ArticlePDF Available

Biological Activities of Plant Pigments Betalains


Abstract and Figures

Abstract Betalains are a family of natural pigments present in most plants of the order Caryophyllales. They provide colors ranging from yellow to violet to structures that in other plants are colored by anthocyanins. These include edible fruits and roots but also flowers, stems, and bracts. The recent characterization of different bioactivities in experiments with betalain-containing extracts and purified pigments has renewed the interest of the research community in these molecules used by the food industry as natural colorants. Studies with multiple cancer cell lines have demonstrated a high chemopreventive potential that finds in vitro support in a strong antiradical and antioxidant activity. Experiments in vivo with model animals and bioavailability studies reinforce the possible role played by betalains in the diet. This work provides a critical review of all the claimed biological activities of betalains, showing that the bioactivities described might be supported by the high antiradical capacity of their structural unit, betalamic acid. Although more investigations with purified compounds are needed, the current evidences suggest a strong health-promoting potential.
Content may be subject to copyright.
Biological Activities of Plant
Pigments Betalains
Departamento de Bioqu
ımica y Biolog
ıa Molecular A, Unidad Docente de Biolog
ıa, Facultad de Veterinaria,
Regional Campus of International Excellence “Campus Mare Nostrum,” Universidad de Murcia, Espinardo,
Murcia, Spain
Betalains are a family of natural pigments present in most plants of the order Caryophyllales. They provide colors ranging
from yellow to violet to structures that in other plants are colored by anthocyanins. These include not only edible fruits and
roots but also flowers, stems, and bracts. The recent characterization of different bioactivities in experiments with betalain
containing extracts and purified pigments has renewed the interest of the research community in these molecules used by
the food industry as natural colorants. Studies with multiple cancer cell lines have demonstrated a high chemopreventive
potential that finds in vitro support in a strong antiradical and antioxidant activity. Experiments in vivo with model animals
and bioavailability studies reinforce the possible role played by betalains in the diet. This work provides a critical review
of all the claimed biological activities of betalains, showing that the bioactivities described might be supported by the high
antiradical capacity of their structural unit, betalamic acid. Although more investigations with purified compounds are
needed, the current evidences suggest a strong health-promoting potential.
Keywords Antiradical, biological activity, cancer, diet, review
Betalains are water-soluble, nitrogen containing pigments
present in most plants of the order Caryophyllales. These are
divided into two groups: the violet betacyanins and the yellow
betaxanthins. Betacyanins present absorbance spectra centered
at wavelengths of around λ
D536 nm. Glycosylation and
acylglycosilation of one or two hydroxyl groups are possible
in betacyanins, and complex pigment structures can be
obtained (Heuer et al., 1994; Strack et al., 2003; Cai et al.,
2005). In contrast, betaxanthins are yellow and no glycosyla-
tion has ever been reported. The absorbance spectra of betax-
anthins are centered at wavelengths of around λ
D480 nm.
Both groups share betalamic acid as the structural and chro-
mophoric unit. It is condensed with amines and amino acids in
betaxanthins and with cyclo-DOPA in betacyanins (Gand
Herrero et al., 2010a). Figure 1 shows the structures for beta-
lamic acid, the betacyanins aglyca (betanidin), and the general
structure for betaxanthins. The structure for the dopamine-
derived betaxanthin (miraxanthin V) is shown and compared
with other known bioactive metabolites such as resveratrol
and the anthocyanidin cyanidin.
Betalains are found not only in edible parts of plants but
also in leaves (Wang et al., 2007), flowers (Gand
et al., 2009b), stems (Schliemann et al., 1996), and bracts
(Heuer et al., 1994). Anthocyanins and betalains are mutually
exclusive and have never been found together in the same
plant (Stafford, 1994; Brockington et al., 2011); in Caryophyl-
lales only the coloration of the Caryophylaceae and Mollugi-
naceae is due to anthocyanins. Among the Caryophyllales
plants, red beet roots (Beta vulgaris) (Hempel and B
1997), the fruits of cacti belonging to the genus Opuntia
(mainly Opuntia ficus indica) (Felker et al, 2008; Osorio-
Esquivel et al., 2011), the dragon fruits from Hylocereus cacti
(mainly Hylocereus polyrhizus) (Wybraniec and Mizrahi,
2002; Wybraniec et al., 2007), and the Swiss chard (Beta vul-
garis) (Kugler et al., 2004) are known edible sources of beta-
cyanins and betaxanthins. Less common sources are the ulluco
tubers (Ullucus tuberosus) (Svenson et al., 2008), fruits of
Eulychnia cacti (Masson et al., 2011), and the berries from
Address correspondence to Dr. Fernando Gand
ıa-Herrero, Departamento
ımica y Biolog
ıa Molecular A, Unidad Docente de Biolog
ıa, Edificio
Facultad de Veterinaria, Universidad de Murcia, E-30100 Espinardo, Murcia,
Spain. E-mail:
Color versions of one or more of the figures in the article can be found
online at
Critical Reviews in Food Science and Nutrition, 56:937–945, (2016)
Copyright c
OTaylor and Francis Group, LLC
ISSN: 1040-8398 / 1549-7852 online
DOI: 10.1080/10408398.2012.740103
Rivina humilis (Khan et al., 2012). Certain Amaranthus spe-
cies are also consumed as cooked or fresh (Amin et al., 2006;
Sang-Uk et al., 2009). Betalain containing beetroot extracts
are used as the additive 73.40 in the 21 CFR section of the
Food and Drug Administration (FDA) in the United States and
under the E-162 code in the European Union to give a pink or
violet color to foods and beverages (Mart
ınez et al., 2006; Pru-
dencio et al., 2008; Junqueira-Goncalves et al., 2011; Gand
Herrero et al., 2012b). Figure 2 shows pictures of edible prod-
ucts containing betalains. New colorants derived from Opuntia
fruit extracts (Mosshammer et al., 2006; Ob
on et al., 2009;
enz et al., 2009) or containing individual pigments (Gand
Herrero et al., 2010b) have been also proposed. The joint pres-
ence of betaxanthins and betacyanins in the same parts of
plants generates orange to red shades depending on the pig-
ment proportion (Schliemann et al., 2001; Kugler et al., 2004;
ıa-Herrero et al., 2005; Felker et al., 2008). Due to their
hydrophilic nature, betalains are accumulated in the vacuoles
of the cells that synthesize them, mainly in epidermal and sub-
epidermal tissues of plants (Wink, 1997). Interestingly, fungi
of the genera Amanita and Hygrocybe (von Ardenne et al.,
1974; Musso, 1979; Stintzing and Schliemann, 2007; Babos
et al., 2011) produce betalain-related pigments.
In recent years, betalains have shown promising bioactive
potential. Early investigations revealed a strong free radical
scavenging capacity of betalains purified from beetroot (Escri-
bano et al., 1998; Pedre~
no and Escribano, 2001). Subsequent
research revealed the existence of an intrinsic activity present
in all betalains that is modulated by structural factors (Cai
et al., 2003; Gand
ıa-Herrero et al., 2010a). Studies with
different cell lines have demonstrated the potential of betalains
in the chemoprevention of cancer (Wu et al., 2006; Sreekanth
et al., 2007), and experiments in vivo have shown that very
low concentrations of dietary pigments inhibit the formation
of tumors in mice (Kapadia et al. 2003; Lechner et al., 2010).
In humans, the plasma concentration of betalains after inges-
tion is sufficiently high to promote their incorporation into
low-density lipoprotein (LDL) and red blood cells, protecting
them from oxidative damage and hemolysis (Tesoriere et al.
2003, 2005). However, most of the biological activities
described have been reported according to studies with plant
extracts with limited or no pigment purification. Although
these studies are useful in identifying potential activities, iso-
lated compounds are necessary to link the effects described
Figure 1 Structures for betalamic acid, the betacyanins aglyca (betanidin), and the general structure for betaxanthins. R
and R
are side residues present in
amines or amino acids. For comparative purposes, the structure for the diphenolic betaxanthin Miraxanthin V is also shown together with resveratrol and
Figure 2 Pictures of the best known sources of betalains: (A) Opuntia ficus-
indica fruit, and (B) Beta vulgaris root. (C) An encapsulated commercial col-
orant based on B. vulgaris root extracts, and (D) a dairy product containing B.
vulgaris extracts.
938 F. GAND
with the structures responsible. In this work, the biological
activities of betalains are exhaustively reviewed, considering
in vitro and in vivo experiments developed since the early
description of its free radical scavenging activity more than a
decade ago.
Free Radical Scavenging and Antioxidant Activities
Since the introduction of a feasible technology to determine
the free radical scavenging potential of molecules and extracts
by the Rice-Evans group (Re et al., 1999), the ABTS (2,2’-azi-
nobis-(3-ethylbenzothiazoline-6-sulfonic acid)) radical assay
has become a standard technique in the evaluation of this
activity. In betalains, the ABTS radical assay has gained rele-
vance with respect to other similar methodologies such as the
DPPH (2,2-diphenyl-1-picrylhydrazyl) radical assay (Brand-
Williams et al., 1995) and the Oxygen Radical Absorbance
Capacity (ORAC) assay (Ou et al., 2001), or the direct reduc-
tion of Fe(III) to Fe(II) through the Ferric Reducing Antioxi-
dant Power (FRAP) assay (Benzie and Strain, 1996). This is
due to the use of a fully aqueous medium, the possibility of pH
variation, and the lack of signal interferences with fluorescent
probes. Antioxidant and antiradical concepts are frequently
not differentiated in the literature. It can be considered that
antiradicals or radical scavengers are antioxidant molecules
measured experimentally in the reduction of a radical. In this
section the term antiradical will be used when the activity has
been assessed through a radical-based assay (ABTS, DPPH, or
ORAC) independent of the original terms used by the authors.
The antioxidant term will be restricted to experiments not
involving stable radicals.
The first investigations that demonstrated a radical scaveng-
ing capacity in betalains were carried out separately with beta-
cyanins and betaxanthins, extracted from Beta vulgaris
(Escribano et al., 1998). Other works demonstrated activities
in betalains purified from different sources: Opuntia ficus-ind-
ica (Butera et al., 2002), B. vulgaris roots grown under axenic
conditions (Pavlov et al., 2002), and plants from Amarantha-
ceae (Cai et al., 2003). In all cases the radical scavenging
activity determined was higher than that detected for other
well-known compounds such as ascorbic acid, catechin, and
Trolox. Glucosylation from betacyanins was demonstrated to
reduce the activity of pigments (Cai et al., 2003) due to the
blockage of one of the hydroxyl groups. However, the pres-
ence of these groups is not necessary to express activity, in
contrast to the results obtained for flavonoids, where the
absence of activity has been described in dehydroxylated com-
pounds trans-chalcone, flavone, flavanone, and isoflavone
(Cai et al., 2006). Thus, an “intrinsic activity” exists in all
betalains studied, which can be enhanced by the presence of
hydroxyl groups, with an increase in terms of Trolox Equiva-
lent Antioxidant Capacity (TEAC) from 2.5 units to 4.0 units
for one hydroxyl group and to 5.8 units for two hydroxyl
groups (Gand
ıa-Herrero et al., 2009a). The last value is higher
than those found for other well-known antioxidants such as
epigallocatechin gallate (EGCG) present in green tea (Rice-
Evans et al., 1996; Stewart et al., 2005). Betalamic acid is the
simplest structure with betalain properties and it also possesses
antiradical and antioxidant activities, with a TEAC value of
2.7 units (Gand
ıa-Herrero et al., 2012a). Thus, betalamic acid
can be considered as the bioactive unit of these pigments
(Fig. 1). Structure–activity relationships in betalains show that
the antiradical activity for the simplest pigments is enhanced
by the connection of the betalain characteristic electron reso-
nance system with an aromatic ring, thus increasing the TEAC
value by 0.4 units. If this is done to form indoline-like sub-
structures, similar to those present in betacyanins, the
enhancement is higher, increasing the TEAC value by
1.6 units (Gand
ıa-Herrero et al., 2010a).
Although the actual contribution of individual pigments is
difficult to be established, the free radical scavenging activity
of betalain containing extracts has been determined in several
cases. The fruits of cactus Hylocereus polyrhizus have
revealed a strong free radical scavenging activity that was
higher in peel extracts than in those obtained from flesh, con-
sistent with a higher content of betalains and flavonoids in the
peel (Wu et al., 2006). Multiple clones of Opuntia plants
showed that the TEAC values for cactus juices were close to
those of red wine and green tea infusions (Stintzing et al.,
2005). In addition to O. ficus-indica, recent attention has been
focused on the fruits of other edible species like O. joconostle
and O. macrorhiza (Moussa-Ayoub et al., 2011; Osorio-Esqui-
vel et al., 2011; Morales et al., 2012). A complex profile of
bioactive substances, including betalains, has been described
in these species, with extracts showing high antiradical and
antioxidant activities. When purified betalain fractions were
obtained, these exhibited higher activity than the flavonoid
and phenolic containing fractions obtained from the same
fruits (Osorio-Esquivel et al., 2011). Betalains from Rivina
humilis were also partially purified and the activity of betacya-
nin- and betaxanthin-rich fractions were assayed (Khan et al.,
2012). This confirmed both antiradical and antioxidant activi-
ties of the pigments and explained the antiradical effect
detected in fruit extracts. In the case of B. vulgaris, extracts
from hairy root cultures have revealed a higher radical scav-
enging potential than intact plants. It has been proposed that
this is due to an increased concentration of phenolic com-
pounds, which may have a synergistic effect with betalains
(Georgiev et al., 2010).
The redox properties of betalains have been studied by
cyclic and differential pulse voltammetry. Reduction poten-
tials were determined for purified indicaxanthin and betanin,
and were higher in the case of the betacyanin (Butera et al.,
2002). Strong antioxidant and antiradical betanin activities
have been properly explained in terms of its electron donor
capacity, calculating the bond dissociation energy and the ioni-
zation potential of the molecule (Gliszczy
Swiglo et al.,
2006). This explains the marked pH dependence found in free
radical scavenging experiments. Increased pH values imply
higher activity and higher TEAC values not only in the case of
betanin. A comparative study with 15 natural and synthetic
betalains showed a common trend in the pH dependence of the
free radical scavenging activity (Gand
ıa-Herrero et al.,
2010a). This indicates the existence of a relevant deprotona-
tion equilibrium in the expression of the activity and common
to all betalains. The same tendency was found for free betala-
mic acid, and it has been linked to its nucleophilic capacity,
determining a pK
value of 6.8 (Gand
ıa-Herrero et al., 2012a).
A similar pH dependence of the free radical scavenging activ-
ity has been described for flavonoids. Deprotonation generates
a phenolate anion in these molecules, which is a better electron
donor and, thus, a more effective scavenger (Madsen et al.,
2000; Muzolf et al., 2008).
Relatedtotheirspectroscopic properties and to their
redox capacity to transfer electrons, betalains have been
used as natural dyes in dye-sensitized solar cells (Zhang
et al., 2008). These are one of the most promising devices
for solar energy conversion due to their reduced production
cost and low environmental impact (Narayan, 2012). The
technology has been assayed with betalain containing
extracts of Beta vulgaris roots (Zhang et al., 2008; Calo-
gero et al., 2010), Hylocereus fruits (Ali and Nayan, 2010),
Opuntia fruits (Calogero et al., 2010; Calogero et al.,
2012), and Bouganvillea bracts (Calogero et al., 2010; Her-
nandez-Martinez et al., 2011). While dye-sensitized solar
cells containing anthocyanins and carotenoids as dyes have
shown overall solar energy conversion efficiencies below
1%, betalain containing cells obtain conversion efficiencies
of up to 1.7% under simulated sunlight conditions, which
is comparable to that of natural photosynthesis (Calogero
et al., 2009; Calogero et al., 2010; Zhou et al., 2011). On
optimizing the performance of solar cell and using purified
betanin instead of raw extracts, the energy conversion effi-
ciencies of the cells have recently been raised to 2.7%
(Sandquist and McHale, 2011; Calogero et al., 2012).
Other Activities In vitro
In addition to their potent antioxidant and free radical scav-
enging activities, and probably in relation with them, the beta-
cyanins betanin and betanidin isolated from B. vulgaris were
able to inhibit the peroxidation of linoleic acid and the oxida-
tion of LDL. The effect was higher than that detected for other
known antioxidants such as a-tocopherol and catechin (Kanner
et al., 2001). These activities were assessed considering differ-
ent oxidizers on linoleic acid emulsions and the oxidative sus-
ceptibility of human LDL obtained from healthy volunteers
and microsomes obtained from turkey muscle tissue. Purified
betalains have been also reported to be scavengers of hypo-
chlorous acid, which is the most powerful oxidant produced
by human neutrophils in inflammatory processes and to
interfere with the activity of myeloperoxidase enzyme respon-
sible for its formation (Allegra et al., 2005).
Although the molecules responsible for plant pigmentation
seem to be erroneously identified (Kugler et al., 2004), aque-
ous extracts of Swiss chard (B. vulgaris) containing betalains
were demonstrated to possess inhibitory activity on the
enzyme acetylcholinesterase (Sacan and Yanardag, 2010).
This enzyme is involved in the processes of neurotransmission
by cleaving the neurotransmitter acetylcholine and its inhibi-
tion has been demonstrated to have therapeutic potential in the
treatment of neurological disorders, including Alzheimer’s
disease (Orhan, 2012).
Recent research with different cancer cell lines has demon-
strated a high chemopreventive potential of betalain contain-
ing extracts. The use of purified pigments in some of the
studies justifies the activities assessed and reinforces the bio-
logical potential of betalains. Extracts of Beta vulgaris have
shown high chemopreventive effect in the induced Epstein–
Barr early antigen activation assay in vitro using a cell line
from lymphoma (Kapadia et al., 1996). Extracts significantly
reduced cells’ viability, with the effect being ascribed to beta-
lains. The activity was compared with other plant extracts,
including an anthocyanin-rich cranberry extract, exhibiting
maximum activity. A limited cytotoxicity of B. vulgaris
extracts has also been demonstrated for human prostate and
breast cancer cell lines (Kapadia et al., 2011).
Extracts obtained from Opuntia ficus-indica proved to be
an effective growth inhibitor and apoptosis inductor in sev-
eral cell lines of immortalized ovarian and cervical epithelial
cells and ovarian, cervical, and bladder cancer cells (Zou
et al., 2005). The effect of the cactus pear solution was dose-
and time-dependent. Cactus extracts were able to inhibit
growth of cancer cells and affect their morphology with con-
centrations of 5% of fruit extract in cell culture. A higher
concentration was necessary (10–25%) for apoptotic effect.
However, the use of raw extracts avoided the identification of
active compounds. Betalains from the berries of Rivina
humilis were tested regarding their effect on hepatocellular
carcinoma cells (Khan et al., 2012). In this case, the betalains
were partially purified and betaxanthins and betacyanins sep-
arately showed dose-dependent cytotoxicity after 24 and
48 hours respectively.
Extracts from the fruit of the cactus Hylocereus polyrhizus
revealed an inhibitory activity of growth of melanoma cancer
cells (Wu et al., 2006). The inhibition was dose-dependent
and higher in peel than in flesh extracts, in relation with a
higher content of flavonoids and betalains. Remarkably, in the
same experiment pure betanin was assayed, revealing a strong
inhibition of the proliferation of melanoma cancer cells. Pure
betanin was also used against a human chronic myeloid leuke-
mia cell line in a different study (Sreekanth et al., 2007). The
940 F. GAND
addition of betanin implied the inhibition of cell growth in a
dose-dependent manner with an IC
of 40 mM after 24 hours
of incubation. Betanin enters the cells and alters the mitochon-
drial membrane integrity. This ultimately leads to the activa-
tion of caspases and nuclear disintegration. The biochemical
alterations were reflected in morphological changes in cells,
which were followed by scanning and transmission electron
microscopy and flow cytometry. Cells entered apoptosis, in a
clear in vitro demonstration of anti-cancer potential of
betalains. Table 1 summarizes all the studies developed with
betalains, which involved cancer cells, identifying the corre-
sponding cell lines.
In addition to the effect on cancer cell lines, B. vulgaris
extracts have been found to dose-dependently suppress the
degradation of tryptophan and the production of neopterin in
human peripheral blood mononuclear cells involved in the
inflammatory response (Winkler et al., 2005). Cells were
obtained from healthy donors and stimulated in the presence
of the betalain containing extracts. The cell response was
reduced by the presence of extracts, suggesting that the B. vul-
garis juice possesses compounds with immunosuppressive and
anti-inflammatory activities. Human red blood cells also
showed increased resistance to hemolysis when they were
incubated with growing concentrations of purified indicaxan-
thin and betanin in vitro (Tesoriere et al., 2005). Furthermore,
LDLs isolated from healthy humans have been demonstrated
to incorporate purified indicaxanthin and betanin after ex vivo
incubation (Tesoriere et al., 2003). LDLs enriched with the
pigments were more resistant than native LDLs to copper-
induced oxidation, with indicaxanthin being more effective
than betanin in the protection from oxidative damage.
In vivo anti-tumor formation activity in mouse skin has
been demonstrated for Beta vulgaris extracts. The extracts
were orally administered to the animals in drinking water after
topical tumor induction (Kapadia et al., 1996). Results showed
a significant decrease in the incidence and number of papillo-
mas found in the mice skin. In the same study, lung tumor for-
mation was induced to mice, and inhibited by the oral
administration of B. vulgaris extracts. A 60% reduction in the
number of mice with adenomas was observed, with an addi-
tional 30% reduction in the number of tumors for the affected
animals. Skin tumor formation induced chemically and pro-
moted by ultraviolet (UV)-based light was also inhibited after
the oral administration of B. vulgaris extracts in mice (Kapadia
et al., 2003). In addition, animals that followed the treatment
also showed a reduced splenomegaly. In the case of induced
tumors in the liver, oral administration of beet extract reduced
the tumor incidence to 40%, showing a potent cancer chemo-
preventive activity in the model animals.
Analogously, chemoprevention against induced esophageal
carcinogenesis was also demonstrated in rats for B. vulgaris
extracts administered orally (Lechner et al., 2010). Results
showed a reduction of 45% in the number of papillomas, limit-
ing cell proliferation, angiogenesis, and inflammation. It was
hypothesized that betalains antioxidant activity reduced the
level of reactive oxygen species to levels that were too low to
stimulate the anomalous proliferation.
In addition, dose-dependent protection of the adverse
effects of g-ray irradiation in rats has been demonstrated in
vivo for Beta vulgaris extracts, and the effect has been
ascribed to betalains (Lu et al., 2009). Orally administered
extracts partially restored the normal biochemical levels of the
altered parameters caused by irradiation, including catalase,
superoxide dismutase, and lipid oxidation activities in liver,
spleen, and kidney. Furthermore, the prevention of decline in
the spleen and thymus index in irradiated mice led the authors
to suggest that betalains could partially restore the immuno-
logical function and improve the pathological status in vivo.
Although the effect of betalains cannot be identified, aque-
ous extracts of Opuntia ficus-indica were able to inhibit tumor
Table 1 Studies with betalain containing extracts involving cancer cells. The cell line identification, the source of betalains, the use of purified pigments, and the
corresponding references are shown in each case
Cell line Betalains source Effect Pigment purification References
Raji (lymphoma, human) Beta vulgaris Reduced cell viability No Kapadia et al., 1996
IOSE (ovarian epithelium, human) Opuntia ficus-indica Growth inhibition No Zou et al., 2005
OVCA420 (ovarian cancer, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
SKOV3 (ovarian cancer, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
TCL-1 (cervical epithelium, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
HeLa (cervical cancer, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
Me180 (cervical cancer, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
UM-UC-6 (bladder cancer, human) Opuntia ficus-indica Growth inhibition No Zou et al., 2005
T24 (bladder cancer, human) Opuntia ficus-indica Growth inhibition, apoptosis induction No Zou et al., 2005
B16F10 (melanoma, mouse) Hylocereus polyrhizus Growth inhibition No Wu et al., 2006
B16F10 (melanoma, mouse) Unspecified (commercial) Growth inhibition Purified betanin Wu et al., 2006
K562 (leukemia, human) Opuntia ficus-indica Growth inhibition, apoptosis induction Purified betanin Sreekanth et al., 2007
PC-3 (prostate cancer, human) Beta vulgaris Growth inhibition No Kapadia et al., 2011
MCF-7 (breast cancer, human) Beta vulgaris Growth inhibition No Kapadia et al., 2011
HepG2 (liver cancer, human) Rivina humilis Reduced cell viability Partial Khan et al., 2012
growth in a nude mouse of ovarian cancer model compared
with untreated animals (Zou et al., 2005). The extract was
administered by injection and compared with the chemo-
preventive agent N-(4-hydroxyphernyl) retinamide (4-HPR),
used in ovarian cancer clinical trials. Both cactus pear extracts
and chemopreventive agent reduced the tumor size in a com-
parable manner. Opuntia fruits’ extracts have also demon-
strated their potential in the protection and recovery of the
liver after damage has been induced. Hepatotoxicity by carbon
tetrachloride (CCl
) in rats was limited in animals fed with the
betalain containing extracts both after and before the damag-
ing treatment (Galati et al., 2005). The oral administration of
the extract promoted liver recovery at histochemical and bio-
chemical levels. The same effect was described for betalain
containing extracts of whole plants of Amaranthus spinosus,
and it has been proposed that the mechanism of hepatoprotec-
tion was due to its antioxidant activity (Zeashan et al., 2008,
Interestingly, other activities have been also described for
betalains. Purified pigments extracted from Portulaca olera-
cea have demonstrated their capacity to reverse induced learn-
ing and memory impairments produced by D-galactose in
mice (Wang and Yang, 2010). In comparison with ascorbic
acid, orally administered betalains showed a more pronounced
effect in ameliorating cognition deficits in mice and restored
the normal biochemical levels of relevant enzymes, being
proposed a neuroprotective effect. Extracts of Amaranthus spi-
nosus and Boerhaavia erecta containing betalains have dem-
onstrated antimalarial activity in an in vivo model assay in
mice (Hilou et al., 2006). The aqueous extracts were able to
inhibit the growth of inoculated parasites in a dose-dependent
manner. These plants are used in the traditional medicine
against Plasmodium falciparum infections (malaria) in
humans. The authors pointed to betacyanins betanin and amar-
anthin as possible molecules responsible for the assessed activ-
ity at the same time that they acknowledged the need to
perform experiments with purified compounds. Opuntia ficus-
indica extracts and its main pigment, indicaxanthin, in a pure
form were demonstrated to reduce the contractility of the ileal
longitudinal muscle obtained from mice (Baldassano et al.,
2010, 2011). The authors propose the usefulness of the finding
in the regulation of intestinal motility in related disorders and
describe the mechanism of action. It implies the inhibition of
phosphodiesterase enzymes and the increase in cAMP levels,
which lead to a decrease in intracellular Ca
which ultimately promotes the smooth muscle relaxation.
Bioavailability studies of betalains after oral administration
in humans indicate that model betalains, betanin, and indicax-
anthin remain in the body and are able to play a health-promot-
ing function, thereby improving the body redox status
(Tesoriere et al., 2004; Frank et al., 2005). Maximum plasma
concentrations are reached three hours after consumption,
with a decline corresponding to first-order kinetics. After this
time period, betalains can be incorporated into the red cells in
vivo (Tesoriere et al., 2005). They are completely eliminated
after 12 hours of ingestion, with a urinary excretion of 76% in
the case of betaxanthin, but highly limited in the case of beta-
nin. This indicates metabolization of the pigment and its trans-
formation to other compounds, including betalamic acid, as
demonstrated in simulated digestion studies (Pavlov et al.,
2005; Tesoriere et al., 2008). However, the inability of part of
the population to metabolize betanin has also been described,
excreting it to a high level in urine. This is known as beeturia,
and although its mechanism is not well understood, its inci-
dence is high in iron-deficient subjects (Watson et al., 1963;
Sotos, 1999; Mitchell, 2001).
The description of the betalains free radical scavenging
activity implied the renaissance of interest in these molecules
by the research community. Since then, multiple articles with
claims regarding the biological activity of betalains have been
published, including studies on the chemoprevention of tumor
formation. Considering their demonstrated safety (Schwartz
et al., 1983; Khan et al., 2011), the recent bibliography indi-
cates the potential of betalains for food, pharmaceutical, and
cosmetic industries.
Promising results have been reported for tumor prevention
in vivo and the possible role played by betalains in the diet.
However, the use of extracts limits the conclusions drawn, the
hypothesis on the mechanisms involved, and the therapeutic
potential of the assays. Currently, studies with purified pig-
ments are scarce but they provide exciting conclusions.
Increasing of these studies would help to establish the actual
role played by betalains alone or in cooperation with other
compounds. Caution must be taken regarding the possible
application of biological activities described in natural mole-
cules, but the results for betalains are promising in terms of
their health-promoting potential.
The authors acknowledge the financial support of Ministerio
de Ciencia e Innovaci
on (MICINN, FEDER, Spain, project
AGL2011-25023 and AGL2014-57431) and Fundaci
oneca, Agencia de Ciencia y Tecnologia de la Regi
on de
Murcia (Plan Regional de Ciencia y Tecnologia 2007/2010,
Programa de Ayudas a Grupos de Excelencia de la Regi
on de
Murcia). F. Gand
ıa-Herrero has a contract with the “Programa
on y Cajal” (MICINN, FEDER, Spain).
Ali, R. A. M. and Nayan, N. (2010). Fabrication and analysis of dye-sensitized
solar cell using natural dye extracted from dragon fruit. Int. J. Integr. Eng.
942 F. GAND
Allegra, M., Furtm
uller, P. G., Jantschko, W., Zederbauer, M., Tesoriere, L.,
Livrea, M. A. and Obinger, C. (2005). Mechanism of interaction of betanin
and indicaxanthin with human myeloperoxidase and hypochlorous acid.
Biochem. Biophys. Res. Commun.332:837–844.
Amin, I., Norazaidah, Y. and Hainida, K. I. E. (2006). Antioxidant activity and
phenolic content of raw and blanched Amaranthus species. Food Chem.
Babos, M., Halasz, K., Zagyva, T., Zold-Balogh, A., Szego, D. and Bratek, Z.
(2011). Preliminary notes on dual relevance of ITS sequences and pigments
in Hygrocybe taxonomy. Persoonia.26:99–107.
Baldassano, S., Rotondo, A., Serio, R., Livrea, M. A., Tesoriere, L. and Mul
F. (2011). Inhibitory effects of indicaxanthin on mouse ileal contractility:
Analysis of the mechanism of action. Eur. J. Pharmacol.658:200–205.
Baldassano, S., Tesoriere, L., Rotondo, A., Serio, R., Livrea, M. A. and Mul
F. (2010). Inhibition of the mechanical activity of mouse ileum by cactus
pear (Opuntia ficus indica, L, Mill.) fruit extract and its pigment indicaxan-
thin. J. Agric. Food Chem.58:7565–7571.
Benzie, I. F. F. and Strain, J. J. (1996). The ferric reducing ability of plasma
(FRAP) as a measure of “antioxidant power”: The FRAP assay. Anal. Bio-
Brand-Williams, W., Cuvelier, M. E. and Berset, C. (1995). Use of a free
radical method to evaluate antioxidant activity. LWT Food Sci. Technol.
Brockington, S. F., Walker, R. H., Glover, B. J., Soltis, P. S. and Soltis, D. E.
(2011). Complex pigment evolution in the Caryophyllales. New Phytol.
Butera, D., Tesoriere, L., Di Gaudio, F., Bongiorno, A., Allegra, M., Pintaudi,
A. M., Kohen, R. and Livrea, M. A. (2002). Antioxidant activities of sicilian
prickly pear (Opuntia ficus indica) fruit extracts and reducing properties of
its betalains: Betanin and indicaxanthin. J. Agric. Food Chem.50:6895–
Cai, Y., Sun, M. and Corke, H. (2003). Antioxidant activity of betalains from
plants of the Amaranthaceae. J. Agric. Food Chem.51:2288–2294.
Cai, Y.-Z., Sun, M. and Corke, H. (2005). Characterization and application of
betalain pigments from plants of the Amaranthaceae. Trends Food Sci.
Cai, Y.-Z., Sun, M., Xing, J., Luo, Q. and Corke, H. (2006). Structure-radical
scavenging activity relationships of phenolic compounds from traditional
Chinese medicinal plants. Life Sci.78:2872–2888.
Calogero, G., Di Marco, G., Caramori, S., Cazzanti, S., Argazzic, R. and
Bignozzi, C. A. (2009). Natural dye senstizers for photoelectrochemical
cells. Energy Environ. Sci.2:1162–1172.
Carlo, A. and Bignozzi, C. A. (2010). Efficient dye-sensitized solar
cells using red turnip and purple wild Sicilian prickly pear fruits. Int. J.
Mol. Sci.11:254–267.
Calogero, G., Yumb, J.-H., Sinopoli, A., Di Marco, G., Gr
atzel, M. and
Nazeeruddin, M. K. (2012). Anthocyanins and betalains as light-harvesting
pigments for dye-sensitized solar cells. Sol. Energy.86:1563–1575.
Escribano, J., Pedre~
no, M. A., Garc
ıa-Carmona, F. and Mu~
noz, R. (1998).
Characterization of the antiradical activity of betalains from Beta vulgaris
L. roots. Phytochem. Anal.9:124–127.
Felker, P., Stintzing, F. C., M
ussig, E., Leitenberger, M., Carle, R., Vogt, T.
and Bunch, R. (2008). Colour inheritance in cactus pear (Opuntia ficus-ind-
ica) fruits. Ann. Appl. Biol.152:307–318.
Frank, T., Stintzing, F. C., Carle, R., Bitsch, I., Quaas, D., Strass, G.,
Bitsch, R. and Netzel, M. (2005). Urinary pharmacokinetics of betalains
following consumption of red beet juice in healthy humans. Pharmacol.
Galati, E. M., Mondello, M. R., Lauriano, E. R., Taviano, M. F., Galluzzo, M.
and Miceli, N. (2005). Opuntia ficus indica (L.) Mill. fruit juice protects
liver from carbon tetrachloride induced injury. Phytother. Res.19:796–800.
ıa-Herrero, F., Escribano, J. and Garc
ıa-Carmona, F. (2009a). The role of
phenolic hydroxy groups in the free radical scavenging activity of betalains.
J. Nat. Prod.72:1142–1146.
ıa-Herrero, F., Escribano, J. and Garc
ıa-Carmona, F. (2010a). Structural
implications on color, fluorescence, and antiradical activity in betalains.
ıa-Herrero, F., Escribano, J. and Garc
ıa-Carmona, F. (2012a). Purification
and antiradical properties of the structural unit of betalains. J. Nat. Prod.
ıa-Herrero, F., Garc
ıa-Carmona, F. and Escribano, J. (2005). Floral fluo-
rescence effect. Nature.437:334–334.
ıa-Herrero, F., Jim
enzar, M., Cabanes, J., Escribano, J. and
ıa-Carmona, F. (2009b). Fluorescence detection of tyrosinase activity
on dopamine-betaxanthin purified from Portulaca oleracea (common purs-
lane) flowers. J. Agric. Food Chem.57:2523–2528.
ıa-Herrero, F., Jim
enzar, M., Cabanes, J., Garc
ıa-Carmona, F.
and Escribano, J. (2010b). Stabilization of the bioactive pigment of
Opuntia fruits through maltodextrin encapsulation. J. Agric. Food.
ıa-Herrero, F., Sim
on-Carrillo, A., Escribano, J. and Garc
ıa-Carmona, F.
(2012b). Determination of beet root betanin in dairy products by high-per-
formance liquid chromatography (HPLC). J. Chem. Educ.89:660–664.
Georgiev, V. G., Weber, J., Kneschke, E.-M., Denev, P. N., Bley, T. and Pav-
lov, A. I. (2010). Antioxidant activity and phenolic content of betalain
extracts from intact plants and hairy root cultures of the red beetroot Beta
vulgaris cv. Detroit dark red. Plant Foods Hum. Nutr.65:105–111.
Swiglo, A., Szymusiak, H. and Malinowska, P. (2006). Betanin,
the main pigment of red beet: Molecular origin of its exceptionally high free
radical-scavenging activity. Food Add. Cont.23:1079–1087.
Hempel, J. and B
ohm, H. (1997). Betaxanthin pattern of hairy roots from Beta
vulgaris var. lutea and its alteration by feeding of amino acids. Phytochem-
Hernandez-Martinez, A. R., Estevez, M., Vargas, S., Quintanilla, F. and Rodri-
guez, R. (2011). New dye-sensitized solar cells obtained from extracted
bracts of Bougainvillea glabra and spectabilis betalain pigments by differ-
ent purification processes. Int. J. Mol. Sci.12:5565–5576.
Heuer, S., Richter, S., Metzger, J. W., Wray, V., Nimtzt, M. and Strack, D.
(1994). Betacyanins from bracts of Bougainvillea glabra.Phytochemistry.
Hilou, A., Nacoulma, O. G. and Guiguemde, T. R. (2006). In vivo antimalarial
activities of extracts from Amaranthus spinosus L. and Boerhaavia erecta
L. in mice. J. Ethnopharmacol.103:236–240.
Junqueira-Goncalves, M. P., Cardoso, L. P., Pinto, M. S., Pereira, R. M.,
Soares, N. F. and Miltz, J. (2011). Irradiated beetroot extract as a colorant
for cream cheese. Radiat. Phys. Chem.80:114–118.
Kanner, J., Harel, S. and Granit, R. (2001). Betalains, a new class of dietary
cationized antioxidants. J. Agric. Food. Chem.49:5178–5185.
Kapadia, G. J., Azuine, M. A., Rao, G. S., Arai, T., Iida, A. and Tokuda, H.
(2011). Cytotoxic effect of the red beetroot (Beta vulgaris L.) extract com-
pared to doxorubicin (adriamycin) in the human prostate (PC-3) and breast
(MCF-7) cancer cell lines. Anti. Cancer Agents Med. Chem.11:280–284.
Kapadia, G. J., Azuine, M. A., Sridhar, R., Okuda, Y., Tsuruta, A., Ichiishi, E.,
Mukainake, T., Takasaki, M., Konoshima, T., Nishino, H. and Tokuda, H.
(2003). Chemoprevention of DMBA-induced UV-B promoted, NOR-1-
induced TPA promoted skin carcinogenesis, and DEN-induced phenobarbi-
tal promoted liver tumors in mice by extract of beetroot. Pharmacol. Res.
Kapadia, G. J., Tokuda, H., Kinoshima, T. and Nishino, H. (1996). Chemopre-
vention of lung and skin cancer by Beta vulgaris (Beet) root extract. Cancer
Khan, M. I., Harsha, P. S. C. S., Giridhar, P. and Ravishankar, G. A. (2012).
Pigment identification, nutritional composition, bioactivity, and in vitro can-
cer cell cytotoxicity of Rivina humilis L. berries, potential source of beta-
lains. LWT Food Sci. Technol.47:315–323.
Khan, M. I., Joseph, K. M. D., Muralidhara, Ramesh, H. P., Giridhar, P. and
Ravishankar, G. A. (2011) Acute, subacute and subchronic safety assess-
ment of betalains rich Rivina humilis L. berry juice in rats. Food Chem. Tox-
Kugler, F., Stintzing, F. C. and Carle, R. (2004). Identification of beta-
lains from petioles of differently colored Swiss chard (Beta vulgaris L.
ssp. cicla [L.] alef. cv. bright lights) by high-performance liquid chro-
matography–electrospray ionization mass spectrometry. J. Agric. Food
Lechner, J. F., Wang, L. S., Rocha, C. M., Larue, B., Henry, C., McIn-
tyre, C. M., Riedl, K. M., Schwartz, S. J. and Stoner, G. D. (2010).
Drinking water with red beetroot food color antagonizes esophageal
carcinogenesis in N-nitrosomethylbenzylamine-treated rats. J. Med.
Lu, X., Wang, Y. and Zhang, Z. (2009). Radioprotective activity of betalains
from red beets in mice exposed to gamma irradiation. Eur. J. Pharmacol.
Madsen, H. L., Andersen, C. M., Jørgensen, L. V. and Skibsted, L. H. (2000).
Radical scavenging by dietary flavonoids. A kinetic study of antioxidant
efficiencies. Eur. Food Res. Technol.211:240–246.
ınez, L., Cilla, I., Beltr
an, J. A. and Roncal
es, P. (2006). Comparative
effect of red yeast rice (Monascus purpureus), red beet root (Beta vulgaris)
and betanin (E-162) on colour and consumer acceptability of fresh pork sau-
sages packaged in a modified atmosphere. J. Sci. Food Agric.86:500–508.
Masson, L., Salvatierra, M. A., Robert, P., Encina, C. and Camilo, C. (2011).
Chemical and nutritional composition of copao fruit (Eulychnia acida Phil.)
under three environmental conditions in the coquimbo region. Chil. J. Agric.
Mitchell, S. C. (2001). Food idiosyncrasies: Beetroot and asparagus. Drug
Metab Dispos.29:539–543.
Morales, P., Ram
ırez-Moreno, E., Sanchez-Mata, M. C., Carvalho, A. M. and
Ferreira, I. C. F. R. (2012). Nutritional and antioxidant properties of pulp
and seeds of two xoconostle cultivars (Opuntia joconostle F. A. C. Weber
ex Diguet and Opuntia matudae Scheinvar) of high consumption in Mexico.
Food Res. Int.46:279–285.
Mosshammer, M. R., Stintzing, F. C. and Carle, R. (2006). Evaluation of dif-
ferent methods for the production of juice concentrates and fruit powders
from cactus pear. Innov. Food Sci. Emerg. Technol.7:275–287.
Moussa-Ayoub, T. E., El-Samahy, S. K., Rohn, S. and Kroh, L. W. (2011).
Flavonols, betacyanins content and antioxidant activity of cactus Opuntia
macrorhiza fruits. Food Res. Int.44:2169–2174.
Musso, H. (1979). Pigments of fly agaric, Amanita Muscaria. Tetrahedron.
Muzolf, M., Szymusiak, H., Gliszczy
Swiglo, A., Rietjens, I. M. C. M.
and Tyrakowska, B. E. (2008). pH-dependent radical scavenging capacity
of green tea catechins. J. Agric. Food. Chem.56:816–823.
Narayan, M. R. (2012). Dye sensitized solar cells based on natural photosensi-
tizers. Renew Sustain Energy Rev.16:208–215.
on, J. M., Castellar, M. R., Alacid, M. and Fern
opez, J. A. (2009).
Production of a red-purple food colorant from Opuntia stricta fruits by spray
drying and its application in food model systems. J. Food Eng.90:471–479.
Orhan, I. E. (2012). Current concepts on selected plant secondary metabolites
with promising inhibitory effects against enzymes linked to Alzheimer’s
disease. Curr. Med. Chem.19:2252–2261.
Osorio-Esquivel, O., Alicia-Ortiz-Moreno,
Alvarez, V. B., Dorantes-
L. and Giusti, M. M. (2011). Phenolics, betacyanins and antioxidant activity
in Opuntia joconostle fruits. Food Res. Int.44:2160–2168.
Ou, B., Hampsch-Woodill, M. and Prior, R. L. (2001). Development and vali-
dation of an improved oxygen radical absorbance capacity assay using fluo-
rescein as the fluorescent probe. J. Agric. Food Chem.49:4619–4626.
Pavlov, A., Kovatcheva, P., Georgiev, V., Koleva, I. and Ilieva, M.
(2002). Biosynthesis and radical scavenging activity of betalains during
the cultivation of red beet (Beta vulgaris) hairy root cultures. Z. Natur-
Pavlov, A., Kovatcheva, P., Tuneva, D., Ilieva, M. and Bley, T. (2005). Radi-
cal scavenging activity and stability of betalains from Beta vulgaris hairy
root culture in simulated conditions of human gastrointestinal tract. Plant
Food Hum. Nutr.60:43–47.
no, M. A. and Escribano, J. (2001). Correlation between antiradical
activity and stability of betanine from Beta vulgaris L roots under different
pH, temperature and light conditions. J. Sci. Food Agric.81:627–631.
Prudencio, I. D., Prudencio, E. S., Gris, E. F., Tomazi, T. and Bordignon-Luiz,
M. T. (2008). Petit suisse manufactured with cheese whey retentate and
application of betalains and anthocyanins. LWT.41:905–910.
Re, R., Pellegrini, N., Proteggente, A., Pannala, A., Yang, M. and Rice-Evans,
C. (1999). Antioxidant activity applying an improved ABTS radical cation
decolorization assay. Free Rad. Biol. Med.26:1231–1237.
Rice-Evans, C. A., Miller, N. J. and Paganda, G. (1996). Structure-antioxidant
activity relationships of flavonoids and phenolic acids. Free Rad. Biol. Med.
Sacan, O. and Yanardag, R. (2010). Antioxidant and antiacetylcholinesterase
activities of chard (Beta vulgaris L. var. cicla). Food Chem. Toxicol.
enz, C., Tapia, S., Ch
avez, J. and Robert, P. (2009). Microencapsulation by
spray drying of bioactive compounds from cactus pear (Opuntia ficus-ind-
ica). Food Chem.114:616–622.
Sandquist, C. and McHale, J. L. (2011). Improved efficiency of betanin-based
dye-sensitized solar cells. J. Photochem. Photobiol. Chem.221:90–97.
Sang-Uk, C., Buk-Gu, H., Yong-Seo, P., Dong-Kwan, K. and Shela, G. (2009).
Total phenolics level, antioxidant activities and cytotoxicity of young
sprouts of some traditional Korean salad plants. Plant Food Hum. Nutr.
Schliemann, W., Cai, Y., Degenkolb, T., Schmidt, J. Y. and Corke, H. (2001).
Betalains of Celosia argentea.Phytochemistry.58:159–165.
Schliemann, W., Joy IV, R. W., Komamine, A., Metzger, J. W., Nimtz, M.,
Wray, V. and Strack, D. (1996). Betacyanins from plants and cell cultures
of Phytolacca americana.Phytochemistry.42:1039–1046.
Schwartz, S. J., von Elbe, J. H., Pariza, M. W., Goldsworthy, T. and Pilot, H.
C. (1983). Inability of red beet betalain pigments to initiate or promote hep-
atocarcinogenesis. Food Chem. Toxicol.21:531–535.
Sotos, J. G. (1999). Beeturia and iron absorption. Lancet.354:1032–1032.
Sreekanth, D., Arunasree, M. K., Roy, K. R., Reddy, T. C., Reddy, G. V. and
Reddanna, P. (2007). Betanin, a betacyanin pigment purified from fruits of
Opuntia ficus-indica induces apoptosis in human chronic myeloid leukemia
cell line-K562. Phytomedicine.4:739–746.
Stafford, H. A. (1994). Anthocyanins and betalains: Evolution of the mutually
exclusive pathways. Plant Sci.101:91–98.
Stewart, A. J., Mullen, W. and Crozier, A. (2005). On-line high-performance
liquid chromatography analysis of the antioxidant activity of phenolic com-
pounds in green and black tea. Mol. Nutr. Food Res.49:52–60.
Stintzing, F. C., Herbach, K. M., Mosshammer, M. R., Carle, R., Yi, W., Sell-
appan, S., Akoh, C. C., Bunch, R. and Felker, P. (2005). Color, betalain pat-
tern, and antioxidant properties of cactus pear (Opuntia spp.) clones. J.
Agric. Food Chem.53:442–451.
Stintzing, F. and Schliemann, W. (2007). Pigments of fly agaric (Amanita mus-
caria). Z. Naturforsch.62c:779–785.
Strack, D., Vogt, T. and Schliemann, W. (2003). Recent advances in betalain
research. Phytochemistry.62:247–269.
Svenson, J., Smallfield, B. M., Joyce, N. I., Sanson, C. E. and Perry, N. B.
(2008). Betalains in red and yellow varieties of the andean tuber crop ulluco
(Ullucus tuberosus). J. Agric. Food Chem.56:7730–7737.
Tesoriere, L., Butera, D., Allegra, M., Fazzari, M. and Livrea, M. A. (2005).
Distribution of betalain pigments in red blood cells after consumption of
cactus pear fruits and increased resistance of the cells to ex vivo induced
oxidative hemolysis in humans. J. Agric. Food. Chem.53:1266–1270.
Tesoriere, L., Butera, D., D’Arpa, D., Di Gaudio, F., Allegra, M., Gentile, C.
and Livrea, M. A. (2003). Increased resistance to oxidation of betalain-
enriched human low-density lipoproteins. Free. Radic. Res.37:689–696.
Tesoriere, L., Butera, D., Pintaudi, M., Allegra, M. and Livrea, M. A. (2004).
Supplementation with cactus pear (Opuntia ficus-indica) fruit decreases oxi-
dative stress in healthy humans: A comparative study with Vit C. Am. J.
Clin. Nutr.80:391–395.
944 F. GAND
Tesoriere, L., Fazzari, M., Angileri, F., Gentile, C. and Livrea, M. A. (2008).
In vitro digestion of betalainic foods. Stability and bioaccessibility of betax-
anthins and betacyanins and antioxidative potential of food digesta. J. Agric.
Food Chem.56:10487–10492.
von Ardenne, R., D
opp, H., Musso, H. and Steiglich, W. (1974). (ber das
Vorkommen von Muscaflavin bei Hygrocyben (Agaricales) und seine Dihy-
droazepin-Struktur. Z Naturforsch.29c:637–639.
Wang, C.-Q., Chen, M. and Wang, B.-S. (2007). Betacyanin accumulation in
the leaves of C3 halophyte Suaeda salsa L. is induced by watering roots
with H
.Plant. Sci.172:1–7.
Wang, C.-Q. and Yang, G.-Q. (2010). Betacyanins from Portulaca olera-
cea L. ameliorate cognition deficits and attenuate oxidative damage
induced by D-galactose in the brains of senescent mice. Phytomedicine.
Watson, W. C., Luke, R. G. and Inall, J. A. (1963). Beeturia: Its incidence and
a clue to its mechanism. Brit. Med. J.5363:971–973.
Wink, M. (1997). Compartmentation of secondary metabolites and xenobiotics
in plant vacuoles. Adv. Bot. Res.25:141–169.
Winkler, C., Wirleitner, B., Schroecksnadel, K., Schennach, H. and Fuchs,
D. (2005). In vitro effects of beet root juice on stimulated and unstimu-
lated peripheral blood mononuclear cells. Am.J.Biochem.Biotech.
Wu, L.-C., Hsu, H.-W., Chen, Y.-C., Chiu, C.-C., Lin, Y.-I. and Ho, J.-A. A.
(2006). Antioxidant and antiproliferative activities of red pitaya. Food
Wybraniec, S. and Mizrahi, Y. (2002). Fruit flesh betacyanin pigments in
Hylocereus cacti. J. Agric. Food Chem.50:6086–6089.
Wybraniec, S., Nowak-Wydra, B., Mitka, K., Kowalski, P. and Mizrahi, Y.
(2007). Minor betalains in fruits of Hylocereus species. Phytochemistry.
Zeashan, H., Amresh, G., Singh, S. and Rao, C. V. (2008). Hepatoprotective
activity of Amaranthus spinosus in experimental animals. Food Chem. Toxi-
Zeashan, H., Amresh, G., Singh, S. and Rao, C. V. (2009). Hepatoprotective
and antioxidant activity of Amaranthus spinosus against CCl
induced toxic-
ity. J. Ethnopharmacol.125:364–366.
Zhang, D., Lanier, S. M., Downing, J. A., Avent, J. L., Lumc, J. and McHalea,
J. L. (2008). Betalain pigments for dye-sensitized solar cells. J. Photochem.
Photobiol. Chem.195:72–80.
Zhou, H., Wu, L., Gao, Y. and Ma, T. (2011). Dye-sensitized solar cells using
20 natural dyes as sensitizers. J. Photochem. Photobiol. Chem.219:188–194.
Zou, D. M., Brewer, M., Garcia, F., Feugang, J. M., Wang, J., Zang, R., Liu, H.
and Zou, C. (2005). Cactus pear: A natural product in cancer chemopreven-
tion. Nutr. J.4:25.
... Los resultados elevados obtenidos de betalaínas en comparación con estudios previos, puede deberse a la suma de betalaínas presentes en epicarpio y mesocarpio de la muestra, lo que contribuyó a mayores contenidos de estos compuestos. El color del fruto de xoconostle le atribuye la presencia de estos pigmentos característicos (betalaínas), lo que lo convierte no solo en un alimento atractivo (Hernández-Fuentes et al., 2015), si no en una fuente de fitoquímicos altamente beneficioso al ser incorporado en la dieta, donde ingestas pequeñas de estos compuestos han reportado efectos quimiopreventivos y reducción de daños oxidativos, confiriendo efectos antioxidantes (Gandía-Herrero et al., 2016;Belhadj et al., 2017). ...
Full-text available
RESUMEN Los frutos de Opuntia spp. (xoconostle) son ampliamente consumidos en varias regiones de México, lo que denota su aceptabilidad gastronómica y accesibilidad, por sabor y bajos costos de adquisición. Sin embargo, su calidad nutrimental, y compuestos bioactivos puede cambiar de acuerdo al estado de madurez, condiciones agroclimáticas y variabilidad genética. genética. Es por ello, que el objetivo de esta investigación fué presentar los datos más recientes del xoconostle Ulapa, cultivado en la región de Ulapa de Melchor Ocampo, Hidalgo, México. Se realizó la evaluación de las propiedades fisicoquímicas como, color, sólidos solubles totales (SST), pH y acidez titulable (AT). Se evaluó el contenido de fenoles totales y flavonoides; adicionalmente se determinó betalaínas totales. Los resultados mostraron un alto contenido de betalaínas y flavonoides en comparación con otras especies previamente estudiadas en México. Las concentraciones de estos fitoquímicos presentes en el xoconostle, lo convierten en una fuente permanente y alternativa de compuestos bioactivos. ABSTRACT The fruits of Opuntia spp. (xoconostle) are widely consumed in several regions of Mexico, which denotes its gastronomic acceptability and accessibility, for taste and low acquisition costs. However, its nutritional quality and bioactive compounds can change according to the state of maturity, agroclimatic conditions, and genetic variability. Therefore, the aim of this research was to present the most recent information of the Ulapa xoconostle, cultivated at the region of Ulapa of Melchor Ocampo, Hidalgo, Mexico. The evaluation of the physicochemical properties such as color, total soluble solids (TSS), pH, and titratable acidity (TA) were carried out. The content of total phenols and flavonoids was evaluated; additionally, total betalains were determined. The results showed a high content of betalains and flavonoids compared to other species previously studied in Mexico. The concentrations of these phytochemicals present in the xoconostle make it a permanent and alternative source of bioactive compounds.
... Furthermore, families belonging to Portulacaceae produce betalains, known as nitrogen-containing plant pigments with limited occurrence in nature [23,24]. This subgroup is natural colorants in the food and cosmetic sectors, although studies have shown their neuroprotective [25], chemoprotective [26], and antimicrobial potential [24]. ...
Full-text available
Portulaca oleracea L., popularly known as purslane, is an herbaceous succulent plant classified as one of the most important invasive weeds in the world. Due to its high nutritional level and wide range of pharmacological effects, involving anti-inflammatory, antibacterial, antioxidant, and antiulcerogenic, purslane is one of the medicinal species listed by the World Health Organization. In addition, purslane produces several phytochemicals, including flavonoids, alkaloids, and terpenoids, which confer different pharmacological activities and make the plant highly attractive for use in the most diverse industries. It has high adaptability to extreme soil conditions, able to grow and spread in environments under drought stress, salinity, and poor nutrients; and has been presented as a potential model plant to study resistance to abiotic stresses. Among other purslane traits of interest to the agriculture sector, is worth to mention phytoremediation and allelopathy, thus being a sustainable alternative in organic agriculture. Here, we report a bibliometric analysis of purslane in vitro tissue culture and genetic modification/editing, and discuss opportunities and limitations to exploit the biotechnological potential of purslane as a source of valuable bio-molecules for many different industries.
... Chlorophyll, anthocyanins, carotenoids and betalains are the major classes of plant pigments being approved and regulated for industrial use (Scotter, 2011;Solymosi et al., 2015). Among them, betalains are plant derived, water soluble, nitrogen-consisting pigments found only in plants from the Caryophyllales order (Gengatharan et al., 2015;Gandía-Herrero et al., 2016). They can be further classi ed into two different groups, betacyanin (red-violet) and betaxanthins (yellow-orange). ...
Full-text available
Betalains are water soluble nitrogenous pigments produce by plants under the Caryophyllales order and has been favoured as a natural colourant in food and pharmaceutical industries due to its high stability towards pH and temperature over a wide range of food. There is a constant search for alternative source and technique for betalain production to meet the growing demand as conventional extraction method requires high quantity of plant material. Thus, this study sought to examine the potential of producing betalain through callus culture of a natural betalain bearing plant, Gomphrena globosa using different plant growth regulators (PGR) and to evaluate the effect of elicitation in enhancing betalain production. Callus induction from different explants showed that the percentage of callus induction (84.00-100.00%) from the leaf and hypocotyl explants was significantly higher than seeds (53.33%). A combination of 0.5 mg/L 2,4-dichlorophenoxyacetic acid (2,4-D) with 1.0 mg/L 6-benzylaminopurine (BAP) were found to be effective inducing pink callus in full strength MS medium. Elicitation with tyrosine was the most effective in enhancing the betacyanin content (red-violet pigments) followed by salicylic acid. The highest betacyanin content, 0.139 ± 0.035 mg/mg FW callus was obtained when 100 µM of tyrosine was supplied. Copper sulphate was found to be effective in increasing the callus size but not the betalain content. The callus size was about 13-fold bigger in MS medium supplemented with 25 µM copper sulphate compared to medium without elicitors. This is the first study reporting an optimised protocol in the production of pigmented callus containing betalain from G. globosa using a combination of PGRs consisting of 2,4-D and BAP. In addition, tyrosine can be used as a suitable elicitor to enhance betalain production which provides an alternative source of betalain for the commercial production of natural colorants.
Pitaya (pitahaya or dragon fruit) belongs to the genus Hylocereus or Selenicereus in the Cactaceae family, and is well-known all over the world, especially in tropical and subtropical regions, for its health-benefiting properties. Although the pitaya traits have been extensively studied, only few studies focus on the molecular mechanisms underlying the formation of fruit quality traits in pitaya. The breeding improvement of pitaya is a complex task due to various factors, including its generation cycle, polyploidy, high heterozygosity, and complex physiological structure. Furthermore, the absence of effective methods and resources exacerbates the challenges in enhancing the breeding of this fruit crop. In recent years, advancements in biotechnology and sequencing technologies have played a vital role in supporting and expediting conventional breeding techniques, as well as providing insights into the molecular mechanisms and evolutionary processes of pitaya. This review puts a spotlight on an overview of the latest developments in pitaya research, including genome, transcriptome, metabolome, and proteome sequencing, functional gene identification and regulatory network analysis, as well as the novel tools, platforms, and programs that have emerged in this field. It paves the way for studying the gene functions and molecular breeding with desirable fruit traits of pitaya, which will help accelerate pitaya breeding progress.
Full-text available
This work aimed to study the natural dye extracted from Indonesian wild plants (Rivina humilis L.) using different solvents. The natural dye was extracted using the maceration method. Three different solvents, namely, aquades, acetone, and ethanol 96%, were used to extract natural dye from Rivina humilis L fruit. The absorbance spectra of the extracted dye were recorded using Ultraviolet-Visible (UV-Vis) spectroscopy. The different spectra of betalain pigment revealed the dye extract’s dependence on the solvent. The functional groups of the extracted dye were analyzed using Fourier transform infrared (FTIR) spectroscopy. The adherence of carbonyl and hydroxyl groups from FTIR spectra indicated that this dye could anchor to a semiconducting material, e.g., TiO2, which was commonly used in dye-sensitized solar cells (DSSC). The electrochemical properties of the extracted pigments were studied through higher occupied molecular orbital (HOMO) and lower unoccupied molecular orbital (LUMO) energy levels. Based on the results, the best performance to construct DSSC was achieved by natural dye adsorption with aquades solvent.
Betalains are plant pigments synthesized in the cells of Caryophyllales (red beets, opuntia, etc.). They are involved in the inactivation of reactive oxygen species and free radicals. The paper summarizes the data on the physical-chemical and pharmacological properties of betalains. Betalains eliminate the consequences of oxidative stress, effectively correct metabolic disorders in diabetes mellitus and abdominal obesity, and reduce the risk of cardiovascular diseases. A betalain-enriched diet has a wide range of anticancer effects. Betalains protect brain dopaminergic neurons from oxidative damage and reduce the severity of neurodegenerative disorders in Alzheimer’s and Parkinson’s disease. However, betalains are not stable enough to resist degradation during processing and storage of plant raw materials. Therefore, developing non-damaging technologies for betalain-containing treatment is highly relevant.
Betalains are water-soluble, nitrogen-containing vacuolar pigment and can be divided into two subclasses: the yellow – orange betaxanthins and the red – violet betacyanin. These pigments can be found mainly in Latin America, but also in some parts of Asia, Africa, Australia and in the Mediterranean area. In this work an overview related with the status of research about betalains extracted from Opuntia spp and the enforces made to evaluate their positive incidence in the human body is provided. Several studies enhance their anticancer, anti-inflammatory and antioxidant properties. They also exhibit antimicrobial and antidiabetic effect. Taking into account these properties, betalains seem to be a promising natural alternative as a colorant to replace the synthetic ones in the food additive industry. In addition, the use of Opuntia spp fruits as possible colorant sources in the Food Industry, may contribute positively to the sustainable development in semi-arid regions.
Full-text available
Sporophores of Hygrocybe species (Agaricales) contain muscaflavin, a pigment recently isolated from Amanita muscaria. Spectroscopic and chemical evidence for its dihydroazepine structure 5a is presented, indicating a close biogenetic relationship to betalamic acid 6a. The chemical nature of the water soluble pigments of Hygrocybe is dis­ cussed
Full-text available
Dragon fruit dye has been prepared and used in the fabrication of DSSC as sensitizer. The properties of dragon fruit dye have been investigated by UV-Vis and FTIR technique. The absorption spectrum shows a peak value of 535 nm. Chemically dragon fruit dye shows present of intermolecular H-bond, conjugate C=O stretching and esters acetates C-O-C stretching vibration, which is due to the component of anthocyanin. On the other hand, the resistivity of TiO2 film on ITO glass before it is used for the fabrication of DSSC is also investigated. The TiO2 sheet resistivity increase from 1 layer = 22.1Ωcm to 2 layers = 369.6 Ωcm. Finally, the efficiency of assemble DSSC was evaluated and simulated using a custom made technique. The result shows fill factor, Pmax and efficiency during the present of halogen lamp are 0.30, 13 μW, 0.22%, respectively. We have successfully showed that the DSSC using dragon fruit as a dye sensitizer is useful for the preparation of environmental friendly and low-cost DSSC.
Full-text available
Copao (Eulychnia acida Phil.) is an endemic arborescent cactus restricted mainly to the semi-arid Coquimbo Region (29°54′28″ S, 71°15′15″ W), Chile. The area of distribution is from sea level to 1200 m.a.s.l. The edible fruit called rumpa is generally round, with green or pink peel and small scales on its surface, showing wide variability in size and weight. The aim of this work was to characterize the rumpa harvested in January 2009 and 2010 in three sectors of Coquimbo Region to determine chemical and nutritional composition in three fractions: pulp with seeds, juice, and peel. The research showed that this fruit is a good natural source of mainly soluble dietary fiber, which has a jellied texture and is present in the three fractions analyzed: 2% for juice, 3% for pulp with seeds, and approximately 5% for peel, making it potentially a good source of hydrocolloids for the food industry. The fruit is also a good source of vitamin C; around 55 mg 100 g-1 in peel, and 30 mg 100 g-1 in pulp with seeds and juice, values considered high compared to 18 mg 100 g-1 for prickly pear (Opuntia ficus-indica [L.] Mill.) The main minerals were: K, Mg, Ca, and P. Total polyphenols and betalain pigments were also determined in the pulp with seeds and pink peel fractions, respectively. The nutritional characteristics, together with its high water content of around 96%, make rumpa a promising raw material for agro-industrial development of natural juices or isotonic drinks. This characterization helps in the recovery of an endemic native species by reducing potential threats to destroy wild populations of E. acid, especially near agricultural areas, and by promoting habitat conservation of the species in the region.
Due to population growth and subsequent growing energy demand, clean energy generation is one of the major challenges today and in the future. Dye-sensitized solar cells (DSSCs) is one of the new approaches to producing clean, endless energy. In this review, the primary function, structure, performance parameters, and key elements of DSSCs are analyzed first, with a focus on the photosensitizer as the crucial part due to electron production. Photosensitizers based on natural resources recently attracted attention because of their economic and environmental advantages, but there are substantial challenges such as low conversion efficiency and long term stability. Also, diverse and, in some cases, contradictory results presented in the literature make conclusions difficult. Here, different challenges about DSSCs based on natural dyes are categorized and discussed according to the published papers, and possible solutions are presented, which could shed light on future investigations. Furthermore, according to different factors, such as average solar radiation and economic/technological aspects, DSSCs based on natural dyes are proposed to be the most promising renewable energy technology for arid to semi-arid, sunny countries like Iran in the future. Accordingly, the performance of DSSCs based on natural dyes manufactured using dye extract of different plants grow in Iranian is summarized then. In recent years, many DSSCs based on natural dyes have been manufactured using different plants grown in different parts of the country. Interestingly, it is possible to extract many natural dyes for DSSCs based on natural dyes from agricultural wastes such as saffron petals and walnut shells, of which thousand tons them are disposed every year.
A method for the screening of antioxidant activity is reported as a decolorization assay applicable to both lipophilic and hydrophilic antioxidants, including flavonoids, hydroxycinnamates, carotenoids, and plasma antioxidants. The pre-formed radical monocation of 2,2'-azinobis-(3-ethylbenzothiazoline-6-sulfonic acid) (ABTS*+) is generated by oxidation of ABTS with potassium persulfate and is reduced in the presence of such hydrogen-donating antioxidants. The influences of both the concentration of antioxidant and duration of reaction on the inhibition of the radical cation absorption are taken into account when determining the antioxidant activity. This assay clearly improves the original TEAC assay (the ferryl myoglobin/ABTS assay) for the determination of antioxidant activity in a number of ways. First, the chemistry involves the direct generation of the ABTS radical monocation with no involvement of an intermediary radical. Second, it is a decolorization assay; thus the radical cation is pre-formed prior to addition of antioxidant test systems, rather than the generation of the radical taking place continually in the presence of the antioxidant. Hence the results obtained with the improved system may not always be directly comparable with those obtained using the original TEAC assay. Third, it is applicable to both aqueous and lipophilic systems.
Betalains (betacyanins and betaxanthins), the main pigments of red beet (Beta vulgaris) roots, showed an antiradical effect when measured by the destruction of the 2,2'-azino-bis(3-ethylbenzthiazoline-6-sulphonic acid (ABTS) free radical generated by the horse-radish peroxidase/hydrogen peroxide-mediated oxidation of ABTS. The antiradical activity of betacyanins was greater than that of the betaxanthins and increased with the pH of the reaction medium. The different antiradical properties shown by both types of betalain is discussed in light of the respective ease with which it is possible to withdraw one electron from their molecules and the stability of their corresponding radicals.