Gs 001

CHAPTER EIGHT
Bioactive Potential of Andean

Fruits, Seeds, and Tubers
David Campos*, Rosana Chirinos*, Lena Gálvez Ranilla†,
Romina Pedreschi‡,1
*Institute of Biotechnology (IBT), Universidad Agraria La Molina, Lima, Peru
†
Universidad Catolica de Santa Maria, Arequipa, Peru
‡
School of Agronomy, Pontificia Universidad Católica de Valparaı́so, Valparaı́so, Chile
1
Corresponding author: e-mail address: romina.pedreschi@pucv.cl
Contents
1. Biodiversity of the Andes 288
2. Fruits 288
2.1 General Information on Composition and Nutritional Value 288
2.2 Typical Use in Old Traditions 290
2.3 Reported Bioactive Potential 291
3. Seeds/Andean Grains 293
4. Tubers and Roots 317
5. Conclusions 331
References 331
Further Reading 343
Abstract
The Andes is considered the longest continental mountain range in the world. It covers
7000 km long and about 200–700 km wide and an average height of about 4000 m. Very
unique plant species are endemic of this area including fruits (e.g., lucuma, cherimoya,
sweet pepino, sauco), roots and tubers (potatoes, sweet potatoes, yacón, chicuru,
mashua, olluco, etc.), and seeds (quinoa, amaranth, tarwi, etc.). These crops have been
used for centuries by the native population and relatively recently have gained the
world attention due to the wide range of nutrients and/or phytochemicals they possess.
In this chapter, main Andean fruits, seeds, and roots and tubers have been selected and
detailed nutritional and functional information is provided. In addition, traditional and
current uses are provided and their bioactive potential is reported based on published
scientific literature.
Advances in Food and Nutrition Research, Volume 84 # 2018 Elsevier Inc. 287
ISSN 1043-4526 All rights reserved.
https://doi.org/10.1016/bs.afnr.2017.12.005
288 David Campos et al.
1. BIODIVERSITY OF THE ANDES

The Andes is considered the longest continental mountain range in
the world. It covers 7000 km long and about 200–700 km wide and an aver-
age height of about 4000 m. It extends from north to south through seven
South American countries: Venezuela, Colombia, Ecuador, Peru, Bolivia,
Argentina, and Chile. Very unique plant species are endemic of this area
including fruits (e.g., lucuma, cherimoya, sweet cucumber, sauco), roots
and tubers (potatoes, sweet potatoes, yacón, chicuru, mashua, olluco,
etc.), seeds and grains (corn, tarwi, etc.), and pseudocereals (quinoa,
kañiwua, etc.) (Brack, 2012). The biodiversity of plant species found in
the Andes is very extensive. Thus, this chapter will present a selection of
fruits, roots and tubers, and seeds.
2. FRUITS
2.1 General Information on Composition and
Nutritional Value
2.1.1 Cherimoya
Cherimoya (Annona cherimola Mill) is a tree cultivated prior and during
the times of the Incas, dating back to 1200 BC in the inner valleys of Peru
and Ecuador. It is a subtropical fruit native to the Andes, with a thick green
peel and a creamy and sweet mesocarp. Its exceptional taste and flavor have
helped to increase its production in tropical and subtropical regions. Main
producers of cherimoya are Spain, Peru, and Chile, and small production
areas exist in California, Israel, and Madeira Island (Portugal) (Gonçalves
Alburqueque et al., 2016). In terms of nutritional composition, it contains
75.1 g water/100 g fresh weight (FW), 1.20 protein/100 g FW, 0.2 fat/100 g
FW, 22.6 g carbohydrates/100 g FW, and 2.4 g fiber/100 g FW. It
presents high content of sugars, moderate content of calcium and phospho-
rus (20 and 63 mg/100 g, respectively), and a good source of thiamine
(0.09 mg/100 g FW), riboflavin (0.16 mg/100 FW), and niacin (1.62 mg/
100 g FW). An ascorbic acid content of 3.30 mg/100 g FW has been
reported (Ministerio de Salud, 2009). Two acetogenins were reported in
the seeds: acetogenins, cherimolin-2, and almunequin (Cortes, Myint,
Dupont, & Davoust, 1993).
Bioactive Potential of Andean Fruits, Seeds, and Tubers 289
2.1.2 Lucuma
Pouteria lucuma Ruiz and Pav. is known as the “Gold of the Incas” or
“lucuma” (Fig. 1) and it is a food of ancient cultivation before the Incas
(Dini, 2011). It is a subtropical fruit from the Andean region of Peru, Chile,
and Ecuador that belong to the Sapotaceae family and to the genus Pouteria.
The chemistry and biological activities of this family were previously
reviewed (Silva, Simeoni, & Silveira, 2009) and as main constituents,
triterpenes, and flavonoids were reported besides their antioxidant, anti-
inflammatory, antibacterial, and antifungal activities. Lucuma mesocarp is rich
in carbohydrates, protein, fiber mainly in the insoluble form, β-carotene,
vitamins, and minerals (Yahia, & Gutierrez-Orozco, 2011). Nutritional value
of lucuma flesh fruit is composed of (g/100 g FW): 72.3% water, 1.5% protein,
25.0% carbohydrates, 0.5% fat, and 1.1% fiber. Reported nutrient contents
correspond to: 16mg calcium/100 g FW, 26 mg phosphorus/100 g FW,
0.4 mg iron/100 g FW, 0.01 mg thiamine/100 g FW, 0.14 mg riboflavin/
100 g FW, 1.96 mg niacin/100 g FW, and 2.2 mg vitamin C/100 g FW
(Ministerio de Salud, 2009). In general, it is considered as a good source of
fiber, minerals, β-carotene, phenolics, and niacin.
2.1.3 Andean Berries

Goldenberry or cape gooseberry (Physalis peruviana L.) is a fruit of the
Andes belonging to the Solanaceae family. It is a golden sphere with a paper
Fig. 1 Andean fruits: (A) lucuma, (B) cherimoya, (C) sweet pepino, (D) goldenberry, and
(E) sauco.
husk, the size of marble with a pleasant taste and great source of vitamin C
(28 mg/100 g FW), vitamin A (0.15 mg/100 g FW), and vitamin E
(0.5 mg/100 g FW) (Ramadan & Morsel, 2003). Main nutritional compo-
nents are (g/100 g FW): 81% water, 13.3% carbohydrates, 1.1% fat, and
0.5% fiber. It also contains different minerals such as iron (1.3 mg/100 g
FW), magnesium (8 mg/100 g FW), potassium (170 mg/100 g FW), phos-
phorus (39 mg/100 g FW), and calcium (12 mg/100 g FW).
Sambucus peruviana H.BK is originary from the Andes but distributed
from Costa Rica to Argentina. It grows in temperate climates at altitudes
between 1200 and 3500 m above sea level (m.a.s.l) (Brack, 2012). The
flowers of S. peruviana contain butyrose composed of a terpene and a resin.
The bark contains an alkaloid known as sambucin together with a resin.
Similarly, the leaves contain an alkaloid (Minagri, 2017).
2.1.4 Sweet Pepino

Pepino (Solanum muricatum Aiton) is an aromatic and sweet fruit, with a juicy
melting flesh, native to the Andean region. It belongs to the Solanaceae
family being close relative of the tomato (Solanum lycopersicum) and the
potato (Solanum tuberosum) (Contreras, Schwab, Mayershofer, González-
Ag€ uero, & Defilippi, 2017). It is composed of water (92 g/100 g), protein
(0.3 g/100 g), carbohydrates (7 g/100 g), and fiber (0.5 g/100 g). It is a
rich source of potassium (117 mg/100 g FW) and contains other minerals
such as calcium (30 mg/100 g FW) and phosphorus (10 mg/100 g FW).
It is an important source of vitamin C (29.7 mg/100 g FW), also contains
vitamin A (29.70 mg/100 g FW) and vitamin B1 (0.04 mg/100 g FW)
(Ministerio de Salud, 2009) (Fig. 1).
2.2 Typical Use in Old Traditions

Annona cherimola’s main use by the population has been as fruit and in desserts
and bakery but the seeds have been used as medicine and pesticides. In
addition, an alcoholic beverage known as “cherimoya wine” is elaborated
(Brack, 2012).
P. lucuma, is mainly consumed in desserts, dairy, and baked products but
can also be consumed fresh. The inedible seed is rich in polyunsaturated fatty
acids with attributes appreciated by the cosmetic industry in terms of skin
regeneration (Rojo et al., 2010). Both cherimoya and lucuma were repre-
sented in ceramics during pre-Inca times.
P. peruviana is consumed fresh but also mainly processed into marmalade
and used in the bakery industry.
S. peruviana can be consumed fresh but it is mainly processed into mer-

melades. It is also used to blue dye. In some places of the Andes, the flowers
are used as refreshment, after immersion in white vinegar with the addition
of water and sugar. The flowers are also used cooked as sudorific and mixed
with soap as patches to favor suppuration (Minagri, 2017). The cooked fruits
are also used to treat mouth infections.
Sweet pepino is consumed as fresh refreshing fruit and with hydrating
properties after physical efforts. People from Moche and Virú cultures
carried sweet pepinos with them during the long walks along the desert
and probably associated to its high potassium content. Its diuretic properties
are well known (Brack, 2012). Also properties against goiter are attributed
and possibly correlated to its good iodine content.
2.3 Reported Bioactive Potential

2.3.1 Cherimoya
Relatively recent works have reported a significant amount of phenolics in
the mesocarp, skin, and seeds of different cultivars of cherimoya (Garcı́a-
Salas, Gómez-Caravaca, Morales-Soto, Segura-Carretero, & Fernández-
Gutierrez, 2015; Gonçalves Alburqueque et al., 2016; Santos et al.,
2016). Garcı́a-Salas et al. (2015) reported a total of 21 phenolic and other
polar compounds in cherimoya pulp, 37 in peel and 22 in seeds. The most
represented family of polyphenols present in cherimoya pulp and peel was
flavan-3-ols. The peel was the richest part of the fruit in terms of phenolics
followed by the seeds and pulp and there were differences between cultivars.
Santos et al. (2016) reported lipophilic and phenolics of the ripe mesocarp of
four cherimoya cultivars (Perry Vidal, Mateus I, Mateus III, and Funchal)
from Madeira Island being the predominant lipophilic compounds: kaurene
diterpenes (42.2%–59.6%), fatty acids (18.0%–35.6%), and sterols (9.6%–
23.7%). Kaur-16-en-19-oic acid was the major lipophilic component of
all cultivars (554 and 1350 mg/kg of dry material). These cultivars presented
a high variety of flavan-3-ols, including galloylated and nongalloylated
compounds. Five phenolic compounds were identified for the first time:
catechin, (epi)catechin-(epi)gallocatechin, (epi)gallocatechin, (epi)afzelechin-
(epi)catechin, and procyanidin tetramer. “Mateus I” and “Mateus III” cultivars
present the highest content of phenolic compounds (6299 and 9603 mg/kg of
dry weight, DW, respectively). Gonçalves Alburqueque et al. (2016) reported a
nutritional composition in terms of proximates and vitamins and bioactive
compounds content in four cultivars of Madeira Island. The peel of Madeira
cultivar showed the highest antioxidant capacity, with an EC50 of 0.97 mg/mL,
and total flavonoids (44.7 epicatechin equivalents/100 g). The most abundant
carotenoid was lutein, with values ranging from 129 to 232 μg/100 g. The
highest L-ascorbic acid content (4.41 mg/100 g) was found in the peel of Perry
Vidal cultivar. In comparison to other Andean fruits, cherimoya presents a
low content of ascorbic acid (4–6 mg/100 g FW) but an intermediate content
of total phenolics and hydrophilic DPPH antioxidant activity (323 83 mg
GAE/100g FW and 23 6 μmol Trolox g1 FW, respectively) (Vasco,
Ruales, & Kamal-Eldin, 2008).
2.3.2 Lucuma
Fuentealba et al. (2016) characterized the primary (e.g., sugars and organic
acids) and secondary (e.g., phenolics and carotenoids) and in vitro antioxi-
dant and antihyperglycemic properties of Rosalia, Montero, and Leiva 1
lucuma biotypes. Significant differences were found in these metabolites
and functional properties related to biotype and ripeness stage. Results
showed significant amounts of sugars (119.4–344 mg total sugars g1 DW)
and organic acids (44.4–30.0 mg g1 DW) and functional associated com-
pounds such as ascorbic acid (0.35–1.07 mg g1 DW), total phenolics
(0.7–61.6 mg GAE g1 DW), and total carotenoids (0.22–0.50 mg
β-carotene g1 DW). Important in vitro antioxidant and antihyperglycemic
properties were also reported for this emblematic fruit.
2.3.3 Andean Berries

For P. peruviana, Ramadan and Morsel (2003) compared the contents of fatty
acids, triacylglycerols, phytosterols, fat soluble vitamins, and β-carotene in
whole berries, seeds, and pulp/peel. Whole berries presented higher lipid
contents (2%) compared to the seeds and (1.8%) and pulp/peel (0.2%) being
linoleic and oleic acid the major fatty acids present. The highest level of phy-
tosterols was found in the pulp/peel (dominated by Ϫ5-avenasterol and camp-
esterol) while in the whole berry and seeds, campesterol, and β-sitosterol were
the dominating ones. Tocopherol, β-carotene, and vitamin K levels were
higher in the pulp/peel oil than the whole berry and seed oils. In terms
of hydrophilic compounds such as phenolics, vitamin C, and hydrophilic
antioxidant capacity, Vasco et al. (2008), classified P. peruviana as a low
containing phenolic fruit, with contents of 87 19 mg GAE/100 g FW com-
pared to other fruits such as Andean blackberry containing 2167 835 mg
GAE/100 g FW. DPPH hydrophilic antioxidant activity is pretty
low (0.7 0.4 μmol Trolox g1 sample FW). But a significant content of vita-
min C was present in this fruit species (58–68 mg ascorbic acid/100 FW)
comparable to other fruits rich in vitamin C such as banana passion fruit and
guava (63–93 and 74–84 mg ascorbic acid/100 FW).
S. peruviana known as sauco is an Andean wild berry rich in anthocyanins
and with a significant content of total phenolics of 33.2 0.5 mg GAE g1
DW and hydrophilic antioxidant activity (154.7 5.7 and 361.3 23.6
μmol TE g1 DW DPPH and ORAC, respectively). This hydrophilic
antioxidant potential is much higher than in other Andean fruits such as
tuna (Opuntia ficus indica), guinda (Prunus serotina), and granadilla (Passiflora
ligularis) but slightly lower than tumbo (Passiflora mollisima, 174.3 2.3 and
205.0 18.1 μmol TE g1 DW DPPH and ORAC, respectively) (Chirinos,
Pedreschi, Rogez, Larondelle, & Campos, 2013). This berry is mainly
processed into marmalade and used in the bakery industry.
2.3.4 Sweet Pepino

Few studies exist related to the bioactive potential of this fruit. Vasco et al.
(2008) reported a medium amount of ascorbic acid in sweet pepino
(20–38 mg/100 g FW) comparable to tomato (18–21 mg/100 FW), lower
than banana passion fruit (63–93 mg/100 FW), physalis (58–68 mg/100
FW), and strawberry (50–66 mg/100 FW) but much higher than other
Andean fruits such as cherimoya (4–6 mg/100 FW) and Andean blackberry
(10–11 mg/100 FW). In terms of total soluble phenolic compounds (mg
GAE/100 g sample FW) and DPPH antioxidant capacity (μmol Trolox g1
sample FW) it presents low levels, 56 15 mg GAE/100 g sample FW and
0.3 0.2 μmol Trolox g1 sample FW, respectively, compared to other
Andean fruits such as Andean blackberry (2167 835 mg GAE/100 g sample
FW and 41 16 μmol Trolox g1 sample FW) and banana passion fruit
(1010 198 mg GAE/100 g sample FW and 70 4 μmol Trolox g1 sample
FW, respectively) (Vasco et al., 2008).
3. SEEDS/ANDEAN GRAINS
The most representative indigenous food grains from the Andean
region in South America are quinoa (Chenopodium quinoa Willd.), cañihua
(Chenopodium pallidicaule Aellen), amaranth (Amaranthus caudatus L.), and
lupin or tarwi (Lupinus mutabilis Sweet) ( Jacobsen, Mujica, & Ortiz,
2003) (Fig. 2). Although quinoa, cañihua, and amaranth are similar to
cereals due to their starchy edible seeds and physical appearance, botani-
cally they are known as pseudocereals which are dicotyledonous species
different to the monocotyledonous true cereals (Poaceae family) and in
Fig. 2 Andean grains: (A) quinoa and (B and C) tarwi.
general, their nutritive value in terms of protein content and quality is bet-
ter than the cereal species (Das, 2016; Shewry, 2002). In contrast, the
Andean lupin (L. mutabilis) is a legume (Fabaceae family) (Haq, 1993),
regionally important due to its high seed protein and oil contents
(Atchinson et al., 2016).
These grains have been domesticated and cultivated since pre-
Columbian times and the increase of scientific evidence in relation to their
nutritional and bioactive characteristics is revealing that Andean grains rep-
resent important sources of macro- and micronutrients along with a myriad
of health-relevant phytochemicals with potential for their application in
worldwide food nutrition and security.
3.1 General Information on Composition and Nutritional Value

3.1.1 Quinoa
Quinoa (C. quinoa Willd.) belongs to the family Amaranthaceae, genus
Chenopodium and species quinoa and it was domesticated around 8000 years
ago on the high plateau of the Andes near Lake Titicaca (Kadereit, Borsch,
Weising, & Freitag, 2003; National Research Council, 1989; Pearsall,
1992). Quinoa has been adapted to harsh climatic conditions of the Andes
and maintains its productivity on rather poor soils and under conditions of
water shortage and high salinity (Ruiz et al., 2014). The production of qui-
noa in Andean countries such as Peru, Bolivia, and Ecuador has increased
over the last 10 years (114,725, 74,382, and 3711 tons in year 2014)
(FAOSTAT, 2014) representing an important commercial crop among
these countries. Quinoa cultivation has spread to other places such as the
United States, China, and Europe indicating that quinoa has been adapted
to different agroecological environments (Schoenlechner, Siebenhandl, &
Berghofer, 2008). This increase is related to the fact that quinoa has impor-
tant nutritional properties which are generally better than conventional
cereals.
A comparative proximal composition of quinoa from Andean countries

is shown in Table 1. Ranges of macronutrient contents vary greatly
depending on the origin, variety/ecotype, environmental conditions, and
geographical area of cultivation as observed by Miranda, Vega-Gálvez,
Quispe-Fuentes, et al. (2012) and Repo-Carrasco-Valencia and Serna
(2011). The quinoa protein content and quality are generally better than
common cereal grains. The protein contents found among Andean quinoa
varieties/ecotypes range from 11.1 to 16.2 g/100 g edible portion (EP)
(Table 2) whereas Nowak, Du, and Charrondière (2016) recently reported
ranges of 9.1–15.7 g/100 g EP for quinoa samples from different origins
which were higher than values found in raw yellow corn (9.42 g/100 g
EP), raw rice (6.61 g/100 g EP), and white hard wheat (11.31 g/100 g EP)
(USDA, 2017). The nutritional value of quinoa protein is comparable to
that of milk since it has all essential amino acids with an adequate balance
close to levels recommended by the WHO/FAO/UNU (2007) for adults,
and contents of cereal-limiting amino acids such as lysine and threonine
are higher than values shown by wheat and maize (Dini, Tenore, & Dini,
2005; Kozioł, 1992; Nowak et al., 2016; Wright et al., 2002). The protein
biological value is similar to beef and higher than rice, wheat, and corn
(Bastidas, Roura, Rizzolo, Massanes, & Gomis, 2016). In addition, the
in vitro gastric digestibility of quinoa protein isolates simulating
physiological conditions is also comparable to that of whey and egg protein
depending on heat treatment parameters and extraction conditions (Ruiz
et al., 2016).
The quinoa protein is mainly composed by globulin and albumin with
low concentrations of prolamins (7%) which are the major proteins in
common cereals such as wheat, barley, and rye (Graf et al., 2015;
Zevallos, Ellis, Šuligoj, Herencia, & Ciclitira, 2012). This has increased
the interest on quinoa as an important natural gluten-free ingredient suitable
for people with celiac disease. In this sense, quinoa has been used for improv-
ing the nutritional quality of several gluten-free formulations such as breads,
pasta, and other bakery/confectionary derivatives showing comparable
functional and physical characteristics as those made with wheat flour but
their commercial production is still low and directed to certain target mar-
kets (Alvarez-Jubete, Auty, Arendt, & Gallagher, 2010; Elgeti et al., 2014;
Schoenlechner, Drausinger, Ottenschlaeger, Jurackova, & Berghofer,
2010). Most quinoa cultivars do not have celiac–toxic prolamin epitopes
but Zevallos et al. (2012) revealed that certain Andean quinoa cultivars
showed celiac–toxic epitopes that can activate immune response of patients
Table 1 Proximate Composition (g/100 g Edible Portion) in Raw Quinoa From the Andean Region
Repo-Carrasco-Valencia and Wright, Pike, Fairbanks, Miranda, Vega-Gálvez, Martinez, Nascimento

Reference Serna (2011) and Huber (2002) et al. (2012) et al. (2014)
Variety/ Varieties: Surumi (sweet variety) and a Ecotypes: NR
Ecotype La Molina 89, Kcancolla, bitter variety Ancovinto, Cancosa, Cáhuil, Faro,
Blanca de Juli, Sajama Regalona, Villarica
Country of Peru Bolivia Chile Argentina
Origin
Sampling Lima, Puno Letanias North-altiplano, center, and south Jujuy
Place
Water 10.8–12.6 8.2–9.0 7.7–15.2 11.3
Ash 3.0–4.8 2.6–3.1 3.2–3.7 2.0
a
Total protein 12.4–13.6 14.8–15.7 11.1–16.2 12.1
Total fat 4.1–6.0 5.3–5.7 5.6–7.1 6.3
Crude fiber 1.7–3.0 NR 1.2–2.8 NR
b
Carbohydrates 60.6–66.6 66.5–69.1 56.5–68.1 57.2c
Total dietary 11.8–14.1 8.8–10.3 7.3–10.2 10.4
fiber
N 6.25.
a
b
Calculated by difference.
c
Only starch.
NR: Not reported.
Table 2 Contents of Minerals (mg/100 g Dry Basis) in Raw Quinoa From the Andean Region
National Institute of Health— Ruales and Nair Miranda, Vega-Gálvez, Quispe-Fuentes, Nascimento et al.
Reference Perú (2009) (1993) et al. (2012) (2014)
Variety/ Sweet quinoa blanca Variety 40,057 Ecotypes: NR
Ecotype Latinreco Ancovinto, Cancosa, Cáhuil, Faro,
Regalona, Villarica
Country of Peru Ecuador Chile Argentina
Origin
Sampling Puno Quito North-altiplano, center, and south Jujuy
Place
Ca 129.5 874 77.1–211.3 49.6
Fe 6.0 81 4.8–7.2 6.16
Mg NR 2620 150.9–166.7 222.1
P 254.5 5350 285.1–526.4 527.6
K NR 1201 1672.8–2325.6 748.6
Na NR 22 17.1–23.4 <LoQ
Zn 3.7 36 2.7–5.0 3.3
Cu NR 10 0.7–1.5 0.66
NR: Not reported. LoQ: limit of quantification.
with celiac disease. Same research group later reported that daily intake of
50 g of quinoa during 6 weeks may be safe for celiac patients (Zevallos
et al., 2014).
The fat contents in Andean quinoa varieties range from 4.1 to 7.1 mg/
100 g EP (Table 1) with higher unsaturated: saturated fatty acid ratios
(4.9–6.2) than most edible oils such as soy-bean oil (3.92), corn oil
(4.65), and olive oil (0.65) (Ruales & Nair, 1993; Valcárcel-Yamani &
Da Silva Lannes, 2012). Linoleic acid (C18:2) is the major unsaturated fatty
acid, followed by oleic acid (C18:1) and linolenic acid (C18:3) whereas
palmitic acid (C16:0) is the predominant saturated fatty acid (Miranda,
Vega-Gálvez, Martinez, et al., 2012; Ruales & Nair, 1993; Villa, Russo,
Kerbab, Landi, & Rastrelli, 2014).
Reported levels of total dietary fiber in quinoa (7.3–14.1 g/100 g EP)
(Table 1) are comparable to those found in wheat (12.2 g/100 g EP;
USDA, 2017) and around 78%–90% of dietary fiber is insoluble and is
mainly composed of galacturonic acid, arabinose, galactose, and xylose
(Lamothe, Srichuwong, Reuhs, & Hamaker, 2015; Repo-Carrasco-
Valencia & Serna, 2011).
Contents of minerals such as calcium, magnesium, and potassium are
superior to cereal grains such as wheat, corn, and rice (Nowak et al.,
2016). As shown in Table 2, Andean quinoa varieties have generally high
concentrations of potassium and phosphorus whereas iron contents
(4.8–81.0 mg/100 g dry basis) are comparable or higher than those found
in wheat (5.0 mg/100 g dry basis), corn (3.0 mg/100 g dry basis), and
rice (0.9 mg/100 g dry basis) (USDA, 2017). Variability on the mineral con-
tent is significantly influenced by the cultivar, origin along with the soil
type, and its mineral composition (Miranda, Vega-Gálvez, Quispe-Fuentes,
et al., 2012).
Vitamins C, B1 (thiamin), B2 (riboflavin), and B3 (niacin) have been
found in several quinoa ecotypes from Chile (Miranda, Vega-Gálvez,
Quispe-Fuentes, et al., 2012) whereas Ruales and Nair (1993) additionally
reported folic acid and vitamin A in quinoa seeds from Ecuador along with a
high content of γ-tocopherol among detected tocopherols with vitamin
E activity. Moreover, γ- and α-tocopherols (797.2 and 721.4 ppm, respec-
tively) have been also detected in Peruvian quinoa seeds (cultivar Huancayo)
(Repo-Carrasco, Espinoza, & Jacobsen, 2003). Levels of vitamin B6 and
total folate found in quinoa can cover the nutritional requirements for chil-
dren and adults and contents of these vitamins along with vitamin B2 are
higher than those from oat and barley ( James, 2009). However, data on
vitamin contents are still limited considering the wide diversity of quinoa
germplasm from Andean countries.
Quinoa also contains bitter compounds called saponins which are con-
centrated in the seed hull and should be removed before consumption
( Jacobsen et al., 2003). Contents may vary between 0.02% and 1.1% and
depending on the concentrations, there are sweet and bitter quinoa varieties
(Bastidas et al., 2016; Filho et al., 2017). Some treatments have been pro-
posed to eliminate saponins without significantly affecting its nutritional
value (Filho et al., 2017). In this regard, an infant food from quinoa
(debittered for saponin elimination) showed a good nutritional quality
and increased the insulin-like growth factor-1 (IGF-1, low levels are related
to malnutrition) in undernourished preschool children after 15 days of intake
compared to the control group (Ruales, Grijalva, Lopez-Jaramillo, &
Nair, 2002).
3.1.2 Cañihua and Andean Amaranth

Cañihua (C. pallidicaule Aellen) and Andean amaranth (A. caudatus L.)
(Amaranthaceae family) are pseudocereals similarly as quinoa and are native
from the Andean region. These crops can be adapted to varying environ-
mental conditions and can be grown in poor and rocky soils (National
Research Council, 1989). Cañihua plant has shown to be even more resis-
tant to frosts and droughts than quinoa (National Research Council, 1989).
Compared to quinoa, no official data exist related to amaranth and cañihua
production but both crops are traditionally cultivated in Peruvian and Boliv-
ian altiplano and represent important sources of proteins and other nutritious
compounds for people from this area (FAOSTAT, 2014).
The macronutrient composition of cañihua and Andean amaranth seeds
is shown in Table 3. Their proximate composition is quite similar to that
shown for quinoa (Table 1), though the nutrient profile and contents
depend on the origin along with the influence of genetic, agronomic,
and environmental factors as previously stated (Miranda, Vega-Gálvez,
Quispe-Fuentes, et al., 2012).
Protein contents in cañihua and Andean amaranth (10.4–13.4 g/100 g
EP) (Table 3) are comparable to values found in conventional cereals such
as wheat (11.31 g/100 g EP, USDA, 2017) and higher than those reported for
rice and yellow corn (6.61 and 9.42 g/100 g EP, respectively; USDA, 2017).
The quality of cañihua protein is comparable to that of quinoa and is parti-
cularly rich in sulfur amino acids, lysine, and aromatic amino acids which
are higher than levels recommended by the WHO/FAO/UNU (2007)
Table 3 Proximate Composition (g/100 g Edible Portion) in Raw Cañihua and Amaranth Seeds From the Andean region
Pseudocereal Cañihua (Chenopodium pallidicaule Aellen) Amaranth (Amaranthus caudatus L.)
Reference Repo-Carrasco-Valencia, de La Cruz, Villa et al. Repo-Carrasco-Valencia, Encina, Binaghi, Nascimento
Alvarez, and Kallio (2009) (2014) Greco, and Ronayne de Ferrer (2010) et al. (2014)
Variety Cupi NR Centenario NR
Country of Peru Peru Peru Argentina
Origin
Sampling Puno Ayabaca- Lima Jujuy
Place Piura
Water 10.4 10.8 11.3 10.5
Ash 4.5 3.1 1.6 2.9
a
Total protein 12.9 12.8 10.4 13.4
Total fat 5.1 7.0 6.7 6.4
Crude fiber 10.1 6.3 NR NR
b c b
Carbohydrates 57.0 59.9 54.9 55.3
Total dietary 22.6 NR 5.1 11.3
fiber
a
N 6.25.
b
c
Only starch.
NR: Not reported.
for adults (Villa et al., 2014). The amaranth protein has also a well-balanced
amino acid pattern but some references have indicated leucine as a limiting
amino acid according to its chemical score (Valcárcel-Yamani & Da Silva
Lannes, 2012). The most dominant proteins in Amaranthus spp. and
Chenopodium spp. are albumins and globulins with low or not detected
levels of prolamins ( Janssen et al., 2017). Therefore, these pseudocereals
may also represent nutritionally important alternatives to rice or other
gluten-free cereals for the design of diets targeted for celiac people (Mota
et al., 2016). The validation of this property through human studies as
investigated for quinoa would be important to establish the safe dose for
daily intake among celiac population (Zevallos et al., 2012).
The fat contents (5.1–7.0 mg/100 g EP) (Table 3) in cañihua and Andean
amaranth are similar to values shown for quinoa (Table 1) and are higher
than those found in common cereals such as rice and wheat (Nascimento
et al., 2014). The fat fraction of cañihua is mostly composed by unsaturated
fatty acids (71.4%) with high levels of linoleic acid (39.2%), followed by
oleic acid (29.8%) and the saturated palmitic acid (22.8%) (Villa et al.,
2014). Similarly, A. caudatus fat is rich in unsaturated fatty acids (73%) being
linoleic (47.8%) and oleic acids (23.7%) the most relevant (Alvarez-Jubete,
Arendt, & Gallagher, 2009).
Andean amaranth and cañihua have shown higher contents of calcium,
iron, and phosphorus than rice and comparable ranges of iron to those of
wheat (Nascimento et al., 2014; Villa et al., 2014). In addition, higher iron
contents than quinoa have been reported in these two pseudocereals (Repo-
Carrasco-Valencia, Encina, et al., 2010). In a study with A. caudatus culti-
vated in Argentina, authors highlighted that this grain exhibited a high
contribution to the recommended daily dietary intake for minerals, and it
was even higher than quinoa in case of calcium, iron, and zinc
(Nascimento et al., 2014). Thermal processing such as boiling can negatively
affect the mineral content of these grains although the in vitro availability is
improved (Repo-Carrasco-Valencia, Encina, et al., 2010).
Unlike quinoa, cañihua and amaranth seeds do not require prior treat-
ment before their use ( Jacobsen et al., 2003). However, Rastrelli, De
Simone, Schettino, and Dini (1996) detected triterpene saponins in cañihua
seeds that may give bitter taste to the grain but in a lesser extent than in the
case of quinoa. Repo-Carrasco-Valencia, Encina, et al. (2010) have indi-
cated that cañihua is commonly consumed without the outer seed coat
where saponins are more concentrated. This reduces the total dietary fiber
content from 25 to 12.6 g/100 g dry basis; however along with amaranth and
quinoa, these pseudocereals are still good sources of dietary fiber (Repo-
Carrasco-Valencia et al., 2009; Repo-Carrasco-Valencia, Encina,
et al., 2010).
Regarding vitamins, major tocopherols found in a Peruvian cañihua
(white cultivar) were γ-tocopherol (788.4 ppm) followed by α-tocopherol
(726.0 ppm) (Repo-Carrasco et al., 2003). In contrast, β- and α-tocopherols
(47.8 and 61.6 ppm, respectively) were major vitamin E isomers found in an
Ecuadorian sample of A. caudatus, and γ- and δ-tocopherol were detected in
lower concentrations (Bruni et al., 2001). Vitamins C and those of
B complex such as niacin, niacinamide, pyridoxine, and riboflavin have been
reported in raw seeds of A. caudatus although the analyzed amaranth sample
was grown in an environment different from the Andean area (Gamel,
Linssen, Mesallam, Damir, & Shekib, 2006). Still limited information exists
about the vitamin contents and profiles of cañihua and amaranth, specifically
the caudatus Andean species.
In consideration to the high nutritional potential of Andean
pseudocereals for the design of gluten-free diets, investigations on the var-
iability of their macro- and micronutrients as affected by the different tech-
nological processes, and the potential interactions with the other food
ingredients are critical and necessary.
3.1.3 Andean Lupin or Tarwi

The genus Lupinus (Fabaceae family) has numerous species but only five are
the most cultivated (Lupinus albus, Lupinus angustifolius, Lupinus luteus,
Lupinus mutabilis, and Lupinus consentenii) in different regions worldwide
such as northern Europe, the Australian plains, and the Andean highlands
(Putnam, 1991). The species L. mutabilis Sweet also known as “tarwi,”
“chocho,” or Andean lupin is an endemic legume traditionally cultivated
in Ecuador, Peru, and Bolivia and first domesticated in northern Peru, most
likely in the Cajamarca region (Atchinson et al., 2016). L. mutabilis has been
used for soil enrichment and as a food crop in the Andean region (Gross
et al., 1988).
The proximate composition of Andean lupin samples compared with that
of soybean (Glycine max) is shown in Table 4. L. mutabilis has on average higher
contents of protein and fat than Mediterranean lupin species and values are
comparable to those found in soybeans (Salmanowicz, 1999). A recent review
has shown that protein contents in L. mutabilis ranged from 32.0 to 52.6 g/
100 g dry basis, similarly fat contents varied from 13.0 to 24.6g/100g dry basis
Table 4 Proximal Composition (g/100 g Dry Basis) in Raw Lupinus mutabilis Seeds and Comparison With Soybean Composition
Legume Lupinus mutabilis Seeds Raw Soybean
(Glycine max)
Reference Carvajal-Larenas, Van Boekel, Koziol, Nout, Gross et al. Carvalho, Chaves, and Pinto USDA (2017)
and Linnemann (2014) (1988) Ricardo (2005)
Cultivar NR cv. Inti cv. Potosi NR
Country of Ecuador Chile NR NR
Origin
Sampling Cotopaxi Province Temuco NR NR
Place
Ash 5.0 NR 4.2 3.4
a
Total protein 41.4 51.0 45.1 39.9
Total fat 23.4 16.0 18 21.8
Crude fiber NR NR 9.4 NR
b c b
Carbohydrates 26.6 14.8 23.3 33.0b
Total dietary NR NR NR 10.2
fiber
N 6.25.
a
b
c
Only oligosaccharides.
NR: Not reported.
(Carvajal-Larenas, Linnemann, Nout, Koziol, & Van Boekel, 2016). These

variations are mainly associated with genetic and agronomic factors. In this
sense, Carvalho et al. (2005) determined that cultivating L. mutabilis under
water stress reduced by half the fat content but increased the total carbohy-
drates. The protein is mainly composed of globulins followed by albumins,
and β-conglutin along with α-conglutin and γ-conglutin were found as the
major globulin storage proteins in L. mutabilis (Santos, Ferreira, & Teixeira,
1997). All essential amino acids recommended by WHO/FAO/UNU
(2007) are present in Andean lupin seeds, the average content of lysine is high
but sulfur amino acids (cystine and methionine) are limiting (Carvajal-
Larenas et al., 2016; Gross et al., 1988). This has been reflected in the low
protein quality (measured by the protein efficiency ratio) of L. mutabilis in
comparison with quinoa and casein proteins; however, this could be over-
come by mixing Andean lupin with other high-protein quality Andean crops
such as quinoa (Gross, Koch, Malaga, de Miranda, & Trugo, 1989).
L. mutabilis proteins have shown good technofunctional properties such
as oil absorption capacity, emulsifying capacity, foam stability, and gelation
properties comparable or even higher than those reported for soybean pro-
teins; therefore, Andean lupin proteins may potentially be used to partly
replace soybean proteins in targeted industrial applications to improve spe-
cific functional characteristics (Moure, Sineiro, Domı́nguez, & Parajó, 2006;
Sathe, Deshpande, & Salunkhe, 1982).
The fat fraction is mainly composed by unsaturated fatty acids (80%)
especially oleic and linoleic acids (42% and 39% of total fatty acids, respec-
tively) whereas the content of linolenic acid is low (3% of total fatty acids)
(Gross et al., 1988).
Andean lupin is also an important source of macro- and microelements.
Ranges of potassium (1130–1400 mg/100 g dry basis), phosphorus
(600–880 mg/100 g dry basis), magnesium (240–330 mg/100 g dry basis),
calcium (120–180 mg/100 g dry basis), iron (5.0–7.3 mg/100 g dry basis),
and zinc (3.4–3.6 mg/100 g dry basis) have been reported in a review by
Carvajal-Larenas et al. (2016). Higher levels of potassium, phosphorus, mag-
nesium, and zinc have been found in raw seeds of L. mutabilis when com-
pared with other leguminous species such as pea (Pisum sativum), lentils (Lens
culinaris), and common beans (Phaseolus vulgaris L.) whereas iron contents
were comparable to those shown by above legumes (Grela, Samoli nska,
Kiczorowska, Klebaniuk, & Kiczorowski, 2017).
Carbohydrates are mainly constituted by oligosaccharides with stachyose
as the major α-galactoside (40%–45% of total oligosaccharides), followed by
raffinose, sucrose, and verbascose (Carvalho et al., 2005; Gross et al., 1988).
These compounds are not digested by human enzymes and are flatulence-
causing agents (Piotrowicz-Cieslak, Michalcyk, & Gorecki, 2004).
Studies related to the determination of vitamin profiles and contents of
Andean lupin are limited. Boschin and Arnoldi (2011) only detected
γ-tocopherol in a cultivar from Ecuador (Andino 450) and levels
(10.3 mg/100 g of seed weight) were generally higher than contents found
in other Lupinus species (L. albus and L. angustifolius). The vitamin E activity
of L. mutabilis was lower (1.03 mg/100 g of seeds) than soybean (1.83 mg/
100 g of seeds) but higher than Phaseolus species (0.30–0.31 mg/100 g of
seeds) and L. culinaris (0.62 mg/100 g of seeds).
L. mutabilis has some nutritional advantages when compared to soybean
since small amounts of trypsin inhibitors, phytates, tannins, and saponins have
been detected in this legume (Schoeneberger, Gross, Cremer, & Elmadfa,
1982). However, L. mutabilis has also toxic and bitter compounds such as
quinolizidine alkaloids (lupanine, D-lupanine, sparteine, 3-hydroxylupanine,
13-hydroxylupanine, and 4-hydroxylupanine as major alkaloids) with total
values that may range from 0.007 to 4.5 g/100 g dry basis (Carvajal-Larenas
et al., 2016). Several treatments have been proposed for eliminating alkaloids
and oligosaccharides in Andean lupin such as breeding techniques, germina-
tion processes, soaking in boiling water followed by a continuous water wash
for up to 15h, and in general these treatments increased the nutritional value
uemes-Vera, Peña-
of L. mutabilis seeds and derivatives (Gross et al., 1988; G€
Bautista, Jimenez-Martı́nez, Dávila-Ortiz, & Calderón-Domı́nguez, 2008;
Villacres et al., 2015).

Andean grains have been cultivated since pre-Inca’s times; therefore ancient
civilizations had already developed different techniques and treatments for
their preparation and consumption. After the Spanish conquest, most of
the indigenous crops including Andean grains were banned and replaced
by European crops such as wheat, oat, and rye (National Research
Council, 1989). However, their cultivation and consumption still remained
in some places and now all these native grains are gaining an increasing inter-
est and demand worldwide due to their important nutritional and functional
properties.
Some traditional preparations and uses of Andean grains by indigenous
population are shown in Table 5.
Table 5 Traditional Preparations and Uses of Andean Grains by Indigenous Population
Andean Grain Preparation/Use
Quinoaa Quispiño: quinoa dough made from raw flour and animal fat and cooked
Pesqhe: salted porridge of sweet quinoa seeds
Tecte: white fermented beverage of quinoa
Tactte: cake with quinoa flour and fried in animal fat
Quinoa milk: vegetable milk
Pop-quinoa: puffed quinoa product
Cooked quinoa and applied in soups, as rice replacer, in juices with fruits
Quinoa flour for applications in soups and baked products
Andean lupin or Tarwib Debittered and boiled beans and consumed in salads and soups
Chicha: fermented beverage of tarwi
Puree with lupin beans
Tarwi bean sauce
Tarwi milk: soaked, cooked, and mashed tarwi
Tarwi flour for baking applications
Cañihua and Andean amaranthc Kañiwako: toasted and milled flour of cañihua
Pop-kiwicha: puffed amaranth product consumed directly or for the formulation of snack bars
Cañihua and amaranth flours for applications in soups, juices, and baked products
a
Jacobsen et al. (2003).
b
FAO (2013).
c
Repo-Carrasco-Valencia, Hellstr€
om, Pihlava, and Mattila (2010).
Many of these preparations and culinary applications were inherited from

ancient times and this traditional knowledge has been the base for the inves-
tigation of technologies targeted to the processing of Andean grains at indus-
trial scale. In this regard, several reports have shown the variability of
applications of Andean grains for the formulation of high-nutritious foods
such as extruded products, cereal-based snacks and bars, and in case of
pseudocereals for the design of gluten-free foods (Diaz et al., 2013, 2017;
Repo-Carrasco-Valencia et al., 2009; Valcárcel-Yamani & Da Silva
Lannes, 2012).

3.3.1 Quinoa
Several health-relevant bioactive compounds such as polyphenols, caroten-
oids, phytoecdysteroids, and saponins have been identified in quinoa.
Table 6 summarizes the reported bioactive compounds and associated
potential functional properties of Andean quinoa.
Polyphenols such as phenolic acids (30.3–59.7 mg/100 g dry basis)
and flavonoids (36.2–72.6 mg/100 g dry basis) have been detected in several
quinoa ecotypes and varieties from Peru, and major phenolic acids were fer-
ulic, vanillic, and p-coumaric acids (Table 6). Quercetin and kaempferol
were the major flavonoids detected in all quinoa ecotypes/varieties whereas
myricetin and isorhamnetin were found only in some varieties (Repo-
Carrasco-Valencia, Hellstr€ om, et al., 2010). Reported phenolic acid con-
tents were similar to levels found in oat, barley, corn, and rice but lower
compared with common cereals like wheat and rye (Repo-Carrasco-
Valencia, Hellstr€ om, et al., 2010). Alvarez-Jubete, Wijngaard, et al.
(2010) and Ranilla, Apostolidis, et al. (2009) also reported quercetin and
kaempferol glycosides as main phenolic compounds in quinoa seeds from
Bolivia and Peru, respectively, and phenolic contents were correlated with
the in vitro antioxidant capacity. Flavonoid contents tended to increase with
sprouting but a significant reduction was produced when quinoa was used in
a bread formula (Alvarez-Jubete, Wijngaard, et al., 2010). However, pheno-
lic variability seems to be related to the type of heat treatment since extrusion
has shown to increase the phenolic content and the ABTS antioxidant
capacity (Repo-Carrasco-Valencia & Serna, 2011). Isoflavones such as daid-
zein (0.7–1.15 mg/100 g dry basis) and genistein (0.05–0.25 mg/100 g dry
basis) have been also reported in Chilean ecotypes with varying concentra-
tions depending on the genetic factors and the geographical origin (Lutz
et al., 2013).
Table 6 Bioactive Compounds and Reported Bioactivity in Quinoa Seeds or Derivatives
Variety/Ecotype and Origin Bioactive Compounds Used Model and Reported Reference
Treatment Bioactivity
Raw seeds Puno-Peru Phenolic acids: Repo-Carrasco-
Ecotypes: 03-21-1181, Cusco-Peru Caffeic acid, ferulic acid, Valencia, Hellstr€
om,
03-21-0093, Witulla, Roja (only p-coumaric acid, et al. (2010)
coporaque, Huaripongo, commercial p-hydroxybenzoic acid,
Ccoito samples) vanillic acid
Varieties: INIA-415, Flavonoids:
Pasankalla, Salcedo INIA and Quercetin, kaempferol,
two commercial samples myricetin, isorhamnetin
Preprocessed seeds for Bolivia Phenolic acids: In vitro hydrophilic Alvarez-Jubete,
partially removing saponins Protocatechuic acid, vanillic antioxidant capacity (DPPH Wijngaard, Arendt, and
acid derivative and FRAP assay) Gallagher (2010)
Flavonoids:
Quercetin glycosides,
kaempferol glycosides
Pretreated seeds for saponin Chile Daidzein Lutz, Martı́nez, and
removing (North- Genistein Martı́nez (2013)
Ecotypes: R49, UdeC9, altiplano,
BO78 center, and
south)
Autoclaved seeds Arequipa- Quercetin derivatives In vitro hydrophilic Ranilla, Apostolidis,
Red quinoa Peru antioxidant capacity (DPPH Genovese, Lajolo, and
assay) Shetty (2009)
In vitro inhibition of the
hyperglycemia-relevant
α-glucosidase enzyme
Raw seeds Andean Carotenoids: trans-lutein, In vitro lipophilic antioxidant Tang et al. (2015)
White, red, black mountains of zeaxanthin capacity (DPPH, FRAP,
South Tocopherols (γ, α, β, δ) and ORAC, and PCL: photo-
America tocotrienols (α, β) induced chemiluminescence
of luminol assays)
Raw seeds Peruvian Betalains: amaranthin, In vitro hydrophilic Escribano et al. (2017)
29 quinoa varieties with altiplano and isoamaranthin, betanin, antioxidant capacity (FRAP,
different colors valleys isobetanin, dopazanthin, ABTS, and ORAC assays)
dopamine-derived
betaxanthin, proline-derived
betaxanthin
Saponins
Pretreated seeds for saponin Bolivia Phytoecdysteroids: In vivo: diet-induced obese, Graf et al. (2014)
removing 20-hydroxyecdysone, hyperglycemic mice model
Red quinoa makisterone A, epi- Decrease of fasting blood
makisterone A, 24(28)- glucose
dehydromakisterone A,
makisterone C
Flavonoids: quercetin
trisaccharides
Unsaturated fatty acids and
amino acids
Quinoa seeds in a formula Bolivia Hypocholesterolemic and In vivo: male Wistar rats Pasko, Barton, et al.
Treatments: LDL lowering effects likely Decrease of serum total (2010)
Control group (C) associated to protein, cholesterol, LDL, Pasko, Zagrodzki,
Continued
Table 6 Bioactive Compounds and Reported Bioactivity in Quinoa Seeds or Derivatives—cont’d
Variety/Ecotype and Origin Bioactive Compounds Used Model and Reported Reference
Treatment Bioactivity
Control group with 31% of unsaturated fatty acids, triglycerides in Q group Barto
n, Chłopicka, and
fructose (F) tocopherols, dietary fiber and Decrease of glucose levels and Gorinstein (2010)
Group fed with quinoa (Q) saponins. plasma total protein in
Group fed with quinoa and Q group
31% of fructose (QF) No inhibition of triglyceride
Time: 5 weeks increase in QF group
Decrease of malondialdehyde
in plasma (Q and QF groups)
Moderate protection of
antioxidant enzymes of
plasma, heart, kidney, testis,
lung, and pancreas
Quinoa flakes in a cereal bar Bolivia Effects likely related to quinoa Human study: 22 subjects Farinazzi-Machado,
Consumption of cereal bars bioactive composition: (18–45 years old) Barbalho, Oshiiwa,
with quinoa for 30 days, vitamin E, phytosteroids, Decrease of plasma total Goulart, and Pessan
equivalent to the intake of polyphenols, saponins, cholesterol, triglycerides, and Junior (2012)
19.5 g of quinoa/day minerals LDL
56.7% of subjects with
reduction of blood glucose
levels, 42.2% of subjects with
reduction of body weight,
and 40.7% with reduced
blood pressure
Quinoa flakes (QF) NR Effects possibly associated to Human study: prospective, De Carvalho et al.
Treatments: quinoa dietary fiber, protein, double-blind study with (2014)
Group 1: consumption of vitamin E, and different 35 women in menopause
25 g of QF bioactive compounds Reduction of plasma total
Group 2. Consumption of cholesterol, LDL, and
25 g of corn flakes triglycerides only in QF
Time: 4 weeks group
Modulation of plasma
oxidative markers: decrease of
TBARS (thiobarbituric acid
reactive substances) and
increase of GSH (reduced
glutathione) only in QF
group
NR: Not reported.
Some quinoa varieties may exhibit yellow–orange and red–violet colors

in their grains and this is mostly associated to their betalain (betacyanins and
betaxanthins) profiles and contents which have shown a strong correlation
with the in vitro hydrophilic antioxidant capacity (Escribano et al., 2017).
Carotenoids such as trans-lutein and zeaxanthin along with tocopherols
and tocotrienols have been detected in black, white, and red quinoa varieties
exhibiting a high correlation with the in vitro lipophilic antioxidant capacity
(Tang et al., 2015). Other quinoa bioactives such as polyhydroxylated ste-
roids named phytoecdysteroids and saponins have been related to hypogly-
cemic and antiinflammatory activities, respectively (Graf et al., 2014; Yao,
Yang, Shi, & Ren, 2014).
Quinoa protein has been described as a source of short-chain bioactive
peptides with in vitro inhibitory effects against hyperglycemia and
hypertension-relevant enzymes (Aluko & Monu, 2003; Vilcacundo,
Martı́nez-Villaluenga, & Hernández-Ledesma, 2017). Moreover, peptides
derived from quinoa protein hydrolysis decreased risk markers of cardiovas-
cular disease by exhibiting antihypertensive activity and antiinflammatory
effects on human umbilical vein endothelial cells (Ravisankar, Gutierrez,
Chirinos, & Noratto, 2015).
According to in vivo studies, short-term consumption of quinoa has
shown moderate protective effects linked to high fructose consumption
by reducing plasma lipid peroxidation and enhancing the antioxidant capac-
ity of blood (Pasko, Barton, et al., 2010; Pasko, Zagrodzki, et al., 2010). In
addition, the potential reduction of risk factors related to cardiovascular dis-
eases and protection against effects of oxidative stress have been also reported
in some human studies (De Carvalho et al., 2014; Farinazzi-Machado
et al., 2012).
3.3.2 Cañihua and Andean Amaranth

In addition to the nutritional potential of cañihua and Andean amaranth,
several studies have highlighted the importance of these pseudocereals as
potential sources of biologically active compounds such as polyphenols,
squalene, phytosterols, and bioactive peptides.
Simple polyphenols such as protocatechuic acids have been detected in
Peruvian A. caudatus grains and total phenolic contents (21.2 mg gallic acid
equivalents/100 g dry basis) were lower than quinoa and wheat (71.7 and
53.1 mg gallic acid equivalents/100 g dry basis, respectively) (Alvarez-
Jubete, Wijngaard, et al., 2010). Total phenolic acids in several cañihua
and amaranth ecotypes from Peru ranged from 18.3 to 40.1 and from
16.8 to 32.9 mg/100 g dry basis, respectively. Major phenolic acid in cañihua
was ferulic acid whereas vanillic, p-hydroxybenzoic, caffeic, and p-coumaric
acids were detected in minor concentrations (Repo-Carrasco-Valencia,
Hellstr€om, et al., 2010). Additionally to these phenolic acids, protocatechuic
and sinapic acids were found in Andean amaranth samples (Repo-Carrasco-
Valencia, Hellstr€ om, et al., 2010). Cañihua ecotypes also had high flavonoid
contents (46.1–144.3 mg/100 g dry basis) (mainly quercetin) comparable to
those found in quinoa (36.2–72.6 mg/100 g dry basis) whereas no flavonoids
were found in amaranth (Repo-Carrasco-Valencia, Hellstr€ om, et al., 2010).
Similarly, Ranilla, Apostolidis, et al. (2009) reported high quercetin deriv-
atives concentrations in an autoclaved cañihua sample from Peru along with
high in vitro antioxidant capacity whereas the phenolic content and antiox-
idant capacity of Peruvian amaranth (autoclaved) was low.
Squalene, a triterpenoid and key intermediate in cholesterol biosynthesis
with potential for cholesterol reduction, has been detected in two varieties
(Oscar blanco and Victor red) of A. caudatus grown in Bolivia and greatly
contributed to the in vitro lipid peroxidation inhibitory activity (Conforti
et al., 2005). In addition, methanol extracts of these amaranth varieties
exhibited in vitro antidiabetic activity through the inhibition of the
α-amylase enzyme (28%–50.5% at 25 μg/mL of concentration). Among
phytosterols detected in the unsaponifiable fat fractions of cañihua and ama-
ranth, stigmasterol has been reported as the main sterol in cañihua whereas
chondrillasterol was the most abundant in Andean amaranth (Bruni et al.,
2001; Villa et al., 2014).
Some animal studies have revealed the antihypercholesterolemic effects
with potential for the prevention of coronary heart diseases of A. caudatus
grains or its plant derivatives. The intake of a diet including extruded
A. caudatus flour (Oscar blanco variety, from Peru) for 21 days significantly
decreased the total cholesterol and LDL-cholesterol concentrations while
the triglycerides contents were 50% lower in hypercholesterolemic rabbits
compared to the control treatment (Plate & Arêas, 2002). Such positive
effects were likely attributed to the dietary fiber, squalene, and protein in
amaranth (Plate & Arêas, 2002). In another study, hydroalcoholic extracts
of A. caudatus (stems, leaves, and flowers) showed antihypercholesterolemic
and antiatherogenic effects associated with the reduction of serum lipids and
oxidative stress markers in high cholesterol-fed rabbits (Kabiri, Asgary, &
Setorki, 2011).
A comprehensive review of the potential functionality of amaranth
grains has evidenced that investigations are more focused on other
Amaranthus species such as Amaranthus cruentus, Amaranthus hypochondriacus,

and Amaranthus esculentus (Caselato-Sousa & Amaya-Farfán, 2012). For
instance, proteins extracted from A. hypochondriacus have shown in vivo
antithrombotic properties whereas its derived peptides have exhibited
angiotensin I-converting enzyme-inhibitory and antioxidant activities
(Sabbione, Rinaldi, Añón, & Scilingo, 2016; Soriano-Santos & Escalona-
Buendı́a, 2015). Andean A. caudatus and cañihua may show similar func-
tional potential; however, this should be validated with in vivo or human
studies considering that phytochemical profiles are variable depending on
the specie and variety.
3.3.3 Andean Lupin or Tarwi

L. mutabilis Sweet is not only an important source of macronutrients and
micronutrients, but also contains several health-relevant compounds such
as isoflavones, carotenoides, alkaloids, and bioactive peptides with potential
biological properties as shown in Table 7.
Isoflavones, mainly genistein derivatives, have been detected in several
Peruvian cultivars in ranges from 14 to 35 mg genistein/100 g seeds in
FW showing a high correlation with the in vitro antioxidant capacity mea-
sured by the DPPH assay (Ranilla, Genovese, et al., 2009). Andean lupin
shows moderate isoflavone contents compared with those found in soybeans
(57–188 mg/100 g FW), however this legume may represent an interesting
isoflavone source (Genovese, Barbosa, Pinto, & Lajolo, 2007; Ranilla,
Genovese, et al., 2009). Aqueous extracts from autoclaved seeds of
L. mutabilis have shown moderate in vitro inhibition against α-glucosidase
enzyme relevant for hyperglycemia management (19%–26%, at 5 mg of sam-
ple dose) but significantly inhibited the hypertension-relevant angiotensin
I-converting enzyme (ACE-I) (52%, at 5 mg of sample dose) (Ranilla,
Apostolidis, et al., 2009). Same authors suggested that this in vitro activity
was likely related to the protein or thermally released peptides, since no cor-
relation was found with the total phenolic contents (Ranilla, Apostolidis,
et al., 2009).
Little information exists related to the carotenoid profile in L. mutabilis
cultivated in the Andean region. In a study with L. mutabilis from Australia,
lutein has been reported as the major carotenoid, followed by zeaxanthin,
β-carotene, and α-carotene, but in lower levels compared to other Lupinus
species (L. luteus, L. albus, and L. angustifolious) (Wang et al., 2008). Envi-
ronmental growing conditions associated to different geographical origins
are known to influence the phytochemical profile in legumes as in other
Table 7 Bioactive Compounds and Reported Bioactivity in Lupinus mutabilis Sweet Seeds or Derivatives
Variety/Cultivar and Treatment Origin Bioactive Compounds Used Model and Reported Bioactivity Reference
Raw seeds Lima- Isoflavones: genistein and a In vitro hydrophilic antioxidant Ranilla, Genovese, and
Cultivars: SLP-1, SLP-4, SCG-22, Peru genistein derivative capacity (DPPH assay) (cotyledons, Lajolo (2009)
H-6, Andares, Yunguyo Flavonoids: flavone derivative seed coats, and hypocotyls).
(Cotyledons, seed coats and
hypocotyls).
Autoclaved seeds Lima- Total phenolic contents In vitro hydrophilic antioxidant Ranilla, Apostolidis,
Cultivars: SLP-1, H6 Peru capacity (DPPH assay) et al. (2009)
In vitro inhibition of the
hyperglycemia-relevant α-glucosidase
enzyme
In vitro inhibition of the hypertension-
relevant angiotensin I-converting
enzyme
Raw seeds Australia Lutein, zeaxanthin, β-carotene, Wang, Errington, and
Cultivar P26961 and α-carotene Yap (2008)
Raw lupin flour. Line 450 Cotopaxi- Effects likely associated to Human study. Oral intake of lupin Fornasini et al. (2012)
Treatments: Ecuador quinolizidine alkaloids and flour with maximum dose of 3.1 mg/kg
G1: intake of lupin flour by healthy proteins (γ-conglutin) body weight of alkaloids
individuals (n ¼ 16). Decrease of blood and insulin
G2: intake of lupin flour by concentration only in G2
individuals with fasting glucose No effect on G1
(>100 mg/dL) (n ¼ 17). No effect in control group (soy intake)
Control: intake of soy flour
(n ¼ 5–7).
Blood analysis at time 0, 60, and
90 min after oral intake
Continued
Table 7 Bioactive Compounds and Reported Bioactivity in Lupinus mutabilis Sweet Seeds or Derivatives—cont’d
Variety/Cultivar and Treatment Origin Bioactive Compounds Used Model and Reported Bioactivity Reference
Cooked and debittered lupin flour Ecuador Effects likely associated to Human study with volunteers Baldeón, Castro,
(CDL) quinolizidine alkaloids and diagnosed with type-2 diabetes Villacres, Narváez, and
Treatments: proteins (γ-conglutin) Oral intake of CDL or purified Fornasini (2012)
G1: intake of CDL (n ¼ 20) alkaloids with maximum dose of
G2: intake of purified lupin alkaloids 2.5 mg/kg body weight of alkaloids
(n ¼ 10) Decrease of blood glucose in G1 and
Blood analysis at time 0, 60, and G2 at 60 and 90 min.
90 min after oral intake No effect on blood insulin in G1 and
G2
Lupin in a snack (LS) Ecuador Human study with type-2 diabetic Baldeón et al. (2017)
Treatments: patients
G1: double dose of metformin Decrease of blood glucose, glycated
(n ¼ 44) hemoglobin (HbA1c), and total
G2: normal dose of metformin cholesterol in G1.
+10 g of LS Decrease of HbA1c and body mass
Time: intake for 6 months index in G2
plant-based foods (Rosales et al., 2011). Thus, more research is needed for
proper characterization of the carotenoid profile of Andean lupin diversity.
Important beneficial effects of consumption of L. mutabilis on type-2 dia-
betes risk factors have been reported by few human studies. The oral admin-
istration of cooked and debittered lupin flour significantly reduced the blood
glucose levels in individuals diagnosed with type-2 diabetes but in a lesser
extent in subjects with dysglycemia and when consumed raw lupin flour
(Baldeón et al., 2012; Fornasini et al., 2012). Moreover, the intake of a
healthy lupin-based snack with the conventional type-2 diabetes treatment
(with metformin) showed good hyperglycemia control in diabetic individ-
uals (Baldeón et al., 2017). Quinolizidine alkaloids and lupin protein frac-
tions may explain such effects. However, the exact in vivo mechanism
along with the potential interaction of the other health-relevant lupin com-
pounds such as oligosaccharides, dietary fiber, and unsaturated fatty acids still
remains unknown.
The bioactivity of other Lupinus species has been more extensively
reported compared with information available for L. mutabilis. The
antiinflammatory, anticarcinogenic, and antimicrobial activities have been
investigated in L. angustifolius, L. albus, L. campestris, and L. luteus (Del
Carmen Millán-Linares, Bermúdez, del Mar Yust, Millán, & Pedroche,
2014; Lampart-Szczapa et al., 2003; Martı́nez-Villaluenga et al., 2009). Fur-
thermore, the hydrolysis of L. luteus proteins produced bioactive peptides
that showed significant inhibitory activities against the angiotensin
I-converting enzyme relevant for hypertension management (Boschin,
Scigliuolo, Resta, & Arnoldi, 2014). L. mutabilis may also show potential
in relation to above beneficial effects; therefore, more research focused
on the in vitro and in vivo bioactivity of Andean lupin is necessary.
4. TUBERS AND ROOTS

4.1 General Information on Composition and Nutritional
Value
4.1.1 Maca
Maca (Lepidium meyenii Walp.) is an Andean crop belonging to the
Brassicaceae family. It grows between 3500 and 4500 m.a.s.l in the Central
Andean Region of Perú. This area is characterized by arid and rocky soils
with high solar light intensity, strong winds, and temperatures below
zero degrees (Flores, Walker, Guimarães, Bsid, & Vivanco, 2003). It is a
nutritionally valuable native Peruvian plant used in the Andean diet
(Leon, 1964). This plant was domesticated at least 2 centuries ago in the
Andean mountains, where natives used its tubers as food and as a folk
medicine.
Regardless of the maca product, postharvest drying of maca is carried out
in an artisanal way in the production places before other type of processing
(extrusion, milling, extraction–atomization, etc.) is applied. Briefly, envi-
ronmental drying is carried out for approximately 60–90 days in extreme
temperature conditions ranging from 10 to 15°C. These extreme environ-
mental conditions together with the handling procedures during harvest and
postharvest can have a significant effect on the final bioactive compounds
content in maca (Yábar, Pedreschi, Chirinos, & Campos, 2011). Recently,
artisanal plastic houses have been implemented to dry the product with a
significant reduction in the drying time to 20–25 days and better preserva-
tion of some bioactive compounds such as glucosinolates (Campos et al.,
unpublished results).
Fresh maca contains 80% water while dehydrated maca contains 8.87%–
11.6% protein, 1.09%–2.2% lipid, 54.6%–60.0% carbohydrates (23.4%
sucrose, 1.55% glucose, 4.56% oligosaccharides, and 30.4% polysaccharides),
8.23%–9.08% fiber, 4.9%–5.0% ash, and an energy content of 663 kJ/100 g
(Dini, Migliuolo, Rastrelli, Saturnino, & Schettino, 1994; Valentová et al.,
2006; Yu & Jin, 2004). It contains a good amino acid profile as compared to
the FAO–WHO standard. It contains 18 amino acids and 7 are essential
(no tryptophan present). A good ratio of saturated/unsaturated fatty acids
(0.76) is present in maca. The phytosterol fraction is composed of 45.5%
β-sitosterol, 27.3% campesterol, 13.6% ergosterol, 9.1% brasicasterol, and
4.5% Δ7,22 ergostadienol. A good content of minerals such as Fe, Mn,
Cu, Zn, Na, K, and Ca with values of 16.6, 0.8, 5.9, 3.8, 18.7, 2050,
and 150 mg/100 g DW, respectively, is reported (Dini et al., 1994).
Maca biodiversity is less compared to other Andean crops such as yacon,
mashua, oca, olluco, etc. (Fig. 3). Thirteen different ecotypes presenting
color differences (white, cream, yellow, pink, red, violet, purple, gray,
and back) have been reported. The most commercial biotype corresponds
to the yellow one. Red maca contains a higher protein and potassium con-
tent but less reducing sugars, Fe and riboflavin than the black one while the
yellow one contains intermediate values (Yllescas, 1994). The color of the
different ecotypes is correlated to the content of bioactive compounds and
biological activity (Clement et al., 2010; Gonzales et al., 2006).
4.1.2 Yacon
Yacon (Smallanthus sonchifolius) is an Andean crop from the humid moun-
tains of Peru and Bolivia and it got extended to the north and south of
Fig. 3 Andean roots and tubers: (A) maca, (B) oca cultivars, (C) olluco, (D) mashua cul-
tivars, and (E) yacon roots.
the humid Andean clives and to the dry inter Andean valleys even up to the
coast. Its original habit corresponds to highlands from the south of Colombia
to the north of Argentina at altitudes of 1800 and 2800 m.a.s.l in temperate
hilly climates. It easily adapts to a diversity of climates and soils from sea level
up to 3500 m.a.s.l (Seminario, Valderrama, & Manrique, 2003).
Yacón production in the Andean region and other countries has
increased due to the presumed medicinal properties of both roots and leaves
(Genta, Cabrera, Grau, & Sanchez, 2005). Yacon roots have a high water
content of 83%–90%, followed by its carbohydrate content representing
90% of the DW of the roots. Fructooligosaccharides (FOS) represent
50%–70% of the carbohydrates and the rest is composed of sucrose, fructose,
and glucose (Asami, Minamisawa, Tsuchiya, et al., 1991; Campos et al.,
2012). Yacon FOS are of low degree of polymerization (2–10) with a glu-
cose molecule in the nonreducing extreme. Its FOS and sugar composition
are dependent on genetic diversity and pre- and postharvest factors.
Very few studies exist regarding the biodiversity of yacon. Grau and Rea
(1997) reported that most of the Yacon germoplasm diversity can be found
in the eastern Andean slopes of Peru and Bolivia. Mansilla, López, Blas,
Chia, and Baudoin (2006) reported the molecular variability of a Peruvian
collection of 30 accessions and concluded that most of the variability is found
in the central region of Peru. Lebeda et al. (2003) analyzed 25 genotypes
cultivated in Checz Republic in terms of morphological, weight, incidence
of plagues and pytopathologies, DNA content, polymorphism of isoen-
zymes, sugars, FOS, and phenolic compounds in leaves. A significant
variation in tuber shape, weight, and content of glucose, fructose, sucrose,
oligofructans, and phenolics was found, the same was concluded for DNA
content. The phenolic compounds and FOS content are dependent on
botanical origin, morphological traits, and polymorphisms of individual
genotypes (Lachman, Fernández, Viehmannová, Šulc, & Èepková, 2007).
Campos et al. (2012) found great variability in the contents of bioactive
compounds and antioxidant activity of 35 yacon accessions. Overall, among
the 35 accessions of yacon, the FOS content was inversely correlated with
reducing sugars, while total phenolic content and antioxidant capacity were
positively correlated.
Yacon roots contain important quantities of phenolic compounds.
Campos et al. (2012) reported values of 7.9–30.8 mg of chlorogenic acid
equivalent (CAE) g1 DM while Yan et al. (1999) reported values close
to 38 mg CAE g1 DM and values of 5.7–3.5 mg of gallic acid equivalent g1
DM. Main identified antioxidant phenolics correspond to chlorogenic and
caffeic acids and their derivatives. A high correlation between AC and TPC
(r ¼ 0.89, P < 0.01) was found (Campos et al., 2012), being indicative that
phenolic compounds are mainly responsible for the antioxidant capacity
of yacon roots.
4.1.3 Oca
Oca (Oxalis tuberosa Molina), also known as oqa, apilla, uncha, ibia, and
cuiba, is a tuber domesticated in the Andean region at altitudes between
2800 and 3000 m.a.s.l (Roca, Ynouye, Manrique, Arbizu, & Gomez,
2007). Oca is mainly grown in the highlands of Ecuador, Peru, and Bolivia,
but is also found in some parts of Chile, Argentina, Colombia, and
Venezuela, and it has been introduced in New Zealand over a century
ago (National Research Council, 1989). The shape of the tuber ranges from
ovoid to cylindrical and claviform forms and its surface color varies from
white to cream, yellow, orange, pink, and purple. More than 1000 acces-
sions exist.
Oca present high water content (>80%) and a very low energy density in
the fresh material. The nutritional value of oca is comparable to potato, but
variation in nutritional properties among genotypes exists. Oca tuber is con-
sidered as a good source of carbohydrates (83%–88.8% of DW; or 10.41% of
FW) and its protein, fat, and fiber contents ranged from 3.0% to 8.4%, 0.5%
to 1.0%, and 1.4% to 5.1% DW, respectively; with a starch content of 56.8%
DW and an rate of insoluble: soluble fiber of 23.6:1. Also, oca tuber presents
as predominant sugars, sucrose, followed by glucose (21% and 3.6%, DW)
and it is a good source of calcium and iron (130.1 and 98.45 mg/100 g, DW)
(Flores et al., 2003; Gross et al., 1989; King & Gershoff, 1987; León-
Marroú, Villacorta González, & Pagador Flores, 2011; Valcárcel-Yamani,
Rondán-Sanabria, & Finardi-Filho, 2013). Gross et al. (1989) reported that
the native Andean tuber: oca, ulluco, and mashua constitute sources of
highly digestible carbohydrates. Oca starch presents ellipsoidal and oval
granules forms and, amylose contents, granule size, and gelatinization tem-
perature that varies between 22.4% and 33.0%, 10 and 100 μm, and 60.8
and 64.0°C, respectively (Cruz, Ribotta, Ferrero, & Iturriaga, 2016;
Hernández-Lauzardo, Mendez-Montealvo, Velázquez del Valle, Solorza-
Feria, & Bello-Perez, 2004; Valcárcel-Yamani et al., 2013). Flores et al.
(2003) reported antinutritional factors such as soluble oxalates which are
degraded by solar exposure of the tubers for few days. Levels of oxalates
in oca ranging from 80 to 220 mg soluble oxalate/100 g FW and concentra-
tion up to 500 μg kg1 are responsible of the bitter taste of the tuber
(Sangketkit, Savage, Martin, Mason, & Vanhanen, 1999; Savage,
Mason, & Vanhanen, 2008). One hundred and sixty-nine cultivars of oca
from the International Potato Center germoplasm bank were evaluated
and the content of oxalates corresponded to 62.7–430.9 mg/100 g FW,
being most of them soluble (48.2 and 396.3 mg/100 g. FW). Sun exposure
for 8 days resulted in a decrease of more than 50% of the oxalate content
(Campos et al., unpublished results). Oxalate content of the oca tubers
remains stable by boiling or steaming, but increased by baking because of
loss of moisture content from the tubers. Bioavailability of oxalate in baked
oca appears to be similar to that in spinach, but the bioavailability varies
among individuals and depends on other constituents of a combined meal
(Albihn & Savage, 2001).
4.1.4 Ulluco
Ulluco (Ullucus tuberosus Caldas) (or olluco, ulluku, melloco, chigua, ruba,
papa lisa o lisas, ulluma, and ulluca) is another important Andean crop. Its
distribution in the Andean Region is similar to oca tuber. It commonly
grows at altitudes of 2800–3800 m.a.s.l, in places somehow protected of
low temperatures but this tuber has adapted to lower altitudes (Roca
et al., 2007; Roca & Manrique, 2005). Tubers varies in shapes (cylindrical,
ovoid, and spherical) and in colors (white, yellow, pink, orange, magenta, or
red or with red spots on a yellow background) (Flores et al., 2003).
Consumer panelists preferred the red tubers (Busch et al., 2000). Some
varieties present a larger quantity of mucilage and so must be boiled before
preparation. The nutritional value of ulluco is also variable, thus presents
carbohydrates, proteins, fat, and fiber contents from 73.5% to 84.2%, 8.5%
to 15.7%, 0.1% to 1.4%, and 0.5% to 5.0% DW, respectively, and among the
main sugars are glucose, fructose, and sucrose (13.1, 11.1% and 6.08% DM,
respectively) (Gross et al., 1989; King & Gershoff, 1987). Also, ulluco starch
(64.96% DW) has been studied. Ulluco starch granules present ellipsoid,
oval, conical, and prismatic shapes, with length up to 32.09 μm, and amylose
content and gelatinization temperature ranged from 20.4% to 26.4% and
62.7–67.8°C, respectively (Cruz et al., 2016; Valcárcel-Yamani et al., 2013).
4.1.5 Mashua
Mashua (Tropaeolum tuberosum Ruı́z & Pavón) (or isañu, cubio, añu, maswa)
is also a tuber crop indigenous to the Andean highlands and is of economic
value as a food and medicinal crop. It grows between 2800 and 4000 m.a.s.l
and its geographical distribution goes from Colombia up to Bolivia (Roca
et al., 2007). It ranks fourth in importance in the Andean region after pota-
toes, oca, and olluco (National Research Council, 1989). Among Andean
tubers, mashua shows the highest yield, grows easily, it is highly tolerant
to cold temperatures and it is unaffected by poor management. Mashua is
also a valuable crop because it is resistant to many insects, nematodes, fungi,
and other pathogens (Grau, Ortega, Nieto, & Hermman, 2003). This resis-
tance is correlated to the content of glucosinolates present in the tuber.
Mashua tubers have shapes ranging from conical to carrot-like and tubers
vary in color from white to yellow with occasionally variants that are purple
or red. Often, the peel is mottled or striped with red or purple; especially
bellow the eyes (National Research Council, 1989). Mashua tubers contain
carbohydrates, starch, protein, fat, and fiber contents of 69.7%–85.8%,
41.35%, 6.9%–15.7%, 0.1%–1.0%, and 4.8%–8.6% DW and a total sugar
content of 27.7% DW. In addition, insoluble fiber (in DW) reaches a value
of 10.55% respect to 6.85% and 7.22% presents in oca and ulluco tubers
(Gross et al., 1989; King & Gershoff, 1987; Valcárcel-Yamani et al.,
2013). Mashua tuber presents important contents of vitamin C in compar-
ison to other tuber crops, with values reported of 480 mg/100 g DW or
77.5 mg/100 g FW (Collazos, 1996; Sperling & King, 1990) as well as carot-
enoids (9 μg β-carotene/100 g DW).
More than 100 accessions have been recognized but probably many
more exist given the great variability of the genotypes in terms of shape
and color. This variability in terms of color and shape seems to be correlated
to the content of bioactive compounds. Among 84 evaluated genotypes,
phenolic compounds varied between 5.5 and 16.7 mg GAE g1 DW,
anthocyanins between 0.09 and 2.68 mg CGE g1 DW and carotenoids

between 0.48 and 15.09 mg β-caroteno/100 g DW. In vitro antioxidant
capacity varied depending on the method ABTS, FRAP, and ORAC
between values of 20.6–128.2, 22.7–173.2, and 35.1–158.8 μmol TE g1
DW, respectively (Campos et al., unpublished results).

4.2.1 Maca
Maca has been grown in Peru for at least 2000 years ago, but little is known
about its origin, in the Andean mountains. Maca is used by the Andean peo-
ple boiled or roasted as a food and also in ethnological medicine as an
antidepressive and injury cicatrizing drug. Maca is consumed fresh or
dehydrated, roots are boiled, oven or pot cooked and as alcoholic beverage
(Quirós & Aliaga-Cárdenas, 1992). It is also used in folk medicine as aph-
rodisiac and to improve fertility in humans and animals, to alleviate meno-
pause symptoms including hot flushes, tender breasts, vaginal dryness,
osteoporosis, etc. (Zhang, Yu, Ao, & Jin, 2006). During the Inca’s times,
soldiers were fed with maca before going to war to increase their energy
and vitality (Quirós & Aliaga-Cárdenas, 1992).
Currently, it is sold in local markets as aphrodisiac, energizer, and pro-
moter of fertility. It is also recommended to cure rheumatism, respiratory
illness, and as laxative. Other medicinal properties attributed to maca include
regulation of hormonal secretion, memory improvement, stimulation of
metabolism, antidepressive activity and fight against anemia, leukemia, can-
cer, and alcoholism (Quirós & Aliaga-Cárdenas, 1992).
4.2.2 Yacon
There is evidence about the use of Yacon by the pre-Incas and Inca cultures.
Chronicles that date back to 1615 mentioned Yacon as one of the 55 plants
cultivated in the Andes (Zardini, 1991). Yacon is traditionally consumed
fresh or dehydrated. Fresh yacon is very refreshing due to its high content
of water. In limited instances and Peruvian locations, it is consumed as jam or
chicha (alcoholic beverage). It can prevent tiredness and cramps due to its
high content of potassium reason why native people consume it during long
walks (a person can consume between 500 and 1000 g fresh Yacon). This
level of consumption does not produce toxic effects, except for few occa-
sions when flatulence can occur due to the high content of FOS
(Seminario et al., 2003). The Andean communities solar dry the tubers to
increase sweetness (Pedreschi, Campos, Noratto, Chirinos, & Cisneros-

Zevallos, 2003).
In Peru and Bolivia, the root is consumed by people with diabetes and
digestive problems but it is not clear if this practice has been recently adopted
or it is a tradition from old times. Andean growers consider Yacon as a skin
rejuvenating agent. In Cajamarca, folk tradition mentioned that during old
times people consumed it before going to bed to retard aging (Seminario
et al., 2003).
4.2.3 Oca and Olluco

Oca and olluco have the same range of culinary uses as do potatoes and they
are used in a large number of traditional recipes. These tubers, are usually
boiled and can be used as a component in many dishes, ranging from soups
and stews to desserts. Oca tubers in the Andean region are often placed in the
sun for a few days, during which they become sweet. Oca is converted into
dry products (cavi or caya, kaya, or khaya) to eliminate the bitter taste,
besides this transformation allows to eliminate soluble compounds such as
oxalates (Hermann & Erazo, 2000). A similar product as “chuño”
(frozen-dehydrated product from potato) can also be elaborated with olluco
(National Research Council, 1989; Roca et al., 2007).
Traditionally ulluco is known for its medicinal properties to treat burns
and to prevent scars. Heil, Bravo, Montoya, Robledo, and Osorio (2017)
evaluated the wound healing activity of aqueous extracts of U. tuberosus
tuber using in vitro models and found that an increase in collagenase activity
of 12% supports the utility of this tuber as an agent for scar treatment. Oca is
used to alleviate gastritis and acne treatment (communication with local
growers).
4.2.4 Mashua
In Bolivia and some parts of Peru, people eat cooked and frozen mashua
soaked in sugarcane syrup (thayacha). Also, mashua tubers are exposed to
the sun to increase their sweetness and reduce odor due to cyanide content
(Roca et al., 2007).
Mashua has many uses in the folk medicine of the Andean region.
Mashua diets are reputed to have beneficial effects on liver and kidneys
and to alleviate prostate and urinary disorders and also it is regarded as a
diuretic agent ( Johns, Kitts, Newsome, & Towers, 1982). Mashua uses
include also treatment of skin ulcers, diabetes, dengue, and malaria fever
and postpartum recovery (National Research Council, 1989; Rea, 1984)
and as antibiotic. Traditional use attributes antilibido and reduction of the

reproductive function of men (Roca & Manrique, 2005). Johns et al.
(1982) showed experimentally that male rats fed with mashua tubers had
significantly reduced testosterone and dihydrotestosterone levels (45%).
However, test animals maintained their capability to impregnate females.
Cárdenas-Valencia et al. (2008) found that after 12 days of treatment
(i.e., one spermatogenic cycle), mashua (1 g/kg) reduced testicular function,
increased epididymal sperm transit rate, and consequently reduced epidid-
ymal sperm number without any effect on testosterone levels.

4.3.1 Maca
Several authors have reported diverse bioactive compounds in maca such as
glucosinolates and derivatives (thiocyanates, nitriles, indoles, etc.), phytos-
terols, phenolic antioxidants, macaenes, macamides, alkaloids, and among
others. Some studies suggest that secondary metabolites found in maca are
responsible of the physiological effects. Scientific literature reports a low oral
toxicity level of maca in animals and in cellular experiments in vitro. Health
claims cover treatment of a wide range of illnesses and medical conditions
including energizer and fertility-enhancer properties, performance on sexual
dysfunctions, osteoporosis, benign prostatic hyperplasia, memory and learn-
ing, and protection of skin against ultraviolet radiation (Gasco, Villegas,
Yucra, Rubio, & Gonzales, 2007; Gonzales, 2012; Rubio, Caldas,
Dávila, Gasco, & Gonzales, 2006; Wang, Wang, McNeil, & Harvey,
2007). Gonzales (2012) reported 2000 different studies that demonstrate
the biological and pharmacological effects of maca consumption (Table 8).
The reported works have been carried out with commercial gelatinized
maca or with maca extracts. In most of the cases, commercial yellow maca
has been used and with less frequency red or black maca. In terms of the used
extracts, aqueous, hydroalcoholic, or with ethyl acetate have been reported.
Thus, different genotypes, products, or extracts will originate differences in
the content of bioactive compounds bioavailable.
Normal male rats treated with maca extracts showed significantly
increased epididymal sperm counts and spermatids, and higher testicular
and epididymis weights (Gonzales, Ruiz, Gonzales, Villegas, & Córdova,
2001). In rats, maca was also effective to prevent the reduction of body
weight and epididymal sperm count induced by high altitude (Gonzales
et al., 2004). Rubio et al. (2006) reported prevention of infertility in rats
treated with lead acetate. Gonzales et al. (2006) evaluated rats fed with three
Table 8 Summary of Some Studies That Evaluate the Consumption of Maca in Animals
Species Property
Rats Increase sperm count and sperm motility
Increase male sexual behavior
Small effect on rat male sexual behavior
Nutritional
Antistress
Prevent testosterone-induced prostatic hyperplasia Reversed
osteoporosis
Neuroprotective effects
Protects against UV radiation
Antioxidant status, lipid, and glucose metabolism
Mice Increase male sexual behavior
Increase embryo survival
Prevent testosterone-induced prostatic hyperplasia
Increase number of offsprings
Improve memory and learning
Guinea Increase number of offsprings
pigs
Fish Nutritional
Increase embryo survival
Bulls Improve sperm quantity and quality
Unaffected mating behavior
Fuente: Adapted from Gonzales, G.F. (2012). Ethnobiology and ethnopharmacology of Lepidium
meyenii (Maca), a plant from the Peruvian highlands. Evidence-based Complementary and Alternative
Medicine, 2012, 193496.
maca ecotypes (yellow, red, and black) and reported that black maca dis-
played a higher beneficial effect on sperm counts and epididymal sperm
motility. In this study, aqueous extracts obtained by severe heat treatment
(boiling conditions for 120 min) were employed and could have affected
negatively some bioactive compounds. Gasco et al. (2007) working with
male rats during 84 days and fed with yellow red and black maca extracts
(obtained at boiling conditions for 60 min), reported improved sperm count
in the epididymis and vas deferens; red maca affected sperm count only in the
vas deferens. The authors concluded that maca acts as a modulator of sperm
count at the reproductive tract level. Clement et al. (2010) reported that
whole maca supplementation improved sperm quantity and quality of bulls
to a certain degree, while mating behavior appeared unaffected. Others stud-
ies have demonstrated that oral administration of aqueous or ethanolic
extracts of Maca hypocotyls improved spermatogenesis in male rats and mice

(Gonzales, Rubio, Chung, Gasco, & Villegas, 2003). The efficacy of maca in
female rat fertility was tested by oral administration of maca extract to mice,
leading to increased litter size and pregnancy rates in adult female mice
(Ruiz-Luna et al., 2005). Gonzales, Córdova, et al. (2001) evaluated the
consumption of maca tablets in normal adults (24–44 years old) and reported
an increase of the semen volume, sperm count, mobile sperm count, and
motility while testosterone levels were not altered. Authors hypothesize that
bioavailable testosterone or testosterone receptor binding might be
increased. Another possibility is that maca may act without the participation
of androgen mechanism. This seems to be supported by the fact that the
weight of seminal vesicle, a target for androgen action, was not influenced
by Maca in adult male rats (Gonzales, Ruiz, et al., 2001). Maca is well known
for its aphrodisiac effects in men and women. Zheng et al. (2000) reported
that maca helped to develop libido and sexual potency in normal rats while
rats with erectile dysfunction improved their sexual function. Cicero et al.
(2002) evaluated three extracts obtained by successive extraction with hex-
ane, chloroform, and methanol from hypocotyls of dried and ground maca.
Results showed that subacute oral administration of maca hexanic and
methanolic extracts improved the copulatory performance of sexually inex-
perienced male rats. Hexanic fraction was the most effective, this fraction
improved the majority of the sexual parameters measured. Double-blind,
placebo-controlled, randomized, parallel trials carried out on men have
reported a widespread increase in sexual desire in the test group
(Gonzales, Córdova, et al., 2003). Gonzales et al. (2002) in a study carried
out with healthy men (21–56 years old), after 8 and 12 weeks of treatment
with maca extract, displayed an increase in sexual desire without changes in
the levels of testosterone and estradiol. The mechanisms of maca’s proposed
improvement of sexual performance have been studied, but have not been
fully clarified so far. There is clearly a need for further studies in this area.
The antiproliferative activity of maca has been reported by Gasco et al.
(2007). Aqueous extract of red maca was administered to rats and inhibited
the prostate hyperplasia induced by testosterone enanthate. The antip-
roliferative effect could be due to the transformation in the colon of
glucosinolates to anticarcinogenic derivatives (Fahey, Zalcmann, &
Talalay, 2001). Others health claims include memory and learning (black
Maca improves scopolamine-induced memory deficits) (Rubio et al.,
2006) and protection of skin against ultraviolet radiation (Gonzales, 2012).
4.3.2 Yacón
The bioactive potential of Yacon is related to the FOS and phenolic anti-
oxidants present. Yacon is a particularly abundant source of β-(2! 1)
fructooligosaccharides (FOS), which are considered prebiotic. FOS are fer-
mented selectively by many bifidobacteria, also lactobacilli, which for their
part are held to be probiotic (Pedreschi et al., 2003).
Genta et al. (2009) studied the effect of the consumption of Yacon syrup
on obese and slightly dyslipidemic premenopausal women during 120 days
at daily doses of 0.14 and 0.29 g of FOS/kg/day. A significant decrease of
body weight, waist circumference, and body mass index was reported. Addi-
tionally, a decrease in fasting serum insulin and homeostasis model assess-
ment index was observed. Syrup Yacon consumption increased
defecation frequency, feeling full. Levels of glucose and lipids were not
altered but a positive effect on LDL cholesterol in serum was observed.
Genta et al. (2005) evaluated the subchronic (4 months) effect of the admin-
istration of a dietary supplement of Yacon flour in normal Wistar rats at
levels of 340 and 6800 mg Yacon flour/kg weight. No toxicological or
nutritional effects were observed. Yacon root consumption showed no
hypoglycemic activity in normal rats and resulted in significantly reduced
postprandial serum triacylglycerol levels in both doses assayed.
de Moura, Caetano, Sivieri, et al. (2012) studied the influence of Yacon
on induced carcinogenesis with 1,2-dimethylhydrazine (DMH) in rats after
4 weeks induction with DMH. Rats were fed with a control diet or with a
diet containing different concentrations of atomized Yacon extract (0.5%
and 1%, respectively) and a symbiotic mixture composed of 1% Yacon
and Lactobacillus casei (2.5 1010 CFU g1) during 13 weeks. Results indi-
cate that Yacon and Yacon plus L. casei intake may reduce the development
of chemically induced colon cancer. Stimulatory effects of yacon FOS on Ca
intestinal absorption, bone mineral retention, and structural properties in the
femoral midshaft of Wistar rats fed ad libitum with diets supplemented with
yacon flour, have also been reported (Lobo, Colli, Eliana, & Tullia, 2007).
The prebiotic effect of yacon FOS was tested in vivo with a guinea pig
model. A diet rich in yacon FOS promoted the growth of bifidobacteria and
lactobacilli, resulting in high levels of short chain fatty acids (SCFAs) in the
cecal material and enhancement of cell density and crypt formation in cae-
cum tissue, being indicative of colon health benefits (Campos et al., 2012).
Even when most of the reported works of the beneficial effects of Yacon are
attributed to the prebiotic effect of FOS, it is also possible that the presence
of other beneficial effect such as fiber and phenolic compounds also
contribute to the health benefits. It has also been reported that yacon roots
are rich in phenolic compounds, mainly chlorogenic (caffeoyl-quinic) acid
and other caffeic acid derivatives (Takenaka et al., 2003; Yan et al., 1999).
4.3.3 Oca, Olluco, and Mashua

Campos et al. (2006) evaluated bioactive compounds: total phenolic com-
pounds, carotenoids, anthocyanin, betaxanthin, and betacyanin content as
well as the antioxidant capacity (ABTS assay) of different genotypes from
four species of edible tubers: native potatoes, mashua, oca, and ulluco. Dif-
ferences in bioactive compounds and activities were reported for the differ-
ent species and genotypes. In general, mashua tubers (purple genotypes)
showed the highest antioxidant capacity, total phenolic, and anthocyanin
content compared to the other crops.
Main bioactive compounds reported in oca correspond to phenolics and
anthocyanins. Contrary to mashua, purple oca presents as main anthocya-
nins: malvidin glucosides (malvidin 3-O-glucoside and 3,5-O-diglucoside),
petunidin, peonidin, and delphinidin (mono- and/or diglicosilates) and only
an acylated anthocyanin (malvidin 3-O-acetylglucoside-5-O-glucoside) was
found (Alcalde-Eon, Saavedra, Pascual-Teresa, & Rivas-Gonzalo, 2004).
Among the main nonanthocyanin phenolic compounds present in six dif-
ferent colored oca (yellow and purple) evaluated by HPLC-DAD: caffeic
and cinnamic acid derivatives as well as flavan-3-ols, flavones, and flavanone
derivatives were found. Based on their UV–vis spectral data, the flavan-3-
ols, flavones, and flavanones detected corresponded to bound forms of cat-
echin, luteolin, and apigenin and naringenin, respectively (Chirinos et al.,
2009). Oca phenolic compounds were fractioned in two main fractions:
an aqueous (Faq) and an ethyl acetate fraction (Fea), where Faq fractions
were the major contributors to the ABTS antioxidant capacity (77%–
82%). Finally, it was reported that a storage protein of oca tuber, named
ocatin, presents antibacterial and antifungal activities (Flores, Alape-
Girón, Flores-Dı́az, & Flores, 2002).
Ulluco is a crop that contains betalains in the form of betaxanthins
(22–96 μg g1 FW) and betacyanins (64 μg g1) with no presence of carot-
enoids or anthocyanins (Campos et al., 2006). Svenson, Smallfield, Joyce,
Sansom, and Perry (2008) identified betalains in red and yellow varieties
of ulluco, using LC–DAD–ESI–MS–MS2 analyses. Both the yellow and
red tubers were rich in yellow betaxanthins, and the most prominent, were
histidine–betaxanthin, arginine–betaxanthin, and glutamine–betaxanthin;
betacyanins were found in red tubers, with roughly 50% of this content
being betanin/isobetanin. Betacyanin levels were up to 70 μg g1 FW in red

tubers and betaxanthin levels were up to 50 μg g1 FW in yellow tubers.
Phenolic compounds of mashua (two purple and one yellow genotype)
have been partially identified through HPLC–DAD. Main compounds pre-
sent correspond to: gallic acid, gallocatechin, procyanidin B2, and
epigallocatechin, also flavan 3-ol derivatives, hydroxycinnamic, and hydro-
xybenzoic acid derivatives, rutin and/or myricetin and proanthocyanidins
derivatives. The proanthocyanidin (prodelphynidines) fractions were the
major contributors to the ORAC antioxidant activity of the mashua tubers
(34.7%–39.2%) (Chirinos, Campos, Costa, et al., 2008). Mashua anthocya-
nins from three purple genotypes were identified by HPLC–DAD and
HPLC–ESI/MS–MS. The main anthocyanins were delphynidins followed
by cyanidins and pelargonidins The two major pigments (56.4%–73.0% total
area range at 520 nm) were identified as acylated delphinidins: delphinidin
3-glucoside-5-acetylrhamnoside and delphinidin 3-sophoroside-5-
acetylrhamnoside (Chirinos et al., 2006). Mashua polyphenol extracts dis-
played good antioxidant properties against oxidative damage in biological
structures rich in PUFA (such as LDL and erythrocytes) (Chirinos,
Campos, Warnier, et al., 2008), also mashua polyphenols (not anthocyanins)
has been evaluated as antioxidant natural on soybean oil (Betalleluz-
Pallardel, Chirinos, Rogez, Pedreschi, & Campos, 2012), and in sacha inchi
oil (high polyunsaturated fatty acids: omega-3, 6% 85%) and in ground
raw pork meat (Chirinos, Pedreschi, Cedano, & Campos, 2015) with pos-
itives results. Also, mashua tubers have glucosinolates and isothiocyanates
( Jonhs & Tower, 1981; Ramallo et al., 2004), which give them a pungent
taste. Ramallo et al. (2004) found only one glucosinolate (p-methoxybenzyl
glucosinolate) in the six domestic varieties analyzed. Its concentration varied
between 36.5 and 90.0 μmol g1 DW; this range is relatively high when
compared with other edible glucosinolate-containing vegetables. “Dark”
colored tuber varieties showed higher levels than “light” colored tuber vari-
eties at normal harvest time. The main glucosinolates detected in cultivated
and different accessions of mashua were aromatic: 4-hydroxybenzyl
glucosinolate (glucosialbin), benzyl glucosinolate (glucotropaeolin), and
m-methoxy benzyl glucosinolate (glucolimnathin) (Ortega, Kliebenstein,
Arbizu, Ortega, & Quiros, 2006). The total amount of glucosinolates
observed ranged from 0.27 to 54.7 μmol g1 of dried tuber tissue. Recently,
Martı́n and Higuera (2016) reported a total content of glucosinolates that
ranged between >3.00 101 and 25.8 μmol g1 DW. p-Methoxybenzyl
glucosinolate was identified as the predominant glucosinolate and the
possible presence of p-hydroxy benzyl glucosinolate, in Colombian mashua

accessions was also reported. Glucosinolates are a group of secondary plant
metabolites (β-thioglycoside Nhydroxysulfates linked with a sulfur β-D-
glucopyranose) and depending on the side chain, glucosinolates can be clas-
sified as aliphatic, aromatic, or indolic. Glucosinolates can be hydrolyzed by
myrosinase. Among the breakdown products, isothiocyanates are account-
able for the anticarcinogenic properties of Brassicaceae (Basten, Bao, &
Williamson, 2002). In particular glucoraphanin, which is an aliphatic
glucosinolate, is the precursor of sulforaphane, the isothiocyanate that shows
the highest potency to induce phase II detoxification enzymes in humans
(Basten et al., 2002). Some mashua extracts rich in thiocyanates have dis-
played anticarcinogenic activity against colon and prostate cancer lines
(Noratto, Cisneros-Zevallos, & Mo, 2004). Given the high content of aro-
matic glucosinolates in mashua it would be interesting to evaluate the anti-
cancer properties of the derivatives of these compounds.
Guimaraẽs (2004) reported that mashua tubers possess proteins to confer
potent antifungal activity against T. harzium. Thus mashua tubers have
potential uses for the control of the green mold. In vitro assays established
an important fungal growth inhibition of the potato pathogen P. infestans
(Martı́n & Higuera, 2016).
5. CONCLUSIONS
Andean fruits, seeds, and tubers represent a rich source of nutritional
and functional compounds. This region is characterized by its biodiversity.
Thus, very marked differences in the content and type of nutrients and bio-
actives are encountered in different genotypes/accessions. A lot of effort has
been invested in the last decades to characterize main nutrients and bioac-
tives and some in vitro bioactive properties in many of these Andean fruits,
seeds, and tubers. However, more in vivo and clinical studies are needed.
Additionally, many of these crops are gaining attention in trend gastronomy,
which open the possibility for these crops to be known and commercialized
internationally. Factors beyond genetics, environmental, and crop manage-
ment on nutritional and bioactive potential, such as postharvest technologies
and processing should be further explored.
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FURTHER READING
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CHAPTER EIGHT

Bioactive Potential of Andean

1. BIODIVERSITY OF THE ANDES

2.1.3 Andean Berries

2.1.4 Sweet Pepino

2.2 Typical Use in Old Traditions

S. peruviana can be consumed fresh but it is mainly processed into mer-

2.3 Reported Bioactive Potential

2.3.3 Andean Berries

2.3.4 Sweet Pepino

Fig. 2 Andean grains: (A) quinoa and (B and C) tarwi.

3.1 General Information on Composition and Nutritional Value

A comparative proximal composition of quinoa from Andean countries

Repo-Carrasco-Valencia and Wright, Pike, Fairbanks, Miranda, Vega-Gálvez, Martinez, Nascimento

3.1.2 Cañihua and Andean Amaranth

3.1.3 Andean Lupin or Tarwi

(Carvajal-Larenas, Linnemann, Nout, Koziol, & Van Boekel, 2016). These

3.2 Typical Use in Old Traditions

Many of these preparations and culinary applications were inherited from

3.3 Reported Bioactive Potential

Some quinoa varieties may exhibit yellow–orange and red–violet colors

3.3.2 Cañihua and Andean Amaranth

Amaranthus species such as Amaranthus cruentus, Amaranthus hypochondriacus,

3.3.3 Andean Lupin or Tarwi

4. TUBERS AND ROOTS

anthocyanins between 0.09 and 2.68 mg CGE g1 DW and carotenoids

4.2 Typical Use in Old Traditions

increase sweetness (Pedreschi, Campos, Noratto, Chirinos, & Cisneros-

4.2.3 Oca and Olluco

and as antibiotic. Traditional use attributes antilibido and reduction of the

4.3 Reported Bioactive Potential

extracts of Maca hypocotyls improved spermatogenesis in male rats and mice

4.3.3 Oca, Olluco, and Mashua

being betanin/isobetanin. Betacyanin levels were up to 70 μg g1 FW in red

possible presence of p-hydroxy benzyl glucosinolate, in Colombian mashua

Filho, A. M. M., Pirozi, M. R., Borges, J. T. D. S., Pinheiro Sant’Ana, H. M.,

Martı́n, J. C., & Higuera, B. L. (2016). Glucosinolate composition of Colombian accessions

Piotrowicz-Cieslak A. I., Michalcyk, D. J. & Gorecki, R. J. (2004). Flatulence-producing

Schoenlechner, R., Siebenhandl, S., & Berghofer, E. (2008). Pseudocereals. In E. K. Arendt

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