Essential oils in the seed quality of Myracrodruon urundeuva Fr. AllemãoAbstract
Aroeira-do-sertão is a native species of the Brazilian semiarid region, with pharmacological properties and potential for reforestation of degraded areas, making the use of healthy seeds essential for seedling production. In the absence of registered chemical products for the treatment of native forest seeds, the development of effective alternative methods for pathogen control, such as the use of essential oils, becomes indispensable. The present study aimed to evaluate the effectiveness of essential oils in reducing pathogen incidence and improving the physiological quality of aroeira seeds. The experiment was conducted at the Phytopathology Laboratory of the Federal University of Paraíba. Seeds of aroeira obtained from the Seed Network of the São Francisco Integration Project were used, and their moisture content was determined. Seeds were treated with essential oils at a concentration of 1%. Treatments consisted of T1: Control; T2: Captan; T3: Cinnamon leaf; T4: Ginger; T5: Palmarosa; T6: Thyme; T7: Sicilian lemon; T8: Peppermint; T9: Oregano; and T10: Tangerine. The seeds were subjected to a health test, using ten replications of ten seeds, and to germination and emergence tests, with four replications of 25 seeds. The experiment was carried out in a completely randomized design. Cinnamon leaf and thyme essential oils reduced the incidence of the fungal genera Aspergillus, Cladosporium, Alternaria, Ulocladium, Pestalotia, and Curvularia in M. urundeuva seeds and did not affect seed physiological quality.
Keywords:
aroeira-do-sertão; alternative control; seed pathology; forest species
Resumo
A aroeira-do-sertão é uma espécie nativa do semiárido brasileiro, com propriedades farmacológicas e potencial para o reflorestamento de áreas degradadas, sendo essencial o uso de sementes sadias para a produção de mudas. Diante da ausência de produtos químicos registrados para o tratamento de sementes florestais nativas, torna-se essencial o desenvolvimento de métodos alternativos eficazes para o controle de patógenos, como o uso de óleos essenciais. O presente trabalho teve como objetivo verificar a eficiência dos óleos essenciais sobre a incidência de patógenos e na qualidade fisiológica de sementes de aroeira. O experimento foi conduzido no Laboratório de Fitopatologia, pertencente a Universidade Federal da Paraíba. Foram utilizadas sementes de aroeira oriundas da Rede de Sementes do Projeto de Integração do São Francisco, e determinado teor de água. As sementes foram tratadas como óleos essenciais na concentração de 1%. Os tratamentos foram T1: Testemunha; T2: Captana, T3: Canela folha; T4: Gengibre; T5: Palmarosa; T6: Tomilho; T7: Limão siciliano; T8: Menta Piperita; T9: Orégano e T10: Tangerina. As sementes foram submetidas ao teste de sanidade, em que foram utilizadas dez repetições com 10 sementes e germinação e emergência, com quatro repetições de 25 sementes. O experimento foi realizado em delineamento inteiramente casualizado. Os óleos essenciais de canela de folhas e tomilho reduziram a incidência dos gêneros fúngicos Aspergillus, Cladosporium, Alternaria, Ulocladium, Pestalotia e Curcularia. em sementes de M. urundeuva e não interferiram na qualidade fisiológica das sementes.
Palavras-chave:
aroeira-do-sertão; controle alternativo; patologia de sementes; espécie florestal
1. Introduction
Myracrodruon urundeuva Fr. Allemão is a forest species found in Northeastern Brazil and belongs to the family Anacardiaceae, commonly known as aroeira-do-sertão (Higa et al., 2019). In traditional knowledge, the periderm exhibits pharmacological, anti-inflammatory, and dermatological properties (Sousa et al., 2020). According to Alves et al. (2020), the indiscriminate use of aroeira results from the significant socioeconomic role of this species. Such overexploitation is jeopardizing not only the preservation of aroeira itself but also the biodiversity of the Caatinga biome (Capo et al., 2022).
Therefore, it is crucial to adopt effective conservation and sustainable management strategies to ensure the continuity and protection of this species, which faces a high risk of extinction (Nascimento et al., 2024). Since this species is mainly propagated through seeds, factors such as the physiological and sanitary quality of M. urundeuva seeds intended for propagation and plant establishment must be carefully considered (Santos et al., 2019). Seeds are effective vehicles for the transmission of phytopathogens, such as M. urundeuva (Silva et al., 2025) and Bixa orellana (Nunes et al., 2025), which may compromise seed germination, seedling emergence, and field growth, thus making the use of healthy seeds essential to prevent or reduce pathogen dissemination (Rosário et al., 2022). For forest seeds, maintaining adequate sanitary and physiological quality is a major challenge (Silva et al., 2021).
The management of seeds associated with phytopathogens by means of treatments that are practical, accessible, low-cost, and highly effective in pathogen control is therefore essential for forest seed production (Costa et al., 2022). However, no chemical products are officially registered for the treatment of forest seeds (Parisi et al., 2019), which underscores the importance of employing chemical, physical, and biological methods to preserve seed quality. In this context, alternative natural fungicides represent a promising strategy. Several studies have already demonstrated the effectiveness of essential oils in inhibiting and suppressing the development of various pathogens (Cutrim et al., 2019; Queiroz et al., 2020; Olinto et al., 2023).
This is explained by the fact that essential oils, secondary metabolites of plants, present desirable characteristics such as low toxicity to humans and higher volatility when compared to synthetic chemical products. They also constitute an alternative to the resistance that may develop in organisms exposed to synthetic chemicals (Alonso-Gato et al., 2021).
Thus, the present study aimed to evaluate the effectiveness of essential oils in reducing pathogen incidence and maintaining the physiological quality of M. urundeuva seeds.
2. Material and Methods
2.1. Experimental site and plant material
The experiment was conducted at the Seed Analysis Laboratory (LAS) and the Phytopathology Laboratory (LAFIT) of the Center for Agricultural Sciences, Federal University of Paraíba, Campus II, Areia, PB, Brazil. Traditional M. urundeuva seeds were obtained from the Seed Network of the São Francisco Integration Project, in Petrolina, PE. Seed moisture content was determined using the oven-drying method at 105 ± 3 °C for 24 h, with four replications of 25 seeds (Brasil, 2009). Seeds were surface-sterilized with 1% sodium hypochlorite for 3 minutes.
2.2. Control of pathogens associated with aroeira-do-sertão seeds using essential oils
Seeds were treated with commercial formulations of essential oils (EOs) at a concentration of 1 mL L−1 in 100 mL of sterile distilled water (SDW), with the addition of Tween 80 at 0.05% (v/v) to facilitate emulsification (Brito et al., 2010). The treatments were: T1: Control (SDW); T2: Captan fungicide (240 g 100 kg−1); T3: Cinnamon leaf (Cinnamomum verum J. Presl); T4: Ginger (Zingiber officinale Roscoe); T5: Palmarosa (Cymbopogon martinii (Roxb.) Will. Watson); T6: Thyme (Thymus vulgaris L.); T7: Sicilian lemon (Citrus limon (L.) Osbeck); T8: Peppermint (Mentha piperita L.); T9: Oregano (Origanum vulgare L.); and T10: Tangerine (Citrus reticulata Blanco) (Table 1). Captan treatment was calculated based on seed weight.
The chromatographic analyses of the essential oils had different origins. At the Department of Chemical Engineering and Food Engineering - Technological Center of the Federal University of Santa Catarina, the oils of Cinnamomum verum, Zingiber officinale, Cymbopogon martinii, Citrus limon and Origanum vulgare were analyzed. At the Department of Chemistry, Institute of Exact Sciences, Federal University of Minas Gerais, only the oil of Mentha piperita was quantified. The remaining oils, Thymus vulgaris and Citrus reticulata, were measured at the Multipurpose Laboratory for Characterization and Analysis, Institute for Research in Pharmaceuticals and Medications at the Federal University of Paraíba. The results they are shown in Table 1.
The seed health test was performed with 100 seeds per treatment, divided into 10 replications of 10 seeds, immersed for 5 minutes in the respective treatments. Fungicide application occurred through direct contact with the seed coat. Treated seeds were placed in 9-cm Petri dishes containing a double layer of sterile filter paper moistened with SDW and incubated at 25 ± 2 °C with a 12 h photoperiod for seven days (Brasil, 2009).
Fungal identification was performed under an optical microscope by comparing morphological structures with descriptions from specialized literature (Seifert et al., 2011). Results were expressed as the percentage of infected seeds.
2.3. Evaluation of seed physiological quality
2.3.1. Germination test
The same treatments were used for physiological quality evaluation. The germination test was conducted with 100 seeds, subdivided into four replications of 25 seeds. Seeds were distributed on two sheets of Germitest® paper, covered with a third sheet, and rolled, moistened with SDW equivalent to 2.5 times the paper dry weight. Rolls were placed in transparent polyethylene bags to prevent water loss and incubated in a BOD-type germination chamber at 25 ± 2 °C with a 12 h photoperiod. Germination was assessed at 24 h intervals for 10 days, recording normal seedlings, hard seeds, and dead seeds.
The first count was performed simultaneously with the germination test, recording seeds germinated on the second day after sowing (Brasil, 2013). Germination percentage (GP) was calculated as GP = (N2/N1) × 100, where N2 = number of germinated seedlings and N1 = number of seeds sown. The germination speed index (GSI) was calculated based on the daily record of germinated seedlings (Maguire, 1962).
2.3.2. Emergence test
The emergence test was carried out in a greenhouse at the Seed Analysis Laboratory (CCA). A total of 100 seeds per treatment, subdivided into four replications of 25 seeds, were sown in polystyrene trays containing sterile substrate and irrigated daily. Emerged seedlings were counted every 24 h. The first count was performed on the third day after sowing (Brasil, 2009). The percentage of emerged seedlings (PE) was determined, and the emergence speed index (ESI) was calculated based on daily records until stabilization on the 14th day, according to Maguire (1962).
2.3.3. Root and shoot length and dry matter
At the end of germination and emergence tests, seedling root and shoot lengths were measured with a millimeter ruler, and results were expressed in centimeters. Plant material was dried in an oven at 65 °C for 48 h and weighed on an analytical balance (0.0001 g precision). Results were expressed in g seedling−1.
2.4. Statistical analysis
The experiment was conducted in a completely randomized design. Data were subjected to Shapiro-Wilk normality and Bartlett’s homogeneity of variance tests. Means were grouped using the Scott-Knott test at 5% probability, with the aid of R® statistical software (R Core Team, 2023). Fungal incidence data were previously transformed using (√y + 1).
3. Results
The mycoflora identified in Myracrodruon urundeuva seeds were: Penicillium sp., Aspergillus spp., Cladosporium sp., Fusarium sp., Alternaria sp., Ulocladium sp., Pestalotia sp., Curvularia sp., and Chaetomium sp. No significant differences were observed among the control, fungicide treatment, and the essential oil treatments for Penicillium sp., Fusarium sp., and Chaetomium sp. (Figure 1).
Incidence of fungi in Myracrodruon urundeuva seeds treated with essential oils. T1: Control (sterile distilled water – SDW), T2: Captan fungicide (240 g 100 kg−1 seeds), T3: Cinnamon leaf, T4: Ginger, T5: Palmarosa, T6: Thyme, T7: Sicilian lemon, T8: Peppermint, T9: Oregano, T10: Tangerine, at a concentration of 1%. Means followed by the same letter on the line do not differ according to the Scott-Knott test (p ≤ 0.05). Data transformed as √y+1.
Among the essential oils, cinnamon leaf and thyme showed performance comparable to the fungicide, controlling the development of Cladosporium sp., Alternaria sp., Ulocladium sp., Pestalotia sp., Curvularia sp., and Aspergillus sp. These treatments reduced the incidence of Aspergillus sp. by 100%. Treatments with palmarosa and peppermint oils also reduced Aspergillus sp. incidence, differing from the control and other treatments.
The efficacy of oregano and ginger oils was observed against Alternaria sp., Curvularia sp., and Ulocladium sp. However, no control was observed for Aspergillus sp. and Pestalotia sp. compared with the control, indicating that these oils were ineffective under the tested conditions. For Cladosporium sp., distinct effects were noted: oregano oil reduced incidence, while ginger oil favored fungal development.
Lemon and tangerine oils were ineffective in reducing the incidence of Aspergillus sp., Alternaria sp., and Cladosporium sp. For Cladosporium sp., tangerine oil stimulated fungal growth, and for Pestalotia sp., it was also ineffective, whereas lemon oil showed lower incidence than the control. Both oils were effective only against Curvularia sp. and Ulocladium sp.
The seeds of M. urundeuva had a moisture content of 14.8%. In the germination test, essential oils did not influence seed performance; ginger, Sicilian lemon, and peppermint treatments presented results similar to the control and fungicide, whereas the remaining oils yielded germination rates below 70% (Table 1).
Regarding dead seeds, thyme (39%), cinnamon leaf (38%), and palmarosa (37%) oils showed the highest percentages (Table 2). The germination speed index did not differ significantly among essential oil treatments and the control.
Physiological quality of Myracrodruon urundeuva seeds treated with essential oils: germination (GE), dead seeds (DS), germination speed index (GSI), shoot length (SL), root length (RL), shoot dry matter (SDM), and root dry matter (RDM).
For biometric parameters, tangerine oil promoted greater shoot and root lengths compared to the control, fungicide, and other treatments. Peppermint and thyme oils showed shoot lengths superior to the fungicide and similar to the control, while for root length, both were equivalent to the control and fungicide.
In terms of shoot dry matter, no significant differences were observed among treatments and the control. However, root dry matter was highest for tangerine (0.0081 g seedling−1) and peppermint oils (0.0071 g seedling−1). Thyme, Sicilian lemon, and oregano oils also showed higher root dry matter compared to the control and fungicide (Table 2).
Seedling emergence showed behavior similar to germination, with essential oils not outperforming the control but differing from the fungicide. Emergence percentages for palmarosa (68%), thyme (66%), Sicilian lemon (68%), and tangerine (77%) were comparable to the control (72%), whereas fungicide, cinnamon leaf, ginger, peppermint, and oregano treatments showed lower percentages (60%, 54%, 62%, 60%, and 60%, respectively).
Significant differences were found for the emergence speed index. The control, palmarosa, and tangerine oils showed faster emergence compared to the fungicide and the other essential oils.
For growth, shoot length was higher in seeds treated with cinnamon leaf, thyme, peppermint, tangerine, and the control. However, for root length, only cinnamon leaf and tangerine oils performed similarly to the control, being superior to the fungicide and other oils.
Regarding shoot dry matter, the control, palmarosa, peppermint, and tangerine oils had higher values than the fungicide and other oils. In contrast, ginger (0.0108 g), palmarosa (0.0085 g), and oregano (0.0085 g) oils resulted in the lowest root dry matter values compared to other treatments (Table 3).
Physiological quality of Myracrodruon urundeuva seeds treated with essential oils: seedling emergence (EM), emergence speed index (ESI), shoot length (SL), root length (RL), shoot dry matter (SDM), and root dry matter (RDM).
4. Discussion
In recent years, there has been a growing awareness of the need to replace chemical fungicides in plant disease control. Thus, natural products such as essential oils (EOs) may represent a promising alternative (Xiao et al., 2021). EOs are secondary plant metabolites with antifungal potential due to their composition, which is rich in secondary compounds such as acids, alcohols, aldehydes, phenols, ketones, terpenes, aromatic phenols, and terpenoids (Sil et al., 2020).
The proportions of these compounds vary depending on the type of oil, as antifungal efficacy depends on both the combination of these components and their concentration. Some compounds may effectively inhibit certain fungal species, while others may not exhibit the same effect (Davari and Ezazi, 2022). In the present study, cinnamon leaf and thyme oils inhibited the growth of Aspergillus sp., while peppermint and palmarosa reduced incidence, and ginger, lemon, oregano, and tangerine did not differ from the control (Figure 1).
The genus Aspergillus sp. is frequently associated with M. urundeuva seeds (Nascimento et al., 2020), with reported incidences as high as 92%, a value greater than that found in this study (78%) (Souza et al., 2021). This high incidence is likely due to harvesting and storage processes, during which pathogens can penetrate seeds, reducing germination or even causing deterioration (Bala, 2017).
Fungi such as Penicillium sp. and Aspergillus sp. are reported to significantly reduce seed germination and seedling growth (Mangwende et al., 2021; Kaya and Karaman, 2023). The genus Fusarium includes species that are important plant pathogens, causing considerable losses in various crops both in Brazil and worldwide. Beyond reducing the quality of seeds and grains, which may become infected, the presence of mycotoxins produced by these species poses a significant risk to food safety and human and animal health (Lu et al., 2021).
The antifungal efficacy of cinnamon leaf oil (eugenol) and thyme oil (thymol) is likely determined by their chemical composition, particularly against Aspergillus sp. (Mutlu-Ingok et al., 2020). The effectiveness of these compounds was observed in this study, where both essential oils reduced the incidence of this genus by 100% (Figure 1). Eugenol, the major constituent of cinnamon leaf oil (71.55%), may act in multiple ways against fungi, including damaging hyphal morphology, disrupting cell membranes, reducing cytoplasmic density, inhibiting mycelial growth, and affecting mitochondria (Zhao et al., 2021). Thymol disrupts the cell wall and plasma membrane of pathogenic fungi, leading to cytoplasmic leakage.
Peppermint oil has also been shown to disrupt fungal cell walls and damage the ultrastructure of hyphae and conidia (Chang et al., 2022). In Caesalpinia ferrea (pau-ferro) seeds treated with thyme oil, a reduction in Aspergillus sp. incidence was also reported, attributed to the presence of carvacrol, which has antimicrobial activity against various microorganisms (Silva et al., 2021).
Compounds such as carvone, present in thyme and peppermint oils, display stronger antifungal activity than compounds such as limonene, found in citrus oils (lemon and tangerine), with inhibition rates of 41–100% against different pathogens (Parikh et al., 2021).
Studies have also demonstrated the biostimulant effects of EOs on seed germination, root development, and shoot growth, whether applied to seeds, leaves, or soil. These effects result from antioxidant compounds that protect tissues from oxidative stress and prevent auxin degradation in plants (Costa et al., 2024; Souri and Bakhtiarizade, 2019; Terzić et al., 2023). In this way, EOs help reduce stress and balance hormonal activity, thereby improving physiological quality (Oğuz et al., 2023). This study observed allelopathic effects of some essential oils, which reduced germination when compared to the control, possibly due to the concentration used (Table 2).
The biostimulant effect of EOs depends on both the type and concentration of oil, as some compounds may also exhibit allelopathic effects, inhibiting germination and plant development by interfering with cell division, membrane permeability, and enzyme activation (Verdeguer et al., 2020; Olinto et al., 2023). This was observed in tomato (Solanum lycopersicum) seeds treated with four concentrations of Alpinia zerumbet oil, where germination decreased as doses increased, reaching only 2% germination at 1% concentration (Almeida et al., 2019).
When assessing the phytotoxic effects of Cymbopogon citratus (lemongrass), Cymbopogon winterianus (citronella), and Lavandula angustifolia (lavender) oils on two species, lettuce (Lactuca sativa) and annoni grass (Eragrostis plana), Pawlowski et al. (2021) found that increasing oil concentrations reduced seed germination in both species and inhibited both shoot and root growth. Nascimento et al. (2021) also demonstrated that EO concentration and/or immersion time affected seed quality, with immersion periods longer than one hour negatively impacting seed vigor. This behavior was observed in the emergence test, where there was a reduction in the percentage when the seeds were treated with essential oils Cinnamon leaf, Ginger, Peppermint, and Oregano (Table 3).
5. Conclusion
Cinnamon leaf and thyme essential oils reduced the incidence of the fungal genera Aspergillus, Cladosporium, Alternaria, Ulocladium, Pestalotia, and Curvularia in M. urundeuva seeds and did not interfere with seed physiological quality.
Acknowledgements
To the Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPq and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES for the research grant awarded for this study.
Data Availability Statement
The entire data set that supports the results of this study was published in the article itself.
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Editor:
Takako Matsumura Tundisi
