researchers to explore alternative sources of antimicrobial agents (Cotugno et al., 2025). Plant-derived compounds have garnered significant attention due to their diverse bioactive constituents and potential therapeutic applications (Pereira et al., 2025). Medicinal plants have been utilized for centuries in traditional medicine systems worldwide, serving as a valuable reservoir of novel antimicrobial compounds with diverse mechanisms of action (Bhattacharjee and Sharma, 2025).
Recent studies demonstrated the efficacy of plant-derived antimicrobial agents against various plant pathogens (Košćak et al., 2025; Pérez-Flores et al., 2025). The significant antibacterial activity of the phenolic-rich plant extracts against the phytopathogenic bacteria, such as R. solanacearum has been reported, highlighting their potential as eco-friendly alternatives to the synthetic bactericides (Yihune and Yemata, 2019). Similarly, the antifungal properties of the plant extracts against phytopathogenic fungi emphasize their applications in sustainable agriculture and forestry (Nguyen et al., 2024). The growing body of evidence supporting the antimicrobial efficacy of plant extracts underscores their potential as natural alternatives to conventional antimicrobial agents in agricultural and pharmaceutical applications (Chaachouay and Zidane, 2024). Despite the promising antimicrobial properties of various plant extracts, comprehensive studies investigating the relationship between their phytochemical composition and biological activities remain limited (Kumar et al., 2023; Murugan et al., 2025). Understanding this relationship is essential for developing standardized plant-based antimicrobial formulations with consistent efficacy and safety profiles. Moreover, identifying specific bioactive compounds responsible for the observed antimicrobial activities could facilitate the development of novel antimicrobial agents with enhanced potency and selectivity.
Plant secondary metabolites, particularly phenolic compounds, have been extensively studied for their antimicrobial potencies (Benalach et al., 2025). These compounds exhibit diverse mechanisms of action against microbial pathogens, including disruption of cell membranes, leading to increased permeability and subsequent cell death, inhibition of microbial enzymes, and suppression of virulence factors (Arzani et al., 2025; De Rossi et al., 2025). A recent study highlighted the significant correlation between the phenolic content of plant extracts and their antimicrobial efficacy against various pathogenic microorganisms (Pérez-Flores et al., 2025). Identifying and characterizing the bioactive compounds in the plant extracts are crucial for understanding their antimicrobial mechanisms and potential applications. Advanced analytical techniques such as high-performance liquid chromatography (HPLC) and gas chromatography-mass spectrometry (GC-MS) have revolutionized the field of natural product studies, enabling the precise identification and quantification of bioactive constituents in the complex plant matrices (Debnath et al., 2025). HPLC analysis is particularly effective for separating and quantifying phenolic compounds, while GC-MS provides valuable information about the volatile and semi-volatile components of plant extracts.
The genus Erica (family Ericaceae) comprises approximately 860 species distributed across Europe, the Mediterranean region, and South Africa (Dias et al., 2015). Various Erica species have demonstrated significant biological activities, including antimicrobial, antioxidant, and anti-inflammatory properties (Guendouze-Bouchefa et al., 2015). Erica verticillata, a species native to the Cape Floristic region of South Africa, has been traditionally used for treating various ailments, including respiratory infections and urinary tract disorders (Ojewole, 2008). Despite its ethnomedicinal significance, comprehensive scientific investigations into the antimicrobial potential and phytochemical composition of E. verticillata remain limited. The objectives of this study were to evaluate the antibacterial and antifungal activities of E. verticillata methanolic extract against selected plant pathogenic microorganisms, and correlate these activities with the phytochemical composition of the extract. This study employs a multifaceted approach, combining in vitro antimicrobial assays with advanced analytical techniques (i.e., HPLC and GC-MS) to provide comprehensive insights into the antimicrobial potential and chemical profile of E. verticillata extract. The findings of this study could contribute to the development of natural antimicrobial agents for sustainable plant disease management and potentially address the growing challenge of antimicrobial resistance in agricultural settings.
Materials and Methods
Source of bacterial and fungal strains
This study utilized various bacterial and fungal strains to evaluate the antimicrobial activity of a methanolic extract obtained from leaves of E. verticillata. The bacterial strains comprised P. carotovorum (GenBank accession number OQ878656), P. atrosepticum (GenBank accession number MG706146), R. solanacearum (GenBank accession number OQ878653), and S. scabiei (GenBank accession number OR437480). These strains were previously isolated from potato plants exhibiting characteristic symptoms of bacterial infections. The fungal strains, including F. oxysporum (GenBank accession number OQ820156) sourced from infected potato tubers, B. cinerea (GenBank accession number MN398400) obtained from infected strawberry fruits, and R. solani (GenBank accession number OQ880457) isolated from infected bean plants.
Preparation of a plant extract
Healthy plant samples of E. verticillata were collected from Jebel Akhdar in Libya. Plants devoid of visible morphological diseases were selected. The plant materials were transported to the laboratory, where they were washed with tap water to remove debris and contaminants. The washed plant materials were dried at room temperature (25±2 °C) in the shade for 10 d until fully desiccated, a method recommended for preserving biologically active phenolic compounds (Dai and Mumper, 2010). Dried samples were ground using a mill to obtain particles of 0.5 mm, and 50 g of the powder were extracted with 500 mL of 80 % methanol in a rotary shaker at 100 rpm for 12 h at room temperature. This method is deemed adequate for extraction of phenolic compounds from plant materials, as evidenced by prior studies conducted by Azwanida (2015), Altemimi et al. (2017). The supernatant; free of plant debris, was obtained via filtration using Whatman No. 1 filter paper, and finally the methanol was evaporated at 25-30 °C in a rotary evaporator Re-52A (Wincom, Jiangsu, China).
In-vitro antibacterial assay
The agar disc diffusion method was used to evaluate the antibacterial potential of the methanolic plant extract (Bashir et al., 2024). Briefly, a single colony of the purified bacterial strain was inoculated into 100 mL of the nutrient broth (NB) medium and incubated overnight at 27 °C. At the end of the incubation period, bacterial growth concentrations were adjusted to 108 CFU/ mL using NB medium. Afterward, 100 μL were transferred and spread on the surface of Glycerol Nutrient Agar (GNA) using a sterile glass spreader. Different extract concentrations were prepared in 10 % DMSO to final concentrations of 300, 500, 700, and 1000 μg/ mL in 100 mL of NB medium. The negative control group of the experiment was adjusted using 10 % DMSO in NB medium without the extract. Aliquots of 15 μL of each extract concentration were added to 5 mm diameter filter paper discs, and then left to dry at 4 °C for 24 h. The plant extract-loaded filter paper discs were placed individually on surface of the seeded plates and incubated at 27 °C for 24 h. The positive control groups were adjusted using amoxicillin as a standard antibiotic with a concentration of 25 μg/ disc. After incubation, the antibacterial activities were quantified by measuring the inhibition zone diameter (mm) using a calibrated ruler in triplicate against the control groups.
The inhibition percentages (%) were calculated based on the obtained inhibition zone diameters using the following formula:

In-vitro antifungal assay
The antifungal effects of the plant extract against some plant pathogenic fungi were evaluated using the radial growth assay on poisoned food procedures, as previously described by Khalil et al. (2024). Briefly, different extract concentrations were added to the potato dextrose agar (PDA) plates to obtain final concentrations of 1000, 2000, 3000, 4000, and 5000 µg/mL. A 5 mm disc of 5-day-old fungal culture was placed at the midpoint of each treated plate. The concentration of each extract was evaluated against the negative control (DMSO-PDA) and the positive control (Copper hydroxide) at a concentration of 1.5 g/ L. All plates were incubated for 7 d at 25 °C. After incubation, the antifungal activities were detected by determining the fungal mycelial growth inhibition percentage, which was calculated according to the following equation:
Mycelial growth inhibition (%) = [(A0 - At)/A0] × 100
Where A0 is the average diameter in gth untreated plates (control), and At is the average diameter of fungal growth in the treated plates (Bashir et al., 2024).
High-performance liquid chromatography analysis
Polyphenolic compounds were identified in the methanol extract using high-performance liquid chromatography (HPLC) (Agilent 1260 Infinity HPLC Series, Santa Clara, CA, USA), which was outfitted with a quaternary pump. The conditions for HPLC were established as follows: A Zorbax Eclipse Plus C18 column (100 mm × 4.6 mm) from Agilent Technologies (Santa Clara, CA, USA), was utilized for separation at 30 °C. The separation process was carried out with a 20 μL injection volume, employing a ternary linear elution gradient of water with 0.2 % H3PO4 (HPLC grade v/v), methanol, and acetonitrile. The detected peaks were analyzed with a VWD detector calibrated to 284 nm, utilizing a range of phenolic compound standards, including gallic acid, catechol, p-hydroxybenzoic acid, caffeine, vanillic acid, caffeic acid, syringic acid, vanillin, p-coumaric acid, ferulic acid, ellagic acid, benzoic acid, o-coumaric acid, salicylic acid, and cinnamic acid. Analysis using HPLC is regarded as a highly accurate and trustfull technique for identifying and examining the polyphenolic compounds in the extract, as supported by several prior studies (Naczk and Shahidi, 2004; Abdelkhalek et al., 2024).
Gas chromatography-mass spectrometry analysis
Gas chromatography-mass spectrometry (GC-MS) analysis is a reliable technique for identifying volatile and semi-volatile organic compounds in plant extracts. The potential bioactive ingredients in the extract were determined using an (Agilent Technologies, Santa Clara, Ca, USA),, which included an Agilent mass spectrometric detector featuring a direct capillary interface and a fused silica capillary column (HP-5MS), with specifications of 30 m × 0.32 mm × 0.25 μm film thickness. The column temperature was initially set at 50 °C with a ramp rate of 5 °C/min and then increased to 230 °C. The temperature was maintained at 290 °C for 2 min. before being increased to the final temperature of 300 °C. The sample injection process and program conditions were executed according to a previously published protocol (Youssef et al., 2021). The bioactive components of the extract were identified through a search of mass spectral libraries (NIST and Wiley) and by comparing the mass spectra and retention times with information available in the Wiley and NIST mass spectral library databases.
Statistical analysis
A one-way analysis of variance (ANOVA) was performed using the CoStat program (version 6.45) from the Cohort program to analyze the data statistically. The Tukey honest significant difference (HSD) test was used to evaluate differences and determine the statistical significance of the procedure at a significance level of p ≤ 0.05. An average value (mean ± standard deviation [SD]) together with its standard deviation was displayed for the data. The letters arranged in descending order demonstrated the link and indicated statistical significance (where; a > b > c). The comparable letters indicated no significant alteration.
Results
Antibacterial activity of Erica verticillata extract
Figure 1 illustrates the antibacterial potency of the methanolic extract of E. verticillata. The data indicated different sensitivity levels among the tested bacterial strains in response to the extract treatments. P. atrosepticum and S. scabiei exhibited the highest sensitivity, even at reduced treatment concentrations (300 µg/ mL), recording inhibition percentages of 33.33 % and 62.48 %, respectively.. The results indicate that these bacterial strains exhibited a notable vulnerability to the bioactive compounds present in the extract. R. solanacearum and P. carotovorum showed moderate sensitivity, with inhibition percentages of 60.51 % and 28.36 %, respectively, when treated with 700 µg/ mL of the extract. This evidence suggests that larger amounts of the extract were necessary to suppress the growth of these bacterial strains effectively. Whereas, R. solanacearum and S. scabiei bacterial strains demonstrated a notable sensitivity to the plant extract treatments at a concentration of 1000 µg/ mL, resulting in inhibition percentages of 63.19 % and 75.05 %, respectively. This discovery highlights the potential of E. verticillata extract as a viable antimicrobial option for combating multi-drug resistant bacteria. The statistical analysis revealed no significant differences in in the inhibition percentages across all the tested concentrations of the plant extract against the bacterial strains (p < 0.05). This finding shows that the antibacterial impact of the methanolic extract reaches an inhibitory limit at a specific concentration; however, adding more concentration does not significantly enhances its ability to inhibit the bacteria.
Antifungal activity of Erica verticillata extract
The antifungal properties of methanolic extract from E. verticillata were evaluated against three significant plant pathogenic fungi: F. oxysporum, B. cinerea, and R. solani. The findings are depicted in Figure 2. The extract had substantial inhibitory effects against all the tested fungal strains, particularly demonstrating notable action against B. cinerea. At 5000 µg/ mL, the extract displayed an impressive fungal growth inhibition rate of 78.2 % against B. cinerea, notably surpassing the positive control (copper hydroxide 1.5 g/L), which exhibited only 58.7 % inhibition. Statistical analysis revealed no significant differences (p < 0.05) in the efficacy of the extract against B. cinerea when administered at concentrations of 3000, 4000, and 5000 µg/ mL. These findings suggest that a lower concentration (3000 µg/mL) effectively achieved maximum inhibitory potential against this fungal pathogen, presenting a cost-effective alternative for potential applications. The extract demonstrated significant activity (p < 0.05) against R. solani at a concentration of 5000 µg/mL, with an inhibition percentage of 70.4 %.
The obtained results revealed a performance level of the extract comparable to that of the copper hydroxide treatment (67.1 %) and displayed a significant improvement over the untreated control. The obtained data suggest that the extract may be a natural substitute for chemical fungicides in managing R. solani infections. Similarly, notable suppression of F. oxysporum growth was recorded after applying plant extract concentrations of 2000, 3000, 4000, and 5000 µg/ mL, yielding inhibition percentages of 52.2 %, 66.9 %, 69.1 %, and 70.7 %, respectively. The values observed were significantly greater than the inhibition obtained through copper hydroxide treatment (34.2 %) at p < 0.05, highlighting the enhanced antifungal properties of the E. verticillata extract compared to the traditional fungicides.
High-performance liquid chromatography (HPLC) analysis for the detection of phenolic and flavonoid compounds
The chemical composition of E. verticillata methanolic extract was analyzed using HPLC to identify and quantify the phenolic and flavonoid compounds present. Figure 3 and Table 1 display the HPLC chromatograms and the obtained quantitative results. The HPLC analysis indicated that gallic acid
(2620.3 µg/ g), caffeic acid (606.5 µg/ g), and syringic acid (580.9 µg/ g) were the predominant phenolic compounds in the methanolic plant extract. The potent antioxidant and antimicrobial characteristics of these phenolic acids may substantially enhance the antibacterial and antifungal effects observed in the extract. Naringenin (198.4 µg/ g) and quercetin (169.4 µg/ g) were identified as the most abundant flavonoid compounds in the extract. Flavonoids exhibit a range of biological activities such as antimicrobial effects, and their inclusion in the extract likely enhances its overall bioactivity. Additionally, other phenolic and flavonoid compounds were identified at lower concentrations, including catechin (143.5 µg/ g), coumaric acid (63.8 µg/ g), kaempferol (54.9 µg/ g), and hesperetin (43.4 µg/ g). The presence of these bioactive compounds together may lead to synergistic antimicrobial effects, improving the overall effectiveness of the extract against the phytopathogenic microorganisms.
Gas chromatography-mass spectrometry (GC-MS) analysis of the extract’s secondary metabolites
Figure 4 and Table 2 summarize the results obtained from the GC-MS analysis. The collected data validated the existence of seven primary compounds, each was characterized by a distinct chemical structure and retention time. Hexadecanoic acid emerged as the predominant compound (26.4 %), observed at a retention time of 26.36 min. This fatty acid exhibits antimicrobial properties and may significantly contribute to the bioactivities observed in the extract. Phorbol was identified as the second most prevalent compound (16.4 %). Phorbol and its derivatives exhibit a range of biological activities, including antimicrobial effects, which may enhance the efficacy of the tested extract against the phytopathogens.
Table 1: Analysis of phenolic and flavonoid compounds of methanolic extract of E. verticillata using high-performance liquid chromatography (HPLC).
|
Compounds |
Area |
Concentration (µg/ g) |
|
Gallic acid |
430.61 |
2620.3 |
|
Caffeic acid |
110.45 |
606.5 |
|
Syringic acid |
123.61 |
580.9 |
|
Ferulic acid |
61.53 |
265.3 |
|
Ellagic acid |
15.97 |
202.6 |
|
Naringenin |
27.29 |
198.4 |
|
Vanillin |
60.89 |
177.8 |
|
Cinnamic acid |
139.25 |
174.1 |
|
Quercetin |
18.72 |
169.4 |
|
Catechin |
8.64 |
143.5 |
|
Daidzein |
33.41 |
136.8 |
|
29.97 |
63.8 |
|
|
Methyl gallate |
16.97 |
63.3 |
|
Kaempferol |
11.84 |
54.9 |
|
Hesperetin |
12.07 |
43.4 |
|
Chlorogenic acid |
Not detected |
Not detected |
|
Pyro catechol |
Not detected |
Not detected |
|
Rutin |
Not detected |
Not detected |
|
Apigenin |
Not detected |
Not detected |
Additional compounds were identified at moderate concentrations, comprising α-sitosterol, octadecenoic acid, tibolone, stigmasterol, and methyl phaseate. Figure 4 illustrates the relative area percentages of these compounds, offering a visual representation of their proportions in the extract. Other compounds, such as columbin and loperamide, were detected at lower concentrations. The variety of secondary metabolites in the E. verticillata extract suggests a complex phytochemical profile, contributing to its extensive antimicrobial effectiveness observed against several bacterial and fungal phytopathogens. The synergistic effect of these compounds probably enhances the extract’s efficacy as a natural antimicrobial agent.
Discussion
Plant pathogens, including bacteria and fungi, are known to cause significant qualitative and quantitative harm to the crop production (Al-Askar et al., 2025). Using plant extracts as alternative antimicrobial medicines has recently attracted considerable attention due to their efficacy, efficiency, and diminished side effects. The current study investigated the antimicrobial properties and chemical composition of the methanolic extract of E. verticillata concerning various phytopathogenic bacteria and fungi. The results indicated notable antibacterial and antifungal properties, as well as a range of bioactive compounds contributing to these observed effects. The results of the antibacterial assay suggested that the extract exhibited differing inhibitory effects against the examined phytopathogenic bacterial strains. The differential sensitivity levels observed among the bacterial strains indicate the specificity of the modes of action of the extract’s bioactive compounds. This observed specificity may be attributed to variations in the cell wall composition and membrane permeability of the tested bacteria, affecting their susceptibility to the antimicrobial agents (Nazzaro et al., 2013). A rrecent study demonstrated that plant extracts can serve as alternatives or adjuncts to conventional antibiotics, as their various compounds possess antimicrobial properties that may help combat antimicrobial resistance, a primary global health concern (Hernández-Bolaños et al., 2025).
The antibacterial assay showed that P. atrosepticum and S. scabiei exhibited high sensitivity to the extract, even at a lowe concentration (300 µg/ mL), which holds considerable importance. P. atrosepticum is responsible for blackleg disease in potatoes, resulting in significant economic losses in global potato production (Czajkowski et al., 2015). Similarly, S. scabiei is responsible for common scab disease in potato and other root crops, leading to diminished marketability and yield (Zhang et al., 2024). The pronounced sensitivity of these pathogens to E. verticillata extract indicates promising avenues for developing natural bactericides to manage these economically significant plant diseases. The observation that certain bacterial strains, specifically R. solanacearum and S. scabiei, which exhibit resistance to conventional antibiotics, demonstrated notable sensitivity to the plant extract at elevated concentrations (1000 µg/ mL) is particularly intriguing. This finding is consistent with recent studies that emphasized the potential of plant-derived compounds in addressing antibiotic-resistant bacteria (Angelini, 2024). The rise of antimicrobial resistance poses a growing challenge to human health and agriculture, making it particularly crucial to develop alternative antimicrobial agents (Salam et al., 2023; Ferraz, 2024). Our findings indicated that E. verticillata extract may represent a significant source of innovative antimicrobial compounds for tackling this issue. The absence of notable variations in inhibition zone diameters across the various extract concentrations suggests a saturation effect, implying that increasing the concentration beyond a specific limit does not improve the antibacterial activity (Al-Askar et al., 2025; Gaber et al., 2025).
In this study, the methanol extract exhibited significant antifungal efficacy against B. cinerea. This finding holds particular importance as B. cinerea is a necrotrophic fungal pathogen affecting many hosts, leading to gray mold disease in more than 200 plant species and causing considerable pre- and post-harvest losses (Hamdy et al., 2024). The significant antifungal activity demonstrated against R. solani and F. oxysporum highlights the extensive range of the antifungal activities of the extract. R. solani is a soil-borne pathogen responsible for various diseases, including damping-off, root rot, and stem canker, which affects many crops (Abo-Zaid et al., 2024). F. oxysporum is a complex of pathogenic strain responsible for vascular wilt diseases affecting various economically significant plants (Shehzadi et al., 2025). Successful suppression of these pathogens by E. verticillata extract indicates promising possibilities for creating natural fungicides with comprehensive disease management approaches (Salem et al., 2025). The exceptional antifungal efficacy of the plant extracts compared to copper hydroxide, is especially significant (Lamichhane et al., 2018). Copper-based fungicides have been extensively utilized in agriculture for many years; however, worries about their environmental persistence, soil accumulation, and adverse effects on non-target organisms have led to the exploration of alternative solutions (Lamichhane et al., 2018). The findings of this study suggest that E. verticillata extract may serve as an eco-friendly alternative to traditional copper-based fungicides, thereby promoting more sustainable agricultural practices. The mechanisms by which the extract exhibits antifungal properties may include the disruption of ergosterol; a crucial component of fungal cell membranes, or the inhibition of essential enzymes involved in the biosynthesis of fungal cell walls (Eliaš, Tóth Hervay, and Gbelská, 2024; Song et al., 2025).. The evidence suggests that numerous compounds derived from plants exhibit antifungal effects through similar mechanisms (Song et al., 2025; Xie et al., 2025). This finding is consistent with a recent study that showed methanolic extracts from various plant sources frequently exhibited potent antifungal effects by disrupting fungal cell wall synthesis or altering cellular metabolism, thereby inhibiting growth and development (Xie et al., 2025). To elucidate these effects, the phytochemical composition of the extract was analyzed.
High-performance liquid chromatography (HPLC) analysis revealed the presence of various phenolic and flavonoid compounds in the extract. The most abundant phenolic compounds identified were gallic acid (2620.3 µg/ g), caffeic acid (606.5 µg/g), and syringic acid (580.9 µg/ g). Additionally, naringenin (198.4 µg/ g) and quercetin (169.4 µg/ g) were noted as the predominant flavonoids. These results provide valuable insights into the potential bioactive compounds that contribute to the extract’s antimicrobial activity. Gallic acid has been documented to exhibit potent antimicrobial properties against various bacterial and fungal pathogens (Flores-Maldonado et al., 2024). The mechanism of action involved disrupting the cell membrane, inhibiting vital enzymes, and chelating metal ions essential for microbial growth (Nasaj et al., 2024; Khwaza and Aderibigbe, 2025). The elevated levels of gallic acid in the extract probably play a crucial role in enhancing its antimicrobial effectiveness. Caffeic acid and syringic acid have shown antimicrobial properties in earlier studies (Ecevit et al., 2022; Beulah et al., 2025). Caffeic acid exhibits the ability to inhibit bacterial growth by disrupting cell membrane integrity and inhibiting essential cellular processes (Beulah et al., 2025). Syringic acid exhibits antimicrobial properties through comparable mechanisms and has been shown to enhance the efficacy of the standard antibiotics when used in conjunction (Ecevit et al., 2022). Naringenin and quercetin are flavonoids that exhibit well-documented antimicrobial properties (Shamsudin et al., 2022; Veiko et al., 2023; Hasnat et al., 2024). Naringenin can inhibit the growth of bacteria and fungi by disrupting cell membrane function and inhibiting essential enzymes (Liu et al., 2025). Quercetin exhibits antimicrobial activities through various mechanisms, including inhibition of DNA gyrase, disruption of cell membrane integrity, and interference with energy metabolism (Liang et al., 2021). The presence of these flavonoids in the extract probably plays a significant role in its wide-ranging antimicrobial activity. A recent study on E. manipuliflora suggest that the genus Erica is rich in vanillic acid, fumaric acid, catechin hydrate, quercetin, and phloridzin dihydrate, all contributing to its antimicrobial potential (Yüksel et al., 2021). This study revealed that E. manipuliflora extract exhibited effectiveness against several pathogenic microorganisms, such as Staphylococcus aureus, Escherichia coli, and Salmonella typhimurium, reinforcing the antimicrobial potential of the Erica genus.
The GC-MS analysis identified about 15 primary compounds in the extract, with hexadecanoic acid and phorbol being the most prevalent metabolites. This study also detected α-sitosterol, octadecenoic acid, tibolone, stigmasterol acid, and methyl phaseate as less prevalent metabolites. Hexadecanoic acid, also known as palmitic acid, is a saturated fatty acid recognized for its antimicrobial potential (Ganesan et al., 2024). The mechanism of action involved disrupting the cell membrane, resulting in increased permeability and ultimately leading to cell death (Nasaj et al., 2024; El-Bilawy et al., 2025; Khwaza and Aderibigbe, 2025). Numerous studies detailed the direct beneficial effects of fatty acids, such as linoleic acid, palmitic acid, and their derivatives in managing phytopathogens (Zheng et al., 2005; Liu et al., 2008; Sumayo et al., 2014; López-Arellanes et al., 2025). Phorbol and its derivatives exhibit a range of biological activities, notably antimicrobial effects (Goel et al., 2007). The high concentrations of these compounds in the extract may play a significant role in its antimicrobial efficacy. The variety of bioactive compounds found in the E. verticillata extract suggests that its antimicrobial activity is likely attributed to the combined effects of several compounds rather than relying on a single active principle (Guimarães and Venâncio, 2022; Vaou et al., 2022; Gadouche et al., 2023; Jabal et al., 2025). This synergism may enhance overall efficacy and reduces the likelihood of resistance development, making plant extracts crucial in sustainable disease management strategies (Ayaz et al., 2019). Before synthetic pesticides were introduced, the botanical pesticides were widely used in subsistence and commercial agriculture for a long time. The active components in botanical pesticides are naturally occurring substances extracted from plants using organic solvents and/or combined with essential pesticide adjuvants (Ngegba et al., 2022). The effectiveness and efficacy of synthetic pesticides in managing crop diseases led to a gradual decline in the use of plant-based pesticide products. However, concerns about the environmental and health risks of the synthetic pesticides to humans have since emerged.(Chowdhury et al., 2024; Shekhar et al., 2024).
Finally, the findings of this study have important implications for sustainable plant disease management. The current antimicrobial efficacy of E. verticillata extracts against significant phytopathogens, and their enhanced performance relative to traditional antimicrobial agents in certain instances, highlights their potential as natural alternatives to the synthetic pesticides. Identifying specific bioactive compounds in the E. verticillata extract establishes a basis for subsequent studies on isolating and characterizing these compounds for potential commercial development as biopesticides. Moreover, comprehending the chemical composition of the extract may enhance the standardization and quality control of the botanical formulations, thereby addressing a significant challenge in the commercialization of plant-derived antimicrobial products.
Conclusions and Recommendations
This study demonstrated that methanolic extract of Erica verticillata leaves possess broad-spectrum antimicrobial activity against economically significant phytopathogens, including antibiotic-resistant bacterial strains (e.g., Ralstonia solanacearum) and fungi (Botrytis cinerea, Fusarium oxysporum). Notably, the extract outperformed the conventional copper hydroxide fungicides in inhibiting B. cinerea, recotrding 78.2 % vs. 58.7 %, respectively at 5000 µg/mL, highlighting its potential as a sustainable alternative to the synthetic agrochemicals. Phytochemical profiling revealed high concentrations of multiple bioactive compounds, including gallic acid (2620.3 µg/ g), hexadecanoic acid (26.4 %), and phorbol (16.4 %), which likely drive these amntimicrobial impacts through synergistic mechanisms, mainly membrane disruption and enzyme inhibition. Based on the strong antimicrobial efficacy demonstrated, it is recommended that E. verticillata methanolic extract be further explored and developed as a natural biopesticide for integrated plant disease management. Its superior activity against key phytopathogens, especially Botrytis cinerea, positions it as a sustainable alternative to conventional agrochemicals.
Acknowledgement
The authors express their sincere thanks to the City of Scientific Research and Technological Applications (SRTA-City) and the Faculty of Agriculture (Saba Basha), Alexandria University, Egypt, for providing the necessary research facilities.x
Novelty Statement
This study examined the bioactive potential of Erica verticillata methanol extract focusing on its phytochemical profile, and assessing its antibacterial and antifungal properties against various phytopathogenic microorganisms. The study provided a comprehensive evaluation of the bioactive compounds present in E. verticillata extract, highlighting their unique antimicrobial efficacy against phytopathogenic organisms for the first time. Moreover, it highlights the biocontrol prospects of the detected bioactive compounds and lays the groundwork for future applications as a natural source of antimicrobial agents for the sustainable management of plant diseases.
Author’s Contribution
Abdallah Khalil, Karrar A. Hamzah, and Shimaa Bashir: Methodology, Investigations, and Writing the original draft. Said Behiry and Ahmed Abdelkhalek: Supervision, Statistical study design and analysis, Molecular data analysis, Revision and editing of the manuscript. All authors approved the final version of the manuscript.
Funding source
No funds, grants, or other support were received during the preparation of this study.
Ethical approval
None-applicable.
Conflict of interests
The authors have declared no conflicts of interest.
References
Aamer, H.A., Elalem, S.F., Al-Askar, A.A., Sharaf, O.A., Gaber, M.A., Kowalczewski, P., Behiry, S. and Abdelkhalek, A., 2024. Antioxidant and antimicrobial activities of Salsola imbricata methanolic extract and its phytochemical characterization. Open Life Sci., 19: 20221011. https://doi.org/10.1515/biol-2022-1011
Abdelkhalek, A., Abdelwahab, E.A., Elalem, S.F., Al-Askar, A.A., Kowalczewski, P.Ł. and Behiry, S., 2024. Phytochemical composition and antifungal effectiveness of Phoenix dactylifera L. rachis extracts. Polish J. Chem. Technol., 26:76–84. https://doi.org/10.2478/pjct-2024-0037
Abo-Zaid, G.A., Darwish, M.H., Ghozlan, H.A., Abdel-Gayed, M.A. and Sabry, S.A., 2024. Sustainable management of peanut damping-off and root rot diseases caused by Rhizoctonia solani using environmentally friendly bio-formulations prepared from batch fermentation broth of chitinase-producing Streptomyces cellulosae. BMC Plant Biol., 24: 760. https://doi.org/10.1186/s12870-024-05441-6
Al-Askar, A.A., Al-Otibi, F.O., Yassin, M.T., Abo-Zaid, G.A. and Abdelkhalek, A., 2025. Batch fermentation and GC-MS analysis of biocontrol agent, Bacillus amyloliquefaciens strain KSAS6 and its impact on soilborne fungus, Sclerotium bataticola. Adv. Life Sci., 12: 260–270. https://doi.org/10.62940/als.v12i1.3583
Ali, D.F.I., El-Nahrawy, S., EL-Zawawy, H.A.H. and Omara, A.E.D., 2025. Effective applications of Bacillus subtilis and B. amyloliquefaciens as biocontrol agents of damping-off disease and biostimulation of tomato plants. Stresses, 5: 9. https://doi.org/10.3390/stresses5010009
Altemimi, A., Lakhssassi, N., Baharlouei, A., Watson, D.G. and Lightfoot, D.A., 2017. Phytochemicals: Extraction, isolation, and identification of bioactive compounds from plant extracts. Plants, 6: 42. https://doi.org/10.3390/plants6040042
Angelini, P., 2024. Plant-derived antimicrobials and their crucial role in combating antimicrobial resistance. Antibiotics, 13: 746. https://doi.org/10.3390/antibiotics13080746
Arzani, V., Soleimani, M., Fritsch, T., Jacob, U.M., Calabrese, V. and Arzani, A., 2025. Plant polyphenols, terpenes, and terpenoids in oral health. Open Med., 20: 20251183. https://doi.org/10.1515/med-2025-1183
Ayaz, M., Ullah, F., Sadiq, A., Ullah, F., Ovais, M., Ahmed, J. and Devkota, H.P., 2019. Synergistic interactions of phytochemicals with antimicrobial agents: Potential strategy to counteract drug resistance. Chem. Biol. Interact., 308: 294–303. https://doi.org/10.1016/j.cbi.2019.05.050
Azwanida, N.N., 2015. A review on the extraction methods use in medicinal plants, principle, strength and limitation. Med. Aromat. Plants, 4: 412–2167.
Baldwin, B.C., 1990. Inhibitors of ergosterol biosynthesis as crop protection agents. Biochem. Soc. Trans., 18: 61–62. https://doi.org/10.1042/bst0180061
Bashir, S., Behiry, S., Al-Askar, A.A., Kowalczewski, P.Ł., Emaish, H.H. and Abdelkhalek, A., 2024. Antibacterial, antifungal, and phytochemical properties of Salsola kali ethanolic extract. Open Life Sci., 19: 20220962. https://doi.org/10.1515/biol-2022-0962
Benalach, L., Boukada, F., Cherifi, K., Latreche, A., Messellem, I. and Bellatreche, M., 2025. Investigating the bioactive potential of Ononis spinosa L. Phytochemical profiling and antibacterial effectiveness evaluation. Nov. Res. Microbiol. J., 9: 82–91. https://doi.org/10.17582/journal.NRMJ/2025/9.2.82.91
Beulah, K.C., Prasanna, A., Karunakar, P., Rao, A.S., More, S.S. and Nair, A., 2025. Exploring caffeine as a disruptor of membrane integrity and genomic stability in Staphylococcus aureus: Functional and in silico analysis. Arch. Microbiol., 207: 28. https://doi.org/10.1007/s00203-024-04230-x
Bhattacharjee, U. and Sharma, I., 2025. Bioactive compounds in managements of microbial pathogenesis. In: Bioact. Ingredients Healthc. Ind. Vol. 2. p. 109–122. Springer. https://doi.org/10.1007/978-981-96-4379-0_4
Chaachouay, N. and Zidane, L., 2024. Plant-derived natural products: A source for drug discovery and development. Drugs Drug Candid., 3:184207. https://doi.org/10.3390/ddc3010011
Chowdhury, S.K., Banerjee, M., Basnett, D. and Mazumdar, T., 2024. Natural pesticides for pest control in agricultural crops: An alternative and eco-friendly method. Plant Sci. Today, 11: 433–450.
Cotugno, S., De Vita, E., Frallonardo, L., Novara, R., Papagni, R., Asaduzzaman, M., Segala, F.V., Veronese, N., Nicastri, E., Morea, A., Farkas, F.B., Lakatos, B., Iatta, R., Putoto, G., Saracino, A. and Di Gennaro, F., 2025. Antimicrobial resistance and migration: Interrelation between two hot topics in global health. Ann. Glob. Heal., 91: 12. https://doi.org/10.5334/aogh.4628
Czajkowski, R., Pérombelon, M.C.M., Jafra, S., Lojkowska, E., Potrykus, M., Van Der Wolf, J.M. and Sledz, W., 2015. Detection, identification and differentiation of Pectobacterium and Dickeya species causing potato blackleg and tuber soft rot: A review. Ann. Appl. Biol., 166: 18–38. https://doi.org/10.1111/aab.12166
Dai, J. and Mumper, R.J., 2010. Plant phenolics: Extraction, analysis and their antioxidant and anticancer properties. Molecules, 15: 7313–7352. https://doi.org/10.3390/molecules15107313
De Rossi, L., Rocchetti, G., Lucini, L. and Rebecchi, A., 2025. Antimicrobial potential of polyphenols: Mechanisms of action and microbial responses. A narrative review. Antioxidants, 14: 200. https://doi.org/10.3390/antiox14020200
Debnath, S., Das, M., Mondal, S., Sarkar, B.K. and Babu, G., 2025. Advances in chromatography: Contemporary techniques and applications. Essent. Chem., 2: 1–27. https://doi.org/10.1080/28378083.2025.2466624
Dias, P., Falé, P.L., Martins, A. and Rauter, A.P., 2015. Digestibility and bioavailability of the active components of Erica australis L. aqueous extracts and their therapeutic potential as acetylcholinesterase inhibitors. Evidence-based Complement. Altern. Med., 2015: 854373. https://doi.org/10.1155/2015/854373
Ecevit, K., Barros, A.A., Silva, J.M. and Reis, R.L., 2022. Preventing microbial infections with natural phenolic compounds. Futur. Pharmacol., 2: 460–498. https://doi.org/10.3390/futurepharmacol2040030
El-Bilawy, E.H., Mamdouh, I., Behiry, S. and Teiba, I.I., 2025. Evaluating the antibacterial efficacy of bee venom against multidrug-resistant pathogenic bacteria: Escherichia coli, Salmonella typhimurium, and Enterococcus faecalis. World J. Microbiol. Biotechnol., pp. 41-40. https://doi.org/10.1007/s11274-024-04248-9
Eliaš, D., Tóth Hervay, N. and Gbelská, Y., 2024. Ergosterol Biosynthesis and regulation impact the antifungal resistance and virulence of Candida spp. Stresses, 4: 641–662. https://doi.org/10.3390/stresses4040041
Erbs, G. and Newman, M.A., 2024. Plant diseases caused by prokaryotes: Bacteria and mollicutes. In: Agrios’ Plant Pathol. Sixth Ed. p. 465–546. Elsevier. https://doi.org/10.1016/B978-0-12-822429-8.00016-9
Ferraz, M.P., 2024. Antimicrobial resistance: The impact from and on society according to one health approach. Societies, 14: 187. https://doi.org/10.3390/soc14090187
Flores-Maldonado, O., Dávila-Aviña, J., González, G.M., Becerril-García, M.A. and Ríos-López, A.L., 2024. Antibacterial activity of gallic acid and methyl gallate against emerging non-fermenting bacilli. Folia Microbiol. (Praha). 70: 127–135. https://doi.org/10.1007/s12223-024-01182-z
Gaber, M.A., El-Messeiry, S., El-Tanbouly, R., Elkashef, A.A., Behiry, S.I., Aboshosha, S.S. and Aamer, H.A., 2025. Eco-friendly management of Fusarium wilt in tomato using Salvia officinalis methanolic extract: In vitro, in vivo, and molecular docking approaches. J. Plant Pathol., pp. 1–14. https://doi.org/10.1007/s42161-025-01910-5
Gadouche, L., Alsoufi, A.S.M., Pacholska, D., Skotarek, A., Pączkowski, C. and Szakiel, A., 2023. Triterpenoid and steroid content of lipophilic extracts of selected medicinal plants of the mediterranean region. Molecules, 28: 697. https://doi.org/10.3390/molecules28020697
Ganesan, T., Subban, M., Christopher Leslee, D.B., Kuppannan, S.B. and Seedevi, P., 2024. Structural characterization of n-hexadecanoic acid from the leaves of Ipomoea eriocarpa and its antioxidant and antibacterial activities. Biomass Convers. Biorefinery, 14: 14547–14558. https://doi.org/10.1007/s13399-022-03576-w
Goel, G., Makkar, H.P.S., Francis, G. and Becker, K., 2007. Phorbol esters: Structure, biological activity, and toxicity in animals. Int. J. Toxicol., 26: 279–288. https://doi.org/10.1080/10915810701464641
Guendouze-Bouchefa, N., Madani, K., Chibane, M., Boulekbache-Makhlouf, L., Hauchard, D., Kiendrebeogo, M., Stévigny, C., Okusa, P.N. and Duez, P., 2015. Phenolic compounds, antioxidant and antibacterial activities of three Ericaceae from Algeria. Ind. Crops Prod., 70: 459–466. https://doi.org/10.1016/j.indcrop.2015.03.053
Guimarães, A. and Venâncio, A., 2022. The Potential of fatty acids and their derivatives as antifungal agents: A review. Toxins (Basel), 14: 188. https://doi.org/10.3390/toxins14030188
Hamdy, E., El-Gendi, H., Al-Askar, A., El-Far, A., Kowalczewski, P., Behiry, S. and Abdelkhalek, A., 2024. Copper oxide nanoparticles-mediated Heliotropium bacciferum leaf extract: Antifungal activity and molecular docking assays against strawberry pathogens. Open Chem., 22: 20240028. https://doi.org/10.1515/chem-2024-0028
Hasnat, H., Shompa, S.A., Islam, M.M., Alam, S., Richi, F.T., Emon, N.U., Ashrafi, S., Ahmed, N.U., Chowdhury, M.N.R., Fatema, N., Hossain, M.S., Ghosh, A. and Ahmed, F., 2024. Flavonoids: A treasure house of prospective pharmacological potentials. Heliyon, pp. 10. https://doi.org/10.1016/j.heliyon.2024.e27533
Hernández-Bolaños, E., Sánchez-Retuerta, V., Matías-Hernández, L. and Cuyas, L., 2025. Promising applications on the use of medicinal and aromatic plants in agriculture. Discov. Agric., 3: 36. https://doi.org/10.1007/s44279-025-00187-7
Jabal, K.A., Pigott, M., Sheridan, H. and Walsh, J.J., 2025. Mediterranean basin erica species: traditional uses, phytochemistry and pharmacological properties. Molecules, 30: 2616. https://doi.org/10.3390/molecules30122616
Khalil, A., Abdelwahab, E.A., Sharaf, O.A., Al-Askar, A.A., Kowalczewski, P., Abdelkhalek, A. and Behiry, S., 2024. Torilis arvensis ethanolic extract: Phytochemical analysis, antifungal efficacy, and cytotoxicity properties. Open Chem., 22: 20240113. https://doi.org/10.1515/chem-2024-0113
Khwaza, V. and Aderibigbe, B.A. 2025. Antibacterial activity of selected essential oil components and their derivatives: A review. Antibiotics 14: 68. https://doi.org/10.3390/antibiotics14010068
Košćak, L., Lamovšek, J., Đermić, E. and Godena, S., 2025. Potential of plant-based agents as next-generation plant growth-promotors and green bactericides against Pseudomonas savastanoi pv. savastanoi. Agronomy, 15. https://doi.org/10.3390/agronomy15040819
Kumar, A., Nirmal, P., Kumar, M., Jose, A., Tomer, V., Oz, E., Proestos, C., Zeng, M., Elobeid, T., Sneha, V. and Oz, F., 2023. Major phytochemicals: Recent advances in health benefits and extraction method. Molecules, 28: 887. https://doi.org/10.3390/molecules28020887
Lamichhane, J.R., Osdaghi, E., Behlau, F., Köhl, J., Jones, J.B. and Aubertot, J.N., 2018. Thirteen decades of antimicrobial copper compounds applied in agriculture. A review. Agron. Sustain. Dev., 38: 28. https://doi.org/10.1007/s13593-018-0503-9
Liang, H., Ma, Z., Sun, R. and Song, L., 2021. Research progress on antibacterial activity of bioactive flavonoids. China Surfactant Deterg. Cosmet., 51: 775–781.
Liu, S., Ruan, W., Li, J., Xu, H., Wang, J., Gao, Y. and Wang, J., 2008. Biological control of phytopathogenic fungi by fatty acids. Mycopathologiam 166: 93–102. https://doi.org/10.1007/s11046-008-9124-1
Liu, Y., Zhu, J., Liu, Z., Zhi, Y., Mei, C. and Wang, H., 2025. Flavonoids as promising natural compounds for combating bacterial infections. Int. J. Mol. Sci., 26: 2455. https://doi.org/10.3390/ijms26062455
López-Arellanes, M.E., López-Pacheco, L.D., Elizondo-Luevano, J.H. and González-Meza, G.M., 2025. Algae and cyanobacteria fatty acids and bioactive metabolites: Natural Antifungal Alternative Against Fusarium sp. Microorganisms, 13: 439. https://doi.org/10.3390/microorganisms13020439
Murugan, S., Senthilvelan, T., Govindasamy, M. and Thangavel, K., 2025. A comprehensive review on exploring the potential of phytochemicals and biogenic nanoparticles for the treatment of antimicrobial-resistant pathogenic bacteria. Curr. Microbiol., 82: 90. https://doi.org/10.1007/s00284-025-04064-w
Naczk, M. and Shahidi, F., 2004. Extraction and analysis of phenolics in food. J. Chromatogr. A, 1054: 95–111. https://doi.org/10.1016/j.chroma.2004.08.059
Nasaj, M., Chehelgerdi, M., Asghari, B., Ahmadieh-Yazdi, A., Asgari, M., Kabiri-Samani, S., Sharifi, E. and Arabestani, M., 2024. Factors influencing the antimicrobial mechanism of chitosan action and its derivatives: A review. Int. J. Biol. Macromol., 277: 134321. https://doi.org/10.1016/j.ijbiomac.2024.134321
Nazzaro, F., Fratianni, F., De Martino, L., Coppola, R. and De Feo, V., 2013. Effect of essential oils on pathogenic bacteria. Pharmaceuticals, 6: 1451–1474. https://doi.org/10.3390/ph6121451
Ngegba, P.M., Cui, G., Khalid, M.Z. and Zhong, G., 2022. Use of botanical pesticides in agriculture as an alternative to synthetic pesticides. Agriculture, 12: 600. https://doi.org/10.3390/agriculture12050600
Nguyen, V.D.H., Nguyen, T.T.T., Huynh, T.N.P., Ho, H.H., Nguyen, A.T.V. and Trinh, L.T.P., 2024. Effective control of Fusarium wilt on tomatoes using a combination of phenolic-rich plant extracts. Eur. J. Plant Pathol., 170: 833–850. https://doi.org/10.1007/s10658-024-02830-3
Ojewole, J.A.O., 2008. Anticonvulsant property of Sutherlandia frutescens R. BR. (variety Incana E. MEY.) [Fabaceae] shoot aqueous extract. Brain Res. Bull. 75:126–132. https://doi.org/10.1016/j.brainresbull.2007.08.002
Pereira, A.G., Echave, J., Jorge, A.O.S., Nogueira-Marques, R., Nur Yuksek, E., Barciela, P., Perez-Vazquez, A., Chamorro, F., Maria, M.B., Carpena, M. and Prieto, M.A., 2025. Therapeutic and preventive potential of plant-derived antioxidant nutraceuticals. Foods, 14: 1749. https://doi.org/10.3390/foods14101749
Pérez-Flores, J.G., García-Curiel, L., Pérez-Escalante, E., Contreras-López, E., Aguilar-Lira, G.Y., Ángel-Jijón, C., González-Olivares, L.G., Baena-Santillán, E.S., Ocampo-Salinas, I.O., Guerrero-Solano, J.A. and Portillo-Torres, L.A., 2025. Plant Antimicrobial compounds and their mechanisms of action on spoilage and pathogenic bacteria: A bibliometric study and literature review. Appl. Sci., 15: 3516. https://doi.org/10.3390/app15073516
Salam, M.A., Al-Amin, M.Y., Salam, M.T., Pawar, J.S., Akhter, N., Rabaan, A.A. and Alqumber, M.A.A., 2023. Antimicrobial resistance: A growing serious threat for global public health. In: Healthc. p. 1946. Multidisciplinary Digital Publishing Institute., Vol. 11. https://doi.org/10.3390/healthcare11131946
Salem, M.Z.M., Hassan, A.G.A., Amer, A.M.E., Abdullah, M.F.G., Ahmed, S.M.A., Mahmoud, M.M., Philip, B. and Behiry, S.I., 2025. Bio-based chemical analysis of extracts from the biomass residues of Ceratonia siliqua and Ziziphus spina-christi with their bioactivities against molecularly identified fungi. Biomass Convers. Biorefinery, pp. 1–17. https://doi.org/10.1007/s13399-025-06651-0
Shamsudin, N.F., Ahmed, Q.U., Mahmood, S., Shah, S.A.A., Khatib, A., Mukhtar, S., Alsharif, M.A., Parveen, H. and Zakaria, Z.A., 2022. Antibacterial effects of flavonoids and their structure-activity relationship study: A comparative interpretation. Molecules, 27: 1149. https://doi.org/10.3390/molecules27041149
Shehzadi, L., Anum, S., Shabbir, M.A., Sajid, A., Anwar, M., Fareed, M.A. and Usman, H.M., 2025. Overview of Fusarium oxysporum f. sp. vasinfectum causing okra wilt and its management. J. Agric. Food, 6: 82–93.
Shekhar, C., Khosya, R., Thakur, K., Mahajan, D., Kumar, R., Kumar, S. and Sharma, A.K., 2024. A systematic review of pesticide exposure, associated risks, and long-term human health impacts. Toxicol. Rep., 13: 101840. https://doi.org/10.1016/j.toxrep.2024.101840
Song, L., Wang, S., Zou, H., Yi, X., Jia, S., Li, R. and Song, J., 2025. Regulation of ergosterol biosynthesis in pathogenic fungi: Opportunities for therapeutic development. Microorganisms, 13: 862. https://doi.org/10.3390/microorganisms13040862
Sumayo, M.S., Kwon, D.K. and Ghim, S.Y., 2014. Linoleic acid-induced expression of defense genes and enzymes in tobacco. J. Plant Physiol., 171: 1757–1762. https://doi.org/10.1016/j.jplph.2014.08.015
Valgas, C., De Souza, S.M., Smânia, E.F.A. and Smânia, A., 2007. Screening methods to determine antibacterial activity of natural products. Braz. J. Microbiol., 38: 369–380. https://doi.org/10.1590/S1517-83822007000200034
Vaou, N., Stavropoulou, E., Voidarou, C., Tsakris, Z., Rozos, G., Tsigalou, C. and Bezirtzoglou, E., 2022. Interactions between medical plant-derived bioactive compounds: Focus on antimicrobial combination effects. Antibiotics, 11: 1014. https://doi.org/10.3390/antibiotics11081014
Veiko, A.G., Olchowik-Grabarek, E., Sekowski, S., Roszkowska, A., Lapshina, E.A., Dobrzynska, I., Zamaraeva, M. and Zavodnik, I.B., 2023. Antimicrobial activity of quercetin, naringenin and catechin: Flavonoids inhibit Staphylococcus aureus-induced hemolysis and modify membranes of bacteria and erythrocytes. Molecules, 28: 1252. https://doi.org/10.3390/molecules28031252
Xie, Q., Peng, F., Wang, X., Du, B. and Yang, Y., 2025. Chestnut flower extract as a natural inhibitor of Fusarium graminearum: Antifungal activity and mechanisms. Pest Manag. Sci., https://doi.org/10.1002/ps.8708
Yihune, E. and Yemata, G., 2019. Antibacterial activity of medicinal plant extracts against Ralstonia solanacearum (Smith) that causes bacterial wilt in hot pepper (Capsicum annuum L.). Acta Sci. Biol. Sci., 41: 45402. https://doi.org/10.4025/actascibiolsci.v41i1.45402
Youssef, N.H., Qari, S.H., Behiry, S.I., Dessoky, E.S., El-Hallous, E.I., Elshaer, M.M., Kordy, A., Maresca, V., Abdelkhalek, A. and Heflish, A.A., 2021. Antimycotoxigenic activity of beetroot extracts against Altenaria alternata mycotoxins on potato crop. Appl. Sci., 11: 4239. https://doi.org/10.3390/app11094239.
Yüksel, A.K., Dikici, E., Yüksel, M., Işik, M., Tozoğlu, F. and Köksal, E., 2021. Phytochemicals analysis and some bioactive properties of Erica manipuliflora Salisb (EMS); Antibacterial, Antiradical and Anti-lipid Peroxidation. Iran. J. Pharm. Res., 20: 422–434.
Zhang, H., Ping, Y., Liu, X., He, X. and Du, C., 2024. Pathogenic factors of plant pathogenic Streptomyces. Potato Res., 67: 621–646. https://doi.org/10.1007/s11540-023-09660-6
Zheng, C.J., Yoo, J.S., Lee, T.G., Cho, H.Y., Kim, Y.H. and Kim, W.G., 2005. Fatty acid synthesis is a target for antibacterial activity of unsaturated fatty acids. FEBS Lett., 579: 5157–5162. https://doi.org/10.1016/j.febslet.2005.08.028