Inhibition of germination by gallic acid in Eugenia brasiliensis Lam. seedsABSTRACT:
Seeds of monoembryonic Eugenia L. species have the ability to regenerate seedlings from cotyledon tissues when fractionated; however, only one seedling develops in each fraction, indicating the possibility of a self-control process during germination, preventing simultaneous germination. One of the likely compounds involved in this control is gallic acid, whose concentration increases in the cotyledons when germination begins. In this study, the possible action of this compound was evaluated by pre-soaking of Eugenia brasiliensis Lam. (grumixama) seeds in gallic acid solutions of different concentrations (5, 10, 20, and 40 g.L-1), as well as direct application of these solutions to the germination substrate (vermiculite). Whole seeds and seeds fractionated in half were used. Gallic acid’s potential to inhibit germination has been confirmed, especially when applied directly to the substrate at concentrations of 20 and 40 g.L-1. The application of gallic acid also caused delay in germination and seedling development, as well as darkening, shortening, and fragility of the radicles. Thus, the relationship between increased gallic acid concentration in the cotyledons of E. brasiliensis seeds and the inhibition of new root and/or seedling formation was confirmed.
Index terms:
gallic acid; phenolic compounds; regeneration of seeds; recalcitrant seed
RESUMO:
Sementes de espécies de Eugenia L., monoembriônicas, têm a capacidade de regenerar plântulas a partir dos tecidos dos cotilédones quando são fracionadas; contudo, apenas uma plântula se desenvolve em cada fração, indicando a possibilidade da presença de um processo de autocontrole durante a germinação, evitando germinações simultâneas. Um dos prováveis compostos envolvidos nesse controle é o ácido gálico, que aumenta sua concentração nos cotilédones quando a germinação se inicia. Neste trabalho, avaliou-se a possível ação desse composto por meio de pré-embebição das sementes de Eugenia brasiliensis Lam. (grumixama) em soluções de ácido gálico de diferentes concentrações (5, 10, 20 e 40 g.L-1), bem como a aplicação direta dessas soluções no substrato de germinação (vermiculita). Foram utilizadas sementes inteiras e sementes fracionadas ao meio. O ácido gálico confirmou seu potencial inibidor da germinação, principalmente na aplicação direta no substrato nas concentrações 20 e 40 g.L-1. A aplicação do ácido gálico causou, também, atraso da germinação e no desenvolvimento das plântulas, bem como escurecimento, encurtamento e fragilidade das radículas. Confirmou-se, assim, a relação entre o aumento da concentração do ácido gálico nos cotilédones de sementes de grumixama e a inibição da formação de novas raízes e/ou plântulas.
Termos para indexação:
ácido gálico; compostos fenólicos; regeneração de sementes; semente recalcitrante
INTRODUCTION
Studies on seeds of different species of Eugenia L. have shown that fractions of these seeds, removed when germination had already begun, can develop new root and seedling. The same occurs when the seed is intact, but the initial germination has been lost (Prataviera et al., 2015; Amador and Barbedo, 2015; Alonso et al., 2024). This development originates from perivascular parenchyma cells, located near the injured region of the cotyledons, which divide periclinally and proliferate, giving rise to a root meristem (Delgado et al., 2022).
On the other hand, each fraction produces only one new seedling, suggesting some self-control of germination so that the reserves are not spent completely in simultaneous germination. This has been demonstrated when only cracks were caused to the seeds, without completing the cut, and germination occurred in only one of the fractions. When the halves were totally separated, both fractions germinated (Amador and Barbedo, 2011; Amador and Barbedo, 2015). It is likely, therefore, that the beginning of germination and formation of a new seedling produces compounds that inhibit a second germination, such as gallic acid, whose concentration is significantly increased when the seed begins germination (Amador et al., 2026).
Gallic acid is a phenol known as a powerful antioxidant, and it seems to act primarily as an inhibitor of germination and the energy transfer system within the cell (Amorim et al., 2020, and cited references). Exogenous applications of gallic acid in wheat seeds have demonstrated antioxidant action resulting in better germination and vigor (Nobre et al., 2019). It also has antifungal properties in vivo and in vitro (Li et al., 2017) and acts on primary root elongation (Xu et al., 2024). Phenolic compounds belong to the secondary metabolism of plants and can act in defense against herbivory, pathogen attacks, and ultraviolet radiation (Manach et al., 2004). They can be found in seeds, leaves, flowers, and fruits (Amador and Barbedo, 2015; Vatten et al., 2005) and are linked to cell and plant growth (Ndakidemi and Dakora, 2003). Flavonoids and tannins are found in seeds of Eugenia species with possible antioxidative protective and metabolic modulation actions (Nicácio et al., 2017; Saber et al., 2024).
In soybean seeds, with the advance of germination and seedling formation, there was an accumulation of phenolic compounds, including gallic acid (Ma et al., 2022). These compounds were also detected in common bean and mung bean seeds (Ross et al., 2009; Pajak et al., 2014).
Thus, in order to better understand the action of gallic acid on the germination of Eugenia seeds, this study aimed to verify the effects of the application of gallic acid solutions, diluted in water in different concentrations, and to evaluate its action on the inhibition of germination and the regeneration capacity of roots and seedlings in seeds of Eugenia brasiliensis Lam. (grumixama). This species belongs to the Myrtaceae family and is a tree native to Brazil with occurrence in the Seasonal Semideciduous Forest, in the Pluvial Forest, in the Mixed Ombrophilous Forest and in the Restinga, from the states of Bahia to Santa Catarina, Brazil (Mazine et al., 2022).
To this end, two experiments were conducted to analyze the effect of direct application of gallic acid solutions on the germination substrate and the effect of pre-soaking of seeds in these same solutions.
MATERIAL AND METHODS
Ripe fruits of E. brasiliensis were harvested manually from five trees planted in the Botanical Garden of São Paulo, São Paulo, Brazil (23°38’27.38” S and 46°37’34.57” W), ten meters apart, on average, and taken to the Seed Laboratory of the Institute for Environmental Research, also in São Paulo. The seeds were extracted from the fruits manually and then washed in running water to remove pulp residues. Finally, they were placed on germination paper until all surface water was absorbed or evaporated (Amador and Barbedo, 2015).
A sample of seeds was separated for the initial evaluations. Seed moisture content (in %, wet basis) and dry mass were obtained by the oven method at 105 °C for 24 hours (Brasil, 2009), in 4 replicates of 5 seeds. The germination test was performed using 4 replicates of 16 seeds each, per treatment. For this purpose, the seeds were sown in a gerbox, filled halfway with vermiculite previously moistened with water or with gallic acid suspensions, as described below, and placed in a Mangelsdorf germinator, in a germination room with continuous white light and constant temperature of 25 °C. Evaluations were carried out weekly for 42 days, recording seeds with primary root protrusion of at least 0.5 cm, as well as those with the presence of seedlings, as described in Inocente and Barbedo (2021), and those with fungal development, as described in Françoso and Barbedo (2016).
The seeds were separated into two groups, one with whole seeds and the other with seeds fractionated in half, in the longitudinal direction, as described in Alonso and Barbedo (2020). The germination of seeds fractionated in half was calculated by the ratio between fractions with the presence of root and/or seedling and the total number of seeds placed to germinate. Four replicates of 16 seeds each were used, per treatment.
Preparation of gallic acid solutions
Gallic acid solutions were prepared by diluting the powder (3,4,5-trihydroxybenzoic acid, 97.5 - 102.5% titration, Sigma-Aldrich) in water, in Erlenmeyer® flasks, at concentrations of 5, 10, 20 and 40 g.L-1, with the aid of a magnetic stirrer with heating (TECNAL TE-085), until homogeneous solutions were obtained. The solutions were analyzed for their water potential by means of a WP4 potentiometer (Decagon Devices, Inc., Pullman), based on the dew point of the air in hygroscopic equilibrium with the seed sample (Delgado and Barbedo, 2012).
Germination of seeds in substrate moistened with gallic acid
Immediately after preparing the gallic acid solutions, they were used to moisten vermiculite, in gerbox, until achieving saturation (130 mL of solution per gerbox). Then, whole seeds and seed fractions were sown in these substrates, including the control moistened with water, and placed to germinate, as previously described.
Germination of seeds pre-soaked with gallic acid solutions
Whole seeds and seed fractions were kept in suspension for two minutes in water for the control treatments, as also done with gallic acid solutions of 5, 10, 20 and 40 g.L-1, in 50 mL beaker, and immediately placed to germinate in vermiculite already moistened with 130 mL of water, in a gerbox, as previously described. The predetermined time of two minutes was based on previous evaluations of the maximum time at which there had been no sedimentation of gallic acid.
Experimental design and statistical treatment
The experimental design for both experiments was completely randomized, with four replicates. The data were subjected to analysis of variance (5%), and the means were compared to each other by Tukey test at 5% probability level (Santana and Ranal, 2004). When necessary, normality was corrected by transforming the data to (x+0.5)0.5.
RESULTS AND DISCUSSION
The formation of roots in the whole seeds of E. brasiliensis, when the substrate was moistened with gallic acid solutions, was reduced from the concentration of 20 g.L-1, at which germination was about a quarter of the value obtained for germination in substrate moistened with water (Table 1). At the concentration of 40 g.L-1, germination was almost totally inhibited. Regarding fractionated seeds, the regenerative capacity of seeds of Eugenia L. species was once again verified, as already demonstrated in several previous studies (Delgado et al., 2022), since root production in the substrate moistened with water occurred both in the halves that contained the embryonic axis and in several halves consisting only of cotyledon tissue, reaching values above 140% (Table 1). Regarding seedling formation, the results were similar to the previous ones, because a reduction in values was observed from the gallic acid concentration of 20 g.L-1, both for whole and fractionated seeds (Table 1).
Seeds with root, seedling, and fungal incidence (%) of Eugenia brasiliensis Lam., whole and fractionated, after application directly on the substrate of the gallic acid solution at different concentrations. Means followed by the same letter do not differ from each other (Tukey, 5%).
The fractionation of Eugenia seeds often results in high efficiency in root production in fractions that do not contain the embryonic axis, but in proportionally lower efficiency for seedling formation in these fractions (Delgado et al., 2022, and cited references), a fact that has not yet been fully clarified. However, the production of roots in parts of cotyledon that do not contain an embryonic axis, an infrequent event under natural conditions, originates from the differentiation of cells in the perivascular parenchyma (Delgado et al., 2022). Considering that, despite the demonstrated ability of these cells to differentiate and produce new roots, such an event occurs only when there is no root formation in the seed (or in its fractions), which reveals the presence of some self-control of germination in the seed itself (or in its fractions), preventing simultaneous germination (Amador and Barbedo, 2015), as also observed in this study. One of the potential substances involved in this autoinhibition of the formation of new roots, when the first one begins its development, is gallic acid, as its concentration increases in embryonic tissues when germination begins (Amador et al., 2026). The results obtained here corroborate this idea, since the exogenous supply of gallic acid effectively inhibited the formation of roots. It should be noted that the effects of the gallic acid concentrations of 20 and 40 g.L-1 on the seeds were not merely osmotic, as can be assumed when solutions are applied to the germination substrate. The water potentials of these solutions (the lowest of which being -0.52 MPa for 40 g of gallium acid per liter) are well above the water availability values that prevent the germination of the seeds of this species (Inocente and Barbedo, 2019).
When the gallic acid solutions were not applied directly to the germination substrate, but used for seed pre-soaking, the effects were much less evident. For whole seeds, there were no effects of these treatments on root formation or seedling formation (Table 2). However, for fractionated seeds, the inhibition of root and seedling formation was evident, again from 20 g.L-1 gallic acid. The fractionation of seeds may have favored the imbibition of gallic acid solutions, leading to inhibition of the formation of new roots and seedlings. In fact, Eugenia seeds, in general, despite being intolerant to desiccation (Delgado and Barbedo, 2012), have great resistance to water excess or deficit conditions (Inocente and Barbedo, 2021). The whole seed, therefore, may not have offered conditions for the entry of the gallic acid solution in the short time of 2 minutes, during the period of suspension of imbibition, but the fractionated seeds may not have offered the same resistance to this entry. However, this effect needs to be further investigated.
Seeds with root, seedling and fungal incidence (%), of Eugenia brasiliensis Lam., whole and fractionated, after pre-soaking in gallic acid solution at different concentrations. Means followed by the same letter do not differ from each other (Tukey, 5%).
It is curious to notice that gallic acid demonstrated another effect on seeds: the increase in the number of seeds with the presence of fungi. In fractionated seeds, both the application of gallic acid solutions directly to the substrate (Table 1) and the pre-soaking of the seeds (Table 2) resulted in a substantial increase in the number of seeds with fungi at concentrations of 20 and 40 g.L-1. To date, there are no reports in the literature regarding the action of gallic acid on seed defense systems. Interestingly, however, gallic acid has been reported as an efficient antifungal compound in plants (El-Nagar et al., 2020). Gallic acid has, among other properties, a powerful antioxidant effect (Li et al., 2023) and, as described by Amorim et al. (2020), the concentration of reactive oxygen species (ROS) in seeds can define stimulating or inhibiting effects on germination, as well as their reaction to attack by microorganisms. Therefore, the gallic acid concentrations used in the present study may have been high to the point of reducing the defense systems of the seeds, thus allowing a higher incidence of fungi.
In addition to the differences observed in the final results of germination, after application or imbibition of the seeds in gallic acid solutions, the effects of these solutions on the temporal distribution of this germination were also verified (Figures 1 and 2). From these results, it was evident that the effects of gallic acid have already occurred in the solution with a concentration of 10 g.L-1. It was also possible to observe the effects of gallic acid solutions on seedling development at the end of the germination test (Figure 3), in which the damage caused to the seedlings is also observed at a concentration of 10 g.L-1. It was found that the higher the concentration of gallic acid solution, the smaller and darker the roots were.
Temporal distribution of germination (in blue) and seedling production (in orange) of whole (A, C, E, G and I) or fractionated (B, D, F, H and J) seeds of Eugenia brasiliensis Lam., after application directly on the substrate of gallic acid solutions at different concentrations (A and B: water; C and D: 5 g.L-1; E and F: 10 g.L-1; G and H: 20 g.L-1; I and J: 40 g.L-1).
Temporal distribution of germination (in blue) and seedling production (in orange) of whole (A, C, E, G and I) or fractionated (B, D, F, H and J) seeds of Eugenia brasiliensis Lam., after pre-soaking in gallic acid solutions at different concentrations (A and B: water; C and D: 5 g.L-1; E and F: 10 g.L-1; G and H: 20 g.L-1; I and J: 40 g.L-1)
Seedlings obtained on the last day of evaluation of the germination test from whole or fractionated seeds of E. brasiliensis, placed on substrate moistened with gallic acid solutions in different concentrations (water, 5 g.L-1, 10 g.L-1, 20 g.L-1 and 40 g.L-1) or after pre-soaking in these solutions.
The results obtained in the present study showed the great potential of gallic acid to inhibit the germination of E. brasiliensis seeds. Therefore, the increase in the concentration of this phenolic compound, when seed germination begins, may be involved in the process that avoids simultaneous germination in the seeds. This may represent an important evolutionary advance for the species, since the seeds, even if they are intolerant to desiccation, they can produce seedlings and colonize the environments even when the first germinations occur under unfavorable conditions for seedling development. Successive germinations from the same seeds, increasing the time these seeds remain in the soil in conditions to produce seedlings (Alonso and Barbedo, 2020), result in a system similar to the seed bank in the soil, but without the use of desiccation tolerance and/or dormancy.
CONCLUSIONS
Application of gallic acid solution, directly on the substrate or in pre-soaking of the seeds of E. brasiliensis, inhibits or delays germination and seedling formation from the concentration of 10 g.L-1, especially at concentrations of 20 and 40 g.L-1.
ACKNOWLEDGMENTS
To the Institute for Environmental Research, for providing a place and materials for this project to be carried out; to the Coordination for the Improvement of Higher Education Personnel (CAPES) for the master’s scholarship granted to Y.T. Cavalcanti; to the Graduate Program in Plant Biodiversity and Environment, for the opportunity given to the first author for her Master’s degree.
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