Physiological soybean seed quality in response to the application of plant bioregulatorABSTRACT:
The present study aimed to evaluate the physiological quality of freshly harvested and stored soybean seeds following the foliar application of nutrients and a plant bioregulator. Three field experiments were conducted in the municipality of Lavras, Minas Gerais, Brazil. Treatments consisted of three soybean cultivars (Desafio RR, M5917 IPRO, and 95R40 IPRO) subjected to foliar application of nine products containing nutrients and plant bioregulators at the phenological stages R1 and R2, in addition to a control treatment. After harvest, the seeds were subjected to physiological analyses, including electrical conductivity, vigor and viability by the tetrazolium test, emergence speed index, and accelerated aging, at three evaluation periods: immediately after harvest, and after three and six months of storage. Data were analyzed by analysis of variance, with effects assessed using the F-test and means compared through 95% confidence intervals. A significant effect of product application was observed for most physiological variables. However, responses varied according to cultivar, product applied, and storage period. The effects were more pronounced in vigor-related traits, particularly in freshly harvested seeds and after three months of storage. Overall, treatments did not show a consistent effect on seed storability.
Index terms:
foliar application; Glycine max (L) Merr.; plant nutrition; seed physiological quality
RESUMO:
Objetivou-se avaliar a qualidade fisiológica de sementes de soja recém-colhidas e armazenadas após aplicação foliar de nutrientes e biorregulador vegetal. Foram conduzidos três experimentos, em campo, no município de Lavras-MG. Os tratamentos constituíram de três cultivares de soja (Desafio RR, M5917IPRO e 95R40 IPRO) com aplicação foliar de nove produtos contendo nutrientes e biorregulador vegetal, no estádio fenológico R1 e R2, além do tratamento controle. Após a colheita das sementes, estas foram submetidas às análises fisiológicas (testes de condutividade elétrica, vigor e viabilidade por tetrazólio, índice de velocidade de emergência e envelhecimento acelerado), em três períodos destinos: recém-colhidas, aos três e seis meses de armazenamento. Os dados foram submetidos à análise de variância, sendo os efeitos avaliados pelo teste F e comparados por meio de intervalos de confiança a 95%. Observou-se efeito significativo da aplicação dos produtos sobre a maioria das variáveis fisiológicas. Entretanto, as respostas variaram em função da cultivar, do produto aplicado e do período de armazenamento. Os efeitos foram mais expressivos no vigor, especialmente em sementes recém-colhidas e após três meses de armazenamento. De modo geral, os tratamentos não apresentaram efeito consistente sobre a armazenabilidade das sementes.
Termos para indexação:
aplicação foliar; Glycine max (L) Merr.; nutrição vegetal; qualidade fisiológica de sementes
INTRODUCTION
Physiological quality is defined as the capacity of a seed lot to generate normal seedlings, as determined by germination, vigor, and longevity (Pirredda et al., 2024). These attributes are fundamental to successful stand establishment and are closely associated with high yield (Ghosh et al., 2026).
Longevity is a prerequisite for maintaining seed performance over time, as deterioration inevitably leads to progressive loss of vigor. Even under relatively favorable storage conditions, such as temperatures of 10 °C and relative humidity below 40%, soybean seeds exhibit relatively limited longevity (López-Rodríguez et al., 2026), underscoring the intrinsic susceptibility of this species to physiological deterioration during storage.
The soybean reproductive stages, pod set, seed filling, and seed composition, are critical due to their strong dependence on photosynthetic efficiency. Reduced photosynthesis limits the synthesis, translocation, and accumulation of photoassimilates, impairing yield and seed physiological quality (Oliveira et al., 2025). Seed production is further determined by the number of flowers per plant and the proportion that develop into pods.
The relevance of seed vigor has been consistently demonstrated, as low-vigor seeds are associated with reduced seedling emergence, diminished leaf area expansion, fewer pods per plant, and ultimately lower soybean yield (Ebone et al., 2020). Although appropriate post-harvest conditioning is essential for preserving seed vigor and viability, it is equally critical to maximize physiological quality prior to storage, since seed deterioration is an irreversible and cumulative process (Salvador et al., 2025).
Plant nutrition plays a fundamental role in determining the physiological quality of seeds. Adequate nutrient availability throughout the crop cycle directly influences embryo development and the accumulation of storage reserves, thereby altering seed chemical composition and affecting seed vigor and longevity (Conceição et al., 2020).
Among the micronutrients required for reproductive development, boron (B) and calcium (Ca) are particularly relevant due to their roles in pollen tube growth, fertilization, and seed set. Cobalt (Co) and molybdenum (Mo) are also important in legumes because of their involvement in nitrogen metabolism and biological nitrogen fixation (BNF), contributing to plant performance and seed quality (Abreu-Junior et al., 2023). The R1 and R2 stages represent strategic windows for foliar application because they encompass the onset and full flowering of soybean. Recent studies have reported effects of Ca and B applications during flowering on reproductive and yield-related traits (Galeriani et al., 2022; Garcia et al., 2025), while micronutrient application at R1-R2 has been associated with changes in seed size, germination, and vigor (Cardoso et al., 2024).
Beyond mineral nutrition, plant bioregulators can modulate physiological and morphological processes that sustain plant growth and development. These organic compounds regulate several physiological pathways (Zahid et al., 2023), influencing processes such as tissue differentiation, organogenesis, senescence, flowering, seed germination, and dormancy. Furthermore, plant growth regulators may reduce fruit abscission, enhance fruit set, and improve fruit retention and quality (Zahid et al., 2023), suggesting potential indirect effects on seed development and subsequent physiological performance (Domingos et al., 2021; Gabri et al., 2024).
Therefore, we hypothesized that foliar application of calcium, boron, and a plant bioregulator during the reproductive stage would improve the physiological quality of soybean seeds and contribute to the maintenance of seed vigor during storage, although the magnitude of these responses would vary among cultivars and storage periods.
Thus, this study aimed to evaluate the physiological quality of soybean seeds produced from plants subjected to the application of calcium, boron, and a plant bioregulator during the reproductive stage, as well as the longevity of these seeds during storage.
MATERIAL AND METHODS
Seed production was conducted in Lavras, Minas Gerais, Brazil, during the 2021/2022 crop season. Climatic data, including air temperature and relative humidity recorded throughout the experimental period, are presented in Figure 1. The experimental area is located at an average altitude of 955 m and features a Typic Dystroferric Red Latosol, according to the Brazilian Soil Classification System (Santos et al., 2018), characterized by a deep profile and high clay content.
Accumulated precipitation and maximum, mean, and minimum air temperatures recorded in Lavras, Minas Gerais, Brazil, during the 2021/2022 growing season.
Three contiguous experiments were conducted simultaneously, each corresponding to one soybean cultivar: Desafio RR, M5917 IPRO, and 95R40 IPRO. The treatments were applied at the reproductive phenological stages R1 (beginning bloom) and R2 (full bloom) (Table 1).
The trial was conducted under field conditions using a randomized complete block design (RCBD) with three replicates. Each experimental plot consisted of four rows, 5 m in length, spaced 0.60 m apart, considering the two central rows as the effective plot area.
At sowing, basal fertilization was applied in the planting furrow at a rate of 300 kg ha⁻¹ of the N-P₂O₅-K₂O (02-30-10) formulation, following soil fertility recommendations for soybean cultivation. Seeds were inoculated with Bradyrhizobium japonicum (strains SEMIA 5079 and SEMIA 5080), applied directly in the sowing furrow at planting.
Sowing was performed mechanically in October. Weed, pest, and disease management were carried out in accordance with the technical recommendations for soybean cultivation (Embrapa, 2020).
Harvesting was conducted manually when plants reached the R8 phenological stage (full maturity), as defined by Fehr and Caviness (1977). Threshing was subsequently performed using a vertical plot thresher (IMACK), with due care taken to minimize mechanical damage. The seeds were then transported to the Central Seed Research Laboratory of the Universidade Federal de Lavras (LCPS/UFLA), where physiological quality assessments were conducted.
For laboratory analyses, seeds retained on 6.5-mm round-hole sieves were selected to ensure uniformity in seed size and reduce variability among samples. The evaluations were performed under a completely randomized design (CRD), with four replicates of 50 seeds per treatment. An initial subsample was used to determine the physiological quality of freshly harvested seeds. The remaining seeds were packaged in multiwall paper bags (1 kg capacity) and stored for three and six months in a cold chamber under controlled environmental conditions (11 ± 2 °C and 55% ± 2 relative humidity).
Freshly harvested seeds were subjected to electrical conductivity, tetrazolium viability and vigor tests (França-Neto et al., 1998), standard germination (Brasil, 2025), seedling emergence speed index (Maguire, 1962), and accelerated aging (Marcos-Filho, 1999; Brasil, 2025). For the tetrazolium test, four replicates of 50 seeds were preconditioned at 25 °C for 16 h. Subsequently, the seeds were immersed in a 0.075% solution of 2,3,5-triphenyl tetrazolium chloride and incubated at 40 °C for 3 h in the absence of light. Tetrazolium viability and vigor were assessed according to the staining pattern and tissue characteristics described by França-Neto et al. (1998).
After three and six months of storage, germination, seedling emergence, and accelerated aging tests were repeated to monitor changes in seed performance over time. The electrical conductivity test was conducted in accordance with Vieira and Marcos-Filho (2020), using a digital conductivity meter (MS TECNOPON® - mCA150), to estimate membrane integrity and seed vigor.
Data for Germination (G), Emergence Speed Index (ESI), and Accelerated Aging (AA) were subjected to tests of homogeneity of variances using Bartlett’s test (Bartlett, 1937) and normality of residuals using the Shapiro-Wilk test (Shapiro and Wilk, 1965). After confirming these assumptions, the data were analyzed by analysis of combined variance (ANOVA) at the 5% probability level (p ≤ 0.05). When significant effects were detected, means were compared using 95% confidence intervals estimated from the residual mean square (MSE) and the Student’s t distribution, with graphical inference performed using the plotCI function (Silva, 2025) and adapted for the present study.
Experimental precision was evaluated using the coefficient of variation (CV), as proposed by Resende and Duarte (2007). All statistical analyses were conducted in the R statistical computing environment (R Core Team, 2026), version 4.3.3.
RESULTS AND DISCUSSION
Experimental precision is a prerequisite for robust statistical inference, as it enables the estimation of trait values with minimal deviation from their true magnitude (Souza et al., 2023). In the present study, the coefficient of variation ranged from 2.95% to 26.33%, which, according to Resende and Duarte (2007), is classified from very low to medium, thereby indicating satisfactory experimental control and reliability of the evaluated parameters.
We observed significant effects of the treatments for electrical conductivity (EC), tetrazolium vigor (TZVig), and tetrazolium viability (TZViab) across all cultivars (Appendix F). Germination differed statistically among treatments for the cultivar 95R40 IPRO in both freshly harvested seeds and after three months of storage. For M5917 IPRO, significant differences were observed throughout all evaluation periods, with mean germination values consistently around 82%, suggesting stable physiological performance across storage intervals.
The emergence speed index (ESI) was influenced by treatments after three months of storage in M5917 IPRO and 95R40 IPRO, and in freshly harvested seeds of M5917 IPRO. In contrast, accelerated aging (AA) was not significantly affected by treatments over storage time, with values ranging from 72.35% to 57.70%. The absence of treatment effects on AA suggests that foliar nutrient and bioregulator applications were insufficient to modify the mechanisms underlying seed tolerance to accelerated deterioration. This response may be associated with the predominance of intrinsic biochemical mechanisms governing seed tolerance to aging. In soybean, resistance to deterioration has been associated with activation of the phenylpropanoid pathway, particularly the accumulation of caffeic acid, which can inhibit lipoxygenase (LOX) activity and contribute to the maintenance of seed quality during aging (Li et al., 2024a).
Seed aging encompasses a complex set of cytological, physiological, biochemical, and physical alterations that culminate in the progressive decline of physiological quality and viability (Nadarajan et al., 2023). Notably, the deterioration rate is not linear or uniform; it varies not only among species but also among genotypes within the same species, depending on environmental conditions during seed development and the intrinsic genetic attributes of the evaluated lines (Pirredda et al., 2024). In soybean, deterioration has been more closely associated with lipid peroxidation, particularly linked to linoleic acid content, than with total oil concentration or oleic acid levels (Silva et al., 2023). Nevertheless, genotype-dependent sensitivity plays a decisive role, as evidenced by the differential responses observed in this study.
For cultivar 95R40 IPRO, the treatments 9, 2 e 8 resulted in EC values of 42.96, 43.15, and 43.46 µS·cm⁻¹·g⁻¹, respectively (Figure 2A), significantly differing from the control. These lower EC values denote superior membrane integrity and reduced solute leakage, which are indicative of enhanced physiological quality and seed vigor (Sharaf et al., 2026).
Ninety-five percent confidence intervals (95% CIs) for electrical conductivity (EC; µS·cm⁻¹·g⁻¹), viability (TZViab; %) vigor (TZVig; %) of freshly harvested soybean seeds from different cultivars subjected to plant bioregulator applications.
The loss of cellular membrane integrity constitutes one of the earliest and most sensitive indicators of seed physiological decline. In soybean, the electrical conductivity test is grounded in the premise that seeds with advanced deterioration exhibit greater solute efflux during imbibition, whereas seeds with preserved structural organization rapidly reestablish membrane functionality, thereby minimizing solute leakage (Li et al., 2024b). Thus, reduced EC values reflect a more efficient membrane reorganization process and greater metabolic competence during the initial stages of germination.
The treatments 9 and 2 contained kinetin, gibberellic acid, and indole-3-butyric acid. The lower EC observed in these treatments may be associated with effects of these compounds and nutrients on seed development and postharvest quality. This interpretation is supported by Jappe et al. (2026), who demonstrated that field application of cytokinin at different phenological stages of soybean modified postharvest grain quality, including electrical conductivity and the amount of leached solids. Notably, cytokinin application at the early reproductive stage (R1) altered grain biochemical characteristics, indicating that the timing of hormonal application may influence subsequent seed quality. Thus, the lower EC observed in treatments containing kinetin may reflect changes in seed development and composition resulting from foliar application during the reproductive phase, although membrane integrity was not directly evaluated in the present study.
Treatment 8, containing boron (B) and molybdenum (Mo) and applied during the reproductive stage, also resulted in low EC. This response may be associated with the role of B in maintaining cell wall structure and membrane stability, while Mo contributes to nitrogen metabolism as a cofactor of nitrate reductase, potentially supporting nitrogen assimilation and seed development (Vera-Maldonado et al., 2024; Cardoso et al., 2024).
Although variations in EC magnitude were observed among cultivars, the overall means were classified as low. The average EC values were 50.94, 59.78, and 57.94 µS·cm⁻¹·g⁻¹ for 95R40 IPRO, Desafio RR, and M5917 IPRO, respectively. According to Prado et al. (2019), EC values are categorized as follows: very high vigor (≤ 70 µS·cm⁻¹·g⁻¹), high (71-90 µS·cm⁻¹·g⁻¹), medium (91-110 µS·cm⁻¹·g⁻¹), and low (≥ 111 µS·cm⁻¹·g⁻¹). Therefore, the EC values obtained in the present study fall within the very high vigor range and remain well below the threshold typically associated with compromised membrane integrity, reinforcing the overall high physiological quality of the evaluated seed lots. The lower EC observed in treated seeds may be associated with the combined effects of the phytohormones and the B and Mo association on seed physiological performance, potentially contributing to membrane stability during imbibition and, consequently, reducing electrolyte leakage.
The tetrazolium test is widely recognized as a robust and reliable method for assessing seed viability and vigor, providing detailed information on tissue integrity and metabolic activity (Silva et al., 2025). For the cultivar M5917 IPRO, the highest Viability Tz values were obtained with treatments 4, 5 and 6 (Figure 2H). These results indicate greater preservation of embryonic tissue integrity and, consequently, enhanced maintenance of physiological potential, an essential prerequisite for rapid and uniform seedling establishment under field conditions.
Regarding plant vigor, treatment performance was cultivar-dependent (Figure 2). For 95R40 IPRO, treatment 1 promoted the highest vigor expression (Figure 2C), suggesting a positive effect on metabolic efficiency and tissue functionality. In M5917 IPRO, however, treatments 4 and 5 were associated with superior vigor performance (Figure 2I), reinforcing the genotype-specific nature of treatment responses. Conversely, in the cultivar Desafio RR, the control treatment exhibited the highest Vigor Tz values (Figure 2F), indicating that product application did not enhance, and may not have contributed to, the maintenance of seed vigor in this genetic background.
The tetrazolium assay allows indirect estimation of respiratory activity in seed tissues, as staining intensity and pattern are directly associated with membrane integrity and dehydrogenase enzyme activity in viable cells (Silva et al., 2026). From this perspective, the differential responses observed among cultivars underscore the decisive role of genotype in determining soybean seed physiological quality. Each cultivar exhibited its maximum response under a distinct treatment, demonstrating a lack of convergence in product efficacy and reinforcing the importance of genotype-specific management strategies.
Seeds of the cultivars 95R40 IPRO and Desafio RR presented high initial germination percentages, which were maintained throughout the storage period (Figure 3), indicating intrinsic physiological stability. Nevertheless, product application did not represent an effective strategy for preserving germination capacity over time, as no significant differences were detected between treated seeds and the control. These findings are consistent with those reported by Domingos et al. (2021), who concluded that foliar application of Ca- and B-based fertilizers does not promote detectable changes in soybean seed physiological quality when evaluated by standard germination tests. Collectively, these results reinforce that improvements in vigor detected by biochemical assays do not necessarily translate into measurable gains in germination performance, particularly when baseline seed quality is already high.
Ninety-five percent confidence intervals (95% CIs) for germination (%), in freshly harvested soybean seeds and after three and six months of storage, from cultivars subjected to plant bioregulator applications.
In contrast, germination of the cultivar M5917 IPRO under treatment 6 was initially low, declined markedly after three months of storage, and remained low after six months of storage. The minimum germination standard required for the commercialization of soybean seeds in Brazil is 80%, according to Normative Instruction No. 45 (Brasil, 2013). Under these conditions, germination fell below the regulatory threshold after treatment 6 application (Figures 3C, 3F, 3I). This response contrasts with previous findings in which foliar application of N and B improved soybean seed germination and seedling growth (Julita et al., 2016). However, the response to B nutrition may vary among soybean genotypes, as different varieties have shown distinct responses to B application in terms of seed physiological quality (Gomes et al., 2020).
Although the nutritional status of the mother plant directly influences seed quality, it is important to note that this fertilizer is recommended primarily to strengthen the cell wall and enhance flower retention, rather than specifically to improve seed physiological performance.
The reduction in germination capacity during seed aging is associated with reserve depletion and progressive loss of cellular integrity. Soybean seed longevity results from the interaction among seed coat characteristics, membrane stability, antioxidant defense systems, and genetic factors. High germination rates are essential for successful crop establishment, whereas reduced germination leads to uneven and less competitive seedlings, ultimately compromising plant stand and grain yield (Reed et al., 2022).
Changes in accelerated aging (AA) classification were observed as physiological quality was evaluated over the storage period (Figure 4). Although no statistically significant differences were detected among cultivars as a function of storage time, M5917 IPRO consistently exhibited lower vigor in freshly harvested seeds, a pattern that persisted throughout storage.
Ninety-five percent confidence intervals (95% CIs) for accelerated aging (%), in freshly harvested soybean seeds and after three and six months of storage, from cultivars subjected to plant bioregulator applications.
Seed vigor is primarily determined by three major factors: genetic background, environmental conditions during seed development, and storage conditions. Among the various contributors to seed deterioration, genetic factors play a pivotal role (Pirredda et al., 2024). Consequently, certain genotypes intrinsically exhibit higher physiological quality and maintain it for longer periods, even when exposed to identical environmental and storage conditions.
The cultivar M5917 IPRO is characterized by a high thousand-seed weight (Scarton et al., 2025), tolerance to mechanical damage (Bomfim et al., 2019), and elevated yield potential (Rodrigues et al., 2026). In this context, potential correlations among these traits may be hypothesized.
TSW presents a positive correlation with grain yield, and oil content has also been recurrently associated with higher productivity (Duan et al., 2023). However, seeds with higher oil content tend to be more susceptible to oxidative rancidity, which constitutes a major factor contributing to quality loss during storage. Additionally, tolerance to mechanical damage has been linked to higher lignin content in the seed coat (Menino et al., 2023). Nevertheless, increased lignin concentration does not necessarily imply superior physiological seed quality, as demonstrated by Castro et al. (2024).
Treatment 3 resulted in the highest estimated emergence speed index (ESI) in seeds of the cultivar M5917 IPRO after three months of storage (Figure 5F). These findings indicate a positive effect of the treatment on maintaining physiological vigor during the storage period.
Ninety-five percent confidence intervals (95% CIs) for emergence speed index, in freshly harvested soybean seeds and after three and six months of storage, from cultivars subjected plant bioregulator applications.
Treatment 3 is composed of cobalt (Co) and molybdenum (Mo), elements of particular importance for soybean due to their role in biological nitrogen fixation. The positive response observed may be related to the role of Co in promoting endogenous ethylene synthesis, which has been associated with enhanced germination and seedling development. In agreement with this mechanism, Abreu-Junior et al. (2023) reported that foliar application of Co and Mo during the reproductive stage of soybean increased germination rates and enhanced seed vigor and growth compared to untreated plants.
Overall, the results indicate that the application of plant bioregulators influences the physiological quality of soybean seeds. However, the magnitude and direction of the response depend on the cultivar, the product applied, and the storage duration.
CONCLUSIONS
Plant bioregulators affected the physiological quality of soybean seeds, with responses varying according to cultivar, product, and storage period.
Effects were more evident in vigor-related traits, particularly in freshly harvested seeds and after three months of storage; however, the magnitude and direction of responses varied among cultivars and treatments.
Bioregulator application did not consistently improve seed physiological quality, and the untreated control showed superior performance for tetrazolium-based viability in the Desafio RR cultivar.
In general, these plant bioregulators do not exert consistent effects on seed storability, indicating that their application does not provide effective protection against deterioration during.
ACKNOWLEDGEMENTS
The authors acknowledge the National Council for Scientific and Technological Development (CNPq) and the Coordination for the Improvement of Higher Education Personnel (CAPES) for their financial support, as well as the Research Support Foundation of the State of Minas Gerais (FAPEMIG) for the doctoral scholarship granted.
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