Hydropriming-induced hydration memory and seedling performance of Cenostigma pyramidale [(Tul.) Gagnon & G.P. Lewis] of different provenances under salinityABSTRACT:
Cenostigma pyramidaleis an endemic Caatinga species, and plays an important role in ecological restoration, but seedling production in semi-arid nurseries is constrained by water deficit and the use of brackish irrigation water. This study evaluated germination, seedling emergence, and early growth ofC. pyramidaleseedlings obtained from seeds harvestedin three populations from the Caatinga, subjected to hydropriming and grown under irrigation with non-saline tap water or brackish groundwater. Seed physiological quality and imbibition patterns were assessed to define population-specific priming protocols (hydropriming, HD). The experiment followed a completely randomized design in a 2 × 2 × 3 factorial scheme, with four replications of 25 seeds, considering the factors: HD cycles, irrigation water quality, and seed lots. After 90 days in the nursery, seedlings irrigated with tapwater showed better growth, whereas brackish water reduced performance in all populations. Hydropriming increased emergence under salinity but did not improve subsequent seedling growth. Differences among populations indicate intrinsic variation in vigor and stress response.
Index terms:
Brazilian semi-arid region; Caatinga; physiological conditioning; salt stress; seedling emergence
RESUMO:
Cenostigma pyramidale é uma espécie endêmica da Caatinga, importante para a restauração ecológica, porém a produção de mudas é limitada, no semiárido, pelo déficit hídrico e pelo uso de água salobra em viveiros. Este estudo avaliou a germinação, a emergência e o crescimento inicial de plântulas deC. pyramidaleobtidas de sementes de três populações na Caatinga, submetidas ao hidrocondicionamento e cultivadas sob irrigação com água tratada ou salobra. A qualidade fisiológica das sementes e os padrões de embebição foram avaliados para definir protocolos específicos de priming (hidrocondicionamento, HD). O experimento foi conduzido em delineamento inteiramente casualizado, em esquema fatorial 2 × 2 × 3, com quatro repetições de 25 sementes, avaliando-se os fatores: ciclos de HD, qualidade da água de irrigação e lotes de sementes. Aos 90 dias em viveiro, mudas irrigadas com água tratada apresentaram maior crescimento, enquanto a irrigação com água salobra reduziu o desempenho em todas as populações. O hidrocondicionamento aumentou a emergência sob salinidade, porém não promoveu ganhos no crescimento posterior. As diferenças entre populações indicam variação intrínseca de vigor e de resposta ao estresse.
Termos para indexação:
Semiárido brasileiro; Caatinga; condicionamento fisiológico; estresse salino; emergência de plântulas
INTRODUCTION
Ecological restoration in the Caatinga, an exclusively Brazilian phytogeographic domain located in the Seasonally Dry Tropical Forests (SDTF), is conditioned by high rainfall irregularity, which limits water availability and reduces the effectiveness of vegetation restoration actions (Dryflor et al., 2016; Silva et al., 2017). In the context of a semi-arid region, this limitation extends to the seedling production phase, as nurseries often depend on lower-quality water sources, including brackish water, which can compromise the initial performance of the species.
Given the stochasticity of water availability regarding quantity and quality, recruitment windows in the field are short and discontinuous. Thus, germination and initial establishment act as functional bottlenecks. This reinforces the need for management strategies aimed at the seed phase, capable of bringing forward and modulating responses to water stress (Rito et al., 2017; Silva et al., 2017; Dantas et al., 2020). In this context, physiological conditioning (priming), especially hydropriming, has been pointed out as a promising alternative to increase the speed and uniformity of emergence and improve performance under limiting conditions, such as water deficit and salinity (Lima et al., 2018).
Among the native species of this ecosystem, the endemic Cenostigma pyramidale [(Tul.) Gagnon & G.P. Lewis], known in Portuguese as catingueira-verdadeira, stands out for its wide ecological plasticity and adaptation to adverse conditions (Gagnon et al., 2016; Dantas et al., 2019; Flora do Brasil, 2020). Due to its multiple uses and utilization in restoration of degraded areas, the sustainable production of the species is strategic for the bioeconomy of the semi-arid region and for conservation through use (Matias et al., 2019). However, its germination and vigor can be severely compromised in years of drought, highlighting the need for more refined protocols for seedling production (Silva et al., 2017; Khan et al., 2022).
In addition to the scarcity of rainfall, nursery practices in the region are constrained by water quality, since water is often brackish due to its origin from wells and reservoirs, which adds an ionic component to the osmotic fluctuations already existing in the substrate (Dantas et al., 2019).
As osmotic stress is common to both salinity and water deficit, understanding the physiological responses of seeds to hydration-dehydration (HD) cycles, characteristics of seasonal soils, becomes fundamental. Prior exposure to these cycles can induce “hydration memory”, which allows seeds to modulate their metabolism and germinate more quickly and uniformly under stresses (Lima et al., 2018; Nascimento et al., 2021).
In this context, hydropriming emerges as a promising technique for simulating, in a controlled and low-cost way, the natural HD cycles (Hora and Meiado, 2016), although studies evaluating its potential to increase tolerance to moderate salinity levels are still scarce in the Caatinga.
In view of this gap, it was hypothesized that hydration-dehydration (HD) cycles induce hydration memory in C. pyramidale seeds, favoring emergence and seedling development under salt stress. The objective was to evaluate the effect of these cycles on seeds of different populations of this species, in order to support seedling production protocols for semi-arid conditions.
MATERIAL AND METHODS
The study was conducted using C. pyramidale seeds, harvested in September 2019, from parent plants of three populations located in different municipalities of the Bahia state: Quixabeira (11°27’57” S; 40°03’04” W; average temperature of 24.1 °C; annual rainfall of 617 mm), Andorinha (10°21’18” S; 39°45’50” W; average temperature of 24.7 °C; annual rainfall of 466 mm) and Senhor do Bonfim (10°35’34” S; 40°01’26” W; average temperature of 23.1 °C; annual rainfall of 768 mm).
In each population, 30 healthy motherplants were randomly selected and fruits were harvested from different partsof the tree crown, at the physiological maturity stage (brownish color), before the onset of natural dispersal.
Seeds of the three populations were kept in separate lots, identified by the municipality of origin, packed in polyethylene bags and stored in a cold chamber (10 ± 3 °C; 60 ± 4% RH) for 30 days, as a procedure for standardizing the post-collection period and maintaining the physiological quality until setting up the experiments in the laboratory and greenhouse. Considering the low water content of the seeds and their orthodox behavior, this storage condition reduced the rate of deterioration and seeds showed no change in vigor.
Prior to the experiments, the physical and physiological quality of the seeds of all populations were initially evaluated.
Thousand-seed weight (TSW): determined by random sampling, using eight replications of 100 seeds for each population (Brasil, 2025).
Water content: determined by the oven method at 105 ± 3 °C, for 24 hours, with two replications of 50 seeds (Brasil, 2025).
Germination test: the lots of each population were subjected to the germination test in four replications of 25 seeds. To prevent fungi contamination during experiments, seeds were disinfected with the commercial fungicide Dithane® NT, with contact action, belonging to the chemical group alkylenobis (dithiocarbamate). The product has mancozeb as its active ingredient, with a multi-site contact mechanism (Brasil, 2024). For the treatment, a solution was prepared with 0.5 g of the product dissolved in 500 mL of distilled water (adapted from Brasil, 2024). The seeds homogenized with the solution until all were uniformly coated.
After disinfection, the seeds were sowed in germination paper rolls and kept in a B.O.D. germination chamber, under a 12-h photoperiod and a constant temperature of 25 °C, with evaluation at 10 days after test setup (Brasil, 2025).
The variables evaluated at the end of the germination test were: percentage of primary root emergence (PRE), normal seedlings (NS), shoot length (SL) and root length (RL), as well as shoot fresh and dry mass (SFM, SDM), root fresh and dry mass (RFM, RDM) and cotyledon fresh and dry mass (CFM, CDM).
Determination of the imbibition curve: carried out using 100 seeds per population, in four replications of 25 seeds. The seeds were weighed on an analytical scale and arranged in Petri dishes (15 cm in diameter) on a double layer of Germitest paper moistened with 15 mL of distilled water, kept on a bench (27 ± 4 °C; 45 ± 3% RH) with weighing every 2 h, until 10% of the seeds showed root protrusion.
Prior to each weighing procedure, excess surface moisture was removed with absorbent paper, and after weighing, the seeds were placed back on the substrate moistened with distilled water. Imbibition (mL) was estimated over time, taking the initial weight of the samples as reference and converting the weight variation into volume of water absorbed (mL).
Hydration-dehydration (HD) cycles: Based on the imbibition curve, the hydration times were defined at 9, 13 and 11 h, corresponding to half of phase I for the populations of Andorinha, Senhor do Bonfim and Quixabeira, respectively. The dehydration curve was obtained by the time required for the seeds to return to the initial mass, using 100 seeds (four replications of 25) previously hydrated in distilled water, distributed in plastic trays at room temperature, with weighing every 60 min after removal of surface moisture with absorbent paper. The definition of HD cycles was based on Nascimento et al. (2021), with adaptations.
Seedling production: the seeds were previously subjected to hydration-dehydration (HD) cycles to check whether hydropriming influences the tolerance of seedlings to salinity. The experimental design was completely randomized, in a 2 × 2 × 3 factorial scheme, with four replications of 25 seeds, considering three factors (i) HD cycles: C0 (without HD) and C3 (three cycles); (ii) irrigation water quality: treated water (TW) and brackish water (BW); and (iii) seed lots from the populations of Quixabeira, Senhor do Bonfim and Andorinha.
The seeds of each population were subjected to HD cycles, in which hydrations had the durations defined in the imbibition curve. Subsequently seeds were sown in polyethylene bags (15 × 20 cm, approximately 1.43 L) containing substrate consisting of soil and goat manure in the proportion 1:1 (v/v). Five seeds were sown in each polyethylene bag at a depth of 1.0 cm. The nursery experiment was conducted, under a 50% shade screen at nursery of Embrapa Semi-arid Region, Petrolina, Pernambuco State, Brazil (09°04’16”S; 40°19’05”W; 379 m).
Chemical characterization of the substrate: the soil used, classified as Argissolo Amarelo (Ultisol) (Santos et al., 2018), was collected in the surface layer (0-20 cm) and sieved to remove plant impurities. The organic component (goat manure), from the same area, was subjected to an aging period under irrigation for one week, aiming at leaching excess ammonia before incorporation into the substrate.
The substrate showed a neutral to slightly alkaline reaction (pH 7.6), absence of exchangeable Al³⁺ and potential acidity, and base saturation of 100% (Table 1). P and exchangeable base contents characterize a chemically fertile substrate, with no limitations of acidity or toxicity by Al and with adequate availability of essential nutrients.
Irrigation of seedlings with water of different salinities: irrigation was carried out up to field capacity, three times a week, using local-supply water (treated water - TW) or brackish water (BW) from an artesian well located in the Caatinga Experimental Field, of Embrapa Semi-arid Region (09°03’53” S; 40°19’51” W).
The brackish water used showed high electrical conductivity (7.19 dS.m⁻¹), alkaline pH (8.50) and high concentrations of Na⁺, Mg²⁺ and Cl⁻, constituting a solution with high potential for inducing osmotic and ionic stress in seeds and seedlings (Table 2), classified as C3S2 (high salinity and medium proportion of sodium) (Dantas et al., 2025). The treated water, classified as C1S1 (low salinity and low risk of sodification), showed low electrical conductivity, pH close to neutrality and low levels of Na⁺ and Cl⁻.
Evaluation of seedling quality: seedling emergence was recorded daily during the 21 days following experiment setup. At 90 days after sowing, the following variables were evaluated: number of leaves (NL), plant height (PH), stem diameter (SD), shoot fresh mass (SFM), root fresh mass (RFM), shoot dry mass (SDM) and root dry mass (RDM).
Statistical analysis: Emergence variables were obtained from the daily counts of emerged seedlings, using the SeedCalc package (Silva et al., 2019) in R software (R Core Team, 2023), with determination of the percentage of final emergence (FE), emergence speed index (ESI), mean emergence time (MTE) and synchrony index (SYNC). The data were subjected to tests of normality (Shapiro and Wilk, 1965) and homogeneity of variances (Levene, 1960), at 5% significance level, and analyzed in R with the ExpDes.pt package (Ferreira et al., 2018), suitable for factorial designs.
RESULTS
Thousand-seed weight (TSW) varied among the lots, being equal to 198.19 g in Quixabeira, 156.54 g in Senhor do Bonfim and 165.29 g in Andorinha, with coefficients of variation of 2.98%, 3.19% and 3.15%, respectively. The initial water content ranged from 7.79 to 9.93%, a typical range for orthodox seeds, with similar values among the three populations (Table 3).
Among the seed lots, Quixabeira showed 100% primary root emergence, similar to Senhor do Bonfim (94%) and higher than Andorinha (86%) (Table 3). Quixabeira also showed a higher percentage of normal seedlings (96%) and superior shoot performance, with SL and SFM comparable to Andorinha and higher than Senhor do Bonfim, along with greater SDM than the other populations, indicating enhanced vigor and establishment potential.
No significant differences were observed between the lots for root growth (RL, RDM, RFM), nor in cotyledon biomass (CDM, CFM), suggesting that these variables are less influenced by the origin of the seeds.
The imbibition curves of the three lots showed a three-phase pattern (Figure 1). The Andorinha lot exhibited the fastest imbibition, with Phase I up to 18 h, Phase II for about 10 h, and Phase III starting at 28 h (Figure 1 A).
Imbibition curves of catingueira-verdadeira (Cenostigma pyramidale (Tul.) Gagnon & G.P. Lewis) seeds from three populations: Andorinha (A), Quixabeira (B) and Senhor do Bonfim (C). The dots indicate the mean values of imbibition (mL) as a function of time (h) and the continuous line represents the fit. The dashed vertical lines delimit phases I (initial rapid imbibition), II (stationary phase) and III (new increment in absorption associated with the beginning of germination). The shaded band corresponds to the 95% confidence interval.
Quixabeira (Phase I up to 22 h; beginning of Phase III after 36 h) and Senhor do Bonfim (Phase I up to 26 h; beginning of Phase III after 36 h) lots showed slower imbibition than Andorinha (Figures 1 B and 1 C). Based on the duration of Phase I, the hydration time for HD cycles was defined as its half (Nascimento et al., 2021), resulting in 9 h for Andorinha, 11 h for Quixabeira, and 13 h for Senhor do Bonfim.
For all emergence variables (FE, ESI, MTE and SYNC), there was a triple interaction between population, hydropriming and water quality (Table 4). Irrigation with treated water (TW) resulted, in general, in higher percentages of emergence than brackish water (BW), except for Quixabeira without hydropriming, in which the differences were inconsistent. Emergence synchronization (SYNC) was mainly influenced by water quality and interactions between factors, and hydropriming had a positive effect on only part of the populations.
Final emergence percentage (FE) and emergence performance indices (ESI, MTE and SYNC) of seedlings of catingueira-verdadeira (Cenostigma pyramidale (Tul.) Gagnon & G.P. Lewis) from three populations, as a function of irrigation with brackish water (BW) or treated water (TW) and the application of hydration-dehydration (HD) cycles of the seeds.
Despite showing the lowest vigor in the germination test in the laboratory (Table 3), the Andorinha population had the best performance in the nursery when subjected to hydropriming. Its hydroprimed seeds reached 87% emergence with treated water (HD-TW) and 60% emergence with brackish water (HD-BW), surpassing the other populations under both conditions. Under these conditions, emergence was faster (higher ESI, lower MTE) and more synchronized, indicating that, for Andorinha, hydropriming acted as an attenuator of the salt stress imposed by irrigation with brackish water (Table 4).
Seedlings of the Senhor do Bonfim population did not respond to hydropriming, with similar emergence percentages under irrigation with treated water (approximately 50%) and with brackish water (approximately 40%). However, higher speed and synchrony of emergence was observed with treated water, especially in seeds without hydropriming (WoHD-TW), compared to irrigation with brackish water (Table 4).
In the Quixabeira population, whose seeds showed high vigor in the laboratory (Table 3), the responses to hydropriming and irrigation water quality were more stable. Seedling emergence, as well as speed and synchrony, was similar between treatments, suggesting greater tolerance to salinity conditions and less dependence on hydropriming in this initial phase of seedling production (Table 4).
Regarding seedling performance, hydropriming did not influence growth, but there was an interaction between populations and water quality for plant height, stem diameter, number of leaves (NL), shoot fresh mass (SFM) and shoot dry mass (SDM) (Table 5).
Irrigation of seedlings with treated water (TW) allowed greater growth in all morphological variables and for all populations evaluated in comparison to irrigation with brackish water (BW). Among the populations, significant differences were detected only under TW and restricted to the characteristics of the aerial part. Under this condition, Andorinha and Quixabeira had the highest values of plant height (statistically similar to each other), while Andorinha surpassed Quixabeira and Senhor do Bonfim in stem diameter, NL, SFM and SDM. Under salinity conditions, the populations did not differ from each other for any of the variables, indicating that salt stress homogenized the performance of seedlings between populations (Table 5).
For the root variables, root fresh mass (RFM) and root dry mass (RDM), there were no significant differences between populations under any water quality, although, in all cases, irrigation with TW promoted values higher than those obtained with BW (Table 5).
DISCUSSION
The initial physiological quality of C. pyramidale seeds varied among the populations, reflecting differences in biometric parameters, initial shoot growth and imbibition pattern, with effects on germination, vigor, emergence and nursery performance. This pattern reinforces that the response to hydropriming and salt stress depends not only on the treatment applied, but also on the intrinsic characteristics of each lot, such as its initial physiological condition and the way the seed resumes metabolism during imbibition. This behavior is consistent with that observed for other native species of the Caatinga with different provenances and subjected to different priming techniques (Ferreira et al., 2022; Medeiros et al., 2022).
The thousand-seed weight (TSW) values of the evaluated populations were among the highest ever reported for the species. The wide variation observed, both between populations and compared to the literature (Lima et al., 2014; Matias et al., 2019), indicates high intraspecific variability in size and accumulation of reserves, possibly associated with genetic, environmental, and provenance factors (Campos et al., 2020; Medeiros et al., 2022; Dutra-Silva et al., 2024; Silva et al., 2025).
In physiological terms, differences in seed size and reserve content can influence the speed of water absorption, the mobilization of reserve compounds, and the ability to sustain initial seedling growth under adverse conditions. Like other biometric characteristics, TSW is an important parameter for assessing the physiological quality of the lot and for planning sowing and seedling production (Acchile et al., 2017; Ferreira et al., 2021).
The imbibition dynamics confirmed the three-phase pattern typical of orthodox seeds, but the duration of the phases and the time to root protrusion varied among populations, indicating differences in the physiological behavior of the lots. These differences are relevant because hydropriming acts precisely in the initial stages of imbibition, when membrane reorganization, respiratory reactivation, and resumption of metabolic processes essential for germination occur. Thus, variations in imbibition kinetics can modify treatment responsiveness, since lots that absorb water at different rates can also differ in the speed at which they reconstitute the functional integrity of membranes and resume germinative metabolism. This variability may be associated with intrinsic differences in seed coat permeability and reserve content (Dantas et al., 2008; Campos et al., 2020), modulating both initial water absorption and hydropriming response (Acchile et al., 2017).
Although the seeds from Quixabeira population showed higher initial physiological vigor, their response to hydropriming was modest. In contrast, seeds of Andorinha responded more intensely to the treatment, especially under saline conditions, which suggests greater sensitivity of germination metabolism to hydropriming for this provenance. This result is consistent with the hypothesis that priming especially favors seeds in which the initial metabolic resumption can be benefited by controlled hydration, with possible improvement in membrane reorganization, in the efficiency of reserve use and in tolerance to osmotic stress. On the other hand, in lots of higher initial physiological quality, such as Quixabeira, the lower response to the treatment may indicate less margin for additional gains, since part of the processes favored by hydropriming may occur naturally with higher efficiency (Ferreira et al., 2021; Lallouche and Kouider, 2024; Hameed et al., 2025).
The Senhor do Bonfim population showed lower responsiveness to hydropriming and more modest performance in part of the variables evaluated, suggesting lower physiological plasticity under the conditions imposed in the nursery. Although biochemical or ultrastructural parameters have not been directly evaluated, this behavior may be associated with lower efficiency in metabolic resumption during imbibition or with a lower ability to maintain cellular functioning under prolonged salt stress (Dantas et al., 2008; Farooq et al., 2015). Thus, the results indicate that hydropriming does not act uniformly between provenances, and its effectiveness depends on the initial physiological state of the lot and the interaction of this state with the cultivation environment.
In C. pyramidale, hydropriming acted mainly as a modulator of performance at emergence, attenuating initial limitations in part of the lots, instead of promoting consistent gains in all of them. In general, the results of this study reinforce that both the hydropriming of the seeds and the quality of irrigation water influence seedling emergence, but the intensity of the response is modulated by the initial physiological quality of each lot (Sousa et al., 2024).
Brackish water reduced emergence and initial growth, a result consistent with the combined effect of the osmotic and ionic components of salinity, widely described for C. pyramidale and other woody species of the Caatinga (Dutra et al., 2017; Matias et al., 2019; Dantas et al., 2019; Santos et al., 2021). In the first stage, salinity reduces the water potential of the substrate, making it difficult for the seeds to absorb water and delaying imbibition and root protrusion. Next, the accumulation of ions, especially Na⁺ and Cl⁻, can compromise ionic homeostasis, interfere with the uptake of nutrients such as K⁺ and Ca²⁺, and affect the structural and functional stability of cell membranes. This set of effects helps explain the lower emergence speed and reduced growth observed under brackish water irrigation.
In addition to osmotic and ionic effects, high salinity often favors increased production of reactive oxygen species (ROS) (Oliveira et al., 2025). At moderate levels, ROS participate in cell signaling associated with germination; however, when produced in excess, they can induce lipid peroxidation, loss of membrane selectivity, protein degradation and damage to energy metabolism (Padilha et al., 2025). Although these parameters were not measured in the present study, the reduction of emergence under brackish water and the lower accumulation of biomass in the seedlings are consistent with a scenario of oxidative stress associated with salt stress, as described for seeds and seedlings of other species (Farooq et al., 2015; Lallouche and Kouider, 2024; Hameed et al., 2025). In this context, the ability to maintain efficient antioxidant systems may be one of the factors involved in the differential response between populations.
The positive effect of hydropriming on emergence, especially in Andorinha and under salinity, may be related to the pre-activation of metabolic processes that favor a more efficient resumption of germination. Among the mechanisms described for priming are the reorganization of membranes during rehydration, the reduction of solute leakage, the faster activation of enzymes associated with the mobilization of reserves, and the strengthening of antioxidant defense mechanisms (Farooq et al., 2015; Lallouche and Kouider, 2024). It is also plausible that the treatment contributes to the initial osmotic adjustment, through the more efficient mobilization of compatible solutes, favoring the maintenance of water absorption even under more negative water potentials. Thus, the better performance of part of the lots after hydropriming is consistent with greater tolerance to osmotic stress at the time of emergence, although these mechanisms have not been directly evaluated in this study.
Although hydropriming favored emergence in part of the treatments, this effect was not maintained in the growth of seedlings at 90 days. In this phase, water quality was the predominant factor, and irrigation with brackish water reduced growth in all populations, besides attenuating the differences between provenances. This result indicates that the main effect of hydropriming was concentrated in the pre-germination phase and in the initial establishment, while the subsequent performance began to depend more on the intensity and duration of stress in the nursery than on the initial benefit promoted by the treatment.
In the nursery phase, treated water promoted greater height, diameter, number of leaves and accumulation of aerial biomass in all lots, while brackish water consistently reduced these attributes. This pattern confirms that, even in species of the semi-arid region with some tolerance to salinity, excess salts limit physiological processes that are fundamental to initial growth, such as water absorption, cell expansion, leaf emergence, and biomass production (Sousa et al., 2024; Hameed et al., 2025). Under continuous exposure to salt, the initial osmotic stress tends to add to the progressive ionic effect, which intensifies limitations to metabolism and can reduce the plant’s ability to sustain active growth for longer periods.
Studies that used seedlings irrigated with brackish and biosaline water (≈ 6 dS.m⁻¹, C2S2), for up to 70 days, reported low or no impact of salinity on growth (Dantas et al., 2019; Dantas et al., 2025). In the present study, however, the combination of more saline water (> 7 dS.m⁻¹, C3S2) and exposure for 90 days resulted in marked reductions in seedling growth. This difference suggests that the intensity and duration of stress were determinant for the expression of the negative effects of salinity, surpassing any initial gains promoted by hydropriming.
High salinity reduces plant growth mainly by decreasing the osmotic potential of the soil solution, making it difficult for plants to absorb water, in addition to causing specific ionic effects, such as toxicity and nutritional imbalances (Richards, 1954). Thus, the higher salinity of the water, combined with the longer time of exposure, probably intensified the mechanisms of osmotic, ionic and oxidative stress, generating cumulative effects on water absorption, ionic balance and seedling growth (Dutra et al., 2017; Dantas et al., 2019; Santos et al., 2021; Hameed et al., 2025). Under conditions like these, any initial differences in vigor between lots tend to lose expression, as the saline environment starts to impose a dominant limitation on development.
Another relevant aspect is that, under high and prolonged salinity, the differences between populations became less evident in the growth of seedlings, indicating that salt stress homogenized the performance among provenances. Thus, nurseries that depend on brackish water should not only select lots of higher physiological quality and, when relevant, use hydropriming to improve emergence, but also adopt mitigation strategies, such as water dilution, periodic use of better quality water, and substrate management to reduce salt accumulation in order to preserve seedling production potential (Oliveira et al., 2019; Khan et al., 2022).
In summary, the results reinforce recommendations for restoration with C. pyramidale in the semi-arid region: (i) prioritize lots with high physiological quality for seedling production; (ii) employ hydropriming as a strategy to improve emergence and its synchronization in responsive lots; and (iii) adopt water management practices that minimize the osmotic and ionic effects of salinity. Although mechanisms such as osmotic adjustment, membrane integrity, ROS production, and antioxidant systems have not been directly measured, the observed pattern is consistent with their participation in the differential response between populations and in the action of hydropriming. The incorporation of multiple lots and parent plants should be considered to maximize genetic diversity and population resilience in restoration plantations, articulating the physiological findings of this study with guidelines on provenance and patterns of native seeds (Khan et al., 2022; Medeiros et al., 2022; Silva et al., 2025) and with the recent literature on hydropriming and salinity tolerance in native species (Ferreira et al., 2022; Hameed et al., 2025).
CONCLUSIONS
The emergence and initial performance of Cenostigma pyramidale seedlings varied among the evaluated populations. Hydropriming favored emergence under salinity in part of the lots, with a more evident effect on the Andorinha population, but did not promote gains in seedling growth at 90 days. Irrigation with brackish water reduced growth in all populations and attenuated the differences between provenances in the nursery phase. Thus, the results indicate that hydropriming can contribute to improving the emergence of responsive lots, while the quality of irrigation water is determinant for seedling growth.
ACKNOWLEDGMENTS
The authors thank the Coordination for the Improvement of Higher Education Personnel (CAPES; Financing Code 001), the Financier of Studies and Projects - FINEP (Grant Agreement No. 01.22.0614.00), the Pernambuco State Science and Technology Support Foundation - FACEPE (Postdoctoral Fellowship - PROCESS No.: BFP-0072-5.02/23), the State University of Feira de Santana (UEFS) for the support and to Embrapa Semi-arid Region.
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DATA AVAILABILITY
Data will be made available as soon as approved at: Dantas, B.F., 2025, Produção de mudas da Caatinga., https://doi.org/10.48432/F1D0B6, Redape, Embrapa.
Data will be made available as soon as approved at: Dantas, B.F., 2025, Produção de mudas da Caatinga., https://doi.org/10.48432/F1D0B6, Redape, Embrapa.
