Plant Foods for Human Nutrition

https://doi.org/10.1007/s11130-019-00744-8

ORIGINAL PAPER

Chemical Composition and Antioxidant Activity of the Main Fruits,

Tubers and Legumes Traditionally Consumed in the Andean Regions
of Ecuador as a Source of Health-Promoting Compounds
Doménica Pérez-Balladares 1 & Michael Castañeda-Terán 1 & María G. Granda-Albuja 2 & Eduardo Tejera 3 &
Gabriel Iturralde 2 & Silvana Granda-Albuja 4 & Tatiana Jaramillo-Vivanco 5,6 & Francesca Giampieri 7,8 &
Maurizio Battino 7,8 & José M. Alvarez-Suarez 1

# Springer Science+Business Media, LLC, part of Springer Nature 2019

Abstract
Thirteen fruits, eight legumes and three tubers consumed in the Andean regions of Ecuador were studied to determine their bioactive
compounds, organic acids, sugars content, total antioxidant capacity, as well as to determine which among them showed the greatest
contribution in terms of antioxidant activity and which compounds contributed to it. Among fruits, taxo fruits (Passiflora mollissima
(Kunth) L.H. Bailey) presented the highest values of total phenolic, carotene content, and total antioxidant capacity. The ají ratón
(Capsicum chinense Jacq) showed the highest content of vitamin C. Taxo showed the highest content of β carotene, whilst lycopene
was identified only in guayaba fruits (Psidium guajava L.) and ají ratón was the principal source of lutein. In legumes, chocho perla
(Lupinus mutabilis Sweet) showed the highest values for both total phenolic and flavonoid content, whilst frejol negro (Phaseolus
vulgaris L.) and frejol canario (Vigna unguiculata (L.) Walp) showed the highest values for FRAP and DPPH assay, respectively.
Between tubers, the jícama (Smallanthus sonchifolius (Poepp.) H. Rob.) had the majors values in terms of total phenolic, flavonoid
content, and total antioxidant capacity. In terms of total antioxidant capacity, taxo fruits have the highest contribution in terms of
total antioxidant capacity, whilst the dendogram shown the occurrence of five distinct groups in which taxo was located in the first
largest group. Our data contributing towards gaining better knowledge about the Andean Ecuadorian diet and the composition of
Andean food in order to estimate dietary intakes of health-promoting components.

Keywords Andean fruits . Andean legumes . Andean tubers . Ecuador . Antioxidant capacity . Bioactive compounds

Doménica Pérez-Balladares and Michael Castañeda-Terán contributed

equally to this work.
Electronic supplementary material The online version of this article
(https://doi.org/10.1007/s11130-019-00744-8) contains supplementary
material, which is available to authorized users.

* Maurizio Battino 4
Departamento de Biotecnología, Universidad de Las Fuerzas
m.a.battino@univpm.it Armadas, Sangolquí, Ecuador
* José M. Alvarez-Suarez
5
jose.alvarez@udla.edu.ec Jardín Botánico de Quito, Quito, Ecuador

1 6
Facultad de Ingeniería y Ciencias Aplicadas, Grupo de Investigación Grupo de Investigación en Biotecnología Aplicada a Biomedicina
en Biotecnología Aplicada a Biomedicina (BIOMED), Universidad (BIOMED), Universidad de Las Americas, Quito, Ecuador
de Las Américas, Quito, Ecuador
2 7
Laboratorios de Investigación, Universidad de Las Américas, Nutrition and Food Science Group, Department of Analytical and
Quito, Ecuador Food Chemistry, CITACA, CACTI, University of Vigo, Vigo, Spain
3
Facultad de Ingeniería y Ciencias Aplicadas, Grupo de
8
Bio-Químioinformática (CBQ), Universidad de Las Américas, Department of Clinical Sciences, Università Politecnica delle
Quito, Ecuador Marche, Ancona, Italy

Abbreviations
TPC Total phenolic content
TFC Total flavonoid content
ACY Total anthocyanin content
FW Fresh weight
TAC Total antioxidant capacity
FRAP Ferric reducing antioxidant power
DPPH 2,2-diphenyl-1-picrylhydrazyl
TCC Total carotenoid content
Introduction
Since ancient times, several types of plants and their products
have played an important role in the development of Andean
cultures, including being used as food and therapeutic agents.
One third of all Ecuadorian plant species has been used by the
country’s different cultures [1]. Currently, the use of plants
and plant products for nutritional purposes continues to be a
distinctive characteristic of the traditional Andean culture that
is practiced daily in many indigenous communities (30% of
the entire population belongs to different indigenous groups)
and urban populations of all social classes in Ecuador [2].
Recently, the fruits consumed by the Andean populations
of South America have been the subject of growing interest,
especially due to their potential health benefits and consumer
interest in new exotic fruits. However, while the properties
related to the health and chemical composition of fruits com-
monly found in North America and Europe have been studied
extensively, in the case of Andean fruits, as in Ecuador, they
are still widely unknown. Ecuador is divided into three conti-
nental regions, which are well defined by their climatic con-
ditions and typical vegetation, namely the coast (the lowlands
regions between the Pacific Ocean and the Andes), the moun-
tains (the Andean region) and the Amazon (east of the Andes
mountain range). This climactic diversity allows for an exten-
sive year-round variety of native and exotic fruit species of
potential interest as sources of bioactive compounds with rel-
evant effects on human nutrition and health.
The interandean region or Sierra is a geographical region of
Ecuador that extends centrally from north to south through the
Andes. It range from 1800 to 6310 m above sea level, where
the inhabited areas extend approximately up to 3600 masl. It is
formed by the provinces of Pichincha, Carchi, Tungurahua,
Chimborazo, Cañar, Azuay, Loja, Imbabura, Bolívar, and
Cotopaxi. It is the home of one of the largest indigenous pop-
ulations in the country, who have maintained their traditions,
such as their nutrition, which is based on a high consumption
of autochthonous fruits of the region, tubers, and legumes.
Some studies have highlighted the composition of some major
fruits consumed in Ecuador [3–5], as well as the possible
therapeutic potential of certain berries in the region [6].
However, despite this, studies related to the chemical
Plant Foods Hum Nutr
composition of the principal fruits, tubers, and legumes con-
sumed by these populations are still limited. The aim of this
study was to determine the principal bioactive compounds and
antioxidant capacity of the main fruits, tubers, and legumes
consumed in the Andes regions of Ecuador to elucidate their
potential as natural sources of bioactive compounds.
Materials and Methods
Samples A total of thirteen fruits, eight legumes, and three
tubers were collected during 2018 during their corresponding
crop season from popular markets in the Andean regions of
Ecuador (in the provinces of Chimborazo, Cotopaxi,
Tungurahua, Pichincha, and Imbabura). (Supplementary
Material 1).
Determination of Total Phenolic Content (TPC), Total
Flavonoid Content (TFC) and Total Anthocyanin Content
(ACY) A hydroalcoholic extract was obtained according to a
previously described method [7] and was used for the TPC,
TFC, and ACY contents and total antioxidant capacity deter-
mination. TPC was determined using the Folin-Ciocalteu col-
orimetric method [8], TFC was determined using the alumi-
num chloride method [9], whilst total ACY content was de-
termined using a modified pH differential method [10].
Results were expressed as milligrams of gallic acid equiva-
lents (GAE) per 100 g of fresh weight (FW) of fruits for TPC,
milligrams of catechin equivalents (Cateq) per gram of FW
for TFC, and milligrams of Pg-3-gluc equivalents (PgEq) per
gram of FW for ACY.
HPLC-DAD Analysis For the HPLC-DAD analysis of vitamin
C, carotenoid, and organic acids content the HPLC system
(Agilent Technologies 1260, Santa Clara, California, USA)
consisted of a Quaternary Pump Agilent Technologies
Infinity G1312B and a Diode Array Detector Agilent
Technologies Infinity G1315C.
Vitamin C Analysis Vitamin C was determined as previously
reported [6]. An Eclipse Plus C18 column (5 μm, 4.6 ×
150 mm) was used as stationary phase and the elution was
performed with 50 mM of KH2PO4 (pH 2.5) in an isocratic
gradient at a flow rate of 1 mL/min for 20 min at 245 nm. Total
vitamin C content was expressed as mg of ascorbic acid per
100 g of FW (mg Vit C/100 g FW).
Carotenoid Analysis Carotenoid content was determined as
previously reported [6]. The stationary phase consisted of an
Eclipse Plus C18 (5 μm, 4.6 × 250 mm) and elution was per-
formed with methanol-isopropanol (35:65, v/v) in an isocratic
gradient at a flow rate of 1 mL/min for 15 min at 450 nm. β-
carotene, lutein, and lycopene contents were quantified using
Plant Foods Hum Nutr
a calibration curve of the corresponding external standard and
the results were expressed as μg per 100 g FW.
Organic Acids Analysis Organic acids content was determined
according the method previously reported [11]. A Zorbax SB-
Aq column (3.5 μm, 4.6 × 50 mm) was used as stationary
phase and elution was performed with 20 mM of NaH2PO4
(pH 2.6)/Acetonitrile (99.5:0.5, v/v) in an isocratic gradient at
a flow rate of 0.8 mL/min for 35 min at 210 nm. An external
standard of lactic, acetic, citric and, oxalic acid were used for
identification and quantification, using the respective calibra-
tion curve and the results were expressed as g/100 g of FW.
HPLC-RI Analysis of Glucose and Fructose Content Glucose
and fructose content was determined according to a previously
reported method [12]. The UHPLC system (Dionex Ultimate
3000 Software Chromeleon) coupled with the refractive index
detector was equipped with a quaternary pump (Dionex
Ultimate 3000 UHPLC Focussed), degasser and ERC RI
520 refractive index detector (RefractoMax, 521, USA) and
the chromatographic data were acquired using the
Chromeleon software. The stationary phase consisted of a
Zorbax NH2 (250 × 4.6 mm, 5 μm) and elution was per-
formed with H2SO4 0.01 N in an isocratic gradient at a flow
rate of 1 mL/min for 55 min. An external standard of glucose
and fructose were used for identification and quantification
using the respective calibration curve and the results were
expressed as g/100 g FW.
Determination of Total Antioxidant Capacity (TAC) TAC of
hydroalcoholic extracts was determined using the Ferric
Reducing Antioxidant Power (FRAP) assay [13] and the
2,2-diphenyl-1-picrylhydrazyl free radical method (DPPH)
[14] and the results were expressed as μmol of Trolox equiv-
alents (TEq) per gram of FW (μmol TEq per g FW) for both
assays.
Statistical Analysis Statistical analyses were performed using
IBM SPSS Statistics for Windows version 20.0. A multivari-
ate analysis (MANOVA) was done for chemical composition
and TAC. The bonferroni correction was used in both cases
for post-hoc analysis adjusting for multiples groups’ compar-
ison. The correlations between the variables were calculated
using Pearson’s coefficient. In all cases a P value <0.05 was
considered as statistically significant.
Results and Discussion
Total Phenolic, Flavonoid Content, Anthocyanin Content and
Vitamin C Content TPC varied greatly (Table 1), ranging in the
different fruits between 118.84 mg of GAE/100 g FW
(guayaba) and 6839.64 mg of GAE/100 g FW (taxo).
According to their TPC, samples were classified into three
categories as follows: low (<100 mg of GAE/100 g FW),
medium (100–500 mg of GAE/100 g FW), and high
(>500 mg of GAE/100 g FW) [3]. Following this classi-
fication, none of the fruits had low values, while 38.46%
were in the medium category and 61.54% in the high
category (Fig. S1A). TFC ranged between 1.14 (babaco)
to 65.32 mg of CE/g of FW (taxo), where taxo showed the
best results for both TPC and TFC assays. In terms of
ACY content, the lowest values were found in uvilla
(0.34 mg PgEq/g FW), while the highest were found in
mortiño (13.50 mg PgEq/g FW). The results here exposed
are in line with those previously reported for some of the
fruits here studied [3, 6, 15, 16], as well as those reported
in fruits from other geographical regions [17].
Regarding vitamin C levels (Table 1), the results ranged
between 13.82 (aguacate) to 564.46 mg of Vit C/100 g of
FW (ají ratón), which is in line with the results previously
reported in some of the fruits here studied from other geo-
graphical regions [17–19] and from Ecuador [3, 6, 7].
According to the classification proposed by Ramful et al.
[20] regarding the Vit C content in fruits, 85% of the fruits
here analysed were classified in the group of high Vit C con-
tent (> 50 mg/100 g of FW), while only one fruit was classi-
fied in the intermedium group (30-50 mg/100 g of FW) and
one in the low content group (< 30 mg/100 g of FW). The
intake on a regular basis of this vitamin is essential in order to
maintain a healthy state and prevent diseases caused by oxi-
dative stress and aging. [21]. According to the Food and
Nutrition Board at the Institute of Medicine of the National
Academies [22], the Recommended Dietary Allowances for
vitamin C for an adult (over 19 years old) is 90 mg for male
and 75 mg for female. According to the total Vit C content in
the fruits here analyzed, 53% of them exceed the recommend-
ed consumption limits for 100 g of FW, which shows the high
contribution of this vitamin in the fruits consumed in this
region.
Legumes also proved to be an important source of bioac-
tive compounds [17, 23]. The chocho perla showed the
highest TPC (1270.74 mg of GAE/100 g FW) and TFC
(4.51 mg of CE/g of FW) values. Following the previous
classification, according to the TPC, 37.5% of total legumes
were in the low category, 50% were in the medium category
and only 12.5% were in the high category (Fig. S1B).
Moreover, ACY content ranged between 0.33 (haba pallar)
and 0.86 mg PgEq/g FW (frejol negro), a range similar to
the content reported in fruits, whereas vitamin C was detected
only in haba pallar, classified in the group of low Vit C content
(Table 1).
Within the tubers, jícama showed the highest TPC
(680.19 mg of GAE/100 g FW) and TFC (6.57 mg of CE/g
of FW). According to the previous classification, taking into
account the TPC, tubers were classified in the medium and
 Plant Foods Hum Nutr

Table 1 Chemical composition and antioxidant capacity of the major fruits, legumes, and tubers consumed in the Andean regions of Ecuador

Common name TAC (μmol TE/g FW) Bioactive compounds

FRAP DPPH TPC (mg GAEq/100 g FW) TFC (mg CatEq/g) ACY Vit C (mg
(mg PgEq/g) FW) Vit C/100 g)

Fruits
Aguacate 16.88 ± 2.25g 39.66 ± 8.16j 455.37 ± 20.20g 6.06 ± 0.79f 0.85 ± 0.08c 13.82 ± 2.46g
e
Ají ratón 52.29 ± 3.53 104.58 ± 10.69g 1298.46 ± 140.51c 10.07 ± 1.54d 0.94 ± 0.11c 564.46 ± 59.84a
Babaco 19.56 ± 2.26g 166.55 ± 28.56f 340.86 ± 11.84h 1.14 ± 0.23h 0.35 ± 0.07e 70.04 ± 12.27d
Granadilla 83.72 ± 2.68d 517.30 ± 57.09d 1611.87 ± 298.82c 45.17 ± 4.18b 0.73 ± 0.11c 35.90 ± 6.35f
Guayaba 78.88 ± 2.37d 739.24 ± 32.22c 118.84 ± 7.22j 3.81 ± 0.61g 0.40 ± 0.08e 253.01 ± 21.96b
Mora de Castilla 131.29 ± 8.22c 55.55 ± 8.65i 126.66 ± 4.93j 8.60 ± 1.15e 2.40 ± 0.68b 52.26 ± 6.27e
Mortiño 237.77 ± 26.15b 1337.63 ± 169.06b 3476.03 ± 190.14b 13.13 ± 1.95c 13.50 ± 3.05a 184.44 ± 22.07c
Naranjilla 21.07 ± 2.31g 108.47 ± 10.96g 727.93 ± 12.47f 3.09 ± 0.37g 0.63 ± 0.09d 81.53 ± 14.02d
Pepino dulce 39.79 ± 2.84f 118.34 ± 18.32g 1420.45 ± 204.67c 6.17 ± 1.07f 0.62 ± 0.07d 291.39 ± 41.23b
Taxo 791.20 ± 20.86a 3361.77 ± 320.08a 6839.64 ± 275.86a 65.32 ± 4.50a 0.62 ± 0.08d 217.05 ± 27.14b
Tomate de árbol 39.63 ± 4.37f 198.05 ± 41.65e 682.20 ± 23.33f 4.66 ± 0.70g 0.35 ± 0.07e 164.43 ± 12.02c
Tuna 15.19 ± 3.18g 39.71 ± 5.38j 620.22 ± 30.05f 4.26 ± 0.72g 0.62 ± 0.09d 63.12 ± 9.02d
Uvilla 19.41 ± 2.19g 78.62 ± 5.91h 255.98 ± 31.73i 1.56 ± 0.27h 0.34 ± 0.06e 121.51 ± 18.03c
Legumes
Chocho ud 87.36 ± 18.55b 80.19 ± 5.22e 2.17 ± 0.46b 0.36 ± 0.08b ud
Chocho perla 118.66 ± 15.53c ud 1270.74 ± 20.437a 4.51 ± 0.45a 0.58 ± 0.05b ud
Frejol blanco 71.74 ± 4.77d ud 81.71 ± 7.12e 0.39 ± 0.04d 0.46 ± 0.09b ud
Frejol canario 198.57 ± 6.89b 27.69 ± 4.75d 173.40 ± 9.22c 2.79 ± 0.74b 0.44 ± 0.08b ud
Frejol de castilla 230.53 ± 3.67b 119.35 ± 20.92a 245.14 ± 23.06b 1.59 ± 0.41c 0.47 ± 0.09b ud
a
Frejol negro 319.97 ± 9.52 47.77 ± 10.45c 195.88 ± 23.25c 3.35 ± 0.42b 0.86 ± 0.08a ud
Frejol panamito 129.19 ± 5.80c 81.16 ± 14.06b 134.35 ± 6.90d 1.69 ± 0.53c 0.58 ± 0.09b ud
Haba pallar 6.36 ± 0.74e 31.55 ± 5.82d 92.55 ± 3.58e 2.76 ± 0.70b 0.33 ± 0.07b 15.53 ± 3.38
Tubers
Jícama 713.19 ± 33.68a 0.60 ± 0.07b 680.67 ± 19.36a 6.57 ± 0.52a 0.60 ± 0.07a ud
Melloco 326.98 ± 7.83b ud 264.82 ± 36.76b 1.55 ± 0.54b 0.62 ± 0.05a 116.86 ± 8.89a
Oca 254.55 ± 4.69c 84.43 ± 17.61a 266.78 ± 31.07b 2.32 ± 0.89b 0.33 ± 0.07b 68.84 ± 10.20b

Values are expressed as means ± standard deviation (SD). Mean values within a column with different letter are significantly different (P < 0.05). Samples
were analyzed in triplicate. ud , undetectable.

high category (Fig. S1C). Although low compared to the
fruits, tubers also showed a certain amount of anthocyanins
and vitamin C. The ACY content in the tubers ranged between
0.33 (oca) and 0.62 mg PgEq/g FW (melloco), while Vit C
ranged between 68.84 (oca) and 116.86 mg of Vit C/100 g FW
(melloco). Although vitamin C is present in tubers, one must
take into account that the high temperatures to which
they are exposed during the cooking processes could
affect the bioavailability of this compound. We were
unable to find scientific reports about the bioactive
chemical composition of the main legumes and tubers
consumed in Ecuador. Nevertheless, our results were
within the range of those previously reported in legumes
and tubers from geographical regions with similar climatic
characteristics to Ecuador [17].
Total Carotene Content (TCC) TCC was also analyzed
(Table S2). β carotene content ranged between 98.61 (mora
de castilla) and 1624.83 μg of β carotene/100 g of FW (taxo),
lutein ranged between 213.77 (pepino dulce) and 5557.11 μg
of lutein/100 g of FW (ají ratón), while lycopene was only
found in guayaba (1038.50 μg of lycopene/100 g of FW). In
these cases, taxo fruits showed the highest values of β caro-
tene content [24], while ají ratón was the principal source of
lutein. In the case of legumes, β carotene was found only in
chocho (141.68 μg of β carotene/100 g of FW) and haba
pallar (73.80 μg of β carotene/100 g of FW), lutein range
between 89.64 (frejol canario) and 291.54 μg of lutein/100 g
of FW (chocho perla), while lycopene was not found in any of
the legumes analyzed. Regarding tubers, jícama showed the
highest values of β carotene (1147.57 μg of β carotene/100 g
Plant Foods Hum Nutr
of FW) followed by oca (57.85 μg of β carotene/100 g of
FW), whilst β carotene was not found in melloco.
The lutein content in tubers ranged between
119.16 (jícama) and 2330.50 μg of lutein/100 g of FW
(melloco), while lycopene was not found in any of the tubers
studied. Although our group has carried out studies about
carotene content in fruits in mora de Castilla, mortiño [6]
and capulí [7], there are very few reports about the carotenoid
content in the fruits, legumes, and tubers here studied.
However, the levels of carotenoid content here reported are,
in some cases, higher than those previously reported in fruits,
legumes, and tubers from other geographical regions [17, 24,
25], which could be related to the plants’ exposure to extreme
climatic conditions [26]. Based on intake recommendations
for carotenoids [22], all the fruits, legumes, and vegetables
here analyzed exceed the recommended consumption limits
for 100 g of fresh weight, which shows the high contribution
of this compound in the regional diet. A large amount of
observational epidemiological evidence suggests that higher
blood concentrations of β carotene and other carotenoids ob-
tained from foods are associated with lower risks of several
chronic diseases [22]. Therefore, the main fruits, legumes, and
tubers produced in the Andean regions of Ecuador could con-
stitute important dietary sources of this type of bioactive com-
pounds that would contribute to the health of the populations
that consume them.
Organic Acids Content The organic acids have been identi-
fied as responsible, in great measure, for the texture and
flavour of fruits, which play a fundamental role in con-
sumers’ acceptance [27]. The results of the organic acids
analysis are presented in Table S3. To the best of our knowl-
edge, this report constitutes the first study on the profile of
organic acids in fruits, legumes, and tubers consumed in
Ecuador. Similar to the previous analysis, the content of
organic acids showed significant differences (P < 0.05) be-
tween the different fruits, legumes and tubers. The main
organic acids identified were lactic, citric, and acetic acid,
while oxalic acid was detected only in six fruits, one le-
gume, and one tuber. Citric acid was detected with the
highest values in naranjilla (11.34 g/100 g FW), while acetic
acid was found in higher concentrations in aguacate (8.21 g/
100 g FW), values which are higher than those previously
reported in fruits from other geographical regions, while the
values for the rest of fruits are within the ranges previously
described [11, 27]. Uvilla fruit showed the highest values
for both acetic (4.26 g/100 g FW) and citric acid (9.64 g/
100 g FW), while the highest values of lactic acid were
found in granadilla (2.69 g/100 g FW). Among legumes,
haba pallar (2.23 g/100 g FW) showed the highest values
of lactic acid, whilst chocho perla showed the highest values
of acetic (2.33 g/100 g FW) and citric acid (1.24 g/100 g
FW). Oxalic acid was identified in lower concentration and
frequency with respect to the other acids, identifying itself
only in in six fruits, one legume, and one tuber. The amount
of organic acids present in the fleshy parts of the fruits of
different species can vary greatly, as can the relative abun-
dance of the individual organic acids. In addition, their con-
tents depend on the fruit’s stage of development and diverse
environmental factors [26]. Therefore, the differences
found here between the different foods and between our
results and previously reported values could be explained,
to a certain degree as mentioned above.
Glucose and Fructose Content Sugar content affects the qual-
ity and flavour of the fruits, thus playing an important role in
how consumers perceive and accept the fruits [12]. The results
of the sugar content are presented in Table S4. Regarding
fruits, the highest values of glucose were found in mortiño
(6.80 g/100 g FW), while guayaba (0.12 g/100 g FW) showed
the lowest values. In legumes, haba pallar (6.02 g/100 g FW)
showed the highest values of glucose, while the lowest values
were found in frejol blanco (0.05 g/100 g FW). The jícama
was the tuber with the highest glucose content (3.04 g/100 g
FW), while oca showed the lowest values (0.72 g/100 g FW).
The glucose values in fruits were in the range of those previ-
ously reported in several fruits [28], however the fruits here
studied could be considered low in glucose content in com-
parison to those previously reported in that database [28]. It is
interesting to highlight the high glucose content of frejol ne-
gro, compared to the other beans studied here, which ranged
between 0.05 and 0.70 g/100 g of FW, which is also higher
than values previously reported in other beans [28].
In terms of fructose content, guayaba was the fruit with the
highest values (3.90 g/100 g FW), while the lowest values
were found in tomate de árbol (0.15 g/100 g FW).
Concerning legumes, haba pallar showed the highest values
(0.22 g/100 g of FW); while melloco was the tuber with the
highest values (2.38 g/100 g of FW). The values of fructose in
the different beans here studied differ to those previously re-
ported in other beans types, where fructose was not found
[28]. These differences could be due to the species and geo-
graphical area where these types of grains grow. Crops in the
Andean regions of Ecuador are located at more than 2800
masl where low temperatures, high levels of radiation, such
as UV, and low oxygen levels could influence the content of
this type of compound [6, 26]. However, compared to the
content of sugars reported for other types of fruits, legumes,
and tubers [28], the results here exposed demonstrate that the
principal fruits, legumes, and tubers consumed in the
Ecuadorian Andes could be considered as low in terms of
sugar content.
Total Antioxidant Activity The TAC results (Table 1) showed a
significant variation between the different fruits, legumes, and
tubers, where between the fruits, the highest values for both
 Plant Foods Hum Nutr

methods used (FRAP and DPPH) were found in taxo fruits
(791.20 μmol TE/g FW for FRAP assay and 119.35 μmol TE/
g FW for DPPH assay), which are in agreement with the
highest TAC values previously reported in this fruit [24]. In
legumes, frejol negro showed the highest values of FRAP
(319.97 μmol TE/g FW), whilst frejol de Castilla showed
the highest ability to scavenge the DPPH radicals
(119.35 μmol TE/g FW), which has been mainly attributed
to the presence of phenolic acids, flavonoids and tannins [23].
Jícama was the tuber with the highest values of FRAP
(713.19 μmol TE/g FW), while the oca tuber showed the
highest capacity to capture DPPH radicals (84.43 μmol TE/g
FW) (Table 1). Since that, the total antioxidant capacity of the
foods from plant origin is given by the interaction of several of
its phytochemicals, a factorial and cluster analysis was carried
out for all the fruits, legumes, and grains here studied and was
used to determine which among them showed the greatest
contribution in terms of antioxidant activity and which com-
pounds contributed to it (Fig. 1a). The factorial analysis was
performed using TPC, TFC, Vit C, FRAP, and DPPH using
varimax rotation. Only one factor was extracted (PC1) which
explained 79.15% of the variance. The Person’s correlation
coefficient between these five variables and the extracted fac-
tor are higher than 0.73 and positive in all cases. This new
variable (PC1) will represent the combined antioxidant prop-
erties. Figure 1 showed that taxo had the highest contribution
followed by mortiño, granadilla, and jícama, as compared to
all the other fruits, legumes, and tubers. Moreover, the

Fig. 1 Factorial analysis and

cluster carried out using the total
phenolic content, total flavonoid
content, vitamin C content,
FRAP, and DPPH assay using
varimax rotation using the values
of all samples of fruits, legumes
and tubers (a). Dendogram
created using Ward’s method and
the rescaled squared Euclidian
distance using the chemical
composition of the different fruits,
legumes and tubers (b)
Plant Foods Hum Nutr
dendogram created using Ward’s method and the rescaled
squared Euclidian distance using the chemical composition
shown the occurrence of five distinct groups in which the
fruits, legumes, and tubers analyzed were organized, where
taxo was located in the first largest group (Fig. 1b).
Conclusions In this study, chemical composition and antioxi-
dant capacity were determined in the main fruits, tubers, and
legumes traditionally consumed in the Andean regions of
Ecuador. The results showed a wide range of variation in
terms of chemical composition and TAC between the different
fruits, legumes, and tubers. According to TPC, there were
three well-defined groups that could be identified showing
high, medium, and low phenolic content and TAC, with the
fruits as the group with the highest values, all classified as
medium or high. The taxo fruit presented the highest values
in terms of antioxidant potential. In this sense, this fruit de-
serves further investigation relating to its chemical composi-
tion, principally to its phenolic profile. To the best of our
knowledge, our research constitutes the first report re-
lated to the content of bioactive compounds and antiox-
idant capacity of the most commonly consumed legumes
and tubers linked to the traditional diet of the Andean
peoples of Ecuador. Given all this, our future research
will be aimed at determining the phenolic profile of
each of the fruits, legumes and tubers consumed in the
Andean region of Ecuador.
Funding This study was funded by the Universidad de Las Américas
[grant number: VET.JMA.17.05], Quito, Ecuador. Access to plant genetic
resources was granted by means of the BFramework agreement for access
to plant genetic resources: MAE-DNB-CM-2017-0072-M-001^ celebrat-
ed between the Environment Ministry, Ecuador and the Universidad de
Las Américas, Ecuador.
Compliance with Ethical Standards
Conflict of Interest The authors declare no conflicts of interest.
References
1. Tene V, Malagón O, Finzi PV, Vidari G, Armijos C, Zaragoza T
(2007) An ethnobotanical survey of medicinal plants used in Loja
and Zamora-Chinchipe, Ecuador. J Ethnopharmacol 111:63–81.
https://doi.org/10.1016/j.jep.2006.10.032
2. Carrillo-Perdomo E, Aller A, Cruz-Quintana SM, Giampieri F,
Alvarez-Suarez JM (2015) Andean berries from Ecuador: a review
on botany, agronomy, chemistry and health potential. J Berry Res 5:
49–69
3. Vasco C, Ruales J, Kamal-Eldin A (2008) Total phenolic com-
pounds and antioxidant capacities of major fruits from Ecuador.
Food Chem 111:816–823. https://doi.org/10.1016/j.foodchem.
2008.04.054
4. Vasco C, Riihinen K, Ruales J, Kamal-Eldin A (2009) Phenolic
compounds in rosaceae fruits from Ecuador. J Agric Food Chem
57:1204–1212. https://doi.org/10.1021/jf802656r
5. Vasco C, Riihinen K, Ruales J, Kamal-Eldin A (2009) Chemical
composition and phenolic compound profile of mortiño (Vaccinium
floribundum Kunth). J Agric Food Chem 57:8274–8281. https://
doi.org/10.1021/jf9013586
6. Alarcón-Barrera KS, Armijos-Montesinos DS, García-Tenesaca M,
Iturralde G, Jaramilo-Vivanco T, Granda-Albuja MG, Giampieri F,
Alvarez-Suarez JM (2018) Wild Andean blackberry (Rubus
glaucus Benth) and Andean blueberry (Vaccinium floribundum
Kunth) from the highlands of Ecuador: nutritional composition
and protective effect on human dermal fibroblasts against cytotoxic
oxidative damage. J Berry Res 8:223–236. https://doi.org/10.3233/
JBR-180316
7. Alvarez-Suarez JM, Carrillo-Perdomo E, Aller A, Giampieri F,
Gasparrini M, González-Pérez L, Beltrán-Ayala P, Battino M
(2017) Anti-inflammatory effect of Capuli cherry against LPS-
induced cytotoxic damage in RAW 264.7 macrophages. Food
Chem Toxicol 102:46–52. https://doi.org/10.1016/j.fct.2017.01.
024
8. Singleton VL, Orthofer R, Lamuela-Raventós RM (1998) Analysis
of total phenols and other oxidation substrates and antioxidants by
means of folin-ciocalteu reagent. Methods Enzymol 299:152–178.
https://doi.org/10.1016/S0076-6879(99)99017-1
9. Dewanto V, Wu X, Adom KK, Liu RH (2002) Thermal processing
enhances the nutritional value of tomatoes by increasing total anti-
oxidant activity. J Agric Food Chem 50:3010–3014
10. Giusti M, Wrolstad RE (2005) Characterization and measurement
of anthocyanins by UV-visible spectroscopy. In: Handbook of food
analytical chemistry. John Wiley & Sons, Ltd, pp 19–31
11. Wang L, Fu H, Wang W, Wang Y, Zheng F, Ni H, Chen F (2018)
Analysis of reducing sugars, organic acids and minerals in 15 cul-
tivars of jujube (Ziziphus jujuba mill.) fruits in China. J Food
Compos Anal 73:10–16. https://doi.org/10.1016/j.jfca.2018.07.008
12. Doyon G, Gaudreau G, St-Gelais D, Beaulieu Y, Randall CJ (2013)
Simultaneous HPLC determination of organic acids, sugars and
alcohols. Can Inst Food Sci Technol J 24:87–94. https://doi.org/
10.1016/s0315-5463(91)70025-4
13. Benzie IFF, Strain JJ (1996) The ferric reducing ability of plasma
(FRAP) as a measure of Bantioxidant power^: the FRAP assay.
Anal Biochem 239:70–76. https://doi.org/10.1006/abio.1996.0292
14. Prymont-Przyminska A, Zwolinska A, Sarniak A, Wlodarczyk A,
Krol M, Nowak M, de Graft-Johnson J, Padula G, Bialasiewicz P,
Markowski J, Rutkowski KP, Nowak D (2014) Consumption of
strawberries on a daily basis increases the non-urate 2,2-diphenyl-
1-picryl-hydrazyl (DPPH) radical scavenging activity of fasting
plasma in healthy subjects. J Clin Biochem Nutr 55:48–55.
https://doi.org/10.3164/jcbn.13-93
15. Gancel A-L, Alter P, Dhuique-Mayer C, Ruales J, Vaillant F (2008)
Identifying carotenoids and phenolic compounds in Naranjilla (
Solanum quitoense lam. var. Puyo hybrid), an Andean fruit. J
Agric Food Chem 56:11890–11899. https://doi.org/10.1021/
jf801515p
16. Espin S, Gonzalez-Manzano S, Taco V, Poveda C, Ayuda-Durán B,
Gonzalez-Paramas AM, Santos-Buelga C (2016) Phenolic compo-
sition and antioxidant capacity of yellow and purple-red Ecuadorian
cultivars of tree tomato (Solanum betaceum Cav.). Food Chem 194:
1073–1080. https://doi.org/10.1016/J.FOODCHEM.2015.07.131
17. Campos D, Chirinos R, Gálvez Ranilla L, Pedreschi R (2018)
Bioactive potential of Andean fruits, seeds, and tubers. In:
Advances in food and nutrition research. Academic Press, pp
287–343
18. Kantar MB, Anderson JE, Lucht SA, et al (2016) Vitamin variation
in Capsicum spp. provides opportunities to improve nutritional val-
ue of human diets. https://doi.org/10.1371/journal.pone.0161464
19. Bazalar Pereda MS, Nazareno MA, Viturro CI (2018) Nutritional
and antioxidant properties of Physalis peruviana L. fruits from the

Argentinean northern Andean region. Plant Foods Hum Nutr 74:
68–75
20. Ramful D, Tarnus E, Aruoma OI, Bourdon E, Bahorun T (2011)
Polyphenol composition, vitamin C content and antioxidant capac-
ity of Mauritian citrus fruit pulps. Food Res Int 44:2088–2099.
https://doi.org/10.1016/j.foodres.2011.03.056
21. Li Y, Schellhorn HE (2018) New developments and novel thera-
peutic perspectives for vitamin C. J Nutr 137:2171–2184. https://
doi.org/10.1093/jn/137.10.2171
22. Compounds I of M (US) P on DA and R (2000) Dietary reference
intakes for vitamin C, vitamin E, selenium, and carotenoids.
National Academies Press (US)
23. Chávez-Mendoza C, Sánchez E (2017) Bioactive compounds from
Mexican varieties of the common bean (Phaseolus vulgaris): im-
plications for health. Molecules 22:1360. https://doi.org/10.3390/
molecules22081360
24. García-Ruiz A, Girones-Vilaplana A, León P, Moreno D, Stinco C,
Meléndez-Martínez A, Ruales J (2017) Banana passion fruit
(Passiflora mollissima (Kunth) L.H. bailey): microencapsulation,
phytochemical composition and antioxidant capacity. Molecules
22:85. https://doi.org/10.3390/molecules22010085
Plant Foods Hum Nutr
25. García-Cruz L, Valle-Guadarrama S, Salinas-Moreno Y, Joaquín-
Cruz E (2013) Physical, chemical, and antioxidant activity charac-
terization of pitaya (Stenocereus pruinosus) fruits. Plant Foods
Hum Nutr 68:403–410. https://doi.org/10.1007/s11130-013-
0391-8
26. Khayyat M, Barati Z, Aminifard MH, Samadzadeh A (2018)
Changes in fruit maturity indices and growth pattern along the
harvest season in seedless barberry under different altitude condi-
tions. J Berry Res 8:25–40. https://doi.org/10.3233/JBR-170166
27. Walker RP, Famiani F (2018) Organic acids in fruits. In:
Horticultural reviews. John Wiley & Sons, Inc., Hoboken, NJ,
USA, pp 371–430
28. United States Department of Agriculture Agricultural Marketing
Service Fruit and Vegetable Division Processed Products Branch
United States Standards for Grades of Extracted Honey
Publisher’s Note Springer Nature remains neutral with regard to jurisdic-
tional claims in published maps and institutional affiliations.