Compatibility and synergistic effects of Mirabilis jalapa nanoemulsion and the entomopathogenic fungus Beauveria bassiana against Spodoptera frugiperda larvae

Compatibilidade e efeitos sinérgicos da nanoemulsão de Mirabilis jalapa e do fungo entomopatogênico Beauveria bassiana contra larvas de Spodoptera frugiperda

Abstract

This study evaluated the compatibility and insecticidal efficacy of Mirabilis jalapa nanoemulsion combined with the entomopathogenic fungus Beauveria bassiana against Spodoptera frugiperda larvae. Botanical pesticides and entomopathogenic fungi are considered environmentally friendly alternatives to synthetic insecticides; however, their combined application requires compatibility assessment to avoid antagonistic effects. Compatibility tests were conducted to determine the effect of nanoemulsion formulations on fungal viability. Larval mortality bioassays were then performed using single and combined treatments. Nanoemulsion formulations included F1 (0.1% and 0.8%) and F2 (0.1%, 0.2%, 0.4%, and 0.8%), while B. bassiana was applied at a density of 2.68 × 10⁷ spores mL⁻¹. The results showed that combined treatments significantly increased larval mortality compared to individual applications (ANOVA, Tukey’s test, p < 0.05) at 7 days after application. The combination also reduced the lethal time (LT₅₀), indicating a synergistic interaction. No inhibitory effect on fungal growth was observed, suggesting compatibility between the nanoemulsion and B. bassiana. In conclusion, the combined application shows potential as an environmentally friendly strategy for controlling S. frugiperda under laboratory conditions. Further studies are required to evaluate its effectiveness under field conditions.

Keywords:
nanoemulsion; Beauveria bassiana; compatibility; mortality; Spodoptera frugiperda

Resumo

Este estudo avaliou a compatibilidade e a eficácia inseticida da nanoemulsão de Mirabilis jalapa combinada com o fungo entomopatogênico Beauveria bassiana contra larvas de Spodoptera frugiperda. Pesticidas botânicos e fungos entomopatogênicos são considerados alternativas ecologicamente corretas aos inseticidas sintéticos; no entanto, sua aplicação combinada requer avaliação de compatibilidade para evitar efeitos antagônicos. Testes de compatibilidade foram conduzidos para determinar o efeito das formulações de nanoemulsão na viabilidade do fungo. Bioensaios de mortalidade larval foram então realizados utilizando tratamentos individuais e combinados. As formulações de nanoemulsão incluíram F1 (0,1% e 0,8%) e F2 (0,1%, 0,2%, 0,4% e 0,8%), enquanto B. bassiana foi aplicada na densidade de 2,68 × 10⁷ esporos mL⁻¹. Os resultados mostraram que os tratamentos combinados aumentaram significativamente a mortalidade larval em comparação com as aplicações individuais (ANOVA, teste de Tukey, p < 0,05) 7 dias após a aplicação. A combinação também reduziu o tempo letal (TL₅₀), indicando uma interação sinérgica. Não foi observado efeito inibitório no crescimento do fungo, sugerindo compatibilidade entre a nanoemulsão e B. bassiana. Em conclusão, a aplicação combinada demonstra potencial como uma estratégia ecologicamente correta para o controle de S. frugiperda em condições de laboratório. Estudos adicionais são necessários para avaliar sua eficácia em condições de campo.

Palavras-chave:
nanoemulsão; Beauveria bassiana; compatibilidade; mortalidade; Spodoptera frugiperda

1. Introduction

Biopesticides have gained increasing attention as environmentally friendly alternatives to synthetic pesticides due to their biodegradability, lower toxicity to non-target organisms, and compatibility with integrated pest management (IPM) programs. However, their application remains limited due to relatively low efficacy, slow mode of action, and the need for repeated applications (Lengai et al., 2020). In addition, essential oil–based pesticides often consist of volatile compounds that are prone to rapid degradation and environmental instability, reducing their persistence and effectiveness under field conditions (Devrnja et al., 2022; Sarmah et al., 2025).

To overcome these limitations, nanoformulation technologies such as nanoemulsions have been developed to improve the stability, bioavailability, and controlled release of bioactive compounds. Botanical nanoemulsions have been shown to enhance insecticidal activity by improving penetration and prolonging residual effects compared to conventional plant extracts (Pan et al., 2023; Ayllón-Gutiérrez et al., 2024). Therefore, integrating nanobiopesticides with biological control agents represents a promising strategy for improving pest management efficiency.

Entomopathogenic fungi (EPF), such as Beauveria bassiana, are widely recognized as effective biological control agents due to their ability to infect insect hosts directly through the cuticle. B. bassiana has been extensively studied because of its broad host range, ease of mass production, and proven pathogenicity against various agricultural pests. However, the effectiveness of fungal biocontrol agents is influenced by formulation, environmental conditions, and interactions with other control components, making compatibility a critical factor in combined applications (Ahmad et al., 2025).

Previous studies have demonstrated that combining EPF with botanical or other biological agents can enhance pest mortality and improve control efficiency. For example, combinations of B. bassiana with other entomopathogenic fungi or plant-derived compounds have shown synergistic effects against several insect pests (Ma et al., 2019; Basit et al., 2021). Recent studies have also reported increased mortality of Spodoptera frugiperda when treated with combined biological and botanical agents (Suryani et al., 2024a, b). These findings highlight the potential of integrated biological approaches in pest management.

Despite these advances, the compatibility between nanoemulsion-based botanical formulations and entomopathogenic fungi remains poorly understood. Active compounds in botanical formulations may negatively affect fungal germination, growth, sporulation, and virulence, potentially reducing their effectiveness in integrated pest management programs (Islam et al., 2010; Folorunso et al., 2024). Therefore, compatibility assessment is essential prior to combined application.

To date, no study has specifically evaluated the compatibility and synergistic effects of Mirabilis jalapa nanoemulsion combined with Beauveria bassiana against Spodoptera frugiperda. Therefore, this study aimed to: (1) evaluate the compatibility between M. jalapa nanoemulsion and B. bassiana, and (2) quantify their effects on larval mortality and lethal time (LT₅₀) of S. frugiperda. It was hypothesized that the nanoemulsion would be compatible with B. bassiana and that their combined application would result in a synergistic interaction, leading to higher larval mortality and reduced LT₅₀ compared to individual treatments.

2. Materials and Methods

2.1. Study period

The study was conducted from January to October 2025 at the Laboratory of Pests and Diseases, Faculty of Agriculture, Hasanuddin University, and the Biology Laboratory, State University of Makassar. All bioassays were performed under controlled laboratory conditions at 27 ± 2 °C, relative humidity of 71 ± 5%, and a photoperiod of 12:12 (L:D) hours.

2.2. Insect rearing

Third-instar larvae of S. frugiperda were collected from maize fields in Takalar and reared on fresh organic maize leaves under laboratory conditions until used in the experiments.

2.3. Preparation of Beauveria bassiana

The fungus Beauveria bassiana was cultured on Potato Dextrose Agar (PDA) medium and incubated for 14–15 days at room temperature until sporulation. Conidia were harvested and suspended in sterile distilled water. Spore density was initially adjusted to 1 × 10⁷ spores mL⁻¹ following Sepe et al. (2020) and subsequently determined using a hemocytometer and serial dilution. The final spore concentration used in the experiment was 2.68 × 10⁷ spores mL⁻¹.

2.4. Preparation of nanoemulsion

Two nanoemulsion formulations (F1 and F2) of Mirabilis jalapa extract were prepared following Suryani and Anggraeni (2014) with minor modifications using a magnetic stirrer and ultrasonic homogenizer. The concentrations tested were F1 (0.1% and 0.8%) and F2 (0.1%, 0.2%, 0.4%, and 0.8%).

2.5. Experimental design

The experiment was arranged in a Completely Randomized Design (CRD) consisting of seven treatments: control (untreated), nanoemulsion alone, B. bassiana alone, and combinations of nanoemulsion formulations with B. bassiana. Each treatment consisted of 10 larvae per replicate with three replicates (n = 30 larvae per treatment).

2.6. Compatibility test

Compatibility between nanoemulsion and B. bassiana was evaluated using a disc diffusion method. A 0.1 mL aliquot of fungal suspension (10⁷ spores mL⁻¹) was evenly spread onto PDA plates using a sterile spreader. Sterile paper discs (5 mm diameter) soaked in nanoemulsion formulations were placed on the agar surface. Inhibition zones were measured at 24, 48, and 72 hours using a ruler. The inhibition index (IF) was calculated as Equation 1:

$\frac{\begin{array}{c}\text{Diameter of Clear Zone}\left(\text{mm}\right)-\\ \text{Diameter of Paper Disc}\left(\text{mm}\right)\end{array}}{\text{Diameter of Paper Disc}\left(\text{mm}\right)}$ (1)

Inhibition levels were categorized as weak (≤5 mm), moderate (6–10 mm), strong (11–20 mm), or very strong (>21 mm) (Katili et al., 2020). Weak inhibition was interpreted as compatibility between the nanoemulsion and fungal growth. All treatments were performed in triplicate and analyzed statistically.

2.7. Larval mortality bioassay

Following compatibility assessment, larval bioassays were conducted. Nanoemulsion is applied to the larval feed, while B. bassiana is applied by dropping the suspension on the dorsal part of the larva (Figure 1). Mortality was recorded daily for 7 days after application (DAA). Mortality percentage was calculated as: (number of dead larvae / total larvae) × 100

Figure 1
Application of nanoemulsion B. bassiana on S. frugiperda larvae.

Mortality data were corrected using Abbott’s formula when control mortality exceeded 5% (Equation 2):

$$CorrectedMortality\left(\%\right)=\frac{\left(\text{Mt}-\text{Mc}\right)}{\left(100-\text{Mc}\right)}x100$$

where Mt = mortality in treatment (%) and Mc = mortality in control (%).

2.8. Lethal time (LT₅₀) analysis

The lethal time required to kill 50% of the larval population (LT₅₀) was estimated using probit analysis in SPSS version 27. Model fit was evaluated based on goodness-of-fit statistics, and 95% confidence intervals were calculated.

2.9. Statistical analysis

Mortality data were analyzed using analysis of variance (ANOVA). Prior to analysis, data were tested for normality (Shapiro–Wilk test) and homogeneity of variance (Levene’s test). When assumptions were met, ANOVA was followed by Tukey’s HSD test at a significance level of p < 0.05. All results are presented as mean ± standard deviation (SD).

3. Results and Discussion

Compatibility tests revealed weak inhibition zones (<5 mm), indicating that M. jalapa nanoemulsion and B. bassiana were compatible. This finding aligns with prior studies showing that B. bassiana and M. anisopliae are compatible with neem oil (Depieri et al., 2005; Gebramariam et al., 2022; Jaber et al., 2018). Larval mortality increased significantly with combined treatments. The highest mortality (100%) occurred in treatment P3 (F2 MJ 0.1 + BB 10⁷) on day 4 post-application, followed by P5 (F2 MJ 0.4 + BB 10⁷) with 86.7%. The combination treatments showed significantly higher mortality than controls (p < 0.05).

The LT₅₀ analysis showed that treatment P5 achieved 50% mortality within 0.4 days post-application, the fastest among all treatments, followed by P1 (0.9 days). This rapid mortality indicates a synergistic interaction between plant-derived bioactive compounds and fungal pathogens (Prastiwi et al., 2023). Synergistic interactions between biological agents and botanical compounds are increasingly highlighted as effective tools for overcoming pest resistance and enhancing control efficacy within integrated pest management frameworks (Bajda and Grigoraki, 2020). Such combinations can also influence insect behavior and physiological responses, leading to faster mortality and reduced fitness, as emphasized in recent insect science reviews (Van Leeuwen et al., 2020). Morphological observations (Figure 2) showed infected larvae darkened and decomposed, with fungal growth on their surfaces confirming infection. B. bassiana infects insects by adhering to and penetrating the cuticle using enzymes such as chitinases, proteases, and lipases, then producing toxins like beauvericin, bassianin, and bassianolide that disrupt tissues and the nervous system (Arnoldi et al., 2022; Wahjono et al., 2024). The ribotoxin BbRib also induces apoptosis via ROS-mediated stress, allowing the fungus to overcome host immune defenses (Ma et al., 2021).

Figure 2
Compatibility assay of M. jalapa nanoemulsion against B. bassiana. Upper row: control discs (Aquades and 96% ethanol) in three replicates (a-c). Lower row: F2 nanoemulsion discs (0.1%, 0.2%, 0.4%, and 0.8%) in three replicates (d-f). No inhibition zone (>5mm) was observed, indicating compatibility between the nanoemulsion formulations and B. bassiana.

Table 1 shows that the highest larval mortality was achieved in treatment P3 (F2 MJ 0.1 + BB 10⁷), reaching 100% mortality on the fourth day after application. High mortality was also recorded in treatment P5 (F2 MJ 0.4 + BB 10⁷), which exhibited a significant mortality rate (86.67%) starting from day two and remaining constant until day seven. Based on this analysis, the combination of the two biocontrol agents demonstrated significantly higher effectiveness compared to the control (P0).

Table 1
Percentage of S. frugiperda larval mortality up to 7 days after application (DAA) following combined treatment of nanoemulsio formulations and B. bassiana.

Treatment	1 DAA	2 DAA	3 DAA	4 DAA	5 DAA	6 DAA	7 DAA
P0 (Etanol 96% + BB 107)	6.67 ± 11.55	20a ± 20.00	20a ± 20.00	26.67a ± 23.09	26.67a ± 23.09	26.67a ± 23.09	26.67a ± 23.09
P1 (F1 MJ 0.1 + BB 107)	53.33 ± 30.55	60ab ± 20.00	80bc ± 20.00	86.67b ± 11.55	86.67b ± 11.55	86.67b ± 11.55	86.67b ± 11.55
P2(F1 MJ 0.8+ BB 107)	26.67 ± 11.55	40ab ± 0.00	40ab ± 0.00	40a ± 0.00	53.33ab ± 23.09	66.67ab ± 11.55	73.33b ± 11.55
P3 (F2 MJ 0.1+ BB 107)	46.67 ± 23.09	53.33ab ± 11.55	86.67c ± 11.55	100b ± 0.00	100b ± 0.00	100b ± 0.00	100b ± 0.00
P4 (F2 MJ 0.2+ BB 107)	46.67 ± 30.55	53.33ab ± 23.09	53.33abc ± 23.09	60ab ± 20.00	60ab ± 20.00	60ab ± 20.00	60ab ± 20.00
P5 (F2 MJ 0.4+ BB 107)	60 ± 20.00	80b ± 20.00	86.67c ± 11.55	86.67b ± 11.55	86.67b ± 11.55	86.67b ± 11.55	86.67b ± 11.55
P6 (F2 MJ 0.8+ BB 107)	46.67 ± 11.55	73.33b ± 23.09	86.67c ± 11.55	86.67b ± 11.55	86.67b ± 11.55	93.33b ± 11.55	93.33b ± 11.55

• Note: Means ± SD followed by different letters in the same column are significantly different at p < 0.05 according to Tukey’s test. DAA: Days after application; MJ: M. jalapa; BB: B. bassiana.

Consistent with preliminary observations the mortality rate of S. frugiperda larvae was higher under combined treatments than under single applications (either M. jalapa or B. bassiana alone). These findings align with previous reports suggesting that the toxic compounds in M. jalapa accelerate pest mortality by suppressing the insect immune system. The immune system of the larvae weakened following exposure to M. jalapa extract. Several studies have shown that M. jalapa leaf phytochemicals—such as terpenoids, flavonoids, alkaloids, phenols, and sterols—exhibit antimicrobial and insecticidal properties. Alkaloids can exert toxic effects on insects by disrupting their digestive, circulatory, and nervous systems, while phenolic compounds are known to cause dehydration and mortality (Saikia and Kundu, 2023; Ardiansyah et al., 2023; Kasmara et al., 2018).

As shown in Table 2, the treatment with the fastest estimated time to cause 50% larval mortality was P5, with an LT₅₀ value of approximately 0.4 days after application. The next fastest treatment was P1 (0.1), indicating that specific formulations were highly effective in controlling S. frugiperda larvae. A lower LT₅₀ value implies higher toxicity and faster larval death, often resulting from the synergistic interaction between plant bioactive compounds and fungal pathogens.

Table 2
Lethal Time (LT₅₀) for S. frugiperda larval mortality under combined treatments.

Treatment	LethalTime (Days)	Lower limit	Upper limit
P0 (Etanol 96% + BB 107)	2.780	1.762	3.805
P1 (F1 MJ 0.1 + BB 107)	0.945	0.083	1.690
P2(F1 MJ 0.8+ BB 107)	3.636	2.091	6.985
P3 (F2 MJ 0.1+ BB 107)	1.313	0.603	1.843
P4 (F2 MJ 0.2+ BB 107)	1.448	0	0
P5 (F2 MJ 0.4+ BB 107)	0.414	0	1.209
P6 (F2 MJ 0.8+ BB 107)	1.002	0.255	1.609

The combination of M. jalapa and B. bassiana exhibited enhanced toxicity, as the combined treatment induced larval mortality more rapidly than single-agent applications. These findings are supported by Prastiwi et al. (2023), who reported that the combination of B. bassiana and neem leaf extract effectively controlled Spodoptera exigua larvae, showing an LT₅₀ of 5.79 days—shorter than B. bassiana alone (7.96 days) or neem extract alone (6.1 days).

Figure 3 illustrates that larvae infected with the combined treatment of M. jalapa and B. bassiana exhibited distinct morphological changes. The infected larvae appeared darkened, decomposed, and covered with fungal growth, confirming successful infection and colonization. B. bassiana employs several complex mechanisms to infect and kill insects. Infection begins with spore adhesion and germination on the insect cuticle, followed by the formation of an appressorium that penetrates the cuticle using hydrolytic enzymes such as chitinase, protease, and lipase. After penetration, the fungus proliferates through the hemolymph and produces toxic secondary metabolites—including beauvericin, bassianin, bassiacridin, bassianolide, cyclosporine, oosporein, and tenellin—that disrupt host tissues and the nervous system, ultimately leading to death (Arnoldi et al., 2022; Wahjono et al., 2024). From a broader perspective, entomopathogenic fungi such as Beauveria bassiana are widely recognized as effective biocontrol agents for invasive and economically important pests, contributing not only to direct mortality but also to long-term population suppression (Vilcinskas, 2019). In previous studies, B. bassiana alone has been reported to cause moderate to high mortality in Spodoptera frugiperda, although its effectiveness can vary depending on environmental conditions and formulation.

Figure 3
Morphology of S. frugiperda larvae after treatment P6 with combined formulations of nanoemulsion and B. bassiana.

In the present study, the combined application with Mirabilis jalapa nanoemulsion resulted in increased and faster larval mortality compared to fungal treatment alone, suggesting improved efficacy. This enhancement may be related to physiological and biochemical mechanisms of fungal infection, including the production of ribotoxin proteins such as BbRib, which induce apoptosis through reactive oxygen species (ROS)-mediated oxidative stress and accelerate host deterioration (Ma et al., 2021).

Furthermore, B. bassiana is known to establish endophytic associations with plants, which may contribute to indirect plant protection and improved pest resistance (Arnoldi et al., 2022). However, despite these promising mechanisms, the results obtained in this study were based on laboratory conditions, and their effectiveness under field conditions may be influenced by environmental factors such as temperature, humidity, and UV exposure.

Therefore, while the integration of nanoemulsion-based botanical formulations with entomopathogenic fungi represents a promising approach, further studies are required to evaluate its consistency, persistence, and practical applicability in agricultural systems (Ahmad et al., 2025).

4. Conclusion

The combined application of Mirabilis jalapa nanoemulsion and Beauveria bassiana significantly increased larval mortality and reduced the lethal time (LT₅₀) of Spodoptera frugiperda compared to individual treatments under laboratory conditions. These findings indicate a potential positive interaction between the nanoemulsion and the entomopathogenic fungus. The results also suggest that the nanoemulsion did not inhibit fungal growth, indicating compatibility between both agents for combined application. However, the interaction type (e.g., synergistic, additive, or antagonistic) was not quantitatively assessed in this study and therefore requires further investigation. Although the combined treatment showed improved efficacy under controlled conditions, its performance under field conditions and potential effects on non-target organisms remain to be evaluated. Future research should focus on detailed interaction analysis, field validation, formulation stability, and ecological safety to support the development of integrated and environmentally sound pest management strategies.

Acknowledgements

The authors gratefully acknowledge the Ministry of Education, Culture, Research, and Technology of the Republic of Indonesia and the Research and Community Service Institute (DPPM), Universitas Negeri Makassar, for financial support through the 2025 Research Grant Program under Decree No. 0419/C3/DT.05.00/2025 and Research Agreement/Contract No. 084/C3/DT.05.00/PL/2025; 2855/UN36.11/TU/2025.

Data Availability Statement

The datasets generated and/or analyzed during the current study are available from the corresponding author upon reasonable request.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  27 July 2026
• Date of issue
  2026

History

• Received
  04 Feb 2026
• Accepted
  25 May 2026