© CAB International 2017. Blackberries (eds H.K. Hall and R.C. Funt) 49
* Corresponding author: Jungmin.Lee@ars.usda.gov
Small-fruit quality is closely related to its production of primary and secondary
metabolites (i.e. type, concentration). Blackberry metabolites continue to
undergo anabolism and/or catabolism within the fruit until harvest, and these
components that remain, and are not degraded by the time the fruit (or fruit
product) reaches consumers, determine blackberry fruit quality. The primary
and secondary metabolite composition of a blackberry deﬁnes its characteris-
tic appearance, taste, and texture. Blackberries are typically purchased for con-
sumption as fresh, individually quick frozen (IQF), or as a further processed
product incorporating them into jams, syrups, wines, teas, juices, concen-
trates, and purees. Many potential health beneﬁts from consuming blackber-
ries or blackberry products are attributed to their metabolites. Metabolites also
directly and indirectly inﬂuence processing regimes, shelf life, and consumer
Blackberries contain dietary ﬁber, vitamin C (ascorbic acid), vitamin A,
vitamin E, potassium, and calcium (for additional nutrition facts, see USDA,
2015), along with the phenolic metabolites that are a source of possible health
beneﬁts. Sensory attributes, typically used to describe the taste and ﬂavor of
blackberries, include fresh fruit, cooked fruit, cooked berry, strawberry, rasp-
berry, vegetal, stemmy, and earthy (Du et al., 2010). In this chapter, a sum-
mary of distinct primary and secondary metabolites crucial to blackberry
quality will be presented focusing on fruit, although all parts of the plant
(leaves, canes, and roots) have historically been used as foods or herbal reme-
dies (Arnason et al., 1981; Hummer, 2010). The concentration ranges for
compounds related to blackberry quality are summarized in Table 4.1.
BlackBerry fruiT qualiTy comPonenTs,
comPosiTion, and PoTenTial healTh BenefiTs
USDA-ARS-HCRU Worksite, Parma, Idaho, USA
50 Jungmin Lee
Table 4.1. Blackberry and blackberry hybrid fruit quality components and
reported concentration ranges (in fresh weight).
(n = sample size) References
Fruit mass 3.8–28.3 g
(n = 19)
Finn et al., 2014; Vrhovsek et al., 2008;
H.K. Hall, personal observation
Calories 43–64 kcal/100 g
(n = 3)
% soluble solids 6.9–16.8
(n = 90)
Fan-Chiang and Wrolstad, 2010; Finn
et al., 2014; Mertz et al., 2007; Thomas
et al., 2005; Vrhovsek et al., 2008;
Wang et al., 2008
Titratable acidity 0.08–2.7 g/100 g
(n = 82)
Fan-Chiang and Wrolstad, 2010; Finn
et al., 2014; Mertz et al., 2007; Thomas
et al., 2005; Veberic et al., 2014; Wang
et al., 2008
(n = 73)
Fan-Chiang and Wrolstad, 2010; Finn
et al., 2014; Mertz et al., 2007; Thomas
et al., 2005; Veberic et al., 2014
Simple sugars 2.6–13.9 g/100 g
(n = 63)
Fan-Chiang and Wrolstad, 2010; Mikulic-
Petkovsek et al., 2012; Veberic et al.,
Organic acids* 0.5–2.9 g/100 g
or (n = 73)
Fan-Chiang and Wrolstad, 2010; Mikulic-
Petkovsek et al., 2012; Vrhovsek et al.,
Vitamin C (ascorbic
1.2–11.9 mg/100 g
(n = 12)
Thomas et al., 2005; Veberic et al., 2014
Anthocyanins 28–366 mg/100 g
(n = 1,306)
Conner et al., 2005; Fan-Chiang and
Wrolstad, 2005; Finn et al., 2014;
Scalzo et al., 2008; Sellappan et al.,
2002; Vasco et al., 2009; Veberic et al.,
2014; Wang et al., 2008
Phenolic monomers 0.7–555 mg/100 g
(n = 22)
Acosta-Montoya et al., 2010; Bilyk and
Sapers, 1986; Gancel et al., 2011;
Sellappan et al., 2002; Vasco et al.,
2009; Veberic et al., 2014
17–27 mg/100 g
(n = 5)
Gasperotti et al., 2010
85–390 mg/100 g
(n = 23)
Gancel et al., 2011; Gasperotti et al.,
2010; Vasco et al., 2009; Vrhovsek
et al., 2008
Carotenoids 0.44–0.59 mg/100 g
(n = 2)
Curl, 1964; Marinova and Ribarova,
*Excluding ascorbic acid, listed separately.
Quality, Composition, and Health Beneﬁts 51
SUGARS AND ORGANIC ACIDS
Two simple quality measurements of blackberries and their hybrids assess the
most inﬂuential categories of their taste perception: sugar and organic acid
content. Typically sweetness is described as percent (%) soluble solids, while
acidity is reported as titratable acidity (for concentration ranges, see Table 4.1).
Sugars reported in blackberry fruits are fructose, glucose, sucrose, and occa-
sionally exceedingly low levels of sorbitol (Fan-Chiang and Wrolstad, 2010;
Lee, 2015; Mikulic-Petkovsek et al., 2012; Wrolstad et al., 1980, 1981). Min-
ute concentrations of sorbitol (sugar alcohol) in processed blackberry products
(e.g. juice) likely originated from processing enzymes or immature (under ripe)
fruit (Fan-Chiang and Wrolstad, 2010; Lee, 2015). Since sorbitol is seldom
found in ripe blackberries (Lee, 2015), the detection of sugar alcohol may be
an indicator of accidental or fraudulent adulteration of products with cheaper
fruit (i.e. apples, pears) juices or concentrates (Lee, 2015; Lee et al., 2012;
Wrolstad et al., 1981). Lee (2015) clariﬁed that recent United States media
claims of blackberries containing high levels of sugar alcohol were inaccurate
and actually the opposite of scientiﬁc ﬁndings.
Blackberry tartness is due to nonvolatile organic acids, including ascorbic
acid (vitamin C), citric acid, isocitric acid, lactoisocitric acid, malic acid, shi-
kimic acid, fumaric acid, and succinic acid (Fan-Chiang and Wrolstad, 2010;
Mikulic-Petkovsek et al., 2012; Veberic et al., 2014; Vrhovsek et al., 2008).
Lactoisocitric acid can be a useful organic acid marker for blackberries, with
the caveat that the amounts in blackberry hybrids (e.g. ‘Loganberry,’ ‘Boysen-
berry’) may be too low to act as an effective indicator. Two distinctive patterns
were observed in the acid makeup of blackberry samples examined by Fan-
Chiang and Wrolstad (2010). The samples’ organic acid proﬁles resembled
either that of ‘Marion’ (higher citric acid levels) or of ‘Evergreen’ (higher isoc-
itric acid levels), suggesting this distinguishing factor might allow the identiﬁ-
cation of cultivars used in commercial blackberry products.
Famiani and Walker (2009) investigated changes in blackberry primary
metabolites during fruit ripening, where they found soluble solids increased
while titratable acidity decreased during the ﬁnal growth stage prior to har-
vest. Ratios of sugars to acids will not be discussed here, since ratios are mis-
leading in terms of apparent ﬂavor; equivalent sugar–acid ratios do not equal
similar taste assessments.
Blackberry fruit phenolics have been thoroughly reviewed by Lee et al. (2012)
and Kaume et al. (2012). Unlike other dark-colored Rubus fruit (i.e. black
raspberry and red raspberry), blackberry pigments are chieﬂy cyanidin-based
anthocyanins (Lee et al., 2012). Though the characteristic black color of intact
52 Jungmin Lee
fresh blackberry fruit is actually from its concentration and types of anthocya-
nins (natural red pigments). Since anthocyanin color is pH dependent, color
linked to its structural form that undergoes transformation with changes in
pH, a slight change in pH makes the red anthocyanin within blackberries turn
a deep purple to black color. However, there are some rare blackberries lacking
anthocyanins, including ‘Snowbank’ (R. allegheniensis; Hummer et al., 2015)
and ‘Clark Gold’ (R. trivalis; US PP14935 P2). While these uncommon fruits
are white to yellow, consumers gravitate towards more commercially available
dark blackberries thought to have high pigment concentrations. Blackberry
anthocyanin levels (see Table 4.1) are actually in the lower ranges of what can
be found in black raspberries (anthocyanin levels ranging from 39 to
996 mg/100ml, n > 1,000; Dossett et al., 2012), or blueberries (anthocyanin
levels ranging from 101 to 400 mg/100 g, n = 37; Lee et al., 2004), but higher
than red raspberries (anthocyanin levels ranging from 6 to 98 mg/100 g,
n = 644; Scalzo et al., 2008).
Blackberries contain acylated and non-acylated anthocyanins, but most
are the non-acylated form. The major anthocyanins that have been reported
are cyanidin-glucoside, cyanidin-rutinoside, cyanidin-xyloside, cyanidin-
malonylglucoside, and cyanidin-dioxalylglucoside (or possibly cyanidin-
hydroxymethylglutaroylglucoside) (Conner et al., 2005; Fan-Chiang and
Wrolstad, 2005; Finn et al., 2014; Jordheim et al., 2011; Lee et al., 2012;
Stintzing et al., 2002a; Veberic et al., 2014, 2015). While the most predomi-
nate pigment in blackberries is cyanidin-glucoside (44–95% of total), the ratios
of subsequent anthocyanins vary with cultivar and genotype. Blackberry
hybrids contain different anthocyanin proﬁles compared to non-hybrids. For
example, hybrids of raspberry and blackberry (i.e. ‘Boysenberry’ and ‘Logan-
berry’) contain cyanidin-sophoroside, as found in red raspberry, but not in any
non-hybrid blackberry (Fan-Chiang and Wrolstad, 2005; Lee et al., 2012).
Some blackberries including ‘Marion,’ ‘Waldo,’ ‘Evergreen,’ ‘Black
Douglass,’ ‘Hull Thornless,’ ‘Chester Thornless,’ and ‘Shawnee,’ contain cyan-
idin-dioxalylglucoside (Fan-Chiang and Wrolstad, 2005; Kolniak-Ostek et al.,
2015; Stintzing et al., 2002a). This identiﬁcation may be, at least
partially, disputed, as an independent group has claimed that the
accepted identity of cyanidin- dioxalylglucoside is actually cyanidin-
hydroxymethylglutaroylglucoside (unconﬁrmed; Jordheim et al., 2011). It
should be noted that while both of those anthocyanins are unique to black-
berries (Fan-Chiang and Wrolstad, 2005; Jordheim et al., 2011; Veberic et al.,
2014), they are not necessarily found in all varieties, and neither was detected
in the new ‘Columbia Star’ (Finn et al., 2014).
The prevailing minor anthocyanin in many blackberries is pelargonidin-
glucoside, as Veberic et al. (2014) found in the cultivars ‘Black Satin,’ ‘ C
Bestrna,’ ‘Chester Thornless,’ ‘Thornless Evergreen,’ ‘Loch Ness,’ and ‘Thorn-
free.’ Although one previous study did not ﬁnd pelargonidin-glucoside when
identical cultivars were tested (Fan-Chiang and Wrolstad, 2005).
Quality, Composition, and Health Beneﬁts 53
Blackberry anthocyanins increase in concentration with fruit maturity
(Acosta-Montoya et al., 2010; Famiani and Walker, 2009). While darker fruit
is an indication of ripeness, post-processing modiﬁcations in appearance are
expected. Freezing, thawing, or storage induced visual color changes of dark
black to hues of red, yellow, or blue are due to slight alterations in pH and deg-
radation of ascorbic acid, anthocyanins, etc. (Stintzing et al., 2002b; Veberic
et al., 2014). Anthocyanin-based color is also affected by the physical form of
water (liquid versus ice) and the chemical state of the fruit itself.
Anthocyanin proﬁles, as well as the presence of sorbitol, mentioned ear-
lier, can point to adulteration in blackberry and non-blackberry based prod-
ucts (Lee, 2015; Lee et al., 2012; Wrolstad et al., 1981). For instance, a black
raspberry freeze-dried powder (~$19 per 100 g), sold as a dietary supplement,
was found to actually contain blackberry powder (~$14 per 100 g); Lee (2014)
conﬁrmed this with repeated purchases over time, and by analyzing blackberry
and black raspberry powdered products from the same anonymous vendor.
Although the fruit powders are similar visually, they have distinct anthocyanin
proﬁles (Lee et al., 2012).
NON-ANTHOCYANIN PHENOLIC MONOMERS
The non-anthocyanin phenolic monomers (range shown in Table 4.1) found in
blackberry fruit are the phenolic acids: ellagic acid, gallic acid, p-coumaric acid
esters, caffeic acid, caffeic acid esters (like neochlorogenic acid), ferulic acid,
ferulic acid esters; the ﬂavanols: catechin and epicatechin; and the ﬂavonol-
glycosides: quercetin-, kaempferol-, isorhamnetin-, and myricetin-glycosides
(Bilyk and Sapers, 1986; Kolniak-Ostek et al., 2015; Lee et al., 2012; Mertz
et al., 2007; Sellappan et al., 2002; Veberic et al., 2014). Acylated ﬂavonol-
glycosides have also been reported in blackberries (Veberic et al., 2014). A
more detailed list of blackberry phenolic monomers can be found in Lee et al.
(2012), but additional work is needed to clarify these phenolic classes in
While ellagic acid is the main phenolic acid seen in blackberries, it is a
challenging compound to analyze, since it has poor solubility in water;
although improved in alcohol, its solubility is enhanced best by increasing
solution pH well above what is normally found in foods (Bala et al., 2006). At
least one study reported ﬂavonol-glycoside levels decreased with fruit ripening
(Acosta-Montoya et al., 2010).
Blackberries, and other Rubus fruit (i.e. red raspberries, cloudberries), are a
rich source of ellagitannins (also known as hydrolyzable tannins, and distinct
54 Jungmin Lee
from the more extensively studied condensed tannins). Red raspberry ellagi-
tannin concentrations, at 94–172 mg/100 g, were found lower than for black-
berries (for range, see Table 4.1), but within the same study their single black
raspberry sample tested in the blackberry range at 330 mg/100 g (Vrhovsek
et al., 2008). The main intact (non-hydrolyzed) blackberry ellagitannins have
been recognized as lambertianin C and sanguiin H-6 (Acosta-Montoya et al.,
2010; Gancel et al., 2011; Gasperotti et al., 2010; Mertz et al., 2007;
Sangiovanni et al., 2013). Kool et al. (2010) did not ﬁnd lambertianin C in
their ‘Boysenberry’ samples, but found sanguiin H-6 as the primary ellagitan-
nin, along with three other supplementary ellagitannins.
Some researchers have broken down (hydrolyzed) ellagitannins before
analysis and reported the ellagitannin subunits as methyl gallate, ellagic acid
derivative, ellagic acid, and methyl sanguisorboate, with mean degree of
polymerization (indication of size) ranging from 1.59 to 1.92 (Mertz et al.,
2007; Vrhovsek et al., 2006, 2008). This points to ellagitannin values deter-
mined by hydrolysis prior to high performance liquid chromatography (HPLC)
separation as offering the closest probable approximation of ellagitannin
concentrations within unprocessed fruit. These remain a difﬁcult group of
compounds to extract, separate, and identify. Beside conventional challenges
in investigating a naturally complex class of compounds, work with them is
further hampered by the lack of available pure commercial standards, no con-
ﬁrmed identiﬁcations as of 2016, and problems keeping these compound in
their native states (Acosta et al., 2014; Acosta-Montoya et al., 2010; Aripitsas,
2012; Gasperotti et al., 2010; Lee et al., 2012; Sangiovanni et al., 2013;
Vrhovsek et al., 2006, 2008).
Ellagitannins are found in all fractions of Rubus fruit, but the highest con-
centrations are in seed fractions (Hager et al., 2008). Ellagitannin levels can
decrease during the fruit-ripening period from red to fully ripe (Acosta- Montoya
et al., 2010). They also are reduced during food processing by high tempera-
ture degradation (e.g. pasteurization), precipitation out of solution, and
hydrolysis to ellagic acid (Gancel et al., 2011). Some reports have likely under-
estimated ellagitannins by incomplete extraction due to inappropriate solvents
and non-optimized extraction techniques (Lee et al., 2012; Lei et al., 2001).
Ellagitannins play several roles within plants, including protecting them
from pathogen attack and inhibiting premature seed germination (Lee et al.,
2012; Lei et al., 2001). To humans, these are the compounds seen as sediments
during wine and juice processing, and they can contribute to turbidity issues
for food products requiring clarity as a quality assessment (Lee et al., 2012).
However, these same sediments (i.e. ellagic acid and ellagitannin) from post-
processing waste are also potential future ingredients for value-added products
(Acosta et al., 2014). No work has yet been published on blackberry ellagitan-
nins’ taste, but work done with wood tannins (same phenolic class as found in
blackberries, but different types of ellagitannins) shows their contributions
range from no detectable ﬂavor to added bitterness and astringency, with
Quality, Composition, and Health Beneﬁts 55
threshold concentration dependent to each speciﬁc compound examined
( Glabasnia and Hofmann, 2006). Additional work is needed for identiﬁcation,
quantiﬁcation, and sensory evaluation of this phenolic class from
ADDITIONAL QUALITY COMPONENTS
Other quality constituents of blackberries are carotenoids, vitamins, minerals,
proteins, ﬁber, and aroma compounds. Aroma compounds reported in black-
berries are esters (ethylacetate), aliphatic alcohols (heptanol, hexenol, hex-
anol, and octanol), terpenes (carveol), aldehydes (hexanal, hexenal, and
benzaldehyde), and ketones (heptanone) were found in R. ulmifolius Schott
(D’Agostino et al., 2015; Perez-Gallardo et al., 2015). Blackberry fruit carote-
noids are lutein, b-carotene, zeaxanthin, and b-cryptoxanthin (Marinova and
Ribarova, 2007). Additional work needs to be conducted on the quality of
components listed in this section to provide a better understanding of similari-
ties and differences among cultivars, ﬁeld treatments, etc. Novelty products
made from food processing by-products, such as juice presscake and black-
berry seed oils, are currently available and are a source of vitamin E (Bushman
et al., 2004; Van Hoed et al., 2011).
POTENTIAL HEALTH BENEFITS
Besides the ﬂavor and color blackberries provide to foods, their naturally high
level of phenolics could have potential health beneﬁts that may help protect
their consumers from some chronic diseases. Blackberry fruit phenolics have
been implicated in providing anticancer, antiproliferative, antineurodegenera-
tive, anti-inﬂammatory, antidiarrheal, antidiabetic, antimicrobial, and antivi-
ral activities (Bakkalbasi et al., 2009; Landete, 2011, 2012; Lee et al., 2012;
Pojer et al., 2013). Phenolics bioavailability, metabolism, and potential health
beneﬁts have been previously well reviewed (Bakkalbasi et al., 2009; Landete,
2011, 2012; Pojer et al., 2013), although the exact mechanisms of how black-
berries may impart protection after ingestion remain unclear. It is clear that
diets rich in fruits and vegetables are valuable in preventing some cancers and
reducing the risk of cardiovascular disease (Basu et al., 2010; Van Duyn and
Pivonka, 2000; Pojer et al., 2013), and blackberries can be an element of that
Blackberry fruit anthocyanins are found in their native forms at low con-
centrations after digestion; the majority are found as protocatechuic acid and
its derivatives (i.e. ferulic acid, hippuric acid, vanillic acid), phenylacetic acid,
phenylpropenoic acid, methylated conjugates, glucuronidated conjugates, and
many other metabolites (Czank et al., 2013; de Ferrars et al., 2014; Felgines
56 Jungmin Lee
et al., 2005; Pojer et al., 2013). Fang (2014) reviewed the absorption route of
cyanidin-glucoside, the chief blackberry anthocyanin. Czank et al. (2013)
demonstrated isotopically labeled cyanidin-glucoside (500 mg) remained in
circulation in male subjects for over 48 hours. Cyanidin-glucoside has been
shown to inhibit proliferation of human lung carcinoma cells, and cancer cell
migration in mice (Ding et al., 2006). Blackberries have also been linked to
providing neuroprotective effects in human neuroblastoma (extracranial solid
cancer) cells (Tavares et al., 2013).
As with the previously mentioned challenges to analyzing blackberry phe-
nolic polymers (i.e. ellagitannins), their size and poor solubility limit their bio-
availability as well (Garcia-Munoz and Vaillant, 2014). Ellagitannins’ health
beneﬁcial effects were well summarized recently (Landete, 2011; Garcia-
Munoz et al., 2014). A post-ingestion assessment of blackberry ellagitannins
in human urine found that the metabolites had been converted into urolithins
by gut microbiota (Garcia-Munoz et al., 2014). Urolithins are associated with
preventing or controlling colon, breast, esophageal, and prostate cancers
( Garcia-Munoz and Vaillant, 2014; Landete, 2011). As each of us has hetero-
geneous gut microbiota, some researchers have classiﬁed individuals into
urolithin A, urolithin B, or non-urolithin (unidentiﬁed metabolites) excreters
(Tomas-Barberan et al., 2014). A review of a variety of gut microbiota metabo-
lizing assorted classes of phenolics was well summarized by Selma et al. (2009).
Additional data on this topic will become available as the number of identiﬁed
gut bacteria grows, and they come to be further studied.
The exact mechanism of how blackberry dietary phenolics beneﬁt human
health is not fully elucidated and more work needs to be conducted to clarify
this. While phenolics are attributed in disease prevention, they are also anti-
nutritive and hinder absorption of certain minerals and proteins (Landete,
2012). Human clinical studies on blackberries alone, not mixed berries, are
limited. A USA human clinical study on the inﬂuence of blackberries on can-
cer processes has been completed, but results are not yet available (US clinical
trial identiﬁer NCT01293617). Additional work is needed to clarify the contra-
dicting reports among in vitro and in vivo work, animal models versus human
trials, length of intervention, types of cells used, etc. (Garcia-Munoz and
Vaillant, 2014; Landete, 2011, 2012).
Antioxidant claims have been deemed scientiﬁcally uncorroborated
( Hollman et al., 2011), and will not be discussed due to the controversial
limitations surrounding in vitro methods (Carocho and Ferreira, 2013; Frankel
and Meyer, 2000). Numerous studies and reviews are available on the lack
of evidence for a relationship between antioxidant activity and human health
(Hollman et al., 2011; Lee et al., 2012). In 2010, the European Food Safety
Authority (EFSA), analogous in the European Union (EU) to the US Food and
Drug Administration (FDA), rejected a petition for labeling food packages with
health claims related to antioxidant activity, citing the lack of scientiﬁc data
from human trials to substantiate such a claim (Gilsenan, 2011).
Quality, Composition, and Health Beneﬁts 57
As many become convinced that blackberries’ quality compounds offer desir-
able potential health beneﬁts, efforts have been conducted to further enhance
the dietary phenolic content of blackberry fruit. Various techniques, including
stimulation of metabolite production with ﬁeld management factors via biotic
elicitors (e.g. Pseudomonas ﬂuorescens) and plant hormones (e.g. methyl
jasmonate), have been attempted (Garcia-Seco et al., 2013; Ramos-Solano
et al., 2014, 2015; Wang et al., 2008). Nutrient regimes have also been shown
to alter blackberry phenolics (Ali et al., 2011). The high perishability of fruit
sold in the fresh market has also created a demand to prolong blackberry shelf
life; positive reports have used calcium in combination with pectin spray and
starch-beeswax coating to prolong the shelf life (Perez-Gallardo et al., 2015;
Sousa et al., 2007).
Growers, processors, and consumers should remain conscious that the
quality components discussed here vary with a plant’s genus, species, cultivar/
genotype, and age; along with environment and management practices such
as growing region and conditions, harvest decisions, fruit maturity indices,
processing methods, and storage. Beside their dietary phenolic content, black-
berries also have other important contributors to nutrition, including vitamin
C, vitamin A, vitamin E, vitamin B6, folic acid, dietary ﬁber, potassium, phos-
phorus, magnesium, calcium, and iron.
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