Scientia Horticulturae 129 (2011) 479–485

Contents lists available at ScienceDirect

Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Effects of cultural cycles and nutrient solutions on plant growth, yield and fruit
quality of alpine strawberry (Fragaria vesca L.) grown in hydroponics
Gianluca Caruso a,1 , Gerardo Villari b , Giuseppe Melchionna c,2 , Stefano Conti c,∗
a
Department of Soil Plant Environmental and Animal Production Sciences, University of Naples “Federico II”, Via Università 100, 80055 Portici (NA), Italy
b
Experimental Station for the Food Preserving Industry, Angri (SA) Branch, Via Nazionale 121/123, 84012 Angri (SA) Italy
c
Department of Arboriculture Botany and Plant Pathology, University of Naples “Federico II”, Via Università 100, 80055 Portici (NA), Italy

a r t i c l e i n f o a b s t r a c t

Article history: Alpine strawberry (Fragaria vesca L.) was grown in hydroponics with the nutrient film technique, in
Received 16 August 2010 order to evaluate the effects of four buffer concentrations (1.3, 1.6, 1.9, 2.2 mS cm−1 ) and two cultural
Received in revised form 8 April 2011 cycles (summer-spring versus autumn-spring) in terms of growth, yield and fruit quality (dry and opti-
Accepted 15 April 2011
cal residues, sugars, acids, antioxidants, mineral composition). The longer summer-spring cycle gave a
correspondingly higher yield than the autumn-spring one. The 1.3 mS cm−1 nutrient solution was the
Keywords:
most effective in terms of overall and spring production. However, the autumn and winter yields were
Alpine strawberry
not affected by the buffer EC. Fruit quality did not change with the cultural cycle, but the berries har-
Nutrient film technique
Fruit production
vested in the spring had higher vitamin C and sucrose content and lower nitrate content compared with
Mineral nutrients berries picked up in the winter. Fruit quality was also improved when the nutrient solution concentration
Sugars increased. From the productive point of view, the cultural cycle choice should be made considering that
Antioxidants 71% of the yield of the more productive summer-spring cycle derived from the spring harvest. Moreover,
as regards the nutrient solution strength, 1.3 mS cm−1 EC should be preferred during the spring season,
whereas the 2.2 mS cm−1 EC proved to be best in the winter in terms of fruit quality.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction Given its technical and economical advantages, the hydroponic

Alpine strawberry (Fragaria vesca L.) is to be considered a niche
product which can provide interesting economic results in a wide
range of climatic areas. It is an ever-bearing crop, characterized by
a lower productivity compared with the more commonly grown
garden strawberry (Fragaria x ananassa Duch.) and by higher pro-
duction costs, owing to the prolonged hand harvest and smaller
fruits. However, due to its intense taste and olfactory characteris-
tics, alpine strawberry fruits are requested both by the fresh market
and by the confectionery industry. Therefore, these fruits have a
high market value which may encourage farmers to increase the
cropped area devoted to this species. Nevertheless, as a conse-
quence of the methyl bromide prohibition for soil disinfection,
the cultural practices previously followed for this crop need to be
improved.
∗ Corresponding author. Tel.: +39 081 7760104x46; fax: +39 081 7755114.
E-mail address: stefano.conti@unina.it (S. Conti).
1
Tel.: +39 081 2539104.
2
Tel.: +39 081 7760104x46.
cultivation of alpine strawberry could be promoted among pro-
ducers. In fact, the soilless system facilitates harvest and results in
earlier production of larger and more uniform fruits as preferred
by the market.
Hydroponic cultivation allows farmers easily to control the
nutrient supply, by adjusting the concentration of the hydroponic
nutrient solution. This factor affects the plant water and salt rela-
tions and influences plant growth and quality. In fact, it was
reported that salinity reduced strawberry vegetative growth but,
at moderate levels, it improved fruit quality and antioxidant con-
tent (Awang et al., 1993; Awang and Atherton, 1995; De Pascale
et al., 2001; Keutgen and Pawelzik, 2007a,b).
The choice of the crop cycle for strawberry hydroponic culti-
vation also plays a crucial role, since the seasonal environmental
factors affect the crop productivity and quality in the following
ways. Plant growth is favoured by increasing daylight and flowering
is enhanced by moderate temperature (Chabot, 1978). Light inten-
sity increase also leads to higher sugar and ascorbic acid contents
in many fruits and vegetables (Lee and Kader, 2000). High temper-
atures enhance the fruit antioxidant capacity (Wang and Zheng,
2001); however, they have a negative effect on soluble solids, titrat-
able acidity, and ascorbic acid contents (Wang and Camp, 2000).

0304-4238/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2011.04.020
480 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485

Table 1
Mean values of PAR, temperature and relative humidity from July 2004 to June 2005, in Portici (Naples, Italy).

2004 2005

Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun
−2 −1
PAR (MJ m d ) 9.8 9.0 6.2 4.2 2.6 2.0 2.7 3.1 5.7 6.9 8.9 9.7
Temperature (◦ C)
Day 34.1 33.9 31.2 28.8 24.5 21.6 20.3 17.2 22.9 26.7 30.3 31.7
Night 18.4 19.8 16.8 15.3 9.6 8.6 3.9 1.9 5.6 10.2 15.0 16.5
Relative humidity (%)
Day 95.5 96.2 97.0 97.3 98.6 99.0 99.7 99.4 98.2 97.2 96.6 96.3
Night 40.0 43.1 45.0 54.9 56.0 60.4 60.6 61.4 53.1 51.6 49.3 46.2

Environmental factors also influence the ratios between soluble
sugars, organic acids and insoluble solids contents of the fruits
(Awang and Atherton, 1995; Montero et al., 1996; Wang et al.,
2002).
Thus, although the effects of these individual factors have
already been documented in the related species F. x ananassa,
there remain little information about their effect on the productive
results of F. vesca.
Given the scarcity of research on the subject, we planned an
experimental protocol that aimed to define both the most effec-
tive nutrient solution strength (within the 1.3–2.2 mS cm−1 EC
range) and cultural cycle (summer-spring versus autumn-spring)
on the yield and quality performances of alpine strawberry grown
in hydroponics in southern Italy.
2. Materials and methods
2.1. Plant materials and growth conditions
Alpine strawberry plants (F. vesca L.) cultivar Regina delle Valli
were grown at the experimental site of the University of Naples at
Portici (NA) in southern Italy, in 2004–2005. Plants were grown
in hydroponics using the nutrient film technique (NFT) under a
300 m2 polyethylene tunnel. The monthly means of PAR, tempera-
ture (day/night) and relative humidity (day/night) recorded at the
plant level are reported in Table 1. The NFT equipment consisted of
rigid PVC gullies (each 12 cm wide, 10 cm deep and 300 cm long),
with a 1% slope. The gullies were at 70 cm above ground level and
each gully was fed by a separate 220 l plastic reservoir tank contain-
ing the nutrient solution (NS). Continuous circulation (3 l min−1 ) of
the NS was provided by a 90 W submerged pump into each reser-
voir tank. The volume of NS in each tank was constantly monitored
and it was completely replaced at weekly intervals.
Plants were exposed to four levels of NS concentration, result-
ing in electrical conductivities (EC) of 1.3, 1.6, 1.9 and 2.2 mS cm−1
(Table 2), in factorial combination with two crop cycles (summer-
spring and autumn-spring). The experimental treatments were
randomized in a split-plot design, assigning the crop cycles to the
main plots and the NS concentrations to the sub-plots. Each treat-
ment included 12 plants and it was repeated three times.
Fresh plants (75 days old) were transplanted on 22 July for
the summer-spring cycle or on 21 October for the autumn-spring
crop. Both crop cycles ended on 10 June in the following year. All
plants were transplanted in 10 cm black plastic pots filled with lapil
(5–6 mm). Pots were placed on the NFT gullies through a pierced
white polyethylene film. The gullies were arranged in double rows
which were spaced by 100 cm. Within each double row the plant
spacing was of 40 cm between the rows and 25 cm along the row.
Fruit harvest began on 16 September and 18 February, for the
summer-spring and for autumn-spring cycles respectively and it
continued until the end of the crop cycle, on 10 June.
2.2. General analytical methods
Ripe, undamaged fruits of regular shape were classified as “mar-
ketable”. At each harvest, the weight and number of marketable ripe
fruits in each plot was recorded. The weight of fruits unsuitable for
the market was also recorded in order to monitor total biomass
production for each treatment. Cumulative plant biomass was cal-
culated as the sum of the above ground plant biomass at the end of
the experiment plus the total fruit production from the beginning
of the harvest period. Dry residue was assessed after dehydration
of the fresh samples in an oven at 70 ◦ C under a vacuum until they
reached constant weight. Leaf area was measured at the end of the
cycle, using a bench top LI-COR leaf area meter.
2.3. Plant water consumption and nutrient uptake
Plant water consumption was monitored during the whole cul-
tural cycle. This was calculated as the difference between the NS
volumes in the hydroponics reservoir at the beginning and at the
end of the weekly cycle.
The uptake of nutrients from the hydroponic solution was
assessed in May when the maximum values of water consumption
were also recorded. Nutrient uptake was estimated as the differ-
ence between the concentration of each nutrient in the fresh NS
compared with the residual concentration in the exhausted solu-
tion after one week, when this was replaced with fresh NS. The
concentration of nutrients in the NS was measured by directly ana-
lyzing samples of the NS using the methods described below for
the analyses of cations and of anions.
2.4. Fruit sample preparation
In order to evaluate the quality of berries produced during the
winter and the spring, marketable ripe fruits were sampled on

Table 2
Chemical composition of the hydroponic nutrient solutions.

EC (mS cm−1 ) Macronutrients (mmol l−1 ) Micronutrients (␮mol l−1 )

N P K Ca Mg S Cl Fe Cu Mn Zn B Mo

1.3 6.1 0.5 2.6 2.0 1.6 1.3 1.0 45.0 20.0 25.0 38.0 12.0 1.0
1.6 7.8 0.7 3.3 2.4 1.8 1.8 1.0 45.0 20.0 25.0 38.0 12.0 1.0
1.9 9.5 0.9 4.1 3.0 2.1 2.4 1.0 45.0 20.0 25.0 38.0 12.0 1.0
2.2 11.3 1.0 4.8 3.6 2.3 2.8 1.0 45.0 20.0 25.0 38.0 12.0 1.0

For all treatments pH was adjusted to 5.8 and NH4 /NO3 ratio was 1/9.
 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485 481

Table 3
Growth indexes and yield results of alpine strawberry grown in hydroponics.

Growth indexes Marketable fruits per plant Productive season (days)

DW (g) Leaf area (cm2 ) Total weight (g) Number of fruits Mean weight (g) Production delay Duration

Crop cycle
Summer-spring 77.3 5644.3 185.2 155.8 1.20 56 268
Autumn-spring 56.6 4401.0 128.2 105.3 1.21 120 113
* * * * n.s. *
EC (mS cm−1 )
1.3 73.6a 5820.6a 180.3a 145.4a 1.25a 91a
1.6 71.6a 5485.4a 168.2b 136.6b 1.24a 90ab
1.9 65.7b 4783.7b 153.8c 129.9c 1.19b 87bc
2.2 56.8c 4001.0c 124.7d 110.4d 1.14c 86c

Growth indexes and yield results are reported on a “per plant” basis. Within each column, significant difference at * p ≤ 0.05, n.s. not significant; means followed by different
letters are significantly different according to the Duncan test at p ≤ 0.05. DW = cumulative dry weight, production delay is the time interval in days between the transplant
in hydroponics and the beginning of fruit production.

20 February (winter harvest) and on 5 May (spring harvest), then
rapidly transferred to the laboratory for analysis.
One hundred grams of berries were homogenized in a 1.0 l War-
ing blender (Waring Laboratory, Torrington, CT, USA) and aliquots
of this raw homogenate were used for the analyses of cations and
carotenoids. The raw homogenate was centrifuged at 10,000 × g
for 30 min at 6 ◦ C in an 5810R Eppendorf refrigerated centrifuge
(Eppendorf HQ, Hamburg, Germany). The resulting supernatant
was passed through a 0.45 ␮m Acrodisc filter (Gelman Sciences,
MI, USA). Samples of this filtered fruit extract were used for anion,
sugar and citrate determinations.
2.5. Optical residue
The optical residue (in ◦ Brix) was measured at 20 ◦ C
with a Bellingham & Stanley, model RFM 81 digital refrac-
tometer on the supernatant obtained from raw homogenate
centrifugation.
2.9. Sugars
For sugar analyses (glucose, fructose, and sucrose) the filtered
fruit extracts were diluted with Milli-Q water and 10 ␮l samples
were injected into a Waters Sugar-pak column at 85 ◦ C. The eluent
was EDTA-Ca (50 mg l−1 ) in water, at a flow rate of 0.5 ml min−1 in
the isocratic mode and the detector was a Waters refractive index
detector.
2.10. Citric and malic acid
For citric and malic acid determination, 10 ␮l samples of the fil-
tered extracts were injected into a Biorad column, model HPX87H
at 35 ◦ C. The eluent was 0.01 N sulphuric acid at a flow rate of
0.6 ml min−1 . The detector was a Waters 486 UV detector set to
410 nm.
2.11. Ascorbic acid

Fifty grams of berries were homogenized with 200 ml of 3% cit-

2.6. Cations
A 4.0 g aliquot of the raw homogenate was digested in a
mixture of 10 ml 65% ultra pure HNO3 and 2.5 ml 95% ultra
pure H2 SO4 in a mineralization vessel. The digested samples
were diluted to 50 ml with ultra pure water (Milli-Q) in volu-
metric flasks. Cation (Ca, Mg, K, Fe, Cu and Zn) concentrations
in the digested samples were determined by atomic adsorp-
tion spectrophotometry (AAS) with a model 1100 Perkin-Elmer
spectrophotometer.
2.7. HPLC analysis
Anions, sugars, organic acids, and carotenoids were determined
by high-performance liquid chromatography (HPLC) using a Waters
600E chromatographic system, managed through the Millennium
software version 3.05.01.
2.8. Anions
Ten-microliter samples of filtered fruit extract were used
for the simultaneous determination of nitrates, phosphates, sul-
phates and chlorides by HPLC. The analytical column was a
Dionex anion exchange column (model AS11, 4 mm × 250 mm,
code P/N 044076) with a Dionex pre-column (4 mm × 50 mm,
code P/N 044078). The eluent was 21 mmol l−1 NaOH at a flow
rate of 1 ml min−1 . The detector was a Dionex pulsed elec-
trochemical detector together with an anion self-regenerating
suppressor.
ric acid in a 1.0 l Waring blender. Aliquots of this homogenate
were transferred to volumetric flasks containing 0.2 M NaH2 PO4 ,
pH 2.14 to a total volume of 100 ml under continuous stirring. After
15 min the samples were centrifuged at 10,000 × g for 30 min at
6 ◦ C in a refrigerated centrifuge, and the resulting supernatant was
passed through a 0.45 ␮m Acrodisc filter prior to HPLC analysis.
Twenty-microliter samples of this cleared extract were injected
into a Polymer Laboratories reverse phase column mod. PLRP-S
100 Å, 5 ␮m (4.6 mm × 250 mm). The column was kept at 30 ◦ C, the
eluent was 0.2 M NaH2 PO4 , pH 2.14 at a flow rate of 0.6 ml min−1 ,
in the isocratic mode and the UV detector was set to 220 nm.
2.12. Carotenoids
One gram of fruit raw homogenate was transferred to a 250 ml
separating funnel with 30 ml of dichloromethane:methanol (2:1)
containing 0.1% butylated hydroxytoluene (BHT) as an antioxidant.
The funnel was put on an orbital shaker for 15 min at 20 ◦ C. At the
end of this time the two phases were allowed to separate. The phase
containing the pigments was collected and the extraction proce-
dure was repeated three times in order to extract the remaining
pigments from the tissues.
The combined extracts were evaporated to dryness on a Rotova-
por rotary evaporator (Büchi Labortechnik AG, Switzerland) under
reduced pressure at below 40 ◦ C.
The dry residue was dissolved in 3.0 ml dichloromethane con-
taining 0.1% BHT and filtered through a 0.45 ␮m polyvinylidene
fluoride Acrodisc filter. Ten-microliter samples of this extract were
injected into a YMC Co. C30 column (5 ␮m, 250 mm × 4.6 mm) kept
482 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485

Table 4
Effect of NS strength on the seasonal distribution of alpine strawberry fruit production.

EC (mS cm−1 ) Fruit production (g) Fruit number

Autumn Winter Spring Autumn Winter Spring

1.3 45.2 E 7.9 F 166.6 A 32.3 E 6.3 F 134.0 A

1.6 43.8 E 9.6 F 143.5 B 34.6 E 7.7 F 119.8 B
1.9 43.4 E 11.0 F 121.7 C 36.0 E 8.6 F 109.8 C
2.2 43.0 E 10.1 F 95.5 D 35.6 E 7.9 F 90.7 D

Data are referred to marketable production on a “per plant” basis. Means followed by different letters are significantly different according to the Duncan test at p ≤ 0.01.

Table 5 tion were significantly larger than in the autumn-spring cycle. The
Water consumption (l−1 d−1 per plant) by alpine strawberry crop grown in
latter effect was a consequence of the higher number of fruits
hydroponics.
(+48%), while the mean weight of fruit was not significantly dif-
EC (mS cm−1 ) September November January March May ferent between the two cultural cycles. Total biomass and leaf
Summer-spring cycle area were significantly reduced at the 2.2 mS cm−1 EC treatment,
1.3 0.02a 0.03a 0.02 0.04 0.15a while the harvest season started 5 days earlier compared with the
1.6 0.02a 0.03a 0.02 0.04 0.14ab 1.3 mS cm−1 EC treatment. Both the total number of fruits and the
1.9 0.01b 0.02b 0.02 0.04 0.13bc
berry mean weight were significantly reduced at increasing NS
2.2 0.01b 0.02c 0.02 0.04 0.12c
n.s. n.s. strength.
Autumn-spring cycle Table 4 shows the seasonal distribution of strawberry fruit
1.3 0.01 0.02 0.13a production for the summer-spring cycle only, since there was
1.6 0.01 0.02 0.12ab
no autumn or winter production for the autumn-spring crop.
1.9 0.01 0.02 0.11ab
2.2 0.01 0.02 0.10b The seasonal factor very strongly affected the number of fruits
n.s. n.s. resulting in the maximum fruit production being recorded in the
Means followed by different letters are significantly different according to the Dun-
spring.
can test at p ≤ 0.05; n.s. no significant difference. The effect of the NS strength on fruit production was also
clearly season-dependent (Table 4). In the autumn and winter, the
total plant production was unaffected by the NS concentration,
at 30 ◦ C. The eluents were: eluent A (methanol/H2 O at 95/5 v/v, con- whereas in the spring the total yield decreased with increas-
taining 0.1% BHT and 0.05% TEA), and eluent B (dichloromethane, ing NS concentration, also in terms of fruit number and mean
containing 0.1% BHT and 0.05% TEA). The following gradient elu- weight.
tion was used: 95% A and 5% B in the beginning, maintained
for 35 min followed by 30% A and 70% B for 40 min. The elu- 3.2. Plant water consumption and mineral nutrient uptake
ent flow rate was 1.0 ml min−1 , and the UV–VIS detector was set
to 450 nm. The maximum water consumption (Table 5) was recorded in
May, when the crops were in their full fruiting stage and they were
2.13. Statistical analysis approaching the end of the cultural cycle. At this time, increasing
the NS strength caused a reduction in the crop water consump-
Data were processed by analysis of variance and mean separa- tion. At the 2.2 mS cm−1 EC treatment, water consumption was
tions were performed through the Duncan multiple range test, with 20% or 23% lower for the summer-spring or the autumn-spring
reference to 0.05 and 0.01 probability levels, using SPSS software plants respectively, compared with water consumption at the
version 15. 1.3 mS cm−1 treatment.
The cultural cycle had a significant effect on the plant water and
3. Results nutrient requirements (Table 6): the summer-spring crop displayed
a higher water and nutrient uptake than the autumn-spring one.
3.1. Vegetative plant growth and fruit production The absorption of all macronutrients (N; P; K; Ca; Mg) increased
with the increasing NS EC in both cultural cycles. However, the plant
Plant growth and fruit production were affected by the strength iron uptake showed a reduction from the 1.3 to the 2.2 mS cm−1
of the nutrient solution in both cultural cycles (Table 3). In the EC treatment, although this micronutrient concentration was kept
summer-spring cycle total biomass, leaf area and fruit produc- constant in the four nutrient buffers (45.0 ␮mol l−1 ).

Table 6
Water (l d−1 ) and nutrient (mg d−1 ) uptake of alpine strawberry grown in hydroponics in May.

Water Nitrogen Phosphorus Potassium Calcium Magnesium Iron

Crop cycle
Summer-spring 0.14 15.7 3.04 19.0 14.4 6.04 0.33
Autumn-spring 0.12 13.2 2.55 16.0 12.1 5.06 0.27
* * * * * * *
EC (mS cm−1 )
1.3 0.14a 11.3d 2.18c 13.7c 10.9b 4.99c 0.33a
1.6 0.13ab 13.7c 2.59b 16.6b 12.3b 5.35bc 0.31a
1.9 0.12c 15.6b 3.16a 19.0a 14.1a 5.81ab 0.30ab
2.2 0.11c 17.2a 3.25a 20.8a 15.6a 6.05a 0.27b

All data are reported on a “per plant” basis. Within each column, significant difference at * p ≤ 0.05; means followed by different letters are significantly different according
to the Duncan test at p ≤ 0.05.
 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485 483

Table 7
Fruit quality indicators of alpine strawberry grown in hydroponics.

DR (mg) OR (◦ Brix) Suc (mg) Glc (mg) Fru (mg) Citr (mg) Mal (mg) Asc (␮g) Lut (␮g) ␣-Car (␮g) ␤-Car (␮g)

Harvest season
Winter 180 12.9 10.8 134 161 74 25.2 547.7 8.75 1.83 2.27
Spring 187 13.7 21.2 131 158 107 30.7 2231.9 2.65 0.93 1.09
* * * n.s. n.s. * * * * * *
EC (mS cm−1 )
1.3 176d 12.6d 11.6c 126b 147c 86b 28.0 1346 5.51b 0.92c 1.47b
1.6 180c 13.0c 14.2bc 133a 159b 90ab 28.3 1404 5.69ab 1.29b 1.68ab
1.9 185b 13.5b 17.6ab 136a 167a 93a 28.1 1415 5.79a 1.57a 1.80a
2.2 194a 14.1a 20.8a 136a 166a 93a 27.4 1395 5.75a 1.75a 1.80a
n.s. n.s.

DR data are per g fw of fruit tissue homogenate; other data are per g dw of fruit tissue homogenate. Within each column, significant difference at * p ≤ 0.05, n.s. not significant;
means followed by different letters are significantly different according to the Duncan test at p ≤ 0.05. Abbreviations: DR, dry residue; OR, optical residue; Suc, sucrose; Glc,
glucose; Fru, fructose; Citr, citric acid; Mal, malic acid; Asc, ascorbic acid; Lut, lutein; ␣-Car, ␣-carotene; ␤-Car, ␤-carotene.

Table 8
Fruit mineral composition of alpine strawberry grown in hydroponics.

Treatment Calcium Magnesium Potassium Iron Copper Zinc Nitrates Phosphates Sulphates Chlorides
(mg) (mg) (mg) (␮g) (␮g) (␮g) (mg) (mg) (mg) (mg)

Harvest season
Winter 1.00 2.09 19.5 54.2 5.15 25.8 0.57 4.26 0.76 1.07
Spring 1.09 1.91 19.6 56.0 4.29 25.5 0.32 3.36 0.58 1.18
* * n.s. n.s. * n.s. * * * *
EC (mS cm−1 )
1.3 1.05 1.97 18.9b 57.3a 4.97a 26.5 0.35b 3.67 0.60b 1.20a
1.6 1.06 2.01 19.6a 57.1a 4.84ab 26.2 0.43ab 3.82 0.66ab 1.17a
1.9 1.05 2.01 19.8a 54.9b 4.65bc 25.5 0.48a 3.86 0.71a 1.11ab
2.2 1.04 2.01 19.9a 51.2c 4.43c 24.5 0.52a 3.82 0.71a 1.04b
n.s. n.s. n.s. n.s.

All data are per g dw of fruit tissue homogenate. Within each column, significant difference at * p ≤ 0.05; n.s. no statistically significant difference; means followed by different
letters are significantly different according to the Duncan test at p ≤ 0.05.

3.3. Fruit quality and chemical composition
Fruit quality indicators were not significantly different between
the two crop cycles. For this reason, here we report the data
referred to the summer-spring cycle only (Table 7).
The comparison between winter and spring fruits revealed
that the factor of “harvest season” had significant effects on
strawberry fruit quality. In particular, spring strawberries had
higher levels of dry residue, optical residue, sucrose and organic
acids. Notably, ascorbate content was as much as four times
higher in spring than in winter fruits. With the exception of
malate and ascorbate content, which did not depend signifi-
cantly on the NS strength, the remaining parameters were all
significantly affected by the NS strength and they increased with
increasing NS strengths. The fruit mineral composition was sig-
nificantly affected by the seasonal factor (Table 8). Strawberries
harvested in the spring contained on average a significantly higher
amount of calcium and chloride, while a lower content in mag-
nesium, copper, nitrates, phosphates and sulphates was recorded.
Notably, the nitrate content of strawberry fruit harvested in the
spring was 44% lower than in the winter.
The variation in NS strength had a diverse effect on the mineral
composition of strawberry fruits. At the highest NS strength potas-
sium, nitrate and sulphate content in the fruits increased, while
iron, copper and chloride content fell significantly with increasing
NS strength. Calcium, magnesium, zinc and phosphate content in
the fruits did not significantly change within this EC range.
4. Discussion
4.1. Plant growth and fruit production
Strawberry plants are very demanding in terms of water supply,
due to their shallow root system, large leaf area, and fruits with
a high water content. Therefore, strawberry plants do not easily
tolerate water deficit or osmotic stress (Razavi et al., 2008; Keutgen
and Pawelzik, 2009; Khayyat et al., 2009). In our research, the range
of nutrient solutions of increasing strengths supplied the plants
with increasing availability of mineral nutrients but, at the same
time, this also caused the plants to be exposed to a range of mild
water deficits.
The 2.2 mS cm−1 EC treatment resulted in the reduction of plant
biomass and leaf area by 23% and 31%, respectively, compared with
the 1.3 mS cm−1 treatment (Table 3). Similar effects on vegeta-
tive growth (D’Anna et al., 2003) as well as the plant morphology
and anatomy (Klamkowski and Treder, 2008) have already been
reported. In particular, the reduction in plant leaf area is already
well documented in F. x ananassa (Klamkowski and Treder, 2006;
Razavi et al., 2008) and it corresponds to the rapid plant adaptation
to water deficit (Munns, 2002).
The increasing NS strengths also affected fruit production
(Table 3): despite the higher availability of nutrients, the total
number of fruits, as well as the mean weight of each fruit was sig-
nificantly reduced by the NS strength increase. Similar effects on
fruit production have also been reported in F. x ananassa, where
fruit production was lower and fruit ripening was accelerated in
salt stressed plants (Gehrmann, 1985; D’Anna et al., 2003).
The comparison between the two different cultural cycles
(Table 3) shows that the longer summer-spring cycle allowed the
plants to develop a larger leaf area and a higher biomass compared
with the shorter autumn spring crop. In addition, total fruit pro-
duction for the summer-spring cycle was higher, because it also
benefited from the yield obtained from autumn to mid winter
which missed in the autumn-spring crop.
The “seasonal factor” had a strong effect on fruit production
and it influenced the plant response to the different NS strengths
(Table 4). According to Adams (2002), temperature and day length
may play a key role in affecting fruit production.
484 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485
In the autumn and in the winter, the cooler temperature and
shorter day caused a lower water consumption (Table 5) and fruit
production was unaffected by increasing NS strengths. In the spring,
instead, the varying NS concentrations did have a significant effect
both on fruit number and on mean fruit weight and hence the total
yield as well.
The crop peak water requirements in May (Table 5) were influ-
enced, in effect, by the NS strength whose increase resulted in
reduced leaf area and in lower water consumption, which are
symptoms of water deficit. A similar response to water deficit was
reported in F. x ananassa where the lower leaf area and also the size
and distribution of stomata contributed to reducing transpiration
(Gehrmann, 1985). The higher water requirements of the summer-
spring crop compared with the autumn-spring crop (Table 6) are in
accordance with the larger leaf area of the summer-spring plants
(Table 3).
Plants fed with higher NS strength had a correspondingly higher
uptake of nutrients (Table 6), showing that the uptake of nutrients
from the hydroponic buffer is enhanced by nutrient availability.
Interestingly, within the same range of NS strengths the increased
absorption of mineral nutrients corresponded to a lower water
consumption (Table 6). These data suggest that, within the exper-
imental range, the uptake of water and mineral nutrients were
independently regulated. While the water deficit effect caused a
decrease in water consumption, within the same range the uptake
of nutrients was positively regulated by their availability, which
increased together with the increasing strength of the NS. Simi-
lar results have been reported for F. x ananassa (Tagliavini et al.,
2005) and for Cucurbita pepo (Rouphael and Colla, 2009). Moreover,
considering that neither dry matter or leaf area increased with the
increasing availability of nutrients (Table 3), the increased min-
eral nutrient uptake may be regarded as “luxury” consumption.
This aspect is of special interest for human health in the case of
nitrate, which plants tend to accumulate in their tissues, but the
same behaviour is exhibited in relation to the other nutrients as
well.
4.2. Fruit quality and chemical composition
For increasing NS strengths, the higher availability of nutrients
resulted in higher fruit dry residue (the dietary fibre components)
and optical residue (the soluble solids) (Table 7). This effect was
also reported by Rouphael and Colla (2009) for C. pepo.
Moreover, the fruit dry residue and optical residue were higher
in the spring than in the winter. This may be a consequence of
the environmental climatic conditions which in the warmer season
allow for an increased biomass accumulation than in the winter.
Similar results have been reported by Rouphael and Colla (2005)
for C. pepo grown in hydroponics.
Similarly, the sugar content of the strawberries also followed the
increasing NS strength (Table 7). This result is in agreement with
previous works on F. x ananassa which reported an increase in sugar
content and in titratable acidity as a consequence of water deficit
or salinity (Klamkowski and Treder, 2008; Awang and Atherton,
1995). However, the improvement of fruit quality under water
deficit or salt stress does not appear to be generally true (Keutgen
and Pawelzik, 2007a).
Sucrose content was significantly higher in the spring than in
the winter, while glucose or fructose contents were not affected by
the seasonal factor (Table 7). These results do not agree with those
published by Ferreyra et al. (2007) who found that glucose, fructose
and sucrose content were higher in the winter than in the spring.
Among the organic acids, which are considered important com-
ponents of strawberry flavour (Montero et al., 1996; Wang et al.,
2002), citrate was more abundant than malate and its content
increased with the increasing NS strength (Table 7).
The seasonal factor very strongly affected the fruit content of
both of these organic acids which was higher in the spring than
in the winter, as well as the ascorbic acid content, which in the
spring was four times higher than in the winter. This latter effect
may be related to the enhanced availability of sugars as precursors
of ascorbate synthesis (Wheeler et al., 1998). These results indi-
cate that strawberry fruit composition and quality indicators are
effectively influenced by the seasonal environmental conditions.
Although quantitatively less abundant than ascorbate,
carotenoids are among the antioxidants present in strawberry
fruits. Interestingly, unlike ascorbate, the quantities of lutein,
␣-carotene, and ␤-carotene in the fruits were found to be higher in
the winter than in the spring (Table 7). Little has previously been
reported about the carotenoid content of strawberries or the influ-
ence of seasonal factors, while, for other species, contrasting data
have been published. In kale (De Azevedo and Rodriguez-Amaya,
2005), in endive and New Zealand spinach (De Azevedo-Meleiro
and Rodriguez-Amaya, 2005) carotenoid content was found to be
higher in the summer than in the winter, while for pakchoi (Bras-
sica rapa L. chinensis) the opposite trend was observed (Hanson
et al., 2009). An explanation was proposed by Rodriguez-Amaya
et al. (2008), who suggested that the actual carotenoid content of
a plant tissue resulted from the contrasting effects of temperature
and light intensity.
The fruit mineral composition did not change with the cultural
cycle but it was affected by the NS strength and by the seasonal
factor (Table 8). Notably, the fruit nitrate content was significantly
higher (+78%) in the winter than in the spring as a consequence
of the poorer light conditions in the winter (Roorda van Eysinga,
1984). This can be considered to be undesirable, since nitrates
are believed to be a potential threat to human health (Lundberg
et al., 2004). The Acceptable Daily Intake (ADI) for nitrate is 222 mg
of nitrate per day for a 60 kg adult, as established by the Joint
FAO/WHO Expert Committee on Food Additives in 2002 (Scientific
Panel on Contaminants, 2008). In the worst case, this limit would
be equivalent to consuming approximately 2.2 kg per day of alpine
strawberry (F. vesca) fruits grown by the hydroponic methods
described in the present work, which would be unlikely.
5. Conclusions
From this work we may draw the following concluding remarks.
As regards the choice of the crop cycle, the summer-spring gave a
higher yield compared with the autumn-spring, with most (71%)
of the total production deriving from the spring harvesting season.
In any case, the overall strawberry fruit quality was not affected by
the cultural cycle.
As far as the nutrient solution strength is concerned, its effect
was related to the harvest season. In the spring, plants fed with
the 1.3 mS cm−1 EC buffer gave the highest production, whereas for
the autumn and winter harvests no productive differences emerged
within the 1.3–2.2 mS cm−1 range. In this last season, the preference
for the highest buffer concentration could be justified by the better
quality of the fruits.
Acknowledgements
Dr. Christopher Latham is thanked for his help with the English
language. We also thank Mr. Rosario Nocerino for his expert assis-
tance with the equipment management.
References
Adams, P., 2002. Nutritional control in hydroponics. In: Savvas, D., Passam, H.C.
(Eds.), Hydroponic Production of Vegetables and Ornamentals. Embryo Publi-
cations, Athens, Greece, pp. 211–261.
 G. Caruso et al. / Scientia Horticulturae 129 (2011) 479–485
Awang, Y.B., Atherton, J.G., Taylor, A.J., 1993. Salinity effects on strawberry plants
grown in rockwool, II. Fruit quality. J. Hortic. Sci. 68, 791–795.
Awang, Y.B., Atherton, J.G., 1995. Growth and fruiting responses of strawberry plants
grown on rockwool to shading and salinity. Sci. Hortic. 62, 25–31.
Chabot, B.F., 1978. Environmental influences on photosynthesis and growth in Fra-
garia vesca. New Phytol. 80, 87–98.
D’Anna, F.D., Incalcaterra, G., Moncada, A., Miceli, A., 2003. Effects of different elec-
trical conductivity levels on strawberry grown in soilless culture. Acta Hort. 609,
355–360.
De Azevedo, C.H., Rodriguez-Amaya, D., 2005. Carotenoid composition of kale as
influenced by maturity, season and minimal processing. J. Sci. Food Agric. 85,
591–597.
De Azevedo-Meleiro, C.H., Rodriguez-Amaya, D., 2005. Carotenoids of endive and
New Zealand spinach as affected by maturity, season and minimal processing.
J. Food Compos. Anal. 18, 845–855.
De Pascale, S., Maggio, A., Fogliano, V., Ambrosino, P., Ritieni, A., 2001. Irrigation with
saline water improves carotenoids content and antioxidant activity of tomato.
J. Hortic. Sci. Biotechnol. 76, 447–453.
Ferreyra, R.M., Viña, S.Z., Mugridge, A., Chaves, A.R., 2007. Growth and ripening sea-
son effects on antioxidant capacity of strawberry cultivar Selva. Sci. Hortic. 112,
27–32.
Gehrmann, H., 1985. Growth, yield and fruit quality of strawberries as affected by
water supply. Acta Hort. 171, 463–469.
Hanson, P., Yang, R., Chang, L., Ledesma, L., Ledesma, D., 2009. Contents of
carotenoids, ascorbic acid, minerals and total glucosinolates in leafy brassica
pakchoi (Brassica rapa L. chinensis) as affected by season and variety. J. Sci. Food
Agric. 89, 906–914.
Keutgen, A.J., Pawelzik, E., 2007a. Modifications of strawberry fruit antioxidant pools
and fruit quality under NaCl stress. J. Agric. Food Chem. 55, 4066–4072.
Keutgen, A.J., Pawelzik, E., 2007b. Cultivar-dependent cell wall modification of
strawberry fruit under NaCl salinity stress. J. Agric. Food Chem. 55, 7580–7585.
Keutgen, A.J., Pawelzik, E., 2009. Impacts of NaCl stress on plant growth and min-
eral nutrient assimilation in two cultivars of strawberry. Environ. Exp. Bot. 65,
170–176.
Khayyat, M., Tafazoli, E., Rajaee, S., Vazifeshenas, M., Mahmoodabadi, M.R., Sajja-
dinia, A., 2009. Effects of NaCl and supplementary potassium on gas exchange,
ionic content, and growth of salt-stressed strawberry plants. J. Plant Nutr. 32,
907–918.
Klamkowski, K., Treder, W., 2006. Morphological and physiological responses of
strawberry plants to water stress. Agric. Conspec. Sci. 71, 159–165.
485
Klamkowski, K., Treder, W., 2008. Response to drought stress of three strawberry
cultivars grown under greenhouse conditions. J. Fruit Ornam. Plant Res. 16,
179–188.
Lee, S.K., Kader, A.A., 2000. Preharvest and postharvest factors influencing
vitamin C content of horticultural crops. Postharvest Biol. Technol. 20,
207–220.
Lundberg, J.O., Weitzberg, E., Cole, J.A., Benjamin, N., 2004. Nitrate, bacteria and
human health. Nat. Rev. Microbiol. 2, 593–602.
Montero, T.M., Mollà, E.M., Esteban, R.M., Lopez-Andreu, F.J., 1996. Quality attributes
of strawberry during ripening. Sci. Hortic. 65, 239–250.
Munns, R., 2002. Comparative physiology of salt and water stress. Plant Cell Environ.
25, 239–250.
Razavi, F., Pollet, B., Steppe, K., Van Labeke, M.C., 2008. Chlorophyll fluorescence as a
tool for evaluation of drought stress in strawberry. Photosynthetica 46, 631–633.
Rodriguez-Amaya, D.B., Kimura, M., Godoy, H.T., Amaya-Farfan, J., 2008. Updated
Brazilian database on food carotenoids: factors affecting carotenoid composi-
tion. J. Food Compos. Anal. 21, 445–463.
Roorda van Eysinga, J.P.N.L., 1984. Nitrate in vegetables under protected cultivation.
Acta Hort. 145, 251–256.
Rouphael, Y., Colla, G., 2005. Growth, yield, fruit quality and nutrient uptake of
hydroponically cultivated zucchini squash as affected by irrigation systems and
growing seasons. Sci. Hortic. 105, 177–195.
Rouphael, Y., Colla, G., 2009. The influence of drip irrigation or subirrigation on zuc-
chini squash grown in closed-loop substrate culture with high and low nutrient
solution concentrations. HortScience 44, 306–311.
Scientific Panel on Contaminants, 2008. Opinion of the Scientific Panel on Contami-
nants in the food chain on a request from the European Commission to perform
a scientific risk assessment on nitrate in vegetables. EFSA J. 689, 1–79.
Tagliavini, M., Baldi, E., Lucchi, P., Antonelli, M., Sorrenti, G., Baruzzi, G., Faedi, W.,
2005. Dynamics of nutrients uptake by strawberry plants (Fragaria x ananassa
Dutch.) grown in soil and soilless culture. Eur. J. Agron. 23, 15–25.
Wang, S.Y., Camp, M.J., 2000. Temperatures after bloom affect plant growth and fruit
quality of strawberry. Sci. Hortic. 85, 183–199.
Wang, S.Y., Zheng, W., 2001. Effect of plant growth temperature on antioxidant
capacity in strawberry. J. Agric. Food Chem. 49, 4977–4982.
Wang, S.Y., Zheng, W., Galletta, G.J., 2002. Cultural system affects fruit qual-
ity and antioxidant capacity in strawberries. J. Agric. Food Chem. 50,
6534–6542.
Wheeler, G.L., Jones, M.A., Smirnoff, N., 1998. The biosynthetic pathway of vitamin
C in higher plants. Nature 393, 365–369.