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Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=lfri20 Food Reviews International ISSN: (Print) (Online) Journal homepage: www.tandfonline.com/journals/lfri20 A Comprehensive Review of Eugenia Pyriformis Cambess: Reported Bioactivities and Health Effects Marina Volpato Dacoreggio, Gabriela Soster Santetti, Heloísa Patrício Inácio, Aniela Pinto Kempka & Renata Dias de Mello Castanho Amboni To cite this article: Marina Volpato Dacoreggio, Gabriela Soster Santetti, Heloísa Patrício Inácio, Aniela Pinto Kempka & Renata Dias de Mello Castanho Amboni (2023) A Comprehensive Review of Eugenia Pyriformis Cambess: Reported Bioactivities and Health Effects, Food Reviews International, 39:5, 2477-2491, DOI: 10.1080/87559129.2021.1958226 To link to this article: https://doi.org/10.1080/87559129.2021.1958226 Published online: 30 Jul 2021. Submit your article to this journal Article views: 350 View related articles View Crossmark data Citing articles: 2 View citing articles https://www.tandfonline.com/action/journalInformation?journalCode=lfri20 https://www.tandfonline.com/journals/lfri20?src=pdf https://www.tandfonline.com/action/showCitFormats?doi=10.1080/87559129.2021.1958226 https://doi.org/10.1080/87559129.2021.1958226 https://www.tandfonline.com/action/authorSubmission?journalCode=lfri20&show=instructions&src=pdf https://www.tandfonline.com/action/authorSubmission?journalCode=lfri20&show=instructions&src=pdf https://www.tandfonline.com/doi/mlt/10.1080/87559129.2021.1958226?src=pdf https://www.tandfonline.com/doi/mlt/10.1080/87559129.2021.1958226?src=pdf http://crossmark.crossref.org/dialog/?doi=10.1080/87559129.2021.1958226&domain=pdf&date_stamp=30 Jul 2021 http://crossmark.crossref.org/dialog/?doi=10.1080/87559129.2021.1958226&domain=pdf&date_stamp=30 Jul 2021 https://www.tandfonline.com/doi/citedby/10.1080/87559129.2021.1958226?src=pdf https://www.tandfonline.com/doi/citedby/10.1080/87559129.2021.1958226?src=pdf A Comprehensive Review of Eugenia Pyriformis Cambess: Reported Bioactivities and Health Effects Marina Volpato Dacoreggioa, Gabriela Soster Santettia, Heloísa Patrício Inácioa, Aniela Pinto Kempkab, and Renata Dias de Mello Castanho Ambonia aDepartamento de Ciência e Tecnologia de Alimentos, Universidade Federal de Santa Catarina, Florianópolis, Brasil; bDepartamento de Engenharia de Alimentos e Engenharia Química, Universidade do Estado de Santa Catarina, Pinhalzinho, Brasil ABSTRACT Eugenia pyriformis Cambess, commonly known as uvaia, is a native Brazilian plant. Among its benefits are the high concentration of vitamin C, carote- noids and sources of phytochemicals such as phenolic compounds. Many of these phytochemicals have been linked to the prevention and management of several chronic and degenerative diseases including cancer, cardiovascular diseases, type 2 diabetes mellitus, obesity, among other disorders. The uvaia compounds started to be isolated and characterized to provide a better understanding of the chemical mechanisms related to the biological activ- ities of their extracts. As the identification of compounds progresses, studies investigating the potential of extract are receiving more attention. Some biological activities identified in the different uvaia compounds are antiox- idants, antimicrobials, and anti-inflammatories. Previous research has pointed out that the uvaia fruit extract can be used as a potent anti- inflammatory for inhibited carrageenan-induced paw edema in rats. In this context, this review summarizes the nutritional composition of uvaia and describes the main biological activities reported for this plant, revealing some of its beneficial effects on health and highlighting how much remains to be elucidated. Phytochemicals Health benifits An!oxidant ac!vity An!microbial ac!vity An!-inflammatory ac!vity Vitamin C Vola!le compounds Carotenoids Phenolic compounds KEYWORDS Uvaia; nutritional composition; phytochemicals; bioactive compounds; antioxidant; anti-inflammatory CONTACT Renata Dias de Mello Castanho Amboni renata.amboni@ufsc.br Departamento de Ciência e Tecnologia de Alimentos, Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina SC 88034-001, Brasil FOOD REVIEWS INTERNATIONAL 2023, VOL. 39, NO. 5, 2477–2491 https://doi.org/10.1080/87559129.2021.1958226 © 2021 Taylor & Francis http://www.tandfonline.com https://crossmark.crossref.org/dialog/?doi=10.1080/87559129.2021.1958226&domain=pdf&date_stamp=2023-07-07 Introduction The growing increase in interest on nutrition in recent decades is expanding the concern about diets and human health. Consequently, diets that emphasize the positive aspects of food are increasingly popular.[1] It is known nowadays, through a series of epidemiological studies, that diets rich in fruits and vegetables bring many health benefits. Fruits and vegetables have been persistently identified as natural sources of antioxidants due to the presence of several compounds known for their antioxidant activity such as vitamins, anthocyanins, flavonoids, and other phenolic compounds.[2] Tropical countries produce a large amount of native and exotic vegetable species which are potentially interesting for the food, pharmaceutical, and cosmetic industries.[3] Their attention is focused on polyphenols that are found in fruits (pulp and peels) and leaves since compounds exhibit high antioxidant activities and are hence effective against reactive oxygen species. Uvaia (Eugenia pyriformis Cambess), of the Myrtaceae family, is a native Brazilian species that can be found in the states of São Paulo to the Rio Grande do Sul, as well as in neighboring countries, Argentina and Paraguay.[4] Its name derives from iwa’ya, from the Tupi vocabulary, and means ‘acid fruit,’ and may be known as uvaia, ubaia, uvalha, orvalha, and uvaieira, among others.[5] Although little consumed in nature, the uvaia pulp is used to obtain products such as juices, liqueurs, beers, jellies, sweets, ice cream, yogurts, presenting a high potential for the agrifood sector. In addition, due to its bioactivity (antimicrobial and anti-inflammatory), which may be related to the high content of vitamin C and antioxidants, uvaia fruits may also be valuable to the pharmaceutical industry.[6–8] The reported bioactivity for uvaia fruits is mainly attributed to the high content of phenolic compounds and secondary metabolites with a high antioxidant capacity.[7] These compounds can protect biological systems against excess free radicals and reactive oxygen species.[9] Thus, when included in the human diet, they contribute to reducing the development of degenerative diseases, cardiovascular diseases, cancer, diabetes, among others.[10] Furthermore, the uvaia fruit also contains other interesting chemical compounds such as minerals, proteins, sugars, volatile compounds, and carotenoids, which can also contribute to human health. However, few studies report the potential application of uvaia as a functional and herbal food, as well as exploring different parts of the plant. In this regard, this review highlights the biological activities and chemical composition already reported for fruits/leaves of uvaia in recent years to arouse interest in further exploration of this plant with latent perspectives in the food and pharmaceutical industry. Figure 1. Fruits and leaves of Eugenia pyriformis Cambess. Fig. 1 shows the Eugenia pyriformis Cambess, especially its fruit, when ripe, and its leaves. 2478 M. V. DACOREGGIO ET AL. Uvaia fruit The uvaia fruit is attractive, displaying a yellow or orange coloration and a pleasant aroma, with the pulp being used, mainly, for the manufacture of juices, jellies, and jams.[11] It has an excellent nutritional value, being a source of vitamin C, and minerals, besides having a high content of secondary metabolites with biological activity, such asflavonoids, phenolic acids, and carotenoids.- [6–8] In the southeast, the species flower in August and September, and harvest occurs in October and November,[12] but it may come early or be delayed depending on climatic conditions, especially rainfall. In the southern state of Rio Grande do Sul, Brazil, flowering occurs in November and December and fruit Harvest occurs between January and February.[11] Botanical description The Myrtaceae family, which belongs to the species Eugenia pyriformis, is one of the most striking within the Atlantic Forest biome. Contained in this family, the genus Eugenia comprises more than a hundred species, among them some native to Brazil of great economic importance due to presenting fruits of nutritional value,[13] such as Eugenia brasiliensis Lam. (Grumixama), Eugenia involucrata DC., (cherry), Eugenia pyriformis Camb. (uvaia), and Eugenia uniflora L. (pitanga). Scalon et al.[14] described the existence of two species, E. pyriformis, and Eugenia uvalha. However, E. pyriformis has been accepted as the botanical name for uvaia, and E. uvalha is considered a synonym of E. pyriformis. Jacomino et al.[11] reported the existence of seven E. pyriformis accessions: Bolívia, Comum, Doce, Doce de Patos de Minas, Pêra, Rugosa, and Rugosa Doce. The uvaia is a medium-sized tree (6 a 13 m) with short trunks, up to 50 cm in diameter, and a rounded or elongated crown. Leaves are simple, opposite, glabrous, subcoriaceous, pinkish-red in color when young, and 4 a 7 cm in length. Flowers are white, usually solitary or grouped into 3, hermaphrodite, tetramerous with a great number of stamens. Fruits are berry-type, rounded, with thin, velvety epicarp and yellow or orange fleshy pulp. The trunk is usually erect and the bark tends to peel off in large pieces.[15] Seeds are large, 1 a 3 per fruit, with high germination capacity.[16] The uvaia is native to the Atlantic Forest occurring throughout its extension in the south, southeast, and northeast Brazil. However, its occurrence has been reported in central-west Brazil, outside the Atlantic Forest biome, which can be found in the states of São Paulo to the Rio Grande do Sul, as well as in neighboring countries, Argentina and Paraguay.[17] Commercial uvaia orchards are rare and fruit production comes predominantly from domestic orchards, which makes it difficult to collect basic data that are essential to estimate crop performance. According to Jacomino et al.[11] 10-year old plants pruned to a canopy height of 2.5 m produce 5 kg of fruit per plant on average. However, some plants may produce up to 50 kg of fruit in a harvest season. The plant is extremely adaptable and can be grown in climates with annual temperatures between 18 to 26 degrees Celsius, resisting frosts of up to minus 4 degrees Celsius. Industrial applications of uvaia Despite the small number of studies, the potential for commercial exploitation of uvaia is comparable to that of other tropical fruits. The physicochemical attributes of uvaia provide opportunities for its use in different sectors, including the food and pharmaceutical industries. Because of its acid and distinctive taste and pleasant aroma, uvaia fruits have a large industrial potential for consumption in the form of jams, juices, ice cream, liqueurs, compotes, conserves, or added to other processed products such as yogurts.[6,12,17,18] In recent years, there has been a growing interest in the combining of sensory attributes and protective effects of native fruits against chronic and degenerative diseases. In the case of uvaia, these effects have been attributed to the antioxidant capacity of phenolics and carotenoids found in its pulp.[11] FOOD REVIEWS INTERNATIONAL 2479 Products including compotes, jam, liqueur, ratafia, ice cream, and ice cream bars have been developed for the Federal Institute of Southern Minas Gerais with the object to retain the sensory and nutritional attributes of fresh fruit. Acceptability scores were high for all products in sensory analysis, especially for recipes that ice cream bars with high pulp content. In terms of granted patents, Araújo et al.[7] verified patent applications related to uvaia. There are currently two Brazilian patents, one for ice cream of uvaia of low-value caloric fortified with iron (BRPI1103266) and the other about the process for obtaining an alcoholic drink based on fruits and natural fruits juice (BRPI0601538). Chemical composition Nutritional composition of uvaia pulp The nutritional value of uvaia is detailed in Table 1. Carbohydrates were the most abundant macro- nutrient, whereas ash, lipids and proteins were detected in lower amounts. Regarding the dietary fiber content, researchers noted variations between 31.86 and 44.10 g 100 g−1 in the total dietary fiber content in different accessions of uvaia fruits. Regular intake of soluble or insoluble fibers can contribute to the reduction of cardiovascular diseases, diabetes, and colon cancer, as well as balancing cholesterol, high blood pressure, and blood glucose levels. Still, it is suggested that eating a 200 g day−1 serving of fresh uvaia fruits could provide 15.97% to 25.68% of the dietary reference intake (DRI) of dietary fiber for men and 24.28 to 39.03% for women, which highlights the fruit when compared to the DRI of other fruits considered as sources of dietary fiber, such as blueberry (3.4%) and avocado (13.6%).[23,24] Table 1. Nutritional value of Eugenia pyriformis cambess. Composition Content (unit per 100 g) Reference Moisture 94.50 g b [19–21] Ashes 0.23 g b [19,21] Lipids 0.52 g a [19,21] Protein 15.82 g a [19,21] Total dietary fiber 31.86–44.10 g a [19,21] Insoluble dietary fiber 22.35–39.05 g a [21] Soluble pectin 0.17–0.18 g GA b [22] Total pectin 0.95 g GA b [22] Total sugars 36.72 g a [21,22] Reducing Sugar 36.54 g a [21] pH 2.96–3.06 b [19,22] Total Titratable acidity 1.17–1.67 g CA b [21,22] Total soluble solids 7.70–10.50 º Brix [19,21,22] Vitamin C 39.5–122.51 mg AA b [20,21] Vitamin A 3.78 mg RE a [19,21] P 10.84 mg b [22] K 99.14 mg b [22] Ca 7.59 mg b [22] Mg 5.65 mg b [22] Cu 0.09 mg b [22] Fe 3.38 mg b [22] Mn 0.16 mg b [22] Zn 0.11 mg b [22] Total phenols 3.48 g GAE a [21,22] Yellow flavonoids 7.30–17.50 mg b [20,22] Total athocyanins 1.13 mg b [20] Total carotenoids 2.07–2.56 mg b [20,22] GA – Galacturonic acid. CA – Citric acid. AA – Ascorbic acid. RE – Retinol equivalent. GAE – Gallic acid equivalent.TE – Trolox equivalent. a Values expressed on the basis of dry weight. b Values expressed on the basis of fresh weight. 2480 M. V. DACOREGGIO ET AL. Macronutrients Carbohydrates According to Silva et al.[24] simple carbohydrates can give 13.57 kcal to 25.97 kcal in 100 g of uvaia pulp, which represents only 0.67% to 1.29% of a 2.000 kcal daily diet−1. In this context, the uvaia fruits have lower energy density than citrus fruits, such as oranges (46 kcal 100 g−1), tangerine (53 kcal 100 g−1), and lemons (29 kcal 100 g−1), as well as non-citrus fruits such as strawberries (32 kcal 100 g−1) and mangoes (60 kcal 100 g−1). The levels of non-soluble carbohydrates, characterized as complex, including cellulose, hemicellulose, and lignin, represent, on average, 30% of the total weight of uvaia fruits. Protein and amino acids Although the uvaia consumption provides a small contribution to the recommended intake of proteins, it may be relevant in combination with other components present, such as vitamins and phenolic compounds. It can also participate in the prevention of various diseases such as chronic constipation and irritable bowel syndrome, in addition to other disorders caused by oxidative stress.- [25,26] The recommended daily protein intake is 0.8 g.kg−1day−1 for an adult, and while the consump- tion of uvaia fruits makes a small contribution to the recommended protein intake, it can provide potential amounts of essential (leucine, lysine, tyrosine, etc.)and non-essential amino acids (gluta- mine, asparagine, alanine, etc.)[23] that are fundamental to the proper functioning of the human body. These amino acids act on cell signaling, gene expression regulation, and protein phosphorylation cascade.[27] Fatty acids The presence of lipids and their respective fatty acids in fruits is responsible for being a precursor of several characteristic plant components, such as volatile compounds and flavoring agents.[28] Studies involving the characterization of the fatty acid profile for both uvaia fruits (pulp, peels, or seeds) and leaves were not found. However, when analyzing the lipid composition of fruits and seeds of the same family of uvaia, Myrtaceae, a high presence of unsaturated fatty acids is observed. Luzia and Jorge[29] demonstrated that the lipid composition of the seeds of jambolan (Syzygium cumini L.) is composed of about 68% of unsaturated fatty acids, including 24% of monounsaturated and 44% of polyunsaturated acids, with oleic acid (C18:1) and linoleic acid (C18:2) as major components. Regarding that the weight of the uvaia fruit is about 8.5 g, with the seeds corresponding to approximately 16% of the total weight of the fruit, the study of the lipid profile of uvaia seeds becomes a relevant parameter to be studied, considering the role of lipids in the maintenance of human health and its aid in the inflammatory response with antimutagenic and anticarcinogenic action.[30] Organic acids Organic acids (citric, tartaric, malic, lactic, succinic, and the sum of the acids) were identified and quantified in the investigated six uvaia accessions (‘Comum’, ‘Rugosa Doce’, ‘Dura’ and ‘Doce de Patos de Minas’). The sum between the accessions ranged from 0.28 to 1.36 g 100 g−1 (f.w.), with the ‘Comum’ and ‘Rugosa Doce’ accessions being more acidic, while ‘Rugosa’ and ‘Pêra’ displayed lower acidity contents. Organic acid profiles were different among uvaia accessions, mainly succinic and malic acid in four of the six accessions, verifying that succinic and malic acids contribute significantly to the peculiar taste attributed to uvaia fruits.[19] Micronutrients Minerals Fruit consumption is generally recommended as it is also a source of minerals. The absence or insufficiency of the ingestion of these micronutrients can cause irreversible diseases such as cretinism, FOOD REVIEWS INTERNATIONAL 2481 iodine deficiency, or infertility, as well as lack of zinc. In contrast, regular intake of minerals such as selenium and zinc stimulate and strengthen the immune system, improving the inflammatory response.[31,32] The amount of minerals present in the uvaia pulp suggests the fruit is a good source of these micronutrients. Silva et al.[22] evidenced the potassium (K) mineral in significant amounts (99.15 mg.100 g−1), similar to those found in watermelon (104 mg 100 g−1) and higher than those found in apples (75 mg 100 g−1). The amount of phosphorus (P) (10.85 mg.100 g−1) was similar to calcium (Ca) (10 mg 100 g−1). On the other hand, the magnesium content (Mg) (5.65 mg 100 g−1) was similar to that found in peach, plum, apple, and Italian grape. Finally, the uvaia pulp has a high iron (Fe) content (3.38 mg 100 g−1), which is of great interest since this microelement plays a fundamental role in hemoglobin and myoglobin, and other metabolic processes. Vitamins Franco[33] demonstrated the presence of vitamins A (30 µg 100 g−1), B1 (40 µg 100 g−1), B2 (40 µg 100 g−1), and B3 (0.5 mg 100 g−1) in his studies with E. pyriformis, conferring an important vitamin content for uvaia pulp. In particular, vitamin C, or ascorbic acid, has distinct physiological functions. The two most important functions are the high antioxidant activity, which protects cell membranes and lipoproteins against lipid peroxidation, and the absorption of plant-derived iron.[34,35] The uvaia pulp can be considered a rich source of vitamin C, ranging from 1.37 mg 100 g−1 to 70 mg 100 g−1. [7,20,24] Higher levels of ascorbic acid have also been reported with values between 110.51 mg 100 g−1 and 122.51 mg 100 g−1.[22] In this sense, consuming 100 g of fruit can provide more than the recommended daily intake of vitamin C for adults. Phytochemicals Carotenoids Carotenoids are the second most abundant natural pigments on earth, with over 750 members.[36] In addition to their plant-specific functions, carotenoids play important roles in human health and nutrition as essential components in human diets. Total carotenoid levels in uvaia pulp vary considerably, with values ranging from 130.7 to 441.26 mg β-carotene/100 g fresh weight.[19,24,37] However, in other studies, the total carotenoid values found were lower, ranging from 1.7 to 32.1 mg β-carotene/100 g fresh weight.[20,22] Table 2. Carotenoids from pulp of Uvaia (Eugenia pyriformis Cambess). Compounds Concentration ReferenceExpressed as µg/g dry matter 9-cis-B-carotene 0.56 [7,19] all-trans-lutein 0.86 [7,19] All-trans-zeaxanthin 0.56–0.78 [7,19] Phytoened 0.06 [7] All-trans-zeinoxanthin 0.56–0.78 [7,19] 5,8-epoxy-B-carotene 0.33 [7,19] 9-cis-B-cryptoxanthin 0.58 [7,19] 9-cis-neoxanthin 0.29 [7] All-trans-neochrome 0.12 [7] Cis-antheraxanthin 0.22 [7] 9-cis-violaxanthin 0.19 [7] 5,6-epoxy-B-cryotoxanthin 0.24 [7] 5,8-epoxy-B-cryotoxanthin 0.36 [7] 13-cis-B-cryptoxanthin 0.22 [7,19] Phytoene 0.06 [19] All-trans -B-cryptoxanthin 5.2 [19] All-trans-alfa-carotene 0.14 [7,19] All-trans-B-carotene 1.71 [7,19] 2482 M. V. DACOREGGIO ET AL. Table 2 shows the individual carotenoids reported for the uvaia pulp. The main individual carotenoid compounds found were all-trans-β cryptoxanthin (5.21 ± 0.18 µg/g dry matter), followed by all-trans-β carotene (1.71 ± 0.11 µg/g dry matter).[7,19] Other authors, when studying other species of the Myrtaceae family, found that the composition of carotenoids is quite similar, highlighting the same predominant compounds.[20,21,23] The roles of carotenoids in human health have been reported by several authors, who confirmed their roles in reducing the risk of cardiovascular disease, oxidative stress, and low-density lipoprotein oxidation (LDL),[38,39] as well as its role in eye health, macular pigment formation, and cancer incidence reduction.[40] Volatile compounds Volatile compounds are part of a diverse group of organic compounds, such as aldehydes, ketones, and other light hydrocarbons, which have been used to extend the shelf life of food products because they have natural preservative properties.[41] These compounds have high vapor pressure under ambient conditions, as well as low molecular weight (<250 Da). In fruits, volatile compounds are responsible for the aroma characteristics, and they include several chemicals.[42] The volatile composition of uvaia pulp has been reported to date by two authors using solid-phase microextraction (SPME).[24,37] Terpenes were the main constituents in both studies, corresponding between 46.75% and 47.46% of the identified compounds. However, the total volatile compounds found ranged from 59 to 77, due to the genetic variability of the sample, as well as edaphoclimatic conditions. Silva et al.[24] when investigating six distinct accessions, reported that the volatile compo- sition varied according to the adhesion of the uvaia fruit, in which only 3 esters (ethyl butyrate, hexyl hexanoate, and 3- hexenyl hexanoate), 3 sesquiterpenes (o-elemene, cubebene and (–δ-cadinene), and 3 aliphatic alcohols and aldehydes (hexanal, 1-hexanol, and nonanal) were reported in all accessions. Uvaia fruits share a common pattern of sweet, fruity citrus aroma. The compounds with the most significant impact on citrus odor characteristics were (p)-bourbonene, 3-hexenol, and 2-transhexenil. The combination of compounds such as ethyl butyrate and hexyl acetate is responsible for the sweet and fruity aroma notes.[37] Besides their flavoring properties, several studies have reported the potential role ofthese volatile compounds in human health, including anti-inflammatory, antioxidant, anti-obesity, and anticancer activities.[41] Terpenoids, for example, the major volatile compounds present in uvaia fruits, cause inhibition of tumor cell proliferation, inducing tumor cell death and consequently inhibit specific types of cancer such as breast and prostate.[42] Its compounds, alone or in combination, also have proven antioxidant, anticancer, and anti-inflammatory activity, as well as an antinociceptive and hepatoprotective effect.[43–46] Phenolic compounds The total content of phenolic compounds and total flavonoids of the uvaia pulp were reported between 126.5 mg and 483.25 mg of gallic acid equivalent (GAE) 100 g−1 and between 0.04 mg and 38.30 mg of quercetin equivalent 100 g−1, respectively.[7,20,24,47–49] However, in other studies, the phenolic com- pound content ranged from 1930 mg to 3028.58 mg GAE 100 g−1 when the uvaia pulp was lyophilized.[20,37] Disparities were also observed in the contents of phenolic compounds, depending on the solvent used for extraction, where Rufino et al.[49] found that most pulp compounds were extracted using hydroethanol, followed by ethanol and water (34.70 mg, 107.14 mg, and 189.41 mg of GAE 100 g−1, respectively). The total phenolic content also varies considerably between different accessions. Six different accessions of uvaia pulp presented contents ranging from 271.67 to 413.02 mg of GAE. 100 g−1.[24] All of these quantitative differences observed for the total phenolic content of uvaia fruits may occur due to cultivation region, edaphoclimatic conditions, year of study, and extraction method. Table 3 shows the individual phenolic profile of the uvaia pulp. The phenolic composition is responsible for 13 compounds, and gallic acid, galloyl-bis-HHDP hexoside, and quercetin are the main ones (Table 3). FOOD REVIEWS INTERNATIONAL 2483 Although the Eugenia pyriformis Cambess has a phenolic profile already described for the pulp, there is a lack of studies reporting the complete individual phenolic profile, by different methodologies and for other parts of the plant. The essential oils obtained by hydrodistillation from different parts of the uvaia plant present different compositions. In leaf oils, the main components are β-pinene, limonene, 1,8-cineol, and caryophyllene oxide, the flower oil has E-caryophyllene and germacrene D as major components and the fruit oil has caryophyllene oxide and limonene as its main components,[50] while Durazzini et al.[51] observed that the five major compounds identified in essential oil from aerial parts are limonene (14.8%), nerolidol (11.0%), a-cadinol (10.3%), caryophyllene oxide (9.9%) and b-pinene (7.1%). Phenolic compounds are secondary metabolites of plants and are critical for their growth and reproduction. Also, they are formed in stressful conditions such as wounds, infections, UV radiation, among others.[9] There is evidence that the consumption of phenolic compounds in foods may decrease the risk of health disorders due to their antioxidant activity. Phenolic compounds are classified as primary antioxidants, which are mainly free radical scavengers that delay or inhibit the initiation step or stop the propagation step of lipid oxidation.[23,52,53] They are known to reduce the risk of heart diseases, cancer, and diabetes. They also inhibit cyclooxygenase (COX) activity, plasma platelet aggregation, and histamine release, as well as exert anti-inflammatory, antiallergic, antiviral, and antibacterial activities.[10] In fact, the main phenolic compounds present in the uvaia pulp (gallic acid and galloyl-bis-HHDP hexoside) are described with neuroprotective,[54] anticancer,[55] anti- inflammatory,[56] hepatoprotective,[57] and gastroprotective[58] properties. Biological activities Because it is considered a source of bioactive compounds, some studies have been conducted with uvaia to correlate its chemical composition with the biological effects observed by in vitro and in vivo models. This information suggests that the fruit and other botanical parts of uvaia have the potential to be used to prevent non-communicable chronic diseases. Species of the Eugenia genera have important pharmacological activities, such as antioxidant, antihypertensive, antidiarrheal, hypothermic, antimi- crobial, and antifungal.[59] However, new research aimed at identifying and quantifying compounds of biological interest, as well as the use of cellular and animal tests to establish safe and efficient dosages, must be carried out to guarantee the health and welfare of consumers. Table 3. Individual phenolic compounds found in pulp of Uvaia (Eugenia pyriformis Cambess). Compounds Concentration ReferenceExpressed as mg/100 g Gallic acid 7.5–29.9 [19,48] Galloyl-bis-HHDP hexoside 19.3 [19] Quercetin 12.4–18.1 [48] Gallic acid derivative 14.8 [19] Trigalloyl acid lactonized 13.8 [19] Gallic acid derivate 11.0 [19] Galloyl hexoside 5.1 [19] Quercetin deoxyhexoside 4.8 [19] Chlorogenic acid 2.72–4.23 [48] Myricetin 2.29–2.54 [48] Dicaffeic acid 2.0 [19] Galloy hexoside isomer 1.8 [19] Ferulic acid 0.25–0.33 [48] Kaempferol 0.27 [48] p-coumaric acid 0.092–0.29 [48] Rutin 0.083–0.11 [48] 2484 M. V. DACOREGGIO ET AL. In vitro and in vivo antioxidant activity It is known that free radicals, especially when in excess, can promote disturbances in the body, as they induce oxidation, causing possible damage to lipids, membranes, and alterations in cell DNA.[60] Also, many diseases like Alzheimer’s, cancer, cardiovascular, diabetes, and even weight gain can be spread or started due to such cell damage.[55,56] For this reason, the elimination of free radicals is essential to prevent or slow the onset of some diseases related to the oxidative process. In this sense, antioxidant compounds are considered important for health maintenance.[61,62] Studies suggest that the uvaia fruit (Eugenia pyriformis Cambess) has a high antioxidant potential when compared to other Brazilian fruits, having many phytochemical constituents with proven activity.[20,21,63–65] In most research, the uvaia pulp, juice, or the whole fruit were used for extraction and quantification of phenolic compounds with antioxidant activity. To prepare the extracts, solvents such as water, ethanol, acetone, and methanol were used. At the same time, the antioxidant capacity was determined by in vitro methodologies standardized in the literature such as 2,2-diphenyl-1-picrylhydraz (DPPH), 2,2ʹ-azinobis (3-ethylbenzothiazoline-6-sulfonic acid) (ABTS), ferric reducing antioxidant power (FRAP), and oxygen radical absorption capacity (ORAC) (Table 4). Rufino et al.[49] analyzed the uvaia fresh pulp by the DPPH method. They noticed that uvaia has a high antioxidant capacity when compared to other fruits such as açaí and mangaba. The same authors proposed a study using fresh pulp and freeze-dried uvaia pulp to determine the antioxidant potential. The DPPH assay yielded results similar to the previous study. However, in this study, two other methodologies were employed (ABTS and FRAP assay), in which the authors observed that in both tests, the lyophilized pulp presented the highest values of antioxidant activity when compared to fresh pulp.[20] Supported by the studies, Silva et al.[22] observed that the lyophilized uvaia pulp presented antioxidant capacity values of 15.45 and 17.31 mmol Trolox 100 g−1, evaluated by ORAC and DPPH, respectively. The authors also reported that uvaia trees produce large fruits with desirable characteristics for in natura commercialization. However, smaller fruits are firmer, sweeter, and have Table 4. Antioxidant activity of Uvaia (Eugenia pyriformis cambes) extracts evaluated in vitro. Part of plant/solvent Assay (s) Response Reference Pulp fresh/methanol:water (50: 50, v/v) and acetone:water (70: 30, v/v) DPPH 3246.5 EC50 (g/g DPPH) [49] Pulp fresh and pulp freeze-dried/methanol:water (50: 50, v/v) and acetone:water (70: 30, v/v) DPPH, ABTS, and FRAP DDPH ranged from 3247 and 276 EC50 (g/g DPPH; while ABTS from 18 and 182 µmol Trolox/g, and FRAP from 35.4 and 408 µmol Fe2SO4/g, respectively for pulp fresh and pulp freeze-dried [38] Whole fruit frozen/ethanol:water (40: 60, v/ v) DPPH 45.49 EC50 (g/g DPPH) [48] Whole fruit freeze-dried/ethanolic extract DPPH DPPH ranged from 50 to 90% (percent inhibition) for differents genotypes of uvaia [64] Pulp fresh/methanol:water (50: 50, v/v) and acetone:water (70: 30, v/v) DDPH and ABTS 170.26 EC50 (g/g DPPH) and 336.26 µmol Trolox/g, respectively [21] Pulp freeze-dried/ethanol:water ORAC and DPPH ORAC and DPPH ranged from 13.79 to 17.09, and from 16.77 to 17.84 mmol Trolox/100 g−1 lyophilized pulp, respectively [19] Uvaia juice/water extraction DPPH Uvaia juice of 1 to 10% (v/v) DPPH assay ranged from 11.06 to 94.93% (percent inhibition) [66] Pulp freeze-dried/acetone:acetic acid: water (70: 1: 29, v/v/v) DPPH and ABTS 7.95 and 11.26 mmol Trolox/100 g−1, respectively [65] Pulp freeze-dried/methanol:water (80:20, v/v) ORAC and DPPH ORAC ranged from 4.45 to 16.12 mmol Trolox/100 g−1, while DPPH ranged from 9.94 to 29.71 mmol Trolox/100 g−1 for six accessions of uvaia [24] Pulp freeze-dried/water extraction ORAC Storage period in three stages, ranging from 220 to 240 mol/ g trolox (d.m) [37] ABTS: 2,2′-azinobis(3-ethylbenzothiazoline-6-sulphonic acid); DPPH: 2,2-diphenyl-1-picrylhydrazyl;+ FRAP: ferric reducing antioxi- dant power; ORAC: oxygen radical absorbance capacity. D.M: dry matter. FOOD REVIEWS INTERNATIONAL 2485 higher levels of natural antioxidants and can be consumed both fresh and processed. The same authors, in a recent study,[24] evaluated different accessions of uvaia (6 accessions) concerning the antioxidant capacity and observed results similar to those found in their previous study, as well as when compared to literature data using the ORAC and DPPH methods. They also reported that the two assays used had a positive correlation, and the DPPH method showed a positive correlation with the phenolic compounds of the six accessions of uvaia. In turn, Haminiuket al.[63] evaluated the antioxidant capacity of different fruits. They noted that jabuticaba and uvaia extracts presented the best results for DPPH free radical scavenging efficiency (72.41 and 71.08%, respectively). Also, in a study conducted by Ramirez et al.[64] in which they evaluated 12 uvaia genotypes, they noted that genetics is directly related to phenolic content and, consequently, to their antioxidant capacity since there was a difference between the genotypes studied. Egea & Pereira-Netto[65] evaluated different fruits during ripening (lemon guava, strawberry guava, gabiroba, uvaia, and butiá), where the progressive reduction in antioxidant activity by the ABTS and DPPH methods for uvaia fruits (12.75–11.26, and 9.69–7.95 mM Trolox g−1, respectively) was noted over time. However, Freitas et al.[37] have not found any difference in the antioxidant activity of the fruits throughout the storage period, suggesting that uvaia can be harvested in the green phase, keeping the same bioactive potential of its ripe fruits. In vivo studies may show the real effectiveness of bioactive compounds in the body. However, they are scarce or nonexistent concerning the ingestion of uvaia fruits or other parts of the plant. Nevertheless, there is a study conducted by Lopes et al.[66] in which rats were fed a high-fat diet to investigate the effects of uvaia juice intake on possible oxidative damage as well as antioxidant enzyme levels and activities. The authors observed that the animal model, when added to the diet with uvaia juice, showed an effective relationship in reducing oxidative stress, hepatic carbonyl protein accumu- lation, and preventing the reduction of antioxidant enzyme activity. Antimicrobial activity A study by Souza et al.[67] evaluated the antimicrobial activity of the leaf and stem of E. pyriformis. They observed that both leaf and stem extracts were effective against Enterococcus faecalis and Staphylococcus aureus (MIC = 62.5 µg mL−1) while the antifungal activity evaluation showed that extracts are promising candidates to control leveduriform fungi strains (Candida albicans ATCC 40175, Candida krusei ATCC 40,174, and Candida parapsilosis ATCC 40038) with MIC values ranging from 7.81 and 62.5 µg mL−1. Agredo,[68] by studying the essential oil of the uvaia fruit, also obtained bacteriostatic activity against the strains of Staphylococcus aureus, Bacillus cereus, and Pseudomonas aeruginosa. Bactericidal activity was also observed against Listeria innocua and Escherichia coli. In contrast, the uvaia extract showed bactericidal activity against L. innocua and a bacteriostatic behavior against S. aureus and Candida albicans. Durazzini et al.[51] investigated the antifungal activity of essential oil from aerial parts of the uvaia. The oil exhibited high antifungal activity against Malassezia furfur (MIC = 30 µg/ mL) and may be considered an ally in the preparation of herbal formulations.The antimicrobial activity of plants is mainly related to the presence of phenolic compounds in leaves and fruits. The uvaia fruits contain flavonoids (quercetin and kaempferol), which are well-known antimicrobial agents. The mode of action of these compounds is related to the formation of complexes with metal ions, limiting their accessibility to microbial metabolism or their reaction with the microbial cell membrane that inactivates essential enzymes.[69] Anti-inflammatory activity A study conducted by Ramirez et al.[7] showed that the uvaia fruit extract (0.5 and 1.0 g kg−1) administered to rats inhibited carrageenan-induced paw edema. The percentage of edema inhibition was about 51% and 43% for the total extracts. This demonstrated that administration of E. pyriformis 2486 M. V. DACOREGGIO ET AL. extract inhibited the edema starting from the first hour and during all phases of inflammation, which is probably inhibition of different aspects and chemical mediators of inflammation. In turn, when evaluating hydroethanolic extracts of the leaves, Theoduloz et al.[70] showed that the presence of flavonoids inhibits the enzyme xanthine oxidase, and it is used in the treatment of gouty arthritis, a disease caused by the accumulation of uric acid in the joints. The anti-inflammatory ability makes it a suitable candidate for consideration as a dietary supplement to reduce stress oxidative and inflammation. Infante et al.[71] have analyzed leaves, seeds, and pulp extracts from Eugenia leitonii D. Legrand, Eugenia involucrata DC., Eugenia brasiliensis Lam., and Eugenia myrcianthes Nied for anti- inflammatory activity. Based on the results, they suggested that the anti-inflammatory activity of these native fruits is related to the modulation of neutrophil migration, through the inhibition of cytokines, chemokines, and adhesion molecules. Other studies have also associated species from the Eugenia genera with white blood cell influx, polymorphonuclear neutrophils migration to the inflam- matory site, and decreasing exudate,[72] which corroborates the preliminary data found so far. In vitro antileishmanial activity Durazzini et al.[51] investigated the in vitro antileishmanial activity of essential oil from aerial parts of Eugenia pyriformis Cambess. The oil showed strong antileishmanial activity against promastigote forms of Leishmania amazonensis (IC50 = 2.16 µg/mL), an important result that also adds value to several studies of natural products that can be applied to the treatment of this neglected disease (leishmaniasis). In turn, Nunes et al.[73] when evaluating the antileishmanial potential of the essential oil of another Eugenia sp. Eugenia piauhiensis Vellaf suggests the essential oil and its major constituent γ-elemene as promising candidates for new drugdevelopment against leishmaniasis. Cytotoxic activity Silva et al.[72] found that the extract of the uvaia leaves was not considered cytotoxic, showing an LC50 greater than 1000 µg mL−1. It can be concluded that this species does not have substances with cytotoxic characteristics in high concentrations. The Eugenia uniflora extract and the Eugenia jambo- lana flavonoid fraction presented low cytotoxicity when a study by Sobral and collaborators,[74] opening the floor for new biological studies. Thus, the absence of this cytotoxicity may be an indicator that the biological system can well tolerate the plant. However, more detailed studies to evaluate the toxicity of this extract are necessary using other models (in vitro and in vivo), in view that Gasca et al.[75] evaluated if the aqueous leaf extract of E. dysenterica exerts a moderate cytotoxic effect on human neuroblastoma, SH- SY5Y, (at concentrations higher than 7.8 mg/ml given for 24 h) cells line. Conclusions Eugenia pyriformis Cambess comprises a promising matrix for biotechnological and pharmaceutical purposes. Both the nutritional value and the phytochemical composition make the fruit of this plant a functional food with a great prospect, given its remarkable biological effects. Different studies indicate promising pharmacological properties, mainly related to their chemical properties, where phenolic compounds present in uvaia pulp, such as gallic acid and quercetin, are described for their antioxidant, antimicrobial, anti-inflammatory activity. However, there is still much to be investigated about this exotic tropical plant, especially concerning other parts of the plant, such as its leaves. To extend the research to the nutritional benefits of uvaia, studies on the bioavailability of the various nutritional components can reveal unique attributes of absorption in the intestine, bringing additional health benefits when compared to other fruits. Thus, further efforts should be made to investigate other FOOD REVIEWS INTERNATIONAL 2487 biological activities, as both fruit and other parts of uvaia can be useful in preventing and treating oxidative stress-related conditions and potential candidates to treat other diseases. Disclosure statement The author declares that there is no conflict of interest. Funding This work was supported in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001. Renata D. M. C. Amboni is granted a fellowship (PQ1D) from CNPq. 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FOOD REVIEWS INTERNATIONAL 2491 https://doi.org/10.5897/ajpp2014.4100 https://doi.org/10.1016/j.foodchem.2011.03.119 https://doi.org/10.1016/j.foodchem.2011.03.119 https://doi.org/10.1016/0378-8741(88)90149-3 https://doi.org/10.1371/journal.pone.0152974 https://doi.org/10.25110/arqsaude.v19i3.2015.5550 https://doi.org/10.1016/j.cbi.2021.109429 https://doi.org/10.1016/j.apjtb.2017.08.007 https://doi.org/10.1016/J.FCT.2017.02.032 Abstract Introduction Uvaia fruit Botanical description Industrial applications of uvaia Chemical composition Nutritional composition of uvaia pulp Macronutrients Carbohydrates Protein and amino acids Fatty acids Organic acids Micronutrients Minerals Vitamins Phytochemicals Carotenoids Volatile compounds Phenolic compounds Biological activities In vitro and in vivo antioxidant activity Antimicrobialactivity Anti-inflammatory activity In vitro antileishmanial activity Cytotoxic activity Conclusions Disclosure statement Funding References