Does the chemical treatment of soybean seeds with cracked coat influence the quality after seed storage?

O tratamento químico de sementes de soja com rasgo no tegumento influencia a qualidade após o armazenamento?

ABSTRACT

The increased incidence of seed coat cracking (SCC) in soybean seeds has been frequently reported in production fields. However, the impact of SCC on seed quality, especially in the context of phytosanitary treatment and during storage, remains largely unexplored. This study evaluates the tolerance of SCC-affected seeds with seed treatment during storage in the seed quality. The experimental design was a completely randomized triple factorial analysis (2 x 4 x 3) involving two seed treatments (with and without), four levels of SCC (0%, 10%, 20%, and 40%), and three storage periods (0, 60, and 120 days). Two soybean cultivars, BMX Desafio and P96R70, were treated with a fungicide and two insecticides and evaluated for germination and vigor. It was found that seed treatment had a negative effect on the BMX Desafio cultivar, observed mainly in lots with SCC and after 60 days of storage. Susceptibility to seed deterioration with SCC was observed in the P96R70 cultivar. Seed vigor deterioration accelerates after 60 days of storage. Despite the impact of SCC on the P96R70 cultivar, its effect on physiological quality during a 120-day storage period is minimal in lots with up to 40% SCC, suggesting that SCC may not be a limiting factor up to this level. The results reinforce the importance of considering genotype, SCC level, and storage time in post-harvest management, in addition to providing input for breeding programs aimed at SCC tolerance.

Index terms:
Glycine max L.; seed quality; seed coat damage; phytotoxicity

RESUMO

O aumento da incidência de rasgo no tegumento de sementes de soja (SCC) tem sido frequentemente relatado em campos de produção. Contudo, o impacto desse defeito na qualidade da semente, especialmente no contexto do tratamento fitossanitário e durante o armazenamento, ainda é pouco explorado. Este estudo avaliou a tolerância de sementes com diferentes níveis de SCC submetidas ao tratamento e armazenamento, com base na qualidade fisiológica. O delineamento experimental foi inteiramente casualizado em esquema fatorial triplo (2 x 4 x 3), envolvendo dois tratamentos de sementes (com e sem), quatro níveis de SCC (0%, 10%, 20% e 40%) e três períodos de armazenamento (0, 60 e 120 dias). Foram utilizadas as cultivares BMX Desafio e P96R70, tratadas com um fungicida e dois inseticidas, sendo avaliadas quanto à germinação e vigor. Observou-se que o tratamento de sementes reduziu a qualidade fisiológica da cultivar BMX Desafio, especialmente em lotes com SCC após 60 dias de armazenamento. Já na cultivar P96R70, a suscetibilidade à deterioração foi mais evidente nos lotes com SCC, com acentuada perda de vigor após 60 dias. Apesar disso, em até 40% de SCC, o impacto sobre a qualidade fisiológica ao longo de 120 dias foi limitado, sugerindo que o defeito não é necessariamente restritivo nesse nível. Os resultados destacam a necessidade de considerar o genótipo, a intensidade do SCC e o tempo de armazenamento no manejo pós-colheita. Além disso, fornecem subsídios para programas de melhoramento voltados à tolerância ao SCC e ao tratamento de sementes.

Termos para indexação:
Glycine max L.; qualidade de sementes; dano no tegumento de sementes; fitotocixidade

Introduction

The quality of soybean seeds is a critical determinant of establishing robust and uniform seedlings, directly impacting crop productivity. The demand for high-performance soybeans and market competitiveness has led to the release of new cultivars, some of which are more susceptible to seed coat cracking (SCC) due to their genetic makeup, seed attributes and environmental conditions during seed filling (Lemes & Catão, 2024).

The emergence of SCC caused concern in the production of the soybean chain because it compromises the seed coat, which protects the embryo from physical damage and pathogens, as well as regulates the germination process. This vulnerability can reduce physiological quality - viability, germination, and vigor - during storage (Marcos-Filho, 2015). Furthermore, the severity of SCC is influenced by environmental and genetic factors, including climatic variations and cultivar susceptibility (Bahry et al., 2015; Teixeira, Zorato, & Meneghello, 2024).

Industrial Seed Treatment (IST) has gained popularity for its benefits, such as precise application of phytosanitary products, logistical efficiency, and process standardization (Reis et al., 2023; Rocha et al., 2024) performed by the seed companies industry, and OnFarm treatment, performed by the producer. The objective of this research was to compare the influence of industrial and OnFarm soybean seed treatment technologies, including the machines and processes, on the occurrence of mechanical damage and the functional quality of treatment. Soybean seeds were subjected to phytosanitary treatments with different processes and machines: (1. By 2028, 58% of soybean seeds in Brazil are expected to undergo IST (França-Neto et al., 2024). However, IST poses challenges, including potential phytotoxicity of the phytosanitary products, a condition where the treatment chemicals can harm the seeds, reducing their physiological quality during storage (Rocha et al., 2025; Toni et al., 2024).

The interaction between SCC and IST remains unclear and warrants urgent and thorough investigation. SCC is linked to genetic factors, such as genes that promote epidermal and hypodermal tissue separation, exposing the parenchyma and potentially increasing phytotoxicity risks while storing treated seeds (Chu et al., 2023)pathogen infection, water absorption and seed storage in soybean. In view of this, a recombinant inbred line (RIL). This phenomenon often occurs under high soil water availability, accelerating cell expansion during seed filling (Lemes & Catão, 2024; Senda et al., 2018). Moreover, early-maturing cultivars are more frequently susceptible to SCC, because the quantitative trait loci for this trait are associated with maturity loci (Kang et al., 2020).

Although SCC may not immediately affect seed quality, its long-term impact on storage durability is concerning (Teixeira, Zorato, & Meneghello, 2024). The application of phytosanitary products to seeds exhibiting seed coat cracking (SCC) accelerates their physiological deterioration, leading to reduced germination and vigor, which in turn compromises crop establishment and final productivity.

The objective of this study was to determine the tolerance threshold of seeds to SCC during storage. Specifically, we aimed to identify the level of SCC that negatively affects seed physiological quality. Establishing this threshold will provide critical information for developing more resistant soybean cultivars, improving seed treatment practices, and creating clear quality guidelines for seed production companies.

Material and Methods

Seeds from two soybean cultivars with reported coat cracking, differing in maturity groups (MG) and coat lignin content, were used: P96R70 (MG 6.7, 50.07 mg g⁻¹ lignin) and BMX Desafio (MG 7.4, 45.92 mg g⁻¹ lignin). Both were produced under identical soil and climate conditions at the Centro de Desenvolvimento e Transferência de Tecnologia (CDTT) in Ijaci, Minas Gerais, Brazil (Figure 1).

Figure 1:
Meteorological data on rainfall and average daily temperatures at the station Lavras, Minas Gerais, Brazil, during the soybean crop cycle of the cultivars P96R70 and BMX Desafio.

All cultivation and post-harvest processes were standardized. Manual harvesting occurred at 15% moisture content, followed by threshing using a stationary electric plot thresher (cylinder and concave). Seeds were naturally dried to 12% moisture, then size-classified (6 mm). After grading, seeds were sorted based on coat cracking presence or absence. Samples with and without cracking were weighed and processed using the IST process.

The seeds were treated using one fungicide and two insecticides (Table 1), supplemented with polymer (Biocroma vermelho®, 100 mL·100 kg^-1 seeds) and a finishing powder (Biogloss®, 200 g ·100 kg^-1 seeds). Untreated seeds were also maintained, creating four distinct sample groups: (1) treated without SCC, (2) treated with SCC, (3) untreated without SCC, and (4) untreated with SCC.

Table 1:
Description of the specific products that constitute the seed treatment.

Active ingredient (a.i.)	Concentration a.i.	Type	Dose of commercial product
Thiamethoxam	350 g· L-1	Insecticide	200 mL·100 kg-1 seeds
Cyantraniliprole	600 g· L-1	Insecticide	60 mL·100 kg-1 seeds
Metalaxyl-M	20 g· L-1	Fungicide	100 mL 100 mL·100 kg-1 seeds
Tiabendazole	150 g· L-1
Fludioxonil	25 g· L-1

Based on the findings of Teixeira, Zorato and Meneghello (2024) four levels of SCC incidence - 0%, 10%, 20%, and 40% - were selected from the four seed samples described above. These levels were established during the setup of seed physiological quality assessment tests, ensuring the exact percentage of seeds with SCC matched the specified incidence, improving analysis accuracy and precision.

The seeds were stored in a controlled environment with alternating temperatures: 30 ± 1 °C under light for 12h and 20 ± 1 °C in darkness for 12h (to simulate storage conditions at seed retailers), using permeable kraft packaging. Evaluations were conducted at three storage periods (SP), to represent the average time that seed lots wait to be sown: immediately after treatment (0 days), 60 days, and 120 days.

Physiological quality assessments were conducted at the Laboratório Central de Pesquisa em Sementes of the Departamento de Agricultura, Escola de Ciências Agrárias de Lavras (ESAL), Universidade Federal de Lavras, using the following tests:

a) Germination: four replicates of 50 seeds per treatment were sown between paper towels moistened with distilled water (2.5 times the substrate weight). Rolls were prepared and placed in a Mangesdorf germinator at 25 ± 2 °C. Results, expressed as the percentage of normal seedlings, followed International Seed Testing Association (ISTA, 2025) guidelines.

b) Accelerated aging: boxes with aluminum mesh were used, each containing 40 mL of distilled water. Seeds were distributed in a single layer on the mesh and placed in a BOD chamber at 41 ± 0.3 °C for 48 hours. After aging, seeds were subjected to the germination test (ISTA, 2025). Normal seedlings were evaluated five days post-sowing, with results expressed as a percentage.

c) Initial seedling development: three replicates of 20 seeds were tested following germination test methodologies (ISTA, 2025). Seedling images were captured four days post-sowing using the GroundEye® system (version S120). Seedlings were placed in the capture module tray for high-resolution imaging. The CIELab color model was used for background calibration (luminosity index: 0-100, dimension “a”: 13.9-46.1, dimension “b”: -57.1 to -40.6). Automated analysis extracted average values for total seedling length (Reis et al., 2022).

The experimental design was a completely randomized three-way analysis (2 x 4 x 3), evaluating two seed treatments (with and without phytosanitary treatment), four levels of SCC (0%, 10%, 20%, and 40%), and three SP (0, 60, and 120 days). Data were analyzed using analysis of variance (ANOVA) with the F-test (p < 0.05). When significant differences were detected (p < 0.05), means were compared using the Tukey test. All statistical analyses were performed in RStudio with the ExpDes package (R Core Team, 2022). For SCC levels showing statistical significance (p < 0.05), regression analyses were conducted to identify the best-fit model, prioritizing the highest coefficient of determination and biological relevance.

Results and Discussion

A three-way interaction was identified among the factors under investigation concerning seed germination. No significant differences in seed treatments (ST) were observed regarding SCC levels. However, for seeds without ST and those exhibiting 40% SCC, a decline of up to 12 percentage points (p.p.) was noted after 60 days of storage (Figure 2).

Figure 2:
Percentage of normal soybean seedlings in the germination test for cultivar P96R70 as a function of different SP, ST, and SCC levels. Averages followed by the same letter, upper case between SCC in the same SP and ST, lowercase between SP in the same ST and SCC level, * between ST in the same SP and SCC level, do not differ by Tukey’s test (p > 0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

Concerning the ST factor, after 120 days of storage, the lot with 40% SCC and without ST demonstrated a decrease in quality, reflected by a 5 p.p. inferiority compared to the equivalent lot with ST. The storage analysis indicated no reduction in germination for lots lacking SCC throughout the storage period (SP), regardless of ST. Nevertheless, a decline was noted at 60 days of storage in lots with more than 10% SCC, particularly affecting those treated with 10% and 40% SCC and the untreated lots with 40% SCC (Figure 2).

Accelerated aging test results revealed a two-way interaction between SCC levels and ST, as well as an isolated effect of the SP factor. The statistical analysis indicated no significant difference in SCC levels between treated and untreated seeds. However, the lots that did not undergo ST showed a marked reduction in vigor, with the most pronounced decline of 11 p.p. observed when comparing the lot with 40% SCC to that with 0% SCC (Figure 3a).

When the storage factor was analyzed independently of the other variables, a reduction in vigor was observed at 120 days (Figure 3b), which was anticipated given the known deterioration process of soybean seeds during storage.

Figure 3:
Percentage of normal soybean seedlings in the accelerated aging test for cultivar P96R70. A - This is a function of different ST and SCC levels. B - As a function of different SP. A -* Averages followed by the same upper case letter between SCC levels in the same ST and lowercase letter between ST in the same SCC levels do not differ according to Tukey’s test (p > 0.05). B - * Averages followed by the same letter do not differ according to Tukey’s test (p > 0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

A three-way interaction between the factors was also observed for seedling length (SL). For the lots with ST, there was a discernible difference in SCC levels at both 0 days and 60 days, though this effect was not observed at 120 days. During the SP, a reduction in SL was observed in all lots, irrespective of the SCC level, from 60 days onwards. However, no such reduction was evident in the lot treated with 40% SCC at 60 days of storage (Figure 4).

Figure 4:
Total length of normal soybean seedlings by germination test for cultivar P96R70 as a function of different SP, ST, and SCC levels. Averages followed by the same letter, upper case between SCC in the same SP and ST, lowercase between SP in the same ST and SCC level, * between ST in the same SP and SCC level, do not differ by Tukey’s test (p > 0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

Conversely, examining the lots without ST revealed a discrepancy in the SCC levels across all storage intervals, accompanied by a decline in SL after 60 days, similar to the observations made in the lots with ST. Moreover, a distinction was observed between the lots with and without ST, specifically in the 20% SCC level, at both 0 and 120 days of storage (Figure 4).

A linear regression analysis of the isolated effect of SCC levels reveals a reduction in the physiological quality of lots for both germination and accelerated aging variables, with an approximate 1.3% and 1.7% reduction in normal seedlings, respectively, for each 10% increase in SCC level in the lot (Figure 5).

Figure 5:
Regression of the percentage of normal soybean seedlings in the germination (G) and accelerated aging (AA) test for cultivar P96R70 as a function of the simple effect of different SCC levels.

No statistically significant difference was observed in the isolated effect of SCC levels (Figure 6a) for the BMX Desafio cultivar. Conversely, a two-way interaction was identified for the storage periods and ST variables affecting germination. Following storage, ST likely caused phytotoxicity, thereby reducing the germination of seed lots with ST at 60 and 120 days by up to 9 p.p. Moreover, storage reduced germination irrespective of ST during storage (Figure 6b).

Figure 6:
Percentage of normal soybean seedlings by germination test for the BMX Desafio cultivar. A -As a function of different SCC. B - As a function of different SP and ST. A - * Averages followed by the same letter do not differ according to the Tukey’s test (p > 0.05). B - * Averages followed by the same upper case letter between SP in the same ST and lower case between ST in the same SP do not differ according to Tukey’s test (p > 0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

The accelerated aging test for the BMX Desafio cultivar revealed an interaction between SCC levels, ST, and storage periods. Despite the SP, SCC levels were insignificant except for 0% at 0 days. However, seed lots’ vigor decreased to up to 19 p.p. at 120 days of storage (Figure 7a). When comparing ST and storage periods, it was observed that vigor was reduced by 8 and 20 p.p., respectively, at 120 days of storage, regardless of seed treatment. However, vigor was better maintained when ST was used (Figure 7b).

Figure 7:
Percentage of normal soybean seedlings in the accelerated aging test for the BMX Desafio cultivar. A - As a function of different SP and SCC levels. B - As a function of different SP and ST. A - * Averages followed by the same upper case letter between SCC levels in the same SP and lower case between SP in the same SCC level do not differ by the Tukey’s test at 5% (p > 0.05). B - * Averages followed by the same upper case letter between SP in the same ST and lower case between ST in the same SP do not differ by Tukey’s test (p > 0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

There was also a three-way interaction between the factors for SL. SCC levels differed after 120 days of storage for the lots with ST, with the lot with 40% SCC having the highest SL. During the storage periods, there was a reduction in SL regardless of the level of SCC from 60 days (Figure 8), as was also observed for cultivar P96R70 (Figure 4).

Figure 8:
Total length of soybean seedlings of the BMX Desafio cultivar as a function of different SP, ST, and SCC levels. Averages followed by the same letter, upper case between SCC in the same SP and ST, lowercase between SP in the same ST and SCC level, * between ST in the same SP and SCC level, do not differ by Tukey’s test (p >0.05). SP = storage period, ST = seed treatment, and SCC = seed coat cracking.

For the lots without ST, there was a reduction in SL after 60 days of storage, and after 120 days, there was no difference between the SCC levels. A decrease in SL was also observed during the storage periods, irrespective of the SCC levels. The reduction in SL due to the ST factor was only observed in the treatments with 40% SCC at 60 days and 20% SCC at 120 days (Figure 8).

The integrity of the coat is essential for maintaining seed quality, as its primary function is to protect the embryo from environmental fluctuations and pathogen attacks (Radchuk & Borisjuk, 2014). In addition, the seed coat affects various metabolic processes, including gas exchange and intercommunication between the internal and external tissues of the seed. This highlights the importance of the seed coat in regulating seed germination and the intrinsic mechanisms that control seed development and dormancy (Marcos-Filho, 2015).

The seed coat is composed of four distinct layers: the cuticle, epidermis (palisade cells), hypodermis (osteosclereids), and parenchyma (parenchymatous cells), which extend from the surface to the interior of the seed (Brzezinski et al., 2022). These layers protect the embryo from mechanical damage, weathering deterioration damage, and pathogens.

SCC is a recent and ongoing problem in Brazil and Argentina caused by genetic and environmental factors that compromise the integrity of the seed’s outer layer. Genetic factors contribute significantly to the presence of SCC, as evidenced by the different levels of susceptibility observed in soybean cultivars due to the influence of maternal genes (Saruta et al., 2019). The genetic basis of SCC in soybean has been evaluated in quantitative trait locus studies and diversity panels, and although candidate genes such as T, IC, Glyma.06G197300, and Glyma.06G202100 have been proposed, no clear underlying mechanism has been identified (Chu et al., 2023).

In addition, some cultivars, especially those with early maturity, are more susceptible to this condition, suggesting the effect of maturity loci, especially the E2 locus for variation in SCC associated with maturity (Kang et al., 2020). These varieties are often exposed to water stress or temperature fluctuations during critical stages of seed development, which may explain the higher incidence of SCC.

Another factor genetic that may be associated with SCC is lignin, an essential component of the cell wall, which contributes to the rigidity and resistance of the seed coat to mechanical damage (Krzyzanowski, França-Neto, & Henning, 2023). It was hypothesized that cultivars with a higher lignin content in the seed coat would demonstrate greater tolerance to SCC and, consequently, maintain their physiological quality during storage (Senda et al., 2017).

However, the results of this study did not corroborate this hypothesis. The cultivar P96R70, which has a higher concentration of lignin in the tegument (4.15 mg g⁻¹ more than BMX Desafio), demonstrated a reduction in physiological quality due to the presence of SCC.

For environmental factors, SCC results in an imbalance in the growth of cotyledon and tegument cells, especially in conditions of heavy rainfall followed by periods of drought. When these seeds reach physiological maturity (R7 stage) in the field, the presence of SCC can make them more vulnerable to moisture deterioration. During storage, this susceptibility increases, and depending on the location of the cracks, the tissues essential for the emergence of the radicle can be compromised, reducing the viability of the seed (Teixeira, Zorato, & Meneghello, 2024).

The findings of this study reveal that the two cultivars exhibited distinct behaviors during the storage period. The presence of SCC negatively affected the germination and vigor of P96R70 seeds. In contrast, the BMX Desafio cultivar showed no significant difference in germination and vigor related to SCC levels.

Existing evidence suggests that SCC significantly impacts the physiological quality of seeds. Seeds with SCC demonstrate an increased capacity to absorb water, accelerating deterioration and reducing physiological quality during storage (Machado et al., 2019). Furthermore, these effects are intensified in seeds stored for prolonged periods, where oxidative stress and fungal contamination - such as by Fusarium sp. and Aspergillus sp. - lead to substantial quality deterioration (Machado et al., 2019). This damage is linked to reduced vigor, particularly after more than 6 months of storage and in lots with 100% of SCC, especially in tropical environments with elevated temperatures (Teixeira, Zorato, & Meneghello, 2024).

However, these outcomes of this study may be attributed to SCC levels and the cultivars’ tolerance to storage, with BMX Desafio showing greater tolerance, as discussed by Nardelli et al. (2025). Additional research is necessary to understand better seed quality about these factors.

Furthermore, the behavior of seedlings in lots with varying levels of SCC was examined. It was observed that plots with a high incidence of SCC exhibited greater SL under specific conditions, which may appear to be contradictory given the preceding discussion. However, the increased permeability resulting from SCC can accelerate initial soaking, thus promoting initial seedling growth (Ma, 2004; Machado et al., 2019)2019.

However, this advantage in initial seedling growth is only temporary. After 60 days of storage, as observed in the P96R70 cultivar, rapid soaking contributes to possible soaking damage and reduced seedling length, thus compromising long-term physiological quality. The natural deterioration of post-harvest seeds, associated with the high permeability of the coat, potentiates this damage (Teixeira, Zorato, & Meneghello, 2024).

While there is a correlation between SCC and moisture content, it is not a causal relationship. Although the process can originate in the cotyledons, prolonged storage and the water content of the seeds at the time of sowing can facilitate the rupture’s propagation to the end of the central cylinder. Such conditions can impact the viability of the embryo cells, which are integral to the emergence of the radicle (Huth et al., 2016).

The deterioration of soybean seeds results in oxidative stress, compromising their physiological quality. This negatively impacts germination by reducing the seeds’ ability to produce normal seedlings and making the seedlings more susceptible to environmental stresses (Jiamtae et al., 2023; Silva et al., 2023).

The findings of this study suggest that seed lots with a high prevalence of SCC are susceptible to accelerated deterioration, particularly at elevated temperatures. Seeds stored at elevated temperatures (approximately 30 °C) exhibit accelerated deterioration, negatively impacting antioxidant activity and physiological quality (Jiamtae et al., 2023; Toni et al., 2024). This effect is particularly pronounced in cultivars that are more sensitive to storage conditions and have SCC, rendering them more susceptible to fluctuations in the external environment, as corroborated by the accelerated aging test (Nardelli et al., 2025).

The cultivars’ response to ST varied throughout storage. For the BMX Desafio cultivar, ST reduced germination at 60 and 120 days, regardless of the presence of SCC. In addition, SL was only affected in lots with SCC. In contrast, the P96R70 cultivar showed no reduction in physiological quality due to ST, showing greater tolerance to the treatment. The deterioration observed in the seed lots was more pronounced over time, particularly in those that underwent phytosanitary treatment, which may have caused phytotoxicity.

This sensitivity of cultivar BMX Desafio is because certain cultivars react unfavorably to specific active ingredients in commercial seed treatments (Rocha et al., 2025; Toni et al., 2024). In the case of the BMX Desafio cultivar, sensitivity to ST resulted in the deterioration of physiological quality. Therefore, selecting appropriate commercial products and doses is crucial to mitigate adverse effects and ensure the effectiveness of ST.

Toni et al. (2024) noted that the decline in germination and vigor of soybean seeds during storage, when treated with thiamethoxam, may be attributed to the natural deterioration of the seeds over time, coupled with the genetic characteristics of the cultivar. The effects of this active ingredient can lead to potential phytotoxicity, whether applied alone or in conjunction with soybean seeds. They found that this phytotoxicity might be linked to the combination of liquid polymer and finishing powder with the fungicide and insecticide. This corroborates the present study, which used thiamethoxam, polymer and finishing powder.

Conclusions

Seed treatment reduced the quality of BMX Desafio, mainly in SCC lots after 60 days of storage. Genetic factors influenced deterioration, with early-maturing varieties like P96R70 being more susceptible. Vigor declined faster after 60 days, but SCC impact up to 40% was limited over 120 days, indicating it is not a restrictive factor at this level. Results highlight the importance of genotype, SCC intensity, and storage in management, supporting breeding and seed treatment strategies for greater tolerance.

Acknowledgements

The authors would like to extend their sincere gratitude to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG), and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for their generous financial support, including scholarships and a research productivity grant. Also, thanks to Embrapa Soja for conducting the lignin content analysis and to the Seedcare Institute for their generous product donations.

Data Availability Statement

Data available upon request to authors.

References

• Bahry, C. A. et al. (2015). Chemical composition and structural characterization of contrasting colors of soybean seed coats. Semina: Ciências Agrárias, 36(3):1913-1926.
• Brzezinski, C. R. et al. (2022). Structural analysis of soybean pods and seeds subjected to weathering deterioration in pre-harvest. Pesquisa Agropecuária Brasileira, 57:e02697.
• Chu, J. et al. (2023). Mining of seed coat cracking related genetic loci and genes in diverse soybean (Glycine max) populations. Plant Breeding, 142(2):170-183.
• França-Neto, J. de B. et al. (2024). Adoção do tratamento de sementes de soja na indústria 21p. Embrapa. (Boletim de Pesquisa e Desenvolvimento, N. 37).
• Huth, C. et al. (2016). Susceptibility to weathering damage and oxidative stress on soybean seeds with different lignin contents in the seed coat. Journal of Seed Science, 38(4):296-304.
• International Seed Testing Association - ISTA. (2025). International Rules for Seed Testing International for Seed Testing, 320p. https://doi.org/10.15258/istarules.2025.F
  » https://doi.org/10.15258/istarules.2025.F
• Jiamtae, P. et al. (2023). Biochemical changes in recombinant inbred lines of soybean (Glycine max L. Merr.) under storage-related stresses. Journal of Crop Science and Biotechnology, 26:467-488.
• Kang, B. K. et al. (2020). High-density linkage map reveals QTL for Type-I seed coat cracking in RIL population of soybean [Glycine max (L.) Merr.]. Euphytica, 216:153.
• Krzyzanowski, F. C., França-Neto, J. D. B., & Henning, F. A. (2023). Importance of the lignin content in the pod wall and seed coat on soybean seed physiological and health performances. Journal of Seed Science, 45:e202345006.
• Lemes, E. M., & Catão, H. C. R. M. (2024). Soybean seed coat cracks and green seeds-predisposing conditions, identification and management. Seeds, 3(1):133-148.
• Ma, F. (2004). Cracks in the palisade cuticle of soybean seed coats correlate with their permeability to water. Annals of Botany, 94(2):213-228.
• Machado, B. R. et al. (2019). Effect of tear/crack on soybean (Glycine max) seed coat, physiological quality and pathology of the seed. Australian Journal of Crop Science, 13(06):988-994.
• Marcos-Filho, J. (2015). Seed physiology of cultivated plants. Associação Brasileira de Tecnologia de Sementes - Abrates.
• Nardelli, A. C. P. et al. (2025). Lignin content, morphoanatomical traits and antioxidant activity as determinants of soybean seed storage tolerance and quality. Acta Scientiarum. Agronomy, in press.
• R Core Team. (2022). R: A language and environment for statistical computing R Foundation for Statistical Computing. Available in: <https://www.R-project.org/>.
  » https://www.R-project.org/
• Radchuk, V., & Borisjuk, L. (2014). Physical, metabolic and developmental functions of the seed coat. Frontiers in Plant Science, 5:1-17.
• Reis, L. V. et al. (2023). Treatment technologies for soybean seeds: Dose effectiveness, mechanical damage and seed coating. Ciência e Agrotecnologia, 47:e013622.
• Reis, V. U. V. et al. (2022). Vigor of maize seeds and its effects on plant stand establishment, crop development and grain yield. Journal of Seed Science, 44:e202244020.
• Rocha, D. K. et al. (2024). Dose effectiveness and quality of soybean seed treatment in Brazilian agriculture as a function of application technology. Journal of Seed Science, 46:e202446018.
• Rocha, D. K. et al. (2025). How do the components used in chemical seed treatment affect physiological quality over the storage period? Bragantia, 84:e20240131.
• Saruta, M. et al. (2019). A QTL associated with high seed coat cracking rate of a leading Japanese soybean variety. Breeding Science, 69(4):665-671.
• Senda, M. et al. (2018). Occurrence and tolerance mechanisms of seed cracking under low temperatures in soybean (Glycine max). Planta, 248(2):369-379.
• Senda, M. et al. (2017). Accumulation of proanthocyanidins and/or lignin deposition in buff-pigmented soybean seed coats may lead to frequent defective cracking. Planta, 245(3):659-670.
• Silva, M. F. D. et al. (2023). The role of the biochemical composition of soybean seeds in the tolerance to deterioration under natural and artificial aging. Heliyon, 9(12):e21628.
• Teixeira, S. B., Zorato, M. D. F., & Meneghello, G. E. (2024). Implications of seed coat cracking on soybean seed quality. Journal of Seed Science, 46:e202446030.
• Toni, J. R. et al. (2024). Quality assessment of soybean seeds submitted to industrial seed treatment and stored in a natural and controlled environment. Journal of Stored Products Research, 108:102372.

• Editor de seção:
  Renato Paiva

Publication Dates

• Publication in this collection
  16 Jan 2026
• Date of issue
  2025

History

• Received
  11 Apr 2025
• Accepted
  28 July 2025