Seed germination is a critical phase in the plant life cycle, as it determines the successful establishment of seedlings. This process is influenced by factors such as physiological maturity, water content, and reserve composition, which vary among species. When exposed to an appropriate substrate and favorable environmental conditions, seeds resume metabolic activities and initiate embryo development (Carvalho & Nakagawa, 2012; Pereira et al., 2022; Nautiyal et al., 2023). Germination begins with water imbibition, during which the seed absorbs water, consumes stored reserves, and prepares for primary root protrusion, marking the transition to nutrient uptake from the environment (Schmidt, 2007; Borges & Toorop, 2015; Aumonde et al., 2019).

Imbibition follows a triphasic model of water absorption, with each phase characterized by distinct physiological and biochemical changes. Phase I (PI) involves rapid water uptake due to the dry seed’s matrix potential, resulting in increased seed mass and initial metabolic activation (Bewley & Black, 1994; Cardoso, 2013; Taiz et al., 2017). Phase II (PII) is marked by a stabilization of water content, with seeds maintaining a constant or slightly increased mass as metabolic processes are activated and cellular structures repaired. Finally, Phase III (PIII) is characterized by a renewed increase in water absorption associated with embryo growth, culminating in the protrusion of the primary root (Castro & Hilhorst, 2004; Cardoso, 2013; Taiz et al., 2017).

The duration and dynamics of the imbibition process vary among species, influenced by seed morphology, chemical composition, and tegument permeability. The seed coat plays a crucial role during imbibition by regulating water entry and acting as a reservoir in the early stages of germination (Silva & Dantas, 2016). Seeds with high oil content, for example, tend to absorb less water compared to starchy or protein-rich seeds, leading to differences in absorption speed and intensity (Bewley et al., 2013). Additionally, factors such as tegument porosity, structure, and pigmentation influence water permeability and hydration efficiency (Marcos-Filho, 2015; Ribeiro- Oliveira & Silva, 2024).

Laboratory germination tests provide valuable insights into the germination potential of seeds under controlled conditions. These tests allow for the evaluation of seed quality and vigor, facilitating comparisons of physiological potential, determining germination rates for sowing and commercial use, and identifying optimal storage conditions (Carvalho & Nakagawa, 2012). Understanding imbibition periods is particularly important, as this phase governs key processes such as tegument permeability, reserve mobilization, and the onset of metabolic activity (Borges et al., 2002).

Monitoring imbibition patterns is essential for optimizing seed germination and maximizing seedling emergence under field conditions. A precise understanding of water uptake dynamics allows for better prediction of germination timing and facilitates the development of species-specific pre-sowing treatments to enhance performance in restoration scenarios (Finch-Savage & Bassel, 2016); Nautiyal et al., 2023).

Moreover, characterizing the imbibition behavior of native forest species used in ecological restoration offers key insights into their functional responses under direct seeding conditions. This is particularly relevant for species with contrasting ecological strategies and functional traits, as variations in seed coat permeability, reserve composition, and dormancy mechanisms can strongly influence germination success and field establishment (Rajjou et al., 2012; Bewley et al., 2013). By integrating these physiological parameters into species selection and sowing strategies, restoration practitioners can design more effective, context-specific interventions to accelerate ecosystem recovery in degraded landscapes.

Variations among the duration of the three imbibition phases (PI, PII, and PIII) can reflect species-specific adaptations that can be quantified under controlled conditions. Understanding these differences provides insights into the germination potential of these species, supporting their optimized use in direct seeding for ecological restoration. Thus, this study aims to characterize the imbibition patterns and germination dynamics of native forest species used in direct seeding to identify and compare variations in the duration of the imbibition phases (PI, PII, and PIII) across species.

Species were selected for direct seeding based on the criteria proposed by Piotrowski et al. (2023), which emphasize small and/or lightweight seeds from different successional groups that exhibit high germination rates under laboratory conditions (>30%) but low emergence and establishment in the field. The selected species included representatives from the families Fabaceae (Mimosa bimucronata [DC.] Kuntze, Senegalia polyphylla [DC.] Britton & Rose, Pterogyne nitens Tul., Poecilanthe parviflora Bentham), Malvaceae (Apeiba tibourbou Aubl., Ceiba speciosa [A.St.-Hil.] Ravenna, Guazuma ulmifolia Lam.), Bixaceae (Bixa orellana L.), Anacardiaceae (Astronium urundeuva [M.Allemão] Engl.), Bignoniaceae (Handroanthus impetiginosus [Mart. ex DC.] Mattos), and Moraceae (Maclura tinctoria [L.] D.Don ex Steud.).

Seeds were obtained from commercial suppliers between August and December 2023 to ensure a diversity of provenances and a representative mix of maternal genotypes. Analyses were carried out between October and December 2023, with seeds stored at 5°C in a cold chamber until the beginning of the experiments. Seed lots were characterized for moisture content using the oven-drying method (105°C for 24 hours), thousand seed weight (TSW), and the number of seeds per kilogram, in accordance with Brazilian seed testing regulations (Brazil, 2009) (Table 1). The seeds of A. urundeuva and G. ulmifolia had their dormancy overcome by soaking the seeds in water for 48 hours, followed by immersion at 90°C and water soak for 1 hour off the heat, as recommended by the Guidelines for Forest Seed Analysis (Brazil, 2013).

Table 1 – Physical characterization of seed species based on seed moisture content, thousand seed weight (TSW), and seed count per kilogram

Source: Authors (2024)

Four replicates of 25 seeds were used to evaluate the imbibition process. The seeds were placed on germination paper, arranged in roll form, and moistened with distilled water at 2.5 times their dry weight. The paper rolls were enclosed in plastic bags to maintain humidity and incubated in a Biochemical Oxygen Demand (BOD) germinator under a 12-hour photoperiod at 25°C, except for Mimosa bimucronata and Apeiba tibourbou, which were maintained at 30°C. Seeds were weighed at two-hour intervals during the first 12 hours of imbibition and then every 24 hours thereafter. Monitoring continued until germination stabilized, defined as the protrusion of the primary root to a length of ≥ 2 mm.

The imbibition curves were generated based on the increase in seed water content (%) over time, modeled using a third-order polynomial equation: A+BX+CX2+DX3. This model was selected due to its close resemblance to a sigmoidal curve, as determined by the relationship between water content percentage and imbibition duration. Data analysis was conducted using R software version 4.2.2 (R Core Team, 2022).

Imbibition monitoring revealed that all species followed the three-phase water absorption model proposed by Bewley & Black (1994) (Figure 1). However, significant differences were observed in the duration of each phase. Phase I (PI), the initial rapid water uptake, varied among species, lasting from 2 to 16 hours, as also described by Marcos-Filho (2015). Specifically, PI was completed within 2 hours for Guazuma ulmifolia and Senegalia polyphylla, 4 hours for Apeiba tibourbou, 6 hours for Astronium urundeuva and Mimosa bimucronata, 8 hours for Ceiba speciosa and Pterogyne nitens, 12 hours for Bixa orellana, Handroanthus impetiginosus, and Poecilanthe parviflora, and up to 24 hours for Maclura tinctoria.

Phase II (PII), characterized by a plateau in water uptake, showed substantial interspecific variation, ranging from 12 hours in A. tibourbou, M. bimucronata, and S. polyphylla to 144 hours in M. tinctoria and H. impetiginosus. While the extended duration of PII in some species may be partly attributed to the activation of DNA and membrane repair mechanisms during early germination, particularly in seeds subjected to prolonged storage (Rajjou & Debeaujon, 2008; El-Maarouf-Bouteau & Bailly, 2008), we hypothesize that this variation is also strongly influenced by functional seed traits.

These traits may include seed coat characteristics such as thickness, permeability, and mechanical resistance, which directly influence the rate and uniformity of water uptake (Upretee et al., 2024; Zhou et al., 2022). Additionally, the chemical composition of seeds—particularly the type and abundance of storage reserves (proteins, lipids, and carbohydrates) and antioxidant metabolites—affects both hydration dynamics and the speed of metabolic reactivation (Ali & Elozeiri, 2017; Soriano et al., 2013). Together, these structural and biochemical traits contribute to species-specific imbibition behaviors and may be key determinants of germination performance under field conditions.

The autoecology of each species, including adaptations to specific environmental conditions, may influence the duration of PII by modulating seed dormancy and germination strategies (Fenner & Thompson, 2005). As proposed by these authors, species with harder seed coats or higher lipid content may exhibit prolonged PII due to slower water absorption or delayed metabolic processes (Bewley et al., 2013; Baskin & Baskin, 2014). Similarly, species from habitats with unpredictable moisture availability may have evolved slower germination processes as a bet-hedging strategy to ensure establishment under favorable conditions (Fenner & Thompson, 2005). Further exploration of these traits, combined with a deeper understanding of their ecological significance, could provide valuable insights into the germination patterns observed in this study.

The primary root emission (PIII) occurred most rapidly in Mimosa bimucronata and Senegalia polyphylla (24 hours; Figure 1-a;b) followed by Ceiba speciosa, Guazuma ulmifolia, and Pterogyne nitens (48 hours; Figure 1-c;d;e). Intermediate durations were observed in Bixa orellana (72 hours; Figure 1-f) and Apeiba tibourbou (96 hours; Figure 1-g). The most prolonged periods for primary root emergence were recorded for Astronium urundeuva and Poecilanthe parviflora (144 hours; Figure 1-h;i) and Handroanthus impetiginosus and Maclura tinctoria (168 hours; Figure 1-j;k).

Figure 1 – Imbibition curves of the 11 studied species. (a) Mimosa bimucronata and (b) Senegalia polyphylla showed primary root emission at 24 h. (c) Ceiba speciosa, (d) Guazuma ulmifolia, and (e) Pterogyne nitens at 48 h. (f) Bixa orellana and (g) Apeiba tibourbou at 72–96 h. (h) Astronium urundeuva and (i) Poecilanthe parviflora at 144 h, and (j) Handroanthus impetiginosus and (k) Maclura tinctoria at 168 h. The red points represent the increase in water content (%) in the imbibition phases (I, II, III), the blue line indicates the trend, the gray area the confidence interval and the arrow indicates the primary root protrusion (PR)

Source: Authors (2024)

The germination process in seeds, in particular the transition through phases I (PI), II (PII), and III (PIII), reflects complex interactions between the physical properties of the seed and its ecological adaptations. The emission of the primary root (PIII) is a critical milestone, marking the beginning of resource acquisition and seedling establishment. Our findings indicate substantial variation in the duration of these phases among the studied species, highlighting potential adaptive strategies for different ecological contexts and implications for direct seeding.

Species such as Mimosa bimucronata and Senegalia polyphylla, which complete PIII within 24 hours, exemplify traits advantageous for early establishment in direct seeding projects. Their short PI and PII phases likely reflect efficient water absorption and rapid metabolic activation, reducing the lag time between hydration and germination. This swift response confers significant adaptive advantages, particularly in environments where water availability is transient or unpredictable. Rapid root emergence allows seedlings to establish early access to soil moisture, anchoring the seedling and reducing desiccation risk (Baskin & Baskin, 2014; Finch-Savage & Bassel, 2016).

The biochemical basis for these differences may lie in the composition of hydrophilic compounds, such as soluble carbohydrates and storage proteins, which facilitate faster hydration and metabolic reactivation (Bewley et al., 2013). Additionally, thinner or more permeable seed coats may expedite water uptake, as observed in species like C. speciosa and P. nitens (Silva & Dantas, 2016).

In contrast, species with prolonged PIII phases, such as Handroanthus impetiginosus and Maclura tinctoria (168 hours), likely represent bet-hedging strategies suited for less predictable environments. These species may rely on slower imbibition rates or mechanisms to delay germination until environmental conditions are optimal, minimizing the risk of seedling mortality during unfavorable periods (Fenner & Thompson, 2005; Dürr et al., 2015). The extended duration of PII in these species may also reflect higher investment in metabolic repair or the mobilization of stored reserves, enhancing seedling vigor once germination occurs (Masetto et al., 2014).

Imbibition is influenced by functional traits, including seed coat permeability, chemical composition, and storage compound type. For example, species with high lipid content, such as Bixa orellana, may exhibit slower water uptake during PI and PII, as lipids require emulsification before metabolism can begin (Bewley & Black, 1994). Additionally, species with thicker seed coats, such as H. impetiginosus, may demonstrate extended PII due to the slower rate of water penetration and oxygen diffusion (Silva & Dantas, 2016).

From the literature, variations in PI duration among forest species further support this hypothesis. For instance, rapid PI completion was observed in Bignoniaceae species such as Jacaranda brasiliana 2 hours (Lima et al., 2018) and Handroanthus chrysotrichus 4 hours (Guollo et al., 2018), while species like Sapindus saponaria (Sapindaceae) required 71 hours (Torres et al., 2020). These differences underline the importance of both intrinsic traits and environmental factors in shaping imbibition dynamics.

In direct seeding, combining fast- and slow-germinating species can enhance restoration outcomes by aligning functional traits with ecological roles. Pioneer species such as M. bimucronata and S. polyphylla, characterized by rapid imbibition and germination, are well-suited for early establishment, soil stabilization, and microsite amelioration. These species can mitigate erosion, improve nutrient cycling, and provide shade, facilitating the recruitment of slower germinating species like M. tinctoria and H. impetiginosus, which contribute to long-term resilience and diversity (Moles & Westoby, 2004; Finch-Savage & Bassel, 2016). Differences in imbibition dynamics reflect distinct germination strategies shaped by environmental pressures. Rapid imbibing species typically exhibit traits such as small seed size, fast establishment, and tolerance to open, exposed environments. In contrast, slow-imbibing species may rely on more stable moisture conditions and often benefit from facilitation by earlier-establishing species (Saatkamp et al., 2019; Larson & Funk, 2016). Recognizing these physiological and ecological complementarities is crucial for optimizing species mixtures especially in contexts of increasing climatic variability and soil degradation.

A notable gap in the literature exists regarding the imbibition and germination processes of the studied species. Understanding these dynamics at the biochemical and ecological levels is critical for optimizing their use in restoration. Future studies should investigate the molecular pathways that control water uptake, metabolic activation, and stress responses during germination, particularly for species with significant variability in imbibition patterns.

This study revealed significant variation in the duration of the triphasic water absorption model (PI, PII, and PIII). Phase I ranged from 2 to 24 hours, Phase II from 12 to 144 hours, and Phase III from 24 to 168 hours, reflecting species-specific differences in hydration efficiency. These variations underscore the influence of functional seed traits, such as seed coat permeability and reserve composition, on germination dynamics.

The results provide valuable insights into the potential application of these species in ecological restoration. We identified rapid imbibition and germination, such as Mimosa bimucronata and Senegalia polyphylla, ideal for early establishment, and slower-imbibing species like Handroanthus impetiginosus and Maclura tinctoria. Understanding these imbibition patterns supports the optimization of direct seeding protocols, enhancing the efficiency and success of restoration efforts while contributing to biodiversity recovery in degraded ecosystems.

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001. We extend our gratitude to Enzo Coletti Manzoli for his valuable assistance in reviewing the analysis scripts, to the Seed and Seedling Laboratory (LASEM, lasemufscar.wixsite.com/lasem) for their technical support and to the Graduate Program in Planning and Use of Renewable Resources (PPGPUR).

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Authorship contributions

1 – Andreliza Roberta Terciotti de Oliveira

Master’s degree in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0001-8441-9922 • andrelizaterciotti@estudante.ufscar.br

Contribution: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Writing – original draft, Writing – review & editing

2 – Mariane Cristina Inocente

PhD in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0001-6456-3396 • mariane.inocente@estudante.ufscar.br

Contribution: Methodology, Supervision, Writing – review & editing

3 – Felipe Bueno Dutra

PhD in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0002-6280-7307 • fbdutra@estudante.ufscar.br

Contribution: Formal analysis

4 – Ana Paula Ponce Shiguehara

Master’s degree in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0002-3591-3286 • anapps@estudante.ufscar.br

Contribution: Writing – review & editing

5 – Danilo Ribeiro da Costa

PhD in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0002-2446-0234 • danilorc.ambiente@gmail.com

Contribution: Formal analysis, Software

6 – Fatima Conceição Márquez Piña-Rodrigues

PhD in Planning and Use of Renewable Resources from the Federal University of São Carlos

https://orcid.org/0000-0001-8713-448X • fpina@ufscar.br

Contribution: Supervision

How to quote this article

Oliveira, A. R. T., Inocente, M. C., Dutra, F. B., Shiguehara, A. P. P., Costa, D. R., & Piña-Rodrigues, F. C. M. (2026). Imbibition patterns in native seeds: insights for direct seeding in restoration ecology. Ciencia e Natura, 48, e90607. DOI: 10.5902/2179460X90607. Available in: https://doi.org/10.5902/2179460X90607