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166 Wild Edible Plants
TABLE 5.5 (Continued)
KAEM-
BERRY C3G C3S C3GAL C3A C3X RUTIN QUERCETIN
Сhokeberry
(A. melanocarpa)
1.69 n.a 125.6 142.43 46.90 n.a 57.50 n.a n.d n.a n.a n.d 261.2 n.a n.a Zheng &
8.61 n.a 239.7 91.66 12.43 91.7 1.7 n.a n.a n.a n.a 70.9 n.a n.a n.a Jakobek
9.63 n.a 371.3 160.9 12.37 n.a n.a n.a n.a n.a n.a n.a n.a n.a n.a Brand et al.,
PFEROL MYRICETIN
Elderberry,
S. nigra
Guelder rose
(V. opulus)
Barberry
(B. vulgaris)
C3G, cyanidin- 3- glucoside; C3S, cyanidin- 3- sambubioside; C3Gal, cyanidin 3- galactoside; C3A,
cyanidin 3- arabinoside; C3X, cyanidin 3- xyloside; n.d, not detected; n.a, not available.
39.06–
486.44
156.70–
334.81
132.17 57.91 0.61 n.d n.d n.a n.a n.a n.a n.a n.a n.a n.a n.a n.a Veberic et al.,
8.6– 13.2 n.a n.a n.a n.a 4−11 6.3– 11.1 n. a n.a n.a n.a 180– 243 n.a 7.1– 23.4 8 Ozrenk et al.,
n.a n.a n.a n.a n.a 8.1 1.1 n.a n.a n.a 13.2 75.2 21.8 21.8 n.a Gundogdu,
n.a n.a n.a n.a n.a 7.3 2.1 n.a n.a n.a 182 62.4 8.9 64.0 n.a Yang et al.,
208.43–
579.17
232.74–
392.86
n.a n.a n.a n.a 14.72–
45.04
n.a n.a n.a n.a 14.44–
43.11
10.38–
24.95
12.79–
74.66
5.4 ANTHOCYANIN PROFILES OF WILD
BERRIES
Anthocyanins are water- soluble avonoid pigments, one of the subclasses of phenolic
compounds, whose presence in berries determines their red, blue, and purple colors
(Kahkonen et al., 2003; Khoo et al., 2017). A variety of research conducted in vitro,
in animal models, in epidemiological and clinical studies in humans have shown
anthocyanin’s advantages for cardiovascular health, protecting the body from infectious
diseases, normalizing metabolism, and lowering the risk of cancer and neurodegenerative
diseases (Khoo et al., 2017; Wallace, 2011; Zaa et al., 2023).
By chemical structure, anthocyanins are glycosides containing the anthocyanidins
as an aglycone part, which are glycosidically bonded with mono- or oligosaccharides.
There areabout 17 known anthocyanidins, while the six most commonly distributed
ones included cyanidin, delphinidin, malvidin, pelargonidin, peonidin, and petunidin
(Table 5.6).
It is considered that the percentages of these six most prevailing anthocyanidins
found in fruits and vegetables are the following: cyanidin (50%), delphinidin (12%),
pelargonidin (12%), peonidin (12%), petunidin (7%), and malvidin (7%) (Giuliani

Antioxidant Compounds in Wild Edible Berries 167
ELLAGIC
ACID
n.a n.a n.a n.a n.a n.a n.a Caruso et al.,
n.a n.a n.a n.a n.a n.a n.a Caruso et al.,
TABLE 5.6 Chemical structure of the six most common anthocyanidins
GALLIC
ACID
CHLOROGENIC
ACID
CAFFEIC
ACID CATECHIN
PROANTHO-
CYANIDIN REFERENCES
Wang,
2003
et al.,
2012
2017
2019
2016
2015
2020
2013
2022
Aglycone R1 R2
Cyanidin – OH – H
Pelargonidin – H – H
Peonidin – OCH
3
– H
Delphinidin – OH – OH
Malvidin – OCH
Petunidin – OCH
3
3
– OCH
– OH
3
et al., 2016). The carbohydrate part are represented more often by galactose, glucose,
arabinose, xylose, and rhamnose bonded to anthocyanidins in the form of mono- , di- ,
or trisaccharides (Veberič et al., 2015). Cyanidin is the most abundant aglycone, and for
many berries it represents up to 100% from the total anthocyanin content. The amounts
of anthocyanidins found in the largest quantities in wild berries are shown in Table 5.7.
Berries belonging to the genus Vaccinium contain glycosides of delphinidin,
malvidin, peonidin, and petunidin. Malvidin glycosides were predominant in bog

TABLE 5.7 Percentage (%) of anthocyanins grouped according to their aglycone with respect to total anthocyanin content in wild berries
BERRY CYANIDIN PELARGONIDIN PEONIDIN DELPHINIDIN MALVIDIN PETUNIDIN REFERENCES
Blackthorn (P. spinosa) 41.59 n.a 58.41 n.d n.d n.d Määttä- Riihinen et al., 2004
Rowanberry (S. aucuparia) 100.0 n.d n.d n.d n.d n.d Koponen et al., 2007
Blackberry (Rubus sp.) 100.0 n.d n.d n.d n.d n.d Veberic et al., 2015
Raspberry (Rubus idaeus) 87.1 12.9 n.d n.d n.d n.d Veberic et al., 2015
Raspberry (Rubus idaeus) 97.71 2.29 n.d n.d n.d n.d Koponen et al., 2007
Cloudberry (R. chamaemorus) 100.0 n.d. n.d n.d n.d n.d Koponen et al., 2007
Strawberry (F. vesca) 42.8 57.2 n.d n.d n.d n.d Veberic et al., 2015
Strawberry (F. vesca) 50.0 50.0 n.d n.d n.d n.d Sondheimer & Karash, 1956
Elderberry (S. nigra) 100.0 n.d n.d n.d n.d n.d Veberic et al., 2015
Elderberry (S. nigra) 100.00 n.a n.d n.d n.d n.d Määttä- Riihinen et al., 2004
Bog bilberries (V. uliginosum) 7.0 n.d 4.3 28.1 40.4 18.9 Lätti et al., 2010
Bog whortleberry (V.uliginosum) 3.7 n.a n.a 30.8 44.0 21.5 Määttä- Riihinen et al., 2004
Bog whortleberry (V.uliginosum) 5.84 n.d 3.31 30.26 45.39 15.19 Koponen et al., 2007
Blueberry (Vaccinium myrtillus) 20.02 n.d 12.50 31.92 14.57 20.99 Bunea et al., 2011
Blueberry (V. myrtillus) 30.3 n.d 5.1 38.7 7.4 14.0 Kähkönen et al., 2003
Blueberry 13.62 n.d 7.59 29.00 31.78 18.00 Wu et al., 2006
Blueberry (V. myrtillus) 30.02 n.d 6.50 36.36 18.07 8.06 Jovančević et al., 2011
Blueberry (V. myrtillus) 23.85 n.d 3.64 45.70 8.02 18.79 Zorenc et al., 2016
Bog bilberries (V. myrtillus) 7.0 n.d 5.0 27.5 42.5 13.5 Andersen, 1987
Cranberry (V. oxycoccus) 46.93 n.d 46.48 1.35 3.75 1.50 Koponen et al., 2007
Cranberry (V. oxycoccus) 46.5 n.d 53.5 n.d n.d n.d Česonienė et al., 2009
Cranberry (V. oxycoccus) 21.75 n.d 78.25 n.d n.d n.d Narwojsz et al., 2019
Lingonberry (V. vitis- idaea) 99.23 n.d 0.77 n.d n.d n.d Koponen et al., 2007
Lingonberry (V. vitis- idaea) 100.0 n.d n.d n.d n.d n.d Kähkönen et al., 2003
Lingonberry (V. vitis- idaea) 98.7 n.a 1.3 n.d n.d n.d Määttä- Riihinen et al., 2004
Rose hip (R. rugosa) 100.0 n.d n.d n.d n.d n.d Koponen et al., 2007
Сhokeberry (A. melanocarpa) 100.0 n.d n.d n.d n.d n.d Zheng & Wang, 2003
Сhokeberry (A. melanocarpa) 100.0 n.d n.d n.d n.d n.d Koponen et al., 2007
newgenrtpdf
168 Wild Edible Plants
n.d, not detected; n.a, not available.

Antioxidant Compounds in Wild Edible Berries 169
bilberries (Vaccinium uliginosum), while peonidin glycosides were the most dominant
anthocyanin in cranberry (Vaccinium oxycoccus).
5.5 DIETARY RECOMMENDATIONS FOR
ANTHOCYANIN INTAKE
It is believed that anthocyanins are not nutrients necessary for regular consumption, and
their deciency in the diet does not cause visible disorders in the body. There is currently no strictly established daily intake of anthocyanins in the European Union and
United States. However, an anthocyanin intake of 50 mg per day is generally considered
a permissible quantity (Da Silva et al., 2019; Stabnikova et al., 2024; Wallace & Giusti,
2015). It was estimated derived from analyses of foods consumed by population of certain country that the daily intake of anthocyanin in the United States consisted of 12.5
mg/ day in 2001– 2002 (Wu et al., 2006) and 47 mg/ day (women, 53; men, 43) in Finland
in 2007 (Ovaskainen et al., 2008). Estimated amounts of different anthocyanidins in
daily diet, %, were as follows: cyanidin, 44.7; delphinidin, 20.7%; malvidin, 15.4%;
petunidin, 9.0%; peonidin, 6.9%, and pelargonidin, 3.3 (USA), and cyanidin, 48.9, and
delphinidin, 27.7 (Finland).
Nowadays, it is widely accepted that including anthocyanin- rich foods like fruits
and vegetables in the daily diet is benecial for maintaining health and well- being giving
protection against chronic diseases (Carlsen et al., 2010). According to a World Health
Organization (WHO) report, 3.9 million deaths in the world in 2017 were associated
with insufcient dietary intake of fruits and vegetables. To support overall health and
diminish the risk of non- communicable diseases, the WHO recommends daily consumption of fruits and vegetables more than 400 g as sources of vitamins, minerals,
dietary ber, and benecial non- nutrient substances, including plant sterols, avonoids,
and other antioxidants (WHO, 2023). Among the sources of anthocyanins in the diet of
US adults aged ≥20 y, berries occupied rst place, accounting for 20% of total anthocyanin consumption, followed by wine (16%), grapes (11%), red/ purple vegetables
(8%), 100% non- citrus juice (6%), yogurt (6%), and other food sources (33%) (Wallace
& Giusti, 2015).
Anthocyanins, when introduced into the human body, are quickly absorbed and
are found in the bloodstream in a few minutes after consumption. Currently, the scientic literature describing the results of studies in humans does not contain information about the toxicity of anthocyanins from food intake (Wallace & Giusti, 2015).
There is an established, by the Joint FAO/ WHO Expert Committee on Food Additives
(JECFA), daily dose of 2.5 mg/ kg of body weight per day for anthocyanins from grapeskin extract, food additive E 163, which is authorized as food additives in the European
Union. However, there are no JECFA specications for other anthocyanins (EFSA,
2013). It was indicated that because toxicological properties of anthocyanins were
studied using fruit extracts containing several anthocyanins, conclusions could be done
only for anthocyanins in general.

170 Wild Edible Plants
5.6 WILD BERRIES IN FUNCTIONAL
FOOD PREPARATION
A signicant amount of research in the elds of food science and technology has been
devoted to the use of plant additives to increase the nutritional value of traditional food
products (Stabnikova et al., 2021, 2024). Berries containing valuable compounds with
antioxidant properties and antimicrobial activity has recently become important in the
food industry and can be widely used for manufacturing of functional products.
Wild berries traditionally have been used for production of jams, jellies, purées,
marmalades, as lling for pies, and could serve as an ingredient for drinks, including
alcoholic ones. In recent decades, anthocyanidins have attracted much attention as a
possible substitute for synthetic colorants in food products, and anthocyanins extracted
from edible berries (grape and grape skin, blackcurrant, and elderberry) are authorized
as food coloring substances (E 163) in the European Union (EFSA, 2013). So, elderberry puree or juice as natural colorants to replace synthetic ones could nd application
in manufacturing of dairy, bakery, and meat products (Domínguez et al., 2021; Haseeb
et al., 2018; Stabnikova et al., 2024; Szalóki- Dorkó et al., 2015). Adding berry juice as a
dye simultaneously increased antioxidant capacity of food products, thereby increasing
their nutritional value and allowing them to be stored longer due to slow down oxidative
processes.
It was proposed to use elderberry (Sambucus nigra L.) juice or puree as colorants
in production of yogurt (Cais- Sokolińska and Walkowiak- Tomczak, 2021; NajgebauerLejko et al., 2021), ker (Du and Myracle, 2018); in preparation of croissant (Da Silva
et al., 2019), gluten- free wafer sheets (Różyło et al., 2019), ber- enriched pasta (SunWaterhouse et al., 2013). Adding elderberry in the form of dry powder or elderberry
vinegar in manufacturing of meat products was effective to reduce oxidative processes
in them (Cordeiro et al., 2020; Jin et al., 2021).
Because anthocyanins present in berries may act as prebiotics, new functional products containing probiotic bacteria added with berries were proposed. It
was shown that addition of pulp from wild rowan, 5%, to the probiotic ice- cream
increased survival of lactic acid bacteria Lacticaesibacillus rhamnosus, increased icecream antioxidant capacity, meanwhile enriched ice- cream was acceptable in terms
of consumer taste (Goktas, 2023). Fruit puree from wild elderberry (Sambucus nigra)
or wild blackthorn (Prunus spinosa), added in quantity of 10% to probiotic yogurts
served a natural colorants and avorings, increased the antioxidant capacity of the
product, and did not affect the viability of probiotic strains Lactobacillus acidophilus
and Bidobacterium animalis ssp. lactis during 4 weeks of cold storage (NajgebauerLejko et al., 2021).
Ice- cream containing 15% of blackthorn puree (Prunus spinosa) rated higher than
the control in terms of color and appearance, and overall acceptability (Ürkek et al.,
2019). Generally, ice- cream is considered to be a vehicle for incorporating healthpromoting ingredients including fruits and fruit- based products such as puree or juice to
increase phenolic content of the product (Soukoulis et al., 2014). Addition (5.33% w/ w)

Antioxidant Compounds in Wild Edible Berries 171
of wild blueberry puree or juice concentrate to soy- milk- based ice- cream increased
its total anthocyanins content and antioxidant capacity, meanwhile overall consumer
acceptability was not affecting (Camire et al., 2006).
5.7 ANTHOCYANINS FROM BERRIES FOR
HUMAN HEALTH IMPROVEMENT
Consumption of fruits and vegetables is recommended for the prevention of cardiovascular disease (Hartley et al., 2013) and weight gain (WHO, 2023), reduction of inam mation (Joseph et al., 2014), the risk of obesity (Stabnikova & Paredes- López, 2024),
diabetes, neurodegenerative diseases (Zhong et al., 2023), and certain types of cancer
(Pem & Jeewon, 2015). It has been shown that the combination of fruits may have an
additive or synergistic effect on antioxidant efcacy and status in humans (Zafra- Stone
et al., 2007).
Anthocyanin- rich berries have traditionally been used in Europe and Asia for
treatment of different diseases. For example, bird cherries found their application in
folk medicine to treat cough, fever, improve eyesight, diseases of the stomach and gastritis (Pasko et al., 2012); rowan berries and tea, syrup, jelly or liqueur from it have been
used to treat respiratory tract, gastrointestinal disorders, fever, infections, colds, u,
rheumatism, and gout (Bobinaitė et al., 2020; Vogl et al., 2013); guelder rose has been
used to treat asthma, coughs and colds, heart troubles, stomach and kidney diseases and
disorders (Kajszczak et al., 2020); chokeberries and bilberries are useful for treatment
of atherosclerosis and chronic venous insufciency (Wallace & Giusti, 2015); elderberry in folk medicine has been used to treat respiratory diseases, wounds, kidney and
eye problems (Sidor & Gramza- Michałowska, 2015); dried barberry fruits are used to
treat diarrhea, fever, sore throat and as an antiarrhythmic and sedative (Abd El- Wahab
et al., 2013; Fatehi et al., 2005); hawthorn extract as a cardiovascular tonic has been
known since at least the rst century AD and it has been widely used in folk medicine
for the treatment of mild heart diseases (Ercisli et al., 2015). Hawthorn extracts nd
applications to treat such age- related diseases as atherosclerosis and arthritis as well as
upper respiratory infections, including colds, bronchitis, and pneumonia (Barros et al.,
2010), as an antispasmodic agent in the treatment of asthma and as a sedative for the
treatment of insomnia (WHO, 2010).
Although berries, berry extracts, and infusions rich in anthocyanins have been used
in traditional medicine since ancient times, anthocyanins themselves are not currently
used for therapeutic purposes in Western medicine, although research in this area is
being extremely intensive (Wallace & Giusti, 2015; Jaiswal et al., 2020).
The benecial properties of anthocyanins have been shown in many studies
conducted in animals and cell culture models, meanwhile there is limited information
in the literature obtained from clinical studies in humans. However, a lot of epidemiological studies conrmed the importance for health of including in the diet foods rich
in anthocyanin, in particular berries, which are the main source of anthocyanins in the

172 Wild Edible Plants
human diet, and, despite the fact that berries contain a variety of substances required for
complete nutrition, their benecial health properties are associated primarily with the
presence of anthocyanins (Yang & Kortesniemi, 2015). There are some clinical trials
that conrmed the positive health effect of regular intake of berries or their extracts on
human health.
A 20- year study involving 2,682 middle- aged or older Finnish men, showed that
the higher intake of berries may be related to reduction of the risk of type- 2 diabetes
(Mursu et al., 2014).
Even a short- term blueberry intake by overweight and obese children increased
the biological antioxidant potential in serum (Giongo et al., 2011). Regular consumption of wild blueberries improves memory, may slow cognitive decline, and reduces
symptoms of depression in older adults (Krikorian et al., 2010); reduces inammation
decreasing cardiometabolic risk in the long term in individuals with features of metabolic syndrome (Kolehmainen et al., 2012).
Effectiveness of a capsulated elderberry extract intake in the treatment of upper
respiratory symptoms have been shown in the meta- analysis of a randomized, controlled clinical trial (Hawkins et al., 2019). Consumption of elderberry extract improved
the respiratory and mental health of people under stressful conditions (Tiralongo et al.,
2016), and reduced the duration of inuenza infection (Zakay- Rones et al., 2004).
Cranberry supplementation signicantly reduced the risk of developing urinary
tract infections, reduced the administration of antibiotics (Luís et al., 2017), and can
be considered as adjuvant therapy for preventing urinary tract infections in susceptible
individuals (Xia et al., 2021).
Consumption of barberry juice by patients with diabetes signicantly reduced
blood pressure, fasting blood sugar, total cholesterol and triglyceride (Lazavi et al.,
2018). Daily intake of barberry by patients with type- 2 diabetes reduced blood lipids,
glucose, and insulin (Shidfar et al., 2012). Daily intake of barberry reduced oxidative
status in patients with metabolic syndrome (Mohammadi et al., 2014).
The positive effect of hawthorn extract on blood pressure reduction in humans has
been shown in randomized clinical trials (Plotnikoff & Dusek, 2018).
Epidemiologic study involving 99 826 women aged 55– 69 who had no cardiovascular diseases (CVD) found that dietary intakes of foods rich in anthocyanidins – strawberry – associated with signicant mortality reduction from CVD (Mink et al., 2007).
Consumption of strawberries or mixture of strawberries, bilberries, lingonberries, and
chokeberries with wheat bread reduced the postprandial insulin response caused by
starch present in bread in healthy females (Törrönen et al., 2013). It was shown that
inclusion of dried lingonberry (Vaccinium vitis- idaea L.) into meals with high glucose
content reduced postprandial glycemic response (Linderborg et al., 2012). Daily intake
of 500 g of cultivated strawberries for 1 month by healthy volunteers improved total
antioxidant capacity and plasma lipid prole, reducing the level of total cholesterol by
8.78%, low- density lipoprotein cholesterol by 13.72%, and triglycerides by 20.80%,
and antioxidant status (Alvarez- Suarez et al., 2014).
It was found that anthocyanins possess low bioavailability, so, anthocyanins from
the berries reach the gut where they are digested producing bioactive metabolites,
which may modulate the gut microbiota. There is evidence that anthocyanins derived
from various berries act as prebiotic substrates for gut bacteria promoting probiotics

Antioxidant Compounds in Wild Edible Berries 173
growth, particularly the benecial bacteria Lactobacillus spp., Bidobacterium spp.,
and Akkermansia spp. and suppressing the growth of harmful bacteria Clostridium
histolyticum and Bacteroides spp. (Faria et al., 2014; Liang et al., 2024; Wang et al.,
2022; Yang et al., 2014). Thus, it is currently reported that berry consumption may
regulate the composition of intestinal microorganisms, meanwhile gut microbiota is
closely associated with health status of human organism modulating immunity, the
host’s energy metabolism, weight gain or loss, obesity and related diseases (Piccioni
et al., 2022). Higher intakes of berries lead to a reduction in proinammatory mediators,
which may be the reason for observed benecial effects in cases of chronic inammation and cancer in the colon (Hosseini et al., 2018).
Finally, despite the evidence of positive results of anthocyanins on human health,
further clinical trials to support berries’ application in the management of chronic
diseases should be provided.
5.8 CONCLUSIONS
Wild berries contain valuable substances including anthocyanins’ consumption, which
helps to prevent some serious diseases, and, so, have high nutraceutical potential.
Despite signicant differences in the content of components that determine the nutritional value of wild berries depending on the variety, climate conditions, soil composition, and stage of ripening, they always remain an invaluable source of antioxidants and
other important substances for maintaining human health.
The pronounced health- promoting properties of wild berries give grounds to recommend them to be included in the everyday human diet.
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