Sanitary and physiological quality of Delonix regia seeds inoculated with Trichoderma asperellum and Saccharomyces cerevisiaeAbstract
Delonix regia is an ornamental tree species that is widely used in Brazilian landscaping. It is propagated by seeds, the quality of which can be compromised by the presence of pathogenic fungi. The use of biological agents, such as Trichoderma asperellum and Saccharomyces cerevisiae, in seed treatment has proven to be a promising alternative to the use of synthetic fungicides, promoting phytosanitary and physiological benefits to seeds. Therefore, the objective of this work was to determine the antagonistic effects of the agents T. asperellum and S. cerevisiae on the sanitary and physiological quality of D. regia seeds. The experimental design used was completely randomized, with treatments distributed in a 2 × 5 + 1 factorial scheme, with two biocontrol agents (T. asperellum and S. cerevisiae), five concentrations (0.0; 0.5; 1.0; 1.5 and 2.0 mL) + one additional treatment (seeds treated with fungicide). Seed health, germination, emergence and initial seedling development were evaluated. The fungi associated with the seeds were from the genera Fusarium sp., Cladosporium sp., Chaetomium sp., Aspergillus sp., Lasiodiplodia sp. and Rhizoctonia sp. The use of T. asperellum is effective in controlling Fusarium sp. in D. regia seeds, although its use may compromise physiological aspects of the seeds, such as germination and vigor. On the other hand, the application of S. cerevisiae was promising for reducing the incidence of microorganisms and, simultaneously, promoting improvements in the physiological quality of seeds. These findings indicate the potential of using biological agents, especially S. cerevisiae, as a sustainable strategy for the treatment of forest seeds.
Keywords:
biological control; flamboyant; forest seeds; fungi inhabiting the soil
Resumo
Delonix regia é uma espécie arbórea ornamental, amplamente utilizada no paisagismo brasileiro. Sua propagação é realizada por sementes, cuja qualidade pode ser comprometida pela presença de fungos patogênicos. A utilização de agentes biológicos no tratamento de sementes, como Trichoderma asperellum e Saccharomyces cerevisiae, tem se mostrado uma alternativa promissora ao uso de fungicidas sintéticos, promovendo benefícios fitossanitários e fisiológicos às sementes. Visto isso, o objetivo neste trabalho foi determinar a ação antagonista dos agentes T. asperellum e S. cerevisiae na qualidade sanitária e fisiológica de sementes de D. regia. O delineamento experimental utilizado foi inteiramente casualizado, com os tratamentos distribuídos em um esquema fatorial 2 × 5 + 1, com dois agentes de biocontrole (T. asperellum e S. cerevisiae), cinco concentrações (0,0; 0,5; 1,0; 1,5 e 2,0 mL) + um tratamento adicional (sementes tratadas com fungicida). Foi avaliada a sanidade das sementes, germinação, emergência e desenvolvimento inicial das plântulas. Os fungos associados às sementes foram dos gêneros Fusarium sp., Cladosporium sp., Chaetomium sp., Aspergillus sp., Lasiodiplodia sp. e, Rhizoctonia sp. estavam. O uso de T. asperellum é eficaz no controle de Fusarium sp. em sementes de D. regia, embora seu uso possa comprometer aspectos fisiológicos das sementes, como germinação e vigor. Por outro lado, a aplicação de S. cerevisiae revelou-se promissora ao reduzir a incidência de microrganismos e, simultaneamente, promover melhorias na qualidade fisiológica das sementes. Esses achados indicam o potencial do uso de agentes biológicos, em especial S. cerevisiae, como estratégia sustentável para o tratamento de sementes florestais.
Palavras-chave:
controle biológico; flamboyant; sementes florestais; fungos habitantes do solo
1. Introdução
Delonix regia (Hook.) Raf, belonging to the Fabaceae family, is a tree species native to India, Africa, Madagascar and northern Australia (Ammar et al., 2025). In Brazil, it stands out for its ornamental value and its exuberant flowering. In addition to these characteristics, it presents accelerated vegetative development and propagation exclusively by seeds (Solomon et al., 2025). This species has adapted satisfactorily to the soil and climate conditions of Northeast Brazil and is commonly used in the afforestation of parks, gardens and urban spaces (Silva et al., 2021).
D. regia seeds play important roles in the natural and artificial regeneration of this species, as well as in the dynamics of ecological succession (Costa et al., 2022). However, the efficiency of these processes is directly related to the viability and physiological quality of the seeds. Therefore, proper seed management becomes essential, both for preserving their reserves and for reducing the incidence of pathogenic fungi, which can compromise germination and favor the spread of diseases (Junges et al., 2016; Araújo et al., 2018).
The control of phytopathogens in seeds is commonly carried out via the use of synthetic chemical pesticides, which are widely used because of their ability to effectively preserve the physiological quality of seeds. However, the indiscriminate use of these inputs can have social and environmental impacts. Furthermore, although chemical seed treatment is widely regulated for agricultural crops, there are still no specific recommendations for forest species (Smiderle and Souza, 2021).
A reduction in the health of forest species seeds can affect their quality, since pathogens can cause their death through the release of enzymes and toxins before and during germination, as well as after seedling emergence (Parisi et al., 2019). Therefore, to obtain high-quality seeds, alternative methods that are viable and effective at controlling pathogenic fungi are essential.
In this context, seed microbiolization has proven to be efficient in controlling pathogens, with studies reporting a reduction in pathogen incidence and improvements in seed sanitary quality (Cruz et al., 2020, 2022). This technique consists of applying live microorganisms or their metabolites to protect seeds. Microbiolization thus emerges as a viable alternative to replace or reduce the use of synthetic pesticides, offering economic and environmental advantages (Sá et al., 2019; Silva and Araújo, 2025).
Among the microorganisms used in microbiolization, fungi of the genus Trichoderma and bacteria of the genus Bacillus stand out. Trichoderma spp. work as biocontrol agents in several pathological conditions (Junges et al., 2017) and are widely used in seed treatment to control pathogens, increasing plant growth and yield (Smiderle and Souza 2022). These microorganisms solubilize insoluble micronutrients in the soil and increase the absorption and translocation of minerals that are not widely available (Missio et al., 2016). During germination, its use favors seedling emergence, initial vigor and resistance to pathogen attack (Gomes et al., 2019).
In addition to fungi, yeasts such as Saccharomyces cerevisiae, which are traditionally used in the production of alcoholic beverages, ethanol and baking, have also demonstrated potential in controlling plant diseases. Several studies have demonstrated the potential of S. cerevisiae in controlling plant diseases, either by direct control of phytopathogenic microorganisms or by activating defense mechanisms in plants, especially those of the genus Saccharomyces (Heling et al., 2017; Souza, et al., 2019).
Therefore, the present study aimed to evaluate the antagonistic effects of the agents T. asperellum and S. cerevisiae on the sanitary and physiological qualities of D. regia seeds.
2. Materials and Methods
The experiment was carried out at the Phytopathology Laboratory of the Center for Agricultural Sciences, Campus II of the Federal University of Paraíba. The fruits of D. regia were collected from mother plants located in the city of Esperança, Paraíba, a tropical climate region with an altitude of 631 m and geographic coordinates of latitude 7° 1’ 37’’ S and 35° 51’ 34’’ W (IBGE, 2009).
After collection, the fruits were selected, and those with mechanical damage or signs of deterioration were discarded to compose a uniform batch. The intact fruits were subsequently manually opened to extract the seeds. The seeds were subsequently selected, excluding those with visible physical damage, for subsequent treatment.
2.1. Preparation and application of treatments
The seeds were disinfected in 1% sodium hypochlorite for three minutes and then washed three times with sterile distilled water (SWD). The treatments consisted of immersing the seeds in a 50 mL container in different solutions. During this period, the seeds were stirred with the aid of a glass rod to ensure homogeneous contact with the treatment.
The treatments used were as follows: T0-Control, treated with ADE; T1-Captana fungicide (240 g of the product for 100 kg of seeds); T2 to T5- Trichodel® (0.5, 1.0, 1.5 and 2.0 mL, respectively); and T6 to T9- S. cerevisiae (0.5, 1.0, 1.5 and 2.0 mL, respectively). All treatment volumes were diluted in 100 mL of SWD.
2.2. Sanity test
Sanitary quality was assessed via the incubation method on filter paper substrate (Blotter Test) in accordance with the rules for seed analysis (Brasil, 2009). The seeds were incubated in Petri dishes (90 × 15 mm) distributed in ten replicates of ten seeds for each treatment, containing a double layer of sterilized filter paper moistened with SWD, and kept for a period of seven days at a temperature of 25 ± 2 °C.
The identification of fungi associated with seeds was performed via stereoscopic and optical microscopy, and the results were compared with those of specialized methods in the literature (Seifert et al., 2011). The results are expressed as a percentage of infected seeds for each fungal genus identified.
2.3. Germination test
The germination test was conducted with four replicates of 25 seeds per treatment, which were distributed on two previously sterilized sheets of Germitest® paper, covered with a third and organized in the form of rolls, previously moistened with SWD, equivalent to 2.5 times the weight of the paper. The rolls were placed in transparent plastic bags to avoid water loss through evaporation.
The test was conducted in a BOD germination chamber regulated at a constant temperature of 25 °C, with a photoperiod of eight hours of light (Brasil, 2013). Evaluations were performed daily until the fifteenth day after sowing, and germinated seeds were considered those whose root system was at least 2 mm in length.
To evaluate physiological quality, the first germination count (FGC) was used to determine the percentage of normal seedlings on the seventh day after sowing; germination percentage (GP), with the seed that produced a primary root of at least 2 mm in length being considered germinated; germination speed index (GSI), determined by counting the number of germinated seeds divided by the number of days elapsed between sowing and germination, according to Maguire (1962); mean germination time (MGT), assessed together with the germination test and calculated via the Labouriau formula (Labouriau, 1983); length of the aerial part (LAG) and of the root system (LRS) using a ruler graduated in centimeters, with the results expressed in cm; dry matter mass of the aerial part (DMAP); and dry mass of the root system (DMRS).
The dry mass was obtained by drying in an oven at 80 °C for 24 hours, according to Nakagawa (1999). After drying, the aerial part and root system were weighed, and the results are expressed in g seedling-1.
2.4. Emergency test
The test was conducted in a greenhouse at an average temperature of 32 °C and a relative humidity of 60%, as measured by a digital thermohygrometer. Four replicates of 25 seeds per treatment were used, sown at a depth of 2 cm in polypropylene plastic trays, containing washed sand as a substrate, previously sterilized in an autoclave. The humidity of the sand substrate was maintained under manual irrigation, with daily watering being carried out according to the loss of humidity of the substrate.
The evaluations were carried out daily until the fifteenth day after sowing, and the number of normal seedlings that emerged at each evaluation was determined (Brasil, 2013). The same variables described in the germination test were considered.
2.5. Experimental design and statistical analysis
The experimental design used was completely randomized, with treatments distributed in a 2 × 5 + 1 factorial scheme, with two biocontrol agents (T. asperellum and S. cerevisiae), four concentrations (0.0; 0.5; 1.0; 1.5 and 2.0 mL) + one additional treatment (seeds treated with fungicide).
Statistical analysis was performed using the R statistical software (R Core Team, 2025) with the emmeans (Lenth and Piaskowski, 2025) and multcomp (Hothorn et al., 2025) packages. The results were subjected to analysis of variance and polynomial regression, with the linear and quadratic models tested. The means of each concentration were compared for each biological agent via the F test (p <0.05), and the treatments were compared with the fungicide via the Dunnett test (p <0.05). The health data were previously transformed into √x + 1.
3. Results
3.1. Effects of biocontrol agents on the health of D. regia seeds
The sanitary analysis of D. regia seeds (Table 1) revealed a high incidence of fungi belonging to the genera Fusarium sp. (42%), Cladosporium sp. (16%), Chaetomium sp. (13%), Aspergillus sp. (15%), Lasiodiplodia sp. (13%) and Rhizoctonia sp. (10%).
Incidence of fungi (%) associated with Delonix regia seeds treated with Trichoderma asperellum - TCH (107 conidia mL-1) and Saccharomyces cerevisiae - SCH (g L-1).
The highest incidence of Fusarium sp. and Cladosporium sp. was observed in the control treatment, without the application of biological agents. Compared with the seeds that did not receive the biological product, the seeds treated with S. cerevisiae (g L-1) at concentrations of 0.5 and 1.0 g L-1 resulted in decreases of 23.8% and 11.90%, respectively, in the incidence of Fusarium sp. (Table 1). Notably, seeds treated with 2.0 g/L S. cerevisiae did not contain Fusarium sp.
Treatments with T. asperellum showed high efficacy, eliminating 100% of the incidence of Fusarium sp., Cladosporium sp. and Chaetomium sp. even at the lowest concentration tested (0.5 g L−1). These results indicate that T. asperellum may be a promising alternative for seed sanitation aimed at large-scale seedling production.
A comparison of the two biocontrol agents (Table 1) revealed that T. asperellum manifested itself positively, reducing the incidence 100% from a dose of 0.5 g; however, with respect to S. cerevisiae, the total reduction in incidence occurred at a dose of 1.0 g L-1. In addition, the occurrence of Aspergillus sp. was observed in the seeds of D. regia, with the action of T. asperellum at the lowest dose (0.5 g L−1) being similar to that of funcigide (Captan®), revealing total elimination of the infection.
3.2. Effects of biocontrol agents on the physiological quality of D. regia seeds
3.2.1. Germination
Analyses of the first germination count, germination percentage and speed index (Figure 1a, 11c) revealed that increasing concentrations of S. cerevisiae significantly favored the physiological performance of the seeds. In addition to controlling fungi, S. cerevisiae may stimulate germination through the production of hormones or enzymes related to the initial metabolism of the seeds.
Average values of first germination count - FGC (A), germination percentage - GP (B) and germination speed index - GSI (C) of Delonix regia seeds treated with Trichoderma asperellum (TCH) and Saccharomyces cerevisiae (SCH). * = The mean differs statistically from the fungicide by Dunnett's test (p <0.05), and means with the same letter at the same concentration do not differ statistically from each other by the F test (p <0.05).
Although the different concentrations of T. asperellum used in the present research were efficient at controlling the incidence of phytopathogenic fungi, they did not efficiently affect the physiological quality of the seeds (Figure 1a, 1b, 1c), which was inferior to the use of S. cerevisiae.
The T. asperellum and S. cerevisiae treatments resulted in increases in the lengths of the shoots (Figure 2a) and roots of D. regia seedlings (Figure 2b), and the regression curves fit the increasing linear model. It is believed that this behavior was due to the decrease in the incidence of fungi associated with these seeds.
Average values of the length of the aerial part - LAP (A), root system length - RSL (B), dry matter mass of the aerial part - DMAP (C) and roots - DMR (D) of Delonix regia seedlings treated with Trichoderma asperellum (TCH) and Saccharomyces cerevisiae (SCH). * = The mean differs statistically from the fungicide by Dunnett's test (p <0.05), and means with the same letter at the same concentration do not differ statistically from each other by the F test (p <0.05).
When different doses of S. cerevisiae were applied, an increase in the accumulation of dry mass was observed, both in the aerial part (Figure 2C) and in the roots (Figure 2D), as the concentrations of the microorganisms increased, with higher values in relation to the use of Captan. On the other hand, the treatments with T. asperellum presented little variation in dry mass, with similar results between the control (concentration 0) and the highest concentration. When the two agents were compared, regardless of their concentration, the dry mass values were greater when S. cerevisiae was used.
3.2.2. Emergency
In terms of vigor, characterized by the first emergence count (Figure 3a) as a function of the different concentrations of S. cerevisiae, the trend of the data was quadratic, reaching a maximum value at a dose of 1.0 g L−1. At the different concentrations of T. asperellum, there were no differences in the values of the first count, with the values being similar to those of the treatment with the fungicide.
First emergence count - FEC (A), emergence - E (B) and emergence speed index - ESI (C) of Delonix regia seedlings from seeds treated with Trichoderma asperellum (TCH) and Saccharomyces cerevisiae (SCH). * = The mean differs statistically from the fungicide by Dunnett's test (p <0.05), and means with the same letter at the same concentration do not differ statistically from each other by the F test (p <0.05).
For the percentage of emergence (Figure 3B), the data related to the different doses of S. cerevisiae fit the increasing linear equation, in which at a dose of 2.0 g L-1, the value obtained was 84%, a value close to that obtained with the fungicide, which was 91%. The data from the treatments with T. asperellum also fit an increasing linear equation, although the values obtained were lower than those obtained with S. cerevisiae.
With respect to the emergence speed index (Figure 3C), the use of S. cerevisiae provided greater value than did the T. asperellum treatment. Although it was efficient at controlling the pathogenic fungi present in D. regia seeds, T. asperellum had a negative effect on characteristics related to emergence.
In terms of the length of the aerial part (Figure 4A), the highest dose of S. cerevisiae resulted in an increase in the number of seedlings from the seeds treated with the fungicide. In the T. asperellum treatments, the greatest length of the aerial parts was associated with the 0.0 and 1.0 g L-1 concentrations.
Lengths of the aerial parts - LAP (A) and root system length - RSL (B) of Delonix regia seedlings originating from seeds treated with Trichoderma asperellum (TCH) and Saccharomyces cerevisiae (SCH). * = The mean differs statistically from the fungicide by Dunnett's test (p <0.05), and means with the same letter at the same concentration do not differ statistically from each other by the F test (p <0.05).
The seedling root length data as a function of S. cerevisiae treatment were adjusted to a linear model, in which the average length varied from 10.09 cm to 12.92 cm between doses of 0.0 and 2.0 g L-1. The use of different concentrations of T. asperellum resulted in a gradual reduction in root growth as the concentration increased, since at the lowest concentration, the root length obtained was 10.09 cm, whereas at the highest concentration, the average length was 7.05 cm (Figure 4B).
For the data on dry matter mass of the aerial part and roots of D. regia seedlings, there was no significant effect, that is, the different doses and different biological agents did not influence this variable.
4. Discussion
The seeds of D. regia presented a high incidence of fungi of the genera Fusarium sp. (42%), Cladosporium sp. (16%), Chaetomium sp. (13%), Aspergillus sp. (15%), Lasiodiplodia sp. (13%) and Rhizoctonia sp. (10%). The composition of the observed mycoflora is similar to that reported by Medeiros et al. (2012) in seeds of Caesalpinia pulcherrima, which belong to the same botanical family, suggesting a certain degree of fungal specificity associated with forest legumes.
Among the pathogens detected, Fusarium spp. stand out as the most common. According to Rosário et al. (2022), this genus is one of the main genera associated with the seeds of forest species and can affect germination, in addition to causing root rot and seedling damping. This pathogen has been described in association with seeds of at least 100 forest species (Santos and Rego, 2011). Quevedo et al. (2020), when studying Luehea divaricate seeds, demonstrated the vertical transmission of Fusarium spp., with symptoms of necrosis in the root system and subsequent death of seedlings, reinforcing its destructive potential in the forest regeneration process.
Another economically important field fungus is Rhizoctonia sp., whose infection typically occurs during seed maturation in the field. This species is associated with damping-off or seedling knocking-off, and can cause significant losses in forest species nurseries.
The genera Aspergillus spp. and Lasiodiplodia spp. are considered “storage fungi”, and when exposed to favorable environmental conditions, such as high humidity and temperature, these microorganisms can produce mycotoxins that compromise seed vigor and germination (Prestes et al., 2019). The presence of these fungi in D. regia reinforces the need for good storage practices and monitoring of sanitary quality throughout the processing process.
In this sense, the sanitary characteristics of forest seeds represent an important factor in seed germination, since losses resulting from deterioration, lesions and abnormalities in seedlings are reported, with fungi being the main organisms responsible for these losses (Santos et al., 2020).
As a promising alternative for the management of these pathogens, the application of biological control agents has proven effective. Studies with Trichoderma spp. indicate its potential for reducing the incidence of pathogenic fungi and promoting plant growth. Carvalho et al. (2011) reported that T. harzianum efficiently reduced the incidence of F. oxysporum f. sp. phaseoli in common bean seeds by 35-51%. Similarly, Junges et al. (2016), when evaluating the effects of Trichoderma spp. in controlling fungi associated with the seeds of three forest species (Penicillium sp., Fusarium sp., and Chaetomium sp.), reported greater efficiency in controlling these fungi in canafístula seeds (Pelthophorum dubium). Farias et al. (2019) tested T. asperellum in cotton seeds (Gossypium sp.) and reported that its use reduced the incidence and severity of F. oxysporum f. sp. vasinfectum (Fov).
In addition to their antagonistic potential, T. asperellum isolates are associated with the promotion of seedling growth. Rabuske et al. (2023) demonstrated that T. asperellum culture filtrates stimulated the initial development of pecan seedlings, in addition to reducing the incidence and severity of stem canker. Similar results were obtained by Santos et al. (2020) and Griebeler et al. (2021), who reported an increase in the root system biomass of Handroanthus serratifolius, Cordia trichotoma and Cedrela fissilis seedlings after treatment with T. asperellum. These beneficial effects on initial growth can increase seedling survival in the field, resulting in greater success in the implementation of forest stands.
Additionally, the yeast S. cerevisiae also has positive physiological effects, which can be attributed to its ability to synthesize indole-3-acetic acid (IAA), a phytohormone involved in stimulating root development and shoot biomass accumulation (Contreras-Cornejo et al., 2022). This effect is related to the regulation of H+-ATPase activity in the plasma membrane of root cells, as described by Olivares et al. (2017), which favors the initial growth of seedlings and more efficient establishment of seedlings in the field.
A relevant aspect of the present study refers to the dose-response behavior of the biological agents evaluated. The linear adjustment observed for sanitary and physiological quality indicates that, within the tested dose range, the effects of T. asperellum and S. cerevisiae were proportional to the increase in the applied dose. This pattern suggests that the maximum potential for biological response from the microorganisms may not have been fully reached, since no saturation point or stabilization of the responses was observed.
From an applied perspective, the predominance of linear responses indicates that the doses evaluated likely represent an initial response range, reinforcing the need for further studies with higher doses and wider gradients. Thus, the results obtained in this study highlight the importance of phytosanitary diagnosis in seeds of forest species, such as D. regia, not only to prevent loss of physiological quality, but also as a basis for the adoption of sustainable management strategies. Although the application of T. asperellum and S. cerevisiae has shown promising potential in controlling phytopathogens and improving seed quality, defining ideal doses depends on future research that will allow maximizing biological efficiency without compromising the physiological performance of the seeds.
5. Conclusions
The use of T. asperellum is effective in controlling Fusarium sp. in D. regia seeds, although its use may compromise physiological aspects of the seeds, such as germination and vigor. On the other hand, the application of S. cerevisiae was promising for reducing the incidence of microorganisms and, simultaneously, promoting improvements in the physiological quality of the seeds.
The predominance of linear responses as a function of the evaluated doses indicates that the tested levels may not have reached the maximum response potential of the biological agents. Thus, future studies evaluating higher doses and wider ranges are needed to define optimal concentrations, contributing to the more efficient and safe use of seed microbiolization of forest species.
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:
Jairo Lizandro Schmitt
