Food Sci. Technol. Res.

, 18 (3), 315 – 324, 2012

Review

Anthocyanins as Functional Food Factors

— Chemistry, Nutrition and Health Promotion —

Takanori Tsuda*

College of Bioscience and Biotechnology, Chubu University, Kasugai, Aichi 487-8501, Japan

Received January 25, 2012; Accepted February 5, 2012

Anthocyanins are widely known as pigments responsible for the rich reds and purples in fruits, veg-
etables and legume seeds. Many flower colors are also attributed to anthocyanins. The physiological func-
tions of anthocyanins as components in food have attracted attention, and research on their bioavailabil-
ity and physiological functionality has progressed over the last 20 years. This review focuses on the health
benefits of anthocyanins. It first describes the chemistry, metabolism, and absorption of anthocyanins and
next summarizes the trends in research on the anti-diabetic and anti-obesity effects of anthocyanins. It
then describes the other health benefits of anthocyanins, namely, the prevention of cancer and improve-
ment of visual and brain functions, and finally discusses the challenges in and prospects of anthocyanin
research.

Keywords: anthocyanin, bioavailability, diabetes, obesity, cancer, vision, brain function

Abbreviations: AMPK, AMP-activated protein kinase; BA, blackcurrant anthocyanins; BBE, bilberry extract;
CS, Caffeoylsophorose; Cy, cyanidin; C3G, cyanidin 3-glucoside; Del, delphinidin; Glut4, glucose transporter 4;
NF-κB, nuclear factor-kappa B; PPAR, peroxisome proliferator activated receptor; RBP4, retinol-binding protein 4;
TNF-α, tumor necrosis factor-alpha; WBP, whole blueberry powder

Introduction and it became possible after the discovery of genes involved

Anthocyanins are widely known as pigments respon-
sible for the rich reds and purples in fruits such as apples,
strawberries, and blueberries, as well as in eggplants, perilla
herbs, and legume seeds. Many flower colors are also at-
tributed to anthocyanins (Harborne and Grayer, 1988). In
food chemistry, anthocyanins have been studied in relation
to changes and stability of colors in foods such as fruits dur-
ing processing and storage and also for their use as natural
colorants. Indeed, many types of anthocyanin food colorants
have been developed and are now available to customize
the appearance of foods. In horticulture, color conversion of
flower pigments has become possible based on new findings
of anthocyanin research. Creation of flowers in new colors
enriches our life; for example, the creation of blue roses is a
noteworthy achievement (Tanaka et al., 2010). Genetic en-
gineering is the key technology for converting flower color,
*To whom correspondence should be addressed.
E-mail: tsudat@isc.chubu.ac.jp
in anthocyanin biosynthesis and elucidation of their expres-
sion mechanisms. In addition to the above properties and
mechanisms, the physiological functions of anthocyanins as
components in food, have attracted attention, and research
on their bioavailability and physiological functionality has
progressed over the last 20 years.
This review focuses on the health benefits of antho-
cyanins. It first describes the chemistry, metabolism, and
absorption of anthocyanins and next summarizes the trends
in research on the anti-diabetic and anti-obesity effects of
anthocyanins. It then describes the other health benefits of
anthocyanins, namely, the prevention of cancer and improve-
ment of visual and brain functions, and finally discusses the
challenges in and prospects of anthocyanin research.
Chemistry, Source, Intake, Metabolism, and Absorption
of Anthocyanins
Anthocyanins are flavonoids that usually exist in plants in
the form of glycosides, and their non-carbohydrate moieties
316
(aglycones) are called anthocyanidins. There are many types
of anthocyanins, which are distinguished according to the
substituent groups on the B ring, type and number of conju-
gated sugars, and the presence or absence of an acyl group.
Six principal types of anthocyanidins (pelargonidin, cyanidin
(Cy), delphinidin (Del), peonidin, petunidin, and malvidin)
are shown in Fig. 1. The hue of an anthocyanin is determined
by the substituent groups on the B-ring; the hue deepens as
the number of hydroxyl groups increases and lightens with
the presence of methoxyl groups. Figure 2 shows the struc-
tural changes of anthocyanins in aqueous solution — they
exist in the form of stable red flavylium cations under acidic
conditions and are converted into unstable colorless pseudo-
bases upon reaction with water under weakly acidic and neu-
tral conditions (Brouillard, 1988).
Fig. 1. Chemical structures of the anthocyanidins.
Fig. 2. Structural changes of glucosyl anthocyanins in aqueous solution (G, glucose).
T. Tsuda
Anthocyanin content varies markedly among plant spe-
cies and cultivars and also within a single species, depending
on harvest time. Anthocyanins are contained in many types
of regularly consumed foods, such as cereals, tubers, roots,
greens, pulses, and fruits. Cy and its derivatives, which pos-
sess two hydroxyl groups on the B-ring, are the most widely
distributed, followed by Del and its derivatives. Although
similar data for foods in Japan are unavailable, the typi-
cal concentrations of anthocyanins in foods commercially
available in the United States have been reported (Wu et al.,
2006). Berries are particularly rich in anthocyanins (Table
1), and daily anthocyanin intake in the United States is esti-
mated at 12.5 mg/person. This figure is thought to be higher
than the corresponding figure in Japan, albeit with significant
differences among individuals.
Recently, results were reported in detail for the European
Prospective Investigation into Cancer and Nutrition study,
wherein the intake of anthocyanidins in 36,037 individuals
(age range, 35 − 74 years) living across 10 European coun-
tries was investigated (Zamora-Ros et al., 2001). According
to this report, the mean daily anthocyanidin intake in men
ranged from 19.8 − 64.9 mg, while that in women ranged
from 18.7 − 44.1 mg. There was a clear north-to-south gradi-
Anthocyanins and Health Benefits 317

protocatechuic acid, syringic acid, vanillic acid, phloroglu-

Table 1. Concentration of anthocyanins in common foods.
cinol aldehyde, phloroglucinol acid, and gallic acid, have
Total anthocyanins been reported in multiple studies as anthocyanin metabolites,
Total anthocyanins
Food (mg/100 g of fresh weight
or form consumed)
per serving (mg) depending on their structure (Gonthier et al., 2003; Aura et
al., 2005; He et al., 2005; Keppler and Humpf, 2005; Borges
1. Apple
et al., 2007; Forester and Waterhouse, 2008, 2010; Avila et
(Fuji) 1.3 1.8
(Gala) 2.3 3.2 al., 2009). These phenolic acids are likely to be produced
(Red Delicious) 12.3 17.0 during anthocyanin metabolism involving enteric bacteria,
2. Blackberry 245.0 353.0 or by chemical conversion, and have also been found in
3. Blueberry humans (Kahle et al., 2006; Vitaglione et al., 2007; Azzini
(Cultivated) 386.6 529.0
et al., 2010). The actions of these metabolites must be taken
(Wild) 486.5 705.0
4. Cherry, sweet 122.0 177.0 into consideration when discussing the health benefits of
5. Chokeberry 1,480.0 2,147.0 anthocyanins. However, whether these metabolites are solely
6. Cranberry 140.0 133.0 responsible for the health benefits of anthocyanins remains a
7. Currant contentious topic. Recently, Lila et al. (2012) aimed to elu-
(Black currant) 476.0 533.0
cidate the metabolic fate of anthocyanins by employing two
(Red currant) 12.8 14.3
8. Elderberry 1,375.0 1,993.0 approaches. The first approach used a human gastrointestinal
9. Grape model for investigating the bioaccessibility of anthocyanins,
(Red grape) 26.7 42.7 which is defined as the potential for a substance to interact
(Concord) 120.1 192.0 with or be absorbed by an organism. The second approach
10. Nectarine 6.8 9.2
employed radio-labeled (13C or 14C) anthocyanins synthe-
11. Peach 4.8 4.7
12. Plum 19.0 12.5 sized by plant cells (Lila et al., 2012). Both approaches are
13. Strawberry 21.2 35.0 crucial for revealing the metabolic pathway and absorption
14. Black bean 44.5 23.1 of anthocyanins in detail. To date, however, only a limited
15. Eggplant 85.7 35.1 number of studies have reported anthocyanins as essential for
16. Red cabbage 322.0 113.0
human health benefits. A clinical ophthalmology study found
17. Red onion 48.5 38.8
possible visual improvement in response to a daily intake of
This table is reproduced from Wu et al. (2006) with permission 50 mg anthocyanins (Nakaishi et al., 2000).
from the American Chemical Society.

Health Benefits of Anthocyanins

ent, and the intake was highest in southern countries. Antho-
cyanidin intake was also high in non-obese older women and
non-smokers and increased in line with educational level and
degree of physical activity. The main sources of anthocyani-
dins were fruits, vegetables, wine, and non-alcoholic bever-
ages. Although fruits like berries are rich in anthocyanins,
vegetables are the likely source for the Japanese population
as they are often included daily in every meal. Considering
the relatively low anthocyanin content in vegetables, creating
new anthocyanin-rich varieties and cultivars may contribute
to stable intake of anthocyanins. Juices rich in anthocyanins
may also serve as a good source.
It is known that a large proportion of anthocyanin gly-
cosides are absorbed directly into the body without losing
their sugar moiety (Tsuda et al., 1999). On the other hand,
protocatechuic acid, a derivative of the B-ring of cyanidin
3-glucoside (C3G), was also found as an anthocyanin me-
tabolite (Tsuda et al., 1999). In recent years, biodegradation
of anthocyanins by enteric bacteria and the resulting pheno-
lic acids has attracted much interest. Phenolic acids, such as
Since their antioxidant effect was revealed, various health
benefits of anthocyanins have been reported. In particular,
marked advancements in research over the last 10 years have
been made regarding the conferred health benefits of antho-
cyanins and the underlying molecular mechanisms. Recent
findings in relation to the preventive and suppressive effects
of anthocyanins against obesity and diabetes are outlined in
the next section.
Obesity and diabetes Suppression of body fat accumu-
lation by anthocyanin intake was first reported by Tsuda et al.
(2003). Supplementing high-fat meals (60% of energy) with
2 g/kg of C3G significantly reduced high-fat meal-induced
body fat accumulation in C57BL/6J mice (Tsuda et al., 2003;
Tsuda, 2008), and impaired lipid synthesis in white adipose
tissue was considered a likely underlying mechanism. It was
also shown that a diet supplemented with C3G significantly
suppresses the elevation of the serum glucose levels induced
by high-fat meals. Prior et al. (2008) reported that supple-
menting a high-fat diet (45% of energy) with anthocyanins
extracted from blueberries inhibited weight gain and body fat
318
accumulation, while supplementation with whole blueberry
powder (WBP) promoted body fat accumulation in C57BL/6
mice. A separate study by the same group also demonstrated
that consumption of blueberry juice caused a significant re-
duction in body weight gain and percentage of white adipose
tissue (epididymal and retroperitoneal fat) in mice fed a high-
fat diet (45% of energy) (Prior et al., 2010a). Similarly, De-
Furia et al. (2009) reported that supplementing a high-fat diet
(60% of energy) with WBP did not reduce body weight gain
significantly in C57BL/6 mice. Contrary to these findings,
interesting results were recently reported by Seymour et al.
(2011), who demonstrated that supplementing a high-fat diet
(45% of energy) with 2% WBP reduced abdominal fat mass
and increased the activity of the adipose tissue and skeletal
muscle peroxisome proliferator-activated receptor (PPAR) in
Zucker fatty rats. However, the same study showed a WBP-
induced body weight gain in Zucker lean rats (Seymour et
al., 2011).
Anthocyanins from berries other than blueberries have
also been tested for their anti-obesity and anti-diabetic ef-
fects. Intake of black raspberry anthocyanins (as a juice or in
powder form) did not significantly reduce body fat accumu-
lation or body weight gain induced by a high-fat diet (60%
of energy) in mice (Prior et al., 2009, 2010b; Kaume et al.,
2012). On the other hand, intake of mulberry water extracts
containing a high concentration of anthocyanins reduced
body weight gain (Peng et al., 2011). Intake of tart cherry
powder also reduced body weight gain and the amount
of retroperitoneal fat in Zucker fatty rats (Seymour et al.,
2009). Intake of extracts of another anthocyanin-rich berry,
chokeberry, also suppressed the increase in epididymal white
adipose tissue and blood glucose level in mice fed a fructose-
rich diet (Qin and Anderson, 2012).
Anthocyanins act on adipocytes, thereby modulating
adipocytokine expression. C3G, or its aglycon Cy, was re-
ported to upregulate the expression of adiponectin, which
increases insulin sensitivity in rat white adipose tissue and in
human adipocytes (Tsuda et al., 2004, 2006). However, these
findings have not yet been supported by in vivo results. The
study by DeFuria et al. (2009), introduced above, demon-
strated the suppressive effect on obesity-associated inflam-
mation in white adipose tissue in the group that received
WBP. More specifically, the degree of upregulation of tumor
necrosis factor-alpha (TNF-α) and monocyte chemoattractant
protein-1 mRNAs was significantly lower in the group fed
a high-fat diet with blueberry supplementation than in the
group without supplementation. It was also shown that a sig-
nificant downregulation of glutathione peroxide 3 level, in-
duced by the high-fat diet, was restored by blueberry intake.
Furthermore, intake of tart cherry powder suppressed the
T. Tsuda
upregulation of proinflammatory cytokines (IL-6 and TNF-α)
associated with obesity, while upregulating PPARα and
PPARγ mRNAs in Zucker fatty rats (Seymour et al., 2009).
Similarly, chokeberry intake increased plasma adiponectin
levels and decreased plasma TNF-α and IL-6 levels (Qin and
Anderson, 2012).
Anthocyanin intake was also shown to reduce increases
in blood glucose level and to improve insulin sensitivity in
type 2 diabetes models. Similar effects were found in stud-
ies using high purity C3G (Sasaki et al., 2007; Tsuda, 2008)
and also bilberry extract (BBE) containing different types
of anthocyanins (Takikawa et al., 2010). Downregulation of
retinol binding protein 4 (RBP4) was shown to constitute a
part of the mechanism underlying such effects of isolated
C3G in KK-Ay mice, a model for type 2 diabetes (Sasaki
et al., 2007). Yang et al. (2005) identified RBP4 as a new
adipocytokine linked to the pathogenesis of type 2 diabetes
and a factor contributing to insulin resistance (Yang et al.,
2005). Although the role of RBP4 in diabetes in humans
remains debatable, the anti-diabetic effect of C3G in mice
can be explained by this finding. More specifically, C3G
intake upregulates the expression of glucose transporter 4
(Glut4), thereby decreasing RBP4 expression. This improves
insulin sensitivity in peripheral tissues and suppresses glu-
cose release induced by elevated gluconeogenesis (Sasaki et
al., 2007). Further, C3G and its metabolite protocatechuic
acid were recently shown to activate PPARγ and to induce
upregulation of Glut4 and adiponectin in murine 3T3-L1
adipocytes and human adipocytes (Scazzocchio et al., 2011).
However, C3G does not appear to serve as a ligand of PPARγ
or to upregulate the expression of adiponectin in vivo (Sasaki
et al., 2007). Thus, the anti-diabetic effect of C3G cannot be
explained by the upregulated expression of PPARγ and adi-
ponectin.
Considering the low C3G content in BBE, it is difficult to
explain its anti-diabetic effect by the mechanism involving
RBP4 downregulation. BBE intake was shown to activate
AMP-activated protein kinase (AMPK) in skeletal muscle
and white adipose tissue and the liver (Takikawa et al.,
2010). BBE-induced AMPK activation leads to upregulation
of Glut4 expression in skeletal muscle and white adipose tis-
sue, as well as suppression of gluconeogenesis in the liver.
At the same time, it also promotes lipid utilization, thereby
suppressing increases in blood glucose level and improving
insulin sensitivity (Fig. 3) (Takikawa et al., 2010). The fact
that BBE contains many types of anthocyanins may be the
key in the AMPK-medicated anti-diabetic effect, and this
point needs to be addressed in future studies.
Another possible mechanism for anthocyanins’ preven-
tive and suppressive effect on diabetes is the inhibition of
Anthocyanins and Health Benefits
Fig. 3. Proposed mechanism for amelioration of hyperglycemia and insulin sensitivity by dietary bilberry extract,
BBE (Takikawa et al., 2010). ACC, acetylCoA carboxylase; ACO, acylCoA oxidase; CPT1A, carnitine palmitoyl-
transferase-1A; G6Pase, glucose-6-phosphatase; PEPCK, phosphoenol pyruvate carboxykinase.
α-glucosidase in the small intestinal endothelium, which in
turn inhibits degradation and absorption of carbohydrates,
thereby ameliorating hyperglycemia. Glycosides, such as
C3G, are generally very weak inhibitors of α-glucosidase
(Iwai et al., 2006), except for acylated anthocyanins (Matsui
et al., 2001a, 2001b, 2002; Terahara et al., 2003, 2009) and
Matsui et al. (2004) revealed that caffeoylsophorose (CS), a
component of acylated anthocyanins, is the molecular struc-
ture responsible for this strong inhibitory activity. Since CS
is absorbed in an intact form to produce caffeic acid and its
conjugates (Qiu et al., 2011), it is possible that absorbed CS
or its metabolites exert an anti-diabetic effect via pathways
that do not include α-glucosidase inhibition.
Other Health Benefits of Anthocyanins: Anthocyanins in
Relation to Cancer, Visual Function, and Brain Function
Cancer The preventive and suppressive effects of
anthocyanins on cancer have been demonstrated in many
studies, and key findings in this area can be found in the
literature (Thomasset et al., 2009; Prasad et al., 2010). In
general, anthocyanins suppress inflammatory pathways as-
sociated with cancer pathogenesis. For example, Del, an
aglycon, binds directly to mitogen-activated protein kinase
4 and phosphatidylinositol 3-kinase to inhibit both enzymes,
thereby suppressing ultraviolet B-induced cyclooxygenase-2
319
expression (Kwon et al., 2009). However, Del does not exist
in the form of the flavylium cation and is unstable under neu-
tral conditions; thus, whether it can exert an action on these
target molecules in vivo remains unclear.
Recent attempts to create purple tomatoes by expressing
high levels of anthocyanins (Butelli et al., 2008; Gonzali et
al., 2009; Povero et al., 2011) are of particular interest. Mar-
tin and colleagues expressed snapdragon genes involved in
anthocyanin biosynthesis in tomato and successfully obtained
anthocyanin-rich tomatoes with intense purple coloration
(Butelli et al., 2008). When cancer-susceptible mice (Trp53-
/-
mice) were fed a diet supplemented with purple tomatoes
or normal tomatoes, the life span extension was significantly
longer in the mice that consumed purple tomatoes than in
those that consumed normal tomatoes (Butelli et al., 2008).
A similar approach may be useful for other health promoting
purposes, as well as for cancer prevention. More precisely,
establishing and cultivating vegetables that contain appropri-
ate anthocyanin compositions and content, sufficient for ex-
erting health benefits, will be useful for health improvement.
Visual function Improvement of visual function as well
as antioxidant effects is a well-known health benefit of antho-
cyanins. A famous anecdote about the effect of anthocyanins
on vision in Japan describes Royal Air Force pilots during
World War II who experienced enhanced vision in the dark
320
after ingesting blueberry jam. However, this anecdote itself
is not evidence-based, and several clinical studies failed to
verify the above claim (Levy and Glovinsky, 1998; Zadok et
al., 1999; Muth et al., 2000; Canter and Ernst, 2004). On the
other hand, bilberry anthocyanins have been shown to ex-
hibit neuroprotective effects in ocular tissue and suppressive
effects on diabetic retinopathy via a reduction of angiogen-
esis (Matsunaga et al., 2010a, 2010b). Furthermore, several
studies have demonstrated that blackcurrant anthocyanins
(BA) improve visual function; for example, BA inhibit tran-
sient myopia, reduce eye fatigue, improve dark adaptation,
and increase retinal blood flow in glaucoma (Nakaishi et al.,
2000; Ohguro et al., 2007; Iida et al., 2010). The first three
benefits have been shown in humans, and effective daily BA
intake was reported as 50 mg (Nakaishi et al., 2000). Two
studies have indicated the possible mechanism underlying
inhibition of myopia. The BA concentration was higher in
ocular tissues (sclera, choroid, ciliary body, retina, iris, and
cornea) than in blood in a rat fed a diet containing BA, and
in particular, concentrations in the sclera and choroid were
100-fold higher than the blood BA concentration (Matsu-
moto et al., 2006). In addition, it was shown that an antho-
cyanin in the concentration range of 10-7 to 10-8 M relaxed
endothelin-1-induced contraction of bovine ciliary smooth
muscle (Matsumoto et al., 2005), which plays an important
role in modulating refraction of the lens through contraction
and relaxation, which in turn control accommodation. Thus,
it is suggested that anthocyanins stimulate the endothelin-1
receptor to induce production of NO, thereby relaxing ciliary
smooth muscle, which in turn flattens the lens and conse-
quently inhibits myopia.
Brain function The favorable effects of intake of an-
thocyanin-rich fruits on memory (Shukitt-Hale et al., 2006;
Krikorian et al., 2010a, 2010b) and on cognitive and mo-
tor function by delaying deterioration of neural function in
aged individuals (Shukitt-Hale et al., 2009a, 2009b) were
studied mainly by the group lead by Joseph. Neural signal-
ing and suppression of neuroinflammation are implicated
in the mechanism for such effects. In Fisher rats, blueberry
intake suppressed an aging-associated increase in nuclear
factor-kappa B levels (NF-κB) (Goyarzu et al., 2004) and
upregulation of TNF-α, IL-1β and NF-κB expression in the
hippocampus (Shukitt-Hale et al., 2008).
Williams et al. (2008) demonstrated that blueberry intake
induced activation of cyclic AMP-response element-bonding
protein and an increase in the level of brain-derived neuro-
trophic factor, in which extracellular signal-related kinases 1
and 2 were involved. Improvement of cerebrovascular blood
flow may also be implicated in the favorable effects of ber-
ries on brain function (Spencer, 2010).
T. Tsuda
Challenges in and Prospect for Anthocyanin Research
This review so far described the health benefits of antho-
cyanins, with an emphasis on their preventive and suppres-
sive effects on obesity and diabetes, as well as their metabo-
lism and absorption, and then outlined the trends in cancer
prevention research and the improvement of visual and brain
function by anthocyanins. Next, challenges in anthocyanin
research are discussed to conclude this review.
Although evidence of the pleiotropic properties of an-
thocyanins, such as the suppressive effect on body fat ac-
cumulation and anti-diabetic action, continue to accumulate,
only a limited number of studies have identified the specific
molecular structures of anthocyanins responsible for their
health benefits. Interestingly, in some cases, intake of a blend
of multiple types of anthocyanins can be more effective than
intake of a single molecular species, suggesting the necessity
of clarifying the most effective molecular species and blend
compositions of anthocyanins for exerting individual health
benefits. For example, black raspberry anthocyanins do
not appear effective for suppressing body fat accumulation
(Prior et al., 2009, 2010b; Kaume et al., 2012), and this lack
of effect may be attributed to the compositional difference
between black raspberry and blueberry anthocyanins, as the
main anthocyanin in black raspberry is cyanidin 3-rutinoside.
If so, the sugar moieties of anthocyanins may influence the
degree of function.
Investigations in the current literature used mainly
anthocyanin-rich extracts, with a few exceptions of isolated
and purified anthocyanins. In fact, anthocyanins, as a part of
a diet, are more likely to be ingested with other dietary com-
ponents such as polyphenols. Nevertheless, it is important
to determine whether an anthocyanin (or anthocyanins) is
solely responsible for exerting health benefits, whether other
dietary components are the main mediators of the benefits, or
whether co-intake of an anthocyanin (or anthocyanins) and
other dietary components is necessary.
Current evidence to support the health benefits of antho-
cyanins in humans is insufficient. Although several studies
showed the favorable effects of anthocyanins on visual func-
tion, they used a small number of subjects, and are thus not
considered thorough. Further investigation is necessary to
address the benefits of anthocyanins in humans.
Understanding anthocyanin metabolites on the basis of
chemical structures of anthocyanins is of particular impor-
tance. The quantity of anthocyanin metabolites produced
from processing by enteric bacteria or chemical reactions
must be taken into account when considering the efficacy
of anthocyanins. On the other hand, the effect of the antho-
cyanins directly absorbed in the form of glycosides, albeit
with their low bioavailability, should not be neglected. Even
Anthocyanins and Health Benefits
though the quantities of absorbed anthocyanins, in the form
of glycosides, or their metabolite are small, it is possible that
the signals triggered by them are amplified in the digestive
tract or on the cell surface, thereby exerting diverse func-
tions. Advancements in research in this field may success-
fully depict mechanisms for health benefits of anthocyanins,
despite their low absorption rates in the body. Nevertheless,
the correlations between beneficial levels of anthocyanin
intake and concentrations of their metabolites have not yet
been clarified. If enteric bacteria-generated metabolites of
anthocyanins are important for promoting health benefits,
differences in the quality and quantity of gut flora will influ-
ence the levels of expected health benefits among individu-
als.
It was only a few decades ago that anthocyanins were
regarded as highly degradable compounds, and the main
research areas were their chemical structures, color stability,
use as food constituents, and changes in foods during stor-
age. Anthocyanins are now recognized as food constituents
with potential health benefits, and research related to these
properties has markedly progressed at the molecular level.
Anthocyanins will continue to attract researchers across
various disciplines, including those involved in the creation
of new flower varieties with novel colors. Research on the
health benefits of anthocyanins will provide information
on underlying molecular mechanisms and absorption and
metabolism. Moreover, once these benefits are proven in
humans, development of foods and dietary supplements in
capsule form can be accelerated to promote the proven func-
tions.
Acknowledgements This review was supported in part by Grants-
in-Aid for Scientific Research (Nos. 23580187 and 22248014) from
the Japan Society for Promotion of Science.
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