Revista Ciência Agronômica, v.

56, e202292842, 2025

Centro de Ciências Agrárias - Universidade Federal do Ceará, Fortaleza, CE Scientific Article
www.ccarevista.ufc.br ISSN 1806-6690

Storage potencial of soybeans cultivars under low temperature1

Kiliany Arcia Moreno2*, Yamid Arley Mera-Velasco3, Edila Vilela De Resende VonPinho4, Heloisa Oliveira Dos Santos4

ABSTRACT - Currently with a more demanding market, is necessary to produce better and high physiological quality cultivars of

soybeans. In front of this new challenge, our objective was to evaluate the physiological quality of 11 soybean cultivars: CD201,

SYN1263, SYN1279, BMX, UFLA1, CA115, CD215, CD202, Conquista, Savana, and BRS820 storage for 12 months. Evaluations

were conducted through physiological like germination and vigor (accelerated aging and controlled deterioration) and isoenzymes

analysis. The seeds were stored under controlled conditions at 10 °C and 10% relative humidity. It was assessed every four months

(0, 4, 8, and 12). 200 seeds per treatment were used for each test, divided into 4 replications of 50 seeds. The number of normal

plants was evaluated on the fifth and seventh days, expressed as a percentage value. Isoenzyme analysis of MDH, ADH, Esterase,

and Catalase was made. The results were interpreted from the presence or absence of bands in the gel. An experimental design in

randomized complete blocks, interpreting data using analysis of variance in a factorial scheme 11 x 4 (11 cultivars and 4 times of

storage and averages compared by the test Scott-Knott 5% of probability and regression analysis. The statistical program used was

Sisvar®. We found that cultivars Savana and Conquista showed low physiological quality, and the cultivars CD 215 and BMX

showed high physiological quality during storage.

Key words: Glycine max. Isoenzymes. Soja. Tolerance. Crop development.

DOI: 10.5935/1806-6690.20250005
Editor-in-Chief: Prof. Alek Sandro Dutra - alekdutra@ufc.br
*Author for correspondence
Received for publication 09/06/2022; approved on 29/01/2024
1
This paper is extracted from the dissertation of the lead author presented to the Postgraduate Program in Engenheria Agronomica, Universidad Federal
de Lavras, Brasil. The authors wish to thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)
2
Departamento de Ciencias Agrarias, Universidad del Cauca (UNICAUCA), Popayán-CAU, Colombia, karcia@unicauca.edu.co (ORCID ID
0000-0002-0438-1782)
3
Departamento de biología, Universidad del Cauca (UNICAUCA), Popayán-CAU, Colombia, ymera@unicauca.edu.co (ORCID ID 0000-0002-5221-8824)
4
Departamento de Agricultura (DAG), Universidad Federal de Lavras (UFLA), Lavras-MG, Brasil, edila@dag.ufla.br (ORCID ID 0000-0003-3932-3926),
heloisasantos@dag.ufla.br (ORCID ID 0000-0003-1384-4969)
 KA.Arcia Moreno et al.

INTRODUCTION there is a need to understand the genetic control for these traits
(MEDEROS-RAMÍREZ; ORTIZ-PEREZ, 2021). Currently,
Soybean is the most important oilseed in the world, there is a concern in selecting soybean genotypes with higher
with Brazil being the second largest producer of this seed, storage potential while preserving their physiological quality
harvesting around 96.2 million tons per season. The quality until planting time. However, there are not many studies
control of soybean is crucial from its collection, chemical that relate different storage tolerance levels among soybean
composition, nutritional quality, vigor, processing, and cultivars (MAVAIEIE et al., 2019). The objective of this
storage (BISHT et al., 2015; MARTINS et al., 2018). research was to study the physiological quality of soybean
The quality of seeds directly reflects the crop seeds from 11 cultivars stored for 12 months in a cold chamber
development, leading to plants with high vigor, population and obtain an isoenzyme expression profile during storage.
uniformity, and absence of seed-transmitted diseases
(SILVA; LAZARINI; SÁ, 2010). Similarly, storage is a
fundamental practice to maintain physiological quality MATERIAL AND METHODS
and ensure the preservation of vigor and viability between
collection and planting (AZEVEDO et al., 2003), linked Study Area
to the preservation of physiological and sanitary quality by
The work was conducted in two stages, one in
reducing contamination, pest incidence, microorganisms,
the field and the other in the laboratory. The field phase
and minimizing deterioration. While the latter cannot
was carried out at the experimental vegetable station of
be prevented, its speed can be minimized through
Hortiagro Semillas Ltda., in the municipality of Ijaci
appropriate procedures of production, harvesting,
(Minas Gerais, Brazil), located 13 km northeast of the
drying, processing, and transport (DELOUCHE, 2002;
city of Lavras, at an elevation of 833 msnm, latitude:
FRANÇA NETO et al., 2010; GARCIA et al., 2004;
21° 9’ 24’’ South, and longitude: 44° 55’ 34’’ West.
KRZYZANOWSKI, 2015; MAVAIEIE et al., 2019).
Seed multiplication of 11 soybean cultivars (CD201,
High temperatures and the activity of SYN1263, SYN1279, BMX potencia, UFLA1, CA115,
microorganisms such as fungi and insects accelerate the MS8400, CD215, CD202, CONQUISTA, SAVANA)
respiratory processes of seeds, favoring the population was carried out under controlled field conditions.
growth of these organisms, thus increasing seed
Subsequently, the seeds were collected at the R8
deterioration over time (MARCOS-FILHO, 2005).
phenological stage (95% maturity and 18% moisture content),
Likewise, enzymes involved in slowing down the damage
and then dried until reaching 12% moisture content.
caused by respiratory processes have been documented.
Circular size sieves ranging from 5.55 mm to 6.35 mm
For example, alcohol dehydrogenase (ADH) is an enzyme
were used for seed processing. The seeds were weighed
related to anaerobic respiration, promoting the reduction
and stored in paper bags under controlled conditions
of acetaldehyde to ethanol. Acetaldehyde accelerates seed
in a cold chamber at the central seed laboratory of the
deterioration, therefore, increased ADH activity provides
Department of Agriculture of the Federal University of
protection against the deleterious action of this compound
Lavras (UFLA), located in Lavras municipality, Minas
(ZHANG et al., 2008). Malate dehydrogenase (MDH)
Gerais, with a latitude of 21º 14” S, longitude of 45º 00”,
plays a significant role in the Krebs cycle, catalyzing
and elevation of 918 msnm.
the conversion of malate to oxaloacetate and producing
NADH, which is a fundamental product in the production The experimental design employed was
of ATP and essential intermediate compounds in cellular randomized complete block design with replications. The
functioning (CARVALHO et al., 2014; MAVAIEIE physiological tests evaluated were germination and vigor
et al., 2019). The intracellular enzyme catalase (CAT) (controlled deterioration and accelerated aging). Each test
found in plant peroxisomes has the ability to transform was evaluated by counting normal plants, measurements
reactive oxygen species into harmless forms, such as the taken on the fifth and eighth day, and results expressed as a
breakdown of hydrogen peroxide (BAILLY et al., 2004). percentage. In each test, 200 seeds per cultivar were used,
Esterase (EST) is an enzyme involved in lipid breakdown divided into 4 replications of 50 seeds each, according to
during the germination process, being relevant in the the Seed Analysis Standard (BRASIL, 2009).
renewal of embryonic axis growth, especially in lipid- Germination and Vigor Test (controlled deterioration
rich seeds like soybean (MAVAIEIE et al., 2019). and accelerated aging)
The physiological quality of soybean seeds can For the physiological tests evaluation, the seeds
vary depending on the genotypes, and this characteristic were treated with the insecticide Vitavax Thiram (200 mL
is important during the selection process in breeding per 100 kg of seeds) and water (300 mL per 100 kg
programs (VERNETTI and VERNETTI, 1983). Therefore, of seeds). These tests were conducted at four different

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 Storage potencial of soybeans cultivars under low temperature

times, 0, 4, 8, and 12 months. Subsequently, they The statistical analysis of the obtained data was
were placed on moistened Germitest paper with water, performed using the Sisvar® program (FERREIRA, 2011),
equivalent to 2.5 times the weight of the dry substrate, through the analysis of variance of each test. The comparison
and kept at a temperature of 25 °C in accordance with of means was done using the Scott-Knott test, with a 5%
RAS (BRASIL, 2009) to ensure uniform humidity. probability. The design used was a completely randomized
Afterwards, each treatment with four rolls of Germitest block design in a factorial design of 11x4 (11 cultivars and 4
paper containing 50 seeds for a total of 200 seeds/treatment evaluation periods: 0, 4, 8, 12 months).
were placed in a germinator regulated at 25 °C, where two
evaluations were carried out by counting the number of
normal plants on the fifth and eighth day after sowing. RESULTS AND DISCUSSION
Accelerated aging test
Physiological Tests
Mini cameras of the “Gerbox” type were used
Significant differences (F3; 30 = 15.826; P = 0.0001,
according to the RAS (BRASIL, 2009), where 42 g of
p < 0.05) were found in the analysis of variance for the
seeds from each treatment were placed. The seeds were
physiological quality of soybean seeds in cultivars stored for
suspended on a grid, without coming into contact with
different periods in germination and controlled deterioration
the bottom of the box which previously contained 40 mL
tests on the fifth and eighth day, respectively. The cultivars
of distilled water. Subsequently, the mini cameras were
Savana, Conquista, and BRS 820 were found to have the
placed in a germination chamber at 42 ºC for 82 h, ensuring
lowest evaluated physiological quality (Table 1). According to
maximum stress. After this time, they were removed and
Baldoni (2013), these cultivars were also reported to have low
the germination test was carried out.
physiological quality when evaluated through germination
Controlled deterioration test tests and accelerated aging, with no significant differences
Aluminum envelopes coated with plastic and found (p > 0.05). This lack of differentiation may be due to
hermetically closed with 42 g of seeds from each the fact that the germination test does not provide information
treatment were used. These were kept in a water bath on the progress or potential for deterioration.
at 40 ०C for 48 h, following the protocol described by The evaluation period analyzed through regression
Marcos-Filho (2005). The seed moisture content was showed significant differences for germination tests (y
adjusted to 15% and placed in a cold chamber at 10 ºC/24 h. = -4,1667x3 + 32x2 - 72,833x + 141; R² = 1) (Figure 1a),
before being transferred to a germination chamber controlled deterioration (y = -9E-13x3 + 0,5x2 - 1,5x + 97;
at a temperature of 42 ºC/48 h. On the fourth day, a R² = 1) (Figure 1b), and accelerated aging (y = 1,8333x3
germination test was performed according to the RAS + 1,5x2 - 45,333x + 134; R² = 1) (Figure 1c). A slight
guidelines (BRASIL, 2009). decrease in the physiological quality of the seeds at 4
Isoenzymatic analysis months of storage can be considered, perhaps due to the
presence of fungus in the seed at the time of collection. This
After being harvested and processed, the seeds decrease, along with the passage of time and exposure to
were stored at -80 ºC and evaluated at 0, 4, 8, and 12 low temperatures in a cold room, caused a decrease in its
months. For gene expression analysis, 50 ground seeds proliferation, allowing the potential vigor of the different
were used with liquid nitrogen and polyvinylpyrrolidone soybean cultivars to be observed at 8 and 12 months.
(PVP) antioxidant and divided into Eppendorf tubes According to Mederos-Ramírez and Ortiz-Perez (2021),
with 100 mg each. Then, 300 uL of ethyl ether + 300 µL an increase in soybean seed germination has been found
of water were added to remove the oil. Subsequently, they after 2 and 3 months of storage at room temperature, due to a
were centrifuged for 20 minutes at 14000 rpm at 4 ºC. reduction in the incidence of field fungi on the stored seeds.
Enzyme extraction was performed with 250 µL of The results demonstrate efficiency in seed viability
buffer (0.2 M Tris HCl pH 8 + 0.1% beta-mercaptoethanol), when stored in a cold chamber, after 6 months of storage, as
followed by vortex agitation and overnight incubation. also observed by Mavaieie et al. (2019) and Martins-Filho
Afterward, the samples were centrifuged at 4 ºC at 14000 rpm et al. (2001), after 8 months of storage in a cold chamber,
for 30 minutes. In the running gel, 50 µL of supernatant the seeds of soybean cultivars were superior compared to
(separating gel - 7.5% polyacrylamide and concentrating those stored under non-controlled conditions). Analysis of
gel - 4.5% polyacrylamide) were applied. The buffer gel/ variance showed interaction between evaluation periods
electrode system used was Tris glycine pH 8.9. Runs were and cultivars on day 7 of the controlled deterioration test.
performed at 150 volts for 6 hours. After electrophoresis, Germination test data and seed image analysis were used
the enzyme was revealed following the protocol for each to perform an analysis of variance, and treatment means
one (ALFENAS, 2006). were compared using the Scott-Knott test, revealing

Rev. Ciênc. Agron., v. 56, e202292842, 2025 3

 KA.Arcia Moreno et al.

Table 1 - Mean germination (Ger), controlled deterioration (DC), and accelerated aging (EA) for the first and second count of seeds
from 11 soybean cultivars

Cultivar Ger. 5 Days Ger. 8 Days DC 5 Days DC 8 Days EA 5 Days EA 8 Days

Savana 86 c 89 c 93 b 95 b 77 a 84 a
Conquista 90 b 93 b 95 b 96 b 75 a 80 a
BRS820 92 b 94 b 97 a 98 a 85 a 88 a
BMX 96 a 96 a 98 a 98 a 80 a 83 a
CD202 97 a 97 a 97 a 98 a 81 a 86 a
UFLA 1 97 a 98 a 98 a 98 a 86 a 91 a
CD 201 97 a 97 a 99 a 99 a 84 a 87 a
SYN 1263 97 a 98 a 98 a 98 a 77 a 82 a
CD 215 97 a 97 a 98 a 99 a 74 a 81 a
CA 115 97 a 98 a 97 a 99 a 63 a 69 a
SYN 1279 98 a 98 a 99 a 99 a 71 a 73 a
CV% 5.05 4.43 2.74 1.84 22.77 19.19
* Means followed by the same letter do not differ from each other according to the Scott-Knott test at a 5% probability

Figure 1 - Physiological quality of 11 soybean cultivars seeds during four storage periods: 0, 4, 8, and 12 months; A. Results
corresponding to the fifth-day Germination tests. B. Eighth-day Germination tests. C. Controlled deterioration test on the fifth day

A.

B. C.

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 Storage potencial of soybeans cultivars under low temperature

significant differences (p ≤ 0.0001) (Table 2). An highlighting their high physiological quality and
interaction was observed in the first three storage periods commercial value in the market (BALDONI, 2013).
(months 0, 4, and 8) in the Savana and Conquista cultivars
with poorer physiological quality. The accelerated aging Seeds of the soybean cultivar BMX potency RR, have
test in different periods showed significant differences superior characteristics and maintain a minimum germination
(p < 0.05) in treatments, with the CA115 cultivar being the rate greater than 80% (JUVINO et al., 2014). The CD201
most affected in its physiological quality after 8 months of cultivar is a conventional cultivar. It has been selected
storage, where a noticeable decline in the physiological in different productivity and high physiological quality
quality of different cultivars can be observed (Table 3). studies, and it has been reported to have a germination
percentage greater than 92% during the first year of
Isoenzyme Analysis
production according to Ludwing et al. (2011), and 88% in
The selected cultivars for isoenzyme analysis, the second year with a productivity of 1,878 kg/ha. Thus, the
ranging from low to high quality, were Conquista, results obtained can be seen in Figure 2, where the isoenzyme
Savana, and the BMX and CD201 varieties, respectively. patterns of esterase, catalase, ADH, and MDH are shown for
Their selection was based on previous research reports, the four cultivars: CD201, BMX, Savana, and Conquista.

Table 2 - Mean values of the interaction of cultivars during the four storage periods

Cultivar 0 months 4 months 8 months

Savana 93 b 95 b 98 a
Conquista 98 a 94 b 91 b
BRS820 99 a 98 a 99 a
BMX 97 a 97 a 100 a
CD202 97 a 98 a 99 a
UFLA 1 99 a 97 a 99 a
CD 201 98 a 98 a 99 a
SYN 1263 99 a 97 a 99 a
CD 215 99 a 99 a 100 a
CA 115 98 a 97 a 100 a
SYN 1279 98 a 99 a 100 a
* Means followed by the same letter do not differ from each other according to the Scott-Knott test at a 5% probability

Table 3 - Mean interaction values at 8 months of storage for 11 soybean cultivars

Cultivar Cultivar X Time (8 Months)
CA 115 17 b
SYN 1279 55 a
BRS 820 70 a
BMX 72 a
SAVANA 73 a
CD 201 73 a
CONQUISTA 77 a
CD 215 77 a
SYN 1363 82 a
CD 202 87 a
UFLA 1 88 a
* Means followed by the same letter do not differ from each other according to the Scott-Knott test at a 5% probability level

Rev. Ciênc. Agron., v. 56, e202292842, 2025 5

 KA.Arcia Moreno et al.

The pattern found for the esterase enzyme shows These differences were maintained throughout the
that the Savana and Conquista cultivars have higher storage period. The enzyme catalase (CAT), which involves
activity of this enzyme at 0 and 4 months, indicating the removal of hydrogen peroxide formed from enzyme
deterioration. It also shows higher activity when there activity and is considered the second line of defense in the
is more deterioration, as it degrades lipids during antioxidant system after the enzyme SOD, showed high
germination. At 8 months, activity increases for all four activity during the 0 and 4-month periods. Similarly, no
cultivars. At 12 months, there is a noticeable decrease differences in enzyme expression were observed between
for the high-quality cultivars, except for the Savana treatments within each storage period. However, in
and Conquista cultivars. Mavaieie et al. (2019) found the 0 and 4-month periods, higher activity was found,
that, in cold storage, seeds have higher esterase activity contrary to what was found by Baldoni (2013), who
compared to seeds stored in uncontrolled conditions found differences in enzyme activity between seeds
throughout storage. Esterase plays a relevant role in collected at the R8 stage with higher expression and
the growth of the embryonic axis and the breakdown seeds collected 15 days after this stage with lower
of lipids in the germination process, especially in oily physiological quality.
seeds like soybeans. Esterase activity decreases after In deteriorated seeds, lower enzyme activity
the sixth month, and according to Veiga et al. (2010), (CAT) was also found, with lower efficiency of
bands may disappear after nine or twelve months of free radical scavenging systems. When the seed
storage. However, the Savana and Conquista cultivars ages, lipid peroxidation increases and enzymatic
maintained their enzymatic activity at 12 months, activity of peroxide-scavenging enzymes decreases
possibly due to their low quality. In contrast, the (BRACCINI et al., 2000; MENEZES et al., 2009).
CD201 and BMX cultivars presented lower enzymatic The expression of the alcohol dehydrogenase (ADH)
activity, higher germination, and vigor (Table 1). enzyme showed activity during the 4 storage periods,

Figure 2 - Isoenzyme analysis of four cultivars (C1: CD201, C2: BMX, C3: Savana, C4: Conquista), representing two high-quality
cultivars and two low-quality cultivars, respectively. Enzyme patterns for a) catalase, b) esterase, c) ADH, and d) MDH

6 Rev. Ciênc. Agron., v. 56, e202292842, 2025

 Storage potencial of soybeans cultivars under low temperature

with higher activity in high-quality cultivars in the 0, 8, AZEVEDO, M. R. D. Q. A. et al. Influência das embalagens
and 12-month storage periods. The ADH enzyme is e condições de armazenamento no vigor de sementes de
involved in the respiratory process and functions to gergelim. Revista Brasileira de Engenharia Agrícola e
remove acetaldehyde in seeds, as well as being relevant Ambiental, v. 7, n. 3, p. 519-524. 2003.
due to its conversion of acetaldehyde to ethanol, which BAILLY, C. et al. Catalase activity and expression in developing
is less toxic and reduces the rate of deterioration. Thus, sunflower seeds as related to drying. Journal of Experimental
seeds are less susceptible to the action of acetaldehyde when Botany, v. 55, n. 396, p. 475-83, 2004.
there is higher ADH enzyme activity (VEIGA et al., 2010). BALDONI, A. Expressão de genes relacionados com a
According to Mavaieie et al. (2019), during storage, the qualidade fisiológica de sementes de soja. 2013. 57 p. Tese
expression of this enzyme is generally higher in seeds (Doutorado em Fitotecnia) - Universidade Federal de Lavras,
stored in cold chambers compared to seeds stored under Lavras, MG, 2013.
uncontrolled conditions, especially at 6 and 8 months,
BISHT, P. et al. Evaluation of relative storage potential of
contributing to the maintenance of quality during cold storage. soybean (Glycine max L. Merril) varieties through accelerated
The expression of these enzymes in the 4 cultivars aging. Environment & Ecology, v. 33, p. 589-591, 2015.
showed higher activity during the four storage periods, with
BRACCINI, A. D. L E. et al. Biochemical changes associated
seeds stored in cold chambers exhibiting higher activity of to soybean seeds osmoconditioning during storage. Pesquisa
this enzyme compared to seeds stored under uncontrolled Agropecuaria Brasileira, v. 35, n. 2, p. 433-447, 2000.
conditions. In general, seeds stored in cold chambers show
elevated activity of malate dehydrogenases (MDH) until the BRASIL. Ministério da Agricultura, Pecuária e Abastecimento.
Secretaria de Defesa Agropecuária. Regras para análise de
end of storage, as also observed by Mavaieie et al. (2019).
sementes. Brasília: Mapa/ACS, 2009. 399 p.
Similarly, Mavaieie (2019) emphasizes that the MDH enzyme
has important physiological functions within the cell during CARVALHO, E. R. et al. Alterações isoenzimáticas em sementes
the Krebs cycle, converting malate to oxaloacetate. Acting in de cultivares de soja em diferentes condições de armazenamento.
respiration, it shows increased staining intensity or number of Pesquisa Agropecuária Brasileira, v. 38, n. 4, p. 967-976. 2014.
bands in seeds subjected to long storage periods, especially in DELOUCHE, J. C. Germinação, deterioração e vigor da semente.
seeds undergoing advanced deterioration. Seed News, ed. 6, nov. 2002. Disponível em: https://seednews.
com.br/artigos/2018-germinacao-deterioracao-e-vigor-da-
semente-edicao-novembro-2002. Acesso em: 10 jan. 2022.
CONCLUSIONS FERREIRA, D. F. Sisvar: a computer statistical analysis system.
Ciência e Agrotecnologia, v. 35, n. 6, p. 1039-1042, 2011.
Soybean seeds stored in cold storage maintain germination
and vigor for indefinite periods of time, with different FRANÇA NETO, J. B. F. et al. Mini-curso Tecnologia Produção
levels of tolerance to cold storage. Isoenzyme de Soja. Tecnologia da produção de semente de soja de alta
qualidade. Informativo ABRATES, v. 20, n. 3, p. 26-32, 2010.
expressions are also maintained and can even increase
when seeds are stored at low temperatures, affecting GARCIA, D. C. et al. Secagem de sementes. Ciencia Rural,
their expression according to the genotype and showing v. 34, p. 603-608, 2004.
different behaviors during different storage periods. JUVINO, A. N. et al. Vigor da cultivar BMX Potência RR de
soja durante o beneficiamento e períodos de armazenamento.
Revista Brasileira de Engenharia Agrícola e Ambiental,
v. 18, n. 8, p. 844-850, 2014.
ACKNOWLEDGMENTS
KRZYZANOWSKI, F. C. Tecnologías para produçao de
To the National Council for Scientific and sementes de soja. Londrina: Embrapa Soja, 2015.
Technological Development (CNPq) for the scholarship
LUDWING, M. P. et al. Populações de plantas na cultura da soja
grant. To the Coordination for the Improvement of em cultivares convencionais e Roundup Ready TM. Revista
Higher Education Personnel (CAPES), Minas Gerais Ceres, v. 58, n. 3, p. 305-313, 2011.
State Research Support Foundation (Fapemig) for
the financial support, and to the University of Cauca MARCOS-FILHO, J. Fisiologia de sementes de plantas
(501100005682) for their assistance. cultivadas. Piracicaba: FEALQ, 2005. 495 p.
MARTINS, E. H., et al. Soybean waste in particleboard
production. Ciencia e Agrotecnologia, v. 42 n. 2, p. 186-194, 2018.

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