Agricultural and Food Science, Vol. 19 (2010): 69-80


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Vol. 19(2010): 69–80.

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© Agricultural and Food Science 
Manuscript received: October 2008 

Effect of substrate type on the field performance 
and chemical composition of highbush  

blueberry cv. Patriot

Ireneusz Ochmian1, Józef Grajkowski1 and Katarzyna Skupień2*
1Department of Pomology, West Pomeranian University of Technology, Szczecin ul. Janosika 8, 

71-432 Szczecin, Poland 
2Department of Plant Raw Materials Processing and Storage, West Pomeranian University of Technology, 

Szczecin, ul. Słowackiego 17, 71-434 Szczecin, Poland, 
*e-mail: kskupien@agro.ar.szczecin.pl

The aim of a 3-year study was to compare the effect of substrate type (peat, sawdust and cocoa husk) on 
the yield, fruit quality, general and phenolic composition, and antioxidant activity of highbush blueberry 
(Vaccinium corymbosum L.) cultivar Patriot. The largest berries and the highest yield were obtained from 
bushes grown in sawdust (3.661 kg ha-1). Berries from bushes grown in cocoa husk had the highest content 
of N (14.25 g kg-1), P (1.17 g kg-1), K (6.51 g kg-1), and Zn (8.78 mg kg-1), as well as P (1.26 g kg-1) and K 
(6.99 g kg-1) in the leaves. Both the leaves and fruits of plants grown in sawdust had higher Cu concentra-
tions (3.89 and 3.01 mg kg-1). Berries from sawdust-grown bushes had the highest soluble solids (13.1%) 
and total sugar (10.80 g 100 g-1) content. The fruits obtained from peat-grown bushes exhibited enhanced 
antioxidant capacity (38.6 µmol Trolox g-1). However, the highest levels of phenolics were noted in berries 
produced in the cocoa husk (208.29 mg 100 g-1) and peat-bed system (174.07 mg 100 g-1). Regardless of the 
type of substrate used, the following phenols were identified (in descending order): anthocyanins>chlorogenic 
acid>flavonols. Delphinidin-glycosides were the most abundant pigments, and cyanidin, peonidin, petunidin, 
and malvidin-glycosides were also detected. Total flavonol content (quercetin derivatives and kaempferol 
3-rutinoside) was not significantly affected by the type of substrate. 

Key- words: Vaccinium corymbosum, yield, firmness, macro- and microelements, phenolics 



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Introduction

Highbush blueberries (Vaccinium corymbosum 
L.), among other members of the Ericaceae fam-
ily, are considered to be a rich source of phenolic 
compounds and are known for their antioxidant 
properties (Kalt et al., 1999, Zheng and Wang, 
2003, Castrejón et al. 2008). Due to their chemical 
composition, blueberries are highlighted as one of 
the healthiest foodstuffs (Kader et al., 1996, Main-
land et al. 2002). Because of its value, the species 
has been cultivated for years in North America. 
The first breeding project was launched in 1908 in 
Florida (Lyrene 1997), and soon afterwards high-
bush blueberries were imported from the USA to 
Europe (Strik 2005).

In Poland, the first experiments involving 
highbush blueberry cultivation were undertaken 
in 1946, while the development of large-scale pro-
duction methods began in 1976. According to the 
results of field trials, Patriot was one of the culti-
vars recommended for growing in Poland (Smo-
larz 2006). A considerable increase in blueberry 
production has been recently observed in Poland. 
In 2006 Poland took 1st place in Europe with re-
gard to the highbush blueberry growing area, and 
blueberry production reached 8,000 t, which gave 
us the second place after Germany (10,000 t) (In-
ternetowa giełda rolna fresh-market 2009).

The highbush blueberry requires specific grow-
ing conditions including the right site, high in or-
ganic matter, appropriate water and air drainage, 
low pH, and adequate moisture (Kozinski 2006). 
On the one hand, this species allows the use of 
land which as a result of its high acidity is not suit-
able for other crops (Ciordia et al. 2006). On the 
other hand, such soils are less and less available, 
which hinders the establishment of new planta-
tions. These factors have necessitated the search 
for and the development of modified soil systems.

An improvement in soil conditions can be 
achieved by using organic mulch materials, like 
peat with low pH, sawdust, or coniferous bark. The 
application of mulches contributes to the loosen-
ing of the higher strata of soil thus improving root 
penetration (Kozinski 2006). In the USA, cotton 

by-products, pecan hulls (Krewer et al. 2002), leaf-
mould compost, pit-coal ash, sludge from sewage 
treatment plants (Black et al. 2002), and conifer 
needle litter (Entrop 2000) are used for mulching 
purposes. From the practical and economic point 
of view, the components used for substrates and 
mulching should be relatively cheap, easily acces-
sible, and should meet the habitat requirements of 
a given species. Therefore, the use of agricultural 
and forest by-products for this purpose seems to 
be fully justified. 

The objective of the present study was to evalu-
ate the influence of three types of substrate (cocoa 
husk, sawdust and peat) on the yield, fruit quality 
(fruit size and firmness), bush growth rate, fruit 
chemical composition (the content of soluble sol-
ids, total sugar, L-ascorbic acid, NO3

- , NO2
-, phe-

nolics, macro- and microelements, titratable acid-
ity, juice efficiency, antioxidant activity), macro- 
and microelement concentrations in the leaves of 
highbush blueberry cv. Patriot.

Materials and methods 

The experiment was carried out in the spring of 
2001 at the Experimental Station Rajkowo near 
Szczecin in heavy soil of neutral reaction (pH 7.1). 
The purpose of field trials was to evaluate highbush 
blueberry cultivation in three different substrates 
placed in the trenches, 35 cm deep and 100 cm wide 
(0.35 m3 medium per m of row length). Acidic muck 
soil (peat), conifer sawdust from a local sawmill, 
and cocoa husk – a by-product obtained from the 
Chocolate Confectionery Plant ‘Gryf’ in Szczecin 
were used as substrates. In vitro potted plants of 
highbush blueberry cv. Patriot were spaced 1.5 m 
apart in a row and 2.5 m between the rows. The 
experiment was conducted in 2005–2007. The 
physical properties of the substrates tested in the 
experiment are shown in Table 1. Soil samples were 
taken from each replicate of substrate and from the 
soil, with a sample stick, in mid-August. Among the 
substrates, peat had the highest and sawdust had 
the lowest field water capacity (44.8% and 31.3% 



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respectively), while cocoa husk had the highest full 
water capacity (85.3%). These differences neces-
sitated the use of varied watering rates.

Fertilization was limited to nitrogen supply 
only, because chemical analyses of both the soil 
and substrates showed a high and/or a medium 
content of other nutrients. Each type of media was 
fertilized with ammonium nitrate on three occa-

sions at a total dose of 30 kg N ha-1. Supplemen-
tal irrigation with acidified water (with H2SO4 up 
to pH of 2.5–3.5 measured in H2O) was applied 
through a T-Tape drip line. The rate of water sup-
ply was adjusted to soil moisture content based on 
the results of tensiometric monitoring expressed 
in pF units (pF – soil suction being the logarithm 
of water height in centimeters), carried out twice 
a week. Measuring tubes (30 cm) were installed 
15 cm below the soil surface, and pF 2.2 was 
adopted as a threshold value for irrigation. Hav-
ing reached the threshold, the soil was irrigated 
to approximately pF 1.0. Due to the wide range 
of substrate reaction, water used for irrigation was 
acidified to a different degree. Water used for peat 
irrigation had higher pH (3.72) because peat reac-
tion was suitable for blueberry cultivation, while 
cocoa husk and sawdust beds were irrigated with 
water with pH of 2.36 to lower the reaction of these 
media (Table 2). Among the substrates tested, peat 
maintained a constant pH level of 3.3–3.5 during 
the field trials, while cocoa husk had the highest 
reaction (pH of 4.6–6.4) (data not shown). pH and 
EC were measured with a multi-function computer 
device CX-741 Elmetron (Zabrze, Poland).

The content of available Ca, Na, Fe, Mn, Zn 
and Cu in the soil and in the substrates was de-
termined by the AAS method (Atomic Absorption 
Spectrometry) prior to sample extraction with 0.1 

Table 1. Water capacity, pH and salinity of the sub-
strates used for blueberry cultivation

Peat Cocoa husk Sawdust
Field water capacitya 
(% vv-1)

44.8 36.9 31.3

Full water capacitya 
(% vv-1)

80.6 85.3 82.6

pHb 3.40 5.00 4.70

Soil salinity  
(g NaCl kg-1)c

0.87 0.35 0.56

a Water capacity of soil was measured with the gravimetric 
method in spring 2005.
bThe reaction of substrates was measured with the conductomet-
ric method in KCl at the end of highbush blueberry vegetative 
season. pH values are the mean for 2005-2007.
cSoil salinity was measured with the conductometric method 
and the values are the mean for 2005-2007.

Table 2. Physicochemical properties of water used in the experiment (on average for 2005–2007).

Raw water*
Acidified water for sawdust 
and cocoa husk*

Acidified water to irrigate peat*

Fe+3 (mg·l-1) Ca+2 (mg·l-1) EC (mS·cm-1) pH
EC  

(mS·cm-1)
pH

EC 
(mS·cm-1)

pH

0.17 94.0 0.80 7.01 2.46 2.36 2.01 3.72
* The values are the mean of 2005-2007. The measurements were carried out three times in each year of experiment. Fe3+and Ca2+ 
were determined by the AAS method (Atomic Absorption Spectrometry), and EC (Electrical Conductance) and pH were measured with 
a conductometric method.

Table 3. The content of available macro- and microelements in the grey brown podsolic soil used in the experiment (an 
average for 2005–2007).

P K Ca Mg Na Fe Mn Zn Cu

(mg·100g-1) (mg·kg-1)

12.41 35.6 70.6 5.67 1.25 102.8 66.5 17.9 2.7



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M HCl. Soil samples for phosphorus and potassium 
analyses were extracted according to the Egner-
Riehm method, and the concentrations of these 
elements were determined by a colorimetric and 
flame atomic emission, respectively. Soil samples 
for magnesium content estimation were extracted 
by the Schachtschabel method, and Mg levels were 
measured by the AAS 

For macro- and microelement analyses in plant 
material, one hundred leaves of plants grown in 
the tested substrates were sampled each year at the 
beginning of August. The leaves were dried, first 
at room temperature in a shaded place and then at 
~45 oC, and were pulverized with a WŻ-1 lab mill. 
Aliquots of berry samples of each harvest in the 
season were packed in polyethylene bags (250 g) 
and kept frozen (–25 oC). Following fruit collec-
tion, all fruit samples for each substrate were com-
bined, thawed at room temperature, and dried (ini-
tial temperature 60–70 oC, final 105 oC). The dried 
fruits were pulverized (WŻ-1). The total content 
of macro- and microelements in leaves and fruits 
was determined according to the Polish Standards. 
After mineralization, total nitrogen content was de-
termined by the Kjeldahl method. The concentra-
tions of K and Ca were measured by flame atomic 
absorption spectroscopy, whereas Mg, Cu, Zn, Fe, 
and Mn content was determined by atomic emis-
sion spectrometry using SAA Solaar. Phosphorus 
content was determined by the Barton method at 
a wavelength of 470 nm, and sulphur content - by 
the turbidimetric method at a wavelength of 490 
nm, with a Marcel s 330 PRO spectrophotometer. 

For plants grown in each substrate, bush growth 
rate was estimated and expressed as the total annual 
shoot length (cm) and the average length of one-
year shoots. Fruit yield was assessed and expressed 
as kg per ha. Additionally, the mean weight of 100 
fruits (g) was determined and fruit size measure-
ments (mm) at the vertical and horizontal axis were 
taken. Fruit firmness was measured along the ver-
tical axis (fruit height) and along the horizontal 
axis (fruit diameter) with a FirmTech 2 apparatus 
(BioWorks, USA) immediately after harvest and 
every 7 days during 21-day storage at 2 oC and 
96% relative air humidity. The firmness of 50 ber-
ries selected randomly from every replicate was 

expressed as a gram-force causing fruit surface to 
bend 1 mm.

Titratable acidity, the content of total sugars, 
soluble solids, L-ascorbic acid, nitrates and ni-
trites, and antioxidant capacity were determined 
in fresh fruit soon after harvest. Titratable acidity 
was determined by titration of the water extract of 
blueberry homogenate with 0.1 N NaOH to the end 
point of pH 8.1, according to PN-90/A-75101/04. 
Total sugar content was determined by the Luff-
Schoorl method. Soluble solids content was deter-
mined in berry juice with the use of an Abbé re-
fractometer (PN-90/A-75101/02). L-ascorbic acid 
content was determined by the iodometric method 
(Samotus et al. 1982). In order to measure juice 
extraction efficiency, fruits were homogenized with 
a blender and heated up to 50 oC. After cooling, 3 
mL of pectinase (Rapidase Super, BE, NC, USA) 
per kg of pulp was added. The pulp was left to 
stand at room temperature for 1 hour, and then it 
was pressed for 10 min at a final pressure of 300 
kPa with a laboratory hydraulic press (Oszmiański 
and Wojdylo 2005). Nitrate and nitrite content was 
measured with a RQflex 10 reflectometer (Merck). 
Fruit antioxidant capacity was evaluated by met-
myoglobin oxidation inhibition by antioxidants 
present in fruit extract and compared to that of 
Trolox (Miller and Rice-Evans 1996). Total anti-
oxidant capacity was expressed as μmol Trolox per 
g fruit tissue. Phenolics composition of blueber-
ries was determined in fruit samples that were kept 
frozen (–32 oC) in polyethylene bags (250–300 g) 
until analyzed. The 2 g aliquots of fruit (after thaw-
ing) were extracted three times with approx. 8 mL 
of 80% MeOH acidified with a glacial acetic acid 
(1 mL of 100% acetic acid per 1 L 80% MeOH) in 
an ultrasonic bath for 15 min. The samples were 
filtered and transferred to the flasks and made up 
to the final volume 25 mL. Further, the extracts 
were centrifuged twice at 12,000x g and 20 μL of 
supernatants were injected into the HPLC system. 
The HPLC apparatus consisted of a Merck-Hitachi 
L-7455 diode array detector (DAD) and quater-
nary pump L-7100 equipped with D-7000 HSM 
Multisolvent Delivery System (Merck-Hitachi, 
Tokyo, Japan). The separation was performed on 
a Cadenza CD C18 (75 × 4.6 mm, 5 mm) column 



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(Imtakt, Japan). Column oven temperature was set 
at 30 oC. The mobile phase was composed of sol-
vent A (4.5% formic acid), pH 2.2) and solvent 
B (acetonitrile). The program began with a linear 
gradient from 0% B to 21% B (0–30 min), followed 
by washing and reconditioning the column. The 
flow rate was 1 mL min-1 and the runs were moni-
tored at the following wavelengths: chlorogenic 
acid at 320 nm, flavonols glycosides (quercetin and 
kaempferol derivatives) at 360 nm, and anthocy-
anin glycosides at 520 nm. The Photo Diode Array 
spectra were measured over the wavelength range 
200–600 nm in steps of 2 nm. Retention times and 
spectra were compared to those of pure standards 
within 200–600 nm. Standards of anthocyanidin 
glycosides and chlorogenic acid were obtained 
from Polyphenols Laboratories (Norway), while, 
kaempferol 3- rutinoside and quercetin glycosides 
from Extrasynthese (France). Except for phenolics, 
the all measurements of bushes, leaves and fruits 
were performed for each year of the experiment. 
Phenolic patterns for berries were determined in 
2006 and 2007.

The results obtained were subjected to statisti-
cal analysis using Statistica 7.1 (Statsoft, Poland). 
The values were evaluated by the Duncan test and 
for phenolics by the Student test. The differences 
between the means at p<0.05 were considered sig-
nificant.   

Results and discussion

Data relating to plant growth, yield, fruit size and 
weight of blueberries cv. Patriot are presented in Table 
4. Three-year observations of highbush blueberry de-
velopment showed that peat was the most stimulating 
medium, superior to sawdust and cocoa husk with 
regard to total annual shoot length and average one-
year shoot length. Kozinski (2006) observed a higher 
growth rate of one-year-old shoots when blueberry 
was grown in soil mixed and mulched with sawdust 
compared to soil mixed and mulched with bark as 
well as in pure mineral soil (as control treatment).

In this experiment the vegetative growth of 
plants was not parallel to their productivity. Among 
the substrates tested, the highest yield was obtained 
from bushes grown in sawdust (3.661 kg ha-1 per 
season on average), whereas plants grown in cocoa 
husk were characterized by the lowest productivity 
(1.315 kg ha-1 per season on average). The values 
are so low due to frost damage in 2006. However, 
despite of unfavorable weather conditions, this spe-
cies allows multi-annual utilization of plantings, 
though its productivity is conditioned by the age 
of bushes and agronomic practices involved, too. 
In a study conducted by Ciordia et al. (2006), the 
yield of southern highbush cultivars grown in plas-

Table 4. Growth vigour, yield, weight of 100 fruits, and fruit size of ‘Patriot’ cv. highbush blueberry depending on sub-
strate type (an average for 2005–2007)

Type of substrate Peat Sawdust Cocoa husk

Total annual shoot length (cm) 401 b 311 a 315 a

Mean length of one-year shoots (cm) 46.0 b 37.7 ab 33.0 a

Mean yield (kg·ha-1) 2,639 b 3,661 c 1,315 a

Mean weight of 100 fruits (g) 140 b 126 a 120 a

Fruit size (mm)
ha 14.50 b 15.15 b 12.15 a

øb 17.85 b 17.85 b 16.35 a

Fruit size reduction between begin-
ning and end of harvest (%)

h -34.3 -37.6 -31.6

ø -46.6 -62.1 -48.9

ha – fruit size measured along fruit height (at vertical axis)
øb - fruit size measured along fruit diameter (at horizontal axis)



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tic tunnels, measured over four years, ranged from 
411.36 to 1990.90 g plant-1. Glonek and Komosa 
(2006) determined an average yield of 4.530–5.557 
kg per bush for 10-year-old highbush blueberry cv. 
Bluecrop, depending on fertilization. Smolarz et al. 
(2006) reported that the productivity of 25-year-old 
‘Bluecrop’ bushes under different fertilization con-
ditions was 2.90–20.77 t per ha.

The highest weight of 100 fruits was found for 
berries originating from bushes bedded in peat (140 
g). Plants grown in sawdust (126 g) and cocoa husk 
(120 g) provided smaller fruits. Strik et al. (2003) 
noted much higher fruit weight for ‘Bluecrop’ ber-
ries, ranging from 130 to 220 g (after conversion) 
and even higher for ‘Berkeley’ berries, between 180 
and 260 g (after conversion). One-fruit weight meas-
ured by Heiberg and Stubhaug (2006) varied from 
1.81 to 2.86 g, depending on cultivar and location. 
‘Patriot’ berries are slightly flattened, which can be 
seen when comparing fruit diameter and height (Ta-
ble 4). The highest dimensions were determined for 
berries from bushes grown in sawdust, whereas the 
lowest for those from bushes grown in cocoa husk. 
However, the largest berries (sawdust) did not have 
the highest weight of 100 fruits, while the small-
est berries (cocoa husk) had the lowest 100-fruit 
weight. Fruit size reduction was observed towards 
the end of the harvest season. Fruit diameter was 

dramatically reduced, particularly in berries from 
bushes grown in sawdust (by 62%), whereas fruit 
height was reduced to a lesser degree (>30%). Under 
field conditions, precipitation shortage or the lack 
of watering at the end of the growing season sub-
stantially contribute to fruit size reduction. Ciordia 
et al. (2006) tested six southern blueberry cultivars 
grown in trenches under tunnels, and observed in 
most varieties a decline in the weight of late-harvest 
berries equal to or less than 1 g, which made them 
unacceptable for the fresh fruit market.

The all berries showed higher values of firmness 
when it was measured along the vertical axis (fruit 
height) compared with the horizontal axis (fruit di-
ameter) (Table 5). As regards fruit hardness at diam-
eter position, berries from bushes grown in cocoa 
husk (the smallest ones) showed the highest bend-
ing resistance (210 G mm-1) and the lowest drop 
in firmness during 21-day cold storage (2 oC, 96% 
humidity). The highest firmness measured along the 
fruit height axis (492 G mm-1) and the lowest decline 
in hardness (by 6.1%) were found for berries from 
plants grown in sawdust (the biggest ones). Berries 
from bushes bedded in peat (~as big as those from 
plants grown in sawdust) showed the lowest firm-
ness measured along the horizontal and vertical axis 
(184 and 440 G mm-1, respectively) and the greatest 
decline in hardness (by 8.7 and 8.18%, respective-

Table 5. Changes of firmness for ‘Patriot’ blueberries after the harvest and during storage in a cold room (2 0C, 96% 
relative air humidity) on average for 2005–2007.
Storage time

 
Substrate

After harvest
7-day  

storage
14-day  
storage

21-day  
storage

Firmness changes after 21-day 
storage (%)

Fruit firmness measured at diameter (G mm-1)

Peat 184 180 175 168 -8.70 a

Sawdust 198 195 189 183 -7.58 ab

Cocoa husk 210 207 200 195 -7.14 b

Mean 197 b 194 ab 188 ab 182 a

Fruit firmness measured at height axis (G mm-1)

Peat 440 435 429 404 -8.18 a

Sawdust 492 490 478 462 -6.10 b

Cocoa husk 472 462 456 442 -6.36 b

Mean 468 c 462 bc 454 b 436 a



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ly). Thus, in terms of handling, these berries would 
be most sensitive to up- and side-pressure.

The substrates tested in this experiment differed 
with respect to nutrient abundance (Table 6). In gen-
eral, cocoa husk was abundant in P and Zn, sawdust 
was particularly rich in Mn and Cu but had a low 
Ca and Mg content, whereas peat had the lowest 
K content. Regardless of the substrate, blueberry 
leaves had higher amounts of total N, Ca, Mg, Fe 
and Mn, compared to fruits (Table 7). The highest 
N content was found in the leaves of plants grown 
in sawdust and in the fruits of plants grown in cocoa 
husk. According to Hanson (2006), the optimal N 
content of blueberry leaves picked in mid- summer 
amounts to 1.7–2.1%. After conversion the above 
data it can be seen that the N content of berry leaves 

was sufficient. Fruit N content was approximately 
half of leaf N content, and the values obtained were 
lower compared to 1.7-2.8% determined by Skupień 
(2004) in four blueberry cultivars. 

In this study, plants cultivated in cocoa husk 
had the highest concentration of P and K, both in 
the leaves and fruits. The leaf P status varying from 
1.11 to 1.26 g kg-1 was lower than that reported by 
Glonek and Komosa (2006) for cv. Bluecrop (0.18-
0.20%), while leaf K levels in cv. Patriot (5.15-6.99 
g kg-1) were similar to those determined by these 
authors in cv. Bluecrop (0.56–0.58%). Fruit K levels 
measured in this study (5.18–6.51 g kg-1) were lower 
compared to the values obtained by Skupień (2004) 
(0.738–0.855%).

Table 6.  The available macro- and microelements content in the tested substrates on average for 2005–2007.

P K Ca Mg Fe Mn Zn Cu

(mg·100g-1) (mg·100g-1)

Peat 13.04 b 29.65 a 170.1 b 28.62 b 202 a 32.03 a 12.65 a 1.81 a

Sawdust 7.54 a 52.49 b 83.4 a 21.94 a 231 ab 56.47 b 25.93 a 7.36 b

Cocoa husk 17.16 c 51.26 b 208.5 b 26.71 b 279 b 24.13 a 51.66 b 1.58 a

Table 7. Total macro- and microelements content in leaf and fruit of blueberry ‘Patriot’ on average for 2005–2007.

Macro- and 
microelementsa

Leaves Fruit

Peat Sawdust Cocoa husk Peat Sawdust Cocoa husk
(g·kg-1)

Total N 21.81 a 23.06 b 21.45 a 12.27 a 12.94 a 14.25 b

P 1.13 a 1.11 a 1.26 b  0.79 a 0.90 a 1.17 b

K 5.74 a 5.15 a 6.99 b 5.18 a 5.35 a 6.51 b

Ca 3.38 b 3.05 b 1.54 a 0.12 b 0.10 ab 0.08 a

Mg 1.71 b 1.48 ab 1.14 a 0.27 a 0.58 b 0.26 a

S 1.73 a 1.60 a 1.48 a 1.38 b 1.05 a 1.32 b

(mg·kg-1)

Cu 2.72 ab 3.89 b 1.75 a 1.53 a 3.01 b 1.87 a

Zn 9.41 a 10.16 b 9.64 a 6.60 a 5.66 a 8.78 b

Fe 58.67 a 58.93 a 52.91 a 23.67 a 21.64 a 22.78 a

Mn 199.89 b 170.46 b 121.04 a 24.49 a 23.03 a 22.73 a
a The values are calculated on dry weight basis



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In cv. Patriot, higher concentrations of Ca, Mg, 
and S were recorded in the leaves of bushes grown 
in peat. Glonek and Komosa (2006) determined the 
following values in blueberry leaves: 0.64–0.78% 
Ca, 0.15-0.16% Mg, and 0.11–0.12% S. A com-
parison of these data with our results shows that Ca 
levels in the leaves of blueberries grown in the tested 
substrates were distinctly lower, whereas Mg and S 
content was similar. Fruit S content in ‘Patriot’ ber-
ries (>1.0 g kg-1) was higher, compared to control 
lingonberries analyzed by Levula et al. (2000) (>700 
mg kg-1). ‘Patriot’ berries showed a lower fruit Ca 
content (0.08–0.12 g kg-1) and a higher fruit Mg con-
tent (0.26–0.58 g kg-1), in comparison with blue-
berries studied by Skupień (2004) (0.127–0.191% 
and 0.016–0.018%, respectively). The low leaf Ca 
and fruit Ca levels observed in this study could re-
sult from better acidification of the substrates used. 
As regards between-substrate differences, ‘Patriot’ 
berries from plants grown in peat had the highest 
Ca content. The highest fruit S levels were deter-
mined for berries originating from cocoa husk and 
peat beddings, whereas berries from plants grown 
in sawdust were richest in Mg.

Plants cultivated in substrates rich in microele-
ments showed elevated concentrations of microele-
ments in the leaves and fruits (Table 7). The high-
est Cu levels were noted in the leaves and fruits of 
‘Patriot’ bushes grown in sawdust (3.89 mg kg-1 and 
3.01 mg kg-1, respectively) which was most abun-
dant in Cu. Interestingly, the leaves and berries of 

plants grown in peat and cocoa husk had similar 
Cu levels as the respective substrates. As regards 
sawdust, leaf Cu and fruit Cu concentrations cor-
responded to the values (3.37–3.88 ppm) observed 
by Glonek and Komosa (2006) for the leaves of 
cv. Bluecrop, whereas fruit Cu content reported by 
Skupień (2004) was much lower (0.138–0.303 mg 
kg-1). The highest Zn concentrations determined in 
‘Patriot’ leaves (sawdust) and fruits (cocoa husk) in 
this study were correlated with Zn abundance in the 
medium. Leaf Zn content at 9.41–10.16 mg kg-1 was 
lower than that reported by Merhaut and Darnell 
(1996) at 27-31 µg g-1, and similar to that obtained 
by Glonek and Komosa (2006) at 7.98-8.38 ppm. 
Fruit Zn content (5.66-8.78 mg kg-1) was higher than 
that determined by Skupień (2004) at 1.081–1.30 
mg kg-1.

The all substrates tested as growing media for 
blueberry cv. Patriot showed high Fe concentrations 
(>200 mg 100 g-1). Leaf Fe content (52.91–58.93 mg 
kg-1) was very close to the range of 53.9-57.7 ppm 
reported by Glonek and Komosa (2006) and higher 
than the 36-37 µg g-1 range determined by Merhaut 
and Darnell (1996). On the other hand, fruit Fe con-
tent in blueberry cv. Patriot was ~50% that of leaf 
Fe content, however it was still higher than 14-15 
mg kg-1 reported for lingonberries by Levula et al. 
(2000). Despite statistical differences between the 
media tested, no significant differences were found 
for leaf and fruit Fe content.

Table 8. Blueberry ‘Patriot’ fruit chemical composition in dependence on the type of substrate (an average for 2005–2007).

Itema Peat Sawdust Cocoa husk

Soluble solids (%) 12.4 a 13.1 b 11.7 a 
Total sugar (g·100 g-1) 9.71 a 10.80 b 9.85 a
Titratable acidity (g citric acid·100 g-1) 2.41 b 2.35 b 1.98 a

L-ascorbic acid (mg·100 g-1) 26.0 b 23.4 a 26.2 b 

Juice efficiency (%) 89.98 a 90.29 a 89.88 a 

N-NO2 (mg·kg
-1) 0.75 a 0.95 b 0.85 ab

N-NO3 (mg·kg
-1) 19.3 a 31.4 b 24.6 ab

TEAC (μmol Trolox·g-1) 38.6 b 35.7 a 35.3 a
aThe values are presented on fresh weight basis
TEAC – Trolox Equivalent Antioxidant Capacity



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The highest Mn levels were recorded in the 
leaves of plants grown in peat and sawdust (Mn-
richest substrates) however the type of substrate did 
not affect fruit Mn content. Leaf Mn concentrations 
determined for cv. Patriot (121.04–199.89 mg kg-
1) were higher than those found for cv. Bluecrop 
(107.6–128.0 ppm) (Glonek and Komosa 2006), and 
both ranges were higher than the value of 44–57 µg 
g-1 reported by Merhaut and Darnell (1996). On the 
other hand, ‘Patriot’ berries had a low Mn content, 
ranging from 21.64 mg kg-1 (sawdust) to 23.67 mg 
kg-1 (peat).

‘Patriot’ berries obtained from sawdust-grown 
bushes had the highest soluble solids (13.1%) and 
total sugar content (10.80 g 100 g-1) (Table 8). In 
the study conducted by Skupień (2004) the soluble 
solids content of blueberry cultivars mulched with 
sawdust was determined within a similar range of 
11.6–13.8%. Prior et al. (1998) observed greater 
variation in soluble solids content (10.0–19.0%), 
affected by the cultivar, geographical location and 
harvest date. Total sugar content found in this inves-
tigation (9.85–10.80 g 100 g-1) corresponds to the 
sum of glucose and fructose reported by Wang et 
al. (2008) for blueberries in a conventional culture 
system (10.90 g 100 g-1). Ostrowska and Ściążko 
(1996) determined a slightly lower total sugar con-
tent of blueberries, ranging from 8.36 g 100 g-1(cv. 
Bluecrop) to 9.57 g 100 g-1(cv. Jersey).

Total acid content observed in this study was 
1.98–2.41 g citric acid 100 g-1 (Table 8). Higher 
acidity was observed in the berries of bushes bed-
ded in peat and sawdust. Rosenfeld et al. (1999) 
determined acid concentrations at 0.82% in ‘Blue-
crop’ berries stored at 4 °C. L-ascorbic acid content 
determined for ‘Patriot’ berries varied from 23.4 mg 
100 g-1 (sawdust) to 26.2 mg 100 g-1 (cocoa husk), 
and was not affected by the type of substrate (Table 
8). Łata et al. (2005) reported vitamin C content of 
16.6–30.6 mg 100 g-1 in blueberries, depending on 
the cultivar and seasonal variations. 

Juice efficiency measured for ‘Patriot’ berries 
was high (89.9–90.9%) and not medium-dependent 
(Table 8). Rossi et al. (2003) obtained a lower juice 
yield for blueberries, amounting to 79–81%, which 
could result from a different extraction procedure.

The nitrite content of blueberries obtained from 
plants grown in all media tested in the study was 
below 1 mg kg-1 (Table 8), which is consistent with 
Polish regulations concerning, among others, apple 
juice, fruit-vegetable juices and banana-containing 
products for babies and young children under three. 
Blueberries are characterized by low nitrate accu-
mulation. The values obtained in this study (Table 
8) confirm that blueberries are nitrate-safe product 
and the amounts ranging from 19.9 mg kg-1 (peat) 
to 31.4 mg kg-1 (sawdust) are fairly below the lev-
els permitted for bananas and vegetables meant for 
special usage (baby food, etc.) up to 200 mg kg-1.

The antioxidant capacity of blueberries is well 
substantiated in the literature (Prior et al. 1998, 
Ehlenfeldt and Prior 2001). In this experiment, 
blueberries showed TEAC values in the range of 
35.3–38.6 μmol Trolox·g-1 (Table 8). Connor et al. 
(2002) found lower antioxidant activity for ‘Patriot’ 
berries, varying from 20 to 29 μmol Trolox·g-1 de-
pending on the year of the experiment and location. 
Ścibisz et al. (2003) observed that early harvested 
berries showed 29.9 μmol Trolox·g-1, whereas late-
harvested berries – 61.8 μmol Trolox·g-1. A likely 
reason is that smaller fruits of late harvest have a 
higher surface:volume ratio. Since in blueberries an-
thocyanins are found in the skin only, the enhanced 
amount of these compounds (apart from other phe-
nolics) in smaller berries contributes substantially to 
their higher antioxidant activity.

The type of substrate exerted a significant in-
fluence on total phenolic content (Table 9). Ber-
ries from bushes grown in cocoa husk showed the 
highest amounts of phenolics (208.29 mg 100 g-1), 
while the lowest value was observed for sawdust-
originating berries (122.43 mg 100 g-1). Connor 
et al. (2002) measured a total phenol content of 
360–569 mg chlorogenic acid equivalents 100 g-1 
in ‘Patriot’ berries. Prior et al. (1998) estimated 
phenolic content at 181.1-390.5 mg 100g-1 in blue-
berry cultivars obtained from different sources. In 
this study, the amount of total phenols was not in 
accordance with TEAC values. Berries from plants 
grown in sawdust and cocoa husk beddings had 
almost equal antioxidant capacity (35.7 and 35.3 
μmol Trolox g-1, respectively), whereas the phenol 
content of berries originating from cocoa husk was 



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79

1.7-fold higher than that of sawdust-grown berries. 
Berries collected from peat-grown bushes showed 
the highest antioxidant activity (38.6 µmol Trolox 
g-1), although they had a medium phenolic content 
(174.08 mg 100 g-1). 

The proportions of phenolics identified in blue-
berries were as follows: for peat – 70.4% anthocy-
anins> 22.7% chlorogenic acid > 6.9% flavonols, for 
sawdust – 85.3% anthocyanins > 8% chlorogenic 
acid > 6.7% flavonols, for cocoa husk – 55.4% an-
thocyanins > 39.7% chlorogenic acid > 4.9% fla-
vonols. The greatest differences between berries 
were observed with respect to chlorogenic acid (Ta-
ble 9). The berries from plants grown in cocoa husk 
bedding were richest in chlorogenic acid (82.79 mg 
100 g-1) whereas, the chlorogenic acid content of 

berries from bushes grown in peat and sawdust was 
~2-fold and ~8-fold lower, respectively. Zheng and 
Wang (2003) determined lower chlorogenic acid 
concentrations in blueberry cv. Sierra - 645.9 µg 
g-1. Regardless of substrate type, delphinidin glyco-
sides dominated among anthocyanins (41.4–52.12 
mg 100g-1), especially delphinidin 3-glucoside and 
delphinidin 3-galactoside (Table 9). Cyanidin, peo-
nidin, petunidin and malvidin-glycosides were also 
detected. Berries originating from peat had a higher 
content of total anthocyanins (122.6 mg 100g-1), 
whereas the lowest amounts of these pigments were 
found in the fruits of sawdust-grown plants (104.45 
mg 100 g-1). Connor et al. (2002) determined 140-
191 mg cyanidin 3-glucoside equivalents 100 g-1 in 
‘Patriot’ berries, depending on the year of the experi-

Table 9. Influence of substrates on phenolic composition of ‘Patriot’ blueberries (an average for 2006–2007).

Peat Sawdust Cocoa husk
mg·100g-1

Chlorogenic acid 39.52 b 9.78 a 82.79 c
Cyanidin -3-arabinoside 5.43 3.85 2.75
Cyanidin-3-galactoside 6.14 5.37 3.56
Cyanidin-3-glucoside 5.90 5.13 3.78
Delphinidin-3-arabinoside 10.94 9.74 9.64
Delphinidin-3-galactoside 15.91 14.21 18.27
Delphinidin-3-glucoside 19.52 17.51 24.21
Peonidin-3-arabinoside 6.92 5.47 3.09
Peonidin-3-galactoside 5.92 5.57 7.19
Peonidin-3-glucoside 6.71 5.20 3.50
Petunidin-3-arabinoside 7.18 2.38 11.67
Petunidin-3-galactoside 1.81 4.97 3.05
Petunidin-3-glucoside 3.92 4.89 6.41
Malvidin-3-arabinoside 9.79 7.45 9.36
Malvidin-3-galactoside 10.56 8.36 4.95
Malvidin-3-glucoside 5.95 4.35 3.88
Sum of anthocyanins 122.6 b 104.45 a 115.31 ab
Quercetin-3-galactoside 5.93 4.01 5.83
Quercetin-3- glucoside 1.44 1.04 1.17
Quercetin-3- ramnoside 2.07 1.46 1.94
Kaempferol-3-rutinoside 2.52 1.69 1.26
Sum of flavonols 11.96 a 8.2 a 10.2 a
Total 174.07 b 122.43 a 208.29 b



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79

ment and the growing area. In an experiment per-
formed by Prior et al. (1998), the total anthocyanin 
content of blueberries varied over a wide range of 
93.1 to 235.4 mg 100 g-1. Quercetin derivatives and 
kaempferol 3-rutinoside were also identified, and 
quercetin 3-galactoside was found to be a predomi-
nant flavonol. The total of quercetin derivatives var-
ied from 6.51 mg 100 g-1 (sawdust) to 9.44 mg 100 
g-1 (peat), and these values were higher compared 
with the data obtained for blueberries of different 
varieties and sites by Häkkinen and Törrönen (2000) 
– 2.2-4.7 mg 100 g-1, but lower than those reported 
by Zheng and Wang (2003) for cv. Sierra (248.7 
µg g-1). However, in the present study the type of 
medium did not significantly affect total flavonol 
content (8.2–11.96 mg 100 g-1).

Conclusions

A three-year study of substrates tested as growing 
media for highbush blueberry cv. Patriot showed 
that the most intensive bush growth was observed 
when plants were grown in peat, while the highest 
yield (on a 3-year average) was attained in plants 
grown in sawdust. The berries of bushes bedded 
in sawdust were largest, but they showed the most 
disadvantageous fruit size reduction as the picking 
period progressed. On the other hand, these berries 
demonstrated the highest firmness measured along 
the vertical axis (fruit height), whereas the berries of 
bushes grown in cocoa husk had the highest firmness 
measured along the horizontal axis (fruit diameter).

Irrespective of the substrate used, the leaves of 
‘Patriot’ plants contained larger amounts of N, Ca, 
Mg, Fe and Mn than the fruits. As regards particular 
media, the concentrations of macro- and microele-
ments in leaves and fruits varied, except for leaf Fe, 
fruit Fe and fruit Mn content which showed no cor-
relation with the substrate. 

Berries originating from sawdust contained the 
highest amounts of soluble solids and total sugars, 
and the lowest L-ascorbic acid levels. For all ber-
ries, juice efficiency was high and not substrate-de-
pendent. The accumulation of nitrates and nitrites in 

blueberry cv. Patriot was low and did not exceed the 
permissible levels for food items designed for babies 
and young children.

Berries originating from peat had the highest 
antioxidant capacity and the highest total anthocy-
anin content. Delphinidin-glycosides were pre-
dominant anthocyanins in all berries. The following 
phenolics were identified in ‘Patriot’ berries (in de-
scending order): anthocyanins > chlorogenic acid > 
flavonols. The proportions of individual compounds 
were substrate-dependent. Berries originating from 
the cocoa husk bedding showed the highest content 
of chlorogenic acid and total polyphenols. On the 
other hand, the amount of total flavonols was not 
affected by the type of substrate.

From the quantitative point of view, the best pro-
ductivity (yield) and consumer attractiveness (fruit-
size) were observed for ‘Patriot’ blueberries grown 
in sawdust. These berries were richest in N, P, K, 
Zn, total sugars and soluble solids. However, their 
most undesirable characteristic was a considerable 
fruit size reduction at the end of the harvest season, 
and lower concentrations of anthocyanins and total 
phenols, compared with berries grown in peat and 
cocoa husk respectively.
Acknowledgments. The study was supported by the grant of 
the Scientific Research Committee No.0395/P06/2004/26.

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	Effect of substrate type on the field performance and chemical composition of highbus hblueberry cv. Patriot
	Introduction
	Materials and methods
	Results and discussion
	Conclusions
	References