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Tamarillo (Solanum betaceum): Chemical composition, biological properties, and

product innovation

Sunan Wang, Fan Zhu

PII: S0924-2244(19)30367-X
DOI: https://doi.org/10.1016/j.tifs.2019.11.004
Reference: TIFS 2653

To appear in: Trends in Food Science & Technology

Received Date: 10 May 2019

Revised Date: 30 October 2019
Accepted Date: 2 November 2019

Please cite this article as: Wang, S., Zhu, F., Tamarillo (Solanum betaceum): Chemical composition,
biological properties, and product innovation, Trends in Food Science & Technology (2019), doi: https://
doi.org/10.1016/j.tifs.2019.11.004.

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1 Tamarillo (Solanum betaceum): Chemical composition, biological properties, and product

2 innovation

3 Sunan Wang a, b Fan Zhu b,*

a
4 Canadian Food and Wine Institute, Niagara College, 135 Taylor Road, Niagara-on-the-Lake,

5 Ontario, LOS 1JO, Canada

b
6 School of Chemical Sciences, The University of Auckland, Private Bag 92019, Auckland 1142,

7 New Zealand

*
8 Correspondence, email: fzhu5@yahoo.com

10

1
11 Abstract

12 Background

13 Tamarillo (Solanum betaceum) is native to South America and is cultivated in several other parts

14 of the world for its valued fruit. The fruit has therapeutic and nutritional properties that can

15 satisfy the demands from health-conscious consumers. There has been increasing interest in

16 using tamarillo for consumption and food and non-food product formulations. Overall, tamarillo

17 is an underutilized, sustainable fruit crop with great potential for value-added products.

18 Scope and Approach

19 This review summarizes the composition of nutritional components and biological properties of

20 tamarillo. The chemical and biological properties of tamarillo are compared to those of common

21 fruits and vegetables. Food and other uses of tamarillo are described. Innovative methods for the

22 development of tamarillo-based products are suggested to maximize the potential of this fruit.

23 Key Findings and Conclusions

24 Tamarillo has a range of nutrients including dietary fibers, polyphenols, vitamins C, A, B6, and

25 E, carotenoids, and potassium and iron. Health effects of tamarillo included antioxidative,

26 antiproliferative, antinociceptive, antiinflammatory, allergenicity, and anti-obese properties.

27 These health effects have been tentatively correlated to certain phytochemicals (e.g., non-starch

28 polysaccharides, carotenoids, and anthocyanins). The properties of tamarillo-derived ingredients

29 mainly depended on the cultivar and origin, plant parts, extraction conditions and analytical

30 procedures. Research is needed to explore the composition-activity relationships and

31 mechanisms underlying a biological activity. An array of tamarillo based products have been

2
32 developed. Overall, tamarillo is a promising “new” fruit crop with potential to be exploited as a

33 source of “healthy” products.

34 Keywords: tree tomato; tamarillo fruit; bioactive phytochemicals; underutilized species;

35 polyphenol; Cyphomandra betacea

36

37

3
38 1. Introduction

39 Tamarillo (Solanum betaceum Cav. syn Cyphomandra betacea Sendt.) is a small

40 flowering perennial tree or shrub species (2–4 m high) of the genus Solanum in the nightshade

41 (Solanaceae) family. Tamarillo is native primarily to South America (Andean regions of

42 Colombia, Peru, Chile, Ecuador and Bolivia). Other important food species in the nightshade

43 family include potato (S. tuberosum), tomato (S. lycopersicum), pepper (Capsicum annuum) and

44 eggplant (S. melongena). New Zealand, Colombia, Australia and USA (California) are the main

45 countries involved in commercial tamarillo cultivation (Morton, 1987; Prohens & Nuez, 2001;

46 Pantoja, Pinto, Lopes, Gandra, & Santos, 2009; Acosta-Quezada et al., 2011; Correia & Canhoto,

47 2012). Increasing agricultural efforts have been done to improve the fruit quality due to its

48 increasing popularity (Mwithiga, Mukolwe, Shitanda, & Karanja, 2007, Correia & Canhoto,

49 2012; Abad, Valencia-Chamorro, Castro, &Vasco, 2017).

50 Tamarillo is also commonly known as tree tomato due to its similar shape to tomato (S.

51 lycopersicum). The fruit presents a smooth egg-shape and is two-end pointed. There is a diversity

52 in the size (4−10 cm long, 3–5 cm in diameter), skin color (e.g., yellow, orange, red, purple)

53 (dark longitudinal pigment stripes may be present or absent) and taste (acetous, sour, astringent

54 or sweet) of tamarillo (Morton, 1987). The presence of chlorophylls, carotenoids and

55 anthocyanins determine the color of the fruits (Acosta-Quezada et al., 2015). Purple or red

56 mucilage coats the seeds inside the fruit (Osorio et al., 2012). The diversity in tamarillo fruit

57 indicates that variation in chemical composition exits among different cultivars. Tamarillo fruit

58 is low in calorie and has minimal fat content. The fruit is a source of dietary fibers, minerals

59 (e.g., potassium, phosphorus, magnesium, calcium, copper, iron, and zinc), vitamins (e.g., B6, C

60 and E), proteins, soluble sugars (e.g., glucose, fructose, and sucrose), as well as organic acids

4
61 (e.g., malic and citric acids) (Bobbio, Bobbio, & Rodriguez-Amaya,1983; Hurtado, Morales,

62 González-Miret, Escudero-Gilete, & Heredia, 2009; Mertz et al., 2010; Osorio, Hurtado, Dawid,

63 Hofmann, Heredia-Mira, & Morales, 2012; Nascimento et al., 2013; 2015; Acosta-Quezada et

64 al., 2015; Espin, Gonzalez-Manzano, Taco, Poveda, Ayuda-Durán, Gonzalez-Paramas, &

65 Santos-Buelga, 2016; Giuffrida, Zoccali, Arigò, Cacciola, Roa, Dugo, & Mondello, 2018). A

66 variety of phytochemicals in tamarillo fruit with significant pharmacological and nutritional

67 properties include non-starch polysaccharides (e.g., pectins), flavonoids, carotenoids and so on

68 (Bobbio et al., 1983, Mertz et al., 2009, Osorio et al., 2012). Different parts of tamarillo tend to

69 have different compositions. For example, tamarillo seeds contain lipids (including ω-3, 6, and 9

70 fatty acids), proteins, vitamin E, polyphenols, minerals, and phytosterols (e.g., β-sitosterol,

71 cycloartenol and dihydrolanosterol) (Achicanoy, Benavides, & Mart´ınez-Correa, 2018)

72 Because of the diverse components, an array of in vitro and in vivo biological activities

73 were reported for tamarillo, such as antioxidative, antinociceptive, anti-inflammatory, anticancer,

74 and antiobesity. There was only one self-reported fruit-triggered allergy (Tiwari & Wolber,

75 2011). Because of the biological activities, as well as the “novel” eating experiences, diverse

76 tamarillo based products have been developed. They include salads, sauces, jellies, ice creams,

77 juices, liqueurs, yogurts and fruit effervescent tablets (Morton, 1987; Villegas-Ruíz, Rodríguez-

78 Armas, Guerrero-Beltrán, & Bárcenas-Pozos, 2013; Acosta-Quezada et al., 2015; Ramakrishnan,

79 Adzahan, Yusof, & Muhammad, 2018). Tamarillo-derived functional ingredients showed good

80 potential as a natural cheese-making coagulants in the dairy industry, antioxidant additives in

81 meat industry, and emulsifiers and foam stabilizers in a variety of other food sectors (Gannasin,

82 Ramakrishnan, Adzahan, & Muhammad, 2012; Castro-Vargas, Benelli, Ferreira, & Parada-

83 Alfonso, 2013; Li et al., 2018a; 2018b). In the non-food sectors, tamarillo-derived ingredients

5
 84 were used in the pharmaceutical industries as antinociceptive agents (Bohs, 1989; Hurtado et al.,

85 2009; Nascimento et al., 2013) and as photosensitizers for fabrication of natural dye-sensitized

86 solar cells in the energy industry (Susanti, Nafi, Purwaningsih, Fajarin, & Kusuma, 2014).

87 However, the information related to nutritional and biological properties as well as the

88 applications of tamarillo is rather scattered.

89 Previous reviews mainly focused on agriculture practices, genetic engineering and

90 biotechnology of tamarillo (Prohens & Nuez, 2001; Correia & Canhoto, 2012). With an aim to

91 support the current trend of tamarillo becoming a new “superfood” for consumers, this review

92 provides a basis for better understanding the nutritional and physiological effectiveness of

93 tamarillo fruit phytochemicals and to facilitate the development of potential food and non-food

94 applications. This review also updates formulations and developments in manufacturing

95 tamarillo based products.

96

97 2. Chemical properties of tamarillo

98 There is a great variation in chemical composition of tamarillo fruit (Table 1). This

99 variation could be due to the differences in maturity, fruits parts, genetics, geographic and

100 environmental conditions. The extraction methods and the analytical procedures used in different

101 studies may also contribute to this variation (Acosta-Quezada et al., 2015).

102 2.1. Proximate composition

103 Among the 23 cultivated tamarillo fruits, the moisture contents of fruit pulp (peeled,

104 seeds removed) ranged from 86.1 to 87.7% (fresh basis), while soluble solids, titratable acidity,

105 and protein content of fruits were 10.9 ̶ 12.1, 1.2 ̶ 1.8 and 4.4 ̶ 9.6 % (fresh basis), respectively

106 (Acosta-Quezada et al., 2015). Fruit puree (pulp, seed and mucilage) derived from golden-yellow

6
107 and purple-red tamarillo fruits (peeled, with seeds) from Ecuador and Spain contained 86 ̶ 92%

108 of moisture, 2.2 ̶ 2.6% of protein, 0.1 ̶ 0.7% of crude lipids and 0.7−0.9% of ash (Romero-

109 Rodriguez et al., 1994; Vasco et al., 2009). The moisture, crude protein, crude fat, carbohydrate,

110 fiber and ash of the edible portion of the fresh tamarillo fruits (pedicel removed) cultivated in

111 Taiwan were 85.7, 2.0, 0.4, 10.8, 2.8 and 1.0%, respectively (Kou et al., 2009). The moisture,

112 crude protein, carbohydrate, fiber and ash contents of the edible portion were 85.2, 1.6, 11.9, 6.0

113 and 1.3 % editable portion of Malaysian tamarillo fruits, respectively (Mutalib et al., 2017).

114 Ecuadorian tamarillo fruits (purple-red and golden-yellow) also had little lipids (0.6 ̶ 0.7%).

115 Tamarillo seeds are a source of lipids. Colombian tamarillo seeds contained 17.3 % dry basis

116 (maximum value, based on the optimum supercritical carbon dioxide extraction conditions)

117 (Achicanoy et al., 2018). A total of 2.8−6.0 % (fresh weight) was found inedible parts of the

118 tamarillo fruit (Kou et al., 2009; Mutalib et al., 2017).

119 2.2. Carbohydrates

120 2.2.1. Simple sugars

121 Dark-red whole fresh tamarillo fruits (with peel and seeds) cultivated in Colombia had a
◦
122 Brix value of 10.0 (Osorio et al., 2007). Golden-yellow and purple-red tamarillo fruits (peeled,

123 with seeds) from Ecuador and Spain showed ◦Brix values of 11−12 (Vasco et al., 2009). Twenty-

124 three tamarillo accessions (13 from Ecuador, 4 from Peru, 2 from Bolivia, 1 from Colombia, 1

125 Portugal origin and 2 New Zealand origins) were analyzed for sugar composition. They

126 represented cultivars of five categories, namely orange, orange pointed, red, red conical, and

127 purple (Acosta-Quezada, et al., 2015). The total soluble sugar contents in the pulp (peeled, seeds

128 removed) of these tamarillo fruits ranged from 28.1 to 52.0 g/100g dry basis. Sucrose (15.5 ̶ 27.1

129 g/100g dry basis) was the predominant sugar, followed by fructose (6.3 ̶ 13.2 g/100g dry basis)

7
130 and glucose (6.3 ̶ 11.0 g/100g dry basis) (Acosta-Quezada et al., 2015). Golden-yellow and

131 purple-red tamarillo fruits (peeled, with seeds) from Ecuador and Spain contained sucrose

132 (1.69−2.59% fresh weight), fructose (0.79−1.69%, fresh weight) and glucose (0.59−1.79% fresh

133 weight) (Vasco et al., 2009).

134 2.2.2. Non-starch polysaccharides

135 Non-starch polysaccharides were isolated from the pulp and mucilage of tamarillo fruit

136 using water and alkaline solutions, followed by fractionation (freeze-thaw and α-amylase

137 treatments) and purification (Fehling precipitation and ultrafiltration) (Nascimento et al., 2013,

138 2015, 2016a). The isolation conditions affected the specific monosaccharide compositions and

139 structure, and physicochemical properties of resulting polysaccharides (Nascimento et al., 2013;

140 2015, 2016a). Tamarillo mucilage refers to the locular tissue surrounding the seeds of the fruit

141 (Nascimento et al., 2016b). The mucilage contained a mixture of highly methoxylated

142 homogalacturonans and type I arabinogalactans (Nascimento et al., 2016b). A comparison study

143 reported that tamarillo mucilage polysaccharides structurally differed from its pulp

144 polysaccharides, in terms of the length of side chains of the pectin and the degree of branching of

145 the xylans. Tamarillo mucilage polysaccharides (highly methoxylated homogalacturonans and

146 type I arabinogalactans) had backbones of a linear (1→5)-linked α-L-arabinan and a linear (1→

147 4)-β-D-xylan. A linear (1→4)-β-D-xylan was only found in mucilage polysaccharides

148 (Nascimento et al., 2016b). Galactoarabino-glucuronoxylan polysaccharide isolated from freeze-

149 dried and defatted tamarillo fruit pulp had backbone and side-chains composed of (1→4)-linked

150 β-D-Xylp, (1→5)-linked α-L-Araf, and (1→4)-linked α-D-GlcpA residues, along with non-

151 reducing end units (α-L-Araf, β-Arap, β-D-Galp, α-D-GlcpA and 4-O-Me-α-D-GlcpA)

152 (Nascimento et al., 2013). High molecular weight hemicellulosic polysaccharides and pectin-

8
153 associated arabinogalactan-protein hydrocolloids were also obtained from the pulp and mucilage

154 of tamarillo fruit, respectively (Gannasin et al. 2015). In the study of Gannasin et al. (2015),

155 water and 1% citric acid were used to extract the seed mucilage hydrocolloids, while 72%

156 ethanol and 20 mM 4-(2-hydroxyethyl)-1-piperazineethanesulfonic acid (HEPES) buffer were

157 used for pulp hydrocolloid extraction. In the studies of Nascimento et al. (2013, 2016a and

158 2016b), pulp without seeds and mucilage was freeze-dried and defatted with chloroform-

159 methanol (1:1). Polysaccharides were obtained by hot-water extraction and ethanol precipitation.

160 In addition to different extraction and purification methods used, Gannasin et al. (2015) and

161 Nascimento et al. (2013) investigated the structure of polysaccharides obtained from red and

162 yellow tamarillo fruits, respectively. Factors explaining the structural differences in

163 polysaccharides in pulp and mucilage of tamarillo fruits deserve to be well researched using the

164 same experimental conditions.

165 2.3. Proteins

166 Protein content of cultivated tamarillo fruit pulp ranged from 4.4 ̶ 9.6% fresh basis

167 (Acosta-Quezada et al., 2015). The crude protein account for 2.2 ̶ 2.5% fresh puree of golden-

168 yellow and purple-red tamarillo fruits from Ecuador (Vasco et al., 2009) and Spain (Romero-

169 Rodriguez et al. 1994).Two interesting proteins isolated from the tamarillo fruit were protease

170 and lectin (i.e., CBL1, 25k) (Xu, Moore, Fountain, & Yu, 1992; Xu, Birtles, Fountain, & Yu,

171 1993; Li et al., 2018a). Tamarillo lectin CBL2 isolated from tamarillo fruit (cultivated in New

172 Zealand) remain to be purified (Xu et al., 1992). Amino acid composition and function of

173 tamarillo fruit proteins were not fully characterized.

174 2.3.1. Proteases

9
175 Tamarillin, an alkaline subtilisin-like serine protease, was derived from tamarillo fruit (Li

176 et al., 2018a, 2018b; 2018c; 2018d). The conditions that produced optimum activity of tamarillin

177 were pH 11 and 60 °C. Tamarillin activity was minimally influenced by

178 ethylenediaminetetraacetic acid (EDTA) (up to 5 mM), p-chloromercuribenzoate (PCMB), and

179 metal ions (Na+, Zn2+, Co2+, Ca2+, and Mn2+), but was inhibited by phenylmethylsulfonyl fluoride

180 (PMSF, a serine protease inhibitor) and Hg2+ (high affinity for anionic sulfur). In different

181 organic solvents (25%, v/v), the enzymatic activity of tamarillin was reduced by varying

182 amounts and solvent concentrations. The activity compared to the control without organic

183 solvent were ranked in the following order: ethanol (∼96%) > methanol (89%) > glycol (85%) >

184 dimethyl sulfoxide (DMSO) (∼77%) > isopropanol (∼76%) > chloroform (∼62%) (Li et al.,

185 2018a). These solvents interacted differently with tamarillin, affecting the hydrogen bonds, the

186 hydrophobic interactions and the charges on the enzyme (Li et al., 2018a). The molecular

187 mechanisms behind solvent-modified dynamics and conformation of tamarillin are still unclear.

188 2.3.2. Lectins

189 Lectin (CBL1 and CBL2 with molecular weights of 25 kDa and > 60 kDa, respectively)

190 fractions were isolated from tamarillo fruits (Xu et al., 1992; 1993). Both lectin fractions

191 demonstrated hemagglutination activities. CBL1 (not CBL 2) was able to produce a precipitin

192 line with anti-CBL1 serum. CBL2 (not CBL1) was able to produce a precipitin line with non-

193 immune rabbit serum (Xu et al., 1992; 1993). Future work will be required to fully characterize

194 the CBL2. The cell walls of tamarillo fruit tissues and the seed coat are the primary storage

195 location of lectin (namely CBL1) (Xu et al., 1992). CBL1 is a heat stable protein

196 comprised mainly of cysteine Cys), glycine (Gly), glutamine or glutamate (Glx), serine (Ser),

197 proline (Pro), aspartate or asparagine (Asx) and to a lesser extent hydroxyproline (Hyp) residues

10
198 (Xu et al., 1992). CBL1 contained 24% carbohydrate (Xu et al., 1992). Xu et al. (1993) reported

199 that CBL1 is a chitin-binding protein and was rich in Cys residue. Purified CBL1 showed no

200 chitinase activity and no mitogenic activity against human lymphocytes (Xu et al., 1992).

201 2.3.3. Allergenic proteins

202 Two IgE binding pathogenesis-related proteins (15 kDa) and subtilisin-like protease (26

203 kDa) were identified as potential allergens of tamarillo fruit (Tiwari & Wolber, 2011).

204 2.4. Lipids

205 Overall, the lipid content of tamarillo fruits is minimal. The crude lipids account for 0.1 ̶

206 0.7 % of puree derived from golden-yellow and purple-red tamarillo fruits from Ecuador (Vasco

207 et al., 2009) and Spain (Romero-Rodriguez et al. 1994). However, tamarillo seeds contained a

208 significant amount of lipids. The maximum extraction yield of tamarillo seed oil was 17.4 %

209 under optimized supercritical CO2 extraction conditions (38.1 MPa and 51°C). Soxhlet extraction

210 had a maximum oil yield of 24.1% (Achicanoy et al., 2018). The tamarillo seed oil was highly

211 unsaturated with only an average level of 13.8% of the fatty acids being saturated. Under

212 optimum supercritical carbon dioxide extraction conditions, the maximum amounts of

213 monounsaturated fatty acids (MUFA) and polyunsaturated fatty acids (PUFA) in seed oil were

214 18.2 and 74.0%, respectively (Achicanoy et al., 2018). Linoleic acid (72.1%) was the major

215 unsaturated fatty acid found in the seed oil, followed by oleic (17.9%) and linolenic (1.9 %)

216 acids. The linoleic acid level of tamarillo seed oil was comparable to that of Brazil passion fruit

217 seed oil (72.7%) and grape seed oil (70–78%) (de Oliveira, Rossi, Gimenes, Jagadevan,

218 Giufrida, & de Barros, 2013, Achicanoy et al., 2018). Palmitic and stearic acids accounted for

219 11.0% and 2.8% of seed oil, respectively. Palmitoleic acid (0.2%) in tamarillo seed oil was minor

220 (Achicanoy et al., 2018).

11
221 2.5. Minerals

222 Tamarillo fruit is a good source of potassium. Among 23 tamarillo accessions from five

223 cultivar groups of different the countries of origin (Ecuador, Peru, Bolivia, Colombia, Portugal

224 and New Zealand), tamarillo fruits contain potassium (the predominant mineral), phosphorus,

225 magnesium calcium, copper, iron, zinc, the quantities being 1868 ̶ 3211, 56 ̶ 226, 48 ̶ 230, 16 ̶ 80,

226 0.1 ̶ 3.3, 0.7 ̶ 2.4, 0.2 ̶ 1.4mg/100g, dry weight, respectively (Acosta-Quezada et al., 2015). The

227 potassium level was 410.6 mg of 100 g editable portion of the ripe tamarillo fruits collected from

228 Malaysia, followed by magnesium, sodium, calcium, and iron (quantities being 25.2, 17.8, 11.2

229 and 0.30 mg of 100 g editable portion, respectively (Mutalib et al., 2017). A total of 16.2 mg/100

230 g calcium of edible portion of fresh fruit cultivated in Taiwan was reported by Kou et al. (2009).

231 The contents of potassium, magnesium, sodium, calcium, iron, zinc and copper in golden-yellow

232 and purple-red tamarillo fruits (peeled, with seeds) from Ecuador and Spain had ranges of

233 347.0−440, 14.0−22.0, 0.1−8.9, 9.2−25.0, 0.2−0.6, 0.2−0.2, and 0.1−0.2 mg/100 g, fresh weight

234 (Vasco et al., 2009). Cultivar types, country of origin, geographic/climate conditions, and

235 growing practices have impacts on the variation in mineral levels of tamarillo fruits (Vasco et al.,

236 2009; Acosta-Quezada et al., 2015; Mutalib et al., 2017).

237 2.6. Vitamins

238 Tamarillo fruit is a good source of vitamin C and vitamin A (β-carotene) at a

239 concentration of 55.9 and 4.8 mg/100 g dry weight, respectively (Mutalib et al., 2017). Ascorbic

240 acid and dehydroascorbic acid were present (38.5 mg/100g) in the edible portion of tamarillo

241 fruits (Taiwan). The content of α and β-carotenes was found to be 181.8 mg/100g edible portion

242 of the fresh fruits (Taiwan) (Kou et al., 2009). Vitamin B1 (0.2 mg/100g) and vitamin B2 (0.1

243 mg/100g) were also present in the editable portion of tamarillo fruits (Kou et al., 2009). Vitamin

12
244 E (γ-tocopherol, 2.1 mg/mg) was the predominant tocopherol in tamarillo seed oils from the

245 optimized supercritical CO2 extraction (Achicanoy et al., 2018). Ascorbic acid of fruit nectar of

246 yellow tamarillo (Ecuador) was thermal-liable in the presence of low levels of dissolved oxygen.

247 Degassing fruit nectars reduced the susceptibility of vitamin C to thermal process (such as

248 pasteurization) (Mertz, Brat, Caris-Veyrat, & Gunata, 2010).

249 2.7. Organic acids

250 Organic acids (malic and citric acid), along with soluble sugars (glucose, fructose, and

251 sucrose) significantly contribute to characteristic taste of tamarillo (Boyes & Strubi, 1997; Vasco

252 et al., 2009). Differences among 23 tamarillo fruits in terms of total organic acids (4.5−8.1g/100g

253 dry basis) and sugar-to-acid ratio (4.4 ̶ 9.4) were significant (Acosta-Quezada et al., 2015). The

254 sugar-to-acid ratio of fruits is critical for consumer acceptance (Coombe, Dundon, & Short,

255 1980). Citric acid (4.0 ̶ 7.5g/100g dry basis) was predominant, followed by the malic acid

256 (0.3−1.0 g/100g dry basis) (Acosta-Quezada et al., 2015). These differences were possibly due to

257 the differences in cultivar types, country of origin, and cultivation conditions of the tamarillo

258 fruits (Acosta-Quezada et al., 2015). A New Zealand grown yellow tamarillo variety had higher

259 levels of organic acids than a red variety (Boyes & Strubi, 1997). In the study of Romero-

260 Rodriguez et al. (1994), the Spanish yellow varieties contained 1.8% citric acid and 0.1% malic

261 acid, while Spanish purple-red varieties contained 1.7% citric acid and 0.1% malic acid. The

262 citric acid content (2.7%) in Ecuadorian purple-red tamarillo fruits was comparable to that in

263 Ecuadorian golden-yellow variety (2.6%). However, an Ecuadorian tamarillo fruit (purple-red)

264 had a higher malic acid level (0.5%) than a golden-yellow Ecuadorian fruit (0.3%) (Vasco et al.,

265 2009). This partially explained the different acidic taste between these two varieties. Further

13
266 studies are needed to investigate the potential use of tamarillo organic acids as acidity regulators

267 or pH control agents for modifying or maintaining pH of food products.

268 2.8. Chlorophylls

269 Total chlorophyll contents of 23 tamarillo accessions from five cultivar groups (orange,

270 orange pointed, red, red conical and purple) ranged from 0.7 to 7.5 mg/100 g dry weight. The

271 chlorophyll a content (0.2−1.0 mg/100 g dry weight) was lower than the chlorophyll b content

272 (0.44−6.6 mg/100 g, dry weight) (Acosta-Quezada et al., 2015). Chlorophylls, along with

273 carotenoids and anthocyanins are responsible for the color of tamarillo fruits. Chlorophyll

274 reduces the chroma and luminosity of the fruits of the orange or red-fleshed cultivars. A higher

275 chlorophyll content contributed to a darker flesh colour in the purple-fleshed cultivars (Acosta-

276 Quezada et al., 2015).

277 2.9. Phenolic compounds

278 Total phenolics content (TPC) of 23 tamarillo accessions from five cultivar groups

279 (orange, orange pointed, red, red conical and purple cultivars) ranged from 2.4 ̶ 6.2 g/100 g dry

280 weight, measured by the Folin-Ciocalteu assay (Acosta-Quezada et al., 2015). Using high

281 performance liquid chromatography coupled with diode-array detection and mass spectrometry

282 (HPLC-DAD-MS), Espin et al. (2018) measured hydroxycinnamoyl derivatives (ranging from

283 60.3 ̶ 421.6 mg/100 g dry weight) of 4 cultivars, namely yellow giant cultivar A (Chaltura),

284 yellow giant cultivar B (Pelileo), giant purple cultivar and New Zealand purple cultivar.

285 According to Vasco et al. (2009), TPC in the peel and the seed-jelly of golden-yellow and

286 purple-red tamarillo fruits from Ecuador ranged from 387 to 620 and 94 to 152 mg gallic acid

287 equivalent/100 g fresh weight. The purple-red variety had a higher level of total soluble

288 phenolics than the golden-yellow variety (Vasco et al., 2009). TPC of extracts from the pulp

14
289 surrounding the seeds of the fruit was higher (by approximately 45%) than that of extracts from

290 peels of tamarillo fruits (cultivated in Colombia) (Hurtado et al., 2009). Refrigeration storage

291 (60-day) reduced the TPC from 1.2 to 0.8 mg gallic acid equivalents/g of a tamarillo fruit

292 product treated with sucrose and sodium benzoate (0.1%) (Villegas-Ruíz et al., 2013).

293 2.9.1. Phenolic acids

294 Hydroxycinnamic acid derivatives were the major types of phenolic acids in the aqueous

295 extracts of golden-yellow and purple-red tamarillo fruits at a concentration of 39 and 61 mg/100

296 g fresh weight, respectively (Vasco et al., 2009). The New Zealand purple tomarillo fruit is

297 high in hydroxycinnamates (421.6 mg/100 g dry pulp)(Espin et al., 2006).Tentatively identified

298 phenolic acids and derivatives in tamarillo included quinic acid, caffeoyl glucoside,

299 dehydrodiferulic acids, 3- and 5-O-caffeoylquinic acids, feruloyl glucoside, rosmarinic acid

300 glucosides, malonyl derivative of rosmarinic acid glucoside, and rosmarinic acid (Espin et al.,

301 2016).

302 2.9.2. Flavonoids

303 Total flavonoid content (TFC) was higher in the peel of tamarillo (3.36 mg rutin

304 equivalents/g, dry weight) than that in the flesh (peeled fruit cultivated in Malaysia) (2.41 mg

305 rutin equivalents/g, dry weight) (Hassan & Bakar, 2013). TFC of macerated fruit was higher than

306 that from other preparations (juice, decoction, and pomace) (Ordóñez et al., 2010). TFC of

307 tamarillo extracts from using various solvents ranged from 1.10 (water extract) to 3.10 (ethanol)

308 mg quercetin equivalents/g (dry weight) (Mutalib et al., 2017). Decreasing polarity of the

309 partition solvent increased the TFC of the samples in the order of ethanol > n-butanol > ethyl

310 acetate > water. Compared to water, ethanol was able to affect the polyphenol oxidases, leading

311 to cell wall degradation and the release of the intracellular components (Mutalib et al. 2017).

15
312 Quercetin (4−6 mg /100 g, fresh weight) and myricetin (1.2−1.4 mg/100 g, fresh weight) were

313 the representative flavonoids found in golden-yellow and the purple-red tamarillo (Vasco et al.,

314 2009).

315 2.9.2.1. Anthocyanins

316 Total anthocyanin content (TAC) was higher in the flesh of tamarillo fruit (4.15 mg/100

317 g, dry weight) than that in the peels (1.36 mg/100 g, dry weight) (Hassan & Bakar, 2013). TAC

318 account for 102.4 ̶ 168.9 mg/100 g of dry pulp from Tamarillo fruits (yellow giant, giant purple

319 and New Zealand purple cultivars) cultivated in Ecuador (Espin et al., 2016). TAC of seed jelly

320 and peelcrude extracts (20.03 mg delphinidin 3-glucoside/L sample) of fresh Colombian ripe

321 tamarillo fruits was higher than the peeling crude extracts 0.20 mg delphinidin 3-glucoside/L

322 sample) (Hurtado et al., 2009). In a pH range 2.0–6.2, both seed jelly and peel crude extracts

323 were shown to be relatively more stable under low acid conditions (pH 2.0–3.4). The seed jelly

324 extracts presented a higher sensitivity to pH changes than peeling extracts. This was due a higher

325 polymeric anthocyanins content and the presence phenolic acids, flavones, flavonols, flavanones,

326 and organic acids stabilising the colour of the peeling extracts (Eiro & Heinonen, 2002; Hurtado

327 et al., 2009). Polymeric pigment index of seed jelly and peel extracts was 0.20 and 1.16

328 (absorbance units) (Hurtado et al., 2009). TAC of macerated fruits was higher than other

329 preparations of fruits (juice, decoction, and pomace) (Ordóñez et al., 2010). Refrigeration storage

330 for 60 days reduced the total monomeric anthocyanins content from 0.91 to 0.61 mg cyanidin-3-

331 glucoside/g tamarillo fruit (Villegas-Ruíz et al., 2013).

332 Peonidin-3-O-glucoside was the most abundant anthocyanin in fresh tamarillo (Portugal)

333 (Gomes, Ghica, Rodrigues, de Souza Gil, & Oliveira-Brett, 2016). Cyanidin-3-rutinoside and

334 delphinidin-3-rutinoside were the most abundant anthocyanins in the peel and flesh surrounding

16
335 the seeds, respectively (Wrolstad & Heatherbell, 1974). Pelargonidin 3-O-rutinoside (77.0 ̶ 78.0

336 mg/100 g dry weight), delphinidin 3-O-rutinoside (21.8 ̶ 87.42 mg/100 g dry weight) and

337 cyanidin 3-O-rutinoside (2.5 ̶ 4.5 mg/100 g dry weight) were major anthocyanin pigments in

338 tamarillo fruit pulp (Espin et al., 2016). New Zealand purple tamarillo contained delphinidin 3-

339 O-glucosyl rutinoside, delphinidin 3-O-rutinoside, cyanidin 3-O-rutinoside and pelargonidin 3-

340 O-rutinoside. The content of anthocyanins in Ecuadorian yellow-purple tamarillo was

341 negligible (Espin et al., 2016). At various pH values (2.0−6.2), anthocyanins structures (i.e,

342 hydroxylation and glycosidation) influenced the colour and stability anthocyanin rutinosides

343 isolated from Colombian tamarillo fruits (et al., 2009). Isolated anthocyanin rutinosides included

344 delphinidin 3-O-(6''-O-α-rhamnopyranosyl-β-glucopyranosyl)-3'-O-β-glucopyranoside,

345 delphinidin 3-O-(6''-O-α-rhamnopyranosyl)-β-glucopyranoside, cyanidin 3-O-(6''-O-α-

346 rhamnopyranosyl)-β-glucopyranoside, and pelargonidin 3-O-(6''-O-α-rhamnopyranosyl)-β-

347 glucopyranoside (Hurtado et al., 2009).

348 Overall, the variations in anthocyanin composition depended on several factors, including

349 cultivation locations (e.g., New Zealand vs. Ecuador), fruit colors (e.g., yellow vs. purple), fruit

350 parts (e.g., flesh vs. peels), extraction and identification methods used, as well as storage

351 conditions (Wrolstad & Heatherbell, 1974; Bobbio, Bobbio, & Rodriguez-Amaya, 1983; De

352 Rosso & Mercadante, 2007; Mertz et al., 2009; Hurtado, Morales, González-Miret, Escudero-

353 Gilete, & Heredia, 2009; Mertz, Brat, Caris-Veyrat, & Gunata, 2010; Villegas-Ruíz et al., 2013;

354 Espin et al., 2016).

355 2.10. Carotenoids

356 In total, 8 free carotenoids and 15 carotenoids esters were identified in Colombian

357 yellow tamarillo using supercritical fluid extraction and supercritical fluid chromatography with

17
358 triple quadrupole mass spectrometry detection (SFE-SFC-QqQ/MS) with C30 column with sub-2

359 micron particles (Giuffrida et al., 2018). In a comparison study of Giuffrida et al. (2018), the

360 partially porous C30 column (UHPLCPDA/MS) provided a better compound resolution and

361 shorter analysis time than the conventional C30 column (HPLC-PDA-MS) in the identification

362 of the carotenoids. The free carotenoids included 13-Z-β-carotene, β-carotene, 9-Z-β-carotene, 3-

363 Z-β-cryptoxanthin, β-cryptoxanthin, 5,8-epoxy-β-cryptoxanthin, lutein, and zeaxanthin. β-

364 Cryptoxanthin-C12:0, zeaxanthin-C12:0, β-cryptoxanthin-C14:0, antheraxanthin-C12:0, β-

365 cryptoxanthin-C16:0, antheraxanthin-C14:0, and antheraxanthin-C16:0 were identified as

366 carotenoid monoesters. The carotenoid diesters included zeaxanthin-C12:0,C14:0, lutein-

367 C14:0,C14:0, zeaxanthin-C14:0,C14:0, lutein-C14:0,C16:0, zeaxanthin-C14:0,C16:0,

368 zeaxanthin-C12:0,C18:0, lutein-C16:0,C16:0, and zeaxanthin-C16:0,C16. SFE-SFC-QqQ/MS

369 analysis also rapidly detected 7 apocarotenoids (apo-8′-zeaxanthinal, apo-10′-zeaxanthinal, apo-

370 12′-zeaxanthinal, apo-14′-zeaxanthinal, apo-15′-zeaxanthinal, apo-12′-carotenal, apo-14′-

371 carotenal) and 2 apocarotenoids esters (apo-10′-zeaxanthinal-C4:0 and apo-8′-zeaxanthinal-

372 C12:0). Similar results were obtained by Bobbio, Bobbio, and Rodriguez-Amaya (1983) and

373 Mertz et al. (2010).

374 A total of 5.76 mg/100g carotenoids was extracted from tamarillo fruit cultivated in

375 China using optimized response surface methodology (RSM) (Yang & Zhao, 2013). Total

376 carotenoids of 23 cultivated tamarillo fruits account for 2.6 ̶ 11.2 mg/100 g of dry pulp (Acosta-

377 Quezada et al., 2015). Thermal and oxidative lability varied among the carotenoids of yellow

378 tamarillo fruit. Zeaxanthin esters were more susceptible to thermal treatment (Mertz et al., 2010).

379 Thermal degradation of tamarillo carotenoids included 5, 8-epoxidation and cis-isomerization

380 (Mertz et al., 2010). Thermal treatment in presence of low levels of dissolved oxygen minimally

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381 influenced the total carotenoid content of yellow tamarillo fruit nectar (Mertz et al., 2010).

382 Microencapsulation improved carotenoid retention (Garcia et al., 2018). Mango peel powder,

383 banana peel powder, freeze-dried powder from yellow tamarillo fruit pulp and maltodextrin were

384 used for microencapsulation which significantly improved the carotenoid content up to 1.3 mg β-

385 carotene/100 g sample (Garcia et al., 2018).

386 2.11. Squalene and phytosterols

387 Using supercritical CO2 extraction under optimized operating conditions, the squalene (a

388 precursor for phytosterol synthesis) in tamarillo fruit seeds was 19.8 mg/mL seed oil. β-sitosterol

389 (3.7 mg/mL seed oil) was the predominant phytosterol in the seed oil, followed by cycloartenol

390 (2.8mg/mL seed oil) and dihydrolanosterol (0.7 mg/mL seed oil) (Achicanoy et al., 2018).

391 2.12. Volatiles

392 According to high resolution gas chromatography coupled to high resolution mass

393 spectrometry (HRGC/HRMS) analysis, pentane-dichloromethane liquid-liquid extracts of red

394 tamarillo fruit mainly contained methyl hexanoate, (E)-hex-2-enal, (Z)-hex-3-en-1-ol, eugenol,

395 and 4-allyl-2,6-dimethoxyphenol as the volatiles (Torrado et al. 1995). Fruity notes of tamarillo

396 fruit were attributed to the volatiles including trans-3-hexenal, cis-3-hexenol, methyl hexanoate,

397 3-hydroxybutanoates and 3-hydroxyhexanoates. Eugenol and 4-allyl-2,6-dimethoxyphenol

398 contributed the spicy notes of the fruit (Torrado et al. 1995). Using headspace solid-phase

399 microextraction (HS-SPME) coupled with gas chromatography-mass spectrometry (GC-MS), α-

400 terpineol, methyl hexanoate, ethyl octanoate, ethyl hexanoate, and 1,8-cineole were identified as

401 the predominant volatile compounds in the golden-yellow fruits (Durant, Rodríguez, Santana,

402 Herrero, Rodríguez, & Gupta, 2013). A sensomics approach, which combines a gentle volatile

403 extraction (solvent-assisted flavor evaporation), gas chromatography-mass spectrometry, and

19
404 sensory analyses (gas chromatography-olfactometry and aroma extract dilution analysis) was

405 applied to identify odor-active volatile compounds of yellow tamarillo fruit (García et al., 2016).

406 Volatile compounds ((Z)-3-hexenal, hexanal, and ethyl butanoate)) in yellow tamarillo

407 contributed to its unique aroma. The herbal-green, fruity, and fresh-mint odor notes of the yellow

408 variety were due to the presence of C6-aliphatic compounds, aliphatic esters, and terpenols,

409 respectively (García, Prieto, Guevara, Malagon, & Osorio, 2016). Other odor-active volatile

410 compounds including (Z)-3-hexenol, ethyl 3-hydroxybutanoate, 2, 3-butanediol and terpinen-4-

411 ol contributed to the green, citric, rancid and cooked earthy notes of yellow tamarillo,

412 respectively (García et al., 2016). Tamarillo fruits are not commonly characterised as very

413 aromatic/fragrant fruits. Breeding programme and genetic means remain to be applied to

414 improve this aspect of the sensory quality of tamarillo fruits.

415 2.13. Alkaloids

416 Pyrrolidine, tropane, calystegins, solasodine and tomatidenol were the alkaloids in the

417 tamarillo plant (Eich, 2008). Alkaloids from tamarillo roots included solacaproine [NN-bis-(4-

418 dimethylaminobutyl) hexanamide], the N-hexanoyl derivative of solamine, solamine (principal

419 component), tropinone, and cuscohygrine, hyoscyamine (atropine), tropine, Φ-tropine, tigloidine

420 (1αH, 5αH-tropan-3β-yl 2-methylcrotonate), and a homologue of solacaproin (Lim, 2013). The

421 absence or a concentration of alkaloids below the detection limit were obtained in the peels and

422 pulp of the golden-yellow and the purple-red tamarillo fruits (Vasco et al., 2009).

423

424 3. Biological activities of tamarillo

425 3.1. Antioxidative properties

20
426 In vitro antioxidant activities of tamarillo fruit extracts were studied intensively using

427 chemistry-based and/or cell-based assays (Hurtado et al., 2009; Kou et al., 2009; Espin et al.,

428 2016; Mutalib et al., 2017) (Table 2). Results of these in vitro screening assays may not fully

429 explain the biological antioxidant activities of tamarillo fruits (Granato et al., 2018).

430 3.1.1. Antioxidative properties based on chemical assays

431 The chemical assays were based on ferric reducing antioxidant power (FRAP), 2,2′-

432 azinobis(3-ethylbenzothiazoline-6-sulfonate) (ABTS), oxygen radical absorbance capacity

433 (ORAC), β-carotene bleaching (BCB), 2,2-diphenyl-1-picrylhydrazyl (DPPH), Trolox equivalent

434 antioxidant capacity (TEAC), copper-induced low-density lipoprotein (LDL) oxidation, and

435 nitrite scavenging properties (Hurtado et al., 2009; Kou, Yen, Hong, Wang, Lin, & Wu, 2009;

436 Ordonnez, Cardozo, Zampini, & Isla, 2010; Acosta-Quezada et al., 2015; Espin et al., 2016;

437 Mutalib et al., 2017). All the studies showed that tamarillo had broad a spectrum of antioxidant

438 potentials. For example, the DPPH based antioxidant activities of 23 tamarillo varieties ranged

439 from 12.2 ̶ 48.7 µmol Trolox/g fresh weight (Acosta-Quezada et al., 2015). Phenolics (such as

440 hydroxycinnamoyl acid and rosmarinic acid and delphinidin 3-rutinoside), chlorophylls,

441 carotenoids, ascorbic acid, and tocopherol were largely antioxidative in tamarillo (Acosta-

442 Quezada et al., 2015; Espin et al., 2016). Results from different studies are difficult to compare

443 due to the methodological variations and the differences in the means of expressing results. For

444 example, the DPPH value were presented as Trolox equivalents (TE) by Acosta-Quezada et al.

445 (2015), while Mutalib et al. (2017) reported DPPH values using IC50 (concentration of the

446 samples needed to decrease the DPPH free radical concentration by 50%).

447 Overall, comparison of results from the same study is meaningful. A comparative study

448 showed different antioxidant capacities in 4 tamarillo cultivars, including yellow giant A and B,

21
449 giant purple and New Zealand purple (Espin et al., 2016). The New Zealand purple cultivar had

450 higher antioxidant values based on the FRAP, ABTS, and ORAC assays (50, 89, 352 µmol

451 Trolox/g, dry weight, respectively) than the other three cultivars (Espin et al., 2016). The golden-

452 yellow tamarillo variety had lower DPPH radical scavenging activity than the purple-red variety

453 (Vasco et al., 2009). It might be due to the fact that the total phenolics in the golden-yellow

454 variety (125 mg gallic acid equivalents/100 g fresh weight) was lower than that of purple-red

455 variety (187 mg gallic acid equivalents/100 g fresh weight) (Vasco et al., 2009). The antioxidant

456 activities of tamarillo products were compared with some common compounds (Hurtado et al.,

457 2009; Mutalib et al., 2017). The TEAC value of tamarillo fruit peel extracts (1.09 mmol

458 Trolox/g) was comparable to that of ascorbic acid (1.09 mmol Trolox/mmol), but lower than that

459 of the extracts from the fruit pulp (1.90 mmol Trolox/g) (Hurtado et al., 2009). Various solvent

460 extracts of tamarillo fruits had a lower antioxidant activity (determined by BCB assay) compared

461 to butylated hydroxyl toluene (BHT). The BCB values ranked as follows: BHT (95.6 %) > crude

462 ethanol extract (79.3 %) > n-butanol fraction (68.6%) > ethyl acetate fraction (57.5%) > water

463 fraction (55.2%) (Mutalib et al., 2017).

464 Compared to ascorbic acid (1.09 mmol of Trolox/mmol), higher TEAC was reported for

465 the anthocyanins isolated from tamarillo fruits, including delphinidin 3-O-(6''-O-α-

466 rhamnopyranosyl-β-glucopyranosyl)-3'-O-β-glucopyranoside (2.20 mmol of Trolox/mmol ),

467 delphinidin 3-O-(6''-O-α-rhamnopyranosyl)-β-glucopyranoside (4.91 mmol of Trolox/mmol),

468 cyanidin 3-O-(6''-O-α-rhamnopyranosyl)-β-glucopyranoside (1.80 mmol of Trolox/mmol), and

469 pelargonidin 3-O-(6''-O-α-rhamnopyranosyl)-β-glucopyranosid (1.48 mmol of Trolox/mmol)

470 (Hurtado et al., 2009). The antioxidant capacity of the above-purified anthocyanins was also

22
471 dependent on the degree of hydroxylation and glycosylation pattern of the rutinosides (Rice-

472 Evans, Miller, & Paganga, 1996; Hurtado et al., 2009).

473 3.1.2. Antioxidative properties based on cell based assays

474 The different cell models used to study the antioxidant properties of tamarillo products

475 were based on neuronal PC12 and hepatocellular carcinoma HepG2 cells (Hurtado et al., 2009;

476 Ordonnez et al., 2010). Pre-treatment of PC12 cells with an ethyl acetate fraction of tamarillo

477 fruit (25, 50, 100 and 200 µg/mL) showed concentration-dependent inhibition of the hydrogen

478 peroxide (H2O2) induced cytotoxicity which resulted in an increasing range of cell viability

479 (66−100%) (Kou et al., 2009). Kou et al., (2009) suggested that the tamarillo phenolics protected

480 the PC12 cells via its antioxidant activity, in particular its free radical scavenging activity. Pre-

481 treatments of HepG2 cells with different fruit preparations (maceration, decoction juice and

482 pomace) at a dose range of 25−100 µg/mL dose-dependently inhibited the H2O2 induced HepG2

483 cell death (Ordóñez et al., 2010). Further studies are needed to identify the phenolic compounds

484 in tamarillo that are responsible for the obtained antioxidant activity.

485 3.1.3. Effect of processing on antioxidative properties

486 The antioxidant properties of tamarillo based products were significantly affected by the

487 processing methods used such as the fractionation methods (Ordóñez et al., 2010; Kou et al.,

488 2009; Mutalib et al., 2017). Tamarillo fruit maceration with higher TFC and TAC had higher

489 antioxidant potential than the other preparations (juice, decoction, and pomace) as evaluated by

490 β-carotene bleaching, DPPH, ABTS and nitrite scavenging assays (Ordóñez et al., 2010).

491 Antioxidant activities of the soluble fractions of different preparations of the fruit (juice,

492 decoction, and maceration) were higher than that of the insoluble fractions (pomace) (Ordóñez et

493 al., 2010). The DPPH scavenging activities of tamarillo extracts had the following ranking due to

23
494 different solvents used: n-butanol fraction (IC50: 0.70 mg/mL) > crude ethanol extract (IC50: 0.80

495 mg/mL) > ethyl acetate fraction (IC50: 1.40 mg/mL) > water fraction (IC50: 1.75 mg/mL)

496 (Mutalib et al., 2017). Kou et al. (2009) found that the scavenging activities against DPPH and

497 ABTS radicals of crude extract and different fractions of tamarillo fruit were ranked in the

498 following order: water fraction (13.5 mg catechin equivalents (CE)/g dry weight) < n-butanol

499 fraction (23.1 mg CE/g dry weight) < crude ethanol extract (28.8 mg CE/g dry weight) < ethyl

500 acetate fraction (61.1 mg CE/g dry weight). Inhibition of LDL oxidation by the ethyl acetate

501 fraction (IC50: 7.66 µg/mL) of the fruit was stronger than DL-α-tocopherol (IC50:9.27 µg/mL),

502 while the n-butanol fraction (IC50:15.02 µg/mL fractions of fruits had a weaker inhibition of

503 LDL oxidation than DL-α-tocopherol (Kou et al., 2009). Overall, suitable processing methods

504 should be used to maximize the antioxidant potential of tamarillo-based products.

505 3.2. Antinociceptive and antiinflammatory properties

506 Anti-nociceptive effects of the polysaccharide fractions of tamarillo fruit pulp were

507 evaluated using two inflammatory pain models (female Swiss mice) based on acetic acid and

508 formalin (Nascimento et al., 2013, 2015) (Table 2). The acetic acid test model is a screening

509 model for assessing the analgesic or anti-inflammatory properties. Acetic acid physiologically

510 stimulates the abdominal constriction. Formalin physiologically stimulates nociception

511 responses. The formalin test model is an acute and tonic pain model for accessing inflammatory

512 and chronic pain states. Examining the differences in the formalin-induced nociceptive

513 behaviours may provide insights into the mechanisms underlying the nociceptive responses that

514 occur in different phases of the formalin test [e.g., neurogenic pain (phase I) and inflammatory

515 (phase II)] (Shibata, Ohkubo, Takahashi, & Inoki, 1989). The former (neurogenic pain) is due to

516 direct chemical stimulation of the nociceptors. The latter phase is a result of inflammation of the

24
517 peripheral tissues. Intraperitoneal administration of a fraction (STK-1000R) (0.01, 0.1, and 1

518 mg/kg) dose-dependently inhibited the acetic acid (0.6%)-induced abdominal constrictions [50%

519 inhibitory dose (ID50): 0.47 mg/kg] and inhibited the formalin (2.5%)-induced inflammation

520 (phase II) (ID50 0.09 mg/kg), with a minimal influence on formalin-induced neurogenic pain

521 (phase I) (Nascimento et al., 2013). In another study, intraperitoneal administration of a

522 polysaccharide fraction (50E) (0.1–10 mg/kg) dose-dependently inhibited the acetic acid (0.6%)-

523 induced abdominal constrictions (ID50 = 0.91 and maximum inhibition of 92% with 100 mg/kg

524 50E), while minimally influencing the formalin (2.5%)-induced inflammation (phase II), with a

525 minimal influence on the formalin-induced neurogenic pain (phase I) (Nascimento et al., 2015).

526 The observed in vivo anti-inflammatory and anti-nociceptive activities evaluated on female

527 Swiss mice were attributed to the galactoarabinoglucuronoxylans (in fraction STK-1000R) and

528 type I arabinogalactans (in fraction 50E) (Nascimento et al., 2013; 2015). Compared to the

529 fraction 50E, STK-1000R had better inhibition of the abdominal constrictions induced by acetic

530 acid in the animal trials (Nascimento et al., 2013; 2015). This difference was attributed to the

531 structural difference between type I arabinogalactans and galactoarabinoglucuronoxylans

532 (Nascimento et al., 2013; 2015). Further studies are necessary to evaluate the molecular

533 interactions between tamarillo bioactive compounds and possible biomarkers that involved in in

534 vivo nociception. Metabolomic approaches to uncover the tamarillo-derived metabolites

535 implicated in the antinociceptive responses in humans are expected.

536 3.3. Anticancer properties

537 Tamarillo fruit extracts were selectively cytotoxic to liver hepatocellular carcinoma

538 (HepG2) and non-hormone dependent breast carcinoma (MDA-MB-231), while being non-toxic

539 to normal mouse fibroblast cells (3T3) (Mutalib et al., 2017) (Table 2). There was variation in

25
540 the cytotoxicity against various cancer cell lines among a crude tamarillo ethanol extract and

541 three other fractions (ethyl acetate, n-butanol, and water fractions). For HepG2 cells, the water

542 fraction had the highest half-maximal inhibitory concentration (IC50:110 µg/mL) value, followed

543 by the ethyl acetate fraction (IC50: 44 µg/mL), the n-butanol fraction (IC50: 26 µg/mL) and the

544 crude ethanol extract (IC50: 10 µg/mL). For MDA-MB-231 cells, the water fraction had the

545 highest IC50 (130 µg/mL) followed by n-butanol fraction (IC50: 96 µg/mL), ethyl acetate fraction

546 (IC50: 82 µg/mL) and crude ethanol extract (IC50: 80 µg/mL) (Mutalib et al., 2017). Mechanisms

547 of action for the selective actions of the tamarillo fruit extracts against the cancer cells remain

548 unknown. Tamarillo phenolic acids and flavonoids could inhibit the growth of certain cancer

549 cells with the cell cycle arresting followed by apoptosis (Roleira, Tavares-Da-Silva, Varela,

550 Costa, Silva, Garrido, & Borges, 2015). In addition to the pro-apoptotic activities, flavonoids

551 could affect different signaling pathways and target DNA topoisomerases of cancer cells (Raffa,

552 Maggio, Raimondi, Plescia, & Daidone, 2017).

553 3.4. Anti-obesity

554 Rats with obesity induced by a high fat diet (Male Sprague Dawley rats, 200−250 g)

555 received a daily dose of tamarillo fruit extracts at three different levels (150, 200, 300 mg/ kg)

556 via oral gavage for 7 weeks (Abdul-Kadir, Rahmat, & Jaafar, 2015) (Table 2). Administering

557 extracts (200 or 300 mg/ kg) to the rats resulted in weight loss, reduced total cholesterol level

558 and increased antioxidant activities of superoxide dismutase and glutathione peroxidase. Anti-

559 obese effect of tamarillo fruits was possibly associated with anti-inflammatory and/or

560 antioxidative effects (Abdul-Kadir et al., 2015). The connections between anti-obesity and the

561 tentative bioactive compounds, such as carotenoids, flavonoids, phytoestrogens, and dietary

562 fibre) were not clear for tamarillo (Abdul-Kadir et al., 2015). Molecular mechanisms responsible

26
563 for the anti-obese property of tamarillo fruits were unclear, though previous in vitro and in vivo

564 studies revealed an array of mechanisms (Trigueros, Peña, Ugidos, Sayas-Barberá, Pérez-

565 Álvarez, & Sendra, 2017). For example, dietary anthocyanins could regulate of the adipocyte

566 function, which was associated with adipocytochine expression and insulin sensitivity. Dietary

567 fibers could limit the energy intake and increase the secretion of anorexigenic peptides. Dietary

568 calcium could influence the fecal fat excretion by the formation of insoluble calcium-fatty acid

569 soaps and/or binding of bile acids, regulate adipocyte lipid metabolism and triacylglycerol

570 storage, and modify adipocyte-uncoupling protein 2 (UCP2) expression associated with calcitriol

571 in modulating adipocyte apoptosis (Trigueros, Peña, Ugidos, Sayas-Barberá, Pérez-Álvarez,

572 Sendra, 2017).

573 3.5. Toxicity

574 Studies on tamarillo toxicity are rare. One self-reported allergy case showed that

575 consuming tamarillo fruit triggered hives within 24 h. IgE binding proteins (15 kDa and 26 kDa)

576 possibly represented the allergenic proteins of tamarrilo fruit (Tiwari & Wolber, 2011). For

577 toxicity level on normal cell lines, tamarillo fruit extracts were largely non-cytotoxic to normal

578 cells (3T3) with an IC50 of > 200.00 µg/mL (Mutalib et al., 2017). In an animal trial, daily

579 treatment of male Sprague Dawley rats with tamarillo fruit extract at a dose of 300 mg/ kg for 7

580 weeks showed non-toxicity (Abdul-Kadir et al., 2015). Therefore, it seems that tamarillo fruit is

581 largely safe to consume, though possible allergy reactions should be better studied.

582

583 4. Comparative studies

584 Comparative studies determine the similarities and differences among the different

585 tamarillo varieties, and between tamarillo fruits and other commonly consumed or reported fruits

27
586 and vegetables. The selected fruits and vegetables may have at least one aspect, which overlaps

587 with tamarillo fruits, such as fruit morphological characters, country of origin, and plant families

588 or species. The identified differences indicate the unique value of tamarillo. While the identified

589 similarities provide a conception of the possible relations among the comparative subjects. This

590 in turn tends to support tomatillo derived product developments based on current applications of

591 those compared fruits and vegetables.

592 4.1. Comparisons of different tamarillo varieties

593 According to Pantoja et al. (2009), there was not much of a difference in proximate

594 composition (except ash and reducing sugar contents) between the Brazilian yellow and the red

595 tamarillo varieties. Approximately 85% more ash was found in the red variety, whereas 45%

596 more reduce sugars was found in the yellow variety (Pantoja et al., 2009). Vasco et al. (2009)

597 found Ecuadorian golden-yellow and purple-red varieties had the same ash content. Only the

598 sucrose content of the golden-yellow variety was 10% higher than that of the purple-red variety.

599 The golden-yellow variety contained a higher level of magnesium, but lower levels of calcium,

600 iron, and sodium than the purple-red varieties (Vasco et al., 2009). Total soluble solids (Brix)

601 and titratable acidity, pH values and vitamin C were similar between golden-yellow and purple-

602 red varieties (Vasco et al., 2009). Compared to purple-red tamarillo varieties, golden-yellow

603 variety had less β-carotene and total soluble phenolic compounds (Vasco et al., 2009). In

604 particular, the golden-yellow variety did not contain anthocyanins (Vasco et al., 2009).

605 Gannasin et al. (2015) also pointed out that only red tamarillo mesocarp contained (polar)

606 anthocyanin-rich dark red seed mucilage and (non-polar) carotenoid-rich orangish pulp. Espin et

607 al. (2016) found that only purple cultivars (not yellow cultivars) contained the anthocyanins.

608 Different varieties exhibited different chemical profiles, which may result in different

28
609 bioactivities and functionalities. In terms of the in vitro antioxidant capacity, greater antioxidant

610 values of DPPH and ORAC assays in purple-red varieties were reported by Vasco et al. (2009)

611 and Mertz et al. (2009), respectively, compared to yellow varieties. Acosta-Quezada et al. (2015)

612 found similar DPPH values between orange and red tamarillo varieties. Espin et al. (2016)

613 reported New Zealand purple variety had higher FRAP, ABTS and ORAC values than two

614 yellow varieties. ORAC and FRAP values of giant purple verity were lower than those of yellow

615 giant varieties. More comparison studies on different tamarillo varieties are necessary in order to

616 increase their utilization efficiency.

617 4.2 Comparison of between tamarillo and other fruits and vegetables

618 4.2.1. Comparison in chemical composition

619 The majority of research only focused on the chemical composition of tamarillo fruit as

620 shown in the section 3. There has been little comparison between tamarillo and other plant

621 sources, especially those in the Solanaceae family such as tomato (Solanum lycopersicum).

622 Tomato as a popular vegetable has a range of health-promoting phytochemicals (e.g.,

623 carotenoids, vitamins C, pro-vitamin A, folic acid) (Dzakovich et al., 2019). All-trans-lycopene

624 (1000 µg/g dry weight), cis-lycopene (130 µg/g dry weight) and all-trans-β-carotene (50 µg/g dry

625 weight) are the major carotenoids in tomato (Gama, Tadiotti, & Sylos, 2009; Dzakovich et al.,

626 2019). In tamarillo, all-trans-β-cryptoxanthin esters and all-trans-β-carotene are the principle

627 carotenoids (Mertz et al., 2010). Properties of tamarillo lectin were compared with that of other

628 plants of the Solanaceae. The molecular mass of purified tamarillo lectin CBL1 (25kDa) was

629 smaller than that of tomato lectin (a glycoprotein of 71 kDa), thorn-apple (Datura stramonium)

630 lectin (disulfide bond linked non-identical subunits of 40 and 46 kDa), and potato lectin (a non-

631 convalently linked dimeric glycoprotein of 100 kDa). Purified CBL1 had an N-terminal sequence

29
632 of Thr-Ser-Glu-Leu-Pro-Gln, while tomato lectin contained methionine as its N-terminal amino

633 acid. The carbohydrate content of CBL1 (24%) was less than that of lectins from potato, tomato

634 and thorn-apple (arabinose and galactose). Lectins from potato, tomato and thorn-apple were rich

635 in hydroxyproline residues, while CBL1 was rich in Cys residues (Allen & Neuberger, 1973;

636 Nachbar, Oppenheim, & Thomas, 1980; Crowley & Goldstein, 1981; Xu et al., 1992; 1993).

637 Some chemical properties of tamarillo were compared to some other fruits. Compared to

638 Andes berry (Rubus glaucus) (moisture content, 94.1%; ◦Brix 9.8), tamarillo fruit had a lower

639 moisture content (89.3%) and a higher soluble solids content (◦Brix 10). The level and

640 composition of anthocyanins differed between tamarillo fruit and other plants such as Andes

641 berry (Osorio, Franco, Castaño, Gonzalez-Miret, Heredia, & Morales, 2007; Garzón, Riedl, &

642 Schwartz, 2009; Osorio et al., 2012; Espin et al., 2016). Tamarillo fruit had a higher anthocyanin

643 content than Andes berry (Garzón, Riedl, & Schwartz, 2009; Espin et al., 2016; Osorio et al.,

644 2007). The anthocyanin-rich extracts of Andes berry mainly contained cyanidin-3-O-α-L-

645 rhamnopyranosyl-(1 →6)-O-β-D-xylopyranosyl-(1→2)-β-D-glucopyranoside (Osorio et al.,

646 2012). This is different from those of tamarillo as described in section 2.9.2.1. The potassium

647 content (410 mg/100g dry weight) of tamarillo fruit was comparable to that of banana pulp

648 (295−463 mg/100 g fresh basis, one of the good sources of potassium) (Sulaiman et al., 2011).

649 The level of polyunsaturated fatty acids (72%) in tamarillo seed oil extracted using supercritical

650 CO2 was comparable to that in passion fruit seed oil (73%) and grape seed oil (70–78%)

651 (Achicanoy et al., 2018). The total sterol concentration of tamarillo seed oil (3.4 − 6.8 mg/mL)

652 was comparable to that of cottonseed, sesame seed, and rapeseed oils (Achicanoy et al., 2018).

653 Therefore, tamarillo fruit was seen to be nutritionally complementary to some other fruits and

654 vegetables.

30
655 4.2.2. Comparison in biological properties

656 The in vitro antioxidant activity of tamarillo fruit was compared with other fruits.

657 Antioxidant activity (0.12 mmol Trolox/kg fruit) of tamarillo (dark-red) fruit was lower than that

658 of Andes berry (1.08 mmol Trolox/kg fruit) (Osorio et al., 2007). Blackberry had a higher

659 antioxidant capacity than tamarillo based on the DPPH assay (Gomes et al., 2017).

660 Electrochemical quantitative index (EI) was used to reflect the antioxidant capacity of different

661 fresh fruits including blackberry, blueberry, physalis, raspberry and tamarillo. Tamarillo (EI,

662 0.035) had a lower antioxidant capacity than blackberry (EI, 0.738), blueberry (EI, 0.474), and

663 raspberry (EI, 0.474) and had a higher capacity than physalis (EI, 0.003) (Gomes et al., 2017).

664 The antioxidant activities of tamarillo extracts (crude extracts, ethanol, n-butanol, ethyl acetate

665 and water fractions) were lower than butylated hydroxytoluene (BHT) (Gomes et al., 2006).

666 Therefore, it seemed that tamarillo was not particular strong in terms of in vitro antioxidant

667 activities.

668 Comparative studies in biological properties between tamarillo and other commonly used

669 food plants of the nightshade family such as tomato are rare. Like tomato, some tamarillo

670 varieties also contained a significant amount of carotenoids. Previous studies showed that

671 carotenoids have some health-promoting properties (Giovannucci, 2005; Mehta, Patani, &

672 Singhvi, 2018). However, it is difficult to compare biological properties between tamarillo and

673 other common plant foods from different studies due to the differences in the experimental

674 design and assay conditions. Future studies should focus on comparisons that show how effective

675 tamarillo is at providing health-promoting benefits.

676 5. Food and non-food uses of tamarillo

677 5.1. Food uses

31
678 Fresh tamarillo fruit is the most commonly used form of the fruit. The fresh fruit is a

679 dietary ingredient in juice, salad, and desserts (Espin et al., 2016). The maceration, decoction,

680 juice and pomace of tamarillo have pleasant flavors (Ordóñez et al., 2010).

681 There is great interest in developing functional ingredients from tamarillo. Based on

682 laboratory observations, crude protease extracts from tamarillo fruit may be used as a natural

683 rennet-like clotting agent in reconstituted skim milk gels. The optimized proteolytic activities

684 were at pH of 5.5−6.5 (Li, Scott, Otter, Zhou, & Hemar, 2018b). Compared to rennet, tamarillo

685 extracts had a broad specific activity in a broad pH range of 5.5−11 on bovine caseins (Li et al.,

686 2018b). Tamarillin, a novel serine protease, is the protease in the extracts. Comparative studies

687 showed that milk gels induced by tamarillin had lower elasticity, higher porosity, less brittle

688 behaviour than rennet-induced gels (Li et al., 2018d). Both rennet (10 and 20%, w/w milk solids

689 content), and tamarillin induced gels had similar morphologies of milk gel networks and large-

690 scale structures (Li et al., 2018d). The application of tamarillin in the commercial cheese is worth

691 further study.

692 Many techno-functional properties such as water-holding and oil-holding capacities,

693 foaming capacity, foam stability, emulsifying activity and emulsion stability are possessed by

694 hydrocolloids. The pulp hydrocolloids from red tamarillo possessed considerable oil-holding

695 (3.3–3.6 g oil/g dry sample) and water holding (25–27 g water/g dry sample) capacities

696 (Gannasin et al., 2016). Preliminary emulsifying tests on the pulp hydrocolloids suggested their

697 potential applications as food emulsifiers. Tamarillo fruit extracts are promising foam stabilisers

698 that can be used in foam based food products (e.g., mousses, marshmallows and meringues)

699 (Gannasin et al., 2016). High-methoxyl-pectin polysaccharides were isolated from tamarillo fruit

700 pulp. The dispersions of the pectins displayed liquid-like, shear-thinning behaviors with low

32
701 apparent viscosity (Nascimento et al., 2016a). Adding modifiers such as sucrose changed the

702 rheological viscoelastic properties of the dispersions (Nascimento et al., 2016a), extending the

703 applications of tamarillo polysaccharides as a potential gelation agent.

704 The antioxidants in tamarillo fruits such as anthocyanins could function as natural food

705 additives that prolong the shelf life of foods by preventing or delaying lipid oxidation (Hurtado

706 et al., 2009; Castro-Vargas et al., 2013). Tamarillo peel extracts minimized the lipid oxidation in

707 cooked beef during its storage at 4 oC (Castro-Vargas et al., 2013). In addition to being a source

708 of food antioxidants, other representative tamarillo-derived products are still under development

709 at the laboratory stage. They included the carotenoids-rich microencapsulates derived from the

710 freeze-dried pulp of yellow tamarillo and colour-and flavour-enriched syrups (osmotic

711 solutions) derived from dark-red tamarillo fruits by osmotic dehydration (Osorio et al., 2007;

712 García et al., 2018).

713 5.2. Other uses

714 Non-food uses of tamarillo are limited. For nutraceutical and pharmaceutical uses,

715 tamarillo antioxidants such as anthocyanin rutinosides could be used as dietary supplements to

716 improve the nutritive value and quality of foods (Hurtado et al., 2009). The fruit pomace could

717 also be a source of antioxidant rich ingredients for the formulations of nutraceuticals or

718 functional foods (Ordóñez et al., 2010). The effectiveness of tamarillo phenolics as

719 chemotherapeutic and anti-inflammatory pain relief agents were reported using in vitro and in

720 vivo models (Nascimento et al., 2013; Abdul-Kadir, Rahmat, & Jaafar, 2015, Mutalib et al.,

721 2017). Clinical trials to confirm these claims remain to be done. The development of tamarillo

722 bioactive derived medicines for pharmaceutical applications may be facilitated through better

723 identification, characterization and extraction of the health promoting compounds.

33
724 For the energy industry, the efficacy of tamarillo fruit extracts as photosensitive dye for

725 dye-sensitized solar cells was reported (Susanti, Nafi, Purwaningsih, Fajarin, & Kusuma, 2014).

726 The photosensitive dye captures the incoming photons of light and uses the energy to excite

727 electrons. The solar energy industry recognizes the benefits of using natural dyes (e.g., minimal

728 environmental impact, non-toxic and eco-friendly) over synthetic dyes. TiO2-based and tamarillo

729 fruit extract derived dye-sensitized solar cells have been fabricated by Susanti et al. (2014). The

730 sensitized solar cells had a maximum efficiency of 0.043%, a circuit voltage of 542.5 mV and

731 short circuited current density of 0.356 mA/cm2 (Susanti et al., 2014). Tamarillo pigments, such

732 as anthocyanins, carotenoids and chlorophylls as photosensitizers, contributed to the solar energy

733 harvesting by converting it to electrical energy with the aid of a semiconducting

734 photoanode. Efficiency comparisons are needed between the tamarillo fruit extract sensitized

735 solar cells with those sensitized with other fruit extracts. Overall, the continued growth of non-

736 food uses of tamarillo products will require diversification and more innovative ideas.

737 5. Conclusions and outlooks

738 Tamarillo is an underutilized species, even though it has therapeutic and nutritional

739 values that can satisfy the demands from health-conscious consumers. Tamarillo fruit is low in

740 lipid content and is a source of proteins, carbohydrates and total dietary fibers, vitamins C, A,

741 B6, and E, minerals including potassium, magnesium and iron, organic acids, volatile aroma

742 compounds, polyphenols (e.g., phenolic acids and anthocyanins) and carotenoids (e.g., β-

743 cryptoxanthin). Tamarillo fruit based products such as extracts showed a range of biological

744 activities including antioxidation, antiinflammation, anticancer, antinociception, antiobesity, and

745 allergenicity. These claimed effects could be due to the components such as polyphenols and

746 their combinations with other bioactive compounds in the fruit. Comparative analysis showed

34
747 that tamarillo fruit can be nutritionally complementary to some other fruits and vegetables such

748 as tomato. Tamarillo fruit has been processed to make a range of ingredients including tamarillin

749 as a natural rennet-like clotting agent, hydrocolloids as emulsifiers and stabilizers, and

750 antioxidants (natural food preservatives). Tamarillo derived ingredients were also used in other

751 applications such as in pharmaceutical and energy industries. It can be concluded that tamarillo

752 is a promising “new” commercial fruit crop with potential to be fully exploited as a source of

753 “healthy” products.

754 For a better understanding and exploitation of tamarillo as a source of food and non-food

755 ingredients/products, the following aspects could be explored in future studies: (1) assessing the

756 genetic diversity of tamarillo in chemical composition; (2) purification and characterisation of

757 components for specific properties, such as texture, taste or specific health claims; (3)

758 optimization of the extraction process for tamarillo-derived nutrients and functional ingredients;

759 (4) selection of appropriate evaluation models to reveal the effective physiological

760 concentrations of bioactive compounds in tamarillo and their combinations, and related

761 mechanisms; (5) metabolomics approaches for determining dietary tamarillo bioactive-derived

762 metabolites implicated in in vivo biological responses; (6) allergenicity and long-term toxicity

763 evaluation; (7) diversification and innovation of food and non-food products; (8) studies of

764 composition, extraction, functional properties, bioactivity, applications of functional ingredients

765 derived from unutilized parts of tamarillo plant such as leaves and stems; (9) improvement of the

766 aroma of tamarillo fruits through targeting volatile compounds and genetic means; (10) functions

767 of non-volatile taste-active components in tamarillo.

768

769 Declaration of conflict of interest

35
770 None.

771 Funding source

772 This research didn’t receive any specific grant.

773 Acknowledgements

774 Mr. Cai Min kindly provided the photo of tamarillo fruits grown in China.

775

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942 relationship of flavonoids and phenolic acids. Free radical Biology and Medicine,

943 20, 933–956.

944 Romero-Rodriguez, M.A., Vazquez-Oderiz, M.L., Lopez-Hernandez, J., & Simal-Lozano, J.

945 (1994). Composition of babaco, feijoa, passion fruit and tamarillo produced in Galicia (North-

946 west Spain). Food Chemistry, 49, 23–27.

947 Shibata, M., Ohkubo, T., Takahashi, H., & Inoki, R. (1989). Modified formalin test:

948 characteristic biphasic pain response. Pain, 38, 347–352.

949 Sulaiman, S. F., Yusoff, N. A. M., Eldeen, I.M., Seow, E. M., Sajak, A. A. B., Supriatno, K. L.,

950 & Ooi, K. L. (2011). Correlation between total phenolic and mineral contents with antioxidant

951 activity of eight Malaysian bananas (Musa sp.). Journal of Food Composition and Analysis, 24,

952 1–10.

953 Susanti, D., Nafi, M., Purwaningsih, H., Fajarin, R., & Kusuma, G. E. (2014). The preparation of

954 dye densitized dolar cell (DSSC) from TiO2 and tamarillo extract. Procedia Chemistry, 9, 3–10.

955 Tiwari, R., & Wolber, F.M. (2011). Identification of Tamarillo (Cyphomandra betacea)

956 allergens. The Journal of Allergy and Clinical Immunology, 127, AB110–AB110.

957 Torrado, A., Suarez, M., Duque, C., Krajewski, D., Neugebauer, W., & Schreier P. (1995).

958 Volatile constituents from Tamarillo (Cyphomandra betacea Sendtn.) fruit. Flavour and

959 Fragrance Journal, 10, 349–354.

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960 Trigueros, L., Peña, S., Ugidos, A.V., Sayas-Barberá, E., Pérez-Álvarez, J.A., & Sendra, E.

961 (2013). Food ingredients as anti-obesity agents: A review. Critical Reviews in Food Science and

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45
Table 1 Chemical composition of tamarillo fruits
Chemical composition Concentration
Proximate composition Pulp (peeled, seeds Peel Puree (pulp + Seed Seed-Jelly Jelly
removed) seed and mucilage)

Moisture (% fresh basis) 86.1 ̶ 87.71 86 ̶ 926

Crude protein (% fresh basis) 4.4 ̶ 9.61 2.2 ̶ 2.56
Crude lipids (% dry basis) 0.1 ̶ 0.76 17.43
6
Ash (% dry basis) 0.7 ̶ 0.9
Total dietary fiber (% dry weight, puree hydrocolloid) 60.3-69.69
Soluble solids content (% fresh basis) 10.9 ̶ 12.11
Titratable acidity (% fresh basis) 1.2 ̶ 1.81

Fatty acid composition

Fatty acids (% seed oil)
Total saturated fatty acids 13.83
Palmitic acid (C16:0) 11.03
Stearic acid (C18:0) 2.83
Total monounsaturated fatty acids 18.23
Palmitoleic (C16:1) 0.33
Oleic (C18:1) 17.93
Total polyunsaturated fatty acids 74.03
Linoleic acid (C18:2) 72.13
Linolenic acid (C18:3) 1.93

Total organic acids and soluble sugars

Total organic acids (g/100 g dry basis ) 4.5 ̶ 8.11
Malic acid 0.3 ̶ 1.01
Citric acid 4.0 ̶ 7.51
Total soluble sugars (g/100 g dry basis ) 28.1 ̶ 52.01
Glucose 6.3 ̶ 11.01
Fructose 6.3 ̶ 13.21
Sucrose 15.5 ̶ 27.11
Ratios of soluble sugars:organic acids 4.4 ̶ 9.41

1
Minor dietary constituents Pulp (peeled, seeds Peel Puree (pulp + Seed Seed-Jelly Jelly
removed) seed and mucilage)

Minerals (mg/100g dry basis)

Potassium (K) 1868 ̶ 32111
Phosphorus (P) 56 ̶ 2261
Magnesium (Mg) 48 ̶ 2301
Calcium (Ca) 16 ̶ 801
Copper (Cu) 0.1 ̶ 3.31
Iron (Fe) 0.7 ̶ 2.41
Zinc (Zn) 0.2 ̶ 1.41
Vitamin (mg/100 g dry basis )
Vitamin A (β-carotene) 4.88
Vitamin C (ascorbic acid) 55.98
Vitamin B1 0.27
Vitamin B2 0.17
Vitamin E (γ ̵ tocopherol) (mg/mL seed oil) 2.13
1
Total Chlorophylls (mg/100g dry weight) 0.7 ̶ 7.5
Chlorophyll a 0.2 ̶ 1.01
Chlorophyll b 0.4 ̶ 6.61
Squalene (mg/mL seed oil) 19.83
Total sterol (mg/mL seed oil) 7.23
β-Sitosterol (mg/mL seed oil) 3.73
Cycloartenol (mg/mL seed oil) 2.83
Dihydrolanosterol (mg/mL seed oil) 0.73

Carotenoids
Total carotenoids (mg/100 g dry mass) 2.6 ̶ 11.21
Free carotenoids
Lutein 0.025
Zeaxanthin 0.15
5,8-Epoxy-β-cryptoxanthin 0.015
13-Z-β-cryptoxanthin 0.025
β-Cryptoxanthin 0.55

2
 13-Z-β-carotene 0.15
β-carotene 2.55
9-Z-β-carotene 0.085
Carotenoid monoesters
β-Cryptoxanthin-C12:0 0.25
β-Cryptoxanthin-C14:0 3.85
β-cryptoxanthin-C16:0 3.15
Carotenoid diesters (mg/100 g)
Lutein-C14:0, C14:0 0.085
Zeaxanthin-C14:0, C14:0 0.15
Lutein-C14:0, C16:0 0.15
Zeaxanthin-C14:0, C16:0 0.085
Lutein-C16:0, C16:0 0.045
Zeaxanthin-C16:0, C16:0 0.15

Pulp (peeled, seeds Peel Puree (pulp + Seed Seed-Jelly Jelly

removed) seed and mucilage)
Phenolic compounds
Total phenolics (g/100 g dry weight1, or mg gallic acid 2.4 ̶ 6.21 387–6206 94–1526
equivalent/100 g fresh weight 6)
Phenolic acids and their derivatives
Caffeoyl glucoside 1.4 ̶ 29.32
3-O-Caffeoylquinic acid 25.0 ̶ 163.62
Dehydrodiferulic acid (I) 0.1 ̶ 21.12
Dehydrodiferulic acid (II) 7.5 ̶ 22.12
5-O-Caffeoylquinic acid 0.5 ̶ 2.62
Feruloyl glucoside 0.2 ̶ 3.02
Rosmarinic acid glucoside (isomer I) 3.3 ̶ 14.92
Rosmarinic acid glucoside (isomer II) 2.0 ̶ 14.62
Rosmarinic acid glucoside (isomer III) 3.6 ̶ 19.72
Malonyl derivative of rosmarinic acid glucoside 1.6 ̶ 6.32
Rosmarinic acid glucoside (isomer IV) 0.9 ̶ 5.72
Rosmarinic acid 12.2 ̶ 121.92
Total hydroxycinnamoyl acids 60.3 ̶ 421.62

3
Total anthocyanins (mg/100 g dry weight) 102.4 ̶ 168.92
Delphinidin 3-O-rutinoside (Dp-3-rut) 21.8 ̶ 87.42
Cyanidin 3-O-rutinoside (Cy-3-rut) 2.5 ̶ 4.52
Pelargonidin 3-O-rutinoside 77.0 ̶ 78.02
Anthocyanin rich extracts (ARE)
Delphinidin-3-O-α-L-(-(1→6)-β-D- 138.04
glucopyranoside-3'-O-β-D-glucopyranoside
(mg Dp-3 rut/100 g ARE)
Pelargonidin-p-coumaroyl-rutinoside 32.04
(mg Dp-3 rut/100 g ARE)
Delphinidin-3-O-(6''-O-α-rhamnopyranosyl)-β- 4934.04
glucopyranoside (mg Dp-3 rut/100 g ARE)
Cyanidin-3-O-(6''-O-a-rhamnopyranosyl)-β- 438.04
glucopyranoside (mg Dp-3 rut/100 g ARE)
Pelargonidin-3-O-(6''-O-α-rhamnopyranosyl)-β- 2276.04
glucopyranoside (mg Dp-3 rut/100 g ARE)

Notes: 1 23 tamarillo accessions (five cultivar groups (orange, orange pointed, red, red conical and purple cultivars) from Ecuador
(Acosta-Quezada et al., 2015); 2 tamarillo fruits (yellow giant, giant purple and New Zealand purple cultivars) from Ecuador (Espin et
al., 2016); 3 Colombian tamarillo seed oil, maximum value of based on optimum supercritical carbon dioxide extraction conditions,
unit of crude oil is % (w/w), dry weight (Achicanoy et al., 2018); 4 Colombian ripe tamarillo fruits (red variety) (Osorio et al. 2012); 5
Colombian fresh yellow tamarillo fruits (lyophilized samples). Free carotenoid, carotenoid monoesters and diesters quantitated by a
coupling of supercritical fluid extraction and supercritical fluid chromatography with triple quadrupole mass spectrometry (SFE-SFC-
QqQ/MS) containing C30 column with sub-2 micron particles (Giuffrida et al., 2018); 6 Golden-yellow and purple-red tamarillo fruits
from Ecuador (Vasco et al., 2009) and Spain (Romero-Rodriguez et al., 1994); 7 Taiwan ripe tamarillo fruits (fruit color was not
specified) (Kou et al., 2009); 8 Malaysian ripe tamarillo fruits (fruit color was not specified) (Mutalib et al., 2017); 9 Fresh red
tamarillo fruit from Malaysia.The fruit puree hydrocolloids obtained from the citric acid, water, HEPES buffer and ethanol
extractions, respectively (Gannasin et al., 2015).

4
5
Table 2 Representative bioactivities of tamarillo
Sample information In vitro/in vivo model systems Observations Tentative key References
(country origin) bioactives

Anti-oxidative
Extracts from the pulp surrounding Assay In vitro TEAC values of pulp was higher Anthocyanin Hurtado et al.
the seeds of fruit and peels. Model TEAC than that of the peel extract and rutinosides and (2009)
Anthocyanin rutinosides purified Doses Not applicable ascorbic acid. other polyphenols
from fresh tamarillo ripe fruits Admi. Not applicable Purified delphinidin 3-O-(6''-O-α-
(fruit color was not specified) Time Not applicable rhamno-pyranosyl)-β-gluco-
(Colombia) pyranoside had the highest TEAC
value among purified samples.
Hydroxylation degree of the
isolated rutinosides determined
TEAC value.

Lyophilized fruits of 23 tamarillo Assay In vitro The DPPH values varied among Chlorophylls, Acosta-Quezada et
accessions from 5 cultivar groups: Model DPPH the 23 tamarillo with a range of polyphenols, and al. (2015)
orange, orange pointed, red, red 12.2 ̶ 48.7 mol TE/g. carotenoids
Doses Not applicable
conical and purple cultivars (13 Wide variation occurred within
from Ecuador, 4 Peru, 2 Bolivia, 1 Admi. Not applicable cultivar groups, but not among
Colombia, 1 Portugal and 2 New Time Not applicable cultivars.
Zealand)

Purple fruits (Giant purple and Assay In vitro New Zealand purple cultivar had Phenolics Espin et al. (2016)
New Model FRAP, ABTS and the highest FRAP, ABTS and (rosmarinic acid
Zealand purple cultivars) and ORAC ORAC values than the other and delphinidin 3-
yellow fruits (Chaltura giant A Doses Not applicable cultivars. rutinoside), and
and Pelileo giant B cultivars) Admi. Not applicable non-phenolics
(Ecuador) Time Not applicable (carotenoids
ascorbic acid
tocopherol)

6
Crude ethanol extract of fresh fruit Assay In vitro Ethyl acetate fraction presented Phenolics Kou et al. (2009)
and its partitioned fractions (ethyl Model DPPH, TEAC the highest DPPH scavenging
acetate, butanol and water Copper-induced LDL activity, TEAC, and copper-
fractions; fruit color was not oxidation, Neuronal induced LDL oxidation.
specified) PC12 cells Pre-treatment of PC12 cells with
(Taiwan) Doses Model dependent ethyl acetate fraction (25, 50, 100
Admi. Model dependent and 200 µg/mL) concentration-
Time Model dependent dependently inhibited hydrogen
peroxide (H2O2) induced cell
death.

Fruit preparations (maceration, Assay In vitro Antioxidant activities of soluble Phenolics, Ordonez et al.
decoction, juice, and pomace; Model β-Carotene bleaching, fruit juice, decoction, and flavonoids (2010)
The fruit pulp is red-orange) nitrite scavenging. maceration were higher than
(Argentina) ABTS, DPPH, insoluble Pomace.
HepG2 cells Fruit maceration, decoction juice
Doses Model depended and pomace (25−100 µg/mL)
Admi. Model depended dose-dependently inhibited H2O2
Time Model depended induced HepG2 cell death.

Ripe tamarillo fruits (Crude Assay In vitro Crude ethanol extracts had the Phenolics, Mutalib et al.
ethanol extract, ethyl acetate, n- Model β-carotene bleaching, highest antioxidant activity vitamins (2017)
butanol, and DPPH assayed by the β-carotene
water fractions; fruit color was not Doses Not applicable bleaching assay.
specified) Admi. Not applicable
(Malaysia) Time Not applicable n-Butanol had the highest DPPH
scavenging activity.

Anti-nociceptive
STK-1000R (Pulp hemicellulose Assay Acetic acid and STK-1000R at 10 mg/kg Hemicellulose Nascimento
polysaccharides from ripe tamarillo formalin tests maximally inhibited the number polysaccharides, et al. (2013)
fruits, orange type) Model Female Swiss mice of writhes (78%) in mice induced alactoarabino-
(Brazil) Doses 0.01−10 mg/kg by 0.6% acetic acid. glucuronoxylan

7
 Admi. Intraperitoneal STK-1000R at 1 mg/kg had the
Time 30 min maximally inhibited (80%)
nociception in mice induced by
2.5% formalin. Antinociceptive
effects via antiinflammatory
mechanisms

50E (pulp pectic polysaccharides Assay Acetic acid and 50E fraction at 100 mg/kg Pectic poly- Nascimento
from ripe tamarillo fruits, orange formalin tests maximally reduced the number of saccharides et al. (2015)
type) Model Female Swiss mice abdominal constrictions by 92% type I arabino-
(Brazil) Doses 1–100 mg/kg in mice treated by 0.6% acetic galactan
Admi. Intraperitoneal acid.
Time 30 min Antinociceptive effects of 50E
could be due to its anti-
inflammatory property.
Mechanism(s) underlying this
effect is unknown.

Anticancer
Ripe tamarillo fruit (crude ethanol Assay MTT assay Water fraction exhibited the Not reported Mutalib et al.
extract, ethyl acetate, Model HepG2, MDA-MB- highest selective cytotoxic to (2017)
n-butanol, and water fractions; fruit 231, 3T3 cancerous cells (HepG2, MDA-
color was not specified) Doses 3.125–200 µg/mL MB-23), while minimal
(Malaysia) Admi. Add to cell culture influencing proliferations of
Time 24 h normal cell (3T3).

Antiobesity
Freshly harvested fruits (fruit color Assay In vivo Obese rats fed with extracts (200 Carotenoids, Abdul Kadir et al.
was not specified) Model Sprague-Dawley rats and 300 mg/ kg) had decreased flavonoids, (2015)
(Malaysia) (male) levels of total cholesterols, blood phytoestrogens,
Doses 150, 200, 300 glucose, triglycerides, and LDL- dietary fibers
mg/kg, rat body C, TNF-α and IL-6 activities, and
weight increased HDL-C, SOD and GPx

8
 Admi. Oral gavages activities, and TAS.
Time 7 weeks
3T3, normal mouse fibroblast; Admin., administration mean; DPPH, 2,2-diphenyl-1-picrylhydrazyl; EC50, half maximal effective
concentration; FRAP, ferric-reducing antioxidant power; GPx: glutathione peroxidase; HepG2, hepatocellular carcinoma; HDL, high-
density lipoprotein; HDL-C, high-density lipoprotein–cholesterol; LDL, low-density lipoprotein; LDL-C, low density lipoprotein –
cholesterol; MDA-MB-23, breast adeno-carcinoma; MTT: 3-(4,5-dimethylthiazol-2yl)-2,5-diphenyltetrazolium bromide ; ORAC,
oxygen radical absorbance capacity; SOD, superoxide dismutase; TAS, total antioxidant status; TEAC, Trolox equivalent antioxidant
capacity.

9
Figure 1 (A) Representative tamarillo fruits from different locations (Osorio et al., 2012; Nascimento et al., 2013); (B) the most
dominant chemical structures of tamarillo anthocyanins (Osorio et al., 2012); Reprinted with permission from Elsevier.

(A)

Red tamarillo fruits Orange-yellow tamarillo fruits Orange-yellow tamarillo fruits

(Colombia) (Brazil) (China)
Osorio et al. (2012) Nascimento et al. (2013)

(B)

R1 R2 R3 R4
OGlc H OH H Delphinidin-3-O-α-L-rhamnopyranosyl-
(Glc: (1 6)-β-D-glucopyranoside-3'-O-β-D-
glucose) glucopyranoside
OH H OH H Cyanidin-3-O-(6''-O-α-
rhamnopyranosyl)-β-glucopyranoside
OH H H H Cyanidin-3-O-(6''-O-α-
rhamnopyranosyl)-β-glucopyranoside
H H H H pelargonidin-3-O-(6''-O-α-
rhamnopyranosyl)-β-glucopyranoside

1
Highlights

 Tamarillo is a source of fiber, vitamin C, potassium, polyphenols and carotenoids.

 Tamarillo has high polyphenol content compared to other tropical fruits.

 Tamarillo has claimed health effects such as antioxidation and antinociception.

 Tamarillo provides innovative ingredients for food, beverage, pharmaceutical, and energy

industries.