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Article

Morinda citrifolia Essential Oil in the Control of Banana Anthracnose: Impacts on Phytotoxicity, Preventive and Curative Effects and Fruit Quality

by
Maysa C. Santos
1,
Luis O. Viteri
1,2,3,
Paulo R. Fernandes
1,
Rosilene C. Carvalho
1,
Manuel A. Gonzalez
1,
Osmany M. Herrera
1,
Pedro R. Osório
1,
Dalmarcia S. C. Mourão
1,
Sabrina H. Araujo
4,
Cristiano B. Moraes
4,
Marcos V. Giongo
1,4,
Wellington S. Moura
5,
Marcos P. Camara
6,
Alex Sander R. Cangussu
2,
Raimundo W. S. Aguiar
1,2,
Eugênio E. Oliveira
1,2,7 and
Gil R. Santos
1,2,4,*
1
Programa de Pós-Graduação em Produção Vegetal, Universidade Federal do Tocantins, Gurupi 77402-970, Brazil
2
Programa de Pós-Graduação em Biotecnologia, Universidade Federal do Tocantins, Gurupi 77402-970, Brazil
3
Programa de Pós-Graduação em Biologia Animal, Universidade Federal de Viçosa, Viçosa 36570-900, Brazil
4
Programa de Pós-Graduação em Ciências Florestais e Ambientais, Universidade Federal do Tocantins, Gurupi 77402-970, Brazil
5
Programa de Pós-Graduação em Biodiversidade e Biotecnologia—Rede Bionorte, Universidade Federal do Tocantins, Gurupi 77402-970, Brazil
6
Departamento de Agronomia, Universidade Federal Rural de Pernambuco, Recife 52171-900, Brazil
7
Departamento de Entomologia, Universidade Federal de Viçosa, Viçosa 36570-900, Brazil
*
Author to whom correspondence should be addressed.
Microbiol. Res. 2025, 16(7), 149; https://doi.org/10.3390/microbiolres16070149
Submission received: 24 May 2025 / Revised: 14 June 2025 / Accepted: 24 June 2025 / Published: 3 July 2025
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Abstract

Bananas, one of the most widely consumed tropical fruits in the world, are susceptible to attack by the anthracnose fungus Colletotrichum musae during the post-harvest period. Currently, fungus control is generally based on the use of chemical products, often applied a few days before harvest, which could lead to a risk of residues in the fruit, thus creating a high demand for fresh and organic fruits. Therefore, essential oils present an emerging alternative for the treatment of anthracnose. Here, we evaluated the chemical composition and potential of Morinda citrifolia essential oil as a preventive and curative measure to control C. musae in bananas, also considering the quality of the fruit. In addition, computational docking analysis was conducted to predict potential molecular interactions between octanoic and butanoic acids and the enzyme Tyrosine tRNA, as a potential target for the M. citrifolia essential oil fungicide actions. We also evaluated the essential oil’s safety for beneficial organisms such as the fungus Trichoderma asperellum and the ladybugs Eriopis connexa Germar and Coleomegilla maculata DeGeer. Initially, in vitro growth inhibition tests were performed with doses of 10.0, 30.0, and 50.0 µL/mL of M. citrifolia essential oil, as well as an assessment of the phytotoxic effects on the fruit. Subsequently, using non-phytotoxic doses, we evaluated the effect of the essential oil as a preventive and curative measure against anthracnose and its impact on fruit quality. Our results showed that octanoic, butanoic, and hexanoic acids were the major compounds in M. citrifolia essential oil, inhibiting the growth of C. musae by interacting with the Tyrosine tRNA enzyme of C. musae. The non-phytotoxic dose on the fruit was 10 µL/mL of noni essential oil, which reduced C. musae growth by 30% when applied preventively and by approximately 25% when applied as a curative measure. This significantly reduced the Area Under the Disease Progress Curve without affecting the fruit weight, although there was a slight reduction in °Brix. The growth of non-target organisms, such as T. asperellum and the insect predators Co. maculata and E. connexa, was not affected. Collectively, our findings suggest that M. citrifolia essential oil is a promising alternative for the prevention and control of anthracnose in banana fruit caused by C. musae, without adversely affecting its organoleptic characteristics or non-target organisms.

1. Introduction

Anthracnose is a widespread post-harvest disease affecting fruit, caused by various species of the Colletotrichum genus. Tropical and subtropical fruits such as bananas, mangoes, papayas, and avocados are particularly susceptible [1,2,3]. The greatest deterioration of the fruit occurs post-harvest, caused by fungal pathogens, impacting the appearance and quality of the fruit, with anthracnose, caused by C. musae, being the most aggressive [4,5,6]. Typical symptoms include black or dark brown lesions with conidial masses on the fruit surface. Infection accelerates ripening and, in advanced stages, leads to rotting, rendering the fruit unfit for consumption or sale [7,8,9]. Consumer decisions are heavily influenced by the fruit’s taste and quality.
Bananas are among the most widely consumed tropical fruits globally, playing a key role in food security and holding significant economic value [10,11]. They are grown in over 135 countries across tropical and subtropical regions, making bananas the most traded fruit in the world and the sixth most valuable food crop [12]. Despite their global significance, most bananas are consumed locally in the countries where they are produced, with only a small percentage exported to cooler regions. During storage and transportation, bananas are highly vulnerable to pathogens—particularly the fungus C. musae, which causes post-harvest losses of 30–40%. Without treatment, losses can rise to as much as 80% [7,13,14]. As a result, bananas often reach the market in poor quality.
Synthetic fungicides are the primary method used to control fungal diseases in fruit during the post-harvest period [15]. In the case of bananas, fungicides such as imazalil, thiophanate-methyl, and chlorothalonil are commonly applied to manage anthracnose and preserve the fruits’ freshness [7]. However, these compounds may offer efficacy, but they often lack important attributes such as selectivity for non-target organisms and rapidity of environmental manipulation [16], making them unsuitable for sustainable management. Furthermore, the short interval between treatment and consumption raises concerns about chemical residues in the fruit. Today consumer demand is shifting toward fresh and organic produce, especially in major banana markets. As a result, alternative approaches—physical, biological, and biotechnological—are being explored to prevent or treat anthracnose [17,18,19]. Among these, essential oils have emerged as promising agents against C. musae. [7,20,21]. This has fueled ongoing research into plant-derived molecules as natural disease control strategies. Any use of chemical or natural products for managing pests should prioritize those that present selectivity to beneficial organisms (pollinators, predators, or parasitoids). Beneficial organisms such as ladybug species, which are natural predators and act as naturally occurring biological control agents of different insect pests [22], and the fungus of the genus Trichoderma spp. that has been effective in controlling several phytopathogenic fungi, nematodes, and pests [23,24,25], represent relevant organisms to be evaluated in terms of the unintended effects of novel alternative pesticides.
In this context, essential oils have been shown as sustainable candidates for the management of diseases [26,27]. Particularly, the essential oil of M. citriflora has stood out as an antimicrobial [28], antifungal [29], antibacterial [30], and antiviral [31] with insecticide [32] properties, along with other promising biological activities in plants [33,34]. Furthermore, essential oils generally pose lower risks to non-target organisms, including humans [35,36,37,38]. M. citrifolia is a perennial, resilient plant well adapted to diverse environmental conditions, making the production of its by-products more accessible and sustainable. Therefore, the present study aimed to evaluate the fungistatic potential of M. citrifolia essential oil for the in vitro and in vivo control of C. musae isolated from ‘Nanica’ bananas. This was studied by combining molecular combinations in silico; in addition, toxicological bioassays were performed to investigate the safety of this essential oil for beneficial organisms such as the fungus T. asperellum and the predatory ladybugs E. connexa Germar and C. maculata DeGeer.

2. Materials and Methods

2.1. Isolation of Pathogen and Molecular Identification

The collection, cultivation, and isolation of the fungus followed established protocols [3]. Briefly, ‘Nanica’ bananas showing anthracnose symptoms were collected in Gurupi, Tocantins State, Brazil (11°43′4″ S, 49°04′07″ W). In the laboratory, the fruits were sanitized, placed in plastic trays, and maintained at 27 °C for seven days. After this period, symptomatic tissue was excised, cleaned, disinfected, and transferred to 9 cm Petri dishes containing potato dextrose agar (PDA), then incubated under a white light spectrum (40 W daylight fluorescent lamp, T10 OSRAM) at 27 °C ± 2 °C under a 12 h photoperiod. Fungi emerging from the lesions were subcultured on fresh PDA plates to obtain pure isolates. Preliminary identification to the genus level was conducted using optical microscopy and classical taxonomic literature [39]. The pathogen was obtained from the Phytopathology Laboratory of the Federal University of Tocantins (UFT, Gurupi, TO, Brazil), which was previously identified by morphological characteristics, as well as a molecular and phylogenetic analysis comparing it with other species, concluding in the identification of the species C. musae, work published in the journal Microbiology [3].

2.2. Extraction and Chemical Characterization of Essential Oil

We collected ripe noni fruits, M. citrifolia L., from trees located in the municipality of Gurupi (11°43′45″ S, 49°04′07″ W, Gurupi, TO, Brazil). The essential oil of M. citrifolia was extracted by an adapted hydrodistillation method [29]. Portions of 500 g of ripe fruit were placed in a two-liter round-bottomed flask containing 1000 mL of distilled water and then boiled for two hours in a Clevenger apparatus. After extraction, the essential oil was collected in the form of a supernatant, placed in an amber bottle, identified, and stored in a refrigerator at 4 °C ± 2 °C for analysis and use.
The chemical composition of M. citrifolia essential oil was analyzed using gas chromatography coupled with mass spectrometry (GC-MS) [27,29]. The analysis was conducted on a Shimadzu GC-210 chromatograph equipped with a QP2010 Plus mass detector, made in kyoto, Japan. A fused silica capillary column (RTX-5MS, 30 m × 0.25 mm × 0.25 μm film thickness) was used under the following conditions: oven temperature programmed from 60 °C to 240 °C at a rate of 3 °C/min; injector temperature at 220 °C; helium as the carrier gas; and splitless injection of a 1 μL sample (1:1000 dilution in hexane). The mass spectrometer operated at 70 eV, with the ion source and interface temperatures set at 200 °C. Compound identification was based on comparisons of mass spectra with those in the NIST and Wiley 229 library databases [40].

2.3. In Vitro Fungistatic Action of Morinda citrifolia Essential Oil on Colletotrichum musae

In vitro bioassays were performed in Petri dishes (90 mm in diameter) containing potato dextrose agar (PDA), evaluating five concentrations (10, 20, 30, 40, and 50 µL/mL), diluted in distilled and sterilized water and polysorbate Tween 80 (10 mg/mL) Dinâmica® (Química Contemporânea Ltd, Indaiatuba, São Paulo, Brazil), according to the adapted methodology [4]. An amount of 200 µL of each concentration was used and distributed evenly over the surface using a Drigalsky loop.
A 6 mm diameter PDA disc containing C. musae mycelium was then placed at the center of each plate. The plates were sealed, labeled, and incubated at 25 ± 2 °C under a 12 h photoperiod for ten days (as previously described). Each treatment included three replicates. Mycelial growth was measured at 48 h intervals across five evaluations, using a digital caliper to record the average diameter from two opposing points. Positive (10 µL/mL methyl thiophanate) and negative (untreated PDA) controls were included. The Mycelial Growth Velocity Index (MGVI) was calculated using the formula described by Oliveira [41],
MGVI = (D − Da)/N,
where D is the current average colony diameter, Da is the diameter from the previous measurement, and N is the number of days since inoculation.

2.4. Molecular Docking

2.4.1. Lingads

The compounds selected for molecular docking analysis were octanoic acid and acetate octan, the major constituents identified in Morinda citrifolia essential oil. The 3D structures of these compounds, in their neutral forms, were constructed using Marvin Sketch 18.10 ChemAxon (http://www.chemaxon.com, (accessed on 2 December 2024).

2.4.2. Target Modeling

Amino acid sequences of Tyrosine tRNA ligase from Colletotrichum sp. were retrieved from the UniProt database (http://uniprot.org). The 3D protein structures were generated using homology modeling with the Swiss Model Workspace (https://swissmodel.expasy.org/), following template selection via the BLASTp tool. Templates were downloaded from the Protein Data Bank (https://www.rcsb.org/), based on quality criteria including experimental method, resolution, R-value, and ligand complexation. The Swiss Model was also used to assess structural integrity and verify amino acid positioning in the active site [42]. The model’s validation was performed through an analysis of Ramachandran plots [43,44], which evaluate the distribution of backbone torsion angles (φ and ψ) to assess stereochemical quality. Additionally, the QMEAN score was used as an overall metric of the model’s quality [45].

2.4.3. Molecular Docking Calculations

Targets and ligands were prepared for molecular docking using AutoDock Tools 1.5.7 [46], following the methodology described by Moura, et al. [47]. Docking calculations were performed with AutoDock Vina [48], generating nine binding poses per ligand, along with their corresponding affinity energy values (kcal/mol). The docking results were analyzed using PyMOL 2.0 [49] and Discovery Studio 4.5 [50] to identify the best ligand binding position within each protein target [47].

2.5. Phytotoxicity of Morinda citrifolia Essential Oil to Banana Fruit

M. citrifolia essential oil solutions were prepared at concentrations of 10, 20, 30, 40, and 50 µL/mL using Tween-80 (10 mg/mL). Ripe and uniform fruits of the cultivar Nanica (Musa cavendishii) were purchased from a local supermarket, washed with the neutral detergent Limpol (Bombril S/A), rinsed with sterile water, and air-dried. Using a hand sprayer, three fruits per treatment were sprayed with 5 mL of each concentration. The positive control was sprayed with thiophanate methyl (10 µL/mL) and the negative control with distilled water. Then, they were stored in an incubation chamber at 25 ± 3 °C and an 85 ± 3% relative humidity. Phytotoxicity was evaluated at 24 and 48 h using the scale of Goes et al. [43], where 0 = no immunity to the consumer, 1 = fruit with mild symptoms (presence of light spots, without restriction to the consumer market), 2 = fruit with moderate symptoms (presence of small visible spots but accepted with restriction by the consumer market), 3 = fruit with severe symptoms (very dark spots, rejected by the consumer market).

2.6. Preventive and Curative Control of Banana Anthracnose Using Morinda citrifolia Essential Oil

Based on the results obtained in the phytotoxicity test, other concentrations were made, starting from the highest concentration that did not cause phytotoxicity after 48 h. Thus, for the preventive and curative tests, the concentrations (1.0, 2.5, 5.0, 7.5, and 10.0 µL/mL of essential oil) were evaluated. For preventive treatment, three banana fruits were sprayed with 5 mL of each concentration of essential oil [44]. The positive and negative controls were sprayed with thiophanate methyl (10 µL/mL) and distilled water, respectively. One hour after flowering, a 6 mm disc of C. musae mycelium was applied to each fruit. The inoculated fruits were placed in separate trays and stored in a humid chamber with an 85 ± 3% relative humidity and a temperature of 28 ± 2 °C for 24 h. After 24 h they were transferred to the incubation room at a temperature of 28 ± 2 °C for the disease’s development. In the curative treatment, bananas were first inoculated with the 6 mm mycelium discs and maintained in a humid chamber with an 85 ± 3% relative humidity and a temperature of 28 ± 2 °C for 12 h. Afterwards, the fruits were removed and sprayed with 5 mL of the essential oil solutions and kept in the incubation room at 28 ± 2 °C. To calculate the Area Under the Disease Progress Curve (AUDPC) for both treatments, evaluations were conducted every 48 h up to 10 days post-inoculation. Disease severity was assessed by measuring lesion diameters (mm) using a digital caliper. The AUDPC values were then calculated from these measurements using the equation proposed by Campbell and Madden [51].

2.7. Effect of Morinda citrifolia Essential Oil on Fruit Quality

Bananas from the preventive treatment were used to assess the effect of M. citrifolia essential oil on fruit quality. Fresh mass loss (FML) was calculated by comparing the initial and final weights (ten days post-inoculation) using a precision digital scale. On the final day of evaluation of disease progression, the Total Soluble Solids concentration (°Brix) in the fruit juice was measured using a manual Atago refractometer with a 0–32 °Brix scale.

2.8. Selectivity of Essential Oil to Microorganism Trichoderma asperellum and Ladybugs Coleomegilla maculata and Eriopis connexa

In the selectivity test for non-target organisms, a concentration of 30 µL/mL of M. citrifolia essential oil was three times higher than the highest concentration used in preventive and curative treatments was applied. The fungus T. asperellum was obtained from the mycotheque of the Phytopathology Laboratory of the Federal University of Toacantis (UFT, Gurupi, TO, Brazil); T. asperellum was initially cultured on potato dextrose agar (PDA). Two days after inoculation, the essential oil solution (30 µL/mL) was sprayed onto the medium’s surface. Distilled water and methyl thiophanate served as negative and positive controls, respectively. Mycelial growth was measured after ten days across all treatments.
The bioassay with ladybugs (C. maculata and E. connexa) was conducted following methods previously described elsewhere [52,53]. Briefly, five adult ladybugs of each species were placed in 9 cm Petri dish arenas lined with dry filter paper impregnated with essential oil (30 µL/mL). The dishes were maintained under laboratory conditions for 24 h. Six replicates were prepared per species, with a sterile water/DMSO (2%, v/v) solution serving as the negative control. After 24 h, insect survival was assessed; individuals unable to walk twice their body length when prodded with a fine hair brush were considered dead.

2.9. Statistical Analysis

The in vitro mycelial growth of Colletotrichum musae was analyzed using nonlinear regression, while preventive and curative treatments, as well as weight loss, were evaluated by linear regression using SigmaPlot 12.0. Total anthracnose severity after ten days and °Brix values were analyzed by one-way ANOVA (p < 0.05). Weight loss comparisons between the control and highest treatment doses were performed using a t-test (p < 0.05), also with SigmaPlot 12.0.

3. Results

3.1. Chemical Composition of Morinda citrifolia Essential Oil

The qualitative and quantitative analysis of M. citrifolia essential oil identified 13 major compounds. The predominant components were octanoic acid (caprylic acid) at 64.03%, butanoic acid (butyric acid) at 10.16%, hexanoic acid (caproic acid) at 8.64%, and methyl octanoate at 5.35% of the total composition. These compounds suggest potential antimicrobial activity of the noni essential oil (Table 1).

3.2. In Vitro Fungistatic Action of Morinda citrifolia Essential Oil on Colletotrichum musae

The mycelial growth of C. musae decreased as the concentration of M. citrifolia essential oil increased. At 50 µL/mL, the essential oil reduced mycelial growth by approximately 75%, nearly matching the effectiveness of the synthetic fungicide (Figure 1). Concentrations of 10 and 30 µL/mL also caused modest reductions in growth, as indicated by the distinct separation of the growth curves between treatments (Figure 1). The curve fitting parameters are provided in Table S1.

3.3. Molecular Docking

The target Tyrosine tRNA ligase of the organism Colletotrichum sp. is T0L8P8, found in the UNIPROT database. From the homology modeling, the 4OJM template was chosen, which presented an identity of 61.62%, and values of 97.46% and 0.78, for Ramachandran favored and QMEAN, respectively. The major compounds present in noni EO complexed with the receptors and formed various types of interactions with varying affinity energies, as indicated by the docking assays performed (Table 2). The octanoic acid ligand showed a better affinity energy with the target of the Colletotrichum sp., with a value of −4.8 kcal/mol. We also observed that acetate octanoate presented a good affinity energy, with a value of −4.7 kcal/mol.
The complex formed between the ligands and the target receptor Tyrosine tRNA ligase of the organism Colletotrichum sp. complex showed interactions between the target’s active site and octanoic acid (Figure 2A) of the following types: conventional hydrogen bond with GLY56 and SER215; alkyl interactions with LEU257, LEU258, and LYS265; and van der Walls interactions with LEU59, GLY214, ASP216, VAL255, PRO256, and PHE266. The ligand acetate octanoate presented interactions of the following types: conventional hydrogen bond with SER215; alkyl interactions with LEU59, LEU258, LYS265, PHE266, and LYS268; and van der Walls interactions with GLY56, GLY214, ASP216, LEU257, and PHE305 (Figure 2B,C).

3.4. Phytotoxicity of Morinda citrifolia Essential Oil to Banana Fruit

Toxicity to the ‘Nanica’ banana was concentration-dependent. Within the first 24 h, concentrations below 30 µL/mL showed no toxic effects; however, after 48 h, only doses of 10 µL/mL or lower remained non-toxic (Table 3).

3.5. Preventive and Curative Control of Banana Anthracnose Using Morinda citrifolia Essential Oil

The lesion diameter increased over time and was influenced by the concentration of essential oil (Figure 3A, Table S2). After ten days, all doses except the lowest reduced disease severity by about 30% during preventive treatment (df = 5; F = 12.35; p < 0.001) (Figure 3B). Similarly, in curative treatment—applied after C. musae’s colonization—severity depended on time and dose (Figure 3C, Table S2). However, only the highest dose achieved a modest 25% reduction in mycelial growth after ten days (df = 5; F = 8.78; p < 0.001) (Figure 3D). This indicates that M. citrifolia essential oil is less effective once fungal colonization has begun.
The Area Under the Disease Progress Curve (AUDPC) was significantly reduced in ‘Nanica’ bananas treated preventively with M. citrifolia essential oil, showing a concentration-dependent effect starting at 2.5 µL/mL (Figure 4A). In contrast, fruit already infected with Colletotrichum musae required concentrations above 7.5 µL/mL to achieve a noticeable reduction in AUDPC (Figure 4B).

3.6. Effect of Morinda citrifolia Essential Oil on the Fruit Quality

Weight loss in ‘Nanica’ bananas increased with higher doses of M. citrifolia essential oil, ranging from 7.95% at 1.0 µL/mL to 31.63% at 10.0 µL/mL (Figure 5A). However, there was no significant difference in weight loss between the highest dose and the control (t = 0.993; df = 4; p = 0.993). Total Soluble Solids (°Brix) also varied significantly across treatments (df = 5; F = 8.38; p < 0.001), with values ranging from 17.5 to 18.47 (Figure 5B).

3.7. Selectivity of Noni Essential Oil to Non-Target Organisms

After 10 days, the mycelial growth of T. asperellum showed no significant difference between plates treated with M. citrifolia essential oil and the negative control (Figure 6A). Likewise, survival rates of C. maculata and E. connexa were 100%, demonstrating the essential oil’s selectivity toward these beneficial organisms (Figure 6B).

4. Discussion

Our results demonstrate that M. citrifolia essential oil, rich in octanoic and butanoic acids, effectively inhibits C. musae, likely through interactions with the fungal Tyrosine tRNA ligase. Concentrations below 10 µL/mL showed no phytotoxic effects on banana fruits while reducing anthracnose both preventively and curatively. Although the treated bananas experienced some weight loss, the highest loss did not differ significantly from the control. Total Soluble Solids (°Brix) were also affected by treatment. Additionally, the essential oil exhibited selectivity, showing no harmful effects on the beneficial fungus Trichoderma asperellum or the predatory insects C. maculata and E. connexa.
M. citriflora essential oil effectively inhibited the mycelial growth of Colletotrichum musae, consistent with previous studies showing that essential oils can suppress fungi causing anthracnose in fruits [4,17,48,49], particularly species of the Colletotrichum genus [7,21,55,56,57,58]. The susceptibility of C. musae, the pathogen responsible for banana anthracnose, to these biomolecules has also been demonstrated [7,20,59]. The antifungal activity of M. citriflora oil is largely attributed to its main compound, octanoic acid, which has shown effectiveness against fungi such as Stagonosporopsis cucurbitacearum [29], Colletotrichum truncatum [60], and Colletotrichum gloeosporioides [61].
This study also confirmed that M. citrifolia essential oil is effective both in preventing Colletotrichum musae infestation and in treating infected banana fruit. The broad biological activity of M. citrifolia is likely linked to the chemical diversity of its compounds and possible synergistic interactions among them [28,56]. Its high content of carboxylic acids—particularly octanoic, butanoic, and hexanoic acids—confers strong antifungal properties [35], with octanoic acid alone accounting for over 80% of the oil’s composition [31,35,62]. Notably, isolated octanoic acid has demonstrated antifungal activity comparable to the full essential oil [29]. Stevens and Hofmeyr [63] reported that octanoic acid enters the cytoplasm of Saccharomyces cerevisiae, dissociates to release H+ ions, lowers cytoplasmic pH, and activates H+-ATPase, ultimately depleting cellular ATP and inhibiting growth. More broadly, volatile organic compounds (VOCs) like these acids can induce oxidative stress by triggering the accumulation of reactive oxygen species (ROS), thereby inhibiting fungal growth [64]. Essential oils are also known to penetrate fungal cell membranes, disrupt the metabolic pathways involved in cell wall synthesis, and impair spores’ germination and development [13,65]. In support of these mechanisms, our in silico analysis revealed that both octanoic and butanoic acids interact with Tyrosine tRNA ligase, an essential enzyme in C. musae. The antifungal and antibacterial potential of plant-derived molecules has previously been attributed to the inhibition of this enzyme [66,67], with recent studies confirming the affinity of these fatty acids for Tyrosine tRNA ligase [27,29,34]. This suggests that their incorporation into fungal lipid membranes may interfere with essential physiological processes at the cellular level.
At higher concentrations, M. citrifolia essential oil exhibited phytotoxicity in banana fruits, consistent with previous reports on both the susceptibility [20,68] and tolerance [69,70] of banana fruit to essential oils. However, lower concentrations—used preventively and curatively—significantly reduced the disease’s severity (as measured by AUDPC) without causing phytotoxic effects. This aligns with studies showing that doses below 10 µL/mL of M. citrifolia oil are non-toxic to rice and maize seeds [35,71]. This selective safety is notable, as essential oils—being plant secondary metabolites—often serve as natural defense compounds and are not universally safe for all plants. Indeed, phytotoxic effects of M. citrifolia oil at similar concentrations have been reported in maize (Zea mays) and melon (Cucumis melo) [29,62,72] Such effects are expected, as these compounds are typically stored in specialized plant structures to prevent self-toxicity [73,74,75].
Although noni essential oil reduced the Area Under the Anthracnose Progress Curve (AACPD), it did not prevent weight loss in banana fruit and also led to a reduction in Total Soluble Solids (°Brix). Weight loss following the application of essential oil is common and is often attributed to the formation of a semi-permeable protective barrier; however, opposite effects have also been reported [76]. The influence of essential oils on fruit quality—particularly °Brix—has been documented [77], though both increases and decreases in this parameter have been observed across different fruits and oil types [78,79,80]. An increase in °Brix is often linked to reduced water loss, resulting in a higher concentration of soluble solids. Conversely, a decrease may result from the metabolic conversion of sugars into other compounds, such as alcohols, during storage or ripening.
In addition, the essential oil of M. citrifolia did not affect the mycelial growth of T. asperellum or the survival of C. maculata and E. connexa ladybugs. This is exactly one of the characteristics of essential oils that make them promising as a source of molecules with fungal potential.

5. Conclusions

Collectively, our results demonstrated that the essential oil of M. citriflora appears to be a potential growth regulator for the fungus C. musae on Nanica banana fruit, without affecting their physical appearance; it could therefore be used as a preventative or curative measure against anthracnose. However, further studies on persistence and stability, as well as selectivity to human cells, should be carried out.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/microbiolres16070149/s1. Table S1: The summary of the nonlinear regression analysis is shown in Figure 1. Table S2: Summary of the linear regression analysis presented in Figure 3; preventive (A) and curative (C).

Author Contributions

Conceptualization, M.C.S., G.R.S., D.S.C.M. and P.R.F.; methodology, M.C.S., G.R.S., D.S.C.M., P.R.F. and R.C.C.; software, R.C.C., S.H.A., P.R.O., M.A.G., O.M.H., E.E.O., C.B.M., M.V.G. and W.S.M.; validation, G.R.S., R.W.S.A., M.P.C. and P.R.F.; investigation, M.C.S., P.R.F., R.C.C., M.A.G., O.M.H., D.S.C.M., P.R.O. and S.H.A.; formal analysis, L.O.V., G.R.S., W.S.M., R.W.S.A., A.S.R.C., C.B.M., M.V.G., M.P.C. and E.E.O.; resources, G.R.S., R.W.S.A., M.P.C., C.B.M., M.V.G. and E.E.O.; writing—original draft, M.C.S., L.O.V., G.R.S. and E.E.O.; writing—review and editing, L.O.V., G.R.S., M.P.C. and E.E.O.; visualization, L.O.V., E.E.O. and G.R.S.; supervision, G.R.S., D.S.C.M. and P.R.O.; project administration, G.R.S., M.P.C. and E.E.O.; funding acquisition, G.R.S. All authors have read and agreed to the published version of the manuscript.

Funding

The authors are highly thankful to the Coordination for the Improvement of Higher Education Personnel—Brazil (CAPES)—Finance Code 001, the National Council of Scientific and Technological Development (CNPq), the Tocantins State Foundation for Research Aid (FAPTO), and the Federal University of Tocantins (CTC-UFT/PROPESQ).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

When requested from the corresponding author, all data will be provided.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. In vitro mycelial growth (mm) of Colletotrichum musae over time exposed to varying concentrations of Morinda citrifolia essential oil. Symbols represent the mean of three replicates; vertical lines indicate the standard error.
Figure 1. In vitro mycelial growth (mm) of Colletotrichum musae over time exposed to varying concentrations of Morinda citrifolia essential oil. Symbols represent the mean of three replicates; vertical lines indicate the standard error.
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Figure 2. Octanoic acid (Red) and acetate octanoate (Green) complexed with Tyrosine tRNA ligase (A) and 2D maps of molecular interactions with amino acids in target’s active site (Yellow) with octanoic acid (B) and acetate octanoate (C) of Colletotrichum sp.
Figure 2. Octanoic acid (Red) and acetate octanoate (Green) complexed with Tyrosine tRNA ligase (A) and 2D maps of molecular interactions with amino acids in target’s active site (Yellow) with octanoic acid (B) and acetate octanoate (C) of Colletotrichum sp.
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Figure 3. Regression analysis of anthracnose severity in ‘Nanica’ banana’s area of infected tissue in (mm) over time (A,C) and after ten days (B,D) when treated with different doses of Morinda citrifolia essential oil as preventive (A,B) or curative (C,D) treatment; and lines covering the same bars (B,D) or the same level not do not differ according to the one-way ANOVA test (p < 0.005).
Figure 3. Regression analysis of anthracnose severity in ‘Nanica’ banana’s area of infected tissue in (mm) over time (A,C) and after ten days (B,D) when treated with different doses of Morinda citrifolia essential oil as preventive (A,B) or curative (C,D) treatment; and lines covering the same bars (B,D) or the same level not do not differ according to the one-way ANOVA test (p < 0.005).
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Figure 4. Area Under the Disease Progress Curve in ‘Nanica’ banana when exposed to different doses of Morinda citrifolia essential oil. Preventive (A) and curative (B) treatment.
Figure 4. Area Under the Disease Progress Curve in ‘Nanica’ banana when exposed to different doses of Morinda citrifolia essential oil. Preventive (A) and curative (B) treatment.
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Figure 5. Weight loss (A) and °Brix (B) of ‘Nanica’ bananas after ten days of preventive treatment with different doses of Morinda citrifolia essential oil. In (A), the horizontal line connecting the highest dose and the control indicates no significant difference by t-test (p > 0.05). In (B), bars sharing the same letter or level do not differ significantly according to one-way ANOVA (p < 0.005).
Figure 5. Weight loss (A) and °Brix (B) of ‘Nanica’ bananas after ten days of preventive treatment with different doses of Morinda citrifolia essential oil. In (A), the horizontal line connecting the highest dose and the control indicates no significant difference by t-test (p > 0.05). In (B), bars sharing the same letter or level do not differ significantly according to one-way ANOVA (p < 0.005).
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Figure 6. Growth of Trichoderma asperellum after 10 days when exposed to Morinda citrifolia essential oil (A) and survival of ladybugs Coleomegilla maculata and Eriopsis connexa after 24 h of exposure to 30 uL/mL of M. citrifolia (B).
Figure 6. Growth of Trichoderma asperellum after 10 days when exposed to Morinda citrifolia essential oil (A) and survival of ladybugs Coleomegilla maculata and Eriopsis connexa after 24 h of exposure to 30 uL/mL of M. citrifolia (B).
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Table 1. Chemical composition of Morinda citrifolia essential oil determined by GC/MS analysis.
Table 1. Chemical composition of Morinda citrifolia essential oil determined by GC/MS analysis.
CompoundsRTIR(%)
2-Heptanone5.0139270.14
Methyl hexanoate5.7879441.08
Ethyl hexanoate7.9969910.5
Hexanoic acid8.68810068.64
Butanoic acid, 4-pentenyl ester10.39510430.19
Methyl octanoate12.75510945.35
Octanoate acetate15.84611633.58
Octanoic acid17.413119864.03
4-Pentenyl hexanoate18.60212254,3
Decanoic acid, methyl ester21.39012880,19
Butanoic acid26.995142010.16
Hexyl octanoate31.93915400.37
Others--1.47
Total--100
TR = Retention time in minutes. IR = Calculated Kovats retention index, determined in relation to the retention time of a series of n-alkanes (C9–C24).
Table 2. Molecular docking results for complexes between major compounds of noni EO and target of Colletotrichum sp.
Table 2. Molecular docking results for complexes between major compounds of noni EO and target of Colletotrichum sp.
LigandAffinity Energy (kcal/mol)
Octanoic acid−4.8
Hexanoic acid−4.4
Butanoic acid−4.5
Methyl octanoate−4.4
Acetate octanoate−4.7
4-Pentene octanoate−4.5
Table 3. Phytotoxicity was observed after 24 and 48 h in banana ‘Nanica’ fruit when exposed to different doses of Morinda citrifolia essential oil.
Table 3. Phytotoxicity was observed after 24 and 48 h in banana ‘Nanica’ fruit when exposed to different doses of Morinda citrifolia essential oil.
Treatment (µL/mL)Scale (24 h)Observed PhytotoxicityScale (48 h)Observed Phytotoxicity
Control
(H2O + tween-80)		0Absent0Absent
Methyl tiophanate 100Absent0Absent
100Absent0Absent
200Absent1Light chlorosis
300Absent2Moderate chlorosis
401.67Light chlorosis3Severe chlorosis
501.67Light chlorosis3Severe chlorosis
Notes based on the scale proposed by [54].
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MDPI and ACS Style

Santos, M.C.; Viteri, L.O.; Fernandes, P.R.; Carvalho, R.C.; Gonzalez, M.A.; Herrera, O.M.; Osório, P.R.; Mourão, D.S.C.; Araujo, S.H.; Moraes, C.B.; et al. Morinda citrifolia Essential Oil in the Control of Banana Anthracnose: Impacts on Phytotoxicity, Preventive and Curative Effects and Fruit Quality. Microbiol. Res. 2025, 16, 149. https://doi.org/10.3390/microbiolres16070149

AMA Style

Santos MC, Viteri LO, Fernandes PR, Carvalho RC, Gonzalez MA, Herrera OM, Osório PR, Mourão DSC, Araujo SH, Moraes CB, et al. Morinda citrifolia Essential Oil in the Control of Banana Anthracnose: Impacts on Phytotoxicity, Preventive and Curative Effects and Fruit Quality. Microbiology Research. 2025; 16(7):149. https://doi.org/10.3390/microbiolres16070149

Chicago/Turabian Style

Santos, Maysa C., Luis O. Viteri, Paulo R. Fernandes, Rosilene C. Carvalho, Manuel A. Gonzalez, Osmany M. Herrera, Pedro R. Osório, Dalmarcia S. C. Mourão, Sabrina H. Araujo, Cristiano B. Moraes, and et al. 2025. "Morinda citrifolia Essential Oil in the Control of Banana Anthracnose: Impacts on Phytotoxicity, Preventive and Curative Effects and Fruit Quality" Microbiology Research 16, no. 7: 149. https://doi.org/10.3390/microbiolres16070149

APA Style

Santos, M. C., Viteri, L. O., Fernandes, P. R., Carvalho, R. C., Gonzalez, M. A., Herrera, O. M., Osório, P. R., Mourão, D. S. C., Araujo, S. H., Moraes, C. B., Giongo, M. V., Moura, W. S., Camara, M. P., Cangussu, A. S. R., Aguiar, R. W. S., Oliveira, E. E., & Santos, G. R. (2025). Morinda citrifolia Essential Oil in the Control of Banana Anthracnose: Impacts on Phytotoxicity, Preventive and Curative Effects and Fruit Quality. Microbiology Research, 16(7), 149. https://doi.org/10.3390/microbiolres16070149

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