Journal of Plant Nutrition

ISSN: 0190-4167 (Print) 1532-4087 (Online) Journal homepage: http://www.tandfonline.com/loi/lpla20

Photosynthetic reaction, mineral uptake, and fruit

quality of strawberry affected by different levels of
macronutrients

Rouhi Shirko, Nazarideljou Mohammad Javad, Mozafari Ali Akbar & Ghaderi
Naser

To cite this article: Rouhi Shirko, Nazarideljou Mohammad Javad, Mozafari Ali Akbar & Ghaderi
Naser (2018): Photosynthetic reaction, mineral uptake, and fruit quality of strawberry affected by
different levels of macronutrients, Journal of Plant Nutrition, DOI: 10.1080/01904167.2018.1462380

To link to this article: https://doi.org/10.1080/01904167.2018.1462380

Published online: 01 Jun 2018.

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JOURNAL OF PLANT NUTRITION
https://doi.org/10.1080/01904167.2018.1462380

Photosynthetic reaction, mineral uptake, and fruit quality

of strawberry affected by different levels of macronutrients
Rouhi Shirkoa, Nazarideljou Mohammad Javada, Mozafari Ali Akbarb,c, and Ghaderi Naserb,c
a
Department of Horticultural Science, Mahabad Branch, Islamic Azad University, Mahabad, Iran; bDepartment of
Horticultural Science, Faculty of Agriculture, University of Kurdistan, Sanandaj, Iran; cResearch Center of Strawberry
Breeding and Improvement, University of Kurdistan, Sanandaj, Iran

ABSTRACT ARTICLE HISTORY

This experiment was conducted to evaluate the effects of three nutrient recipes Received 2 April 2017
containing different concentrations of macronutrients in the vegetative stage Accepted 4 December 2017
[Vegetative Nutrient Solution: (VNS-I, VNS-II, and VNS-III)] and two nutrient KEYWORDS
recipes in reproductive stage [Reproductive Nutrient Solution: (RNS-I and RNS-II)], Fruit firmness; Paros; shelf
on physiological and biochemical parameters of strawberry ‘Paros’ in a soilless life; TSS/TA; vitamin C
system. The results indicated the significant effects of nutrient solution on the
photosynthetic capacity, yield, minerals uptake and titratable acidity of
strawberry fruits. In the other hand fruit total soluble solids affected by
vegetative nutrient solution. The highest yield was obtained under application of
lowest level of minerals. Vegetative growth including leaf number and leaf area,
as well as total chlorophyll were the highest as potassium (KC), calcium (Ca2C),
and nitrogen (N) concentrations were increased. The highest content of N and
Ca2C uptake were in VNS-III £ RNS-I formulation. Additionally, the highest
vitamin C was in VNS-I formulation. Moreover, the most firmed fruits and the
highest post-harvest shelf life of fruits were produced in VNS-III £ RNS-I
formulation.

Introduction
Limited land and water resources for agricultural crops production and increasing human food needs
require extension of soilless culture systems (Putra and Yuliando 2015). This limitation is especially
important in countries with limited water resources such as Iran. In soilless systems, mineral availabil-
ity in comparison to other parameters (e.g., light, day and night temperature, and relative humidity) is
the main factor affecting yield and quality of crops production.
Strawberry is an important resource for antioxidant compounds, phenolics, organic acids, minerals,
and anthocyanin (Mandave et al. 2014). Recently, the total area used for the production of strawberry
in soilless systems has increased in Iran. The Kurdistan province is the main site for strawberry pro-
duction in field systems; however, the productivity in this system is low because of limitations of water
resources. This, and the high performance potential of soilless systems has led to the design of an
experiment based on Caruso et al. (2011), who used different levels of nutrients before and after flower-
ing to evaluate the quality and productivity of strawberry ‘Paros’.
Nutrient solution management and control of pH, electrical conductivity (EC), O2, and temperature of
solution are among the most important factors affecting successful production in soilless cultures (Gorbe
and Calatayud 2010). A direct or indirect effect of nutrients on fruit quality is anticipated. However, it is chal-
lenging to decide the critical and optimum level of an element or which ion interactions are consequential for

CONTACT Nazarideljou Mohammad Javad nazarideljou@yahoo.com Department of Horticultural Science, Mahabad Branch,
Islamic Azad University, P.O. Box 59135-433, Mahabad, Iran.
© 2018 Islamic Azad University of Mahabad
2 R. SHIRKO ET AL.

the development of fruit quality (Nestby et al. 2005). Quality of strawberry is based primarily on color, tex-
ture, and fruit flavor (do Nascimento Nunes 2009). In addition to temperature, time, and post-harvest condi-
tion, strawberry quality is mainly affected by minerals. In terms of nutrition physiology and nutrient uptake,
the concentration of individual mineral elements as well as the maintenance of a balance between nutrients is
very important (Heckman et al. 2003). The supply of different nutrients in a soilless system affected growth,
yield, and fruit quality of tomato “Durinta” (Krauss et al. 2006). Furthermore, plant growth and productivity
were increased in response to an increase in nutrient solution concentration in lettuce “Acephala” production
(Fallovo et al. 2009). Nutrient deficiency leads to reduced production (Fageria 2008), and high concentration
of minerals lead to ion toxicity and/or affect osmotic potential causing abiotic stress, subsequently decreasing
mineral uptake (Barker and Pilbeam 2015; Mitra 2015). Furthermore, availability of minerals during different
stages of growth affect crop yield and quality (Şirin 2011; Resh 2012). Although, the proper ratio of cations
and anions in nutrient solution has been documented (Steiner 1961; Jones Jr, 2004), it is not the same for dif-
ferent plants and at all developmental stages. In some cases, concentrations of nutrients are even more impor-
tant than the ratio between minerals. For instance, Neocleous and Savvas (2015) in a study on strawberry
“Camarosa” reported that EC of nutrient solution is more important than balancing cations ratio while bal-
ance of cations has been disrupted [especially calcium (Ca2C) compared to potassium (KC) and magnesium
(Mg2C)]. Additionally, in some cases commercial producers use mineral concentrations at toxicity levels due
to the absence of practical research, which can lead to environmental pollution. Caruso et al. (2011) and Lee
et al. (2015) reported that an increase in concentration of elements in the nutrient solution causes a decrease
in yield and quality of strawberry. Considering several studies on horticultural crops nutrition, most of these
researches are commercial reports and some have merely focused on the study of a single element (Choi et al.
2009), a few elements (Bautista et al. 2009), the ratio between KC and Ca2C (Torre et al. 2001), and the ratio
between nitrogen (N) and Ca2C (Voogt 2002). Therefore, a comprehensive study is needed to evaluate the
effect of different levels of macronutrients in both vegetative and reproductive phases on productivity and
quality attributes of strawberry ‘Paros’ in a soilless system. To this aim, we performed a study to select the
appropriate formula of macronutrients for vegetative and reproductive stages of strawberry to produce fruits
with high quality indices.

Materials and methods

This study was performed in a hydroponic greenhouse with polyethylene cover during winter and
spring of 2013. Greenhouse conditions were controlled [(photoperiod 12 hr; day/night temperature
25/18 § 2
C; relative humidity 65 § 5%)]. Strawberry seedlings were cultivated in 4-L pots. The cul-
ture medium was a mixture of perlite and cocofibre (30:70 V/V).
Three nutrient formulas during vegetative phase or before first flowering (VNS-I, VNS-II, and VNS-
III) and two formulas during reproductive phase or after flowering (RNS-I and RNS-II) were studied
on strawberry ‘Paros’ parameters in a factorial experiment basis on completely randomized design
with three replications. The concentrations of macro- and micro-elements were calculated (Table 1)
based on Caruso et al. (2011). The pH of the nutrient solution was adjusted to 6.5.

Table 1. Macronutrients concentration during vegetative and reproductive stages.

Vegetative stage Reproductive stage

Macronutrients (mmol L¡1) VNS-Iy VNS-II VNS-III RNS-Iyy RNS- II Micronutrients (mmol L¡1)

N 3.93 4.29 4.64 5.7 6.07 Fe 45

P 0.65 0.81 0.97 0.97 0.97 Cu 20
K 1.28 1.41 1.53 2.17 2.3 Mn 25
Ca 1.5 1.62 1.75 2.5 2.5 Zn 38
Mg 1.44 1.65 1.85 1.85 1.85 B 12
Mo 1
y
VNS: Vegetative Nutrient Solution; yyRNS: Reproductive Nutrient Solution.
 JOURNAL OF PLANT NUTRITION 3

Leaf number and leaf area

The leaf number of every replication was counted at the end of the experiment and plant leaf area was
measured using a leaf area meter (Leaf Area Meter AM 200; UK).

Photosynthetic pigments
In order to measure the photosynthetic pigments (Chl a, Chl b, and total Chl), fully expanded mature
leaves were sampled and dissolved in 80% acetone, and then centrifuged. The absorbances of the sam-
ples were measured using a spectrophotometer (Perkin Elmer, Lambda 25, UV/VIS Spectrophotome-
ter; Perkin Elmer, Waltham, MA) in wavelengths of 663.2 and 646.8 nm. The amount of pigments was
calculated based on Lichtenthaler and Wellburn (1983).

Chlorophyll a D .12:25£A663:2 / -- .2:79£A646:8 /; Chlorophyll b D .21:21£A646:8 / -- .5:1£A663:2 /;

Total Chlorophyll ða C bÞ D .7:15£A663:2 C 18:71£A646:8 /

Fruit yield
The mature fruits were counted during the experiment (winter and spring) and weighed after harvest
and considered as g plant¡1.

Fruit quality indexes

TA, TSS, and TSS/TA
Strawberries were cut into small pieces and homogenized in a grinder. The extract was titrated to pH
8.1 using 0.1 M sodium hydroxide (NaOH). Titratable acidity (TA) was expressed as g of citric acid
per 100 g of strawberry weight. The total soluble solids (TSS) were determined using a digital refrac-
tometer (Atago Co. Ltd., Tokyo; Japan) at 20
C and expressed as a percentage (Hernandez-Munoz
et al. 2008).

Fruit firmness
Fruit firmness was determined by peeling the fruit at two equatorial sites and measuring firmness by
means of a fruit firmness tester equipped with an 8-mm plunger tip, using five fruits from each treat-
ment. Values were expressed in Newton.

Protein assay
1 g of fresh leaf and fruit tissue was ground in 5 mL of 0.05 M Tris-hydrochloric acid (HCl) buffer, pH
7.5. The extraction was centrifuged for 25 min in 10000 £ g at 4
C (Hermle Z216 MK; Germany). The
supernatant was used to determine the protein concentration. 0.1 mL of extraction and 5 mL of Biuret
reagent were mixed and vertexed immediately. Afterwards, the absorbance of samples was read in
595 nm (Perkin Elmer UV/VIS Lambda 25; Waltham, MA) based on Bradford (1976) with some
modifications.
For preparing biuret reagent, 0.1 g coomassie brilliant blue G250 was added to 50 mL of ethanol
95%. Hundred milliliters of phosphoric acid 85% was added and mixed. The solution volume reached
1 L using distilled water. Protein was determined comparing standard curves.

Vitamin C
Briefly, 1 g of strawberry was extracted using 5 cc of tri-chloroacetic acid 10%. The extraction was cen-
trifuged in 3500 rpm (Hermle Z216 MK; Germany). 0.5 cc of supernatant was mixed with 1 cc of 2, 4-
4 R. SHIRKO ET AL.

dinitrophenylhydrazine (6M) reagent. The mixture was put in warm bath (37
C) for 3 hr. Afterwards,
0.75 cc sulfuric acid (H2SO4) 65% was added and put again in warm bath (30
C) for 30 min. The
absorbance of samples was read using spectrophotometer (Perkin Elmer, Lambda 25, UV/VIS) at
520 nm wavelength. The content of ascorbic acid was measured based on a standard curve and
expressed as mg 100 g¡1 (Omaye et al. 1979).

Determination of leaf mineral concentration

Fully expanded mature leaves were sampled and washed using distilled water. Samples were put in
oven (70
C) for 48 hr. Samples were extracted using chloric acid. K concentration was measured using
flame photometric method based on Chapman and Pratt (1962). Ca and Mg concentrations were mea-
sured using atomic absorbance method based on Perkin-Elmer (1964). The Kjeldahl method was used
for determination of total N based on Bremner et al. (1996). P was determined using the calorimetric
method by Chapman and Pratt (1962). Determination of micronutrients including iron (Fe), zinc
(Zn), and copper (Cu) was done using atomic absorbance method (Perkin-Elmer 1964).

Data analysis
Data analysis was done using SAS (V9.2) software (SAS Inc., Cary, NC). Means comparison was calcu-
lated according to the Duncan’s multiple range test at 1% and 5% levels.

Results
Growth parameters
There was a high correlation between growth parameters and nutrition formula. The highest leaf num-
ber (81.7 § 1.5) and leaf area (5414 § 24) were produced in VNS-III £ RNS-I and VNS-I £ RNS-II,
respectively (Table 2). Probably, the increased leaf number and leaf area in these nutrition formulas
are related to better uptake of N in these formulas (Table 3).
Healthy plants contain 3–4% N in their above-ground tissues, which is the most among other
nutrients. The role of N in cell growth and increasing leaf number and area is of outstanding impor-
tance among other effects. Additionally, N is a component of energy transfer compounds such as aden-
osine triphosphate (ATP) which allows cells to conserve and use the energy released in metabolism.
Probably, the role of N in the structure of proteins, amino acids, enzymes, and chlorophyll could be
the reason of high leaf number and leaf area produced in VNS-III £ RNS-I and VNS-I £ RNS-II for-
mulas, which is consistent with Lineberry et al. (1944) and Preusch et al. (2004).
Total chlorophyll was significantly affected by treatments (p < .05). The highest content of total
chlorophyll (1.29 § 0.029 mg g¡1 FW) was produced in treatment with VNS-II at vegetative stage and
RNS-II at reproductive stage (Table 2). According to the results, total chlorophyll changes was in line
with increase of nutrient solution EC in VNS-III compared to VNS-I which is related to N, Ca, and K
uptake in nutrient solution (Figure 1A and 1B).
According to the results, the highest content of total chlorophyll was related to RNS-II formulation.
Probably, increase in content of K in nutrient solution (KNS) and N in nutrient solution (NNS) has led
to increase in K and N uptake which consequently affected total chlorophyll (Tables 1, 2, and 3).
Photosynthesis is chief among the important and key biological processes. Photosynthesis produces
organic compounds by complex molecular and biochemical reactions. Minerals such as N, K, manga-
nese (Mn), Fe, and Ca are directly or indirectly involved in photosynthesis processes (Taiz and Zeiger
2010). The effect of efficient application of NNS in vegetative and reproductive stages increased photo-
synthetic pigments. N is a component of chlorophyll, protein, and amino acids structure (Taiz and
Zeiger 2010); therefore increase in N uptake positively affects the amount of chlorophyll in plant leaves
(Evans 1989). Sinclair and Horie (1989) reported that application of N fertilizer increased leaf chloro-
phyll and there was a linear relation between leaf chlorophyll and leaf N content.
Table 2. Effect of nutrition formula at different growth phases on photosynthetic capacity of strawberry ‘Paros.’

Growth stages Leaf number per plant Leaf area (cm2 plant¡1) Chlorophyll a (mg g¡1 FW) Chlorophyll b (mg g¡1 FW) Total Chlorophyll (mg g¡1 FW)

Vegetative
VNS-Iy 78 § 1.53ayyy 5089 § 185a 0.63 § 0.037 0.44 § 0.033 1.07 § 0.057b
VNS-II 51.8 § 3.16b 3517 § 372b 0.6 § 0.047 0.53 § 0.041 1.13 § 0.077ab
VNS-III 77.5 § 2.32a 4880 § 173a 0.69 § 0.026 0.55 § 0.036 1.24 § 0.031a
Statistical significance (p-values) <.0001 <.0001 ns ns .017
Reproductive
RNS-Iyy 71.7 § 3.57 4787 § 142 0.58 § 0.031 0.48 § 0.03 1.06 § 0.052
\RNS-II 66.6 § 5.5 4204 § 405 0.7 § 0.017 0.53 § 0.036 1.23 § 0.034
Statistical significance (p-values) .005 <.0001 .002 ns .002
Vegetative £ Reproductive
VNS-I RNS-I 75 § 1.15b 4765 § 64.4b 0.55 § 0.029 0.42 § 0.023 0.98 § 0.021b
RNS-II 81 § 1.16a 5414 § 248a 0.71 § 0.016 0.45 § 0.07 1.15 § 0.09a
VNS-II RNS-I 58.3 § 2.33c 4334 § 149c 0.51 § 0.038 0.46 § 0.055 0.97 § 0.043b
RNS-II 45.3 § 1.45d 2700 § 51.6d 0.69 § 0.04 0.6 § 0.019 1.29 § 0.029a
VNS-III RNS-I 81.7 § 1.5a 5263 § 30.7a 0.68 § 0.041 0.57 § 0.047 1.25 § 0.067a
RNS-II 73.3 § 2.72b 4497 § 48.5bc 0.7 § 0.039 0.54 § 0.064 1.24 § 0.017a
Statistical significance (p-values) .0006 <.0001 ns ns .022
y
VNS: Vegetative Nutrient Solution; yyRNS: Reproductive Nutrient Solution; yyy: Data are shown as the means of three replicates § S.D. Values followed by same letters in a column are not significantly different
according to Duncan tests at 5% level; ns: no significant difference.
JOURNAL OF PLANT NUTRITION
5
 6
R. SHIRKO ET AL.

Table 3. Effect of nutrient solution formula on mineral uptake in strawberry ‘Paros.’

Growth stages N (%) P (%) K (%) Ca (%) Mg (%) Fe (mg kg¡1) Cu (mg kg¡1) Zn (mg kg¡1)

Vegetative
VNS-Iy 1.87 § 0.048ayyy 0.35 § 0.014 1.57 § 0.076b 1.34 § 0.088b 0.27 § 0.011b 65.7 § 1.9 9.71 § 0.22 21.5 § 0.47a
VNS-II 1.79 § 0.048b 0.34 § 0.011 1.78 § 0.063a 1.46 § 0.076b 0.26 § 0.024b 62.8 § 1.79 9.49 § 0.14 20. 2 § 0.21b
VNS-III 1.94 § 0.037a 0.32 § 0.005 1.80 § 0.03a 1.72 § 0.071a 0.33 § 0.022a 66.2 § 2.01 9.36 § 0.31 21.3 § 0.56a
Statistical significance (p-values) 0.0046 ns 0.0007 0.005 0.054 ns ns 0.0007
Reproductive
RNS-Iyy 1.81 § 0.047 0.32 § 0.005 1.7 § 0.027 1.52 § 0.082 0.3 § 0.024 63.9 § 1.55 9.51 § 0.16 21.1 § 0.37
RNS-II 1.91 § 0.023 0.35 § 0.01 1.73 § 0.08 1.5 § 0.084 0.28 § 0.01 65.8 § 1.56 9.53 § 0.22 21 § 0.43
Statistical significance (p-values) 0.011 0.0067 ns ns ns ns ns Ns
Vegetative £ Reproductive
VNS-I RNS-I 1.79 § 0.023c 0.32 § 0.006 1.71 § 0.057bc 1.44 § 0.06bc 0.28 § 0.02 68.4 § 2.05a 9.6 § 0.18 22.5 § 0.28a
RNS-II 1.95 § 0.068a 0.38 § 0.006 1.43 § 0.077d 1.25 § 0.16c 0.26 § 0.01 63.0 § 2.53ab 9.8 § 0.46 20.6 § 0.39b
VNS-II RNS-I 1.68 § 0.024c 0.34 § 0.005 1.65 § 0.052c 1.31 § 0.05c 0.25 § 0.046 60.6 § 0.74b 9.3 § 0.17 20.6 § 0.21b
RNS-II 1.89 § 0.017ab 0.35 § 0.023 1.91 § 0.03a 1.62 § 0.06ab 0.28 § 0.026 65.0 § 3.27ab 9.7 § 0.17 19.9 § 0.24b
VNS-III RNS-I 1.97 § 0.07a 0.32 § 0.008 1.74 § 0.032bc 1.82 § 0.07a 0.37 § 0.03 62.8 § 2.86ab 9.6 § 0.46 20.1 § 0.09b
RNS-II 1.89 § 0.017ab 0.33 § 0.006 1.86 § 0.012ab 1.62 § 0.1ab 0.30 § 0.009 69.6 § 0.73a 9.1 § 0.47 22.6 § 0.26a
Statistical significance (p-values) .0024 ns .0003 .029 ns .044 ns <.0001
y
VNS: Vegetative Nutrient Solution; yyRNS: Reproductive Nutrient Solution; yyy: Data are shown as the means of six replicates § S.D. Values followed by same letters in a column are not significantly different
according to Duncan tests at 5% level; ns: no significant difference.
 JOURNAL OF PLANT NUTRITION 7

Figure 1. Relationship between total chlorophyll and leaf nitrogen content (A); relationship between total chlorophyll and potassium
and calcium leaf in the strawberry ‘Paros’ (B); Z: Pearson correlation coefficients, (N D 18).

High N uptake increased the number and leaf area as the base compartment of photosynthesis. In
the absence of enough N, situation will influence photosynthesis reversely (Macit et al. 2007). Thus,
high levels of N, due to improved photosynthesis components such as photosynthetic pigments,
increase the rate of photosynthesis products, which ultimately affects vegetative parameters such as the
number and leaf area (Novoa and Loomis 1981). Probably the biosynthesis of many enzymes and plant
sugars directly depends on the plant leaves (Turhan and Eris 2005). There are many low molecular
weight N organic compounds which are essential for cellular metabolism of vitamins, cofactors, hor-
mones, pigments chlorophyll, and phytochrome (Marschner 2011).

Nutrient uptake
The highest and lowest absorption of N in leaves were resulted in VNS-III £ RNS-I: 1.97 § 0.07 and
VNS-II £ RNS-I: 1.68 § 0.024 solutions, respectively (Table 3).
The content of P in leaves (PL) was not significantly affected by interaction effects of treatments;
hence, the highest content of PL was in VNS-I £ RNS-II (Table 3).
The highest absorption of K, Ca, and Mg was in VNS-III formulation with cations ratio of K:Ca:Mg
[1:1.17:0.75]. The highest amount of KC uptake was recorded in interaction effect of VNS-II £ RNS-II
and VNS-III £ RNS-II solutions, whereas KC absorption was decreased as concentration of minerals
at reproductive stage was increased. Further, the highest Ca2C uptake was observed in VNS-III £
RNS-I (Table 3).
N is the chief among minerals in plant nutrition and its deficiency is considered as one of the limiting
factors for growth and development. N deficiency may occur in soilless culture with cocofibre medium or
closed systems. A lack of N in leaves negatively affects fruit size, quality, and yield. The deficiency level of
N in strawberry leaf is about 1.8% (Morgan 2006). The excess amount of N in strawberry leaf promotes
vegetative growth and delays flower induction (Sharma and Shyan 2009). Castellanos-Moralles et al.
(2010) reported that the level of applied N in nutrient solution had a significant effect on TA, glucose,
and fructose concentrations as well as Brix grade in strawberry fruits “Aromas,” where 6 mm.L¡1 N level
produced more acidic fruits with higher levels of glucose and fructose compared to 3 mm.L¡1 N level.
K is involved in almost all metabolic processes of plants. K plays an important role in photosynthe-
sis, carbohydrate production, nitrate reduction, ammonium ions consumption in amino acids struc-
ture, and protein synthesis. K is vital to maintain the cell turgor pressure. Plants deficient in K produce
low dry matter due to accelerate respiration rate. K strengthens root growth and plays a role in cell
division (Barker and Pilbeam 2015). In strawberry fruits, K is highly needed in order to ensure the fruit
size, quality, and taste as the main indexes of productivity and marketability. Deficiency level of K in
strawberry leaf is about 1% and ideal range is 1.2–2% (Morgan 2006).
8 R. SHIRKO ET AL.

Phosphorus deficiency is not very common in hydroponic systems compared to N and K deficien-
cies; however, it can greatly reduce the yield and plant growth. Phosphorus deficiency causes a decrease
in cell division, purple old leaves, and increase the anthocyanin (Marschner 2011).
The highest content of Fe in leaf (FeL) (69.6 § 0.73 mg kg¡1) was in VNS-III £ RNS-III formulation
with the highest concentration of minerals, which was not significantly different compared to the VNS-
I £ RNS-I with lowest concentration of minerals (Table 3). Generally, Fe absorption is directly related
to the amount of cations and anions in nutrient solution. Further, Fe involves in antioxidant enzymes
such as catalase (CAT) and peroxidase (POD).
The absorption of Cu2C was not significantly affected by nutrient formula, but Cu2C showed a dif-
ferent reaction to nutrients concentration during vegetative and reproductive stages. The highest
absorption of Cu2C at vegetative and reproductive stages was in VNS-I and RNS-II, respectively
(Table 3).
The highest amount of absorbed ZnC2 was in VNS-III £ RNS-III (Table 3). Zn is the activator of
many enzymes and is involved in alcohol dehydrogenase and superoxide dismutase structure (Mitra
2015). Despite the constant concentration of micronutrients in nutrient solution, changes in the
macro-elements, particularly cations such as KC and Ca2C in different formulas during the vegetative
and reproductive stages, have affected the absorption of microelements (Table 3). Therefore, the bal-
ance of elements in nutrient solution is a determining factor in mineral uptake and productivity, espe-
cially in the case of Fe and Cu, which are involved in respiratory and photosynthetic electron transport
chain cytochromes (Marschner 2011).

Fruit yield
Strawberry yield, as the most important indicator of strawberry production, is the result of balancing
the relative contribution of yield components (e.g., genotype, management, and environmental condi-
tions). Fruit yield was significantly affected by different nutrition formulas during vegetative and repro-
ductive stages (p < .05). The highest yield (678 § 30.62, 656.3 § 13.7, and 634 § 26.5) was in VNS-I £
RNS-I, VNS-I £ RNS-II, and VNS-III £ RNS-I, respectively (Figure 2A). According to the main
results, concentration and balanced ratio of nutrients in VNS-I formula during vegetative stage, and
RNS-I and RNS-II during reproductive stage are more appropriate formulas for producing high yield
of strawberry ‘Paros’ (Figure 2A).
Balanced nutrition, in particular the solution formula, at vegetative and reproductive stages
increased the number of flowers, fruits, fruit set, and ultimately increased fruit yield (Figure 2A and

Figure 2. Effect of nutrient solution formula on yield (A) and yield attributes (flower number, fruit number, and fruit set) (B) of straw-
berry ‘Paros’; Vertical bars represent §standard error (n D 3).
 JOURNAL OF PLANT NUTRITION 9

2B). The moderate level of N in solution, balanced the absorption of minerals and increased yield
(Latef and Chaoxing 2011). Continued increase in minerals concentration including N and increase in
solution EC, did not affect fruit yield positively. Kirimi et al. (2011) reported that increases in N con-
centrations reduced the tomato fruit yield. N is a key element in the growth and production of plants,
but excess use of this element causes adverse effects on plant growth (Zhao et al. 2005) and accelerates
sensitivity of plants to pests and diseases.
Total yield of plant depends on growth and development conditions during plant lifecycle. Hence,
the role of minerals and their concentration is determinative. Determinative role of minerals in plant
productivity, including N (Zhao et al. 2005), P (Barker and Pilbeam 2015), K (Husain et al. 2014), Ca,
B (W ojcik and Lewandowski 2003), Zn (Gill and Tuteja 2011), and Fe (Zelitch 2012) have been
reported. Therefore, proper plant nutrition is an important factor in improving the productivity.

Fruit quality
The balance of ions in nutrient solution is an effective strategy to achieve high quality production. In
this study, fruit quality including TSS, TA, TSS/TA, vitamin C, and post-harvest quality (firmness and
shelf life) were significantly affected by treatments. The highest TSS/TA ratio as the fruit taste index
affected by ratio of sugar (TSS) and acidity (TA) was produced in VNS-I £ RNS-II and VNS-III £
RNS-I solutions (Table 4). Increase in content of TSS happens as fruit development progress. Accord-
ing to Shaw et al. (1998), TSS, is more dependent on environmental conditions during production and
less on heredity and genetics of plant. According to (Centeno et al. (2011), high soluble solids in
tomato fruits produces high conversion of starch to sugar. N increased TSS, which are likely to produce
enzymes and proteins effective in respiration process which leads to increase in assimilates content.
The effect of N fertilization on TA and sugar content of fruit are inconsistent and vary from year to
year (Haynes and Goh 1987).
Fertilizer consumption increases respiration and reduces plant organic acids. Additional acids accu-
mulate in vacuole. Acid accumulation is under close control of genes (Causse et al. 2002); however,
nutrition also plays an important role in organic acids and sugar variation. Citric acid, malic acid, and
ascorbic acid are the main organic acids in strawberry (Pelayo et al. 2003). Ratio of soluble solids to
TA is considered as an indicator of strawberry fruit ripening (Kim et al. 2003). Furthermore, Keutgen
and Pawelzik (2007), in an experiment on strawberry “Elsanta,” found that fruit soluble solids
decreased and protein increased in salinity stress conditions. TSS and/or sugar to acid ratio determine
(s) strawberry fruit taste. Sugars, organic acids, and soluble pectins are the main constituents of TSS in
strawberry. Soluble solids are qualitative indices that are directly related to quality of edible fruits and
most of the consumers tend to ripe fruits with high soluble solids. There is a linear relation between
total sugar and TSS (brix). Various strawberry varieties show similar treatment regarding changes dur-
ing different developmental stages.
Furthermore, fruit proteins were significantly affected by treatments (p < .05). There was no signifi-
cant effect on leaf proteins composition as applied by reproductive formula, whereas RNS-I £ VNS-I
formulation that had the lowest concentration of nutrient solution produced the highest amount of
leaf proteins (Table 4).
Vitamin C content was significantly affected by treatments. The highest content of vitamin C was
produced in VNS-I formulation and RNS-1 (Figure 3A and 3B). Probably, low EC, in particular, low N
content in VNS-I formulation (Table 1) has positively affected the vitamin C content.
Ascorbic acid (vitamin C) is an ubiquitous molecule in plants. It functions as an antioxidant, an
enzyme cofactor (functions in photosynthesis), and as a precursor for oxalate and tartrate synthesis
(Scartezzini et al. 2006). Strawberry is rich in vitamin C, which represents antioxidant activities
(Meyers et al. 2003). The content of vitamin C increases as strawberry ripens (Nunes et al. 2006). Fur-
ther, Fe and Zn as well as their interaction effect, increased the vitamin C content in strawberry fruit.
Vitamin C reduces under application of N fertilizers, which may be related to change in cell size and
subsequent accumulation of assimilates. In this condition, plant starts to absorb water in order to keep
the turgor pressure, which results in relative dilution effect (Lee and Kader 2000). Moreover Haynes
 10
R. SHIRKO ET AL.

Table 4. Effect of nutrient solution formula on fruit quality and protein content of strawberry ‘Paros’.

Growth stages Total soluble solid (%) Titrable acidity (mg citric acid 100 g¡1 Fw) TSS/TA (%) Fruit protein (mg g¡1 Fw) Leaf protein (mg g¡1 Fw)

Vegetative
VNS-Iy 9.1 § 0.1ayyy 0.95 § 0.03b 9.7 § 0.23a 0.40 § 0.008 0.43 § 0.013a
VNS-II 8.0 § 0.28b 1.01 § 0.02a 7.9 § 0.38b 0.39 § 0.013 0.39 § 0.008b
VNS-III 8.2 § 0.18b 0.90 § 0.03b 9.2 § 0.28a 0.38 § 0.008 0.40 § 0.012b
Statistical significance (p-values) .006 .002 .001 ns .001
Reproductive
RNS-Iyy 8.4 § 0.21 0.96 § 0.035 8.8 § 0.32 0.40 § 0.004 0.41 § 0.012
RNS-II 8.5 § 0.26 0.95 § 0.017 9.0 § 0.38 0.37 § 0.009 0.40 § 0.009
Statistical significance (p-values) ns ns ns 0.03 ns
Vegetative £ Reproductive
VNS-I RNS-I 9.2 § 0.17 1.003 § 0.028a 9.2 § 0.09ab 0.41 § 0.005 0.45 § 0.009a
RNS-II 9.1 § 0.06 0.896 § 0.018b 10.2 § 0.18a 0.38 § 0.002 0.40 § 0.014b
VNS-II RNS-I 7.9 § 0.06 1.039 § 0.037a 7.6 § 0.234c 0.40 § 0.007 0.40 § 0.009b
RNS-II 8.1 § 0.61 0.992 § 0.018a 8.2 § 0.77bc 0.38 § 0.027 0.37 § 0.003c
VNS-III RNS-I 8.1 § 0.06 0.832 § 0.001b 9.7 § 0.069a 0.39 § 0.005 0.37 § 0.002c
RNS-II 8.3 § 0.38 0.97 § 0.021ba 8.6 § 0.25bc 0.36 § 0.011 0.42 § 0.007b
Statistical significance (p-values) ns .0006 .025 ns .0001
y
VNS: Vegetative Nutrient Solution; yyRNS: Reproductive Nutrient Solution; yyy: Data are shown as the means of three replicates § S.D. Values followed by same letters in a column are not significantly different
according to Duncan tests at 5% level; ns: no significant difference.
 JOURNAL OF PLANT NUTRITION 11

80 A 80 **
B
a†

Vitamin C (mg 100g-1 Fw)

ab

Vitamin C (mg 100g-1 Fw)

70 b 70
60 60
50 50
40 40
30 30
20 20
10 10
0 0
NS-I NS-II NS-III NS-I NS-II
Vegetative Growth Reproductive

Figure 3. Vitamin C content in strawberry fruit ‘Paros’ as affected by different nutrition formula during vegetative (A) and reproductive
(B) stages. Dissimilar letters and error bars indicate significant difference basis on Duncan’s multiple test range (p < .01) and (Mean §
SE, n D 6), respectively.

and Goh (1987) reported that increasing rates of N increases concentration of polyphenols and reduces
those of ascorbic acid. According to Mozafar (1993), N increased the amount of vitamin C.
Treatments affected strawberry ‘Paros’ fruit firmness as well as strawberry fruits shelf life signifi-
cantly. The most firmed fruits were produced in VNS-III £ RNS-I formulation (Figure 4A). Addi-
tionally, the highest shelf life of strawberry fruits ‘Paros’ were resulted in VNS-III £ RNS-I
formulation (Figure 4B). Probably the high concentration of minerals during vegetative growth
period, especially the high concentration of Ca2C and also balanced ratio of cations during reproduc-
tive stage, has affected fruit firmness and shelf life positively. In general, plants with low or moderate
vegetative growth, tend to have firmer fruits (Nestby et al. 2005). Fresh fruits production is still under
study in order to preserve their fresh-like quality during prolonged periods. Physical and chemical
changes may cause fruit softening. Conversion on protopectin to water soluble pectin decrease in cel-
lulose crystallinity and thinning of cell wall (Ward et al. 1989). Ca can interact with pectin polymers
to form a cross-linked polymer network that increases mechanical strength, subsequently decreasing
senescence process (Shafiee et al. 2010).

Figure 4. Fruit firmness (A) and shelf life (B) of strawberry ‘Paros’ as affected by different nutrition formula. Dissimilar letters and error
bars indicate significant difference basis on Duncan’s multiple test range (p < .01) and Mean § SE (n D 6), respectively; VNS: Vegeta-
tive Nutrient Solution; RNS: Reproductive Nutrient Solution.
12 R. SHIRKO ET AL.

Taking together and basis of obtained results, although the VNS-I £ RNS-I produced the highest
yield, however this parameter did not show a significant difference with VNS-III £ RNS-I formula,
which produced the firmest fruits. Further, VNS-III £ RNS-I is one of the formula that increased N
uptake and leaf number; thus the formula VNS-III £ RNS-I is proposed as the best among the studied
solutions for producing the highest yield in strawberry ‘Paros’ fruits with high quality attributes.

Acknowledgment
The authors would like to thank Julian Verdonk for his valuable comments and for proofreading this manuscript.

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