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ARTICLE • J. Seed Sci. 47 • 2025 • https://doi.org/10.1590/2317-1545v47296278 linkcopy

Open-access Germination requirements of Diplusodon glaziovii and Barbacenia pungens seeds, endangered species of Brazilian Campo Rupestre

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    ABSTRACT:

    The endemic species Diplusodon glaziovii Koehne (Lythraceae) and Barbacenia pungens Mello-Silva (Velloziaceae) are threatened with extinction due to anthropogenic actions in the Rupestrian Grassland ecosystem. In this context, propagation protocols may assist in the conservation of these species. Therefore, this study aimed to understand the germination requirements of D. glaziovii and B. pungens. Four different temperatures were tested, as well as the germinative response of seeds to growth regulators. For D. glaziovii, germination was faster and achieved higher percentages at 25 °C, whereas for B. pungens, it occurred at 30 °C. In D. glaziovii, fluridone (FLU) reduced germination percentage by 30% at the concentration of 150 µM, and paclobutrazol (PBZ) decreased germination speed at 100 µM. In B. pungens, abscisic acid (ABA) reduced germination speed at all concentrations tested, while PBZ decreased both germination percentage and speed by approximately 30% at 150 µM. These results confirm the absence of physiological dormancy in these species and may contribute to their conservation.

    Index terms:
    biodiversity conservation; Campo Rupestre; hormones; inhibitors; temperature

    RESUMO:

    As espécies endêmicas Diplusodon glaziovii Koehne (Lythraceae) e Barbacenia pungens Mello-Silva (Velloziaceae) encontram-se ameaçadas de extinção em decorrência de ações antrópicas no ecossistema de Campo Rupestre. Nesse contexto, protocolos de propagação podem auxiliar na conservação dessas espécies. Assim, objetivou-se com esse estudo compreender as exigências germinativas de D. glaziovii e B. pungens. Quatro diferentes temperaturas foram testadas, bem como a resposta germinativa das sementes a reguladores de crescimento. Para D. glaziovii, a germinação foi mais rápida e com maior porcentagem a 25 °C, e para B. pungens a 30 °C. Em D. glaziovii, o fluridone (FLU) diminuiu em 30% a porcentagem de germinação na concentração 150 µM e o paclobutrazol (PBZ) diminuiu a velocidade de germinação na concentração 100 µM. Em B. pungens, o ABA diminuiu a velocidade de germinação em todas as concentrações utilizadas, enquanto o PBZ diminuiu em cerca de 30 % a porcentagem e a velocidade de germinação na concentração 150 µM. Esses resultados confirmam a ausência de dormência fisiológica nessas espécies e podem contribuir para sua conservação.

    Termos de indexação:
    conservação da biodiversidade; Campo Rupestre; hormônios; inibidores; temperatura

    INTRODUCTION

    The Campo Rupestre is an open ecosystem characterized by a seasonal climate with rainy summers and dry winters, high vapor pressure deficit, and high radiation. The average annual temperature ranges from 20 to 22 °C, with maximum temperatures reaching 36 °C during the dry season and minimum temperatures ranging from 0 to 4 °C (Silveira et al., 2016; Fernandes et al., 2020, 2021; Miola et al., 2021). These areas exhibit diverse and predominantly grassy vegetation (Silveira et al., 2016; Mucina, 2018). However, due to anthropogenic activities such as open-pit mining, annual anthropogenic fires, expansion of livestock farming, invasive species, excessive exploitation of ornamental plants, and uncontrolled urbanization, these areas have experienced fragmentation, resulting in the loss of numerous plant species. Additionally, conservation efforts in Campo Rupestre have been neglected in conservation programs (Silveira et al., 2016; Le Stradic et al., 2018; Miola et al., 2019; Caminha-Paiva, 2021).

    The species Diplusodon glaziovii Koehne (Lythraceae) e Barbacenia pungens (N. L. Menezes & Semir) Mello-Silva (Velloziaceae) are endemic species of the Serra do Espinhaço, listed as critically endangered in the Red Book of the Flora of Brazil (Silva et al., 2005). Both are part of the set of target species that make up the Territorial Action Plan for the Conservation of Espinhaço Mineiro, which, within the scope of the Pro-Species Project, is one of the instruments to improve the conservation status of species at greatest risk of extinction in Brazil. Considering the scarcity of information about these species, elucidating the germination requirements, as the presence of dormancy and hormone sensitivity, is essential for conservation programs and seedling propagation. Additionally, seed dispersal is the primary means of colonizing new areas, and the distribution of germination in the environment is crucial for community maintenance (Footitt et al., 2013).

    Seed germination is influenced by multiple factors, such as temperature, that affect water absorption, reactivating biochemical processes, and determining the germination velocity (Ramos et al., 2018; Silva et al., 2017). Thus, in the early stages of germination, water absorption under optimal temperatures is important for the seed coat to expand, enabling radicle protrusion (Fu et al., 2021). Therefore, the range temperature also offers a niche for seed germination and seedling establishment.

    Temperature can interact with genes that regulate physiological dormancy, altering the metabolism of key hormones involved in this process: abscisic acid (ABA) and gibberellins (GAs) (Footitt et al., 2013). Thus, the balance between the hormones ABA and GA will induce dormancy or germination, respectively (Leubner-Metzger, 2016). Therefore, for studying germination requirements and hormonal sensitivity, it is essential to use inhibitors of the respective hormones, fluridone (FLU) and paclobutrazol (PBZ) (Desta and Amare, 2021; Worarad et al., 2017). It is well documented that some species of campo rupestre can perform secondary dormancy or conditional dormancy, and the use of germination regulators offers a deep discussion of germination requirements and sensitivity of GA and ABA (Garcia et al., 2020).

    FLU inhibits phytoene desaturase (PDS3), catalyzing the desaturation step of carotenoids, which are the main precursors in ABA biosynthesis (Dong et al., 2015; Kim et al., 2019; Ali et al., 2022). PBZ, on the other hand, is a repressor of plant growth, interfering in the biosynthesis of GAs. It inhibits the oxidation of ent-kaurene into ent-kaurenoic acid through the oxidation of cytochrome-dependent oxygenase (P450), directly affecting the synthesis of GA, as this is synthesized through the terpene pathway. The blocking in the GAs production leads to a greater accumulation of precursors from the terpenoid pathway, which will be diverted to the synthesis of ABA (Desta and Amare, 2021).

    Thus, it is hypothesized that: (i) Constant temperatures between 20-30 °C favor the germination of seeds of D. glaziovii and B. pungens; (ii) The germination percentage and germination speed index of seeds of D. glaziovii and B. pungens are not reduced by exogenously applied ABA or PBZ. The present study aims, therefore, to provide the first report on the germination requirements of seeds of the species D. glaziovii and B. pungens, to propose protocols for the propagation of these endangered species. This study also aims to investigate whether the seeds of these two species present some level of physiological dormancy through testing the germination in the presence of regulators and hormone inhibitors.

    MATERIAL E METHODS

    Different individuals of species populations in the municipalities of Gouvea (18° 31’ 50’’ S and 43° 53’ 45’’ W) and Diamantina 18° 17’ 53’’ S and 43° 44’ 24’’ W), Minas Gerais, Brasil (Figure 1-a, b) were identificated and the seeds of Diplusodon glaziovii Koehne (Lot FPMZB/BS 534) (Figure 1-c) and Barbacenia pungens (N.L.Menezes & Semir) Mello-Silva (Lots FPMZB/BS 434 and 435) (Figure 1-d) were collected in January and July 2023, respectively. The climate is Cwb according to the Köppen classification (adapted for Brazil´s climate), characterized as humid temperate with a dry winter and summer rainfall (Alvares et al., 2013).

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    Figure 1
    Where the species occur (a, b) Diplusodon glaziovii (c) and Barbacenia pungens (d). (source: Equipe PAT Espinhaço Mineiro).

    The experiments were conducted at Universidade Federal de Lavras, Lavras, Brazil. The first experiment assessed the effects of different temperatures on germination parameters of D. glaziovii and B. pungens. The seeds were disinfected with 1.5% sodium hypochlorite (NaClO) for 15 minutes, triple-washed with deionized water, and then sown in Petri dishes lined with germination paper moistened with deionized water. The entire experiment was carried out in germination chambers with 12 hours of light at 40 μM photons m-2 s-1 at constant temperatures of 20, 25, and 30 °C, and an alternating temperature of 35/15 °C.

    In the second experiment, seeds were treated with different concentrations of ABA, FLU ({1-methyl-3-phenyl-5-[3-(trifluoromethyl)phenyl]-4(1H)-pyridinone}), and PBZ ({1-methyl-3-phenyl-5-[3-(trifluoromethyl)phenyl]-4(1H)-pyridinone}) to understand their sensitivity to these compounds. ABA was dissolved in potassium hydroxide (KOH) (Ni and Bradford, 1992), PBZ in ethanol (Tyler et al., 2004) and FLU in acetone (Fong et al., 1983) and then each solution was diluted in distilled water at concentrations of 50, 100, 150, and 200 µM. The seeds were disinfected and transferred to microtubes containing 2 mL of the respective hormone or inhibitor. Deionized water was used as control (0 µM). After 24 hours of imbibition in the dark, the germination test was performed with the optimal temperature defined in the previous experiment.

    Germination (2 mm protruding radicle) was monitored every 24 hours. Were evaluated the germination percentage, germination speed index (GSI) according to Maguire (1962), and average time for 50% germination to occur (T50), according to Ranal and Santana (2006). Independent experiments were conducted for each species, temperature, and hormone/inhibitor in a completely randomized design. Assay 1: consisted of four treatments (temperatures) with four replications of 25 seeds each. Assay 2: consisted of five treatments (concentrations) with five replications of 20 seeds each. Data were analyzed using the R software with ExpDes.pt package (Ferreira et al., 2011) and subjected to the Shapiro-Wilk normality test, analysis of variance (ANOVA), and Tukey’s test at a 5% probability level.

    RESULTS

    The temperatures significantly affected the germinability of D. glaziovii (Figure 2a-c). The temperature of 25°C resulted in a higher percentage and germination speed index, as well as a lower T50. In contrast, the alternating temperature of 35/15°C delayed the onset of germination by approximately five days and reduced the germination percentage by about 50%. On the other hand, B. pungens exhibited the highest germination performance at 30°C (Figure 3a-c), showing an approximately 40% increase in germination speed and a 30% decreased in T50 compared to the other temperatures tested. The temperature of 20 °C and the alternating temperature of 35/15 °C delayed the onset of germination but did not significantly affect the overall germination percentage.

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    Figure 2
    Cumulative germination (a), germination speed index - GSI (b) and time to 50% germination - T50 (C) of the species Diplusodon glaziovii at different temperatures. Data are means ± standard errors (n= 4). Significant differences (p <0.05) are indicated by different letters.

    thumbnail of Figure 3
    Figure 3
    Cumulative germination (a), germination speed index - GSI (b) and time to 50% germination - T50 (C) of the species Barbacenia pungens at different temperatures. Data are means ± standard errors (n= 4). Significant differences (p <0.05) are indicated by different letters.

    For D. glaziovii, ABA did not significantly influence seed germination parameters (Table 1). PBZ significantly affected the speed and time until 50% germination, without impacting the final percentage (Table 1). At concentrations of 100 and 150 µM, seeds exhibited a lower germination speed index (GSI) and a higher T50. On the other hand, FLU had a significant impact on the final percentage and germination speed at a concentration of 150 µM (Table 1). For B. pungens, final germination under the influence of ABA was not significant (Table 2); however, an average delay of 30% was observed speed and an increase T50 of the seeds to germinate at the evaluated concentrations. PBZ had a significant effect on the tested concentrations, with 150 µM reducing 30% germination percentage and delaying germination speed (Table 2). FLU did not show a significant effect on the studied variables at any of the concentrations (Table 3).

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    Table 1
    Cumulative germination (G%), germination speed index - GSI and time to 50% germination - T50 of the species Diplusodon glaziovii and Barbacenia pungens under different concentrations of abscisic acid (ABA). Data are means ± standard errors (n= 5). Significant differences (p <0.05) are indicated by different letters.

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    Table 2
    Cumulative germination (G%), germination speed index - GSI and time to 50% germination - T50 of the species Diplusodon glaziovii and Barbacenia pungens under different concentrations of paclobutrazol (PBZ). Data are means ± standard errors (n= 5). Significant differences (p <0.05) are indicated by different letters. Ns= non-significant data.

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    Table 3
    Cumulative germination (G%), germination speed index - GSI and time to 50% germination - T50 of the species Diplusodon glaziovii and Barbacenia pungens under different concentrations of fluridone (PBZ). Data are means ± standard errors (n= 5). Significant differences (p <0.05) are indicated by different letters.

    DISCUSSION

    This study provides the first report of the seed germination requirements of Diplusodon glaziovii and Barbacenia pungens, subjected to different temperature conditions and concentrations of abscisic acid, fluridone, and paclobutrazol. The higher and faster germination of D. glaziovii at 25 °C indicates that this temperature is closest to optimal. For B. pungens, otherwise, the highest germination speed was set at 30 °C.

    These species germinate within specific temperature ranges which may be associated with the geographic distribution of the species, as there is a correlation with their microhabitats. Barbacenia pungens plants are more associated to rocks (Figure 1d) where temperatures are generally higher than in more shaded microenvironments, where D. glaziovii fruits were collected. It is known that temperature is the most influential microclimatic variable in the soil, it affects both speed and the final percentage of germination, as well as the subsequent protrusion of the radicle and seedling development (Cochrane et al., 2015; Dürr et al., 2015). Therefore, temperature controls the velocity of water uptake by the seeds, leading to biochemical reactivation that shapes the germination process, including the release of dormancy in some seeds (Silva et al., 2017; Ramos et al., 2018; Yan and Chen, 2020).

    The signal generated by temperature in seeds also directly influences their metabolism and signaling of ABA and GA, controlling the balance of these two hormones. In colder seasons, the action of ABA supersedes that of GAs, while the opposite occurs in warmer seasons (Yan and Chen, 2020), resulting in a seasonal pattern for the optimal time of year for germination to occur. However, the sensitivity of seeds to hormones is also fundamental to the germination process (Finkelstein, 2010; Miransari and Smith, 2014). The seeds of D. glaziovii did not show sensitivity to different concentrations of ABA. The sensitivity of certain seeds to ABA gradually decreases in post-maturation, suggesting that this factor controls dormancy and will impact seedling emergence, potentially leading to germination before complete embryo maturation and seed dispersal in the environment (Schramm et al., 2012). Otherwise, in B. pungens, ABA only affected the germination speed, delaying it. Therefore, both species were dispersed without physiological or conditional dormancy. This is an important trait that helps improve biodiversity recovery and protection because the seeds of these species do not need to be stored after dispersion to overcome dormancy. Therefore, B. pungens was more sensitive to exogenous ABA than D. glaziovii, indicating that the former could be tested for secondary dormancy in future work.

    FLU can reverse the inhibition of germination by ABA (Goggin et al., 2009). As observed in the species B. pungens, there were higher germination parameters when exposed to FLU. When GA biosynthesis is blocked, ent-kaurene oxidation occurs, increasing ABA availability (Desta and Amare, 2021). As a result, higher levels of terpenoid pathway precursors accumulate and are diverted, promoting increased ABA synthesis. This dynamic could decrease germination parameters, as observed in D. glaziovii and B. pungens when treated with PBZ, since GA is needed not only for germination induction, but also for cell elongation at the onset of germination and afterwards. However, the germination percentage of D. glaziovii was not affected by PBZ, indicating absence of a potential de novo GA synthesis, resulting in seed germination induction (Vieira et al., 2017). However, in this work, the germination response to FLU in D. glaziovii were not consistent, and the treatments, mainly at 150uM concentration, could be toxic for the seeds, which explains the reduced germination in this tested dose.

    Climatic factors and abrupt temperature changes can interfere with the processes that facilitate germination, as germination characteristics for environmental colonization are specific to each species (Maleki et al., 2024; Oliveira and Garcia, 2011). Breed et al. (2013) emphasizes the need to understand the environment to be revegetated, as climate changes can directly impact this practice. Therefore, by understanding the ideal germination temperature of the species, it is possible to propose a suitable environment for seeding. Considering the specificity of the seeds of species endemic to the Campo Rupestre, further research into the physiology of these seeds is needed, includingD. glazioviiandB. pungens, microendemic species of the Diamantina Plateau, in the Espinhaço Mineiro, whose populations have so far only been found outside protected areas and are under constant threat from anthropogenic actions, especially fire and mining. Thus, it is crucial to understand the impacts of temperature and hormonal sensitivity on their physiological responses, ensuring success in the prospecting and propagation of native species (Garcia et al., 2020). Understanding seed germination requirements is crucial to the success of revegetation efforts, promoting better adaptation of native species to local conditions (Oliveira et al., 2015; Carvalho et al., 2018). Therefore, seed quality and methods to optimize germination are essential for propagation and revegetation efforts (Salazar et al., 2015).

    CONCLUSIONS

    We conclude that D. glaziovii and B. pungens do not present physiological dormancy and they were not sensitive to exogenous ABA. However, the inhibitors influenced the germination of species. The seeds of D. glaziovii germinate better at constant temperature of 25 °C and B. pungens of 30 °C. These results can help on propagation efforts of revegetation programs. The use of PBZ and FLU slightly influenced germination parameters, i.e., germination velocity; however, other experiments are encouraged to better describe this. Indeed, the responses of the application of exogenous PBZ and FLU, were consistent for confirm the absence of primary dormancy in both species.

    ACKNOWLEDGEMENTS

    The authors express their gratitude to the entire team at PAT Espinhaço Mineiro. The support with biological materials was essential, as well as the valuable interactions and knowledge exchanges during the completion of this work. This research was funded by the Global Environment Facility (GEF) through Project 029840 - Estratégia Nacional para a Conservação de Espécies Ameaçadas de Extinção - Projeto Pró-Espécies: Todos contra a Extinção. The Pro-Species Project is coordinated by the Ministry of the Environment and Climate Change (MMA) and implemented by the Brazilian Biodiversity Fund (Funbio), with WWF as the executing agency. The authors received the award of a grant from CAPES (Fundação Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) - JTLC (grant 88881.689011/2022-01). EMB received productivity grant from National Council for Scientific and Technological Development (CNPq, grant 307846/2022-9).

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    • DATA AVAILABILITY
      Additional data will be made available by the authors upon reasonable request

    Edited by

    • Editor:
      Bárbara França Dantas

    Data availability

    Additional data will be made available by the authors upon reasonable request

    Publication Dates

    • Publication in this collection
      08 Dec 2025
    • Date of issue
      2025

    History

    • Received
      25 Apr 2025
    • Accepted
      11 Nov 2025
    Creative Common - by 4.0
    This is an open-access article distributed under the terms of the Creative Commons Attribution License
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