Plant Growth Regulation 30: 37–47, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

37

Modification of the response of saline stressed tomato plants by the

correction of cation disorders

M. Carvajal, A. Cerdá & V. Martı́nez

Dpto. Fisiologı́a y Nutrición Vegetal. Centro de Edafologı́a y Biologı́a Aplicada del Segura. CSIC. P.O. Box 4195,
30080 Murcia, Spain

Received 14 April 1999; accepted in revised form 26 July 1999

Key words: glucose phosphate synthase, invertase, mineral nutrition, salinity, water relations

Abstract
The interactions between NaCl and other major nutrients have been generally observed in plants. Decreases of
nutrient uptake under saline conditions normally appear in tomato plants grown under saline conditions. In this
work, the effect of increased external Ca, K and Mg concentrations under saline conditions has been investigated.
Tomato plants (Lycopersicon esculentum, Mill) were grown in a greenhouse, in 120 L capacity containers, filled
with continuously aerated Hoagland nutrient solution. Treatments were added to observe the combined effect of
two NaCl levels (30 and 60 mM) and three levels of Ca, K and Mg (in mM ratios of 4:6:1, 7:9:2 and 10:12:3;
treatments C1 , C2 and C3 respectively) on growth, fruit yield and water relations. Saline treatments decreased the
growth, which was partly restored with the C2 treatment and totally with the C3 treatment. A good association
was observed between the electric conductivity of the medium and the water or osmotic potential of the leaves,
independent of the type of treatment (salinity or cation ratio). Salinity at 30 and 60 mM NaCl reduced the fruit yield
compared with that obtained at 0 mM NaCl. However, there was an increase, as a consequence of the application
of treatments C2 and C3 , in each saline treatment. At a high salinity level (60 mM), the ratios Na/K, Na/Ca and
Na/Mg in young leaves decreased as a consequence of cation treatments. Higher concentrations of sugars in leaves
and fruits were obtained after increasing the salinity and cation concentrations. Also, sucrose phosphate synthase
activity in leaves and fruits was increased after the treatments, but there was no measurable invertase activity in
fruits. Therefore, the concentrations of Ca, K and Mg in the nutrient solution could be important factors in the
hydroponic culture of tomato grown under saline conditions.

Abbreviations: Gs – Stomatal conductance; SPS – Sucrose phosphate synthase

1. Introduction in growth and yield reductions of tomato crops [21].

The disorders in plant nutrition produced by salinity
may result from the effect on either nutrient availab-
ility, uptake, transport or partitioning within the plant.
The main disruptive effect on cations are in potassium,
calcium and magnesium concentrations. Potassium is
the most prominent inorganic plant solute, and as such,
makes a major contribution to the low osmotic poten-
tial in the stele of the roots, that is a prerequisite
for the maintenance of pressure-driven turgor [22].
Sodium-induced K+ deficiency has been implicated
Decreased K+ uptake, caused by Na+ is a competitive
process, which occurs regardless of whether or not the
solution is dominated by Na+ . Culture experiments
have shown the deleterious effects associated with
reduced uptake and translocation of K + by tomato
plants grown in high Na+ , which can frequently be
alleviated by the addition of K+ to the substrate [30].
The presence of adequate Ca2+ in the substrate influ-
ences the K+ /Na+ selectivity by shifting the uptake
ratio in favour of K+ at the expense of Na+ . Improve-
ment in Ca2+ mediated membrane integrity invariably
38
leads to reduction of K+ leakage from root cells and a
more favourable root-K+ status [3]. As a result, the
beneficial effects of supplemental Ca2+ on the K+
status of salt-stressed plants are often more evident in
root tissue than in the shoots [21]. Calcium deficiency
symptoms generally arise from differences in its alloc-
ation in the growing regions of the plant. Therefore,
competitive sinks such as leaves, fruits and meristems,
exert their influence on calcium movement independ-
ently [5]. However, one of the causes of blossom-end
rot in tomato fruits has been identified as a decrease
in the uptake of calcium by roots and an increase in
the resistance to transport inside the fruit [19]. Growth
and yield reductions of Na-salinised tomato are more
generally ameliorated by increases in Ca2+ [1]. Very
little information is available about the behaviour of
magnesium in saline environments. In recent stud-
ies, we found that increases in NaCl reduced leaf
Mg2+ concentration in citrus [28] and that interactions
of Mg2+ with Ca2+ , K+ and Na+ should be taken into
account with tomato plants grown under hydroponic
culture [4].
High levels of Na+ and Cl− in the apoplast alter
aqueous and ionic thermodynamic equilibria, which
result in hyperosmotic stress. Thus, it becomes vital
for NaCl-exposed plants to re-establish cellular ion
homeostasis for proper metabolic functioning and
growth. In other words, they have to adapt to the saline
environment. Osmotic adjustment helps plant cells to
withstand salt stress and water deficits by maintain-
ing sufficient turgor for growth [33]. In addition, the
osmotic adjustment involves the transport, accumu-
lation and compartmentation of inorganic ions and
organic solutes [14].
Tomato plants exposed to salinity exhibited
changes in their leaf carbohydrate metabolism and
carbon partitioning patterns, enhancing sugar load-
ing into the phloem and/or unloading into the fruits,
thereby maintaining dry matter accumulation in fruits
even as leaf expansion was inhibited [11]. The abil-
ity of the plant to synthesise and accumulate sucrose
in leaves and fruits is mainly determined by the con-
certed actions of sucrose phosphate synthase (SPS,
EC 2.3.1.14) and invertase (EC 2.3.1.26) [13]. In
terms of dry matter production, fruit yield is related
to the supply of recently produced sucrose, which
is transported through the phloem, and to the par-
titioning of the carbon assimilates to different plant
organs [17].
In this work, the effect of the increase in the bal-
anced concentrations of Ca, K and Mg on water rela-
tions, nutrient concentration, yield and sugar metabol-
ism has been investigated in order to observe whether
the compensating effect of those cations is important
for plant growth and fruit quality.
2. Material and methods
2.1 Growth conditions and experimental design
Seeds of tomato (Lycopersicon sculentum, Mill) cv.
Daniella were germinated in peat trays. On 24 March
1998, after removing all peat from the root system,
two uniform seedlings were transplanted into each of
the 36 continuously-aerated 120 L capacity contain-
ers with the following macronutrients (mmol L−1 ): 6
KNO3 ; 4 Ca(NO3 )2 , 1 NH4 (H2 PO4 ), 1 MgSO4 and
the micronutrients (µmol L−1 ) were applied in the
form of 20 Fe-EDTA, 25 H3 BO3 , 2 MnSO4 , 2 ZnSO4 ,
0.5 CuSO4 and 0.5 H2 MoO4 . The solutions were
made using deionized water. During the experiment,
the volume of the nutrient solutions was maintained
by adding deionized water daily and recording the
volume of water added. All solutions were analysed
weekly and readjusted to initial concentrations when
necessary. The pH was kept within the range of 6.0 to
6.5 by adding H2 SO4 or KOH.
The experimental design consisted of 3 salinity
levels, made by the addition of 0, 30 or 60 mmol
NaCl L−1 (S0 , S1 and S2 , respectively) in a factorial
combination with 3 levels of the ratio Ca:K:Mg (in
mM: 4:7:1, 7:9:2 and 10:12:3; treatments C1 , C2 and
C3 respectively), giving 9 treatments with 5 replic-
ates each. Cation treatments were added on the day
of transplantation such that the nitrate and phosphate
concentrations were not altered. Therefore, for C2
the macronutrients were applied as (mmol L−1 ) 7
Ca(NO3 )2 , 1 NH4 H2 PO4 , 2 MgSO4 , 4.5 K2 SO4 and
C3 as 7 Ca(NO3 )2 , 1 NH4 H2 PO4 , 3 MgSO4 , 3 CaCl2 ,
6 K2 SO4 . The solutions were brought to their corres-
ponding salinity levels 5 days after transplanting by
adding 30 mmol NaCl L−1 to the S2 and S3 treatments.
On the following two days, a further 30 mmol L−1 was
added to the S3 treatment.
2.2 Plant measurements
On 23 April, one plant from each tank was clipped (to
bear 6 clusters) and the other plant was removed after
the measurement of leaf water relations. The different
plant organs were dried at 65 ◦ C for two to four days.
 39

Chemical analyses of plant organs were carried out

after digestion with HNO3 -HClO4 (2:1). Magnesium,
Ca, K and Na concentrations were determined by
atomic absorption spectrometry (Perkin-Elmer 5500,
USA).
The other plant of each tank was maintained until
all fruits were harvested. The first harvest of ripe fruit
was on 6 June and fruit was harvested weekly from
this date until the end of the experiment (21 July).
The fruits of each cluster were separately collected.
Total weight and number of fruits, weight per fruit,
and the number of fruits with blossom-end rot (BER)
were recorded. Leaves and fruits from this plants were
also harvested for sugars and enzymatic analysis.

2.3 Leaf water relations

On 30 April, before the plants were clipped, the leaf

water potential (9w ) of the most recent fully expan-
ded leaves was measured between 10:00 h and 12:00 h
using the pressure chamber technique [31]. The same
leaves were then put into plastic bags and rapidly
frozen with liquid nitrogen. They were subsequently
thawed and pressed to extract the cell sap. The osmotic
potential (9π ) of the leaf sap was calculated after
measuring sap osmolarity using an automatic freez-
ing point depression osmometer (Digital Osmometer,
Roebling, Berlin). Turgor potential was estimated
as the difference between leaf water potential and
osmotic potential. Adaxial stomatal conductance to
water vapour exchange (Gs) was measured in one
leaf per plant with a LI-COR model 1600 diffusion
porometer between 10:00 and 12:00 h.
2.4 Soluble sugars analysis
Organic solutes were extracted from leaf sap and fruit
juice with 80% ethanol. The extracts were filtered
through sep-pak C18 cartridges to remove the colour.
The total sugar concentration was determined by the
anthrone method [16].
2.5 Fruit quality
At the middle of the fruit harvest, three fruits per
plant were taken to determine some quality paramet-
ers; soluble sugars by the anthrone method (see above)
and firmness by the resistance to penetration. Colour
was determined with a tristimulus colour spectropho-
tometer, Minolta CM-508i (Osaka, Japan) to obtain
the absorption spectra from which L∗ a∗ b∗ values were
Figure 1. Stomatal conductance, Gs, and water uptake expressed
per G of root from tomato plants treated with three different
Ca:K:Mg concentrations (C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and
three saline levels. Data are mean ± SE (n = 5).
calculated using illuminant D65 and 10◦ observer
according to the CIELAB 76 convention [24].
2.6 Enzymatic activities
SPS activity was monitored by the production of UDP.
Leaves and fruit samples were extracted with 50 mM
Hepes (NaOH) buffer, pH 7.0, containing 20 mM
MgCl2 , 1 mM EDTA, 1 mM DTT, 1 mM PMSF,
10 mM isoascorbic acid and 2% (w/v) PVP. The crude
extract was desalted by centrifugation through a 1-
ml column of Sephadex G-25. The reaction mixture
contained 50 mM Hepes (NaOH) buffer, pH 7.5, with
10 mM fructose-6-P, 10 mM UDP glucose and 10 mM
MgCl2 in a test tube. After 20 min incubation the reac-
tion was stopped by heating the test tube in a boiling
water bath for 2 min. Afterwards, UDP formation was
assayed by adding PEP/NADH in Hepes (NaOH) buf-
fer, pH 7.5. The initial absorbance was read at 340 nm.
The oxidation of NADH was started with 2 U pyruvate
kinase and 2 U L-lactate dehydrogenase. When the
40
Table 1. Electric conductivity of each cation treatment at each
saline level

Treatments EC (dS m−1 )

NaCl Ca:K:Mg

0 mM 4:6:1 1.98
7:9:2 2.86
10:12:3 3.89

30 mM 4:6:1 4.82
7:9:2 5.45
10:12:3 6.74

60 mM 4:6:1 7.68
7:9:2 8.30
10:12:3 9.21

reaction was completed (1h), the absorbance was also

recorded.
Invertase, acid and neutral, activity was assayed
by measuring by the concentrations of glucose and
fructose produced. Aliquots (100 µl) of the enzyme
preparation (see above) were incubated for 45 min
at 30 ◦ C with 50 mM Hepes (NaOH) buffer, pH 7.5
and 5.5 for the neutral and acid activities respectively,
both containing 50 mM sucrose. The reaction was
stopped by adding 1 M NaOH. To deproteinize the
samples, 1 M FeCN and 1 M ZnSO4 were added and
the precipitates were separated by centrifugation. The Figure 2. Water potential, 9w , osmotic potential, 95 and turgor of
concentrations of glucose and fructose were meas- tomato leaves treated with three different Ca:K:Mg concentrations
ured before and after the reaction took place with the (C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are
mean ± SE (n = 5).
anthrone reagent [16].

3. Results A similar effect of all treatments was also observed

Stomatal conductance of youngest recent fully expan-
ded leaves was measured at midday (Figure 1). A
significant decrease was observed when the NaCl con-
centration was increased from 0 to 60 mM. In the same
way, decreases appeared due to the increase in the
cation concentration. The result was a linear decrease
as the electric conductivity of the nutrient solution
was increased (the electric conductivity of the nutrient
solution is shown in Table 1). Water uptake was also
measured at the same time as the stomatal conductance
(Figure 1) and similar effects of the treatments were
found, that is a significant decrease in uptake occur-
ring after the osmotic potential of the nutrient solution
was raised.
on leaf water potential (9w ) and leaf osmotic potential
(95 ) (Figure 2). A good association was observed,
as well, between those parameters and the electric
conductivity of the nutrient solution (Table 1). When
turgor was calculated, it was increased when salin-
ity was applied (Figure 2) but no significant changes
were observed within the two levels of NaCl. Also,
no variation was found among the different levels of
cations.
Nutrient composition was analysed in the roots,
stems and leaves, divided into old, medium and young
leaves. Apart from the changes in the concentration
of Na+ and Cl− , the only differences produced by
the treatments were in the roots and young leaves
and only with respect to the concentrations of K+ ,

Figure 3. Cation concentration in mmol per Kg of dry weight of tomato roots treated with three different Ca:K:Mg concentrations (C1 :4:6:1;
C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are mean ± SE (n = 5).
Ca2+ , and Mg2+ . In roots, the concentration of Na
increased as a higher concentration of NaCl was added
to the nutrient solution (Figure 3). However, at both
levels of NaCl (30 and 60 mM), when the cation bal-
ance was increased, the concentration of Na decreased
significantly. The concentration of K (Figure 3) only
increased at 0 mM NaCl when the levels of Ca:K:Mg
were 7:9:2. Calcium concentration remained constant
in roots at 0 mM NaCl when the cation balance
was increased. Nevertheless, a significant decrease
appeared with salinity, which was restored when the
cation balance was increased. There were no changes
of Mg concentration (Figure 3) with the saline treat-
ments, but it increased in a similar way to Ca2+ when
the levels of Ca:K:Mg were increased from 4:6:1 to
10:12:3.
In young leaves, the effect of NaCl on the concen-
tration of Na was similar to that detected in roots, that
is an increase, when enhancing the concentration of
41
NaCl, but a decrease when increasing the concentra-
tion of cations (Figure 4). Potassium (Figure 4) was
decreased after the application of both concentrations
of NaCl, but a increase was observed when the con-
centration of cation was increased, with no significant
differences between the levels of Ca:K:Mg, 4:7:1 and
10:12:3. Concentrations of both Ca and Mg did not
change with cation treatments at 0 mM NaCl, but
a large decrease occurred at 30 and 60 mM NaCl,
which was partly restored by increasing the cation
levels. The concentration of Cl− was always increased
proportionally after increasing the NaCl concentra-
tions in the nutrient solution (data not shown). The
ratio between cations was calculated in roots and
young leaves (Figure 5) and it can be seen that at
a high NaCl concentration, the ratios Na/Ca and
Na/Mg decreased as the cation balance was increased.
However, the ratio Na/K did not change with the
treatments.
42
Figure 4. Cation concentration in mmol per Kg of dry weight of tomato young leaves treated with three different Ca:K:Mg concentrations
(C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are mean ± SE (n = 5).
Plant growth (Figure 6) was measured and
expressed as fresh weight (g plant−1 ). No significant
changes occurred as a consequence of the increase in
cation concentration within the 0 mM NaCl treatment,
either in shoots or in roots. In shoots, there was a
decrease when the saline treatments 30 and 60 mM
were applied, which was partially restored to control
values with Ca:K:Mg, 10:12:3. In roots, there was
only a decrease in fresh weight after the treatment of
60 mM NaCl. That decrease was also restored by the
cation concentration Ca:K:Mg, 10:12:3.
At 0 mM NaCl the production of marketable fruits
(Figure 7) was significantly increased as the cations
were increased. As the salt concentration increased,
a progressive decrease was observed. However, in
those cases, the fruit production increased slightly
when the cation levels were increased. The production
of unmarketable fruits was higher, but very variable
within each of the 30 and 60 mM NaCl-treatments.
Total soluble sugars were determined in leaves
(Figure 8) and the result was a linear increase as the
electric conductivity of the nutrient solution increased.
Therefore, there were higher concentrations with the
saline treatments and with the increase in cation levels.
Fruit quality was determined, but there were no
alterations with NaCl/cations treatments, in fruit firm-
ness, or colour in terms of L∗ , a∗ or b∗ values (data not
shown). However, when the soluble sugars content in
fruit juice was analysed (Figure 8), an increase was
observed, with both concentrations of NaCl, which
was greater when the concentration of cations was
increased, with no significant differences between the
levels of Ca:K:Mg, 4:6:1 and 7:9:2.
SPS, and acid and neutral invertases were ana-
lysed in both leaves and fruits. In leaves, a higher
SPS activity was observed after the saline treatments
(Figure 9). Furthermore, within those saline treat-
ments, greater activity was observed after increasing

Figure 5. Cation relations in mmol per Kg of dry weight of tomato
young leaves, treated with three different Ca:K:Mg concentrations
(C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are
mean ± SE (n = 5).
the cation concentration. In fruits, a slight increase
was observed progressively with the saline and cation
treatments. The neutral invertase activity in leaves did
not vary much with the treatments (Figure 10). How-
ever, the acid invertase activity strongly increased in a
similar way for both saline treatments, but no effect
of the cation treatments was observed. Both inver-
tase activities were non-detectable in fruits (data not
shown).
4. Discussion
Plants grown under salt stress conditions alter their
stomatal resistance due to the increased electric con-
ductivity in the soil or nutrient solution. Therefore,
43
Figure 6. Shoot and root weight, in G fresh weight, of tomato plants
treated with three different Ca:K:Mg concentrations (C1 :4:6:1;
C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are mean ±
SE (n = 5).
gas exchange in their leaves is reduced. This reduction
has been attributed to salt damage to the photosyn-
thetic tissue, to stomatal effects with the consequent
restriction of the CO2 availability for carboxylation or
to the acceleration of senescence [26]. In our plants,
there was a reduction in Gs after NaCl treatments and
after the increase in the concentration of cations in the
nutrient solution. This indicates that the reduction was
directly related to the osmotic pressure of the nutrient
solution and was independent of the nature of the salts,
which produced the increase in the osmotic pressure.
The results obtained for Gs are in agreement with the
measurements of water uptake, which was reduced
with all treatments and was also directly related to the
osmotic potential of the nutrient solution.
Osmotic adjustment involves the net accumulation
of solutes in the cells in response to a fall in the water
potential of their environment. As a consequence of
this net accumulation, the cell osmotic potential is
lowered which, in turn, attracts water into the cell
and tends to maintain turgor pressure [2]. The reduc-
tions observed in 95 , when treatments were applied,
44
Figure 7. Fruit production of tomato plants treated with three dif-
ferent Ca:K:Mg concentrations (C1 :4:6:1; C2 :7:9:2; C3 :10:12:13)
and three saline levels. Data are mean ± SE (n = 5).
were closely related to reductions in 9w , which had
a slight effect on turgor only related to saline treat-
ments. The fact that there were few changes in turgor,
in this tomato crop, in response to the treatments
of our experiment indicated a good level of osmotic
adjustment [12].
Leaf nutrient disorders produced by salinity have
been widely studied in tomato plants (for review see
Cuartero and Fernandez-Muñoz, [6]). Large amounts
of Na+ and Cl− decreased the concentrations of
K+ , Ca2+ and Mg2+ , giving high values of the
ratios Na+ /K+ , Na+ /Ca2+ and Na+ /Mg2+ [23]. In
our experiments, the deficiencies of those ions pro-
duced by moderate salinity (30 or 60 mM) were only
observed in roots and young leaves. But those defi-
ciencies were partially compensated by the higher
cation treatments. Therefore, the ratios Na+ /K+ ,
Na+ /Ca2+ and Na+ /Mg2+ were increased to a lesser
extent. In the same way, this compensation for the
disorders gave an increase in shoot and root biomass
production although, shoots were more affected by
salinity than roots. In some cases, the effect of the
Figure 8. Sugar concentration of leaf sap and fruit juice of
tomato plants treated with three different Ca:K:Mg concentrations
(C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are
mean ± SE (n = 5).
cations in saline-treated roots was translated into an
increase in biomass, greater than that of the control
without salinity. The increases in plant biomass were
related to the marketable fruits, in which the reduc-
tion in yield by salinity was partially compensated
by the cation treatment. The importance of appropri-
ate cation activity ratios in the root zone, on yield
of tomato plants grown under saline conditions, has
been reported previously [32]. The nearly complete
restoration of the nutrient deficiencies by cation treat-
ments within the same saline level, was not translated
into a full restoration of plant biomass and fruit yield.
Therefore, the toxic effect of Na+ or Cl− was still
present. The accumulation of high concentrations of
Cl− and Na+ in the leaves of salt-treated plants for the
osmotic adjustment was produced at a lower energy
cost than the accumulation of organic solutes. In
other words, the salinity-induced yield reduction was
probably associated with the toxic effect of the accu-
mulation of Cl− and Na+ . Therefore, the restoration
of fruit yield by cation treatments, within each saline

Figure 9. Sucrose phosphate synthase activity, SPS, in tomato
leaves and fruits treated with three different Ca:K:Mg concentration
(C1 :4:6:1; C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are
mean ± SE (n = 5).
treatment, could possibly be achieved by using other
ions for the osmotic adjustment, thus, eliminating the
toxic effect of the excessive accumulation of Cl− or
Na+ . At 0 mM NaCl, an increase of fruit yield was
obtained by the application of the cation treatments.
This result could be due to the beneficial effect of
the low concentrations of Cl− present in the treatment
of Ca:K:Mg, 10:12:3, in which part of the Ca was
applied as CaCl2 (see materials and methods). There-
fore, the Cl− concentration of the nutrient solution was
6 mM, which has been reported to increase crop yield
in cotton [25] or citrus [15].
Characteristics, such as soluble solids, sugars,
acidity and pH are important quality parameters for
both fresh-market and processing tomatoes [6]. It is
well known that the total soluble solids content, the
most important criterion for tomato paste processing,
increases with salinity, although yield reduction is also
expected [29]. In our samples the enhancing of the
total soluble solids did not produce any significant
alteration to the fruit firmness or the colour. However,
45
Figure 10. Acid and neutral invertase activity in tomato leaves
treated with three different Ca:K:Mg concentrations (C1 :4:6:1;
C2 :7:9:2; C3 :10:12:13) and three saline levels. Data are mean ±
SE (n = 5).
there was a significant increase in the total soluble
sugars after the NaCl and cation levels were increased,
which means that a higher quality was obtained.
About 50% of tomato dry matter consists of sugars,
of which fructose is the most important one contribut-
ing to sweetness. Sucrose is present but rarely exceeds
0.1% of the fresh weight [9]. It has been reported that
young fruits have starch as a large pool of hexoses
[10], which is transformed into sucrose, glucose and
fructose according to the developmental stage [27]. In
our ripe fruits, invertase activity was non-detectable,
but SPS activity was measurable. It has been suggested
that, in spite of the apparent presence of extracellular
leaf invertase, the sucrose may be taken up directly
by pericarp cells for compartmentalisation as soluble
hexoses in the vacuole [8]. However, the fact that in
our ripe tomato fruits, neither invertase activity nor
sucrose were detectable, led us to consider the possib-
ility that all sugar transport from the leaves to the fruits
was in the form of hexoses. This was also demon-
strated by Damon et al. [8], who showed that hexoses
46
were the principal form of sugar in the apoplast sap of
tomato. The fact that very high acid invertase activity
was measured in leaves of plants treated with 30 or
60 mM NaCl could demonstrate that large amounts of
those hexoses were playing an important role in the
osmotic adjustment. It has been reported that the SPS
activity is low throughout tomato fruit development
[7] including in ripe fruits [20]. However, the role of
SPS in the re-synthesis of sucrose during fruit develop-
ment is still not clear [18]. In our plants, SPS activity
in fruits was low compared to that of leaves. However,
the fact that only slight amounts of sucrose were found
in some saline-treated fruits (data not shown) does not
enable us to shed any light on the possible role of SPS
in fruits.
To summarise, we can conclude that the concentra-
tion of cations in plants grown under saline conditions
could be an important factor in tomato production in
greenhouses using hydroponic systems. When saline
water is available, higher concentrations of the ratio
Ca:K:Mg (in mM: 7:9:2 or 10:12:3) increased growth
and fruit yield, although the toxic effect of Na+ or
Cl− was still present. Furthermore, the higher quality
of the tomato fruits can, to some extent, compensate
the decreases in production. Further improvement of
tomato yield and quality will require further research
into the effects of plant mineral nutrition on the regu-
lation of fruit number and size as well as the regulation
of water relations of the plants and sucrose metabolism
and transport within tomato fruits.
Acknowledgements
The authors wish to thank Alicia Aragon for her tech-
nical assistance in cation determination, Román López
for the greenhouse maintenance, Dr Abel Piqueras
and Pilar Flores for their help determining sugars
and enzymatic activities and Dr David Walker for
the English correction of the manuscript. This work
was funded by CICYT (PETRI)-Spain (project n◦
95-0174-OP).
References
1. Al-Harbi AR (1995) Growth and nutrient composition of
tomato and cucumber seedling as affected by sodium chloride
salinity and supplemental calcium. J Plant Nutr 15: 341–358
2. Blum A, Munns R, Passioura JB and Turner NC (1996)
Genetically engineered plants resistant to soil drying and salt
stress: How to interpret osmotic relations? Plant Physiol 110:
1050–1053
3. Cachorro P, Ortiz A and Cerdá A (1994) Implications of cal-
cium nutrition on the response of Phaseolus vulgaris L. to
salinity. Plant Soil 159: 205–212
4. Carvajal M, Martínez V and Cerdá A (1999) Influence of mag-
nesium and salinity on tomato plants grown in hydroponic
culture. J Plant Nutr 22: 177–190
5. Clarkson DT (1984) Calcium transport between tissues and its
distribution in the plant. Plant Cell Environ 7: 449–456
6. Cuartero J and Fernandez-Muñoz R (1999) Tomato and salin-
ity. Scientia Horticul 78: 83–125
7. Dali N, Michaud D and Yelle S (1992) Evidence for the
involvement of sucrose phosphate synthase in the pathway
of sugar accumulation in sucrose-accumulating tomato fruits.
Plant Physiol 99: 434–438
8. Damon S, Hewitt J and Bennett AB (1988) Sink metabolism
in tomato fruit. Plant Physiol 87: 731–736
9. Davis JN and Kempton RJ (1975) Changes in the individual
sugars of tomato fruit during ripening. J Sci Food Agric 26:
1102–1110
10. Dinar M and Stevens MA (1981) The relationship between
starch accumulation and soluble solids content of tomato
fruits. J Am Soc Hortic Sci 106: 415–418
11. Gao Z, Sagi M and Lips SH (1998) Carbohydrate metabol-
ism in leaves and assimilate partitioning in fruits of tomato
(Lycopersicon esculentum M.) as affected by salinity. Plant Sci
135: 149–159
12. Greenway H and Munns R (1980) Mechanisms of salt toler-
ance in nonhalophytes. Ann Rev Plant Physiol 31: 61–69
13. Guan HP and Janes HW (1991) Light regulation of sink
metabolism in tomato fruit. I. Developmental changes in car-
bohydrate metabolizing enzymes. Plant Physiol 96: 922–927
14. Hajibagheri MA, Harvey DMR and Flowers TJ (1987) Quant-
itative ion distribution within root cells of salt-sensitive and
salt-tolerant maize varieties. New Phytol 105: 367–379
15. Heller J, Shalhevet J and Goel A (1973) Response of a cit-
rus orchard to soil moisture and soil salinity. In: Hadas A,
Swartzendruber D, Rijtema PE, Fuchs M and Yaron B (eds)
Physical Aspects of Soil Water and Salts in Ecosystems.
Berlin, Heidelberg, New York: Springer-Verlag, pp 409–419
16. Hewitt BR (1958) Spectrophotometric determination of total
carbohydrate. Nature 182: 246–247
17. Ho LC (1988) Metabolism and compartmentation of imported
sugars in sink organs in relation to sink strength. Ann Rev
Plant Physiol Plant Mol Biol 39: 355–378
18. Ho LC (1996) The mechanism of assimilate partitioning and
carbohydrate compartmentation in fruit in relation to the
quality and yield of tomato. J Exp Bot 47: 1239–1243
19. Ho LC, Belda R, Brown M, Andrews J and Adams P (1993)
Uptake and transport of calcium and the possible causes of
blossom-end rot in tomato. J Exp Bot 44: 509–518
20. Islam MS, Matsu T and Yoshida Y (1996) Carbohydrate con-
tent and the activities of sucrose synthase, sucrose phosphate
synthase and acid invertase in different tomato cultivars during
fruit development. Sci Hortic 65: 125–136
21. Lopez MV and Satti SME (1996) Calcium and potassium-
enhanced growth and yield of tomato under sodium chloride
stress. Plant Science 114: 19–27
22. Marshner H (1995) Mineral Nutrition of Higher Plants. Lon-
don: Academic Press, 889 pp
23. Martínez V, Cerdá A and Fernandez FG (1987) Salt tolerance
of four tomato hybrids. Plant Soil 97: 233–242
24. McLaren K (1980) Food colorimetry. In: Walford J (ed) Devel-
opments in Food Colours.I. Applied London, UK: Science
Publishers Ltd, pp 27–45

25. Pasternak D (1987) Salt tolerance and crop production – a
comprehensive approach. Ann Rev Phytopatol 25: 271–291
26. Pessarakli M (1994) Handbook of Plant and Crop Stress. New
York – Basel – Hong Kong: Marcel Dekker
27. Robinson NL, Hewith JD and Bennett AB (1988) Sink meta-
bolism in tomato fruit. Plant Physiology 87: 727–730
28. Ruiz D, Martínez V and Cerdá A (1997) Citrus response to
salinity: growth and nutrient uptake. Tree Physiol 17: 141–150
29. Saranga Y, Zamir D, Marani A and Rudich J (1991) Breeding
tomatoes for salt tolerance: field evaluation of Lycopersicon
germplasm for yield and dry-matter production. J Am Soc
Hortic Sci 116: 1067–1071
47
30. Song JQ and Fujiyama H (1996) Difference in response of rice
and tomato subjected to sodium salinization to the addition of
calcium. Soil Sci Plant Nutr 42: 503–510
31. Turner NC (1988) Measurement of plant water status by the
pressure chamber technique. Irrigation Sci 9: 289–308
32. Willumsen J, Petersen KK and Kaack K (1996) Yield and
blossom-end rot of tomato as affected by salinity and cation
activity ratios in the root zone. J Hort Sci 71: 81–98
33. Zimmermann U (1978) Physics of turgor and osmo-regulation.
Ann Rev Plant Physiol 29: 121–148