Influence of endophytic bacteria on seed germination and initial growth of camu-camu treeAbstract:
Camu-camu (Myrciaria dubia [Kunth.] McVaugh) is an Amazonian fruit species of high nutritional and economic value, mainly due to its high vitamin C content and bioactive compounds. However, its propagation presents limitations, including low rooting rates in cuttings, genetic variability from sexual propagation,and high contamination in micropropagation systems. In this context, endophytic bacteria emerge as a sustainable alternative to stimulate plant growth through direct mechanisms, such as phytohormone production and biological nitrogen fixation, and indirect mechanisms, including the suppression ofphytopathogens. This study aimed to evaluate the effect of endophytic bacteria on seed germination and early growth of M. dubia. Seeds were surface-sterilized and inoculated in bacterial suspensions prepared in saline solution and maintained at 28°C for 120 minutes, under a completely randomized design with five treatments: Bacillus safensis, Bacillus altitudinis, , Bacillus pumilus,Brevundimonas sp.,and a non-inoculated control. The variables evaluated were germination, plant height,stem diameter, number of leaves, root length, and germination speed index. No significant differences were observed for germination, height, diameter, and number of leaves. However, root length showed significant increase, with B. safensis outperforming B. pumilus and the other treatments. The germination speed index was also higher for B. safensis compared with B. pumilus. Theseresults indicate that Bacillus strains have potential to enhance root development and seedling vigor,contributing to sustainable strategies for camu-camu propagation.
Index terms
Amazon; Bacillus safensis; plant growth-promoting bacteria; bioeconomy; Myrciaria dubia
Resumo:
O camu-camuzeiro (Myrciaria dubia [Kunth.] McVaugh) é uma frutífera amazônica de elevado valor nutricional e econômico, destacando-se pelo alto teor de vitamina C e compostos bioativos. Entretanto, sua propagação apresenta limitações, comobaixa taxa de enraizamento por estaquia, variabilidade genética oriunda da propagação sexuada e elevada contaminação em sistemas de micropropagação. Nesse contexto, o uso de bactérias endofíticas surge como uma alternativa sustentável para estimular ocrescimento vegetal por mecanismos diretos, como produção de fitohormônios e fixação biológica de nitrogênio, e indiretos, como a supressão de fitopatógenos. O objetivo deste estudo foi avaliar o efeito de bactérias endofíticas na germinação e na promoção do crescimento inicial de plântulas de M. dubia. As sementes foram desinfestadas e inoculadas em suspensões bacterianas em solução salina, mantidas a 28 °C por 120 minutos, em delineamento inteiramente casualizado, com cinco tratamentos: Bacillus safensis, Bacillus altitudinis,Bacillus pumilus, Brevundimonas sp. e controle sem inoculação. Foram avaliadas as variáveis germinação,altura, diâmetro, número de folhas, comprimento radicular e índice de velocidade de germinação. Não houve diferenças significativas para germinação, altura, diâmetro e número de folhas. Entretanto, observou-se efeito significativo para ocomprimento radicular,com superioridade de B. safensis em relação a B. pumilus e aos demaistratamentos. O índice de velocidade de germinação também foi superior para B. safensis. Os resultados indicam que cepas de B. safensis apresentam potencial para promover odesenvolvimento radicular e o vigor inicial de plântulas, contribuindo para estratégias sustentáveis de propagação do camu-camuzeiro.
Termos para indexação
amazônia; Bacillus safensis; promotoras de crescimento; bioeconomia; Myrciaria dúbia
Introduction
Camu-camu tree [Myrciaria dubia L. (Kunth.) McVaugh] is a native fruit tree species of high nutritional value and economic potential in the Amazon region, standing out for its richness in bioactive compounds present in the pulp, peel, and seeds, such as flavonoids and anthocyanins, which are widely used in the pharmaceutical, food, and cosmetic industries (GRIGIO et al., 2024). The pulp exhibits exceptionally high vitamin C contents, reaching up to 7,355 mg of ascorbic acid (GRIGIO; DURIGAN; CHAGAS, 2019).
However, its consumption remains limited due to its high acidity and astringency, associated with elevated concentrations of phenolic compounds (CHAGAS et al., 2023).
Native to the western Amazon basin, the species occurs in floodplain areas and along riverbanks, playing a relevant ecological and socioeconomic role, especially for riverside communities; however, commercial production faces constraints related to the scarcity of high-quality propagative material (PINEDO et al., 2013).
Endophytic bacteria inhabit plant tissues without causing apparent damage to the host plant, establishing beneficial symbiotic relationships in different organs, including seeds (GUHA et al., 2024). These microorganisms contribute to the maintenance of seed viability, improvement of germination, increased tolerance to early stresses, and formation of the seedling microbiome, and colonization is influenced by the interaction between plant and bacterial genotypes (GUHA et al., 2024).
These bacteria act as plant growth-promoting bacteria (PGPB) by producing phytohormones essential for early plant development (HOSSAIN et al., 2023). Auxin, especially indole- 3-acetic acid, promotes root development and nutrient uptake and is produced by genera such as Arthrobacter, Azospirillum, Bradyrhizobium, Bacillus, Pseudomonas, and Paenibacillus (HOSSAIN et al., 2023).
The production of gibberellins by microorganisms such as Sphingomonas spp. stimulates shoot elongation, whereas Bacillus amyloliquefaciens stands out for abscisic acid production, regulating dormancy and early growth (HOSSAIN et al., 2023). Cytokinins synthesized by Pseudomonas resinovorans and Paenibacillus polymyxa promote cell division, and some bacteria modulate ethylene levels, influencing germination and stress responses, such as Pseudomonas fluorescens, Variovorax paradoxus, Paenibacillus lentimorbus, and Enterobacter cloacae (HOSSAIN et al., 2023). Therefore, the objective of this study was to determine the effect of endophytic bacterial strains on germination and plant growth promotion in camu-camu seeds.
Material and methods
Seed Collection
All seeds used in the experiment were obtained from LM30 and BQ12 genotypes that belong to the germplasm bank of Serra da Prata Experimental Field (CESP), Embrapa Roraima, municipality of Mucajaí, and were collected on February 7, 2025, ensuring genetic uniformity and adequate physiological quality for the conduction of the analyses.
The seeds were previously selected manually to ensure uniformity in size, integrity, and absence of visible damage, such as cracks or signs of deterioration (Figure 1).
After this screening, only seeds considered viable and homogeneous were selected in order to reduce experimental variability.
Subsequently, the seeds were immersed in water for 24 h, a procedure that assisted in initial hydration and standardization of germination conditions.
Camu-camu seeds (Myrciaria dubia) previously selected for uniformity in size, integrity, and viability.
Under a laminar flow hood, the seeds were subjected to a disinfestation process, adjusted based on the methodology described by Araujo et al. (2015). The procedure consisted of immersion in 70% ethanol for 1 min, followed by immersion in a 1.5% sodium hypochlorite solution, with addition of three drops of neutral detergent, for 10 min.
Origin of the Bacteria
The bacteria used in the experiment, Bacillus altitudinis, Bacillus safensis, Bacillus pumilus, and Brevundimonas spp., were identified in Myrciaria dubia tissue cultures, as reported in studies conducted at Embrapa Roraima and the Federal University of Roraima (UFRR). Regarding the origin of isolation, these microorganisms were obtained directly from the explants used in micropropagation and were isolated from nodal segments cultivated in vitro.
It is noteworthy that the selection of these microorganisms was justified by their performance observed in previous assays and by the variation in responses according to the tissue in which they act. Isolates that exhibit favorable results in explants may not respond similarly in seeds, since each plant organ presents distinct physiological and microenvironmental conditions (Table 1).
Bacteria collected during the micropropagation process of camu-camu tree (Myrciaria dubia) and identified by 16S rRNA sequence analysis.
For culture preparation, each isolate was initially reactivated on nutrient agar (NA) medium and incubated at 28 °C for 48 h, as the bacteria exhibited rapid growth. From these colonies, a single colony was transferred to tubes containing 10 mL of liquid DYGS medium, selected for promoting the growth and physiological activation of endophytic bacteria. The cultures were maintained under constant agitation (120 rpm) at 28 °C for additional 48 h.
Cell concentration was determined by counting colony-forming units (CFU) using serial dilutions and plating 100 μL onto NA medium, in triplicate. After 48 h of incubation, only plates containing between 30 and 300 colonies were considered for calculating cell density and adjusting the suspensions. Based on these values and on preliminary tests conducted with different concentrations (107, 108, and 109 CFU mL-1), a concentration of 109 CFU mL-1 was established as providing the best results, including greater explant survival, uniform colonization, and more consistent physiological responses throughout in vitro cultivation.
The bacterization time was also defined experimentally, since there are no specific recommendations for inoculating these strains in camu-camu tissues. Preliminary assays demonstrated that immersion of the explants for 60 min under continuous agitation at 120 rpm and 28°C provided the best balance between efficient bacterial adhesion and preservation of plant tissue integrity.
Shorter times resulted in insufficient colonization, whereas longer periods increased oxidation and physiological stress.
Thus, the bacterization process was carried out in Erlenmeyer flasks containing 500 mL of standardized bacterial suspension (109 CFU mL-1) in 0.85% NaCl solution, ensuring complete immersion of the seeds and homogeneous conditions during inoculation.
At the end of the 60 min period, the explants were immediately transferred to in vitro culture media, preventing excessive bacterial multiplication in the culture environment and ensuring treatment standardization according to the experimental design.
This procedure follows the protocol adopted by Ludwig et al. (2013), who applied microbiolization to rice seeds (Oryza sativa) to evaluate the effects of rhizobacteria, using seed immersion in bacterial suspension to ensure initial and uniform colonization of the material.
The seeds were sown in a nursery under 50% shading. Planting was carried out in plastic trays measuring 55 × 38 × 9 cm, using washed sand as substrate. Throughout the experiment, irrigation was performed daily to maintain adequate moisture.
At the end of the experiment, 90 days after sowing, no new seedling emergence was observed for a continuous period of 20 days, characterizing the end of the germination phase. At this stage, the initial growth parameters of camu-camu seedlings were evaluated as follows: (a) Plant height (cm): measured from the base of the stem to the last fully expanded leaf using a ruler; (b) Stem diameter (mm): measured at the lower portion of the stem using a digital caliper; (c) Germination rate (%): percentage of emerged seedlings relative to the total number of seeds sown; (d) Number of leaves: counted as the number of fully expanded leaves; and (e) Primary root length (cm): measured from the stem base to the tip of the primary root using a ruler.
The experimental design was completely randomized, with five treatments: Bacillus altitudinis, Bacillus safensis, Bacillus pumilus, Brevundimonas, and a control treatment (without inoculation), with five replicates.
Each replicate consisted of 25 seeds, totaling 125 seeds per treatment.
To verify the assumptions of analysis of variance (ANOVA), data were tested for normality of residuals using the Shapiro–Wilk test (p > 0.05), homogeneity of variances using the Bartlett test (p > 0.05), and independence of errors using the Durbin–Watson test (p > 0.05). When assumptions were met, data were subjected to analysis of variance using the F-test (p < 0.05). Significant variables were compared using Tukey’s test (p < 0.05). Statistical analyses were performed using R software, version 4.2.2 (R Core Team, 2023).
Results and discussion
It was observed that seeds subjected to the different treatments with plant growth-promoting bacteria did not differ statistically from the control treatment (without inoculation) (p>0.05). Inoculation with the tested strains did not promote expressive increases in these initial growth traits, demonstrating that the bacteria did not cause negative effects on plant growth or germination.
The genus Bacillus is widely recognized as one of the most promising groups in plant growth promotion, since several species exhibit high capacity for rhizosphere colonization and contribute to the availability of essential nutrients to plants, such as phosphorus, iron, and nitrogen, in addition to producing beneficial phytohormones, such as indole-3-acetic acid. Moreover, different bacterial genera, including Azotobacter, Burkholderia, Erwinia, Flavobacterium, Micrococcus, Pseudomonas, Serratia, and Bacillus, also exhibit consistent plant growth-promoting properties (BIZJAK-JOHANSSON, 2025).
It has been reported that bacteria showing promising performance as plant growth promoters under in vitro conditions may not necessarily enhance plant development under greenhouse (in vivo) conditions (BIZJAK-JOHANSSON et al., 2025).
The absence of immediate effects on the evaluated variables, such as plant height, stem diameter, germination rate, and number of leaves, is attributed to the fact that seedlings germinated from camu-camu seeds prioritize root growth before noticeable development of stems and leaves. It was observed that bacterial inoculation significantly influenced primary root length (p< 0.05) (Figure 2A).
Evaluation of primary root length (A) and germination speed index (GSI) (B), showingsignificant effect for the Bacillus safensis treatment. Colored bars represent the means of each treatment (Bacillus altitudinis, Bacillus safensis, Bacillus pumilus, Brevundimonas sp., and Control), with error bars indicating the standard deviation of the means. Means followed by the same lowercase letter do not differ significantly according to Tukey’s test at the 5% probability level.
Inoculation with B. safensisshowed the highest values for root growth (Figure 2A) and germination speed index (GSI) (Figure 2B), differing statistically only from B. pumilus (p <0.05).
These results reinforce the hypothesis that certain endophytic bacteria, especially those belonging to the genus Bacillus, may promote growth preferentially in the root system rather than in the shoot. In addition, they indicate that B. safensis enhances initial seed vigor, possibly through growth-promoting mechanisms associated with phytohormone production, such as auxins, and phosphorus solubilization, thereby accelerating metabolic processes related to germination and root development (PRASAD et al., 2022).
These effects occur because roots represent the main site of bacterial colonization and the first organ to interact with metabolites released by bacteria (SANTOYO et al., 2016). Bacillus strains commonly concentrate in the rhizosphere and root tissues shortly after inoculation, directly stimulating root elongation and branching through phytohormone production, especially indole-3-acetic acid (IAA), and phosphorus solubilization, which promote early root system development and improve water and nutrient uptake (JEYANTHI et al.,2018).
In addition, seed endophytes have shown similar benefits in several plant species, although responses are often more evident in roots than in other organs (VERMA et al., 2021). Among plant growth-promoting rhizobacteria (PGPR), Bacillus spp. stand out due to multiple physiological mechanisms, including the production of ACC deaminase enzyme, which reduces endogenous ethylene levels and improves tolerance to abiotic stress (CHERIF-SILINI et al., 2016).
The combined effects of auxin (IAA) production and ACC deaminase activity stimulate root development and mitigate the inhibitory effects of ethylene, further reinforcing these responses (KHAN et al., 2016). Thus, as described for plant growth-promoting bacteria, the greater sensitivity of roots helps explain the early manifestation of the responses observed in this study (VERMA et al., 2021).
In experiments with seeds of Metrosideros excelsa (Myrtaceae), antimicrobial solutions improved the germination index and promoted seedling root development (KHOLOSTIAKOV et al., 2025). Other studies have reported that inoculation effects may be more strongly expressed in root system vigor than in germination itself (BELLINO et al., 2018).
Furthermore, other researchers have demonstrated that seed-transmitted bacteria and fungi accumulate in roots, where they stimulate the formation of denser and more active root hairs, increasing the absorptive surface area and the efficiency of essential nutrient uptake (VASANTHAKUMARI et al., 2018). Similarly, studies on seedlings of Pinus montezumae and Pinus patula reported that inoculation with strains of the genera Serratia, Buttiauxella, and Bacillus resulted in greater root elongation, increased root number, and enhanced root system diameter, without causing significant effects on shoot height (OROZCO-MOSQUEDA et al., 2021).
During early developmental stages, seedlings allocate most of their energy resources to root system growth, as roots ensure the uptake of water and nutrients necessary for survival (OROZCO-MOSQUEDA et al., 2021). Accordingly, Orozco-Mosqueda et al. (2021) state that the effects of bacterial inoculation tend to be more evident in roots than in shoots during the early stages of growth, because only after establishment of a functional root system can the plant allocate more energy to stem and leaf development.
In addition, Vasanthakumari et al. (2018) emphasize that bacterial colonization is strongly influenced by the specific interaction between the bacterial strain and the host plant species, which may result in localized and distinct responses among different plant organs.
The increase in root length is highly relevant considering the well-known difficulty of rooting in the camu-camu tree, even when hormonal inducers are applied (CHAGAS et al., 2012). Thus, although no significant differences were observed for the other variables, the performance of B.safensis highlights its potential as rooting promoter during the early developmental stages of this species.
Seed inoculation with endophytic bacteria has proven to be a promising strategy for the early development of M. dubia seedlings under nursery conditions. In in vitro studies, isolates from the genera Bacillus, Methylobacterium, and Brevundimonas have been reported to exhibit biotechnological potential that favors seedling rooting (SILVA et al., 2025). In addition, these strains display strong antagonistic activity against phytopathogenic fungi, with inhibition rates exceeding 70%. Therefore, seed inoculation with endophytic bacteria may simultaneously promote plant growth and provide protection against pathogens, representing a sustainable alternative for the production of more vigorous and healthy camu-camu seedlings.
Similar results have been obtained in other plant species. Negi (2023) reported that inoculation of turmeric (Curcuma longa) with B. safensis promoted significant increases in rhizome length and width, attributed to auxin production and phosphorus solubilization.
Likewise, Sharma et al. (2025) observed that application of B. safensis to ginger (Zingiber officinale) resulted in more developed rhizomes, with average length of 21.74 cm and width of 11.57 cm, compared with the control, which reached 18.94 cm and 9.68 cm, respectively, in addition to higher nitrogen and potassium uptake and lower incidence of rot.
In the present study, the Pearson correlation matrix among seedling morphological variables revealed that all traits were positively correlated. Correlation coefficients ranged from moderate to very strong (0.75 to 0.91), indicating that increases in one variable tend to be accompanied by increases in others. Plant height showed strong correlations with root length (0.75), stem diameter (0.83), and number of leaves (0.88), demonstrating that taller individuals also tend to be more vigorous and exhibit greater leaf production (Figure 3).
Pearson correlation matrix among the morphological variables evaluated in seedlings, showing moderate to very strong positive correlations among plant height, root length, stem diameter, and number of leaves. Higher values indicate stronger associations among growth traits.
These results reinforce that seedling growth occurs in an integrated manner, as discussed by González-Paleo et al. (2024), who emphasize the importance of phenotypic integration among roots, stems, and leaves as a mechanism that optimizes resource allocation and promotes greater physiological stability during early development. Complementarily, the results are also consistent with the findings of Shi et al. (2020), who demonstrated that early morphological variables tend to exhibit high correlations in crop plants, serving as reliable predictors of vigor and establishment capacity.
As observed by these authors, the association between greater structural robustness particularly reflected by stem diameter and increased leaf production suggests that these attributes may indicate higher photosynthetic potential and improved early seedling performance.
Conclusion
Inoculation with Bacillus safensis promoted increase in root length and germination speed index (GSI) of camu-camu seeds, confirming its potential as a plant growth-promoting bacterium.
The beneficial effects of B. safensis initially manifested in the root system, characterizing a positive plant–microorganism interaction, highlighting the potential of this species for applications in propagation programs and sustainable management of the camu-camu tree.
Data Availability:
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