Drought-tolerance mechanisms in rumberry, a native Brazilian fruit species

Mecanismos de tolerância à seca em cambuí, uma espécie frutífera nativa do Brasil

Abstract

The objective of this work was to characterize the physiological and biochemical responses related to drought tolerance in rumberry (Myrciaria floribunda) plants. The experiment was carried out on plants aged about 2.5 years, which were grown in a greenhouse and subjected to two water conditions, as follows: well-watered soil (at field capacity) and water stress (suspension of irrigation for 30 days), with ten replicates per treatment. The following parameters were evaluated: number of leaves; relative water content; electrolyte leakage; photosynthetic pigment concentrations; photochemical efficiency of photosystem II; osmolyte contents; malondialdehyde concentration; and the antioxidant enzymatic activity of superoxide dismutase, catalase, and ascorbate peroxidase. The water stress regime resulted in reduced the relative water content, the potential and effective efficiency of photosystem II, total soluble proteins, and superoxide dismutase activity. No significant changes were observed for the number of leaves, electrolyte leakage, photosynthetic pigment concentrations, and total soluble sugar content, or malondialdehyde concentrations. Conversely, water stress induced increases of total soluble amino acids, free foliar proline, and the activity of catalase and ascorbate peroxidase. The activation of osmoprotective and antioxidant mechanisms, associated with the maintenance of cellular integrity, confers tolerance of rumberry to water stress.

Index terms:
Myrciaria floribunda; antioxidant enzymes; chlorophyll fluorescence; drought tolerance; osmolytes; pomology.

Resumo

O objetivo deste trabalho foi caracterizar as respostas fisiológicas e bioquímicas de tolerância à deficiência hídrica em plantas de cambuí (Myrciaria floribunda). O estudo foi realizado com plantas de cerca de 2,5 anos de idade, cultivadas em casa de vegetação e submetidas a dois tratamentos hídricos, conforme a seguir: solo bem irrigado (em capacidade de campo), e deficiência hídrica (suspensão da irrigação por 30 dias), com 10 repetições por tratamento. Os seguintes parâmetros foram avaliados: número de folhas; teor relativo de água; extravasamento de eletrólitos; teores de pigmentos fotossintéticos; eficiência fotoquímica do fotossistema II; teores de osmólitos; teor de malondialdeído; e atividade das enzimas antioxidantes superóxido dismutase, catalase e ascorbato peroxidase. A deficiência hídrica causou a redução do teor relativo de água, da eficiência potencial e efetiva do fotossistema II, dos teores de proteínas solúveis totais e da atividade da superóxido dismutase. Não foram observadas alterações significativas quanto ao número de folhas, ao extravasamento de eletrólitos, aos teores de pigmentos fotossintéticos, e quanto aos teores de açúcares solúveis totais e de malondialdeído. Em contrapartida, a deficiência hídrica induziu o aumento dos teores de aminoácidos solúveis totais, de prolina livre foliar e da atividade das enzimas catalase e ascorbato peroxidase. A ativação de mecanismos osmoprotetores e antioxidantes, associada à manutenção da integridade celular, confere tolerância ao estresse hídrico em cambuí.

Termos para indexação:
Myrciaria floribunda; enzimas antioxidantes; fluores-cência da clorofila; tolerância à; seca; osmorreguladores; fruticultura.

Introduction

Climate change is already adversely affecting global temperature and precipitation patterns, with negative consequences for agriculture due to rising temperatures and declining rainfall concentrations (Karimi et al., 2018). In Brazil, several regions such as the Semiarid region in the Northeast (Santos et al., 2017), already experience low-precipitation rates and are expected to suffer further declines according to projected climate shifts (Ferreira et al., 2018).

Among the species not yet cultivated on a commercial scale, but with considerable economic potential is Myrciaria floribunda (H. West ex Willd.) O. Berg (Myrtaceae), popularly known as rumberry or guavaberry in English, and as cambuí in Portuguese (García et al., 2021). This species occurs spontaneously across the phytogeographic domains of the Amazon, Caatinga, Cerrado, and Atlantic Forest biomes (Lemos et al., 2018).

Rumberry is agroextractively exploited in the Atlantic Forest (Alvares-Carvalho et al., 2025) and its commercial-scale cultivation could reduce the extractive pressure on natural populations (Oliveira et al., 2021). As this species is still in the process of domestication, all aspects related to its cultivation remain dependent on further researches (Santos et al., 2022). Studies on morphoanatomy, floral biology, and reproductive phenology show well-defined reproductive patterns that support the productive management of rumberry (Oliveira et al., 2021), while physicochemical analyses, bioactive compound profiling, and volatile compound assessments show fruits with high functional quality and broad variability among accessions, highlighting the potential for selecting superior genotypes (García et al., 2021). In addition, research involving Brazilian native species of the Myrtaceae family, such as Eugenia uniflora and Campomanesia adamantium, reports their pronounced physiological plasticity under drought stress (Junglos et al., 2016; Louzada et al., 2026).

Although specific studies are still needed, there is evidence that rumberry tolerates seasonal variations in water availability, as it naturally occurs in drier environments such as the Caatinga biome (Lemos et al., 2018). This ecological pattern suggests that the species may have developed adaptive strategies, considering that the successful plant development depends on complex physiological and biochemical adjustments during periods of low precipitation (Huang & Jin, 2025). Expected responses include osmolyte accumulation, enhanced antioxidant enzymatic activity, maintenance of the photosynthetic apparatus, and preservation of cell membrane integrity, which are mechanisms that support tolerance to water stress (El-Saadoni et al., 2024).

Therefore, the objective of this work was to characterize the physiological and biochemical responses associated with drought tolerance in rumberry plants.

Materials and Methods

The experiment was carried out from February to March 2021 in a greenhouse in the Campus of Engineering and Agricultural Sciences of the Universidade Federal de Alagoas (CECA/UFAL), in the municipality of Rio Largo (09°28'02"S; 35°49'43"W; at 127 m altitude), in the state of Alagoas, Brazil.

The seedlings used in the experiment were obtained from seed collected from ripe fruit in the Cambuí Germplasm Bank from CECA/UFAL. Seed were sown in trays filled with vermiculite and, shortly after emergence, the seedlings were transplanted into 230 cm³ tubes containing Bioplant Plus substrate (Bioplant Misturadora Agrícola, Nova Ponte, MG, Brazil), and maintained in a 50% shaded nursery for six months, at room temperature, with daily irrigation by micro-spraying for 1 hour. Subsequently, the plants were transferred to 2 L plastic bags filled with a 3:1 mixture of soil and cattle manure for an additional six-month period, maintained under full sun, and irrigated daily. Finally, the seedlings were transplanted into 20 L plastic buckets containing the same soil mixture, kept outside and irrigated once a day, without temperature or relative humidity control, and fertilized every 6 months with 50 g of NPK formula 04-14-08 per plant, until the plants reached 50 cm height at 2.5 years of age.

Meteorological data were recorded using an automatic weather station WS-GP1 model (DELTA-T Devices, Cambridge, England) installed inside the greenhouse. Parameters monitored included air temperature, relative humidity (RH), solar radiation, and vapor pressure deficit (VPD) (Figure 1). In addition, the soil moisture at 20 cm depth was measured every three days, using an SM-200 probe (DELTA-T Devices, Cambridge, England) (Figure 1).

Figure 1
Meteorological data in greenhouse and soil moisture: A, air temperature and relative humidity during the experimental period; B, mean diurnal variation of air temperature and relative humidity; C, solar radiation and vapor pressure deficit (VPD), during the experimental period; D, mean diurnal variation of solar radiation and VPD; E, soil moisture in pots used for the cultivation of rumberry (Myrciaria floribunda), under two water conditions - well-watered (WW) and water stress (WS) -, over 30 days of evaluation. Vertical bars represent the standard error of the mean.

The experimental design was completely randomized, with two treatments: well-watered (WW) soil, where soil moisture was maintained at field capacity; and water stress (WS) imposed by completely suspending irrigation for 30 days. For the experiment, 2.5-year-old plants were use, which have grown in 20 L containers. Each treatment included ten replicates. Leaf wilting in the WS treatment was considered the peak stress condition, which was observed 30 days after the experiment began. At this point, the experiment was terminated, and the number of leaves was recorded.

Leaves used for physiological and biochemical analyses were collected on day 30 from the middle third of each plant, ensuring they were fully expanded and in good phytosanitary conditions. Leaf sampling was carried out between 11:00 h and 13:00 h. Leaves were stored in thermal boxes with ice and immediately transported to the laboratory, in order to determine the relative water content (RWC), electrolyte leakage (EL), photosynthetic pigments, and osmolyte analyses. For analyses of malondialdehyde (MDA) content and antioxidant enzyme activity, the samples were stored in thermal containers filled with liquid nitrogen.

Measurements of RWC, EL, and photosynthetic pigments concentrations were performed on the same day as the samplings. Osmolytes, MDA, and antioxidant enzyme analyses were conducted approximately six months later, using samples previously stored at -80 °C in an ultra-freezer. The number of leaves for all plants were recorded when plants under WS conditions exhibited significant drought symptoms.

The RWC was determined by sequentially weighing the fresh mass (FM), turgid mass (TM), and dry mass (DM) of six leaf discs of about 8 mm diameter, according to the equation: RWC = [(FM - DM)/(TM - DM)] × 100 (Barros et al., 2023).

The photosynthetic pigment concentrations were determined following the methodology recommended by Barros et al. (2023). Chlorophyll a and chlorophyll b contents were expressed in (mg g^-1 FM), while carotenoid contents were expressed n (μmol g^-1 FM). Total chlorophyll (chlorophyll a + chlorophyll b) and the chlorophyll a/b ratio were subsequently calculated.

Throughout the 30-day period, the photochemical efficiency of photosystem II (PSII) was assessed every three days on fully expanded le aves, located in the middle third of the plant and in good phytosanitary conditions, using a PAM-2500 modulated chlorophyll fluorometer (Walz, Germany). Measurements of effective quantum yield of PSII (ΦPSII) were performed on three leaves per plant, between 9:00 h and 11:00 h, whereas maximum quantum efficiency of PSII (F_v/F_m) was measured around noon on only one leaf per plant, which was previously dark-adapted for approximately 30 min using leaf clips (Barros et al., 2023).

The extraction of total soluble amino acids, total soluble sugars, and free foliar proline was performed following the protocol described by Bieleski & Turner (1966). Protein extraction was carried out using the precipitated and preserved leaf material obtained during the previous extraction process. Protein quantification (μg g^-1 DM) was performed according to the Bradford method (Bradford, 1976). The contents of total soluble amino acids (μmol g^-1 DM), total soluble sugars (μg g^-1 DM), and proline (μmol g^-1 DM) were determined following the procedures described by Santos et al. (2019).

Lipid peroxidation was quantified by measuring the concentration of reactive substances to 2-thiobarbituric acid (TBA), expressed as malondialdehyde (MDA) equivalents (Barros et al., 2023), and results were expressed in (nmol g^-1 FM).

The EL was assessed in six leaf discs of 8 mm diameter, using a benchtop conductivity meter Tecnopon µCA-150. Measurements were based on the ratio between initial conductivity (IC) and final conductivity (FC), calculated by the equation: EL = (IC/FC) × 100 (Barros et al., 2023).

Protein content in the leaf extracts used for enzyme activity assays was quantified using the Bradford method (Bradford, 1976). The enzymatic activities of superoxide dismutase (SOD, units mg^-1 protein), ascorbate peroxidase (APX, units min^-1 mg^-1 protein), and catalase (CAT, units min^-1 mg^-1 protein) were determined according to the methodology described by Barros et al. (2023).

All statistical analyses were performed using Statistica software, version 13.0 (TIBCO Software Inc., Palo Alto, CA, USA). Shapiro-Wilk’s and Levene’s tests were employed to assess the assumptions of normality and homogeneity of variances, respectively. Data of ΦPSII and F_v/F_m were analyzed using repeated-measures analyses of variance with F-test (p<0.05). For the remaining variables, paired t-tests were applied (p<0.05), except for RWC, chlorophyll a/b ratio, free foliar proline, and CAT antioxidant enzyme activity, for which normality and/or homogeneity assumptions were not met; these variables were analyzed using the paired Wilcoxon’s test (p<0.05). All statistical analyses were performed using Statistica software, version 13.0 (TIBCO Software Inc., CA, USA).

Results and Discussion

Plants of rumberry subjected to water stress for 30 days maintained their initial number of leaves and exhibited 29.78% reduction of RWC, in comparison with well-watered plants (Figure 2). The maintenance of the number of leaves in rumberry, even under a marked reduction of RWC, indicates that the species did not activate the classical mechanisms of senescence and leaf abscission typically associated with WS (El-Saadony et al., 2024). In this context, the observed leaf retention suggests that the species sustains its photosynthetically active surface even under WS, which is physiologically relevant, as it preserves tissue functionality during the stress period and may potentially favor a more efficient resumption of growth after rehydration (Negin et al., 2023).

Figure 2
Number of leaves (A) and relative water content (B) of rumberry (Myrciaria floribunda) grown under two water conditions, well-watered (WW) and water stress (WS), on the 30^th day of stress. Vertical bars represent the standard error of the mean. Means for the number of leaves (A), followed by equal letters, do not differ significantly by paired t-tests, at 5% probability. Means for relative water content, followed by equal letters, do not differ by the paired Wilcoxon’s test, at 5% probability.

Chlorophyll a, chlorophyll b, total chlorophyll, the chlorophyll a/b ratio, and carotenoid concentrations were not affected by WS (Figure 3). In contrast, Junglos et al. (2016) reported a reduction of photosynthetic pigment contents in Campomanesia adamantium (Myrtaceae) after 12 days of irrigation suspension, which differs from the response observed in rumberry, in which photosynthetic pigment stability was maintained even after 30 days of irrigation suspension. In this context, the maintenance of pigment concentrations in rumberry suggests the existence of physiological mechanisms involved in the protection of the photosynthetic apparatus. In general, the maintenance of pigment stability under drought conditions has been associated with the attenuation of oxidative stress, as plants activate protective mechanisms capable of limiting the damage, mediated by the exacerbated production of reactive oxygen species (ROS), including increased proline accumulation and enhanced antioxidant enzyme activity (Rajput et al., 2021; Suárez-Salazar et al., 2024). Functionally, the preservation of chlorophylls and carotenoids ensures improved photochemical efficiency (Zafar et al., 2021).

Figure 3
Chlorophyll a (A), chlorophyll b (B), total chlorophyll (C), chlorophyll a/b ratio (D), and carotenoids (E) contents in leaves of rumberry (Myrciaria floribunda) grown under two water conditions - well-watered (WW) and water stress (WS) -, on the 30^th day of stress. Vertical bars represent the standard error of the mean. Means followed by equal letters do not differ significantly, by the paired Wilcoxon´s test, at 5% probability.

In the present study, from the 21^st day of irrigation suspension, ΦPSII decreased by 7.8%, reaching 27.8% by the 30^th day. Similarly, F_v/F_m values declined by 21.7%, starting from the 24^th day, reaching 29.4% reduction at the 30^th day, in comparison with the control treatment (Figure 4). Lin et al. (2017) also observed that WS negatively affected the photochemical efficiency parameters in potted guava (Psidium guajava L.) plants after five days of stress, suggesting that rumberry may exhibit a greater tolerance than this species. This greater drought-tolerance capacity may be associated with enhanced efficiency of PSII photoprotective mechanisms, as evidenced by the maintenance of chlorophyll b, suggesting improved regulation of energy flow, and of carotenoids, which act directly in excess energy dissipation and in the attenuation of oxidative stress (Simkin et al., 2022).

Figure 4
Effective quantum yield of PSII (ΦPSII) (A) and maximum quantum efficiency of PSII (F_v/F_m) (B) in leaves of rumberry (Myrciaria floribunda) grown under two water conditions - well-watered (WW) and water stress (WS) -, over 30 days of evaluation. Vertical bars represent the standard error of the mean. Arrows indicate the beginning of statistically significant differences between treatments at 5% probability.

The WS condition can lead to photoinhibition, thereby reducing photochemical efficiency (Cintra et al., 2020). In the context of this investigation, however, photoinhibition should not be interpreted as damage, but rather as a protective mechanism that enables the dissipation of excess thermal energy (Takahashi & Badger, 2011), since no degradation of photosynthetic pigment was observed in rumberry plants under the WS regime.

The reduction of 15.6% of total soluble protein content was observed under WS, in comparison with the WW treatment, while total soluble amino acids and free foliar proline increased by 33.9% and 1,226.6%, respectively (Figure 5). The total protein and amino acid concentrations observed suggest that this may represent a tolerance mechanism in rumberry, as protein degradation catalyzed by protease enzymes and the inhibition or arrest of protein synthesis result in increased concentrations of soluble amino acids and decreased protein concentrations (Krasensky & Jonak, 2012). Changes in osmolyte concentrations are frequently observed in plants with greater drought tolerance (Zafar et al., 2021).

Figure 5
Total soluble proteins (A), total soluble amino acids (B), total soluble sugars (C), and free foliar proline (D) in leaves of rumberry (Myrciaria floribunda) grown under two water conditions - well-watered (WW) and water stress (WS) -, on the 30^th day of the experiment. Vertical bars represent the standard error of the mean. Means followed by equal letters do not differ significantly, by the paired t-tests, at 5% probability. For free proline (D), means followed by equal letters do not differ, by the paired Wilcoxon’s test, at 5% probability.

No significant differences were detected for total soluble sugar content between WS and WW plants (Figure 5), suggesting that soluble sugars were not the main osmoprotective mechanism mobilized by the species in response to WS conditions. Similar findings were reported by Bascuñán-Godoy et al. (2015), who found no significant changes of soluble sugar concentrations in Myrceugenia exsucca and Luma chequen under WS conditions.

Additionally, rumberry plants under WS showed about 1,227% increase of proline content, in comparison with the WW treatment (Figure 5), indicating a strong metabolic adjustment to dehydration. Proline accumulation promotes water retention in the cytoplasm, contributing to the maintenance of cell turgor and membrane stabilization, thereby reducing EL, in addition to acting in ROS scavenging and antioxidant protection of cells (Rajput et al., 2021; Suárez-Salazar et al., 2024). Furthermore the proline accumulation contributes to the survival of rumberry under WS conditions. This response, together with the accumulation of total soluble amino acids and the maintenance of total soluble sugars, reinforces the role of osmolytes in osmotic adjustment and in minimizing cellular dehydration (Huang & Jin, 2025).

Alterations were observed in the activity of the enzymatic antioxidant system: SOD activity decreased by 45.51%, while CAT and APX enzyme activities increased by 141.66% and 88.99%, respectively (Figure 6). Vieira et al. (2022), when evaluating seed of Campomanesia xanthocarpa (Myrtaceae) subjected to desiccation, observed a reduction of the SOD activity and a marked increase of the activities of the antioxidant enzymes CAT and APX after 24 hours, a pattern similar to that observed in the present study with rumberry plants subjected to WS for 30 days. This response profile indicates a reconfiguration of the antioxidant activity favoring enzymes involved in hydrogen peroxide (H₂O₂) scavenging - the main byproduct of photoinhibition - reinforcing the hypothesis that rumberry plants mobilize efficient biochemical strategies to limit the oxidative damage under WS conditions (Rajput et al., 2021).

Figure 6
Antioxidant enzyme activity and cell membrane stability. Superoxide dismutase (SOD) activity (A), catalase (CAT) activity (B), ascorbate peroxidase (APX) activity (C), MDA content (D), and electrolyte leakage (E), in leaves of rumberry (Myrciaria floribunda) grown under two water conditions - well-watered (WW) and water stress (WS) -, on the 30^th day of the experiment. Vertical bars represent the standard error of the mean. Means followed by equal letters do not differ significantly, by the paired t-tests, at 5% probability. For the CAT activity (B), means followed by equal letters do not differ, by the paired Wilcoxon’s test, at 5% probability.

Neither MDA content nor EL were significantly affected by WS conditions (Figure 6), indicating that the oxidative damage to cellular membranes was not pronounced. In contrast, Zafar et al. (2021) reported an increased MDA content and EL in Syzygium cumini (Myrtaceae) subjected to water stress, in a pot experiment conducted under greenhouse conditions, evidencing membrane destabilization under similar conditions. In the present study, the absence of MDA accumulation and the lack of increase in EL can be explained by the increased activity of the antioxidant enzymes CAT and APX, together with proline accumulation, which could contribute to the stabilization of cellular membranes and to the reduction of oxidative stress in rumberry plants.

In the present study, rumberry was found to exhibit different mechanisms of drought tolerance. This behavior indicates a high potential for the cultivation and domestication of the species, especially for fruit production systems established in areas subjected to water stress. In addition, the ability of the plant to maintain its physiological functioning under water-deficit conditions also favors sustainable extractivism (Lemos et al., 2018), since more tolerant natural populations tend to show greater persistence, productivity, and stability even in drier years.

Conclusions

• 1. Rumberry (Myrciaria floribunda) shows tolerance to water stress through coordinated physiological and biochemical adjustments.

• 2. The maintenance of the number of leaves and photosynthetic pigment concentrations, associated with a moderate reduction in photosystem II photochemical efficiency, indicates physiological adjustment under water stress.

• 3. Cellular membrane stability, together with increased proline and total soluble amino acid contents and enhanced catalase and ascorbate peroxidase activities, indicates the biochemical response of the species to water stress.

Data availability statement

Data available upon request: research data are only available upon reasonable request to the corresponding author.

Acknowledgments

To Coordenação de Aperfeiçoamento de Pessoal de Nível (CAPES), for financing, in part, this study (Finance Code 001); and to Universidade Federal de Alagoas (UFAL), specifically to the Graduate Program in Agronomy (Plant Production), to Embrapa Alimentos e Territórios, and to Universidade Federal de Sergipe (UFS), for their institutional support and collaboration.

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Declaration of use of AI technologies

During the preparation of this work, the author(s) used ChatGPT (OpenAI) in order to improve the fluency, readability, and overall language of the manuscript. After this use, the author(s) reviewed and edited the content as needed and take(s) full responsibility for it.

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• Chief editor:
  Edemar Corazza

• Edited by:
  Mírian Baptista

Publication Dates

• Publication in this collection
  14 Sept 2026
• Date of issue
  2026

History

• Received
  07 June 2025
• Accepted
  29 Jan 2026