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EFFECTS OF CALCIUM AND SALINITY

STRESS ON QUALITY OF LETTUCE IN
SOILLESS CULTURE
a a a
Eva Borghesi , Giulia Carmassi , Maria C. Uguccioni , Paolo
a a
Vernieri & Fernando Malorgio
a
Dipartimento di Biologia delle Piante Agrarie, University of Pisa,
Pisa, Italy
Accepted author version posted online: 22 Aug 2012.Published
online: 28 Feb 2013.

To cite this article: Eva Borghesi , Giulia Carmassi , Maria C. Uguccioni , Paolo Vernieri & Fernando
Malorgio (2013) EFFECTS OF CALCIUM AND SALINITY STRESS ON QUALITY OF LETTUCE IN SOILLESS
CULTURE, Journal of Plant Nutrition, 36:5, 677-690, DOI: 10.1080/01904167.2012.721909

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 Journal of Plant Nutrition, 36:677–690, 2013
Copyright  C Taylor & Francis Group, LLC

ISSN: 0190-4167 print / 1532-4087 online

DOI: 10.1080/01904167.2012.721909

EFFECTS OF CALCIUM AND SALINITY STRESS ON QUALITY

OF LETTUCE IN SOILLESS CULTURE

Eva Borghesi, Giulia Carmassi, Maria C. Uguccioni, Paolo Vernieri,

and Fernando Malorgio
Downloaded by [University Library Utrecht] at 10:24 10 August 2013

Dipartimento di Biologia delle Piante Agrarie, University of Pisa, Pisa, Italy

2 Lettuce (Lactuca sativa L.) var. ‘Lollo rossa’ was grown in a floating hydroponic system. Six
saline treatments were used, adding different concentrations of calcium chloride (CaCl2 ) to the
nutrient solution (mol m −3): 0, 2.5, 5, 10, 15, 20, which, respectively, corresponds to an electrical
conductivity of 2.5, 3, 3.5, 4.4, 5.4, 6.3 dS m −1. In plants subjected to moderate salinity stress
the growth was not affected and yields were not statistically significant; however, the data shows
a slight decline in 20 mol m −3 CaCl2 treatments. Nitrate contents in the leaf decreased when the
concentration of CaCl2 in nutrient solutions increased. Other quality parameters were positively
affected by treatment with CaCl2 , especially at moderate concentrations such as 5 and 10 mol m −3.
In fact, both the content of phenols and antioxidant power increased at moderate salt concentrations
and reduced in the treatment with 20 mol m −3 CaCl2 . The content of chlorophyll and carotenoids
were not affected by treatment even at high salt concentrations. The use of CaCl2 did not result in
nutrition imbalances in plants either. In fact, the ratio K/Na remained unchanged in the various
treatments, as well as the contents of microelements.

Keywords: floating system, leafy lettuce, Lactuca sativa L., calcium chloride, antioxidant
capacity, ABA

INTRODUCTION
The effect of dietary factors on health status has been recognized since
antiquity (Shaidi, 2004).
Recognition of the importance to health of consuming more fruits
and vegetables has been heightened by recent strong evidence that cancer
rates could be reduced by 30% with higher intakes of vegetables and fruits
(Gray-Donanld, 2004). Lettuce is an important agricultural commodity avail-
able world-wide throughout the whole year (Rico et al., 2008). The lettuce

Received 12 May 2010; accepted 23 May 2011.

Address correspondence to Eva Borghesi, Dipartimento di Biologia delle Piante Agrarie, University
of Pisa Viale delle Piagge, 23, 56100 Pisa, Italy. E-mail: evaborghesi@libero.it

677
 678 E. Borghesi et al.

composition is a responsible for beneficial effects on health; in particular the

micronutrients such as polyphenoles and carotenoides are involved in many
antioxidant processes that play a role in the prevention of cancer and heart
disease (Hart and Scott, 1995; Szeto et al., 2004). Indeed, epidemiological
studies have suggested a link beetween the consumption of polyphenol-
rich foods or beverages and the prevention of these diseases (Scalbert and
Willansom, 2000).
In general, it possible that the consumption of fresh vegetal play the part
of defensive role against this pathology, primarily, because they contain an
elevated quantity of vegetal fiber and many secondary metabolites (Kaur and
Kapoor, 2001).
Green leafy vegetables are a major source of dietary carotenoids which
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can act as lipholic antioxidants and reduce the incidence of cataract and
macular degeneration (Moller et al., 2000).
Climatic conditions, like temperature or light, have a strong influence
on the concentration of bioactive compounds (Weston and Barth, 1997;
Lefsrud et al., 2005).
Cultural techniques, including nutrient and water availability, have an
influence on these compounds, in particular, a floating system can increase
food quality (Diaz et al., 2005), allowing to obtain first quality in hygienic
characteristics. A floating system also could be adopted to use the beneficial
effects of salinity on leafy vegetables in order to enhance quality character-
istics such as reduction of nitrate leaf content (Mengel and Kirkby, 1987;
Maynard et al., 1976; Gonnella et al., 2003). An excessive amount of nitrate
in edible plants may represent a potential risk to human health (Gangolli
et al., 1994). In fact, the content of nitrate (NO3 ) depends on environmental
factors (light, temperature), genetic factors (cultivar and species) and cul-
tural techniques (Gonnella et al., 2002b; Minotti and Stanley, 1973; Bloom,
2002; Malorgio et al., 1995).
Incrementing the nutritional value of the products by enrichment with
the nutritive substances useful for human health is a goal reachable using
hydroponic cultural techniques (Malorgio et al., 2009).
Positive effects of moderate salinity in fruit quality have been described
by numerous authors for horticultural soilless growing crops such as tomato
and muskmelon (Mavrogianopoulos et al., 1999; Cuartero and Fernandez-
Muñoz, 1999; De Pascale et al., 2001), but for hydroponically grown lettuce
crops results are rare in literature (Andriolo et al., 2005). However, sev-
eral studies indicated that a moderate salt stress might enhance tomato
carotenoids content and antioxidant capacity and, consequently, improve
their nutritional value (Petersen et al, 1998).
In general, salinity stress has a strong impact on the commercial yield of
horticultural crops grown hydroponically (Sonneveld et al, 1999). Salinity
affects the crop during vegetative and reproductive stages and therefore
 Effects of Calcium and Salinity Stress 679

causes reductions in both dry biomass and crop yield (Aslam et al., 1993).
Specific ion concentrations in plant tissues may increase or decrease due to
salinity stress, resulting in ion toxicity or nutrient deficiency (Bernstein et al.,
1974). These disorders may result from the effect of salinity on nutrient
availability, competitive uptake, transport or partitioning within the plant
(Grattan and Grieve, 1999). Elevated sodium chloride (NaCl) levels in the
root medium reduce the nutrient assimilation, especially of potassium (K)
and calcium (Ca), resulting in ion imbalances of K, Ca, and magnesium
(Mg) compared to sodium (Na), as well as in negative effects on enzymes
and membranes (Khan et al., 2000; Keutgen and Pawelzik, 2009).
Metabolic imbalances caused by ionic toxicity, osmotic stress and nutri-
tional deficiency under salinity may also lead to oxidative stress (Zhu, 2002).
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Furthermore, saline stress conditions cause a rapid production of hor-

mone abscisic acid (ABA) to help plant survival (Zhang et al., 2006).
With regards to salinity, lettuce is generally considered a moderately sen-
sitive crop although different authors report different levels of salt tolerance,
probably depending on genotype, plant age and experimental conditions
(Cramer and Spurr, 1986; De Pascale and Barbieri, 1995; Tesi et al., 2003).
Calcium has been shown to ameliorate the adverse effects of salinity on
plants (Reiss and Beale, 1996). For example, Ca additions to soils or as foliar
sprays can sometimes correct disorders caused by Na-induced Ca deficiencies
(Grattan and Grieve, 1999).
Calcium plays an essential role in plant development and overall plant
health because it is a structural component of cell walls and it is necessary
for cell growth and division. In lettuce an increase of calcium in the leaf
tissues can increase photosynthetic capacity and also chlorophyll synthesis
(Reiss and Beale, 1996; Fallovo et al., 2009). In addition, increasing the cal-
cium content in the leafy vegetables could further improve their nutritional
benefits to the consumers considering that calcium is the mineral nutrient
most commonly deficient in modern diets (Grusak, 2002).
The goal of this work was to determine the effect of calcium salinity on
yield and quality of hydroponically grown lettuce plants. In particularly, the
objective of this work was to assess the effect of calcium chloride on leaf
quality characteristics.

MATERIALS AND METHODS

Plant Material and Growing System
The experiment was conducted in a heated greenhouse located in Pisa,
Italy (lat. 43◦ 40 N), on lettuce grown in hydroponic cultures.
Lettuce (Lactuca sativa L.) var. ‘Lollo rossa’ was grown in a floating hy-
droponic system during the spring 2008 (May–June). The seeds were sown
 680 E. Borghesi et al.

in 260-cell plug-trays measuring 0.32 × 0.54 m, and filled with perlite and
vermiculite. After emergence all trays containing seedlings were placed in
floating tanks (volume = 60 L), with the following nutrient solution com-
position (mol m−3): 7.0 nitrogen (N)-NO3 , 7.0 N-ammonium (NH4 ), 1.3
phosphorus (P), 7.4 K, 5.0 Ca, 1.2 Mg, 8.0 chloride (Cl), 9.0 Na. Micronu-
trients were added at Hoagland’s concentration [in µmol L−1: 40 boron
(B), 40 iron (Fe), 1 copper (Cu), 5 zinc (Zn), 10 manganese (Mn)]. Well-
water used for the formulation of the nutrient solution was already very hard
with a Na concentration of 9 mol m−3, Ca concentration of 1 mol m−3, Mg
concentration of 1 mol m−3, and Cl concentration of 8 mol m−3.
Six saline treatments were used in the test, adding different concentra-
tions of CaCl2 (mol m−3) to nutrient solution: 0, 2.5, 5, 10, 15, 20, which
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corresponds to the respective electrical conductivity (EC) of 2.5, 3, 3.5, 4.4,

5.4, 6.3 dS m−1.
A complete randomized block experimental design was adopted with
three replicates for each treatment.
The oxygenation of the nutrient solution was obtained by insufflating
air with air pumps in every tank.

Measurements
When leaves reached commercial maturity, i.e. 6–10 cm in length, plants
of each replicate were harvested, and the fresh weight (FW) of leaf material
was determined. Plant yield was expressed as leaf biomass produced per
square meter (kg m−2).
After harvesting chlorophyll, carotenoids, anthocyanins, and phenolic
compounds were measured in the lettuce leaves.

Total Chlorophyll and Carotenoid Contents

Total chlorophyll and carotenoids were extracted using methanol 99.9%
as solvent. Samples were kept in a dark cold room at 4◦ C for 24 hours.
Quantitative determinations were carried out immediately after extraction.
Absorbance readings were measured at 665.2 and 652.4 nm for chlorophyll
pigments and 470 nm for total carotenoids. Chlorophyll and carotenoid
concentrations were calculated by Lichtenthaler’s formula (Lichtenthaler,
1987).

Anthocyanin and Polyphenol Contents

Anthocyanin and polyphenol content were determined spectrophoto-
metrically with a Perkin Elmer Lambda UV 35, UV-Vis spectrophotometer
(Perkin Elmer, Beaconsfield, UK). Samples of the frozen leaf tissue (100 mg)
were ground in pre-chilled mortar and were extracted into methanolic
 Effects of Calcium and Salinity Stress 681

hydrochloric acid (HCl; 1%). Samples were incubated overnight at 4◦ C in

darkness. The concentration of cyanidin-3-glucoside equivalents was deter-
mined spectrophotometrically at 535 nm (Kho et al., 1977). The concentra-
tion of gallic acid equivalents was determined at 320 nm (Ke and Saltveit,
1989).

Nitrate Content
Nitrate content was measured with a Shimadzu UV-Vis 1204 spectropho-
tometer (Shimadzu, Tokyo, Japan) using the salicylic-sulfuric acid method
(Cataldo et al., 1975). Each sample (100 mg of DW) was extracted with
30 mL of distilled water for two hours at room temperature, under continu-
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ous shaking, then centrifuged at 4000 rpm for 15 min. The supernatant was
recovered and 200 µl were added to 800 µL of 5% salicylic acid in sulfuric
acid, then 30 mL of sodium hydroxide (NaOH) 1.5 N were slowly added.
The spectrophotometer readings were taken at 410 nm.

Antioxidant Capacity
Antioxidant capacity was measured using the ferric-reducing antioxidant
power (FRAP) method developed by Benzie and Strain (1996). The samples
were extracted using methanol 99.9%. A small quantity of the methanol
extract (0.1 mL) was added to 0.9 ml of FRAP reagent and mixed. The
mixture was then kept at 20◦ C for four minutes, and the absorbance was
measured at 593 nm. The FRAP reagent consisted of 1 mM 2,4,6-tripyridyl-2-
triazine (TPTZ) and 2 mM ferric chloride in 0.25 M sodium acetate (pH 3.6).

ABA Content
ABA was determined by an indirect ELISA based on the use of DBPA1
monoclonal antibody, raised against S(+)-ABA (Vernieri et al., 1989).
The ELISA was performed according to the method described by Walker-
Simmons (1987), with minor modifications.
Leaf samples (100 mg FW) were collected, weighted, frozen in liquid
nitrogen, then stored at -80◦ C until analysis. ABA was measured after ex-
traction in distilled water (water:tissue ratio = 10:1 v:w) overnight at 4◦ C.
Plates were coated with 200 µL per well ABA-4 BSA conjugate and incu-
bated overnight at 4ºC, then washed three times with 75 mM PBS buffer,
pH 7.0, containing 1 g L−1 BSA and 1 mL L−1 Tween 20, keeping the third
washing solution for 30 min at 37◦ C. Then 100 µL ABA standard solution or
sample and, subsequently, 100 µL DBPA1 solution (lyophilized cell culture
medium diluted in PBS buffer containing 10 g L−1 BSA and 0.5 mL L−1
Tween 20, at a final concentration of 50 µg mL−1) were added to each
well and competition allowed to occur at 37◦ C for 30 min. Plates were then
 682 E. Borghesi et al.

washed again as described above and 200 µL per well of secondary antibody
(alkaline phosphatase-conjugated rabbit anti-mouse (Sigma Aldrich, Milan,
Italy) in PBS buffer containing 10 g L−1 BSA and 0.5 mL L−1 Tween 20,
at a final dilution of 1:2000) was added and incubated for 30 min at 37◦ C.
Plates were washed again and 200 µL per well p-nitrophenyl phosphate were
added and incubated for 30 min at 37◦ C. Absorbance readings at 415 nm
were obtained using a MDL 680 Perkin-Elmer microplate reader. For each
treatment, four independent samples were assayed in triplicate.

Macronutrient and Micronutrient Determination

Leaves were oven dried for three days in a 70◦ C. The oven-dried-ground
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samples were wet ashed in a mixture of nitric-perchloric acids (HNO3 :

HClO4 ).
Concentration of K, Na, Ca, Mg, Fe, Mn, Zn, and Cu was estimated by
atomic absorption spectrophotometry (Model Varian AA240 FS, Varian, Palo
Alto, CA, USA).

Statistical Analysis
The experimental design was completely randomized. Data were sub-
jected to one-way analysis of variance (ANOVA). The means were separated
using the least significant difference (LSD) test for P = 0.05 or P = 0.01.

RESULTS AND DISCUSSION

The lettuce was harvested after 5 weeks from the sowing when the plants
were high as 6–10 cm. The yield was influenced from the saline treatments
(Table 1). Indeed when the plants were subjected to moderate salinity stress
the growth was not affected and in the yields there were not significant
statistic differences (Table 1) but the data shows a slight decline in 20 mol
m−3 CaCl2 treatments, (3227 g m−2 in 0 mol m−3 CaCl2 and 2200 g m−2 in

TABLE 1 Yield and percent dried weight of Lactuva sativa var. ‘Lollo rossa’ cultivated in floating system
with nutrient solution with diferrent CaCl2 concentration (0, 2.5, 5, 10, 15, 20 mol m−3). Values are
means (n = 3). Data were subjected to one-way ANOVA and different letters in the same column mean
that values are statistically different P < 0.05

Treatment (CaCl2 mol m−3) EC (dS m−1) Yield (g m−2 FW) Dried weight (g kg−1)

0 2.5 3227 a 3.05 a

2.5 3 3405 a 3.30 a
5 3.5 3624 a 2.80 a
10 4.4 3255 a 2.90 a
15 5.4 3209 a 3.60 a
20 6.3 2200 b 3.80 a
 Effects of Calcium and Salinity Stress 683
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FIGURE 1 ABA content (ng g−1 FW) in leaves of Lactuca sativa L. var. ‘Lollo rossa’ cultivated in floating
system with nutrient solutions with different CaCl2 concentrations (0, 2.5, 5, 10, 15, 20 mol m−3). Values
are means with standard errors (n = 3). Data were subjected to one-way ANOVA and different letters
mean that values are statistically different P < 0.05.

20 mol m−3 CaCl2 treatments). Also the percentage of dried weight did not
result significantly different (Table 1).
The ABA content in the lettuce leaf showed that the plants cultivated
with nutrient solutions with an EC up till 4 dS m−1 were not stressed by
treatments with CaCl2 (Figure 1). On the contrary, the contents of ABA in
the treatments 15 and 20 mol m−3 CaCl2 were higher than in the treatments
with 0 mol m−3 (1483.2 to the confront 613.7 ng g−1 FW). Exposure of plants
to salinity is known to induce a proportional increase in ABA concentration
that is, in most cases, correlated with leaf or soil water potential (Zhang et al.,
2006). Salinity up-regulates the biosynthesis of the plant stress hormone ABA
(Jia et al., 2002; Xiong and Zhu, 2003).
Nitrate content in the leaves decreased when the concentration of
CaCl2 in nutrient solutions was increased (Figure 2). Nitrate content of
the treatment 0 mol m−3 CaCl2 was double compared to the treatments
with salinity; nitrate values varied from 1010 mg NO3 kg−1 FW in the con-
trol to 484.75 mg NO3 kg−1 FW in the leaf of the plants cultivated with
20 mol m−3 CaCl2 (Figure 2). Starting from the 5 mol m−3 CaCl2 treat-
ments (corresponding to an electric conductibility of 3.4 dS m−1, i.e. a mod-
erate salinity), the nitrate content values were reduced but the reduction
stayed constant even for increased concentrations of CaCl2 in the nutrient
solution.
The reduction could be caused by the substitution of NO3 by Cl in the
vacuole-like osmolite (Blom-Zandstra and Lampe, 1983). Kafkafi et al. (1992)
found that Cl from CaCl2 and not potassium chloride (KCl), inhibited NO3
uptake in melon and tomato and concluded that the effects of NaCl and KCl
are similar, but that nitrate inhibition by Cl is much more pronounced at
 684 E. Borghesi et al.
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FIGURE 2 Nitrate content in leaves of Lactuca sativa L. var. ‘Lollo rossa’ cultivated in floating system
with nutrient solutions with different CaCl2 concentrations (0, 2.5, 5, 10, 15, 20 mol m−3). Values are
means with standard errors (n = 3). Data were subjected to one-way ANOVA and different letters mean
that values are statistically different P < 0.05.

the lower salinity range if the counter ion were Ca rather than a monovalent
cation.
The nitrate values decreased in accord with Gonnella et al. (2002a) in
celery plants cultivated in floating system with the substitution of the nutrient
solution in the last three days of cultivation with a CaCl2 solution.
Nevertheless, the values of nitrate were always lower than the maximum
level permitted by European legislation (Commission Regulation, EC No.
188/2006; European Union, Brussels).
Table 2 shows the mean values of total chlorophyll, total carotenoids,
phenols and anthocyans contents in the leaf of lettuce as a function of the
salinity treatments. No significant difference among treatments was observed
for chlorophyll total and carotenoids. It is reported in the literature that one
of the major factors inducing leaf senescence is the decrease of chlorophyll
under saline conditions (Chen et al., 1991). In experiments on lettuce with

TABLE 2 Contents of chlorophyll, carotenoids, phenols and anthocyans in the leaves of Lactuca sativa
var. ‘Lollo rossa’ cultivated in floating system with diferrent CaCl2 concentration (0, 2.5, 5, 10, 15,
20 mol m−3). Values are means (n = 3). Data were subjected to one-way ANOVA and different letters in
the same column mean that values are statistically different P < 0.05

Treatment CaCl2 EC Chlorophyll Carotenoids Phenols Anthocyans

(mol m−3) (dS m−1) (g kg−1 FW) (g kg−1 FW) (g kg−1 FW) (g kg−1 FW)

40 2.5 1.01 b 0.83 a 97.50 c 0.12 ab

2.5 3 0.93 b 1.13 a 126.71 abc 0.19 a
5 3.5 0.95 b 1.10 a 137.46 ab 0.15 ab
10 4.4 1.07 b 1.00 a 158.90 a 0.22 a
15 5.4 1.28 a 1.13 a 151.60 a 0.22 a
20 6.3 0.89 b 0.80 a 112.22 bc 0.08 b
 Effects of Calcium and Salinity Stress 685
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FIGURE 3 Power antioxidant ferring reducing (Frap) of leaf of Lactuca sativa L. var. ‘Lollo rossa’
cultivated in nutrient solutions with different CaCl2 concentrations (0, 2.5, 5, 10, 15, 20 mol m−3). Values
are means with standard errors (n = 3). Data were subjected to one-way ANOVA and different letters
mean that values are statistically different P < 0.05.

salinity stress, the chlorophyll of the leaves started to reduce at 60 mol m−3
salinity concentration (Kaya et al., 2002). The same results for chlorophyll
contents could be attributable to the effects mitigated by Ca on the effects
of high salt on chlorophyll content.
The phenol content increased with the salinity treatments, in particular
with moderate salinity. In fact, the higher values were observed in treatments
with 10 and 15 mol m−3 CaCl2 (158.9 µg g−1 FW) but in the treatments with
an EC value of 6.4 dS m−1 the values of phenols decreased (112.22 µg
g−1FW).
The same trend was observed in FRAP values (Figure 3), which reached
a peak in antioxidants in the range of 2.5 to 15 mol m−3of CaCl2 treatments
(63.62 and 60.68 µM Fe2+g−1 FW at 2.5 and 15 mol m−3of CaCl2 ). While
in the 20 mol m−3 treatments, where the EC value was higher, antioxidants
decreased (42.62 µM Fe2+g−1 FW).
The anthocyanin totals declined in the treatments with high contents
of CaCl2 in the nutrient solution but in the other treatments there were no
statistically significant differences (Table 2).
An increase of compounds resulting from secondary metabolism is due to
the capacity of the plant to adapt itself to a moderate saline stress (D’Amico
et al., 2003). It is known that environmental factors and agronomic prac-
tices such as irrigation, fertilization, and salt stress, can affect both PPO
(polyphenol oxidases) and POD (peroxidases) activities. The effect is usu-
ally an increase in both activities and the appearance of specific isoform
and/or isoenzymes in response to stress situations (Tomás-Barberán and
Espı́n, 2001). Moreover, Frohmmeyer et al. (1997) demonstrated in parsley
 686 E. Borghesi et al.

TABLE 3 Macro- and microelements in the leaf of Lactuca sativa L. var. ‘Lollo rossa’ cultivated in a
floating system with diferrent CaCl2 concentrations (0, 2.5, 5, 10, 15, 20 mol m−3). Values are means
(n = 3). Data were subjected to one-way ANOVA and different letters in the same column mean that
values are statistically different P < 0.05. ns = not significant

Macro- or micronutrient (mg kg−1 DW)

Treatment
(CaCl2 mol m−3) Ca Mg K Na Cu Mn Fe Zn

0 10905 b 11694 b 55223 n.s. 8948 n.s. 38 a 310 a 2346 ab 189 n.s.
2.5 10591 b 9122 bcd 61388 8252 21 b 151 b 1693 bc 158
5 11682 b 9716 bc 65354 7906 16 bc 148 b 1853 bc 155
10 11099 b 6403 d 81435 5847 26 c 104 b 1024 c 132
15 11500 b 8399 cd 69705 6113 8 bc 93 b 1606 bc 149
20 53648 a 17481 a 66562 6102 8c 107 b 3216 a 132
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protoplasts that calcium is one of the signaling pathways of chalcone synthase

and PAL used by UV light.
In grapes, it was observed that anthocyanin accumulation and phenolic
metabolism (PAL) were induced by betains and calcium in soil (Yokotsuka
et al., 1999). In addition, the presence of calcium chloride could aid in
maintaining membrane integrity in some crops (Kukura et al., 1998).
Calcium concentration in lettuce leaves increased in response to higher
Ca concentrations in the hydroponic solution, and indeed at 20 mol m−3
of CaCl2 the concentration was five times higher than the concentration at
15 mol m−3 (Table 3); whereas in the other treatments there were not any
significant differences. Concentrations of Cu and Mn significantly decreased
with high salinity concentrations in the nutrient solution (Table 3). In saline
and sodic soils, the availability of micronutrients (Cu and Zn) is particularly
low and it often causes nutrient deficiencies (De Pascale and Barbieri, 1997;
De Pascale et al., 2001). Copper concentrations decreased in maize grown
under saline stress conditions in both soil and solution cultures (Rahman
et al., 1993; Izzo et al., 1991).
The Mg concentration in the lettuce leaves was higher in the treatments
with high EC values (Table 3). These results are in contrast to those reported
from other authors. In fact in the literature, it was indicated that high con-
centrations of Ca in the substrate often result in increased leaf Ca along
with a marked reduction in leaf Mg; it has been reported often that Ca also
induced Mg deficiency in sesame crops (Nassery et al., 1979; Bernstein et al.,
1974).
Iron concentrations were higher in the treatments with higher concen-
trations of CaCl2 but in the other treatments there were no significant dif-
ferences (Table 3). Reports on the influence of salinity on the iron con-
centration in plants are as inconsistent as those that concern Zn and Mn
concentrations. Salinity increased the Fe concentration in the shoots of pea,
tomato, soybean, and decreased its concentration in the shoots of barley and
 Effects of Calcium and Salinity Stress 687

corn (Dahiya and Singh, 1976; Hassan et al., 1970; Maas et al., 1972; Grattan
and Grieve, 1999).
The contents of K and Na in the leaf were similar for all treatments
(Table 3). The K/Na ratio did not change when the electric conductivity
increased. In general, Na induced K deficiency in growth and yield reduc-
tions of various crops and also affected the uptake of K in a competitive
process. The presence of Ca in the substrate influences the K/Na selectivity
by shifting the uptake ratio in favour of K at the expense of Na (Grattan and
Grieve, 1999).

CONCLUSION
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The cultivation of lettuce (Lactuva sativa L. var ‘Lollo rossa’) with a float-
ing system and use of moderate salinity stresses supplied by calcium chloride
was not observed to effect the yield. However, it did influence characteristics
like phenol and antioxidant contents, in particular when the electric conduc-
tivity of the nutrient solution was lower than 6.4 dS m−1. The use of salinity
could increase another important quality factor for the lettuce, namely the
reduction of nitrate contents in leaves. These results suggest that the use of
Ca in the nutrient solution could be a method to reduce some physiological
unbalances due to salinity such as the uptake of micro- and macroelements.

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