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Chayote (Sechium edule) Phytochemical and pharmacological approaches

Authors:
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47
Chayote (Sechium edule (Jacq.) Swartz)
Oscar Andrés Del Ángel Coronel,
1
Elizabeth León-García,
2
Gilber Vela-Gutiérrez,
3
Javier De la Cruz
Medina,
2
Rebeca García Varela,
4
and Hugo S. García
2
1
Instituto Tecnológico Superior de Huatusco, Av. 25 Poniente 100, Col. Reserva Territorial, Huatusco, Ver. 94100, Mexico
2
UNIDA-Instituto Tecnológico de Veracruz, M.A. de Quevedo 2779, Col. Formando Hogar, Veracruz, Ver. 91897, Mexico
3
Facultad de Ciencias de la Nutrición y Alimentos, Universidad de Ciencias y Artes de Chiapas, Lib. Nte. Pte. No. 1150, Ciudad Universitaria, Col. Lajas Maciel, CP
29000, Tuxtla Gutiérrez, Chiapas, Mexico
4
CIATEJ, Autopista Mty-Aeropuerto, Km 10, Parque PIIT, Vía de Innovación 404, Apodaca, NL 66629, Mexico
47.1 Introduction
Chayote (Sechium edule (Jacq.) Swartz) is a plant native to
Mexico and Latin America. It was rst domesticated by the
Aztecs and Mayas in pre-Columbian times. This plant is a
member of the Cucurbitaceae family and is mainly pro-
duced as a non-traditional export crop. Because of the wide
diversity of chayote plants found in Mexico (predomi-
nantly the states of Veracruz, Puebla, Chiapas, and Oaxaca)
and in Guatemala, this region is thought to be their
geographic origin. Today, chayote is cultivated through
tropical and subtropical regions of the world (Newstrom,
1991; Lira Saade, 1996; Bisognin, 2002; Cadena Iñiguez
et al., 2006b). Chayote takes several alternative names
depending on the region, such as cidrayote,chayotte,
chiote,cho-cho,choko,chow-chow,christophene, custard,
hayatouri,huisquil, mango squash, mirliton,sayote, vege-
table pear, and xuxu, among others (Newstrom, 1991;
Aung et al., 1990; Lira Saade, 1996; Lim, 2012).
There is a wide range of forms, colors, sizes, and avors
(bitter, neutral, and slightly sweet) among the S. edule fruit;
however, other physiological and morphological variations
have been reported (Cadena Iñiguez et al., 2006a, 2006b,
2008; Avendaño Arrazate et al., 2012). The whole plant is
of nutritional importance and has many traditional medi-
cine applications (Lim, 2012; Cadena Iñiguez et al., 2010;
Lombardo-Earl et al., 2014). However, only the fruit is of
economic importance for trade and export purposes in
several countries, including Mexico, Costa Rica, Brazil,
Puerto Rico, Canada, and the USA, with an increasing
demand (Cadena Iñiguez et al., 2007; Abdelnour and
Rocha, 2008; Olguín Hernández et al., 2013). It is also
produced and traded at a smaller scale in countries like
England, France, and Spain (Jiménez Hernández et al.,
2007).
The neutral avor and softness of chayote fruit make it
particularly suitable for the food industries. Although
chayote is mainly used for culinary purposes, the taste
may vary depending on the strain and use. Additionally,
the stems, leaves, and tuberous sections of the adventi-
tious roots are also commonly consumed and are of
nutritional relevance as food and feed (Morton, 1981;
Aung et al., 1990; Rao et al., 1990; Booth et al., 1992;
Barrera Marín, 1998; Gajar and Badrie, 2001; Cadena
Iñiguez and Arévalo Galarza, 2011).
47.2 Culinary Uses
Chayote is used for human consumption in many coun-
tries; nevertheless, preparation may be different depend-
ing on the region. According to Aung et al. (1990),
preparation is often dictated by the custom and taste of
a particular ethnic group. For example, in Mexican and
Latin American households and restaurants, the fruit may
be boiled, baked, stuffed, mashed, fried, scalloped, or
pickled, but the main method of consumption is in a
broth (Booth et al., 1992; Lim, 2012). In some cases it can
be eaten raw in salads and salsas (Morton, 1981).
In France, chayote fruit is served as a substitute for
artichoke hearts. In Jamaica and Puerto Rico, the matured
fruit are halved, boiled, and served; the seeds are also
consumed. In New Zealand, chayote is preserved in the
same manner as dill pickles. In India, chayote is eaten
together with other vegetables in a curried dish. In south-
eastern Asia, chayote is consumed similarly to French
fries (Morton, 1981; Aung et al., 1990).
Corresponding author.
979
Fruit and Vegetable Phytochemicals: Chemistry and Human Health, Second Edition. Edited by Elhadi M. Yahia.
© 2017 John Wiley & Sons, Ltd. Published 2017 by John Wiley & Sons, Ltd.
C47 05/01/2017 14:24:32 Page 980
On the other hand, chayote is also employed in the baby
food industry to formulate purees, juices, sauces, pasta
dishes, and jams because of its neutral taste (Gajar and
Badrie, 2001; Cadena Iñiguez and Arévalo Galarza, 2011).
Chayote fruit are an alternative ingredient for stews and
desserts (Gajar and Badrie, 2001); however, the stems,
leaves, and tuberous sections of the adventitious roots are
also consumed. In order to increase chayote shelf life and
availability, dehydration of the fruit has been studied in
Mexico and other countries (Lira Saade, 1996).
After the vine has grown for 2 years, the roots become
the most nutritive part of the plant, in the form of tubers.
In Mexico, Costa Rica, and Indonesia the tuber root is
considered a delicacy. In addition, over-mature tubers can
be employed as a source of starch, with a yield of 10% to
25%. Chayote tuber starch is easy to digest and is recom-
mended for infants and paralyzed patients. It has been
reported as a potential excipient for controlled release of
pharmaceuticals, and as a good substitute for potato in
food dispersions where a high viscosity is needed (Mor-
ton, 1981; Cruz León and López Rueda, 2005; Garzón,
2006a, 2006b; Jiménez Hernández et al., 2007). In Central
America and India it is used as food for humans and
fodder for animals (Lira Saade, 1996; Barrera Marín,
1998).
47.3 Health Effects
The use of natural products in traditional medicine has
been a common practice around the world (Diré et al.,
2007a). For chayote plant, several pharmacological and
medicinal uses have been reported; it is recommended in
controlled diets (Bisognin, 2002) to reduce the health risks
related to diabetes (Gajar and Badrie, 2001; Diré et al.,
2007b) and obesity (Yang et al., 2015a). Decoctions made
from leaves or fruits are used as a diuretic, to reduce
burning during urination, to dissolve kidney stones, in
other renal diseases, and as a remedy for pulmonary
ailments (Aung et al., 1990; Barrera, 1998; Cadena Iñiguez
et al., 2007). Extracts of chayote leaves and seeds are
considered to be effective when consumed several times
daily, to lower blood pressure and dissolve urinary calci-
cations (Morton, 1981; Lira Saade, 1996). An emulsion
of the seeds is given to relieve intestinal inammation
(Morton, 1981). In addition, strong anti-inammatory
and cardiovascular modifying properties of chayote fruit
and leaves have been conrmed by several pharmaco-
logical studies. Chayote extract intakes are suggested for
arteriosclerosis since it promotes vascular relaxation,
lowers hypertension, and reduces the risk of coronary
heart diseases (Aung et al., 1990; Lira Saade, 1996; Ibarra-
Alvarado et al., 2010; Lombardo-Earl et al., 2014). Other
valued properties related to chayote have been explored;
antitumoral activity on cancer cell lines (Cadena Iñiguez
et al., 2013) and hepato-protective and coronary heart
disease protective effects have been determined (Lira
Saade, 1996; Barrera Marín, 1998). These benets may
be associated with its potent antioxidant activity (Firdous
et al., 2012).
Nevertheless, some species of chayote, such as Sechium
compositum, which is closely related to the cultivated
chayote (S. edule), not only have pharmacological appli-
cations in humans but also are used as medicine for
animals. It is used against lice infestation in animals
(Lira and Caballero, 2002).
Few negative side effects have been conveyed for cha-
yote manipulation. Morton (1981) reported skin irritation
and numbness caused by prolonged periods of chayote
manipulation. Jensen and Lai (1986) reported a single case
in which a pregnant 18-year-old white Cuban woman
suffered severe hypokalemia (reduction in total body
potassium) due to a high intake of chayote infusion;
the infusion produced an excessive diuretic effect which
lowered her potassium levels.
47.4 Proximate Analysis of Chayote
The main phytochemical compounds of S. edule extracts
determined in many pharmacological studies have been
identied as peroxidases, alkaloids, avonoids, phenols,
polyphenols, saponins, steroids, triterpenes, and tannins
(Ibarra-Alvarado et al., 2010; Firdous et al., 2012; Nou-
medem et al., 2013; Lombardo-Earl et al., 2014; Chao
et al., 2014).
A compendium of proximate composition data of cha-
yote is shown in Table 47.1. For comparison, we provide
in Table 47.2 our laboratory data regarding proximate
composition, specically for S. edule (Jacq.) Swartz var.
virens levis fruits, which is the main chayote export variety
from Mexico to the United States and Europe. In the
following sections, we provide an overall discussion of the
different components and chayote core nutrients.
47.5 Moisture, Carbohydrate, and
Caloric Content
Because of the high moisture content (c.8995%), cha-
yote fruit falls in the category of the eshy vegetables and
fruits. The chayote plant has a relatively high caloric
content, especially in the young stems and tuber roots;
however, that does not apply in the fruit. Carbohydrates
are the main energy reserve and can be of two types:
available carbohydrateswhich include starch and sug-
ars; and non-available carbohydratessuch as crude ber
and different dietary ber constituents (Modgil et al.,
980 Fruit and Vegetable Phytochemicals
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Table 47.1 Proximate analysis reported for chayote by various authors
Component Seed Fruit Leaves Stems Roots
Calories (kcal) 2631
e),f)
60
e),f)
79
e),f)
Moisture (%) 8994.71
a),e),f),h),i)
91.8
d) (raw)
92.9
d) (cook)
89.7
e),f)
71.079.7
c),e),f)
Carbohydrates (%) 4.26.0
c),e),f)
3.37.7
c),e),h)
0.32.1
c),d) (raw)
2.4
d) (cook)
0.34.7
e),f)
17.8
e),f)
Total solids (%) 4.485.3
a),i)
––
Soluble sugars (%) 4.2
e)
3.58.4
f),g)
––0.030.6
c),e)
Reducing sugars (%) 5.326.42
g)
––
Non-reducing sugars (%) 0.470.77
g)
––
Starch (%) 1.9
c),e)
0.21.56
c),e),g)
0.7
c)
0.7
e)
13.649
c),e),j)
Fats (%) 8.529.1
i)
0.0380.3
b),e),f),h),i)
0.0160.4
c),d) (raw. cook)
0.4
e),f)
0.2
e),f)
Protein (%) 5.5
e),f)
0.161.1
e),f),h),i)
3.0
d) (raw)
2.8
d) (cook)
4.0
e),f)
0.4
c) (as albumen)
2.0
e),f)
Ash (%) 0.393.65
a),e),h)
1.1
d) (raw)
0.4
d) (cook)
1.2
e)
1.0
e)
2.3
c) (as minerals)
Crude ber (%) 0.47.6
a),e),f),h),i),g)
1.1
d) (raw)
0.9
d) (cook)
1.2
e),f)
0.4
e),f)
Hemicelluloses (g/100 g) 6.167.55
g)
––
Cellulose (g/100 g) 16.4217.28
g)
––5.6
c)
Lignin (g/100 g) 0.230.267
g)
––
Carotenes (mg/100 g) –– 1.0
d)
––
Pectin (%) 1.5
a)
––
Sources:
a) Flick et al., 1977, 1978 (green variety of fresh chayotes purchased in New Orleans market);
b) Akihisha et al., 1986 (chayotes purchased locally in Japan);
c) Aung et al., 1990 (analysis of edible enlarged storage root, tender apical shoots, and fruit esh of a light-green type of chayote on a wet weight basis);
d) Booth et al., 1992 (S. edule shoots, raw and cooked);
e) Lira Saade, 1996;
f) Barrera Marín, 1998;
g) Modgil et al., 2004 (g/100 mg dry weight basis);
h) Cadena Iñiguez and Arévalo Galarza, 2011;
i) Del Ángel Coronel, 2015 (values from Mexican export chayote fruit S. edule var. virens levis);
j) Cadena Iñiguez et al., 2011 (from chayote root tuber).
Table 47.2 Proximate analysis of fruits of Sechium edule var. virens levis, the main Mexican export variety (wet weight basis)
Component (g/100 g) Horticultural maturity of fruit
a)
Sprouting or germinated fruit
b)
Moisture 94.70 ±0.02 93.59 ±0.18
Ash 0.32 ±0.16 0.65 ±0.01
Protein 0.16 ±0.006 0.04 ±0.01
Fats 0.13 ±0.004 0.74 ±0.35
Crude ber 1.14 ±0.96 0.92 ±0.01
Carbohydrates 3.54 ±0.01 4.05 ±0.01
Total solids (%) 4.48 ±0.49
a) Horticultural maturity considered when the fruit has reached 18 to 21 days post-anthesis.
b) Sprouted fruits 1317 days postharvest, when the new plantlet is clearly visible.
Source: Del Ángel Coronel, 2015.
47 Chayote (Sechium edule (Jacq.) Swartz) 981
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2004). In Table 47.1, the main source of energy is provided
by young stems and tuber roots, which exhibit more
caloric content than fruit alone. However, the highest
carbohydrate content is found in chayote roots (tuberous
roots) and is essentially represented by starch. According
to Cadena Iñiguez et al. (2011) and Jiménez Hernández
et al. (2007) these data can be compared to those for
potato and maize starches, because they exhibit similar or
higher yields (49%), purity (8998%), and physico-
chemical, rheological, and molecular characteristics. Cha-
yote tuber starch had higher viscosity peak and ow
properties, lower gelatinization temperature, but higher
enthalpy, than potato, and a similar retrogradation rate.
The authors concluded that chayote tuber starch could be
an alternative for starch isolation, with better character-
istics than commercial potato and maize starches.
47.6 Dietary Fiber
High ber diets have been proven to be health benecial.
Individuals with high dietary ber intake appear to have a
signicantly lower risk of developing coronary heart
diseases, strokes, hypertension, diabetes, obesity, and
gastrointestinal diseases, among other chronic and degen-
erative diseases. Increasing ber intake can also lower
blood pressure and cholesterol levels in serum, as well as
impact positively on digestion rate, colon pH, and body
weight reduction (Anderson et al., 2009; Bernaud and
Rodrigues, 2013).
Dietary ber (crude ber) content in chayote fruits
ranges from 0.47.6% when raw to 2.2% when cooked
(Flick et al., 1977; Cadena Iñiguez and Arévalo Galarza,
2011). Chayote fruits have a higher ber content com-
pared to the stems, leaves, and roots (Aung et al., 1990;
Lira Saade, 1996; Modgil and Modgil, 2004).
47.7 Amino Acids and Protein
Content
The total protein content in chayote fruits ranges from
34% when raw to 2.8% after cooking. Nevertheless, the
protein content in chayote fruits is too low to be consid-
ered a primary source of protein when compared to that
from foods of animal origin. The amino acid composition
of chayote proteins is summarized in Table 47.3. Leaves
have higher amino acid content than fruits or seeds, and,
with the exception of cysteine and methionine, chayote
leaves provide the main essential amino acids (Flick et al.,
1977, 1978; Rao et al., 1990).
On the other hand, chayote seeds were considerably
richer in all amino acids than fruits, specically lysine,
arginine, aspartic acid, serine, glutamic acid, alanine,
valine, isoleucine, leucine, and phenylalanine. Methionine
was also found in chayote seeds; however, the concentra-
tion in fruit esh was probably too low for detection (Lira
Saade, 1996; Lim, 2012).
47.8 Total Lipid Content and Fatty Acid
Prole
The lipid content in chayote fruits was very low, ranging
from 0.038% to 1% (Flick et al., 1977; Akihisha et al., 1986;
Lira Saade, 1996; Barrera Marín, 1998; Cadena Iñiguez
and Arévalo Galarza, 2011; Del Ángel Coronel, 2015).
However, the percentage may vary depending on the
maturity stage and variety. Del Ángel Coronel (2015)
measured lipid contents of 0.13 ±0.004% in fruits during
their horticultural maturity stage (18 to 21 days post-
anthesis), but when the fruit sprouted these values
increased to 0.74 ±0.35%. Chayote seeds reached their
maximal values 5 days postharvest (29.1%); they then
progressively decreased and reached their minimum
value of 0.96% after 29 days postharvest.
Table 47.3 Amino acid proles of chayote protein (dry weight
basis)
Amino acid Content (mg/g)
Flesh Seed Leaves
Lysine 0.421 1.527 5.93
Histidine 0.229 0.669 2.20
Ammonia 0.624 0.736
Arginine 0.544 2.498 4.10
Aspartic acid 1.452 2.105 1.24
Threonine 0.641 0.876 2.78
Serine 0.731 1.550 1.03
Glutamic acid 1.973 5.247 12.15
Proline 0.688 0.972 6.00
Glycine 0.648 0.956 11.01
Alanine 0.799 1.570 11.36
Half cysteine (cysteic acid) 0.035 0.097
Valine 0.987 1.744 8.34
Isoleucine 0.696 1.297 5.25
Leucine 1.208 2.694 9.89
Tyrosine 0.502 0.755 2.61
Phenylalanine 0.747 1.810 5.10
Methionine (methionine sulfone) 0.270 0.6
Sources: Flick et al., 1977, 1978; Rao et al., 1990; Lira Saade, 1996.
982 Fruit and Vegetable Phytochemicals
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In addition, differences in lipid contents have been
observed depending on the variety. Wild-type chayotes,
such as nigrum levis (0.07%), nigrum xalapensis (0.03%),
nigrum spinosum (0.08%), and nigrum maxima (0.04%)
(Cadena Iñiguez and Arévalo Galarza, 2011), exhibit a
lower lipids content than commercial strains (S. edule var.
virens levis).
Del Ángel Coronel (2015) identied six lipid families in
chayote fruit mesocarp by gas chromatography, throughout
its shelf life at 25 °C, from harvest to senescence, including
horticultural maturity and sprouting (Table 47.4). The
author reported palmitic acid as the predominant fatty
acid (39.6%), followed by linolenic (18%), stearic (16.8%),
oleic (11.2%), linoleic (8.87%), and palmitoleic acids (3.15%).
However, in chayote leaves, Rao et al. (1990) observed that
linolenic acid was the predominant fatty acid (42.176.7%),
followedbypalmitic(13.738.5%) and linoleic (5.715.3%)
acids (Table 47.5). In addition, these authors reported a total
lipid content in chayote leaves of 2.32%, of which 40.2% was
non-polar lipids (NL), 30.8% glycolipids (GL), and 29%
phospholipids (PhL).
The total lipid contents in stems and roots have been
reported by Lira Saade (1996) and Barrera Marín (1998) as
having values of 0.4% and 0.2% respectively. Similarly to
chayote fruit, these values are too low to be considered
signicant. No data were found for fatty acid proling in
tuber roots or stems.
47.9 Minerals
Chayote fruits contain from 1.1% to 3.65% of ash on a dry
weight basis (Tables 47.1 and Table 47.2); this includes 21
minerals (Table 47.6). Although the macronutrient con-
tents of chayote fruits are adequate, the microelements
are low. Only the potassium and phosphorus contents can
be compared with those of cucumber (Cucumis sativus)
or eggplant (Solanum melongena) (Flick et al., 1978; Lira
Saade, 1996; Modgil et al., 2004; Cadena Iñiguez and
Arévalo Galarza, 2011).
The overall observations determine that the shoots are
more nutritious than the fruits, since they have a higher
Table 47.4 Fatty acid prole of chayote fruit during postharvest period
Postharvest days Free fatty acids (% area GC diagram)
C16:0 C16:1 C18:0 C18:1n9c C18:2n6c C18:3n3
1 35.9
bcd
1.44
bc
5.82
c
5.21
d
17.9
a
33.6
a
5 30.9
d
2.56
bc
10.7
bc
10.2
bcd
12.3
ab
24.9
ab
9 37.4
bcd
2.94
bc
21.6
a
13.5
abc
8.17
bc
13.1
d
13 57.9
a
2.97
bc
22.1
a
6.24
d
4.84
c
3.39
e
17 35.7
cd
3.91
b
21.4
a
8.49
cd
7.86
bc
22.5
bc
21 34.1
d
9.05
a
17.0
ab
18.7
a
5.03
c
16.0
d
25 44.1
b
1.27
c
19.6
a
13.3
abc
6.88
c
14.7
d
29 41.5
bc
1.07
c
16.6
ab
14.5
ab
8.05
bc
16.1
cd
Mean 39.6 ±8.4 3.15 ±2.5 16.8 ±5.8 11.2 ±4.5 8.87 ±4.3 18.03 ±5.9
ae
Means with same letter are not statistically different at P0.05 by Tukey.
Source: Del Ángel Coronel, 2015.
Table 47.5 Fatty acid composition of chayote leaves
Lipid class Fatty acid (% area GC diagram)
14:0 16:0 16:1 16:2 16:3 18:0 18:1 18:2 18:3 20:0 22:0
Non-polar lipids 0.0 38.5 0.2 0.3 0.6 3.2 1.8 11.2 43.7 0.5 0.0
Glycerolipids 0.5 13.7 0.0 0.0 0.0 1.4 1.7 5.7 76.7 0.2 0.1
Phospholipids 0.0 33.8 1.9 0.3 0.0 3.0 3.1 15.3 42.1 0.5 0.0
Source: Rao et al., 1990.
47 Chayote (Sechium edule (Jacq.) Swartz) 983
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iron and phosphorus content and are a good source of
calcium (Morton, 1981; Booth et al., 1992). Other ele-
ments were also found, but in low concentration. It is
important to note that the mineral content may vary with
location; for example, the zinc values of US-grown cha-
yotes were c. 6.4 mg/100 g, while Brazilian-grown fruits
were ca. 45.7 mg/100 g (Flick et al., 1977).
47.10 Vitamins
The immature chayote fruits have good levels of vitamins
C and E, as well as folates (vitamin B
9
)atc. 93.5 mg/100 g
in raw fruit (Table 47.7); however, when the fruits are
boiled and diced, the value is reduced to 14.4 mg/100 g.
Ascorbic acid content (vitamin C) in S. edule may range
from 6.4 to 7.7 mg/100 g, depending on the variety
(Cadena Iñiguez and Arévalo Galarza, 2011). For exam-
ple, when comparing S. edule var. virens levis, the main
export variety, the ascorbic acid values were
6.76 ±0.16 mg/100 g, while for varieties nigrum xalapen-
sis,nigrum spinosum,nigrum levis,albus minor,albus
dulcis,albus levis, and the wild relative, the values were
6.53 ±0.53, 4.95 ±0.49, 6.65 ±0.18, 7.82 ±0.42,
7.42 ±1.27, 7.75 ±0.22, and 3.99 ±0.16 mg/100 g, respec-
tively (Cadena Iñiguez et al., 2011).
Preliminary reports on carotene content demonstrated
that in chayote fruit they were either absent in tissue or
present in a concentration too low to be detected. These
data agree with those most recently reported by Cadena
Iñiguez et al. (2011), where no carotenoids were detected
in ve of the eight varieties of S. edule fruits evaluated;
only yellow fruits had a carotenoid amount, recorded at
Table 47.6 Mineral composition of chayote (dry weight basis)
Trace mineral Fruit
(mg/100 g)
Leaves and stem
(mg/100 g)
Root
(mg/100 g)
Total (ether extract) 0.60.62% ––
N 2.872.9 ––
P 4.030.0 4570 34
Al 0.853 3059.19
Ca 12.019.0 58 7
Cd 0.002 ––
Ce 0.004 ––
Cl 14.64 ––
Co 0.0033 ––
Cr 0.0172 0.18
Cu 0.0870.123 ––
Fe 0.200.779 24.63 0.8
K 125.0128.79 81316
Mg 12.015.40 2667.01
Mn 0.0494 ––
Na 1.772.0 ––
Mo 0.008 ––
Rb 0.098 ––
Se 0.028 ––
Sn 0.016 ––
V 0.0059 ––
Zn 0.0640.74 1.29
Sources: Flick et al., 1977, 1978; Rao et al., 1990; Booth et al., 1992; Hedges
and Lister, 2009; Lim, 2012.
Table 47.7 Vitamins content in chayote fruits
Vitamins Units Content Vitamins Units Content
Vitamin A (retinol) μg 0.001 Folate, DFE mcg DFE 93
Vitamin A, RAE μg RAE 0.0 Vitamin B
12
mg/100 g 0.0
Vitamin A, UI UI 0.0 Vitamin B
12
, added mg/100 g 0.0
Vitamin B
1
(thiamin) mg/100 g 0.025 Vitamin C (ascorbic acid) mg/100 g 7.7
Vitamin B
2
(riboavin) mg/100 g 0.029 Vitamin E (α-tocopherol) mg/100 g 0.12
Vitamin B
3
(niacin) mg/100 g 0.47 Vitamin E, added mg/100 g 0.0
Vitamin B
5
(pantothenic acid) mg/100 g 0.249 Vitamin K (phylloquinone) μg 4.1
Vitamin B
6
(pyridoxine) mg/100 g 0.076 β-Carotene μg 0.0
Choline, total mg/100 g 9.2 α-Carotene μg 0.0
Vitamin B
9
(folic acid) μg 0.0 β-Cryptoxanthin μg 0.0
Folate, total μg 93 Lycopene μg 0.0
Folate, food μg 93 Lutein +zeaxanthin μg 0.0
Sources: Flick et al., 1977; Hedges and Lister, 2009; Cadena Iñiguez and Arévalo Galarza, 2011.
984 Fruit and Vegetable Phytochemicals
C47 05/01/2017 14:24:34 Page 985
levels of 0.0108 mg/100 g for albus minor, 0.0056 mg/
100 g for albus dulcis, and 0.0042 mg/100 g for albus levis.
The authors noted that the carotenoids content in yellow
varieties might be attributed to a possible adjustment of
the mevalonic acid route, since all natural terpenoid
compounds, such as cucurbitacins and carotenoids, bio-
synthesize the mevalonic acid by means of the acetyl CoA
route through a common intermediary. In the metabolic
cucurbitacin route, isoprene units are used as a precursor
of different metabolites; this mechanism allows the plant
to synthesize metabolites in larger quantity when
required, utilizing the existing isoprene units, and reduc-
ing the synthesis of other compounds. In this context,
chemical plasticity would allow S. edule varieties to adjust
cucurbitacin contents in order to synthesize a large
quantity of carotenoids in yellow fruits. Nevertheless,
even though the yellow fruits were the only strains
with carotenoid contents, they had lower chlorophyll-a
and chlorophyll-bcontent than green and wild type fruits.
47.11 Phenolic Compounds
Flavonoids and phenolic acids are two compounds in the
phenolic group; this group includes more than 9000
chemical species produced widely across the plant
kingdom. They have a variety of purposes, such as pro-
tecting against fungal diseases and insect attack, and
imparting taste; some also provide aroma and color.
Structurally, all phenolics contain at least one phenol
ring and at least one hydroxyl group. Because of these
properties phenolic compounds are considered excellent
free radical scavengers with an antioxidant activity
(Hedges and Lister, 2009; Al-Abd et al., 2015; Yang
et al., 2015b). For some plant species, in this case chayote,
avonoids and other phenolic compounds are the main
bioactive compounds of hydro-alcoholic extracts of fruits,
roots, and aerial parts. Ordoñez et al. (2003) found the
highest phenolic compound content in tincture/macer-
ated extracts (2.06, 2.81, and 5 mg/g in leaf, stem, and
seed, respectively), followed by leached/uid extract (0.44,
1.41, and 0.56 mg/g in leaf, stem, and seed, respectively)
and alcoholature (0.15, 0.06, and 0.13 mg/g in leaf, stem,
and seed, respectively). The highest total amount of
avonoids was found in the leaves (35 mg/10 g of dried
part), followed by roots (30.5 mg/10 g) and nally by
stems (19.3 mg/10 g). In these structures, eight avonoids
were detected: three C-glycosyl and ve O-glycosyl a-
vones (Lim, 2012). These compounds are implicated in
several pharmacological reports and traditional medicine
uses previously mentioned.
Lombardo-Earl et al. (2014) found several polyphenolic
compounds, specically avonoids and phenylpropa-
noids, in hydro-alcoholic extracts. Acetone and methanol
extracts obtained from the roots of S. edule, when identi-
ed by MS-PDA-HPLC, uncovered cinnamic derivative
compounds such as cinnamic acid methyl ester, coumaric
acid, and vitexin, which have been demonstrated to have
important pharmacological activity. Their results
revealed an antihypertensive and vasorelaxing effect
from the chayote root extracts and their fractions,
when administered intravenously in anesthetized rats
treated with angiotensin II (AG II) as a potent hyper-
tension inducer. The data obtained conrmed the effects
on different cardiovascular parameters, such as lowering
blood pressure and displaying changes on electrocardio-
graphic recordings. The authors propose that the inter-
action of the extracts produces an AG II and calcium
antagonism. The activity of avonoids and cinnamic acid
derivatives of chayote root extracts can be associated with
an AG II receptor blocking action, as well as obstructing
the secondary messenger system initiated by AG II, which
promotes the efux of sarcoplasmic calcium that activates
the store operated channels, allowing more calcium to
enter the cell and form a calciumcalmodulin complex
that produce vascular contraction.
According to Wu et al. (2014) and Yang et al. (2015a),
the polyphenol components of S. edule shoots attenuated
hepatic lipid accumulation that results in fatty liver, seen
in vitro; these results may have applications for stages of
metabolic syndrome such as obesity, hypertension, and
diabetes. In animal models, these authors observed that
the S. edule shoots extracts were able to lower body
weight, reduce adipose tissue fat, and regulate hepatic
lipid contents (e.g. triglycerides and cholesterol). Addi-
tionally, oleic acid induced lipid accumulation in HePG2
cells was inhibited, AMP-activating protein kinase
(AMPK) activation was enhanced, and a decrease in
numerous lipogenic-related enzymes was produced,
such as fatty acid synthase (FAS), sterol regulator ele-
ment-binding proteins (SREBPs), e.g. SREBP-1 and
SREBP-2, and HMG-CoA reductase (HMGCoR) pro-
teins; and there was increased expression of CPT-I (car-
nitine palmitoyltransferase I) and PPARα(peroxisome
proliferators activated receptor α), which are critical
regulators of hepatic lipid metabolism. The authors con-
cluded that the polyphenol extract of S. edule shoots can
prevent fatty liver. Other main active components of S.
edule shoot extracts are caffeic acid and hesperetin. These
results suggest that S. edule shoots have potential for
developing nutraceutical foods for preventing and
remedying fatty liver.
Firdous et al. (2012) have reported hepatoprotective
activity of chayote fruit ethanolic extracts and its ethyl
acetate and n-butanol fractions against carbon tetra-
chloride (CCl
4
) induced hepatotoxicity in rats. This was
effected by reducing the levels of aspartate amino-
transferase (AST), alanine aminotransferase (ALT),
47 Chayote (Sechium edule (Jacq.) Swartz) 985
C47 05/01/2017 14:24:34 Page 986
alkaline phosphatase (ALP), total bilirubin, and hepatic
lipid peroxidation, and increasing the levels of antioxidant
markers such as hepatic glutathione (GSH), catalase
(CAT), superoxide dismutase (SOD), and total protein
in a dose-dependent manner, which was conrmed by
histopathological examination. When the CCl
4
was dosed
in rats, the authors observed a rise in levels of AST, which
indicate liver damage, similar to viral hepatitis. Serum
ALP and bilirubin levels were used to determine hepatic
cell function. The increase of ALP levels in serum is due to
overload in synthesis in the presence of increasing biliary
pressure. Bilirubin is one of the most useful clinical
parameters to detect the severity of a necrotic process
and its accumulation is a measure of hepatocyte binding,
conjugation, and excretory capacity. Effective control of
bilirubin levels and alkaline phosphatase activity, by dif-
ferent doses of the extract and fractions, points towards an
early improvement in the secretory mechanism of hepa-
tocytes. A reduction in total protein in serum, observed in
the CCl
4
treated animals, may be associated with a
decrease in the hepatic capacity for protein synthesis.
Hence, a decline in total protein content may be consid-
ered as a useful index for severe cellular dysfunction in
chronic liver diseases. The administration of S. edule
fruits as ethanolic extracts and their different fractions
revealed hepatoprotective activity against the toxic effect
of CCl
4
. These extracts seem to offer protection and
maintain the functional integrity of hepatic cells. Treat-
ment of rats exposed to CCl
4
treated with ethanolic
extracts of chayote fruit (200 mg/kg) and their fractions
(200 mg/kg) exhibited almost normal liver architecture by
reducing changes such as Kupffer cell hyperplasia, inam-
matory cells, apoptosis, microvascular fatty changes, and
centrilobular necrosis. All of the above observations
indicate that the S. edule ethanolic extracts and their
fractions preserved the structural integrity of the hepato-
cellular membrane and showed a dose-dependent pro-
tective effect. The authors concluded that this
hepatoprotective effect may be caused by its potent anti-
oxidant activity and/or by scavenging free radicals and
inhibiting lipid peroxidation. Phytochemical screening of
chayote fruit ethanolic extracts showed the presence of
carbohydrates, avonoids, saponins, glycosides, tannins,
and proteins, whereas the ethyl acetate and n-butanol
fractions indicated the presence of avonoids and saponin
glycosides, respectively.
47.12 Sterols
The total content of sterol was reported by Akihisa et al.
(1986) in aerial sections of a mature plant (16 mg/100 g
extracted with MeOH under reux) and pericarp (38 mg/
100 g extracted with CH
2
Cl
2
in a Soxhlet extractor) of
chayote purchased locally in Japan. Out of the total sterol
mixture obtained, 23 sterols were identied as authentic
compounds by gas liquid chromatography (GLC) on an
OV-17 glass capillary column, and further by high-reso-
lution mass spectra (MS) and proton nuclear magnetic
resonance (1H NMR) and carbon-13 nuclear magnetic
resonance (13C NMR) spectroscopy for some isolated
sterols (Table 47.8).
47.13 Triterpenes and Cucurbitacins
Akihisa et al. (1988) focused on the compositions of the
triterpene alcohol fraction of the unsaponiable lipids
from S. edule seed, leaves, and stems of the mature plants
by GLC and MS. Fourteen triterpene alcohols were iden-
tied in both aerial sections and pericarp (Table 47.9).
Table 47.8 Sterol composition of Sechium edule
Sterols Aerial part
(%)
Pericarp
(%)
Cholesterol (cholest-5-enol) 0.3 0.5
24-Methylcholesta-5,22-dienol 0.1
Desmosterol (cholesta-5,24-dienol) 0.3 0.1
24-Methylcholesterol 0.2 0.1
24-Methylencholesterol +24-
methylcholesta-7,22-dienol
0.5 0.1
24-Ethylcholesta-5,22-dienol 0.6 0.1
24-Ethylcholesta-5,22,25-trienol 0.1
24-Ethylcholesta-8,22-dienol 0.8 1.1
24-Methylcholest-7-enol 2.9 4.3
24-Methylencholest-7-enol 0.3 0.5
24-Ethylcholesterol +24-ethylcholesta-
8,22,25-trienol +24-ethylcholesta-5,25-
dienol
1.5 1.4
24-Ethylcholesta-7,22-dienol 59.5 23.0
24-Ethylcholesta-8,25-dienol 1.0 2.4
24-Ethylcholesta-7,22,25-trienol 6.7 15.2
28-Isofucosterol (24Z-
ethylidenecholesterol)
0.2 1.7
24-Ethylcholest-7-enol +24-ethylcholesta-
7,25-dienol
22.8 46.2
28-Isoavenasterol (24E-ethylidenecholest-
7-enol)
0.4
Avenasterol (24Z-ethylidenecholest-7-enol) 1.4 1.1
Peposterol (24-ethylcholesta-7,24-dienol) 0.2
Others 0.9 1.5
Source: Akihisa et al., 1986.
986 Fruit and Vegetable Phytochemicals
C47 05/01/2017 14:24:35 Page 987
According to Cadena Iñiguez et al. (2011), cucurbita-
cins are tetracyclic triterpenoids found in S. edule plant
and fruits; these compounds provide a characteristic
bitter avor (Yang et al., 2015b). Structurally, cucurbita-
cins are characterized by the tetracyclic cucurbitane
nucleus skeleton (triterpenes) based on isoprene units.
Triterpenes are accordingly C30 compounds. Cucurbita-
cins are derivatives of the triterpene hydrocarbon cucur-
bitane named 19-(10 9-β)-abeo-5 alpha-lanostane (also
known as 9-β-methyl-19-nor-lanosta-5-ene), with a vari-
ety of oxygen substitutions (Alghasham, 2013). Different
cucurbitacin compounds exhibit antitumor proliferation
inhibition capacity, and induce apoptosis alone or syner-
gistically with other proven anticancer chemicals and
cytokines (Alghasham, 2013; Kim and Kim, 2015).
Because of their hydrophobic properties, polymeric
micellar systems exhibited a better antitumor efcacy
because of their solubilization and targeting capacity.
The varieties with the highest cucurbitacins com-
pounds were the wild relative (1.456 mg/g), followed
by nigrum levis (0.66 mg/g), nigrum xalapensis
(0.195 mg/g), nigrum spinosum (0.190 mg/g), and nally
virens levis (commercial variety, 0.116 mg/g). The yel-
low varieties had lower concentrations than the domes-
ticated green varieties, and much lower than the wild
fruit, with values ranging from 0.027 to 0.088 mg/g
(Cadena Iñiguez and Arévalo Galarza, 2011; Cadena
Iñiguez et al., 2011). Therefore, the bitter avor of
wild relative chayote may be based on a higher content
of triterpenic compounds (cucurbitacins), while the
slightly sweet avor in yellow chayote fruits may be
related to higher amounts of soluble solids, lower tri-
terpenics, and acidity. Cadena Iñiguez and Arévalo
Galarza(2011)displayacompletelistofcucurbitacins
registered in eight varieties of chayote, including Cu
B,
Cu
E, Cu
P, and Cu
Q1 glycosides, dihydrocucurbita-
cin-Q1, dihydroisocucurbitacin-I, glycocucurbitacin-I,
dihydrocucurbitacin-D, isocucurbitacin-D, dihydroiso-
cucurbitacin-E, hydrocucurbitacin-E, isocucurbitacin-
B, dihydroisocucurbitacin-B, cucurbitacin-L, cucurbita-
cin-E, and cucurbitacin-B.
Additionally, Cadena Iñiguez et al. (2011) remarked
that higher cucurbitacins content may also be related to
the viviparous behavior of chayote fruits; the seed of wild
relative chayote is not viviparous, unlike that of the
domesticated green and yellow varieties. This may be
attributed to a certain adjustment in the content of
cucurbitacin and gibberellins (GAs); since cucurbitacins
have been recognized as antagonists of GA, a hormone
related to the germination process, particularly in S. edule,
this might suggest that the higher the cucurbitacin con-
tent in the fruits, the lower the GA content, and conse-
quently the lower precociousness in germination. In this
regard it is noteworthy that the wild relative, the only wild
variety, registered the highest cucurbitacin contents and
does not display viviparity.
47.14 Antioxidant and Antiradical
Activity
The content of various antioxidant substances in acid
hydrolysates of green and yellow chayote have been
characterized by Chao et al. (2014); they observed signi-
cantly higher levels of myricetin, especially in yellow
chayote (Table 47.10). Sulaiman and Ooi (2013) found
that the ethyl acetate extract of S. edule showed the
highest 1,1-diphenyl-2-picrylhydrazyl (DPPH) free radi-
cal scavenging activity (951.73 ±29.14 mM Trolox equiv-
alents/g extract) among seven cucurbit fruit vegetables
(Benincasa hispida,Cucurbita maxima,Lagenaria sicera-
ria,Luffa acutangula,Momordica charantia,Sechium
edule and Trichosanthes cucumerina). Additionally, the
highest correlation (r=0.99) was observed between the
total phenolic contents and DPPH values of chayote.
Gallic acid was identied as both the main and most
active antioxidant constituent in the ethyl acetate extract
of S. edule, followed by caffeic acid and isoquercetin.
Similarly, Ordoñez et al. (2006) investigated the anti-
oxidant capacity of water and ethanolic extracts of S. edule
leaves and seeds against the coupled oxidation of
Table 47.9 Triterpene alcohols composition of Sechium edule
Triterpene alcohols Aerial part
(%)
Pericarp
(%)
Euphol (euphfa-8,24-dienol) 1.6
Tirucallol (tirucalla-8,24-dienol) 1.2
Cycloartanol (9β,19-cyclolanostanol) +
taraxerol (D-friedoolean-14-enol)
0.4 1.4
β-Amyrin (olean-12-enol) 28.5 33.5
Butyrospermol (eupha-7,24-dienol) 1.0
Isomultiorenol (D:C-friedoolean-8-enol) 1.3 2.4
24-Methylene-24-dihydrolanosterol (24-
methylenelanost-8-enol)
0.2 1.8
α-Amyrin (urs-12-enol) 31.5 11.0
Cycloartenol (24-dehydrocycloartenol) 31.5 8.9
Lupeol (lup-20(29)-enol) 4.6 23.4
24-Methylene-24-dihdyroparkeol (24-
methylenelanost-9(11)-enol)
1.1
24-Methylenecycloartanol 15.0 9.6
Multiorenol (D:C-friedoolean-7-enol) 3.5 1.0
Others 3.8 1.1
Source: Akihisa et al., 1988.
47 Chayote (Sechium edule (Jacq.) Swartz) 987
C47 05/01/2017 14:24:35 Page 988
β-carotene and linoleic acid, as well as the possible anti-
oxidative effect, with a view to its use as a natural preserv-
ative in foods or functional food. The results showed that
the highest antioxidant activity was obtained with an
alcoholic based extraction of dry leaves and seeds (values
from 80% to 91%), whereas stem extracts showed the
lowest antioxidant activity (values between 30% and
65%). Consistently, there was a strong inhibitory activity
by β-carotene bleaching (antioxidant activity values of
90%) as well as a strong reducing effect by a linoleate
model. The bioactive components of S. edule extracts can
act as primary and secondary antioxidants by donating
electrons and scavenging free radicals. Therefore, they can
reduce or inhibit the lipid peroxidation, which makes the
extract suitable as a biopreservative in food or as health
supplement and functional food, since it alleviates oxida-
tive stress. Finally, the authors suggested that the phenolic
compounds avonoids such as avonol in the extracts
could be responsible for the antioxidant effects; however,
this is not predicted based on total phenolic content.
Monroy-Vázquez et al. (2009) and Cadena Iñiguez et al.
(2013) examined the in vitro antiproliferative activity of
crude ethanolic extracts from eight variety groups of S.
edule. The extracts were assessed for their possible anti-
tumoral activity by cell nuclei with crystal violet staining
on three cancer cell lines: human cervical carcinoma
(HeLa), mouse lung brosarcoma (L929), and mouse
macrophage leukemia (P388). The effect of the extracts
on tumor cell line proliferation depended on the variety
group, dose, and cell line. The commercial (S. edule var.
virens levis and nigrum spinosum) and wild-type chayote
exhibited an antiproliferative activity against all tested cell
lines. On the other hand, the S. edule var. albus dulcis and
albus minor extracts did not have a signicant effect on
L929 and P388 cells. The half-maximum inhibitor con-
centration (IC
50
) of these extracts ranged from 0.5 to
0.9 mg/mL; these were the most effective doses to validate
the antitumor properties of chayote against HeLa, L929,
and P388 cell lines. These antiproliferative effects may be
associated with the sechiumin protein of S. edule, which
was reported by Wu et al. (1998) as an inhibitor of protein
synthesis of HeLa cells.
Other biological effects of chayote extracts have been
studied, including the biochemistry of blood and diabetes
in female Wistar rats using radiopharmaceuticals such as
technetium-99m 99mTcO4Na(Diré et al., 2007b). The
distribution, uptake, retention, and elimination of radio-
pharmaceuticals depend on several factors, such as blood
ow, tissue metabolism, and the elements binding to
blood. However, the data suggest that S. edule extracts
were capable of normalizing the pancreatic uptake of
99mTcO4Na in treated animals. In comparison to the
diabetic animals, the extracts were capable of reducing
the radiopharmaceutical uptake in pancreas of treated
rats. Likewise, chayote extracts were altered by radio-
labeling blood elements and the biodistribution of
Table 47.10 Antioxidant contents in chayote leaves of the yellow and green varieties
Antioxidant Units Chayote, green
leaves
Chayote, yellow
leaves
Polyphenols
a)
mg GAE/g dw 2.62 ±0.52 0.63 ±0.18
Flavonoids
b)
mg QE/g dw 4.27 ±0.14 1.87 ±0.23
Flavonols
b)
mg QE/g dw 1.00 ±0.22 2.22 ±0.42
Quercetin μg/g dw 64.86 ±2.86
Myricetin μg/g dw 756.13 ±49.99 1010.54 ±31.05
Morin μg/g dw 194.99 ±6.89 404.38 ±82.33
Kaempferol μg/g dw 36.36 ±5.85
Anthocyanidins unit/g dw 1.42 ±1.09 1.12 ±0.05
DPPH scavenge
c)
IC
50
μg/mL 1801.56 ±3.16 1503.96 ±2.57
TEAC methanolic μmol Trolox/g dw 21.70 ±2.91 18.09 ±0.80
TEAC ethanolic μmol Trolox/g dw 2.32 ±0.29 1.28 ±0.24
ORAC hydrophilic μmol Trolox/g dw 49.95 ±7.94 120.29 ±6.91
ORAC lipophilic μmol Trolox/g dw 38.97 ±0.27 34.30 ±0.82
a) Total phenolics expressed as milligrams gallic acid equivalents (GAE) per gram dry weight (dw).
b) Total avonoids and total avonols expressed as milligrams quercetin equivalents (QE) per gram dry weight (dw).
c) IC
50
: half-maximal inhibitory concentration.
Source: Chao et al., 2014.
988 Fruit and Vegetable Phytochemicals
C47 05/01/2017 14:24:35 Page 989
99mTcO4Na in pancreas. This effect could be related to the
presence of compounds with oxidant properties, which
could be produced by liver metabolism of chayote extracts
and by the generation of advanced glycation end products
(AGEs) in diabetics. These active metabolites could act
directly or indirectly by generation of reactive oxygen
species. The oxidative effect may induce a decrease in the
level of glucose and globulin fractioning. Moreover, the
same protective effect was observed against the lethal
action of stannous chloride (SnCl
4
) using colonies of E.
coli (AB1157). Stannous ions could be oxidized and,
consequently, avoid the generation of active oxygen spe-
cies (Diré et al., 2007a).
On the other hand, methanolic extractsof S. edule leaves,
seeds, and stems have also been reported as potential
antibacterial agents, displaying a large spectra of activities
against both Gram-positive (Ordoñez et al., 2003) and
Gram-negative bacteria (Noumedem et al., 2013). All
methanolic extracts showed inhibitory effects against 29
tested multidrug resistant Gram-negative bacteria (100%
activity against clinical strains of Providencia stuartii,
Pseudomonas aeruginosa,Klebsiella pneumoniae,Escher-
ichia coli,Enterobacter aerogenes, and Enterobacter cloa-
cae), and were particularly active against four Gram-
positive bacteria (Staphylococcus aureus,Enterococcus
faecalis,Streptococcus agalactiae,andStreptococcus pyo-
genes). The extracts were more active than chlorampheni-
col. Additionally, the antibacterial properties of chayote
leaves were related to the presence of a large variety of
secondary metabolites, present in small quantities.
47.15 Concluding Remarks
The chayote plant is an important source of various
bioactive compounds, with potential pharmacological
activities attributed to its phytochemical composition.
The whole chayote plant is nutritionally important for
the human diet. Although the edible parts of S. edule
are relatively low in proteins, vitamins, fatty acids, and
sodium content compared to other vegetables, the
immature fruit possess important levels of vitamins
such as folate and vitamins C and E. The macro-
nutrient content of the fruits is adequate; however,
the micronutrients exhibit low levels compared to
other cucurbit fruits. It is a good source of potassium
and phosphorus. The shoots are more nutritious than
the fruits because of their high levels of iron and
phosphorus and because they represent a good source
of carotenes and calcium. The protein contents in raw
sections have a tendency to diminish after cooking
(from 34% to 2.8%). Additionally, chayote has a high
caloric and carbohydrate content; however, it seems to
be more starch or crude ber based than sugars,
especially in young stems, root, and seed but not in
esh. Consequently, the dietary ber content in cha-
yote fruits is similar to those reported for many veg-
etables, but without their high caloric content, which
makes them suitable for controlled diets.
In addition, chayote fruits, leaves, and roots have also
shown several health benets. The number of studies that
prove their diuretic, anti-inammatory, and hypotensive
properties is increasing. Treatments for kidney stones
include chayote to help their elimination, and it is used as
a complementary therapy in the treatment of athero-
sclerosis and hypertension; it can relieve intestinal and
cutaneous inammation and promote ulcer cauteriza-
tion. The antibacterial and antioxidant properties of the
chayote plant have also been reported, showing that S.
edule contains a large variety of secondary metabolites,
which are involved in many of the plants pharmaco-
logical effects.
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992 Fruit and Vegetable Phytochemicals
... In this regard, our results coincide since a diminution of BP in our experimental group was observed three months after treatment. Such an effect can be attributed to the presence of flavonoids with C-glycosidic, O-glycosidic or quercetin, with vasodilator and antidepressant effects [33]. In addition, it has been reported that Zucker rats with characteristics similar to MetS such as obesity, dyslipidemia, insulin resistance, and hypertension, improve with chronic high doses of quercetin [34]. ...
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