Foliar application of salicylic acid on photochemical responses and growth of Hymenaea courbaril under floodingABSTRACT
Flooding, depending on exposure time, can be stressful and impair plant morphophysiology, but the foliar application of salicylic acid (SA) is an efficient physiological management to alleviate these negative effects. We aimed to evaluate the responsiveness of Hymenaea courbaril L. to flooding and the potential of SA to alleviate the associated stress. Seedlings received foliar applications of four SA concentrations (0, 100, 200, and 400 mg L-1), were exposed to two water regimes (control and flooded), and were evaluated at 15 and 30 days. The treatments were arranged in a subplot scheme, with water regimes assigned to plots and SA doses to subplots. The evaluation periods were treated as time-structured plots. Although flooding decreased photochemical activity in photosystem II, especially at 15 days, it did not negatively affect seedling growth. Foliar application of SA at varying doses increased chlorophyll indices and stimulated seedling growth, as reflected in biomass production, but did not affect photochemical efficiency. H. courbaril seedlings are physiologically responsive to flooding, but this did not reflect in growth through adjustment mechanisms. Foliar application of SA at a dose between 242-266 mg L-1 stimulates the growth of H. courbaril seedlings, regardless of exposure time and water regimes.
Key words:
chlorophyll a fluorescence; ‘jatobá’; exposure time; photosystem II; plasticity
HIGHLIGHTS:
Exposure time to flooding altered the photochemical performance of Hymenaea courbaril L. seedlings.
Foliar application of salicylic acid favored seedling growth.
Flooding did not harm the growth of H. courbaril L. seedlings.
RESUMO
O alagamento em função do tempo de exposição pode ser estressante e prejudicar a morfofisiologia das plantas, mas a aplicação foliar de ácido salicílico (AS) tem demonstrado ser um manejo fisiológico eficiente para aliviar esses efeitos negativos. Objetivamos avaliar a responsividade de Hymenaea courbaril L. ao alagamento e o potencial do AS em aliviar o efeito estressante dessa condição. As mudas receberam aplicação foliar de quatro concentrações de AS: 0, 100, 200 e 400 mg L-1, e foram expostas a dois regimes hídricos: controle e alagamento, e avaliadas aos 15 e 30 dias. Os tratamentos foram arranjados em esquema de subparcelas, alocando os regimes hídricos nas parcelas e as doses de AS nas subparcelas. Os períodos de avaliação foram considerados parcelas subdivididas no tempo. Embora o alagamento reduziu as atividades fotoquímicas no fotossistema II, especialmente aos 15 dias, não afetou negativamente o crescimento das mudas. Aplicação foliar de AS em doses variadas favoreceu o aumento dos índices de clorofilas e estimulou o crescimento das mudas, refletindo em produção de biomassa, mas não apresentou efeito sobre a eficiência fotoquímica. As mudas de H. courbaril são responsivas fisiologicamente ao alagamento, mas não refletiu no crescimento devido mecanismos de ajustes. A aplicação foliar de AS em dose entre 242-266 mg L-1 estimulou o crescimento das mudas de H. courbaril independente do tempo de exposição e dos regimes hídricos.
Palavras-chave:
fluorescência da clorofila a; ‘jatobá’; tempo de exposição; fotossistema II; plasticidade
INTRODUCTION
In recent years, global climate change has been a cause for concern, as it has negatively affected agricultural and forestry activities. This is because it has caused water fluctuations, including periods of excess water in many regions, characterizing a condition of hypoxia or anoxia. Flooding is a condition that, depending on the function of exposure time and the sensitivity of each species, can be stressful.
This condition can impair aerobic root respiration in response to oxygen limitation (Jethva et al., 2022; Eerdekens et al., 2024; Seymen et al., 2024), affecting energy production, which can compromise the efficiency of photochemical activities in photosystem II (PSII) reaction centers, resulting in lower photosynthetic capacity (Ji & Hyun, 2023; Santos et al., 2023; Veen et al., 2025). However, survival capacity and the appearance of adjustment mechanisms to flooding vary with exposure intensity and the phenotypic plasticity of each species.
In this context, within the field of stress physiology, studies aimed at establishing practices to alleviate the negative effects of adverse conditions have increased. According to Saracho et al. (2021) and Santos et al. (2023), salicylic acid (SA), a phytohormone and phenolic compound, enhances the activity of antioxidant enzymes and non-enzymatic compounds, which stabilize the photochemical apparatus and stimulate plant growth.
Among the species of interest in recovery programs of degraded areas or integrated production systems, Hymenaea courbaril L. (‘jatobá’, Fabaceae) is a tree and fruit species found in several regions of South America. The leaves, fruits, and seeds have pharmacological potential (Matos et al., 2023). In addition to their timber value, seedlings can be included in reforestation programs for native forests due to their rapid growth (Ferreira et al., 2016).
H. courbaril occurs from humid forests and flooded environments to dry seasonal forests, such as Cerrado and Caatinga (Oliveira et al., 2011; Locosseli et al., 2016), being frequent in dryland forests and high plains, as well as in clayey and poor soils (Duarte et al., 2016). However, there are still gaps in studies examining associations between SA doses and flooding for the species under study.
We hypothesized that (i) flooding is a stressful condition for H. courbaril and its negative effects vary depending on the time of exposure of the seedlings, but that (ii) the application of SA at specific concentrations modulates chlorophyll indices, improving photochemical performance and stimulating growth, acting to induce tolerance of the species to flooding. We aimed to evaluate the responsiveness of H. courbaril seedlings to flooding and the potential of SA to alleviate the stress.
MATERIAL AND METHODS
The experiment was conducted under nursery conditions with 30% shading screen on top and sides, at the Faculty of Agricultural Sciences (22° 11’ 51.5” S, 54° 56’ 04.3” W) of the Federal University of Grande Dourados (UFGD), Dourados - MS, Brazil.
Ripe fruits of H. courbaril were collected from ten matrices in a remaining area of Cerrado, in Dourados-MS. After manual processing of the fruits, the seeds were subjected to overcoming dormancy by mechanical scarification of the embryo and subsequent immersion in water for 24 hours (pre-soaking), as described by Souza et al. (2015). Subsequently, the seeds were sanitized with sodium hypochlorite for 5 min and washed in running water.
Sowing was carried out in 290 mL tubes previously filled with commercial Tropstrato® substrate. At 15 days after emergence, when the seedlings reached an average height of 15 cm, they were transplanted into 7-L plastic pots filled with Oxisol (Soil Survey Staff, 2022) from 0-20 cm depth + coarse sand (commercially sourced) in a 3:1 (v/v) ratio. The mixture exhibited the following chemical properties, as shown in Table 1.
Chemical attributes of soil used in experiment with Hymenaea courbaril L. seedlings exposed to water regimes and foliar application of salicylic acid
Irrigation was carried out daily maintaining 70% of the water retention capacity (WRC) according to Souza et al. (2000). When seedlings reached an average height of 25.0 cm, received application of salicylic acid (SA) according to treatment: 0, 100, 200, and 400 mg L-1, via foliar application repeated in five applications in 10 days, during 08:00-10:00 hours. For this, SA (99% P.A.) was previously diluted in 20 mL of ethyl alcohol (concentration in the spraying solution: 2.5%) and distilled water was added; 1.0 mL of adjuvant LI 700 was added to facilitate the adhesion of solution on abaxial and adaxial surface of leaves of H. courbaril (Santos et al., 2023), until reaching the runoff point (10 mL per plant, based on pre-test).
At the end of 10 days after the application of SA doses, the seedlings were exposed to two water regimes: i) control (CK) - non-flooded, being irrigated daily maintaining 70% WRC and ii) flooded (FL) - were placed in plastic pools throughout the evaluation period maintaining continuously the water level 5.0 cm above the substrate, as shown in illustration (Figure 1).
Schematic illustration of the foliar application of salicylic acid (SA) in Hymenaea courbaril L. seedlings exposed to flooding, evaluated at 15 and/or 30 days. Source: Illustration adapted using Gemini AI
The evaluations of chlorophyll indices, chlorophyll a fluorescence, and growth (height and stem diameter) were assessed at 15 and 30 days after seedlings were exposed to two water regimes, while total dry mass was evaluated only at 30 days.
The experimental design was completely randomized, with four replicates, and each experimental unit consisted of a pot with two plants. The treatments were arranged in a subplot scheme, with water regimes assigned to plots and SA doses to subplots. The evaluation periods were treated as time-structured plots.
During the experimental period, the pool water was changed weekly to prevent insect proliferation. The pool water temperature was 19.9 and 22.3 ºC at 15 and 30 days, respectively.
The evaluations of chlorophyll indices and chlorophyll a fluorescence were performed on fully expanded leaves located in the middle third of the plants, during 08:00-10:00 hours.
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a) Chlorophyll indices: using a Falker CLF 1030 chlorophyll meter (Falker Chlorophyll Index, FCI), chlorophyll a, b, and total were measured.
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b) Chlorophyll a fluorescence: the leaves were subjected to dark conditions using adapter clips for 30 min. After this period, using a portable fluorometer (OS-30p; Opti-Sciences Chlorophyll Fluorometer, Hudson, NY, USA) under a flash of 1,500 µmol m-2 s-1, initial (F0) and maximum (Fm) chlorophyll a fluorescence and photochemical potential quantum efficiency in photosystem II (Fv/Fm) were determined. From the F0 and Fm data, absorbed energy conversion efficiency (Fv/F0) and the maximum basal yield of non-photochemical processes (F0/Fm) were calculated.
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c) Initial growth: plants were removed from the flooded condition, and plant height (H, cm) was measured, in which the established criterion was the distance from the collar to the apical bud, using a graduated ruler. The stem diameter (D, mm) was measured with a digital caliper inserted 5.0 cm above the substrate level, and the number of leaves was counted.
The seedlings were removed from the plastic pots and washed to remove excess substrate from the roots. The materials were placed in Kraft® paper bags and dried in an oven with forced-air circulation at 60 ± 5 ºC for 72 hours, and the total dry mass (g per plant) was weighed on a scale with thousandth precision (0.0001 g).
The data were subjected to analysis of variance. When significant by the F test (p ≤ 0.05), the means as a function of the isolated effects and the interaction were compared by the Bonferroni t-test for water regimes and evaluation periods (p ≤ 0.05) and regression analysis testing the linear and quadratic models for salicylic acid doses (p ≤ 0.05), using the SISVAR software.
RESULTS AND DISCUSSION
H. courbaril seedlings are physiologically responsive to flooding, with a reduction in photochemical activity in PSII, but this did not negatively affect the growth and production of photoassimilates, demonstrating adjustment mechanisms by physiological plasticity. Conversely, we partially rejected our initial hypothesis, as exogenous SA application did not mitigate the negative effect of flooding on photosystem II energy efficiency but did stimulate plant growth.
The chlorophyll a, b, and total indices in H. courbaril seedlings were influenced by the interaction between water regimes and SA doses (Table 2). The initial chlorophyll a fluorescence (F0), photochemical potential quantum efficiency (Fv/Fm), and maximum basal yield of non-photochemical processes (F0/Fm) were influenced by water regimes and evaluation periods isolated. The maximum fluorescence (Fm) was not influenced by factors under study. The absorbed energy conversion efficiency (Fv/F0) was influenced by water regimes, SA doses, and evaluation periods isolated.
Summary of F test of isolated effect or interaction of water regimes, salicylic acid doses, and evaluation periods on chlorophyll a, b, total indices, chlorophyll a fluorescence, and photochemical processes in Hymenaea courbaril L. seedlings, evaluated at 15 and 30 days
The chlorophyll a index showed a linear response to flooding, with the highest value (28.96 FCI) at 400 mg L-1 SA, whereas the CK plants showed a quadratic response, with the maximum value (29.05 FCI) at 167.50 mg L-1 SA (Table 3). Regarding the chlorophyll b index, in both water regimes, the adjustments were quadratic, with maximum values (11.19 and 9.21 FCI) in seedlings with 282.85 mg L-1 and without SA in the CK and FL conditions, respectively. The chlorophyll total index was highest and lowest (38.54 and 34.72 FCI) at 183.00 and 213.00 mg L-1 SA in CK and FL conditions, respectively.
Regression equations and coefficients of determination of chlorophyll a, b, and total index in Hymenaea courbaril L. seedlings cultivated with doses of salicylic acid and exposed to two water regimes (CK: control; FL: flooded)
Although the role of SA in the biosynthesis of photosynthetic pigments is little known and discussed, it was possible to demonstrate that this phytohormone modulates chlorophyll levels in both water regimes. Still, it did not directly reflect on the activities in PSII. Similarly, Santos et al. (2022) observed an increase in chlorophyll levels in Schinus terebinthifolia Raddi seedlings. In addition, without SA, flooded plants had a higher chlorophyll index than CK plants, indicating a mechanism for maintaining metabolic processes, and that exogenous SA application increased it under both water regimes.
The reduction in chlorophyll index observed at the highest salicylic acid dose (400 mg L⁻1) under control conditions indicates a physiological threshold above which SA ceases to act as a stimulatory modulator and begins to exert an inhibitory effect on H. courbaril seedlings. We believe that, although SA is widely described as a key regulator of tolerance to abiotic stress, its effects are strongly dependent on dose and the environmental context.
Contradictory evidence in the literature indicates that the effects of salicylic acid on the photosynthetic apparatus are not universal but are strongly dependent on the interactions among dose, species, and environmental context. Some studies report increased chlorophyll content and improved photosynthetic efficiency under abiotic stress conditions, attributing a protective role to SA in maintaining chloroplast integrity (Arruda et al., 2023; Sperdouli et al., 2024). Conversely, other studies demonstrate that such benefits are reduced or nonexistent in plants not subjected to stress, or even reversed into negative effects when the compound is applied at high concentrations (Fátima et al., 2023; Decsi et al., 2025).
This apparent paradox suggests that SA acts within an optimal physiological range, outside of which it can trigger responses associated with secondary stress, including redox imbalance and pigment degradation, characterizing a hormetic-type response in which low doses exert a stimulatory effect. At the same time, supraoptimal levels result in physiological inhibition (Khan et al., 2015). Additionally, the activation of defense-related metabolic pathways, often induced by SA, can impose a significant energy cost on plants, redirecting resources from primary metabolism to secondary metabolism (Miao et al., 2024).
The higher F0 values were observed in flooded plants and at 30 days (Table 4). The highest values of the Fv/Fm and Fv/F0 ratios occurred in CK plants at 30 days, demonstrating that flooding impaired the activity of the photochemical apparatus, as represented by the PSII reaction centers. Under stress conditions, such as flooding, the overproduction of reactive oxygen species (ROS) and the presence of toxic compounds from fermentative metabolism (Jethva et al., 2022; Veen et al., 2025) alter leaf metabolism.
Initial fluorescence (F0), photochemical potential quantum efficiency in photosystem II (Fv/Fm), absorbed energy conversion efficiency (Fv/F0), and maximum basal yield of non-photochemical processes (F0/Fm) in Hymenaea courbaril L. seedlings exposed to two water regimes (CK: control; FL: flooded) and two evaluation periods (15 and 30 days)
The decrease in photochemical yields can be associated with the fact that, under stressful conditions, plants may direct energy toward other adjustment mechanisms, such as increased activity of antioxidant enzymes and increased levels of amino acids. Although we did not evaluate it in our study, the activity of protective metabolism helps combat ROS and alleviate damage to the photosynthetic apparatus (Zhang et al., 2023; Choudhary et al., 2024; Xu et al., 2024), as reflected in the increase in F0/Fm.
In addition, it is observed that over the days, i.e., exposure time increases, H. courbaril increases the values of Fv/Fm and Fv/F0 and reduces F0/Fm, indicating an advanced physiological stage of the seedlings, which reflects greater photosynthetic capacity and carbohydrate accumulation throughout their life cycle, especially since they are perennial. Exogenous application of SA influenced the Fv/F0, with a quadratic model significant, but without satisfactory mathematical adjustment (ŷ = 0.9997 - 0.0031*x + 0.00007*x2; R2 = 0.55), with a higher value without and with 400 mg L-1 SA.
The plant height, stem diameter, and total dry mass (TDM) were influenced by doses of salicylic acid (Table 5).
Summary of F test of isolated effect or interaction of water regimes, salicylic acid doses, and evaluation periods on the growth of Hymenaea courbaril L. seedlings, evaluated at 15 and/or 30 days
The maximum estimated values were 42.50 cm, 5.96 mm, and 5.21 g per plant with 242.16, 266.66, and 252.00 mg L-1 SA, respectively, regardless of water regimes (Figure 2). From these results, we reinforce that H. courbaril was physiologically responsive. Still, its plasticity conferred tolerance to these conditions, maintaining photoassimilates production and accumulation, which positively reflected in the increase in TDM, similar to that observed by Saracho et al. (2021). Foliar application of SA may have contributed to CO2 assimilation and the efficiency of carbohydrate production, favoring the expansion of growing organs, as reflected in height and stem diameter, reinforcing the results of our study on H. courbaril.
Regression equations and coefficients of determination of height (A), stem diameter (B), and total dry mass (C) of Hymenaea courbaril L. seedlings grown with doses of salicylic acid, evaluated at 30 days
Salicylic acid regulates cellular antioxidant mechanisms and, as a signaling hormone, promotes the growth, development, and differentiation of plant cells and tissues (Hasanuzzaman et al., 2022). Furthermore, SA increases dry biomass, especially at higher concentrations. This occurs because SA increases the synthesis of carbohydrates, proteins, and lignin (Shao et al., 2018; Saracho et al., 2021), thereby promoting greater tolerance to flooding-induced water stress.
We believe that the time of exposure to flooding evaluated was insufficient for the plant to exhaust its physiological or biochemical adjustment mechanisms, which ensured its plasticity and investment in morphological changes, such as the formation of lenticels and other stress symptoms. In addition, the lower salicylic acid participation, especially as a mitigating agent, is due to conditions that were not stressful over 30 days. We suggest new studies evaluating longer periods of exposure to stress and assessing the activity of protective metabolism using enzymatic and non-enzymatic compounds, aiming to understand adjustment strategies.
CONCLUSIONS
1. Hymenaea courbaril L. seedlings respond to flooding by altering the performance of photochemical activities in photosystem II, but this does not impair initial growth up to 30 days of evaluation.
2. Exogenous application of salicylic acid did not mitigate the effect of flooding depending on the function of exposure times in chlorophyll indices and chlorophyll a fluorescence, but at doses between 242-266 mg L-1 stimulated seedling growth.
1
Financing statement:
Acknowledgments:
The authors thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), for granting the scholarships, and the Fundação de Apoio ao Desenvolvimento do Ensino, Ciência e Tecnologia do Estado de Mato Grosso do Sul (FUNDECT), for financial support.
Data Availability Statement:
The original contributions presented in the study are included in the article, and further inquiries can be directed to the corresponding author.
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* - Significant at p ≤ 0.05 by F test ± standard deviation