The initial vigor of seeds under cold stress influences the performance of rice seedlingAbstract
Rice is a summer crop of great importance in Brazil and is subject to abiotic cold stress at sowing. With the use of high vigor seeds, it is possible for seedlings to have a capacity to recover from this type of stress, tolerating the stress and ensuring initial establishment, uniformity and consequently greater productivity. Therefore, the objective was to evaluate the recovery capacity of rice seedlings from seeds with contrasting vigor, after periods of cold stress. The study was carried out at the Seed Analysis Laboratory of the State University of Santa Catarina. The batches used were from the cultivar SCS122 Miura, from the 2018/2019 harvest and physiological characterization was carried out. Then, two batches were selected (low and high vigor) that were subjected to two, three, four and five days of cold at 10 °C followed by periods of post-stress recovery (exit and five days) at 25 °C. Control was maintained in the germinator at a constant 25 °C. After each period, normal and abnormal seedlings and dead seeds, shoot and root length and seedling dry mass were evaluated. A completely randomized design was used with four replications under a three-factorial scheme. It was observed that when subjected to two days of stress, the batch with greater vigor showed recovery capacity. The lower vigor batch had its performance drastically reduced after stress. It is concluded that the performance of rice seedlings under cold stress is favored by the initial vigor of the seed.
Keywords:
initial establishment; abiotic stress; Oryza sativa; physiological potential
Resumo
O arroz é uma cultura de verão de grande importância no Brasil e está sujeito ao estresse abiótico pelo frio na semeadura. Com a utilização de sementes de alto vigor é possível que as plântulas tenham capacidade de recuperação desse tipo de estresse, tolerando o estresse e garantindo estabelecimento inicial, uniformidade e consequentemente maior produtividade. Portanto, objetivou-se avaliar a capacidade de recuperação de plântulas de arroz a partir de sementes com vigor contrastante, após períodos de estresse por frio. O estudo foi realizado no Laboratório de Análise de Sementes da Universidade do Estado de Santa Catarina. Os lotes utilizados foram da cultivar SCS122 Miura, da safra 2018/2019 e foi realizada caracterização fisiológica. Em seguida, foram selecionados dois lotes (baixo e alto vigor) que foram submetidos a dois, três, quatro e cinco dias de frio a 10 °C, seguidos de períodos de recuperação pós-estresse (saída e cinco dias) a 25 °C. O controle foi mantido no germinador a temperatura constante de 25 °C. Após cada período foram avaliados plântulas normais e anormais e sementes mortas, comprimento da parte aérea e raízes e massa seca das plântulas. Utilizou-se delineamento inteiramente casualizado com quatro repetições em esquema trifatorial. Observou-se que quando submetido a dois dias de estresse, o lote com maior vigor apresentou capacidade de recuperação. O lote de menor vigor teve seu desempenho drasticamente reduzido após o estresse. Conclui-se que o desempenho das plântulas de arroz sob estresse pelo frio é favorecido pelo vigor inicial da semente.
Palavras-chave:
estabelecimento inicial; estresse abiótico; Oryza sativa; potencial fisiológico
1. Introduction
Climate change can drastically affect agriculture, with extreme temperatures, water deficits or floods, becoming a challenge for the cultivation and success of crops (Medeiros et al., 2022). For rice cultivation, temperature is one of the most important climatic elements, both for the growth and development and productivity of the cereal (Embrapa, 2013), presenting an optimal range of 20 to 35 °C for germination and development (Yoshida, 1981).
In the main rice growing areas in Brazil, the occurrence of temperatures below the optimum range is quite common, especially in the initial establishment - in the South of Brazil, irrigated rice sowing occurs between the months of September and November, which present temperatures of 15 to 22ºC and 20.6ºC, respectively (INMET, 2024a, b). Exposure of plants to cold results in changes to multiple physiological, biochemical, molecular and metabolic processes (Bajwa et al., 2014). Depending on the duration, exposure to stress and the stage of plant development, these damages can be irreversible (Chang et al., 2010) and compromise the success of the crop.
In the vegetative phase, seedlings subjected to prolonged exposure to low temperatures, for several days or weeks, may present necrosis and even die. While shorter or intermittent exposure can lead to leaf chlorosis, uneven initial establishment, delayed growth, short stature and reduced tillering (Zhang et al., 2014), indirectly compromising crop productivity.
In Brazil, the South region stands out in national cereal production, with the state of Rio Grande do Sul (7,142,801 tons) as the largest producer, followed by Santa Catarina with 1,176,285 tons and the state of Paraná, with 148,916 tons (IBGE, 2023). As it is common for high temperatures (> 25ºC) to occur during the flowering period (January to March) (INMET, 2020), which can affect productivity (ideal range of 20 to 25ºC; Embrapa, 2013), early sowing has been recommended as a strategy to adapt to climate change (Singh et al., 2011). This management also makes it possible to avoid high temperatures in critical periods, as by anticipating sowing, the reproductive phase will coincide with the time of greatest solar radiation, which is essential for obtaining high yields (Grohs et al., 2017).
Early sowing can expose rice seeds to suboptimal temperatures, which can compromise seedling development and reduce cultivar productivity. To mitigate the impact of this type of abiotic stress on initial establishment, the use of seeds with high vigor should be considered (Silva et al., 2016). High vigor seeds develop more quickly and uniformly and tolerate a wide range of environmental conditions, as well as having the capacity to develop stronger and more vigorous seedlings that can establish themselves more satisfactorily in the face of abiotic and biotic stresses (Marcos-Filho, 2015; Zonzen et al., 2025; Garcia and Coelho, 2021), providing an initial advantage in the use of water, light and nutrients (Mielezrski et al., 2008).
Despite this, it has not been studied how much the cold period (10 ºC) can compromise the establishment of seedlings in the initial stages and whether vigor can favor this condition of expression in the crop.
In view of the above, the objective of this work was to evaluate the performance of rice seedlings, originating from vigorous seeds, after being subjected to cold stress (10 °C).
2. Material and Methods
The study was carried out at the Seed Analysis Laboratory located at the Agroveterinary Sciences Center (CAV) of the State University of Santa Catarina (UDESC), in Lages, SC. We used batches of seeds from the SCS122 Miura rice cultivar produced in the 2018/2019 harvest in the state of Santa Catarina. The cultivar obtained by the Agricultural Research and Rural Extension Company of Santa Catarina (EPAGRI) is considered promising and has high yield potential (estimated at 9.6 t ha-1), the result of crossing PR122 and SCSBRS Tio Taka and backcrossing with SC354 (Epagri, 2018).
A completely randomized design was used, forming a three-factor scheme with four replications, the first factor being: seed vigor level (high or low); second: imposition or not of cold stress (10 °C); and third: times after stress at 25 °C (exit and five days). To this end, four lots of seeds of the cultivar were evaluated – with the interpretation of physiological analyses, two were chosen, of low and high vigor (Figure 1).
Experimental design and evaluations carried out after cold stress in rice seeds (SCS122 Miura). Characterization of batches (contrasting vigor), root protrusion, exposure to cold at different periods and recovery and variables evaluated after each stress cycle, in a completely randomized design with four replications.
Initially, the samples were homogenized and subdivided (replications) into four subsamples (350 g) (Brasil, 2009; Coelho et al., 2010) to carry out physiological analyses. Prior to the physiological tests, the method of overcoming dormancy was used in an oven with air circulation at 50 °C for 96 hours (Brasil, 2009). All analyzes were conducted under a completely randomized design with four replications and carried out following instructions from the Rules for Seed Analysis (RAS).
The germination test was conducted using four replications of 100 seeds for each batch, distributed on a roll of germitest paper (moistened by the equivalent of 2.5 times the weight of the paper, with distilled and deionized water). The rolls were placed in plastic bags, taken to the Mangelsdorf-type germinator and kept at 30 ± 2ºC for eight days (Brasil, 2009). The first count was carried out on five days and the final count on the eighth day, after sowing (Brasil, 2009). At the end of the test, the number of normal seedlings, abnormal seedlings, dead, hard and dormant seeds was recorded. The germination result was expressed as a percentage and average number of normal seedlings.
The vigor by the cold test was carried out using four repetitions of 100 seeds for each batch, distributed on a roll of germitest paper and moistened by the equivalent of 2.5 times the weight of the dry paper, with distilled and deionized water. The rolls were placed in plastic bags, taken to Biochemical Oxygen Demand (BOD) and kept at 10 ± 2 ºC for seven days (Cicero and Vieira, 2020). After this period, the rolls were transferred to the Mangelsdorf type germinator at 25 ± 2 ºC where they remained for another seven days. At the end of the test, the number of normal seedlings, abnormal seedlings and dead seeds was recorded. The percentage of vigor was expressed as a percentage and estimated by the average result of the number of normal seedlings.
The evaluation of the length of shoots and roots was carried out by assembling rolls of germitest paper, which were previously moistened 2.5 times their dry weight, with twenty seeds and four replications for each batch. The seeds were distributed in two rows in the upper third, maintaining a similar distance between them. These rolls were taken to the Mangelsdorf type germinator and kept at 25 ± 2 ºC for seven days. After this period, the length of the shoot and root of all normal seedlings was measured. Measurements were taken with the aid of a digital caliper (10-3). Average results were expressed in mm seedling-1, according to the method proposed by Krzyzanowski et al. (2020).
In the seedlings used to measure length, the endosperm was also separated to determine the dry mass in an oven at 80 °C for 24 hours (Krzyzanowski et al., 2020). And the seedling dry mass (SDM) was expressed in mg seedling-1.
In addition, the reserve mobilization rate (RMR) was carried out. To avoid the interference of the seed coat in the parameters related to mobilization, the dry mass of the seed coats of each batch was carried out. Then, they were dried in an oven to obtain the seed dry mass (SeDM). The RMR, which represents how much was actually mobilized for the seedling, was expressed as a percentage and calculated using the Equation (1):
Through the physiological characterization described previously, two lots of the cultivar SCS122 Miura, of low and high vigor, were selected. After selecting the two lots, four replications were set up with 100 seeds per treatment, distributed on a roll of germitest paper (moistened by the equivalent of 2.5 times the weight of the paper, with distilled and deionized water). The rolls were placed in plastic bags, taken to the Mangelsdorf type germinator and kept at 25 ± 2 ºC for four days, reaching the “S2” point – described by Marcos-Filho (2015), as phase III of the germination process, which is characterized by the resumption of embryo growth and root protrusion, with ≈ 2.0 mm.
Then the seeds were subjected to two, three, four and five days of cold at 10 °C followed by periods of five days at 25 °C in the germinator (recovery). Control was also maintained in the germinator at a constant 25 °C. After each period (stress + recovery), the number of normal and abnormal seedlings and non-germinated seeds (hard, dead or dormant), shoot and root length and seedling dry mass were evaluated, as previously described.
The means were subjected to analysis of variance (F test) and when significant, they were compared using the Tukey test at a 5% probability of error. As well as principal component analysis (PCA) between the variables studied. All statistical analyzes were performed using the R software (R Core Team, 2020).
3. Results and Discussion
All evaluated lots of the cultivar SCS122 Miura produced in the 2018/2019 harvest showed germination above 80%, as recommended by current legislation for the commercialization of certified rice seeds (Brasil, 2013). Batches 1 and 3 showed the highest germination percentages (93%) and did not differ statistically from each other. While lot 4 had the lowest value, 86% (Table 1). This result shows the importance of carrying out vigor tests, as the batches with the highest germination are not always the most vigorous batches.
Physiological characterization of germination and emergence of rice seeds from four lots of the SCS122 Miura cultivar 2018/2019 harvest. Germination (G), cold vigor test (CVT), seedling dry mass (SDM), shoot length (SHL), root length (RL), seedling length (SL) and reserve mobilization rate (RMR).
An important characteristic of this study that must be highlighted is that although the seeds used were all from the SCS122 Miura cultivar, they were different batches, that is, a defined and homogeneous quantity of seeds with similar quality atributes. Therefore, they may differ in terms of sowing times, soil and climatic conditions of production, incidence of pests and diseases, cultural and nutritional management and harvest time. These characteristics can directly influence the quality of the seeds produced (Marcos-Filho, 2015), as observed in the physiological results (Table 1).
In the cold vigor test, a large variation was observed between the tested lots, from 36 to 75%. Lot 2 had the lowest cold vigor (36%). Lots 1 and 3 had 56%, not statistically different from each other. Lot 4 stood out with 75%, presenting the highest percentage of vigor according to the cold test (Table 1). Lots of seeds from the same cultivar, and with similar germination, may present different performances in relation to agronomic behavior due to differences in seed vigor - such differences can be noticed in the field or during storage, these variations can be noticeable and analyzed through vigor tests (Marcos-Filho, 2015; Sampaio et al., 2025).
In terms of seedling performance, it was found that batch 4 had a greater length of shoots, roots and seedlings, and consequently, greater dry mass, standing out from the other batches (Table 1). Andrade et al. (2019) also observed that seeds of corn hybrids with greater vigor give rise to seedlings with greater accumulation of dry mass, total seedling length, aerial part and root.
Furthermore, lot 4 also differed significantly from the others, presenting a higher reserve mobilization rate (RMR), reaching 15.16% (Table 1). Seeds with greater vigor have higher RMR, that is, greater efficiency in the hydrolysis of seed reserve components and mobilization for seedlings, which favors the development of seedlings with superior performance (Padilha et al., 2020; Padilha et al., 2025). It is noteworthy that RMR is strongly correlated with seed vigor (Delgado et al., 2015). As well, for Padilha et al. (2021), bean seeds with greater vigor, showed greater mobilization of reserves due to the action (greater activity) of the alpha-amylase enzyme even under saline stress.
Through the various vigor tests carried out, it was possible to identify the physiological potential of the batches by carrying out a joint analysis of the results, and not just a vigor test to have more reliability (Marcos-Filho, 2013). Thus, based on this physiological characterization, batch 4 was selected and identified as having high vigor, showing 75% vigor due to cold, and batch 2, with 36% vigor using the same test, as having low vigor (Table 1).
Regarding the analysis of main components, it was possible to highlight the influence of factors (different levels of vigor, with or without stress, period of stress and recovery) on the physiological performance of seedlings and/or seeds. The first component (PC1) explains 99.7% of the data variation, showing an association between the variables: normal seedlings (NS_%), shoot length (APL_cm), root length (RL_cm) and dry mass (DMS_g) (Figure 1). Furthermore, the association of normal seedlings, root length, area length and seedling dry mass with seeds of high vigor, lower stress intensity and longer recovery time after stress is highlighted – in contrast, the variable abnormal seedlings was concentrated in more unfavorable scenarios (Figure 2).
Principal component analysis (a) presenting the association between the study variables and the post-stress recovery period (b) association between the variables and the condition with and without stress (c) association between the variables and the vigor groups and (d) association between the variables and the different periods of cold stress (10 ºC).
At the end of the stress period, regardless of the days of stress, the batches did not form normal seedlings (Figure 1a; Table S1, Supplementary Material). The stress probably caused a delay in metabolic reactions, mainly the action of essential enzymes in the mobilization process, such as the esterase enzyme, which leads to a lower accumulation of dry mass, due to the reduction or paralysis of the metabolism of reserve products (Mertz et al., 2009), directly affecting the formation of seedlings. Furthermore, low temperatures can also inhibit the activity of many oxidoreductive enzymes, such as catalase, leading to the accumulation of hydrogen peroxide and the production of free radicals, which can compromise the mobilization of reserves and seedling development (Sun et al., 2010).
In general, there was an increase in the percentage of normal seedlings after five days of recovery after stress. In addition, variables such as dry mass and shoot length approached the factor, indicating better physiological recovery performance (Figure 2a). The variable abnormal seedlings were associated with the period of exit immediately after stress, possibly influenced by the initial effect of stress on the physiological quality of the seedlings (Figure 2a). There was also a significantly superior performance of the high vigor lot over the low one, in the control and when subjected to abiotic stress, suggesting that the characteristics of shoot and root length, dry mass and normal seedlings are associated with the non-stressed condition and high vigor, and the abnormal seedlings, the stressed condition and low vigor (Figure 2b, 2c).
When subjected to two days of stress, it was observed that the high vigor lot had a high capacity to recover from stress, where variables such as area length, dry mass and normal seedlings were associated with the stress period (Figure 2d; Table S1, Supplementary Material). The batch with low vigor, after being subjected to stress, was unable to recover, especially in four and five days of cold stress, showing a greater association with the variable abnormal seedlings (Figure 2c; Table S1, Supplementary Material).
Under low temperatures, low vigor seeds can experience a significant delay in emergence, resulting in weak seedlings and stand failures. As high vigor seeds, they tolerate a wide range of environmental conditions and produce seedlings with greater dry mass and length, resulting in better initial development of the crop and, consequently, high productive potential (Krzyzanowski et al., 2018).
Periods of cold at 10º C for more than two days cause irreversible damage to rice seedlings, but high-vigor seeds have a greater capacity for recovery (Figure 2c, 1d; Table S1, Supplementary Material). In this way, it became evident that depending on the intensity of the stress, the formation of normal seedlings can be delayed or suppressed (Ali and Elozeiri, 2017). Furthermore, it is important to highlight that tolerance to cold stress is considered a multigenic trait, which involves changes in gene expression, membrane structure and function, water content, expression of cold-regulated proteins (such as dehydrins), lipids and primary and secondary metabolites (Basuchaudhuri, 2014; De Freitas et al., 2019), reflecting the seedling's performance in the face of stress.
The seedlings evaluated were in phase III of the germination process. The beginning of this phase is marked by the protrusion of the primary root. At the same time, DNA synthesis, mitochondria synthesis and repair, protein synthesis, gene expression, and an increase in cellular respiration also occur (Kerbauy, 2019), triggering a significant mobilization of soluble compounds for growing seedlings (Taiz et al., 2017). This process, in turn, is highly dependent on the formation of enzymes and their activity (Cheng et al., 2015). Therefore, as thermal stress harms the entire plant metabolism, affecting protein stability and enzymatic reactions, reducing the fluidity of membranes, causing the interruption of several metabolic reactions, in addition to the disruption of homeostasis and ionic regulation (Taiz et al., 2017), it ultimately interferes with the formation of normal seedlings.
The effects of these changes on seed metabolism, and consequently on seedlings in formation, can also be seen in other variables analyzed, such as anormal seedlings. The cold affected the formation of normal seedlings by compromising the structures of the seedlings, causing an increase in the number of abnormal seedlings (Figure 2b; Table S1, Supplementary Material).
The high rate of abnormal seedlings observed may have been the result of a delay in metabolism caused by low temperatures, since after stress there was a reduction in the number of abnormal seedlings over the five days (Figure 2a; Table S1, Supplementary Material).
After being subjected to stress, the high vigor batch showed a reduction of up to 81% of abnormal seedlings over the days. While the low vigor batch had a reduction of only 39% (Figure 2c; Table S1, Supplementary Material). This reduction in the number of abnormal seedlings indicates the time that the metabolism used to reorganize itself after the stress, aiming to resume the growth of structures, subsequently making many seedlings normal. Therefore, there is a significant efficiency of high vigor seeds in resuming growth, thus giving the seedlings the ability to recover from cold stress within a certain duration.
An abiotic stress, when causing some disturbance in the plant cell, signals the seedlings of changes in environmental conditions and induces a response through the alteration of existing routes or the activation of stress response routes (Taiz et al., 2017). Furthermore, seedlings have several physiological, morphological and metabolic mechanisms to minimize or avoid damage, such as osmotic adjustment, adaptation of the root/shoot ratio and memory (Taiz et al., 2017). Therefore, it is possible that vigorous seeds gave rise to seedlings with faster and more efficient responses in signaling and activating mechanisms in the face of cold stress than less vigorous seeds, making up an intrinsic characteristic of vigor.
Severe cold stress directly affects the structure and function of cell membranes, causing a change from the liquid crystalline phase to the solid phase (gel formation), affecting membrane permeability and inducing damage to plant tissues, which can lead to seedling death (Ansari et al., 2019). Therefore, the survival of a plant cell highly depends on the integrity of its membrane, which was possibly significantly affected under the conditions tested.
Regarding the percentage of dead seeds, only the seed vigor level was significant (Table 2). In other words, there was no effect of the stress period or post-stress on the percentage of dead seeds. The low vigor batch had a higher percentage, around 10%, while the high vigor batch had approximately 5% (Table 2).
As there was no effect of the stress period or post-stress, it can be stated that cold stress only caused physiological changes in the formation of rice seedlings and did not trigger seed death. Therefore, the percentage of dead seeds was probably something intrinsic to the batch and was possibly more related to the production conditions of these seeds and their physiological aspects.
The production environment and the interaction of the genotype with the environment can directly influence the quality of the seeds produced, as after physiological maturation, the seeds remain in the production field until the point of harvest – becoming susceptible to inadequate conditions of high temperature, humidity and precipitation, which in turn trigger seed deterioration, reducing physiological quality and even causing death (Szareski et al., 2018).
Differences in vigor between seedlings are, in most cases, noticeable, however it is important to analyze numerical values to facilitate the segregation of vigorous seedlings from those that are not (Nakagawa, 1999). Due to this, the measurement of the average length of normal seedlings or parts thereof is done considering that the batches that present the highest average values are the most vigorous (Nakagawa, 1999).
Furthermore, the work generated information that helps to understand that the cold directly affected the seedlings, since the classification of normal, abnormal and dead seeds further evaluates the issue of the presence or absence of structures, while measurement evaluates the performance of these seedlings under such conditions.
In terms of shoot length, it was observed that although the high vigor batch stood out due to low vigor (in control and under stress), the batches showed similar performance in the post-stress period, both differing statistically from the control (Figure 2a, 2b, 2c; Table S1, Supplementary Material).
However, although cold stress affected the growth of the aerial part of both seedlings originating from seeds with high vigor and those with low vigor, there was a tendency for growth to stabilize (Figure 2c; Table S1, Supplementary Material). This reinforces the greater metabolic efficiency of the high-vigor batch in reorganizing itself after stress. Possibly due to water deficit or osmotic stress caused by the cold, which may affect cellular metabolism and the transfer of abscisic acid (Wu et al., 2024). After the protrusion of the primary root (phase III), the demand for water increases substantially (Taiz et al., 2017), as water is essential to support the growth of roots and shoots, due to the high intensity of cell division and elongation, and tissue synthesis and differentiation (Marcos-Filho, 2015). In this type of abiotic stress, seedlings may have problems absorbing water, and consequently suffer water deficit - related to changes in the permeability of the cells' plasma membrane, with physiological and biochemical changes (Feng et al., 2025). Due to this, the growth of the aerial part may have been affected.
For root length, the high vigor batch also stood out in control and post stress (Figure 2a, 2b, 2c; Table S1, Supplementary Material). Rice roots are more sensitive to low temperatures than leaves (Xiao et al., 2014), as roots produce more reactive oxygen species (ROS) (Hsu and Hsu, 2019). This is due to the process of evolution, where these structures have adapted to more stable temperatures (Źróbek-Sokolnik, 2011) and root proteins have faster responses to stress compared to leaf and stem proteins (Lee et al., 2009). In general, it was observed that seedlings from high and low vigor lots reduced their length by up to 50% when subjected to stress (Table S1, Supplementary Material). Genetic factors can influence seedling responses to cold stress. Kanagatov et al. (2025) reported that transgenic rice plants express the ICE1 gene and show greater tolerance to cold stress than wild-type plants, with increased proline, reduced malondialdehyde and greater peroxidase activity, with gene expression more pronounced in leaves than in roots. As well, Amirova et al. (2024) observed cold tolerance in rice plants linked to three QTLs (qPSST-3, qPSST-7, qPSST-9), demonstrating that the presence of three confers tolerance, while the absence of one, sensitivity.
In addition to the greater sensitivity to this type of stress, the reduction in root length can be attributed to the decrease in cell division and elongation, in addition to the low respiration rate and high oxidative stress that occurs at low temperatures (Hussain et al., 2016). All of this limited growth and may have impaired water absorption, affecting seedling performance.
Once again, it was possible to observe that the duration of stress is decisive in the physiological response. After five days of stress, the root length of the seedlings, from both batches, was stabilized throughout the post-stress period, indicating how severe it was (Figure 2d; Table S1, Supplementary Material).
In general, the seedling dry mass results showed that the controls stood out with the highest accumulations, with no recovery in any batch, differing from the control in all periods of stress and recovery time (Figure 2a, 2b, 2d; Table S1, Supplementary Material).
The batches subjected to stress showed similar performance, with a slight increase in post-stress dry mass. Corroborating the other results already observed in this work, which indicate the greater efficiency of the metabolism of high-vigor seeds in reorganizing themselves post-stress, when compared to those with low vigor.
Considering that most of the mobilization of seed reserves occurs after radicle protrusion and this process is highly influenced by enzymatic activities, which are in turn regulated, among several factors, mainly by temperature (Cheng et al., 2015). It is possible that with the occurrence of stress, temperature interfered with the activity of the enzymes α and β-amylase, mainly responsible for hydrolysis and mobilization of reserves in seeds, resulting in a reduction in mass accumulation, and resulting in seedlings with lower dry mass.
Furthermore, stress decreases the rate of cell initiation in leaves and roots, and consequently decreases cell division and elongation, which reduces the length of these structures, and leads to a reduction in root length - resulting in less accumulation of mass in these seedlings (De Freitas et al., 2019).
According to Źróbek-Sokolnik (2011), there are two theories that help to understand the primary response of plants to abiotic cold stress. The first states that low temperatures induce a change from the liquid crystal phase to the crystal (gel) phase of cell membranes (Lyons, 1973 p. 459 apud Źróbek-Sokolnik, 2011 p. 119) - this change in the physical state of the membrane impairs its proper functioning. The second theory states that low temperatures cause a sudden increase in the concentration of free calcium ions in the cytosol (Minorsky, 1989 p. 123, apud Źróbek-Sokolnik, 2011 p. 121), which causes induced gene expression and results in changes in the content or distribution of hormones, mainly abscisic acid.
Through the study, it was observed that the seedlings' response is dependent on the duration of the stress (two days: high vigor seedlings demonstrate recovery capacity; five days: significantly affected seedlings).
It is noteworthy that high vigor seedlings were also affected by the cold, but showed a significantly higher response than those with low vigor, in all variables analyzed. Furthermore, they have a strong tendency to recover after stress, limiting the duration of the stress.
Therefore, after analyzing the physiological and performance variables, a more in-depth study of which biochemical mechanisms are associated with these observed responses is necessary, such as hormonal regulation, enzymes and gene expression. Since the ability to tolerate abiotic stress by high vigor seeds is possibly influenced by internal regulatory mechanisms, still little elucidated in rice cultivation.
4. Conclusion
High vigor seeds show a superior response to cold stress (10ºC), ensuring better formation of normal seedlings, better performance and recovery after stress, ensuring uniformity and satisfactory initial development. Being able to tolerate the stress condition for up to two days, without compromising post-stress performance.
Supplementary Material
Supplementary material accompanies this paper.
Table S1
This material is available as part of the online article from https://doi.org/10.1590/1519-6984.301963
Acknowledgements
The authors would like to thank the financial support of FAPESC (PAP/FAPESC/2025TR1628). The corresponding author (Coelho, C.M.M.) thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for productivity scholarship.
Data Availability Statement
All original data can be made available.
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Editor:
Takako Matsumura Tundisi
