27
ABSTRACT
Tannins are a group of polyphenolic compounds synthesized and accumulated by higher plants as secondary metabolites. 
They are divided into hydrolysable tannins and proanthocyanidins and are found in many plant tissues in which they occur in 
diverse structures and amounts. This review provides a brief background on tannin distribution in plants, and summarizes 
the current literature on tannins in strawberries, raspberries, blueberries, currently the most commonly cultivated and 
consumed berries, and chokeberries, which have become popular in the last decades. The effects of processing and storage 
on tannin composition and levels in processed products are also provided. 
Key words: ellagitannins, proanthocyanidins, strawberry, raspberry, blueberry, chokeberry
Agricultura 17: No 1-2: 27-36 (2020)
https://doi.org/10.18690/agricultura.17.1-2.27-36.2020
A Review of Tannins in Berries
Barbara BERNJAK, Janja KRISTL*
University of Maribor, Faculty of Agriculture and Life Sciences, Pivola 10, 2311 Hoče, Slovenia
*Correspondence to: 
E-mail: janja.kristl@um.si                                                                                                          Published in 2021
INTRODUCTION 
Tannins belong to the complex phenolic compounds, 
defined as phenolic derivatives synthesized by higher plants 
as secondary metabolites. More than 8000 tannin compounds 
have been isolated and chemically characterized (Krzyzowska 
et al., 2017). Tannins are phenylpropanoid compounds 
often condensed to polymers of variable length (Swanson, 
2003), which have molecular weights between 300 and 3000 
daltons (Da) (Mena et al., 2015). They are unstable and can 
be converted into various compounds when the plant cells 
are damaged (for example, during processing of plant raw 
materials) (Izawa et al., 2010). Tannins are synthesized by 
many plant species and can be found mainly in roots, stems, 
bark, leaves, buds and seeds, where they take up 5 to 10% 
of dry vascular plant material (Barbehenn and Constabel, 
2011). 
Structurally, tannins are divided into two classes: 
hydrolysable and condensed tannins (Izawa et al., 2010). 
Hydrolysable tannins are water-soluble (Izawa et al., 2010), 
and include ellagitannins, gallotannins and also more 
complex tannins. The basic structure of hydrolysable tannin 
is a carbohydrate whose hydroxyl groups are esterified with 
phenolic acids, such as gallic acid (Khanbabaee and Van Ree, 
2001).
The second group are proanthocyanidins (condensed 
tannins). They are the most abundant polyphenols in 
plants (Lamy et al., 2016). Condensed tannins are polymers 
of 2-50 flavonoid units, which are not susceptible to 
hydrolysis (Khanbabaee and Van Ree, 2001). Biosynthesis of 
proanthocyanidins requires products from the shikimate and 
acetate/malonate pathways. The starter and extension units of 
proanthocyanidins are generated via the flavonoid pathway, 
which shares the same upstream pathway with anthocyanidins, 
the substrates for anthocyanin synthesis. The key enzymes 
involved in this process are well known. Anthocyanidins can 
be catalysed by anthocyanidin reductase to produce flavan-
3-ols, important substrates for proanthocyanidin synthesis. 
Recent advances in understanding the molecular genetic 
basis of proanthocyanidin biosynthesis were described by 
Dixon and Sarnala (2020). 
The majority of condensed tannins are water-soluble, 
exceptions are some molecules with higher molecular weight 
(Izawa et al., 2010). They are more resistant to microbial 
degradation and also show stronger antiviral, antifungal, 
and antibacterial activities (Krzyzowska et al., 2017). It was 
found that condensed tannins, from some legume species, 
can be beneficial for cattle and other ruminants, because they 
reduce the risk of flatulence caused by high-protein diets and 
internal parasite loads (Constabel et al., 2014). More than 

28
90% of total tannins on the market are condensed tannins 
(Khanbabaee and Van Ree, 2001; Filgueira et al., 2017). 
Tannins in foods cause astringent sensations and a bitter 
taste, and play an important role, due to their potential 
beneficial effects on human health (Lamy et al., 2016). The 
highest amounts of tannins have been reported in coffee, 
cocoa, chocolate, green and black tea, red wine, nuts, legumes, 
cereal grains, fruits and vegetables (Crozier et al, 2009; 
Smeriglio et al, 2017). Berries are characterised by high levels 
of micronutrients, vitamin C, folates (Di Vittori et al., 2018), 
polyphenols like anthocyanidins, which are responsible for 
the intense colour in many berries and many other antioxidant 
phytonutrients (Milivojević et al., 2011). Pre-harvest factors 
such as cultivar, cultivation practices, environment and plant 
age have a significant impact on commercial, organoleptic 
and nutritional quality in berries (Alvarez-Suarez et al., 2014). 
Light exposure is an important environmental factor with 
positive effects on flavonoid synthesis in raspberries (Wang et 
al., 2009), strawberries (Anttonen et al., 2006), grapes (Matus 
et al., 2009), and blueberries (Uleberg et al., 2012). However, 
blueberries accumulate high levels of anthocyanins even 
when grown in shaded sites, suggesting that the stimulation of 
flavonoid biosynthesis is cultivation and cultivar dependent 
(Di Vittori et al., 2018). In such fruits the biosynthesis of 
various flavonoids depends on the developmental stage of the 
fruit, while environmental factors have little influence. The 
content of anthocyanins in strawberries is more dependent 
on the ripening stage, while the accumulation of flavonols 
and proanthocyanidins is more sensitive to environmental 
factors (Carbone et al., 2009). The effects of light exposure 
and light wavelengths on flavonoid biosynthesis in fruits are 
described in detail in the review by Zoratti et al. (2015). 
This paper reviews the distribution of tannins in plants with 
special attention to strawberries, raspberries and blueberries, 
which are the most widely grown and consumed berries 
in the EU (Oliveira et al., 2019), as well as in chokeberry, 
which has become popular in recent decades. The effects of 
processing and storage on the composition of ellagitannins 
and proanthocyanidins and their levels in berry products are 
also provided.
DISTRIBUTION OF TANNINS IN PLANTS 
The most common composition of tannins in nature is 
a mixture of proanthocyanidins and hydrolysable tannins 
with the latter being present in lower quantities. There are 
some exceptions such as the Acacia sp. and Terminalia sp. 
species, which are an important source of condensed and 
hydrolysable tannins, and a few dicotyledons species (Furlan 
et al., 2010). 
Tannins in plants occupy up to 20% of the dry weight 
(DW), ranking them after cellulose, hemicellulose and 
lignin. The synthesis of tannins in plants is often associated 
with defence responses against microbial pathogens, harmful 
insects, herbivores (Furlan et al., 2010) and UV-A or UV-B 
radiation. Polyphenols are stored in vacuoles and cell walls 
(Fraga-Corral et al., 2020). Because of this, tannins have been 
found in many plant tissues: wood, bark, roots, leaves, fruits, 
and seeds (Sieniawska and Baj, 2017). 
The occurrence of tannins within plants varies widely 
among tissues, organs, cell types (Constabel et al., 2014) and 
varies among different species of the same genus (Prida and 
Puech, 2006). The accumulation of tannins is usually specific 
to certain cell types. During the development of O. viciifolia 
leaves, the condensed tannins shift from the abaxial to the 
adaxial side of the leaf and also into specialized cells in the 
epidermis (Lees et al., 1993). Specialized cells, which contain 
condensed tannins, have also been found in the phloem 
parenchyma in young poplars. In this case, condensed 
tannins accumulate in the hypodermal cells of older stems 
and in the epidermal cells of young stems (Kao et al., 2002). 
The accumulation tendency of condensed tannins into the 
epidermal layers has a protective function against pathogens 
and UV stress (Close and McArthur, 2002). 
Considerable quantities of tannins are present in the roots 
of some woody plants. In P . tremuloides root tips, condensed 
tannins are localized in the cortex, lateral root cap and 
epidermal tissues (Kao et al., 2002), while in eucalyptus and 
jack pine they are found in a region between the suberized 
zone and root tip (Mckenzie and Peterson, 1995). Condensed 
tannins in seeds may contribute to the endosperm and 
embryo protection by blocking the movement of molecules 
that oppose seed dormancy prior to early germination and 
stress by serving as physical and chemical barriers (Lepiniec 
et al., 2006). High tannin content has been reported in 
seed coats of Phaseolus vulgaris and other beans (Jin et al., 
2012), many nuts, sorghum cereal seeds and barley (Prior 
and Gu, 2005). Fruits containing high amounts of tannins 
continue to accumulate them until fully ripened, giving 
them a corresponding astringent taste, while fruits with low 
amounts of tannins stop their production at an early stage of 
development (Akagi et al., 2009).
TANNINS IN BERRIES
The term “berry” refers to small fruits growing in wild 
shrubs that can be bitter or sweet, with a juicy mesocarp, have 
an intense red, blue or purple colour or in some cases also a 
white colour (Hidalgo and Almajano, 2017). Berry fruits are 
usually used because of their special taste and attractive colour. 
They primarily include strawberry, blackberry, blueberry, 
raspberry, cranberry, elderberry, mulberry and currants, 
which are the most abundant berries worldwide (Manganaris 
et al., 2014; Di Vittori et al, 2018). Lesser known species 
also classified as berry fruits are: huckleberry, chokeberry, 
lingonberry, boysenberry, olallieberry, gooseberry, barberry, 
dewberry, juneberry, tayberry and exotic berries: açai berry, 
goji berry, physalis, cloudberry, pineberry and salmonberry 
(Leafy place, 2019).
Strawberry
More than 20 ellagitannins have been determined in fruits, 
leaves and roots of strawberry (Fragaria x ananassa Duch.) 
(Gasperotti et al., 2013; Karlińska et al., 2021). They consist of 
glucose esterified with hexahydroxydiphenic acid (HHDP) 
and gallic acid (Aaby et al., 2012). Their content varies with 
A Review of Tannins in Berries

29
cultivar, plant growth stage, and plant part. The synthesis and 
accumulation of ellagitannins are the most intensive in the 
leaves, however, a decreasing tendency in their content was 
observed throughout plant development (Karlińska et al., 
2021). The growth stage is reported to have a greater effect on 
ellagitannin content in strawberry morphological parts than 
the cultivar (Karlińska et al., 2021), however, the opposite 
was reported for strawberry fruits of different cultivars 
grown in Italy (Gasperotti et al. 2013). The ellagitannin 
level of mature strawberry fruits varies from 6.53 to 52.38 
mg/100 g FW (fresh weight) (Buendia et al. 2010; Nowicka 
et al., 2019; Karlińska et al., 2021). Studies have shown that 
the monomeric ellagitannins in strawberry fruits, roots and 
leaves are pedunculagin, casuarictin and potentillin. The 
main product of potentillin dimerization is agrimoniin, 
which has been found to prevail in strawberry fruits (Aaby et 
al., 2012; Nowicka et al., 2019), leaves and roots (Karlińska et 
al., 2021). Immature fruits have shown the highest contents 
of agrimoniin and those tended to decrease and reach lower 
values in fully mature fruits (Gasperotti et al., 2013; Aaby 
et al., 2012; Fecka et al., 2021), and even less agrimoniin 
was detected in overripe fruits (Gasperotti et al., 2013). Its 
mean values for ‘Syrena’ , ‘Pandora ISK’ , ‘Selvik’ , and ‘Elvira’ 
cultivars vary from 0.17 mg/g in green fruits to 0.118 mg/g 
in pink and 0.111 mg/g FW in red fruits (Fecka et al., 2021). 
Agrimoniin was proposed a s chemotaxonomic marker 
for Fragaria (Okuda et al., 1992). The levels of agrimoniin 
did not show large differences between 27 cultivars grown 
in Norway (Aaby et al. 2012) with the average value of 8.8 
mg/100g FW , however, higher variability was observed for 
90 cultivars grown in Poland, with the values in the range 
from 3.60 to 30.79 mg/100 g FW (Nowicka et al., 2019). 
Woodland strawberries (Fragaria vesca) contain higher 
contents of ellagitannins and ellagic acid conjugates when 
compared to cultivated strawberries (Gasperotti et al., 2013). 
The fragariin, sanguiin H-2, ellagitannin with MW 1718, and 
β-ellagitannin with MW 1718 are generally present in much 
lower concentrations. The ratio of monomeric and dimeric 
ellagitannins in leaves and roots is relatively equal, whereas in 
fruits the dimeric forms predominate (Karlińska et al., 2021). 
The majority of ellagic acid in strawberry fruits is found 
in a bound form as a part of ellagitannins and constitutes, 
together with conjugated derivatives, less than 5% of total 
phenolics (Buendia et al. 2010).
Aaby et al. (2012) reported that the most abundant 
class of polyphenolic compounds in mature strawberry 
fruits are anthocyanins, which account for 41% of the 
total phenolic content, followed by flavan-3-ols (28 %) 
and ellagitannins (14%). However, Buendia et al. (2010) 
found proanthocyanidins to be the predominant phenolic 
compounds in Spanish strawberry cultivars with the values 
ranging from 0.539 to 1.632 mg/g FW . Similar findings 
were reported for strawberry fruits (Fragaria x ananassa 
Duch.) grown in Trentino in which proanthocyanidins 
represented between 54.8 and 77.4% of polyphenolic 
compounds (Gasperotti et al., 2015). Proanthocyanidin 
profile in strawberry fruit is complex. The terminal flavan-
3-ol in oligomers is (epi)catechin, and the extension units 
are (epi)catechin (60-70% of the proanthocyanidins) and 
(epi)afzelechin (propelargonidin). It has been reported that 
the degree of polymerization varies within cultivars. It is 
proposed that cultivars characterized with higher contents of 
monomers and dimers can potentially have better biological 
activity, because they can be better absorbed than larger 
proanthocyanidins oligomers (Buendia et al. 2010). Many 
potential health effects of ellagitannins have been listed in the 
literature. Their effect on the brain’s hippocampus is reducing 
the effect of aging in spatial orientation (Shukitt-Hale et 
al., 2007). They also influence the gastric epithelial cells 
by inhibiting the inflammatory response to TNF through 
NF-B dependent and independent mechanisms (Fumagalli 
et al., 2016), reduce the level of specific biomarkers for 
cardiovascular diseases, inflammation, blood pressure, lowers 
LDL cholesterol, controls glycaemia and promote antitumor 
activity in esophageal, lung and colon cancers (Desjardins, 
2014). A more detailed discussion of potential biological 
functions of ellagitannins and their metabolites is provided 
in the review paper by Landete (2011).
Strawberries as well as other Rosaceae fruits are stored 
frozen or processed in purees, juices, syrups and jams, 
because of their short shelf life. It has been reported that 
the transfer of ellagitannins from fresh fruits to products 
depends on the technological process used. The production 
of purees seems to be superior to the production of juices, 
since 56 to 92% of the total ellagitannins present in fresh 
fruits are transferred to purees, while they are lost in juice 
production, were 65 to 90% are present. The losses have been 
attributed to the high molecular weight compounds (1871-
2038 Da) that remain in pomace from unclarified juice 
production (Milczarek et al. 2021). It has been proposed 
that in the course of fruit processing, high molecular weight 
ellagitannins are depolymerized or degraded to conjugates, 
and that some ellagitannins could be exposed to hydrolysis 
leading to elevated ellagic acid levels in pomaces (Oszmiański 
and Wojdylo, 2009; Milczarek et al. 2021).
Raspberry
A major class of polyphenols in raspberries are ellagitannins 
which represent 53.5% to 75.9% of the total polyphenol 
content, with 100% being the sum of ellagitannins, 
anthocyanins, flavanols and flavonols (Mullen et al., 2002a; 
Sójka et al., 2016). Ellagitannins in raspberries are a mixture 
of monomeric and oligomeric tannins, which structure 
is characterized by ellagic and gallic acid moieties and 
sanguisorboyl linking ester group (Vrhovsek et al., 2006). 
Free ellagic acid represents a small fraction of the total ellagic 
acid released during hydrolysis of plant material, mainly 
from sanguiin H-6 and lambertianin C (Määttä-Riihinen et 
al., 2004; Mullen et al., 2002a). Relatively large variations in 
free ellagic acid (1.98-5.24 mg/kg FW) and total ellagitannins 
content (94.15-326 mg/100 g FW) have been reported for 
fruits of different cultivated raspberry (R. idaeus) cultivars 
(Milivojević et al., 2010; Bobinaité et al., 2012; Smeriglio et 
al., 2017; Vrhovsek et al., 2008), with higher levels of free 
ellagic acid detected in wild Rubus berries (12.71 mg/kg), 
which are usually characterized with smaller fruits of a more 
intensive colour (Çekiç and Őzgen, 2010).
The ellagitannin profile of raspberries usually consists of two 
A Review of Tannins in Berries

30
compounds. The main ellagitannin present in red (R. idaeus) 
and black (R. occidentalis) raspberries is the dimer, sanguiin 
H-6 with values ranging from 135 to 1743 mg/100 g DW and 
1537 mg/100 g DW , respectively (Sparzak et al., 2010; Kula et 
al., 2016). The trimer, lambertianin C, which consists of six 
HHDP , three glucosyl and three galloyl moieties, occurs in 
much lower concentrations with notable differences between 
certain cultivars (Mullen et al., 2003; Kula et al., 2016). The 
sanguiin H-6 (5% of DW) and also free ellagic acid (1% of 
DW) dominate in shoots of R. idaeus which are commonly 
used in folk medicine as herbal remedies (Krauze-Baranowska 
et al., 2014). Proanthocyanidins are B-type polymers and are 
composed of catechin, epicatechin and epiafzelechin. Their 
content in raspberries amounts to 79 mg/100 g FW (Gu et al., 
2004) and prevail in seeds and in insoluble parts of the skin.
The total content of ellagitannins and proanthocyanidins 
in maturing raspberry fruit decreases as maturity approaches 
(Beekwilder et al., 2005). Beside cultivar and stage of maturity, 
the content of phenolic compounds, including tannins, 
depends on cultivation practices, environmental conditions, 
and storage time (Bobinaité et al., 2012; Mazur et al., 2014).
Raspberries are perishable soft fruits and are preserved by 
deep freezing or processed into juice, jam or syrup. The fruits 
are also preserved by various drying techniques such as hot 
air-drying, freeze-drying, microwave-drying, and hot pump-
drying. The advantages and disadvantages of these methods 
are discussed in the review paper prepared by Piccolo et 
al. (2020). The authors concluded that the freeze-drying 
method is most likely the best choice for the preservation of 
bioactive compounds. Industrial processing of raspberries 
into juice causes losses of bioactive compounds. This is a 
consequence of the processing conditions which result in 
a polyphenol transformation or degradation and changes 
in fruit morphological characteristics. An interesting study 
was published by Sójka et al. (2016) reporting that raspberry 
fruit juice, when compared to fresh fruits, retains on average 
11.8% of ellagitannins. The majority of the ellagitannins 
remains in the press cake, especially in its seedless fraction, 
which is characterized by high levels of ellagitannins and 
proanthocyanidins. The phenolic composition of juices 
depends on the processed cultivar. The juice prepared from 
certain cultivars retains more anthocyanins and others 
more ellagitannins, when compared to fresh fruits. Further 
information highlighting the steps in juice processing, 
which cause significant losses and compositional changes 
of ellagitannins, are provided by Howard et al. (2012). 
The authors also discussed the possible mechanisms for 
ellagitannins losses during the processing into juices, purees, 
and canned products. Processing of raspberries into jams 
does not affect the ellagic acid glycoside content, which 
remains quite stable, while an increase in free ellagic acid 
content is usually observed, probably because of its release 
from ellagitannins with thermal treatment (Zafrilla et al., 
2001). However, the content of total ellagic acid in raspberry 
jams is much lower, 23-36% of those being present in the 
unprocessed berries (Koponen et al., 2007; Zafrilla et al., 
2001). When raspberries are frozen within 3 hours of 
harvest and stored at -30 °C for short time (4-5 days), no 
discernible difference in lambertianin C and sanguiin H-6 is 
observed, whereas storage of fruit at 4 °C for 3 days resulted 
in an increase in both ellagitannins (Mullen et al., 2002b). 
Therefore, consummation of freshly picked, freshly frozen or 
fresh commercial fruits, provides the intake of similar levels 
of phytochemicals. Prolonged storage in the freezer (one 
year) maintains total phenolics content, although ellagic acid 
decreases (De Ancos et al., 2000; Tϋrkben et al., 2010).
Blueberry
The highest amount of polyphenols in blueberries 
(Vaccinium cormymbosum L.) represents flavonoids, 
especially 25 individual anthocyanins (1000 mg/100 g 
FW), while flavonols, flavan-3-ols and proanthocyanidins, 
follow in considerably lower amounts. Blueberries are 
almost devoid of ellagitannins, while proanthocyanidins 
are present in a varying amounts with different degrees of 
polymerization; from monomeric flavanols to oligomers 
(degree of polymerization (DP) 2-10), and to high molecular 
weight polymers (DP > 10) (Gu et al., 2004). Oligomers in 
blueberries consist of catechin and epicatechin units that 
are singly linked [B
1
 (monomer) through B
8
 (octamer)] 
(Prior et al., 2001). Terminal units of blueberry polymeric 
proanthocyanidins consists of epicatechin and catechin, 
while extension units consist of epicatechin (Gu et al., 2002). 
The total amount of tannins in blueberries is on average 160 
mg/100 g FW (Diaconeasa et al., 2015). Highly polymerized 
forms of proanthocyanidins dominate in blueberry extracts, 
accounting for 72% of the total extractable proanthocyanidins 
(DP > 10) (Hellström et al., 2009).
The contents of proanthocyanidins in blueberries grown 
in Sweden varied from 13.9 in the ‘Camelia’ to 19.8 mg/100 
g FW in the ‘Duke’ cultivar (Liu et al., 2020). Much higher 
levels were reported by Smeriglio et al. (2017) and Gu et al., 
(2004) for cultivated blueberries (highbush) 87-274 and 179 
mg/100 g FW , respectively, and wild berries (lowbush) 311-
335 mg/100 g FW . However, the literature data is inconsistent. 
Prior et al. (2001) found that wild blueberries contain higher 
levels of total procyanidins compared to cultivated berries. 
Blueberry processing into products results in remarkable 
losses of proanthocyanidins. The losses of proanthocyanidins 
are caused during the large number and complexity of 
processing steps. Clarified blueberry juices preserve on 
average 38% of monomers, 58% of dimers, 24% of trimers, 
20% of tetramers and less than 11% of pentamers of 
proanthocyanidins present in frozen berries. Octamers were 
not detectable. Similar losses of compounds were reported for 
unclarified juices (Rodriguez-Mateos et al., 2014). Regarding 
the total levels of proanthocyanidins, nonclarified and 
clarified juices retained 19-36% and 23-47% of that in frozen 
berries (Brownmiller et al., 2009; Howard, et al., 2012). 
The greatest retention of proanthocyanidins is expected in 
simple canning processes in which thawed berries are covered 
with water or syrup and then pasteurized. In such processes, 
the berries remain intact and the enzymatic degradation 
is limited. Oligomers with DP > 3 were retained less when 
compared to proanthocyanidin monomers and dimers 
(Howard et al., 2012). The losses could be a consequence 
of a preferential binding of the large-molecular-weight 
proanthocyanidins to cell-wall polymers, which occurred 
A Review of Tannins in Berries

31
after cell disruption by heating and mixing. However, it is 
also possible that in response to thermal treatment, larger 
oligomers were depolymerized to monomers and dimers 
(Brownmiller et al., 2009).
Rodriguez-Mateos et al. (2014) studied the changes in 
proanthocyanidins content during the baking of a product 
with freeze-dried wild blueberry powder. They found no 
differences in total proanthocyanidin content. During 
the whole process, the content of lower molecular weight 
oligomers (dimers and trimers) increased by 36 and 28%, 
while nonamers and decamers completely disappeared. 
Proanthocyanidin content in blueberry products declined 
during 6 months of storage at 25 °C. Blueberries canned in 
water retained 38%, blueberries canned in syrup retained 
29%, while purees and juices retained less than 11% of 
proanthocyanidins. Larger oligomers in processed products 
were less stable than monomers and dimers (Brownmiller et 
al., 2009; Howard et al., 2012).
Chokeberry
The genus Aronia is represented by two species Aronia 
melanocarpa (Michx.) Elliot (black chokeberry) and Aronia 
arbutifolia (L.) Pers. known as red chokeberry. Aronia 
melanocarpa is the predominant commercial chokeberry 
cultivar and has gained increased popularity due to its 
assumed health-promoting effects, which have been reviewed 
by Sidor et al. (2019) and Kokotkiewicz et al. (2010) together 
with the berries pharmacologically relevant constituents. 
Chokeberries are very valuable as a food ingredient and are 
used in the food industry mainly for the production of juice, 
jam and wine, and as a natural colorant. 
The fruits have an astringent taste due to a high tannin 
content (Wu et al., 2004). They mainly contain polymeric 
proanthocyanidins (DP > 10) which account for about 66% 
to 82% of total polyphenolic compounds (Oszmiański and 
Wojdylo, 2005, Denev at al., 2018; Wu et al., 2004). Even 
higher amounts are present in smaller fruits (Wangensteen et 
al., 2014) and chokeberry leaves, for which 22% higher levels 
of proanthocyanidins were reported when compared to ripe 
fruits (Teleszko and Wojdylo, 2015). According to literature, 
their levels in fruits vary from 522 mg/100 g FW to 3671 
mg/100 g FW (Denev et al., 2018; Wu et al., 2004; Taheri et 
al., 2013), however, no significant differences were reported 
between black, red and purple (Aronia prunifolia) coloured 
chokeberry fruits (Taheri et al., 2013). Proanthocyanidins 
predominated in the berry flesh (70%) followed by the 
skin (25%) and kernels (5%) (Mayer-Miebach et al., 2012). 
An increasing trend in their content was observed with 
prolonged harvest time (Poyraz Engin and Mert, 2020). 
Chokeberry proanthocyanidins have been identified 
exclusively as procyanidin B-type, containing epicatechin 
as the main monomer unit (Oszmiański and Wojdylo, 
2005). Cultivated and wild A. melanocarpa fruits have a 
similar oligomeric proanthocyanidin composition (Sueiro 
et al., 2006). Different chokeberry varieties contain 80-95% 
extractable proanthocyanidins (Hellström et al., 2009; Taheri 
et al., 2013), while ellagitannins have not been detected 
(Kähkönen et al., 2001). In contrast, the chokeberry leaves, 
which are not used as functional food, are characterized by 
high proportions of flavanols (Teleszko and Wojdylo, 2015). 
The total proanthocyanidin content ranges from 
approximately 1408-1579 mg/100 g DW for chokeberry juice 
to 8192-9586 mg/100 g in pomace (Oszmiański and Wojdylo, 
2005; Rodríguez-Werner et al., 2019) and depends on genetic 
attributes, harvest date, cultivation location and practice, 
processing and storage. Their levels remain stable upon 
blanching, and then increase by 11% after enzyme treatment, 
probably due to the disruption of cell wall polysaccharides 
and proteins to which polymeric procyanidins are bound. The 
higher losses of about 40% of proanthocyanidins occurred 
during the pressing operation after which the majority of 
them remains in the pomace (Mayer-Miebach et al., 2012). 
Oszmiański and Lachowicz (2016) reported that the content 
of procyanidin polymers in juices prepared from crushed 
fruits before processing was higher by over 62% higher than 
in juices prepared from non-crushed berries. Juices stored 
for 6 months at 25 °C, retained more than 90% of the total 
proanthocyanidins (Wilkes et al., 2014). They are quite stable 
as no degradation was noticed after heating purees up to 100 
°C for 20 min (Mayer-Miebach et al., 2012).
CONCLUDING REMARKS
Berry fruits are important sources of tannins. Their 
content and chemical composition depend on the species, 
variety, cultivation practice, and treatment before and 
after harvest. Ellagitannins are found in strawberries and 
raspberries, but are less common in other berry fruits. The 
major class of tannins in blueberries and chokeberries are 
proanthocyanidins, while strawberries are characterized by 
both ellagitannins and proanthocyanidins. Chokeberries are 
characterized with the highest content of condensed tannins 
among 100 plant foods investigated. All these berries can be 
consumed fresh or processed into purees, juices, syrups, and 
jams, due to their short shelf life. They also can be preserved 
by deep freezing or by different drying techniques. Currently, 
cold storage or freeze-drying is the most effective strategy to 
preserve the colour and polyphenol content in berries and 
their products. Tannins are lost during processing to varying 
degrees, depending on the production technology. In general, 
processes comprised of more steps (e.g. juice production) 
result in the greatest losses. As large amounts of bioactive 
compounds are annually discharged in food by-products, 
challenges exist to improve the most critical steps and to retain 
these compounds in berry products. During processing and 
storage of berry products, the tannin composition is altered. 
What exactly happens to the various compounds belonging 
to the class of tannins during these processes is poorly 
understood and requires further consideration.
REFERENCES
Aaby, K., Mazur, S., Nes, A., & Skrede, G. (2012). Phenolic 1. 
compounds in strawberry (Fragaria × ananassa Duch.) 
fruits: Composition in 27 cultivars and changes during 
A Review of Tannins in Berries

32
ripening. Food Chemistry, 132, 86-97.
Akagi, T., Katayama-Ikegami, A., Suzuki, Y ., Y oshida, J., 2. 
Y amada, M., Sato, A., & Y onemori, K. (2009). Expression 
balances of structural genes in shikimate and flavonoid 
biosynthesis cause a difference in proanthocyanidin 
accumulation in persimmon (Diospyros kaki) fruit. 
Planta, 230, 899-915.
Alvarez-Suarez, J.M., Giampieri, F., Tulipani, S., Casoli, 3. 
T., Di Stefano, G., González-Paramás, A.M., Santos-
Buelga, C., Busco, F., Quiles, J.L., Cordero, M.D., 
Bompadre, S., Mezzetti, B., & Battino, M. (2014). One-
month strawberry-rich anthocyanin supplementation 
ameliorates cardiovascular risk, oxidative stress 
markers and platelet activation in humans. The Journal 
of Nutritional Biochemistry, 25, 289-294.
Anttonen, M.J., Hoppula, K.I., Nestby, R., Verheul, M.J., 4. 
& Karjalainen, R.O. (2006). Influence of fertilization, 
mulch color, early forcing, fruit order, planting date, 
shading, growing environment, and genotype on the 
contents of selected phenolics in strawberry (Fragaria x 
ananassa Duch.) fruits. Journal of Agricultural and Food 
Chemistry, 54 (7), 2614-2620.
Barbehenn, R.V ., & Constabel, C.P . (2011). Tannins in 5. 
plant-herbivore interactions. Phytochemistry, 72 (13), 
1551-1565. 
Beekwilder, J., Hall, R.D., & de Vos, C.H. (2005). 6. 
Identification and dietary relevance of antioxidants 
from raspberry. BioFactors, 23, 197-205.
Bobinaité, R., Viskelis, P ., & Venskutonis, R. (2012). 7. 
Variation of total phenolics, anthocyanins, ellagic acid 
and radical scavenging capacity in various raspberry 
(Rubus spp.) cultivars. Food Chemistry, 132, 1495-1501.
Brownmiller, C.R., Howard, L.R., & Prior, R.L. 8. 
(2009). Processing and storage effects on procyanidin 
composition and concentration of processed blueberry 
products. Journal of Agricultural and Food Chemistry, 
57, 1896-1902.
Buendia, B., Gil, M.I., Tudela, J.A., Gady, A.L., Medina, 9. 
J.J., Soria, C., López, J.M., & Tomás-Barberán, F.A. (2010). 
HPLC-MS analysis of proanthocyanidin oligomers and 
other phenolics in 15 strawberry cultivars. Journal of 
Agriculture and Food Chemistry, 58, 3916-3926.
Carbone, F., Preuß, A., De Vos, R., D’ Amico, E., 10. 
Perrotta, G., Bovy, A.G., Martens, S., & Rosati, C. 
(2009). Developmental, genetic and environmental 
factors affect the expression of flavonoid genes, enzymes 
and metabolites in strawberry fruits. Plant Cell and 
Environment, 32 (8), 1117-1131.
Çekiç, C., & Őzgen, M. (2010). Comparison of 11. 
antioxidant capacity and phytochemical properties of 
wild and cultivated red raspberries (Rubus idaeus L.). 
Journal of Food Composition and Analysis, 23, 540-544.
Close, D.C., & McArthur, C. (2002). Rethinking the role 12. 
of many plant phenolics – protection from photodamage 
not herbivores? Oikos, 99, 166-172.
Constabel, C.P ., Y oshida, K., & Walker, V . (2014). 13. 
Diverse ecological roles of plant tannins: plant defense 
and beyond. In: A. Romani, V . Lattanzio & S. Quideau 
(Ed.), Recent Advances in Polyphenol Research (Chapter 
5) [E-reader version]. Retrieved from: https://www.
researchgate.net/publication/269277107_Diverse_
Ecological_Roles_of_Plant_Tannins_Plant_Defense_
and_Beyond
Crozier, A., Jaganath, I.B., & Clifford, M.N. (2009). 14. 
Dietary phenolics: chemistry, bioavailability and effects 
on health. Natural Product Reports, 26, 1001-1043.
De Ancos, B., Ibaῆez, E., Reglero, G., & Cano, M.P . 15. 
(2000). Frozen storage effects on anthocyanins and 
volatile compounds of raspberry fruit. Journal of 
Agricultural and Food Chemistry, 48 (3), 873-879.
Denev, P ., Kratchanova, M., Petrova, I., Klisurova, D., 16. 
Georgiev, Y ., Ognyanov, M., & Y anakieva, I. (2018). 
Black chokeberry (Aronia melanocarpa (Michx.) Elliot) 
fruits and functional drinks differ significantly in their 
chemical composition and antioxidant activity. Hindawi 
Journal of Chemistry, 2018, Article ID 9574587.
Desjardins, Y . (2014). Human health effects of strawberry: 17. 
a review of current research. Acta Horticulturae, 1049, 
827-838.
Diaconeasa, Z., Ranga, F., Rugina, D., Leopold, L., 18. 
Pop, O., Vodnar, D., Cuibus, L., & Socaciu., C. (2015). 
Phenolic content and their antioxidant activity in 
various berries cultivated in Romania. Food Science and 
Technology, 72 (1), 99-103.
Di Vittori, L., Mazzoni, L., Battino, M., & Mezzetti, B. 19. 
(2018). Pre-harvest factors influencing the quality of 
berries. Scientia Horticulturae, 233, 310-322.
Dixon, R.A., & Sarnala, S. (2020). Proanthocyanidin 20. 
biosynthesis – a matter of protection. Plant Physiology, 
184 (2), 579-591.
Fecka, I., Nowicka, A., Kucharska, A.Z., & Sokól- 21. 
Łetowska, A. (2021). The effect of strawberry ripeness 
on the content of polyphenols, cinnamates, L-ascorbic 
and carboxylic acids. Journal of Food Composition and 
Analysis, 95, 103669.
Filgueira, D., Molde, D., Fuentealba, C., & Garcia, D.E. 22. 
(2017). Condensed tannins from pine bark: a novel 
wood surface modifier assisted by laccase. Industrial 
Crops and Products, 103, 185-194.
Fraga-Corral, M., Garcia-Oliveira, P ., Pereira, A.G., 23. 
Lourenço-Lopes, C., Jimenez-Lopez, C., Prieto, M.A., 
& Simal-Gandara, J. (2020). Technological application 
of tannin-based extracts. Molecules, 25, 614, 1-27.
Fumagalli, M., Sangiovanni, E., Vrhovsek, U., Piazza, 24. 
S., Colombo, E., Gasperotti, M., Mattivi, F., De Fabiani, 
E., & Dell´Agli, M. (2016). Strawberry tannins inhibit 
IL-8 secretion in a cell model of gastric inflammation. 
Pharmacological Research, 111, 703-712. 
Furlan, C.M., Motta, L.B., & Cursino dos Santos, D.Y .A. 25. 
(2010). Tannins: What do they represent in plant life? In: 
G.K. Petridis (Ed.), Tannins: Types, Foods Containing, 
and Nutrition (pp. 251-263). São Paulo, Brazil. Nova 
Science Publishers. 
Gasperotti, M., Masuero, D., Guella, G., Palmieri, L., 26. 
Martinatti, P ., Pojer, E., Mattivi, F., & Vrhovsek, U. 
(2013). Evolution of Ellagitannin Content and Profile 
during Fruit Ripening in Fragaria spp. Journal of 
Agricultural and Food Chemistry, 61, 8597-8607.
Gasperotti, M., Masuero, D., Mattivi, F., & Vrhovsek, 27. 
U. (2015). Overall dietary polyphenol intake in a 
A Review of Tannins in Berries

33
bowl of strawberries: The influence of Fragaria spp. In 
nutritional studies. Journal of Functional Foods, 18, Part 
B, 1057-1069.
Gu, L., Kelm, M.A., Hammerstone, J.F., Beecher, G., 28. 
Holden, J., Haytowitz, D., Gebhardt, S., & Prior, R.L. 
(2004). Concentrations of proanthocynidins in common 
foods and estimations of normal consumption. The 
Journal of Nutrition, 134, 613-617.
Gu, L., Kelm, M., Hammerstone, J.F., Beecher, G., 29. 
Cunningham, D., Vannozzi, S., & Prior, R.L. (2002). 
Fractionationa of polymeric procyanidins from lowbush 
blueberry and quantification of procyanidins in selected 
foods with an optimized normal-phase HPLC-MS 
fluorescent detection method. Journal of Agricultural 
and Food Chemistry, 50, 4852-4860.
Hellström, J.K., Törrönen, A.R., & Mattila, P .H. (2009). 30. 
Proanthocyanidins in common food products of plant 
origin. Journal of Agricultural and Food Chemistry, 57 
(17), 7899-7906.
Hidalgo, G.I., & Almajano, M.P . (2017). Red fruits: 31. 
extraction of antioxidants, phenolic content and radical 
scavenging determination: a review. Antioxidants 
(Basel), 6 (1), 310-322.
Howard, L.R., Prior, R.L., Liyanage, R., & O Lay, 32. 
J. (2012). Processing and storage effect on berry 
polyphenols: challenges and implications for bioactive 
properties. Journal of Agricultural and Food Chemistry, 
60 (27), 6678-6693.
33. Izawa, K., Amino, Y ., Kohmura, M., Ueda, Y ., & Kuroda, 
M. (2010). Human – Environment interactions – Taste. 
In: H. Liu & L. Mander (Ed.), Comprehensive natural 
products II (Chapter 4.16.5.1) [E-reader version]. 
Retrieved from: https://www.sciencedirect.com/science/
article/pii/B9780080453828001088?via%3Dihub
34. Jin, A., Ozga, J.A., Lopes-Lutz, D., Schieber, A., 
& Reinecke, D.M. (2012). Characterization of 
proanthocynidins in pea (Pisum sativum L.), lentil 
(Lens culinaris L.), and faba bean (Vicia faba L.) seeds. 
Food Research International, 46, 528-535.
35. Kähkönen, M.P ., Hopia, A.I., & Heinonen, M. (2001). 
Berry phenolics and their antioxidant activity. Journal 
of Agricultural and Food Chemistry, 49(8), 49076-4082.
36. Kao, Y .Y ., Harding, S.A., & Tsai, C.J. (2002). Differential 
expression of two distinct ohenylalanine ammonia-
lyase genes in condensed tannin-accumulating and 
lignifying cells of quaking aspen. Plant Physiology, 130, 
796-807.
37. Karlińska, E., Masny, A., Cieślak, M., Macierzyński, 
J., Pecio, L., Stochmal, A., & Kosmala, M. (2021). 
Ellagitannins in roots, leaves, and fruits of strawberry 
(Fragaria × ananassa Duch.) vary with developmental 
stage and cultivar. Scientia Horticulturae, 275, 109665.
38. Khanbabaee, K., & Van Ree, T. (2001). Tannins: 
classification and definition. Natural Product Reports, 
18, 641-649. 
39. Kokotkiewicz, A., Jaremicz, Z., & Luczkiewicz, M. 
(2010). Aronia plants: a review of traditional use, 
biological activities, and perspectives for modern 
medicine. Journal of Medicinal Food, 13(2), 255-269.
40. Koponen, J.M., Happonen, A.M., Mattila, P .H., & 
Tőrrőnen, A.R. (2007). Contents of anthocaynins and 
ellagitannins in selected foods consumed in Finland. 
Journal of Agricultural and Food Chemistry, 55, 1612-
1619. 
41. Krauze-Baranowska, M., Gtód, D., Kula, M., Majdan, 
M., & Hatasa, R. (2014). Chemical composition and 
biological activity of Rubus idaeus shoots – a traditional 
herbal remedy of Eastern Europe. BMC Complementary 
Medicine and Therapies, 14, 480.
42. Krzyzowska, M., Tomaszewska, E., Ranoszek-Soliwoda, 
K., Bien, K., Orlowski, P ., Celichowski, G., & Grobelny, J. 
(2017). Tannic acid modification of metal nanoparticles: 
possibility for new antiviral applications. In: E. 
Andronescu & A. Grumezescu (Ed.), Nanostructures for 
oral medicine (Chapter 12) [E-reader version]. Retrieved 
from: https://www.sciencedirect.com/science/article/
pii/B9780323477208000134 
43. Kula, M., Majdan, M., Gtód, D., & Krauze-Baranowska, 
M. (2016). Phenolic composition of fruits from different 
cultivars of red and black raspberries grown in Poland. 
Journal of Food Composition and Analysis, 52, 74-82.
44. Lamy, E., Pinheiro, C., Rodrigues, L., Capela e Silva, 
F., Silva Lopes, O., Tavares, S., & Gaspar, R. (2016). 
Determinants of tannin-rich food and beverage 
consumption: oral perception vs. psychosocial aspects. 
In C.A. Combs (Ed.), Tannins: Biochemistry, food 
sources and nutritional properties, [E-reader version] 
(pp. 29-58). Retrieved from: file:///C:/Users/uporabnik/
Downloads/Proofs_ChapterID_37173_6x91.pdf
45. Landete, J.M. (2011). Ellagitannins, ellagic acid and 
their derived metabolites: A review about source, 
metabolism, functions and health. Food Research 
International, 44(5), 1150-1160.
46. Leafy place. (2019). 23 Types of Berries: List of berries 
with their picture and name. Retrieved from https://
leafyplace.com/types-of-berries/.
47. Lees, G.L., Suttill, N.H., & Gruber, M.Y . (1993). 
Condensed tannins in sainfoin.1. A histological and 
cytological survey of plant tissues. Canadian Journal of 
Botany, 71, 1147-1152.
48. Lepiniec, L., Debeaujon, I., Routaboul, J.-M., Baudry, 
A., Pourcel, L., Nesi, N., & Caboche, M. (2006). Genetics 
and biochemistry of seed flavonoids. Annual Review of 
Plant Biology, 57, 405-430.
49. Liu, J., Hefni, M.E., & Witthőft, C.M. (2020). 
Characerization of flavonoid compounds in common 
Swedish berry species. Foods, 9(3), 358.
50. Määttä-Riihinen, K.R., Kamal-Eldin, A., & Tőrrőnen, 
A.R. (2004). Identification and quantification of 
phenolic compound in berries of Fragaria and Rubus 
species (Family Rosaceae). Journal of Agricultural and 
Food Chemistry, 52, 6178-6187.
51. Manganaris, G.A., Goulas, V ., Vicente, A.R., & Terry, 
L.A. (2014). Berry antioxidants: small fruits providing 
large benefits. Journal of the Science of Food and 
Agriculture, 94(5), 825-833.
52. Matus, J.T., Loyola, R., Vega, A., Peῆa-Neira, A., Bordeu, 
E., Arce-Johnson, P ., & Alcalde, J.A. (2009). Post-
veraison sunlight exposure induces MYB-mediated 
transcriptional regulation of anthocyanin and flavonol 
A Review of Tannins in Berries

34
synthesis in berry skins of Vitis vinifera. Journal of 
Experimental Botany, 60(3), 853-867.
53. Mayer-Miebach, E., Adamiuk, M., & Behsnilian, D. 
(2012). Stability of chokeberry bioactive polyphenols 
during juice processing and stabilization of a polyphenol-
rich amterial from the by-product. Agriculture, 2(3), 
244-258.
54. Mazur, S.P ., Nes, A., Wold, A.-B., Remberg, S.F., & Aaby, 
K. (2014). Quality and chemical composition of ten red 
raspberry (Rubus idaeus L.) genotypes during three 
harvest seasons. Food Chemistry, 160, 233-240.
55. McKenzie, B.E., & Peterson, C.S. (1995). Root browning 
in Pinus banksiana and Eucalyptus pilularis Sm.1. 
Anatomy and permeability of the white and tannin 
zones. Botanica Acta, 108, 127-137.
56. Mena, P ., Calani, L., Bruni, R., & Del Rio, D. (2015). 
Bioactivation of High-Molecular-Weight Polyphenols by 
the Gut Microbiome. Diet-Microbe Interactions in the 
Gut, 73-101.
57. Milczarek, A., Sójka, M., & Klewicki, R. (2021). Transfer 
of ellagitannins to unclarified juices and purees in the 
processing of selected fruits of the Rosaceae family. 
Food Chemistry, 344, 128684. 
58. Milivojević, J., Maksimović, V ., Nikolić, M., Bogdanović, 
J., Maletić, R., & Milatović, D. (2010). Chemical and 
antioxidant properties of cultivated and wild Fragaria 
and Rubus berries. Journal of Food Quality, 34, 1-9.
59. Milivojević, J.M., Nikolić, M.D., Dradišić Maksimović, 
J., & Radivojević, D.D. (2011). Generative and fruit 
quality characteristics of primocane fruiting red 
raspberry cultivars. Turkish Journal of Agriculture and 
Forestry, 35, 289-296.
60. Mullen, W ., McGinn, J., Lean, M.E.J., MacLean, M.R., 
Gardner, P ., Duthie, G.G., Y okota, T., & Crozier, A. 
(2002a). Ellagitannins, flavonoids, and other phenolics 
in red raspberries and their contribution to antioxidant 
capacity and vasorelaxation properties. Journal of 
Agricultural and Food Chemistry, 50, 5191-5196.
61. Mullen, W ., Stewart, A.J., Lean., M.E.J., Gardner, P ., 
Duthie, G.G., & Crozier, A. (2002b). Effect of freezing 
and storage on the phenolics, ellagitannins, flavonoids, 
and antioxidant capacity of red raspberries. Journal of 
Agricultural and Food Chemistry, 50, 5197-5201.
62. Mullen, W ., Y okota, T., Lean, M.E.J., & Crozier A. 
(2003). Analysis of ellagitannins and conjugates of 
ellagic acid and quercetin in raspberry fruits by LC-
MSn. Phytochemistry, 64(2), 617-624.
63. Nowicka, A., Kucharska, A.Z., Sokól-Łetowska, A., & 
Fecka, I. (2019). Comparicon of polyphenol content 
and antioxidant capacity of strawberry fruit from 90 
cultivars of Fragaria × ananassa Duch. Food Chemistry, 
270, 32-46.
64. Okuda, T., Y oshida, T., Hatano, T., Iwasaki, M., Kubo, 
M., Orime, T., Y oshizaki, M., & Naruhashi, N. (1992). 
Hydrolysable tannins as chemotaxonomic markers in 
the rosaceae. Phytochemistry, 32(9), 3091-3096.
65. Oliveira, M., Rodrigues, C.M., & Teixeira, P . (2019). 
Microbiological quality of raw berries and their 
products: A focus on foodborne pathogens. Heliyon, 5 
(12), e02992. 
66. Oszmiański, J., & Lachowicz S. (2016). Effect of the 
production of dried fruits and juice from chokeberry 
(Aronia melanocarpa L.) on the content and antioxidative 
activity of bioactive compounds. Molecules, 21, 1098.
67. Oszmiański, J., & Wojdylo, A. (2005). Aronia 
melanocarpa phenolics and their antioxidant activity. 
European Food Research and Technology, 221, 809-813.
68. Oszmiański, J., & Wojdylo, A. (2009). A comparative 
study of phenolic content and antioxidant activity of 
strawberry puree, clear, and cloudy juices. European 
Food Research and Technology, 228(4), 623-631.
69. Piccolo, E.L., Martìnez-Garcìa, L., Landi, M., Guidi, 
L., Massai, R., & Remorini, D. (2020). Influences of 
postharvest storage and processing techniques on 
antioxidant and nutraceutical properties of Rubus 
idaeus l.: A mini-review. Horticulturae, 6(4), 105.
70. Poyraz Engin, S., & Mert, C. (2020). The effects of 
harvesting time on the physicochemical components of 
aronia berry. Turkish Journal of Agriculture and Forestry, 
44(4), 361-370.
71. Prida, A., & Puech, J.-L. (2006). Influence of geographical 
origin and botanical species on the content of extractives 
in American, French, and East European oak woods. 
Journal of Agricultural and Food Chemistry, 54, 8115-
8126.
72. Prior, R.L., & Gu, L. (2005). Occurrence and biological 
significance of proanthocynidins in the American diet. 
Phytochemistry, 66, 2264-80.
73. Prior, R.L., Lazarus, S.A., Cao, G., Muccitelli, H., 
& Hammerstone, J.F. (2001). Identification of 
procyanidins and anthocyanins in blueberries and 
cranberries (Vaccinium spp.) using high-performance 
liquid chromatography/mass spectrometry. Journal of 
Agricultural and Food Chemistry, 49, 1270-1276.
74. Rodriguez-Mateos, A., Cifuentes-Gomez, T., George, 
T.W ., & Spencer, J.P .E. (2014). Impact of cooking, 
proving, and baking on the (poly)phenol content of wild 
bluberry. Journal of Agricultural and Food Chemistry, 
62, 3979-3986.
75. Rodríguez-Werner, M., Winterhalter, P ., & Esatbeyoglu, 
T. (2019). Phenolic composition, radical scavenging 
activity and an approach for authentication of Aronia 
melanocarpa berries, juice and pomace. Journal of Food 
Science, 84(7), 1791-1798.
76. Shukitt-Hale, B., Carey, A.N., Jenkins, D., Rabin, B.M., 
& Joseph, J.A. (2007). Beneficial effects of fruit extracts 
on neuronal function and behaviour in a rodent model 
of accelerated aging. Neurobiology of Aging, 28, 1187-
1194.
77. Sidor, A., Drożdżyńska, A., & Gramza-Michatowska, 
A. (2019). Black chokeberry (Aronia melanocarpa) and 
its products as potential health-promoting factors – An 
overview. Trends in Food Science & Technology, 89, 45-
60.
78. Sieniawska, E., & Baj, T. (2017). Tannins. In: 
S. Badal & R. Delgoda (Ed.), Pharmacognosy 
(Chapter 10) [E-reader version]. Retrieved from: 
https://www.sciencedirect.com/science/article/pii/
B978012802104000010X?via%3Dihub
79. Smeriglio, A., Barreca, D., Bellocco, E., & Trombetta, D. 
A Review of Tannins in Berries

35
(2017). Proanthocyanidins and hydrolysable tannins: 
occurrence, dietary intake and pharmacological effects. 
British Journal of Pharmacology, 174 (11), 1244-1262.
80.  Sójka, M., Macierzyński, J., Zaweracz, W ., & Buczek, 
M. (2016). Transfer and mass balance of ellagitannins, 
anthocyanins, flavan-3-ols and flavonols during the 
processing of red raspberries (Rubus idaeus L.) to juice. 
Journal of Agricultural and Food Chemistry, 64(27), 
5549-63.
81. Sparzak, B., Merino-Arevalo, M., Vander Heyden, Y ., 
Krauze-Baranowska, M., Majdan, M., Fecka, I., Gtód, 
D., & Bączek, T. (2010). HPLC analysis of polyphenols 
in the fruits of Rubus idaeus l. (Rosaceae). Natural 
Product Research, 24(19), 1811-1822.
82. Sueiro, L., Y ousef, G.G., Seigler, D., De Mejia, E.G., 
Grace, M.H., & Lila, M.A. (2006). Chemopreventive 
potential of flavonoid extracts from plantation-bred 
and wild Aronia melanocarpa (Black chokeberry) fruits. 
Journal of Food Science, 71, 480-488.
83. Swanson, B.G. (2003). Tannins and polyphenols. In: 
B. Caballero (Ed.), Encyclopedia of food sciences and 
nutrition (Second edition) [E-reader version]. (pp. 5729-
5733). Retrieved from: https://www.sciencedirect.com/
sdfe/pdf/download/eid/3-s2.0-B012227055X011780/
first-page-pdf
84. Taheri, R., Connolly, B.A., Brand, M.H., & Bolling, B.W . 
(2013). Underutilized chokeberry (Aronia melanocarpa, 
Aronia arbutifolia, Aronia prunifolia) accessions are rich 
sources of anthocyanins, flavonoids, hydroxycinnamic 
acids, and proanthocyanidins. Journal of Agricultural 
and Food Chemistry, 61, 8581-8588.
85. Teleszko, M., & Wojdyto, A. (2015). Comparison 
of phenolic compounds and antioxidant potential 
between selected edible fruits and their leaves. Journal 
of Functional Foods, 14, 736-746.
86. Tϋrkben, C., Sariburun, E., Demir, C., & Uylaşer, V . 
(2010). Effect of freezing and frozen storageon phenolic 
compounds of raspberry and blackberry cultivars. Food 
Analytical Methods, 3, 144-153.
87. Uleberg, E., Rohloff, J., Jaakola, L., Trôst, K., Junttila, 
O., Häggman, H., & Martinussen, I. (2012). Effects of 
temperature and photoperiod on yield and chemical 
composition of northern and southern clones of bilberry 
(Vaccinium myrtillus L.). Journal of Agricultural and 
Food Chemistry, 60(42), 10406-10414.
88. Vrhovsek, U., Giongo, L., Mattivi, F., & Viola, R. (2008). 
A survey of ellagitannin content in raspberry and 
blackberry cultivars grown in Trentino (Italy). European 
Food Research and Technology, 226, 817-824.
89. Vrhovsek, U., Palchetti, A., Reniero, F., Guillou, C., 
Maduero, D., & Mattifi, F. (2006). Concentration and 
mean degree of polymerization of Rubus ellagitannins 
evaluated by optimized acid methanolysis. Journal of 
Agricultural and Food Chemistry, 54, 4469-4475.
90. Wang, S.Y ., Chen, C.T., & Wang, C.Y . (2009). The 
influence of light and maturity on fruit quality and 
flavonoid content of red raspberries. Food Chemistry, 
112(3), 676-684.
91. Wangensteen, H., Bräunlich, M., Nikolic, V ., Malterud 
K.E., Simestad, R., & Barsett, H. (2014). Anthocyanins, 
proanthocyanidins and total phenolics in four cultivars 
of aronia: Antioxidant and enzyme inhibitory effects. 
Journal of Functional Foods, 7(1), 746-752. 
92. Wilkes, K., Howard, L.R., Brownmiller, C., & Prior, R.L. 
(2014). Changes in chokeberry (Aronia melanocarpa 
L.) polyphenols during juice processing and storage. 
Journal of Agricultural and Food Chemistry, 62, 4018-
4025.
93. Wu, X., Gu, L., Prior, R.L., & McKay, S. (2004). 
Characterization of anthocyanins and proanthocyanidins 
in some cultivars of Ribes, Aronia and Sambucus and 
their antioxidant capacity. Journal of Agricultural and 
Food Chemistry, 52(26), 7846-7856.
94. Zafrilla, P ., Ferreres, F., & Tomás-Barberán, F.A. (2001). 
Effect of processing and storage on the antioxidant 
ellagic acid derivatives and flavonoids of red raspberry 
(Rubus idaeus) jams. Journal of Agricultural and Food 
Chemistry, 49(8), 3651-3655.
95. Zoratti, L., Karppinen, K., Escobar, A.L., Häggman, 
H., & Jaakola, L. (2014). Light-controlled flavonoid 
biosynthesis in fruits. Frontiers in Plant Science, 5, 534.
A Review of Tannins in Berries

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Tanini v jagodičevju – pregled
IZVLEČEK
Tanini so skupina polifenolnih spojin, ki jih kot sekundarne metabolite sintetizirajo in akumulirajo višje rastline. Delimo jih 
na hidrolizabilne tanine in proantocianidine. Prisotni so v številnih rastlinskih tkivih v obliki različnih struktur in v različnih 
količinah. Pregledni članek ponuja kratek pregled o porazdelitvi taninov v rastlinah in povzema trenutno znane izsledke o taninih 
v jagodah, malinah, borovnicah, ki so najpogosteje gojeno in konzumirano jagodičevje ter aroniji, ki je postala priljubljena v 
zadnjih desetletjih. Povzeti so tudi učinki predelave in skladiščenja na taninsko sestavo in njihova vsebnost v izdelkih.
Ključne besede: elagitanini, proantocianidini, jagode, maline, borovnice, aronija
A Review of Tannins in Berries