Characterization of strawberry cultivars for resistance to arthropod pests mediated by leaf trichomesABSTRACT
Strawberry (Fragaria × ananassa) is a nutritionally rich pseudo-fruit valued for its distinctive flavor, aroma, and high antioxidant and mineral content. It is consumed both fresh and as an ingredient in various food products. However, strawberry crops are highly susceptible to arthropod pests, particularly the two-spotted spider mite (Tetranychus urticae) and the strawberry aphid (Chaetosiphon fragaefolii), often leading growers to rely heavily on chemical control, with associated environmental and health risks. Genetic resistance offers a sustainable alternative. Identifying potential parental lines with elevated resistance can support the development of new resistant cultivars. In this study, 41 strawberry genotypes from the State University of Londrina collection were evaluated for resistance in choice and no-choice bioassays against T. urticae and C. fragaefolii, and leaf trichomes (glandular and non-glandular) were quantified. Correlations between trichome densities and past performance were analyzed. Significant variability in pest resistance was observed among genotypes, primarily associated with glandular trichomes. The cultivars Camino Real, Ceres, and the experimental genotype RVDA44 exhibited the highest resistance, limiting pest oviposition, survival, and movement, and represent promising candidates for commercial cultivation and as parental lines in breeding programs aimed at developing new resistant cultivars.
Index terms:
Fragaria × ananassa; genetic resistance; glandular trichomes
RESUMO
O morango (Fragaria × ananassa) é um pseudofruto nutricionalmente rico, valorizado por seu sabor característico, aroma e alto teor de antioxidantes e minerais. É consumido tanto in natura quanto como ingrediente em diversos produtos alimentícios. No entanto, a cultura do morangueiro é altamente suscetível a pragas artrópodes, especialmente o ácaro-rajado (Tetranychus urticae) e o pulgão-do-morangueiro (Chaetosiphon fragaefolii), levando frequentemente os produtores a dependerem do controle químico, com riscos ambientais e à saúde. A resistência genética surge como alternativa sustentável. A identificação de genitores com maior resistência pode subsidiar o desenvolvimento de novas cultivares resistentes. Neste estudo, 41 genótipos de morangueiro da coleção da Universidade Estadual de Londrina foram avaliados quanto à resistência, por meio de bioensaios com e sem chance de escolha, frente a T. urticae e C. fragaefolii, e os tricomas foliares (glandulares e não glandulares) foram quantificados. Foram analisadas correlações entre a densidade de tricomas e o desempenho dos genótipos. Observou-se variabilidade significativa na resistência às pragas, principalmente associada aos tricomas glandulares. As cultivares Camino Real, Ceres e o genótipo experimental RVDA44 apresentaram maior resistência, limitando a oviposição, sobrevivência e movimentação das pragas, sendo promissores para cultivo comercial e como genitores em programas de melhoramento visando o desenvolvimento de novas cultivares resistentes.
Termos para indexação:
Fragaria × ananassa; resistência genética; tricoma granular
Introduction
Fragaria × ananassa Duch., the predominant strawberry species in global cultivation, is an interspecific hybrid derived from a cross between Fragaria chiloensis (L.) Mill. and Fragaria virginiana Mill. This species is highly valued for its organoleptic and nutraceutical properties and is consumed both fresh and in processed forms. Rich in minerals, vitamins, and antioxidant compounds, strawberries are considered functional food with the potential to promote health benefits (Giampieri et al., 2012; Newerli-Guz et al., 2023).
In Brazil, strawberry cultivation is predominantly carried out by smallholder farmers, representing an important source of income for thousands of families (Oliva et al., 2025). However, the production is associated with high costs due to dependence on imported seedlings, labor-intensive practices, and the heavy use of agricultural inputs (Resende et al., 2020a; Rutz et al., 2023). This situation is further exacerbated by the occurrence of pests, such as the two-spotted spider mite (Tetranychus urticae Koch) and the strawberry aphid (Chaetosiphon fragaefolii Cockerell). These pests thrive under protected cultivation environments with low relative humidity and high temperatures, conditions that can significantly reduce crop productivity (Figueiredo et al., 2013; Musaqaf, N. et al., 2022, Musaqaf, S. et al., 2022).
Resistance to arthropod pests in strawberries is often associated with morpho- anatomical characteristics, such as the presence of glandular and non-glandular trichomes, which can hinder the movement and feeding of insects and mites (Benatto et al., 2019; Resende et al., 2020b). Previous studies have reported different levels of resistance among commercial cultivars, such as ‘Camino Real’, as well as among wild materials, including Fragaria vesca, highlighting the existence of genetic variability that can be exploited in breeding programs (Milenković et al., 2014; Resende et al., 2020a).
Strawberry breeding programs aim not only to improve productivity and fruit quality but also to incorporate resistance to pests and diseases (Zeist & Resende, 2019; Nascimento et al., 2023). However, the narrow genetic base of Fragaria × ananassa limits rapid breeding progress, reinforcing the importance of exploring available germplasm, including wild species of the genus Fragaria, as sources of resistance genes (Shanks & Moore, 1995; Bac-Molenaar et al., 2019).
Figueiredo et al. (2013) reported resistance of the cultivar ‘Camino Real’ to the two-spotted spider mite and associated this resistance with the presence of glandular trichomes on the leaflet surface. Similarly, Resende et al. (2020b), while evaluating intraspecific hybrids, observed variability in resistance to the two-spotted spider mite and confirmed the resistance of the Camino Real cultivar. Musaqaf et al. (2022) evaluated infestations of C. fragaefolii and Aphis gossypii in mixed populations of wild strawberry plants (Fragaria vesca) and observed differential levels of attack, demonstrating variability among plants in resistance to aphid species. Benatto et al. (2019) further revealed an intrinsic relationship between the number of glandular and non-glandular trichomes in the cultivar Albion and the feeding behavior of C. fragaefolii, suggesting that trichomes act as a physical barrier that interferes with aphid feeding.
These studies confirm the presence of variability within the genus Fragaria in resistance to major pests and link this resistance to morphological and anatomical traits as well as chemical compounds. Identifying such traits in F. × ananassa germplasm is therefore essential for breeding programs aiming to develop cultivars with enhanced pest resistance.
The objective of this study was to evaluate 41 strawberry genotypes for resistance to the two-spotted spider mite and the strawberry aphid, in support of pre-breeding efforts. The identification of cultivars exhibiting resistance to these pests may contribute to the development of commercial strawberry cultivars with improved pest resistance.
Material and Methods
Experimental site
The experiments were conducted in a greenhouse and in the Entomology and Electron Microscopy Laboratories at the State University of Londrina (UEL), Londrina, PR, Brazil (23°19’44”S; 51°12’17”W), at an average altitude of 582 m above sea level (a.s.l.). According to the Köppen climate classification, the regional climate is humid subtropical, characterized by year-round rainfall and hot summers. Mean summer temperatures surpass 22 ºC, while precipitation in the driest month exceeds 30 mm.
Forty-one strawberry genotypes from the UEL germplasm collection were evaluated, including commercial cultivars, commercial cultivars no longer cultivated, pre-commercial cultivars, experimental genotypes, and one F. vesca accession (Table 1). Plant propagation was carried out using runners, which were rooted in cell trays and subsequently transplanted into 3.5 dm³ polypropylene pots containing a 1:1 (v/v) mixture of soil and commercial substrate. Plants were fertilized and maintained under regular irrigation in a protected environment until they were established.
On average, ten mother plants of each cultivar/genotype were maintained in the collection. Clonal plants used in the resistance bioassays were obtained from these mother plants. During the spring/summer of 2023, under conditions of high temperature and long photoperiod, stolon production was stimulated. Stolons were collected and transplanted into 56-cell trays filled with Carolina Soil® substrate and maintained in a greenhouse under controlled conditions (27 ± 4 °C; 70 ± 10% RH) for rooting and seedling development. Genotypes producing fewer stolons were transferred to a phytotron to stimulate vegetative propagation.
Foliar fertilizations were applied every 15 days using Nutran® fertilizer at a dose of 1 L ha-1. For fungal disease prevention, iprodione and mancozeb were applied alternately at 10-day intervals. Pest control was carried out using acephate and abamectin on an alternating 15-day schedule until the final seedling formation stage. For crown borer control, thiamethoxam was applied at a concentration of 1 mL L-1. All insecticide and acaricide applications were suspended at least 30 days prior to the establishment of the aphid and mite bioassays to avoid residual effects on pest performance and to ensure that the evaluations reflected only plant resistance traits.
Approximately 60 days after rooting, seedlings were transplanted into 3.5 dm³ polypropylene pots filled with a 1:1 (v/v) mixture of subsoil and Carolina Soil® substrate. For pH correction, 25 g of dolomitic lime was applied per pot, and the substrate was supplemented with the equivalent of 1500 kg ha-1 of 4-14-8 fertilizer. The pots were maintained in the greenhouse under controlled temperature and humidity for plant establishment. Irrigation was performed manually, and fungal disease control was conducted using copper-based products. Additional applications of Nutran® (1.5 L ha-1) were made at 15-day intervals. Forty-five days after transplanting, plants were selected for resistance bioassays against mites and aphids. At this stage, young fully expanded leaflets were also collected for trichome density analysis.
Bioassays with the two-spotted spider mites (Tetranychus urticae)
Resistance of strawberry genotypes to the two-spotted spider mite was evaluated using non- preference bioassays adapted from Weston and Snyder (1990). Mites were reared on common bean plants (Phaseolus vulgaris), cultivar Águia, maintained in insect-proof cages (1.0 m × 1.0 m × 1.0 m) under greenhouse conditions (27 °C ± 4 °C; 70% ± 10% RH; 12-hour photoperiod).
To standardize mite age, once the colony was established, adult female mites were transferred to bean plants that were completely free from any arthropod pests. After 24 hours, the mites on the leaflets were killed, and eggs were allowed to hatch, resulting in a synchronized mite population.
Two bioassays were conducted to evaluate the resistance of the genotypes to the two-spotted spider mite: a no-choice test and a mite walking test, both under controlled conditions (T= 25 ± 2 ºC).
No-choice test
The experiment followed a randomized block design with nine replications. Treatments consisted of 41 strawberry genotypes (Table 1). Young, fully expanded leaflets were collected from strawberry plants maintained in a greenhouse, with an approximate leaf area of 9.61 cm².
The experiment was conducted as follows: a thin layer of sponge was placed at the base of a 5-mm diameter Petri dish and moistened with distilled water (Figure 1). The sponge maintained leaflet hydration and prevented the mites from escaping. A sheet of filter paper was placed over the sponge layer, on which leaf discs (9.61 cm²) from each genotype were positioned. The leaf discs were randomly arranged with the abaxial surface facing upward. Each Petri dish contained nine strawberry leaflets, with each leaflet representing one experimental unit. One five-day-old adult female two-spotted spider mite from the reared colony was released onto each leaflet.
After 24 hours, the females were removed, and the number of eggs laid was recorded. At 96 and 120 hours after oviposition, the number of hatched mites was counted, and the egg-to-hatch ratio was calculated.
Experimental setup of the no-choice test with two-spotted spider mites on leaf discs of strawberry genotypes.
Mite walking assay
A method adapted from Weston and Snyder (1990) was used in a completely randomized design with five replications. Young expanded leaflets (50 cm²) from each genotype were fixed with the abaxial surface facing upward on White paper mounted on a polystyrene plate. A 9-mm-diameter brass washer was placed at the center of each leaflet (Figure 2). Ten adult female mites were transferred simultaneously onto each leaflet using a fine brush.
Experimental setup of the mite walking assay adapted from Weston and Snyder (1990) to evaluate the resistance of strawberry genotypes to the two-spotted spider mite.
Distances traveled by the mites were measured at 10, 20, 30, and 40 minutes after release using a millimeter ruler. If the mites moved off the leaflet, the distance was measured from the washer center to the leaflet edge. If mites remained on the washer, the distance was recorded as zero. The weighted mean distance traveled by the ten mites per leaflet was calculated for statistical analysis.
Bioassays with the strawberry aphid (Chaetosiphon fragaefolii)
Strawberry plants of the 41 genotypes at the pre-flowering stage were subjected to resistance bioassays against aphid Chaetosiphon fragaefolii. The experiment employed a completely randomized block design with six replications, resulting in a total of 252 pots, with one plant per pot. Pest control was suspended 30 days before bioassay initiation. Plants were transferred to a greenhouse equipped with insect-proof mesh and an antechamber to prevent external infestations and were irrigated daily.
Adult aphids were collected from strawberry plantations in Jaboti, PR, Brazil (23º44’34” S, 50º04’33” W; 560 m a.s.l.). The adults were released onto strawberry plants of the cultivar Cristal to establish and maintain laboratory colonies. The plants were grown in plastic pots and maintained in rearing cages (2 m × 2 m × 2 m) covered with aphid-proof mesh and kept in a greenhouse under controlled conditions (22 °C ± 3 °C, 70% ± 5% RH, and a 12-h photoperiod).
For infestation, ten third- and fourth-instar nymphs from the stock population were transferred to the terminal trifoliate of each plant, preferably on the abaxial surface (Figure 3). Nymphs were carefully removed from leaflets under a binocular stereomicroscope (ECZ-Black, Biofocus) and transferred to Petri dishes lined with filter paper lightly moistened with distilled water to minimize stress. Nymph removal was conducted in a climate-controlled room (20±3 °C; 80% ± 5% RH). After the required number of nymphs was obtained for each block, they were transferred to the plants using fine-bristle brushes moistened with Aloe vera extract diluted in distilled water.
Evaluations were conducted 15 days after infestation. Nymph abundance and population density were assessed at ten-day intervals over a 60-day period, resulting in six evaluations per genotype. One leaflet from the upper third of each plant was collected at each evaluation. The sampled leaflets were transported in thermal boxes to the Entomology Laboratory. Nymphs were counted under a stereomicroscope within a 2 cm² area on the abaxial surface of each leaflet. Mean values were calculated using weighted data from all evaluations.
Population density was evaluated following a methodology adapted from Heie (1993), in which aphids were counted on the first fully expanded leaf across its three leaflets. Winged, wingless, and nymphal aphid counts were recorded. A total of five evaluations were conducted, excluding the first due to low infestation levels. The final mean was calculated using the values from these five evaluations.
Leaf trichome density analyses
Trichomes were classified following Toscano et al. (2001) based on the presence or absence of apical glands: those with glands were designated as glandular (non-tector), and those without glands as non-glandular (tector). Fully expanded young leaves from each genotype were sampled to quantify trichome density. Circular leaf discs (10 mm diameter) were obtained using a leaf punch and divided into four sections, which were fixed with carbon tape in the sample holder compartment for scanning electron microscopy (SEM) analysis at 100 × magnification using a Hitachi High-Tech TM3000 microscope (tungsten filament, low vacuum, 15 kV). Trichome density was quantified on both adaxial and abaxial surfaces of the leaflets and expressed as the number of trichomes per square millimeter (mm²) area. Final mean values were calculated across all sections and surfaces of the leaf discs.
Statistical analyses
The data were tested for normality and homogeneity of residuals using the Shapiro-Wilk and Hartley tests, respectively. When the assumptions were met, the data were subjected to analysis of variance (ANOVA). Means were compared using the Scott-Knott clustering test at a 5% significance level (p < 0.05). Oviposition data that did not meet these assumptions of normality and homogeneity were transformed using √ (x + 1).
All analyses were performed using the SISVAR statistical software (Ferreira, 2019). Pearson correlation coefficients (r) between resistance parameters and trichome density were calculated and tested using the t-test (p ≤ 0.05).
Results and Discussion
Leaf trichome density in strawberry
Analysis of glandular trichome density (Table 2) grouped the 41 evaluated genotypes into eight distinct categories. The cultivar Camino Real exhibited the highest density, followed by Ventana and Ceres, confirming prior findings that linked this genotype to elevated glandular trichomes and enhanced resistance to pest arthropods (Resende et al., 2020b). Conversely, the cultivars Vila Nova, Toyonoka, Seascape, Pircinque, Oso Grande, Oso Bola, Monterey, Florida Festival, Florida Sensation, Campinas, and the wild species Fragaria vesca showed the lowest densities, traits generally associated with higher susceptibility (Lourenção et al., 2000).
Leaf trichome density of 41 strawberry genotypes, including glandular trichomes (GT), non-glandular trichomes (NGT), and total trichomes (TT = GT + NGT). The average distance traveled (D) by Tetranychus urticae females was recorded at 10, 20, 30, and 40 minutes after release onto leaflets, and the total average distance traveled (TADT) was calculated. The number of eggs laid (NE) was assessed 24 hours after release; the number of eggs hatched (N) was evaluated at 96 and 120 hours after oviposition; and the number of live mites (AM) was recorded at 24 and 48 hours after oviposition.
For non-glandular trichomes (Table 2), the genotypes were clustered into four groups. The highest densities were observed in Camino Real, Ceres, Cristal, Dover, and genotype RVDA44, while Campinas, Florida Sensation, Toyonoka, Tudla Milsey, Palomar, F. vesca, and genotypes RVMJ001 and RVMJB002 had the lowest. Non-glandular trichomes act as physical barriers, hindering pest movement, oviposition, and feeding (Benatto et al., 2019).
Beyond morphology, chemical compounds in leaf tissues also contribute to resistance. Alkaloids from Zanthoxylum schreberi reduce the survival and fecundity of spider mites, interfere with their detoxification enzymes, and demonstrate that chemical defenses can act synergistically with trichomes (Rincón, Rodríguez, & Coy-Barrera, 2024). Total phenolic compounds similarly correlate negatively with spider mite performance (Hata et al., 2019; Sousa et al., 2021), suggesting that resistant genotypes may also possess foliar chemical variations, reinforcing their natural defense.
Considering total trichome density (glandular + non-glandular), Camino Real exhibited the highest density, followed by Ceres, Ventana, and RVDA44, whereas Campinas, Toyonoka, F. vesca, and RVMJ001 showed the lowest values. These results support the role of trichome density in resistance against mites and insects, although other factors, such as exudate composition, also play a key role (Simmons & Gurr, 2005; Bleeker et al., 2011).
The functional significance of trichome data is supported by studies in strawberry and other species. Benatto et al. (2018) demonstrated that higher trichome densities increase superficial exploration time and reduce feeding by the aphid Chaetosiphon fragaefolii. Our results further confirm the role of trichomes as constitutive defense mechanisms and highlight their potential for use in breeding programs aimed at enhancing pest resistance.
Choice and no-choice tests with the two-spotted spider mite
Choice and no-choice tests (Table 2) revealed significant variation in the behavior and development of Tetranychus urticae among the evaluated strawberry genotypes. The cultivars Camino Real and Ceres, followed by Albion, Burkley, Cristal, Ventana, Fragaria vesca, and genotype RVDA44, exhibited lower oviposition, reduced hatching rates, and lower mite survival, indicating strong resistance. In contrast, cultivars Toyonoka, Monterey, Vila Nova, Seascape, Jônica, and genotype RVMJ001 were the most susceptible, showing higher levels of oviposition, hatching, and survival.
The reduced attractiveness and oviposition observed in Camino Real and Ceres are consistent with previous findings, which demonstrate that high densities of glandular and non-glandular trichomes limit spider mite movement and oviposition (Figueiredo et al., 2013). Moreover, increased trichome density has been shown to negatively affect egg hatching and survival (Resende et al., 2020b), highlighting these traits as key physical and chemical resistance barriers and supporting the occurrence of antixenosis and antibiosis mechanisms.
Additional studies reinforce the role of trichomes as a constitutive defense in strawberry. Benatto et al. (2018) demonstrated that cultivars with higher trichome density significantly altered the feeding behavior of the aphid Chaetosiphon fragaefolii, increasing non-probing time and reducing phloem feeding. Similarly, Musaqaf, S. et al. (2022b) reported lower oviposition and survival of T. urticae on strawberry cultivars with higher trichome density. In line with these findings, Esteca et al. (2017) observed that new strawberry genotypes with higher trichome levels exhibited moderate resistance to spider mites, even when compared with widely used cultivars such as IAC T-0104, IAC 12, and IAC Princesa Isabel. These results are consistent with the present study, in which genotypic variation was reflected in differing degrees of resistance.
Therefore, the results of this study confirm that genotypes such as Camino Real, Ceres, and RVDA44 possess traits associated with spider mite resistance, primarily due to higher densities of glandular and non-glandular trichomes, reinforcing the importance of incorporating these traits into breeding programs (Figure 4). In contrast, more susceptible genotypes, such as Toyonoka and Monterey, can serve as contrasting controls in future evaluations, facilitating the selection of more resilient and adaptable materials.
Density of non-glandular trichomes on the abaxial surface of leaflets of strawberry cultivars/genotypes: (A) Ceres, (B) Camino Real, (C) RVDA44, (D) Toyonoka, (E) Monterey, and (F) Alpina.
Evaluations with the strawberry aphid
Resistance to the strawberry aphid (Chaetosiphon fragaefolii) varied significantly among the genotypes evaluated (Table 3). Overall, the cultivars Camino Real and Ceres, along with Fragaria vesca and the experimental genotypes RVDA16 and RVDA44, exhibited the lowest infestation levels and were classified as resistant. Conversely, the cultivars Monterey, Toyonoka, Portola, Seascape, and the genotype RVMJB002 were highly susceptible, supporting the formation of large nymph populations. These results highlight the importance of genetic variability in aphid (C. fragaefolii) resistance (Bernardi et al., 2012; Benatto et al., 2018).
Number of live adult strawberry aphids and nymphs on leaflets of 41 strawberry genotypes, assessed every ten days after the release of females. Live adults (LA) were recorded at six evaluations (LA1-LA5), with the total average number of live adults represented as TLA. Live nymphs were recorded at the same six evaluations (N1-N6), with the total average number of live nymphs represented as TLN.
The observed resistance mechanisms can be partially explained by leaf morphology. Both glandular and non-glandular trichomes have been associated with aphid resistance, as they reduce mobility, increase non-probing time, and limit phloem feeding (Benatto et al., 2018).
Beyond morphology, physiological and biochemical factors also play a crucial role. Low nymph survival in certain strawberry genotypes may be associated with plant nutritional quality and antibiosis effects that negatively affect pest development (Benatto et al., 2019; Painter, 1951). Additionally, secondary metabolites such as phenols, flavonoids, and glycosides have been shown to contribute to defense against aphids (Resende et al., 2020c). Trindade et al. (2022) further demonstrated that exogenous application of salicylic acid in strawberry can increase trichome density and reduce T. urticae populations, suggesting that induced mechanisms can complement constitutive resistance.
Overall, the results of the present study confirm that resistance to the green aphid in strawberry is multifactorial, involving both physical barriers (trichomes) and potentially nutritional and biochemical defenses (phenols and flavonoids). The identification of resistant genotypes, such as Camino Real and Ceres, is particularly relevant for breeding programs, as these genotypes can reduce dependence on insecticides and support the adoption of sustainable integrated pest management strategies.
Correlation analysis
Correlation analysis indicated negative relationships between plant defense traits (trichomes) and mite biological variables (Figure 5). Specifically, correlations involving glandular trichomes were stronger (−0.72 to −0.79) than those involving non-glandular trichomes (−0.56 to −0.67). A similar pattern was observed for aphid biological variables (Figure 6). Here, glandular trichomes showed stronger correlations with nymph and adult numbers (−0.69 and −0.70, respectively) compared with non-glandular trichomes (−0.58 and −0.61).
Pearson correlation matrix of traits evaluated in strawberry genotypes for resistance to the two-spotted spider mite (Tetranychus urticae). Colors indicate the strength and direction of correlations: red, positive; orange, intermediate; and blue, negative. Trait abbreviations: GT, glandular trichomes; NGT, non-glandular trichomes; TT, total trichomes; D, total average distance traveled by mites; NE24H, number of eggs oviposited 24 h after female release; N96H and N120H, number of eggs hatched after 96 and 120 h of oviposition; AM24H and AM48H, number of live mites 24 and 48 h after oviposition.
Pearson correlation matrix of traits evaluated in strawberry genotypes for resistance to the aphid Chaetosiphon fragaefolii. Colors indicate the strength and direction of correlations: red, positive; Orange, intermediate; and blue, negative. Trait abbreviations: GT, glandular trichomes; NGT, non-glandular trichomes; TT: total trichomes.
These results emphasize the key role of glandular trichomes as a primary morphological defense, with non-glandular trichomes providing supplementary protection.
Conclusions
The results indicate that Camino Real and Ceres are the most resistant cultivars to Tetranychus urticae and Chaetosiphon fragaefolii, showing reduced oviposition, hatching, survival, and pest development. Genotype RVDA44 also demonstrated notable resistance, especially by limiting mite movement. In contrast, Monterey, Toyonoka, RVMJ001, and RVMJB002 were highly susceptible. The association between trichomes and reduced pest colonization highlights their role in plant defense, supporting the use of resistant genotypes to improve integrated pest management and sustainability.
Data Availability Statement
Data available upon request to authors.
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Editor de seção:
Renato Paiva0000-0001-5107-0285
