Abstract
Many species of woody plants from the Cerrado (tropical savanna in central Brazil) can produce fleshy fruits. Still, their chemical composition is primarily unknown, due to the lack of commercial interest or regular access to said species. The present study provides the centesimal composition of seven native fruit species from the Cerrado: Lobeira (Solanum lycocarpum Solanaceae), Angelim-rasteiro (Andira humilis Fabaceae), Abacaxi-do-Cerrado (Ananas ananassoides Bromeliaceae), Araçá-do-campo (Psidium grandifolium Myrtaceae), Caraguatá (Bromelia balansae Bromeliaceae), Pequi (Caryocar brasiliense Caryocaraceae) and Araticum (Annona crassiflora Annonaceae). Fruit samples were collected from protected areas of the Cerrado in southeastern Brazil. The centesimal composition was analyzed without distinguishing between flesh, seeds, and peel. All fruits presented a moisture content ranging from 55.2% to 83.0%, and low ash contents (less than 4.5%). The Lobeira, the Angelim-rasteiro, and the Abacaxi-do-Cerrado stood out for having a higher percentage of protein (15.9%, 16.2%, and 10%, respectively). In contrast, the Araçá-do-campo, the Caraguatá, and the Araticum had high levels of carbohydrates (23.8%, 21.6% and 35.9%, respectively). Results indicate these fruits’ potential for the food industry, as well as for the development of new food products.
Keywords:
Carbohydrates; Fruit traits; Fruit chemical composition; Lipids; Proteins; Cerrado biodiversity
HIGHLIGHTS
Cerrado fruits show high protein and carbohydrate contents
Cerrado native species offer potential for innovative food products
First description of the composition of the fruits from the Andira humilis and the Psidium grandifolium
1 Introduction
The Cerrado is the second largest Brazilian ecosystem, covering around 2 million km2 (Oliveira-Filho & Ratter, 2002), being described as a tropical savanna with a great plant diversity (estimated in 10,000 species) (Myers et al., 2000; Oliveira-Filho & Ratter, 2002). Around 40% of these plant species are endemic, making the Cerrado a biodiversity hotspot with a high demand for animal and plant species conservation (Myers et al., 2000).
About half of the woody plants from the Cerrado produce fleshy fruits. However, not many fruit species have their chemical composition described in the literature, which constrains the assessment of their potential use. The Araticum (Annona crassiflora Mart.), the Baru (Dipteryx alata Vogel) and the Pequi (Caryocar brasiliense Cambess.) are among the most popular fruits of the Cerrado, being widely used for human consumption in Brazil, with a market value of US$0.31 kg-1, US$5.53 kg-1 and US$2.29 kg-1, respectively (prices from November 16, 2023) (Monteiro et al., 2022; Pereira et al., 2022; Silva et al., 2020). Considering the large number of fruit species available in the Cerrado, it is likely that several other species with potential economic value are being overlooked.
The basis for identifying edible fruits comes from observations of animal feeding habits. Carnivorous mammals often include fruits in their diets (Draper et al., 2022). An ongoing study in the Cerrado identified at least 22 fruits consumed by wild canids such as the Maned-wolf (Chysocyon brachyurus Illiger) and the Crab-eating fox (Cerdocyon thous Linnaeus) (Feliciano, 2024). Given the importance of these fruits for Cerrado’s fauna along with their potential for human use, the present study aimed to characterize seven species in terms of their macrocomponents, comparing the findings with the literature’s data, when possible. Of the seven fruits assessed in this study — Lobeira (Solanum lycocarpum A. St.-Hil., Solanaceae), Abacaxi-do-Cerrado (Ananas ananassoides Baker L.B.Sm, Bromeliaceae), Caraguatá (Bromelia balansae Mez, Bromeliaceae), Pequi (Caryocar brasiliense, Caryocaraceae), Araticum (Annona crassiflora, Annonaceae), Angelim-rasteiro (Andira humilis Mart. ex Benth., Fabaceae), and Araçá-do-campo (Psidium grandifolium Mart. ex DC, Myrtaceae) — most have limited or no available data regarding their macronutrient composition. Specifically, there is a notable absence of information concerning the macronutrients of the latter two species. Taking into account the lack of compiled information in the literature, the description of the abovementioned botanical species was detailed herein to support future authors.
2 Materials and methods
2.1 Study site
The present study utilizes the botanical term “fruit” to encompass all types of diaspores, regardless of their origin or structure (e.g., true fruits, pseudo-fruits, arils, and seeds). Fruits from seven different species, which comprise part of the diet of wild canids, were collected between January and May of 2023 (summer/autumn) from native plants in the Cerrado at the Itirapina Ecological Station (22º15’ S, 48º00’ O, elevation 770 m), located in the municipality of Itirapina, State of São Paulo, Southeastern Brazil. The region experiences an average annual rainfall of 1320 mm and a mean temperature of 22 °C, with a climate classified as Cwa (subtropical with a dry winter), according to the Köppen classification, exhibiting two distinct seasons throughout the year. The predominant soils in the region are Quartzarenic Neosols, characterized by being deep, hydromorphic, very acidic, and with low fertility (Sassaki et al., 1999). Smaller proportions of Red-Yellow Latosols, which are deep, well-drained, sandy or sandy-clayey, and acidic, are also found in this area (Sassaki et al., 1999).
2.2 Sampling and sample preparation
The fruits were harvested directly from the plants, selecting only the ripest, undamaged specimens. Upon collection, the fruits were stored in polyethylene containers under refrigeration for three days before the chemical analysis. For the larger fruits, such as the Lobeira and the Araticum, a single specimen weighing approximately 500 g was used for the analysis. For the smaller fruits, such as the Abacaxi-do-Cerrado, the Caraguatá, the Pequi, the Angelim-rasteiro, and the Araçá-do-campo, two to four fruits were combined to obtain a total sample weight of 300 g for each species. The sampling was performed on one to four individual plants per species.
The maturity stages of the fruits varied: the Lobeira, Araçá-do-campo, Araticum, and Angelim-rasteiro were collected at stage 3 (ripe), while for the Abacaxi-do-Cerrado and Caraguatá fruits, half of the batch was at stage 3 (ripe) and the other half at stage 2 (almost ripe). The Pequi fruits were collected at stage 1 (green).
For the determination of the macrocomponents, raw fruit samples were weighed, and cut and the undesirable parts were removed (e.g., the large seeds from the Pequi, Angelim-rasteiro and Araticum, as well as the peduncle). The samples were then ground using a domestic blender (OLIQ501-220A, Oster). In the case of the Abacaxi-do-Cerrado, 40% of the peel was removed to facilitate handling, due to its sharp texture. For the Lobeira, Araçá-do-campo, Caraguatá and Araticum, water was added to 35.5% of the fruit mass (on average) to aid in homogenization. The mass of both the wet samples and the added water was measured on an analytical balance (Radwag AS.R2 Plus Series, Synergy Lab, Brazil).
To establish the potential nutritional value of each fruit, the whole specimen was harnessed, including peels and seeds. The present analysis sought to account for the nutrients located in all fruit parts, including those often discarded by the industry. These parts are known to contain significant nutritional compounds such as carotenoids, enzymes, oils, vitamins as well as other bioactive compounds (Kumar et al., 2020; Kumoro et al., 2020), and can constitute up to 30% of the total fruit mass (Kumar et al., 2020).
2.3 Centesimal composition
The moisture and ash contents were determined following the AOAC method (Association of Official Analytical Chemists, 2006). Approximately 3 g of the sample was placed in pre-treated porcelain crucibles (cleaned and dried at 105 °C for 4 hours) and dried in an oven (TE-395, Tecnal) at 105 °C for 12 hours. After drying, the samples were cooled in a desiccator with silica. The moisture content was calculated using Equation 1:
For the determination of ash content, dried samples were incinerated in a muffle furnace (SSFM - Sinergia Científica) at 550 °C for 12 hours (with a ramp rate of 3.33 °C min−1). After cooling to room temperature, the ash mass was measured on an analytical balance (SHIMADZU ATY224), and the ash percentage was calculated using Equation 2:
Both analyses were conducted in triplicate.
The lipid content was determined using the Bligh & Dyer (1959) method. Approximately 3.0 g of dried, powdered sample was homogenized with a solvent mixture of chloroform, methanol, and deionized water (in a 1:2:3 ratio) for 20 minutes in a rotary shaker (Vortex Basic 772, Sinergia Científica). After separation, sodium sulfate was added to the chloroform phase to remove the residual water, with 5 mL of this phase being transferred to Petri dishes and dried in an oven (TE-395, Tecnal) at 80 °C for 1 hour. The lipid content was calculated by multiplying the lipid mass by 4, according to Equation 3:
This assessment was performed in triplicate for the Lobeira, Caraguatá, Araticum and Abacaxi-do-Cerrado fruits, with duplicates for the Pequi and Araçá-do-campo due to sample limitations.
The crude protein content was determined using the Dumas combustion method (DUMAS NDA 701, Velp Scientifica), adopting a conversion factor of 6.25. Approximately 200 mg of crushed dry sample was combusted at 950-1000 °C under an oxygen atmosphere, and the nitrogen content was quantified by thermal conductivity. Calibration was done using rice flour solutions in eight different concentrations. The protein content was corrected for blank values (n = 5). Carbohydrate content was calculated by the difference between the total mass and the sum of the protein, ash and lipid contents.
2.4 Comparisons with commercial species in the literature
To find previous information available on the chemical composition of fruit species to be included in the present study, the search engines Google Scholar, Elsevier Science Direct, SciELO, and Capes Cafe were used with the keywords “Cerrado fruits”, “Chemical composition”, “Analysis of centesimal composition”, “Nutritional composition” (in English as well as in Portuguese), “Solanum lycocarpum”, “Psidium”, “Bromelia balansae”, “Annona crassiflora”, “Andira humilis”, “Ananas ananassoides” and “Caryocar brasiliense”. This data is compiled in Table 1.
Centesimal composition of the fruits based on the present study’s as well as on literature’s data.
To compare the values obtained regarding the centesimal composition of Cerrado fruits with the species of fruits that are commonly commercialized, the literature was searched for data on apples (Malus domestica Borkh, Rosaceae), bananas (Musa spp. Rumphius, Musaceae) and avocados (Persea americana Mill., Lauraceae).
3 Results
The analyzed fruits presented a high moisture content ranging from 55.2% (Araticum) to 83% (Pequi). The protein content, an important macromolecule for nutrition, corresponded to a range of values from 3.4% (Araçá-do-campo) to 16.2% (Angelim-rasteiro). Additionally, the fruits also showed low amounts of total lipids, ranging from 2.5% to 3.5%, with higher values for the Abacaxi-do-Cerrado (3.5%). Values obtained for ash (1.9% to 4.5%) were similar to those found in the literature for almost all the fruits tested in this study (Table 1). For carbohydrates, the values found ranged from 6.4% (Pequi) to 35.9% (Araticum), which is in line with the literature’s findings for Cerrado fruits of the same or similar type (Table 2).
Centesimal composition percentage of Cerrado fruit pulps for the plant families analyzed in this study, based on a compilation of data collected from the literature (Camargo, 2014; Maruyama et al., 2019).
4 Discussion
The present study contributes to the developing of information concerning species that are still scarcely (Abacaxi-do-Cerrado, Caraguatá) or not studied at all (Araçá-do-campo, Angelim-rasteiro), promoting knowledge related to the composition of available fruit pulps, and further expanding the data on other species (Lobeira, Araticum, and Pequi) (see Table 1).
It was found that the moisture content of the fruits reported in the literature is similar to the values obtained for the Lobeira, the Caraguatá and the Abacaxi-do-Cerrado (Table 1), regardless of the difference in the fruit samples preparation (e.g. inclusion of peels and seeds in the present study’s samples). These fruits are fleshy and, except the Caraguatá, present a larger pulp:exocarp ratio. Therefore, disregarding the pulp, the remaining matter will represent a small contribution to the total moisture content. In addition, the available information on the other fruit’s moisture values (Araçá-do-campo, Aracticum and Pequi) differed when compared with literature’s data (Cordeiro et al., 2013; Lima et al., 2007; Lima & Portari, 2019; Pereira et al., 2018; Roesler et al., 2007; Schiassi et al., 2018), probably due to the greater influence of including the peel along with the pulp in the present study (Dias et al., 2020).
The presence of peel significantly increased the values of ashes and proteins, when compared to the values found in the literature. It is well known that fruit peels and seeds have a considerable concentration of proteins and ashes or, sometimes, even carbohydrates and lipids (Dias et al., 2020). Regarding protein content, the Lobeira, Abacaxi-do-Cerrado, and Angelim-rasteiro presented high values compared to the literature (Table 1), which may be related to the inclusion of seeds and peels in the sample (Pereira et al., 2019; Roesler et al., 2007). The values found for the Abacaxi-do-Cerrado might be a good example of how the peel and seeds can have different protein concentrations from the fruit pulp, since the values obtained herein were higher than in studies in which only the pulp was used; namely, the present study found: 10% of protein, compared to 0.5% to 4.8% protein found by de Ancos et al. (2016) and Paula-Filho et al. (2016). Another interfering factor might be the fruit’s degree of ripeness, given that an unripe fruit can often have higher protein concentrations than a ripe fruit (Lufu et al., 2020; Shi et al., 2014). It's worth noting, however, that a considerable part of this high protein concentration may be due to secondary nitrogen compounds that play a defensive role in immature fruit and can have an unpalatable, unpleasant taste or even some toxicity (Nelson et al., 2023). As some of these proteins can be degraded or modified during the ripening process, there may be a change in the protein concentration between unripe and ripe fruit (Shi et al., 2014). Therefore, the protein values found herein for the Pequi fruit (which included unripe specimens in the sample) should be interpreted with caution. No reference value was found for the Angelim-rasteiro or the Araçá-do-campo and, in the case of the latter, it was compared with related species of the same genus (Lima & Portari, 2019; Pereira et al., 2018). This study is the first to analyze the composition of the fruit of these two species.
Like most other fruits (Jordano, 1995), the Cerrado species sampled herein presented low total lipid levels, in general ranging from 2.5% to 3.5%. (Table 1) The exception was the Pequi, known to have a high lipid content of around 40% (Cordeiro et al., 2013; Lima et al., 2007). In the current study, the value found for Pequi differs from the expected (Table 1), probably due to the degree of fruit ripeness, which is known to influence its chemical content (Geocze et al., 2021; Lufu et al., 2020; Rodrigues et al., 2015). The Pequi and some of the Araçá-do-campo fruits used in this study had not completed their ripening process yet. It is important to emphasize that unripe fruits, such as those from the Myrtaceae family, are often used as food by animals such as parrots, parakeets, and insects (Gilardi & Toft, 2012). Thus, the present data coupled with the literature considers the alterations of the chemical components in fruits that may be eaten by animals, in regards to the selection between ripe and unripe fruits in the sampling process.
The ash represents the inorganic or mineral content (sodium, zinc, potassium, magnesium, manganese, calcium, iron, phosphorus, etc.) present in the sample, remaining after the organic part has been removed. The values obtained for ash content were similar to those found in the literature for almost all the fruits tested in this study. The only exception was the Abacaxi-do-Cerrado, whose values (4.5%) were much higher than those found in the literature (0.8%, according to Paula Filho et al., 2016). Since the present study used the fruit pulp along with the fruit peel for the analysis, while most of the previous studies used as a comparison did not, this may be one of the factors that explain such discrepancy. This assumption is reinforced by the fact that fruit peels tend to have a high concentration of minerals and vitamins, often higher than those found in the pulp (Dias et al., 2020).
Finally, regarding the carbohydrates, the Lobeira, Caraguatá and Pequi presented values similar to those from the literature (Cordeiro et al., 2013; Lima et al., 2007; Lima & Portari, 2019; Pereira et al., 2019;). For the Araçá-do-campo, the high values of carbohydrates obtained may be related to differences in the fruit’s degree of ripeness and storage time, as already established in the literature with others (Prasanna et al., 2007). For the Araticum, a high carbohydrate count was expected (Arruda & Pastore, 2019), but even higher values were found (Table 1), potentially due to the natural variations in the carbohydrate content among plant populations or harvesting sites, as well as storage times, use of different parts of the fruit for the analysis, and the difference between the studies concerning the use of fresh or dried pulp (Yahia et al., 2019). For the Abacaxi-do-Cerrado, the explanation for the low carbohydrate values is probably related to the variations obtained in the ash and protein values, since once the carbohydrates have been calculated by difference, the alterations in the values obtained for the other components can generate a final divergence in the carbohydrate count.
It is worth mentioning that the sugar composition of each fruit differs and can influence the availability of carbohydrates. The Lobeira tends to have a higher concentration of sucrose in its composition, while reducing sugars are less representative (Pereira et al., 2019). For fruits like the Abacaxi-do-Cerrado, Araticum, Pequi, Araçá-do-campo, and Caraguatá the opposite is observed, with a higher presence of fructose and glucose, along with a lower concentration of non-reducing sugars (Cordenunsi et al., 2010; Damiani et al., 2011; Oliveira et al., 2017; Santos et al., 2009).
Factors such as the maturation degree of the fruit (Geocze et al., 2021; Lufu et al., 2020; Rodrigues et al., 2015), both the storage period and conditions, along with the seasonal climate as well as the soil conditions (Lufu et al., 2020) may influence the chemical composition of the fruit and could be related to the discrepancies found in the proportions of protein, lipids, carbohydrates, ash and water between the current study and the literature. As the present study was carried out in a different location than the studies used to compare the data (Table 1), variations related to local physical characteristics are to be expected and may also influence results.
4.1 Comparisons with commercial species
The moisture values obtained in this study ranged from 55.0% to 82.9%, higher than bananas (28.0%) (Aurore et al., 2009) and usually lower when compared to apples (range 4.0% to 80.0%; Vendruscolo et al., 2008) and avocados (73.2% to 88.9%; Mooz et al., 2012). The Cerrado fruits analyzed herein contain more protein, ranging from 3.4% to 16.2%, while the conventional fruits values range from 1.1% to 5.8%, with apples having the highest concentrations (Aurore et al., 2009; Mooz et al., 2012; Vendruscolo et al., 2008). In terms of carbohydrates, the fruits from the Cerrado presented values between 6.3% and 35.9% (Table 1), contrasting with the 5.6% to 83.8% from conventional fruits, with avocados showing the lowest values (5.6% to 11.5%) (Aurore et al., 2009; Mooz et al., 2012; Vendruscolo et al., 2008).
The values obtained for ash (1.8% to 3.5%) and lipid content (2.4% to 3.5%) from the Cerrado fruits (except the Abacaxi-do-cerrado) are close to the values for conventional fruits, including bananas (a mean of 2.0% and 0.3% for ash and lipids, respectively) as well as apples (a mean of 2.3% and 4.1%, respectively) (Aurore et al., 2009; Vendruscolo et al., 2008), whereas the avocado, which is known to be rich in lipids, have lipids values ranging from 4% to 14.7% (Mooz et al., 2012).
Regarding the comparative analysis of the Cerrado fruits’ and commercial fruits’ composition, it is important to take into account the family of the species, since the chemical composition is often a conservative trait along the phylogenetic context (Jordano, 1995). Therefore, the hypothesis that fruits from closer phylogenetic groups have similar chemical composition was reinforced by the values obtained in the present study, when compared to those found in other taxa from the same plant families (Table 2).
The Cerrado fruits evaluated herein show carbohydrate concentrations close to the values found for fruits already exploited by the industry, while about protein, the wild fruits presented an equal or greater nutritional potential than the conventional fruits. Hence, it is recommended that further studies investigate the potential use of these fruits in further detail. In addition, these fruits may represent an increase in food and nutritional diversity, favoring food and nutritional security plans, especially in developing countries such as Brazil (Asprilla-Perea et al., 2022).
In contrast to the other fruits examined, it is suggested that the A. humilis fruits may induce nausea and dysentery, particularly when consumed in large quantities, due to the presence of toxic compounds in its peel and seeds. Nevertheless, this claim is grounded on only two sources that are mostly based on traditional knowledge and did not report how the information was validated (Kuhlmann, 2021; Mattos, 1979), given the lack of comprehensive data on this species in the literature. Consequently, further investigations regarding the potential of A. humilis for human use are worthwhile. Furthermore, according to traditional knowledge, this fruit is believed to be profitable to the pharmaceutical industry, particularly due to the vermifuge properties attributed to its peel and seeds (Mattos, 1979). On the other hand, the Araçá-do-campo (P. grandifolium), which has also been described for the first time in terms of its macrocomponents has shown itself to be a fruit with great potential for human consumption, standing out for its availability of water and carbohydrates, analogous to other Psidium fruits already described in terms of pulp composition (Pereira et al., 2018).
Overall, Cerrado fruits tend to present significant moisture (64.47 ± 14.74), carbohydrate (20.27 ± 13.94) and lipid (21.37 ± 11.35) content, along with lower values for protein (5.74 ± 2.39) and ash (0.96 ± 0.34), based on data available for all families included in the studies of Camargo (2014) and Maruyama et al. (2019) (Tables 1 and 2). The present study found similar values to those evaluated by Camargo (2014) and Maruyama et al. (2019), but the current study stands out for its greater protein count findings, mainly due to the contribution of the Solanaceae, Bromeliaceae, and Fabaceae species, also presenting lower lipid values when compared to the other two studies. However, the available previous literature reviews (Camargo, 2014; Maruyama et al., 2019) do not include data about species from the Solanaceae and Bromeliaceae families, neither from the species sampled by the present study (except for the Caryocar brasiliense), highlighting the scarcity of chemical composition assessments regarding neotropical fruits in an ecological context. The fruits from this study can be eaten by mammals, birds, and reptiles (Kuhlmann, 2021), possibly being an important resource for animals, given the diverse nutritional value and chemical compositions of such fruits.
5 Conclusion
Most of the specimens analyzed herein presented high moisture and carbohydrate contents, comparable to the values found in conventionally marketed fruits. However, some fruits, such as the Lobeira, the Angelim-rasteiro, and the Abacaxi-do-Cerrado, stood out for their high protein count. Most species show potential for human use (except the Angelim-rasteiro, due to its prospective toxicity), by being consumed in natura or by taking part in industry or food components. All of the abovementioned fruits are also important diet components of the Cerrado’s native fauna and bear the potential to increase the economic value of Cerrado areas by harvesting native fruits.
Acknowledgements
We would like to thank the Brazilian Federal Agency for Support and Evaluation of Graduate Education (CAPES) for the scholarship to RDF and Proap CAPES (Finance Code 001) for funding the research, as well as the Neotropical Grassland Conservancy for the grant that made it possible to carry out the chemical analyses, the Fundação Florestal and the Estação Ecológica/Experimental from Itirapina for allowing the study to be conducted in their protected area, and the Laboratório de Inovação de Alimentos (LINA) from Unicamp for making the composition analyzes possible.
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Cite as:
Feliciano, R. D., Sviech, F., Prata, A. S., & Christianini, A. V. (2025). Physicochemical characterization of seven fruits from the Cerrado. Brazilian Journal of Food Technology, 28, e2024092. https://doi.org/10.1590/1981-6723.09224
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Funding:
Funding for fieldwork was provided by Proap CAPES (Funding Code 001) and the Neotropical Grassland Conservancy grant.
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Edited by
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Associate Editor:
Juliano Lemos Bicas.
Publication Dates
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Publication in this collection
30 May 2025 -
Date of issue
2025
History
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Received
03 Sept 2024 -
Accepted
28 Feb 2025