Scientia Horticulturae 112 (2007) 282–289

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Effects of NaCl stress on red raspberry

(Rubus idaeus L. ‘Autumn Bliss’)
Damianos Neocleous a,*, Miltiadis Vasilakakis b
a
Agricultural Research Institute, Ministry of Agriculture, Natural Resources and Environment, P.O. Box 22016, 1516 Nicosia, Cyprus
b
Department of Horticulture and Viticulture, Aristotle University of Thessaloniki, Thessaloniki 540.06, Greece
Received 13 March 2006; received in revised form 30 October 2006; accepted 5 December 2006

Abstract
The increasing interest in the cultivation of red raspberry (Rubus idaeus L.) in warmer climates and in the mild-Southern parts of Europe
requires better understanding of the salinity effects especially in the primocane-fruiting type cultivars. The aim of this work was to study the
response of the ‘Autumn Bliss’ primocane-fruiting cultivar in elevated NaCl concentrations.
The experiment was carried out under natural greenhouse conditions. ‘Autumn Bliss’ cultivar was grown in pots containing perlite. The plants
were irrigated with Hoagland nutrient solution containing 0, 5, 10, 15, 20 and 30 mM NaCl. Plant growth, photosynthetic parameters, water
relations, chloride and sodium content, leaf chlorophyll content and fluorescence were determined.
The results of this work indicate that increasing salt concentration in the nutrient solution leads to: (a) a decline in the photosynthetic rate and in
stomatal conductance, although there is no reduction in intercellular CO2; (b) the reduction of leaf chlorophyll content and fluorescence; (c) to a
decrease in transpiration rate, leaf water potential, relative water content and plant water consumption, however, the water use efficiency is not
affected; (d) the reduction of: cane length and diameter, percentage of green leaves, leaf area, leaf expansion rate and relative growth rate, while the
number of leaves remain unaffected and the percentage of dead leaves increases; (e) a decrease of the plant fresh and dry weight and to an increase
of the root/shoot ratio; (f) a linear increase of chloride and sodium content in the plant.
Quantifying the response of raspberry in elevated NaCl concentrations provides that the reduction in photosynthesis is most probably due to
damages in the photosynthetic apparatus rather than from factors affecting stomatal closure. Besides, a pronounced toxic effect of Cl was clearly
demonstrated and this points towards that the salt injury effect could be attributed to Cl toxicity (uptake and translocation). As a consequence, the
decrease photosynthetic capacity and salt toxicity in the plant tissues restricted plant growth under saline conditions.
# 2007 Elsevier B.V. All rights reserved.

Keywords: Rubus idaeus; Red raspberry; Salinity; NaCl; Photosynthetic parameters; Water relations; Growth

1. Introduction Salt resistance includes both avoidance and tolerance

Salinity is one of the most important factors limiting fruit
growth and production of several horticultural crops. Most of
the salt stress in nature is due to sodium chloride salts (Levitt,
1972). Salinity can damage the plant through its osmotic effect,
which is equivalent to a decrease in water activity, through
specific toxic effects of ions and by disturbing the uptake of
essential nutrients (Laüchli and Epstein, 1990; Marschner,
1995; Dorais et al., 2001). In general, enzymes and metabolic
activities in plants are highly influenced by both amount and
type of salts (Flowers et al., 1977).
* Corresponding author. Tel.: +357 22 403115; fax: +357 22 316770.
E-mail address: d.neocleous@arinet.ari.gov.cy (D. Neocleous).
0304-4238/$ – see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2006.12.025
mechanisms (Levitt, 1980). The former may operate through
active extrusion of ions by specific pumps, passive exclusion of
ions due to membrane impermeability, or dilution by rapid
growth associated with an increase in water content. An
unavoidable consequence of growth in a solution containing a
high salt concentration is the development of osmotic stress,
which is followed by a loss of turgor. Tolerance to osmotic
stress may operate either through dehydration tolerance, which
permits the cell to survive without growing when the turgor
decreases, or by avoiding dehydration through osmoregulation,
which includes an increase of solute concentration in the cell
and consequently rehydration. The solutes may be either salt
ions, which can be sequestered in the vacuole and osmotically
balanced by organic solutes in the cytoplasm, or organic
substances. The latter occurs when salt ions are prevented from
 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289
entering the cells (Tal, 1984). The first strategy is typical to
halophytes. Salt-sensitive glycophytes in which group most
agricultural and horticultural crops belong respond to salinity
with profound decrease in vegetative and reproductive growth
(Greenway and Munns, 1980). These responses are highly due
to osmotic effects. The inability of osmoregulation may result
from either an insufficient uptake of salt ions or a lack of
synthesis of organic solutes, which leads to growth disruption
occasioned by reduced water uptake as well as reduced water
potential in the soil, resulting to physiological water stress. In
addition, salinity can cause injury by inorganic ions, which are
absorbed by the cell, are not compartmentalized, and are
accumulated at toxic levels in plant tissues. On the other hand, it
can antagonize the uptake of essential nutrients, hence inducing
deficiencies of ions already in adequate amounts in the soil
(Wahome, 2003). Halophytes exhibit a higher concentration of
Na and Cl ions in the cytoplasm than in the vacuole. However,
the sensitivity of glycophytes to salt is due to their inability to
keep the Na and Cl ions outside the cytoplasm (Flowers et al.,
1977). In glycophytes, reduction in the accumulation of Na and
Cl ions in shoots and leaves under saline conditions occurs
through varying root permeability to different ions and
selectivity of the ion uptake. Exclusion of Na in leaves occurs
through accumulation of the ions in the roots and re-
translocation of the ions from the leaves to the roots, where
they are subsequently excreted into the soil medium
(Marschner, 1993).
The largest areas of saline soils in the world are saline
because they occur in the arid and semiarid zones of the world
(Epstein, 1972). In such areas, long periods of drought coincide
with high temperatures. The increasing interest in the
cultivation of red raspberry in warmer climates and in the
mild-Southern parts of Europe (Vasilakakis, 1997; Oliveira
et al., 1996, 1999; Rosati et al., 1999; Neocleous et al.,
2005a,b), where the water salinity is one of the major limiting
factors in agricultural production, a better understanding about
the plant–salt relationship physiology is required since there is
lack of information in the literature in comprehensive
understanding of the principal physiological mechanisms of
salt damage in red raspberries and especially in primocane-
fruiting type cultivars.
Since, no information was found on salinity effects on red
raspberry, the aim of this work was an effort to investigate the
relative contributions of primary (toxic) and secondary
(osmotic) effects of salt to physiological, biochemical and
growth parameters in raspberry plants.
2. Materials and methods
The experiment was conducted at the Agricultural Research
Institute of Cyprus (long. 328E, lat. 358N). Red raspberry plants
(Rubus idaeus L. ‘Autumn Bliss’) were grown in a greenhouse
under soilless (perlite) cultivation. Young secondary shoots at
the fourth true leaf stage (15 cm height) were removed from the
plants and the rooting system was carefully washed. The plant
material was uniform and transferred to 18 L plastic pots filled
with perlite. The pots were covered with polyethylene film to
283
avoid evaporation from perlite surface. One cane per pot was
allowed to grow in a greenhouse under natural illumination and
temperature conditions for 45 days during the period February–
April 2005. The average temperature was 22 8C, the mean
global solar radiation 16.8 MJ m2 and the mean relative
humidity 51%.
The plants were supplied with Hoagland nutrient solution
(Hoagland and Arnon, 1950) containing 0, 5, 10, 15, 20 and
30 mM NaCl from planting until the end of the experiment (45
days). The nutrient solution plus the added NaCl was applied to
the plants when needed according to moisture sensors
embedded in the root environment and at sufficient rates until
drainage, to facilitate two tasks: (a) a constant build-up of NaCl
salts and (b) replenishment of various elements, in the root
zone. The six concentrations of NaCl were arranged in a
randomized complete block design with four replications. Each
replication consisted of three sample plants. Data were
analyzed using the means procedure of SAS/ASSIST: analysis
of variance (SAS Institute, Cary, NC, USA).
At the start of the experiment and before any treatment
began, eight plants were harvested to determine initial dry
weight. The following measurements were performed on the
youngest fully expanded leaf, 2 h after the sunrise (a) 7-days
intervals: (i) photosynthetic rate, stomatal conductance and
transpiration rate using Li-6200 (Li-Cor, Lincoln, NE-USA),
(ii) water use efficiency was estimated as the quotient of the
photosynthetic rate over the transpiration rate, (iii) leaf water
potential using plant moisture system SKPM 1400 (Skye
Instruments Ltd, Wales, UK) and (iv) leaf chlorophyll
fluorescence using chlorophyll fluorometer OS-30p (Opti-
Sciences, Hudson, USA). The OS-30p measures chlorophyll
fluorescence parameters F 0 and F m, where F 0 is the initial
fluorescence, F m the maximum fluorescence and the variable
fluorescence F v is calculated as: F v ¼ F m  F 0 ; (b) 15-days
intervals: (i) leaf chlorophyll content. To estimate the leaf
chlorophyll content three leaf disks of 2.5 cm2 area were
weighed and ground in 10 cm3 of N,N-dimethylformamide.
After 48 h, storage in the dark at 5 8C, the absorbance of the
supernatant was measured at 647 and 664 nm with a Campesc
M350 UV-Vis spectrophotometer (Campesc Ltd., Cambridge,
UK) and chlorophyll content – a, b and a + b – was computed
according to Moran (1982), (ii) relative water content
calculated as: RWC = (Lfw – Ldw)/(Ltw  Ldw), where Lfw is
the leaf fresh weight, Ldw the leaf dry weight and Ltw is the leaf
turgid weight (Barrs and Weatherley, 1962). Plant water
consumption was recorded weekly by weighing the pots before
and after fertigation (at the end of drainage) assuming that
evaporation was zero.
At the end of the experiment (45 days from planting), the
following aspects were determined: (a) cane length, (b) diameter
of the cane (below the top), (c) number of leaves, percentage of
green and dead leaves, (d) leaf area (for the top 10 nodes) using
leaf area meter CI-203 (CID, Inc., Vancouver, USA). At the end
of the measurements, the plants were harvested and the roots
were carefully washed with rainwater to remove any perlite
particles and surface ions before chemical analyses. Plant fresh
and dry weight (after drying at 68 8C) was recorded. Chloride and
284 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289

sodium content was measured separately for roots, top and lower
part of the stem, new and older leaves. Chloride was determined
by titration against silver nitrate and sodium by flame photometer
Sherwood model 420 (Sherwood Scientific Ltd., Cambridge,
UK). Relative growth rate was calculated as: RGR =
(ln W2  ln W1)/(t2  t1) (Blackman, 1919), where W1 and W2
are plant dry weights at times t1 and t2. Leaf expansion rate
calculated as: LER = dAl/dT, where dA1 is leaf area at time
T2  leaf area at time T1, and dT is T2  T1. RGR and LER were
taken over the entire treatment period.

3. Results

3.1. Visual symptoms

The injury caused by NaCl, after 1 month of salt application,

began as leaf tip necrosis then spreading in the middle of the
leaves. Specifically, the symptoms were observed in the lower
and older leaves, which finally abscised. By contrast, younger
growing leaves did not show any symptoms of necrosis. The Fig. 1. The effect of different NaCl concentrations on leaf chlorophyll content
symptoms were more obvious at 20 and 30 mM salt (A) per unit area (B) per cent fresh weight of ‘Autumn Bliss’ red raspberry
cultivar. Data are mean values over a 45-day treatment period. Bars represent
concentrations.
S.E. values and refer to the total chlorophyll content a + b.

3.2. Photosynthetic parameters

Salt concentrations 15 mM NaCl in the nutrient solution

reduced significantly the photosynthetic rate and stomatal
conductance, although there was no reduction in intercellular
CO2 concentration (Table 1). Moreover, sodium chloride
application reduced leaf chlorophyll content and fluorescence,
and this reduction was statistically significant at concentrations
10 mM NaCl (Figs. 1 and 2).

3.3. Water relations

The transpiration rate declined significantly at salt concen-
trations in the solution 15 mM NaCl (Table 1). The leaf water
potential, the relative water content and plant water consumption
decreased with increasing salinity, however, for relative water
content and plant water consumption, this reduction was
significant only at 30 mM NaCl concentration (Table 2). Water
use efficiency did not show any significant differences with
salinity compared to the control (0 mM NaCl) (Table 2).
Fig. 2. The effect of different NaCl concentrations on leaf chlorophyll fluor-

escence (Fv/Fm , Fv/Fo ) of ‘Autumn Bliss’ red raspberry cultivar. Data are mean
values over a 45-day treatment period. Bars represent S.E. values. *Fv/Fm is
the variable fluorescence over the maximum fluorescence. **Fv/Fo is the
variable fluorescence over the initial fluorescence.
3.4. Vegetative growth
The cane length was reduced significantly at 30 mM NaCl
concentration, but the cane diameter and the number of leaves

Table 1
The effect of different NaCl concentrations on photosynthetic rate, stomatal conductance, intercellular CO2 and transpiration rate in the youngest fully expanded leaf
of ‘Autumn Bliss’ red raspberry cultivar
NaCl (mM) Photosynthetic rate Stomatal Intercellular Transpiration rate
(mmol CO2 m2 s1) conductance (cm s1) CO2 (ppm) (mmolH2O m2 s1)
0 8.94a 0.40a 224a 2.87a
5 8.69a 0.38ab 219a 2.79a
10 8.31ab 0.38ab 217a 2.81a
15 7.72b 0.36abc 216a 2.50b
20 7.08c 0.33bc 216a 2.47b
30 6.62c 0.32c 216a 2.22c
Values within the body of the table represent mean measurements over a 45-day treatment period. Means within a column for each NaCl concentration followed by
different letters (a–c) are significantly different according to Duncan’s multiple range test at P < 0.05 level.
 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289 285

Table 2
The effect of different NaCl concentrations on leaf water potential, relative water content, plant water consumption and water use efficiency in the youngest fully
expanded leaf of ‘Autumn Bliss’ red raspberry cultivar
NaCl Leaf water Relative water Plant water consumption Water use efficiency
(mM) potential (MPa) content (RWC) (%) (ml plant1 day1) (mmol CO2/mmolH2O)
0 1.28a 74.6a 327a 3.17ab
5 1.38b 72.8ab 314ab 3.20a
10 1.39b 73.7ab 311ab 3.03b
15 1.45b 72.8ab 271ab 3.16ab
20 1.47b 72.3ab 261ab 3.02b
30 1.59c 70.4b 246b 3.12ab
Values within the body of the table represent mean measurements over a 45-day treatment period. Means within a column for each NaCl concentration followed by
different letters (a and b) are significantly different according to Duncan’s multiple range test at P < 0.05 level.

Table 3
The effect of different NaCl concentrations on growth characteristics of ‘Autumn Bliss’ red raspberry cultivar after a 45-day treatment period
NaCl (mM) Cane length (cm) Leaf number (n) Cane diameter (mm) Green leaves (%) Dead leaves (%) Leaf area (cm2)
0 89.5a 25.9a 4.71abc 87.3a 12.7b 1632a
5 84.0ab 23.7a 5.31ab 88.8a 11.2b 1415ab
10 82.2ab 22.8a 5.36a 86.8a 13.2b 1386abc
15 77.5ab 23.2a 4.92abc 84.0ab 16.0ab 1353bc
20 74.9ab 22.7a 4.64bc 81.0b 19.0a 1194bc
30 68.5b 21.8a 4.56c 80.5b 19.5a 1147c
Means within a column for each NaCl concentration followed by different letters (a–c) are significantly different according to Duncan’s multiple range test at
P < 0.05 level.

were not affected by salinity compared to the control (Table 3).
The percentage of green leaves decreased, whereas the
percentage of dead leaves increased above 15 mM NaCl
salinity concentration (Table 3). A significant reduction in the
leaf area (for the top 10 nodes) was observed at concentrations
15 mM NaCl (Table 3). The relative growth rate and the leaf
expansion rate linearly decreased with salinity (R2 = 0.98 and
0.84, respectively) (Fig. 3). Moreover, the fresh and dry weight
decreased significantly at salt concentrations of 15, 20 and
30 mM NaCl resulting at 34%, 45% and 48% total plant dry
weight diminution, respectively. The fresh to dry weight ratio
did not change significantly with salinity, however, the roots to
shoots ratio (on dry or fresh basis) increased significantly at the
30 mM NaCl concentration (Table 4).
3.5. Elemental concentration

The chloride and sodium ions increased significantly in

various parts of the plant with salinity and as a consequence the
total chloride and sodium concentration in the plant tissue
increased proportionally to salinity in a linear trend (R2 = 0.93
and 0.96, respectively) (Fig. 4A and B). Specifically, the
chloride ions were predominantly concentrated in the new and
older leaves whereas; sodium ions were concentrated mostly in
the roots and lower part of the stem (Fig. 4C and D).

4. Discussion and conclusion

The physiological, biochemical and growth responses of red

Fig. 3. The effect of different NaCl concentrations on (A) relative growth rate
(B) leaf expansion rate of ‘Autumn Bliss’ red raspberry cultivar. Data are mean
values over a 45-day treatment period.
raspberry (R. idaeus L.) to salinity were examined in this study.
The visual symptoms of salinity observed in the current study
corroborate with those found by Banuls and Primo-Millo
(1995) for citrus and Weber and Reimann-Phillip (1989) for
Rosa multiflora. The leaf injury could be a result of the
accumulation of toxic levels of Cl and Na, ion imbalance,
nutrient deficiencies and water stress. The transport of Cl ions
occurs mainly in the transpiration stream (Wahome, 2003),
which explains the high concentration of these ions in leaves
and the occurrence of salt injury. Chloride ions are absorbed at
286 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289

Table 4
The effect of different NaCl concentrations on plant fresh weight, dry weight, fresh over dried weight and roots to shoots ration (on dry and fresh basis) of ‘Autumn
Bliss’ red raspberry cultivar after a 45-day treatment period
NaCl Plant fresh Plant dry Fresh/dry Roots/shoots Roots/shoots
(mM) weight (g) weight (g) weight (n) (on dry basis) (n) (on fresh basis) (n)
0 550a 153a 3.60a 1.93b 2.38b
5 505a 135a 3.72a 2.20ab 2.85ab
10 469ab 127ab 3.73a 1.86b 2.51b
15 366bc 101bc 3.61a 1.72b 2.53b
20 309c 84c 3.67a 2.07ab 2.68ab
30 299c 79c 3.80a 2.93a 3.35a
Means within a column for each NaCl concentration followed by different letters (a–c) are significantly different according to Duncan’s multiple range test at
P < 0.05 level.

higher rates than Na ions (Marschner, 1993). As a result, the
concentration of Cl is higher than that of Na in plant tissues and
possibly the cause of leaf damage in plants (Greenway and
Munns, 1980) like in citrus (Bar et al., 1998), and this seems to
be the case in the current study with raspberries.
The NaCl application had a negative effect on the
photosynthetic rate and stomatal conductance, but the
intercellular CO2 was not statistically affected, although there
was a tendency to be decreased. This indicates that the
reduction in stomatal conductance had not diminished
intercellular CO2 and the photosynthetic rate reduction was
not simply depended from stomatal conductance and inter-
cellular CO2, but possibly from other non-stomatal factors. The
reduction in photosynthesis impaired with non-stomatal factors
has been reported in other salt-sensitive plants as well (Dowton,
1977; Garcia-Legaz et al., 1993). Besides Steduto et al. (2000)
reported the lack of stomatal closure observed in mild-salinity
experiments, and Heuvelink et al. (2003) reported that stomatal

Fig. 4. The effect of different NaCl concentrations on (A) plant Cl content as a sum of chloride content in different plant parts (B) plant Na content as a sum of sodium
content in different plant parts (C) chloride content in different plant parts (D) sodium content in different plant parts of ‘Autumn Bliss’ red raspberry cultivar, after a
45-day treatment period. Bars represent S.E. values.
 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289
closure is a very expensive reaction and may be the last ‘defense
mechanism’. To sum up, the reduction in photosynthesis was
mostly linked to non-stomatal factors, such as the accumulation
of toxic ions, primarily chloride and water imbalance.
The leaf chlorophyll content and fluorescence were reduced
with salinity. Previous researches reported that salinity could
increase chlorophyllase activity (Sivstev et al., 1973), or this
may be caused by the depressive effect of salinity on the
absorption of some ions, such as Mg and Fe, which are involved
in the chloroplast formation (Hanafy et al., 2002). This
reduction in leaf chlorophyll content could be an important
factor connected with photosynthesis reduction. Moreover, the
leaf chloroplyll fluorescence declined simultaneously with
salinity and with the leaf chlorophyll content. This suggests that
the reduction in photosynthesis may be due to damages in the
photosynthetic apparatus rather than from factors affecting
stomatal closure. In accordance, Feierabend et al. (1992)
reported that Cl ions at toxic levels can cause blockage in the
electron transport within photosystem II, and Saied et al. (2003)
found that the dark reactions in photosynthesis were relatively
more impaired by salinity than the light reactions.
The transpiration rate decreased with salinity, while the
intercellular CO2 although tended to decrease was not
statistically affected. This indicates that stomata were partially
closed, and this could be correlated with water stress and the
accumulation of ions in guard cells. Also, structural and
metabolic modifications in leaves may be associated with a
tendency to minimize transpiration rate (Lüttge and Smith,
1984).
Water use efficiency generally did not change with salinity
compared to the control, and this suggests that salinity affects
both photosynthesis and transpiration in an analogous pattern.
This is accordance with Percival et al. (1998), who reported that
in ‘Heritage’ raspberry plants a balance between water loss and
CO2 uptake exists.
The leaf water potential was decreased with increasing
salinity, but RWC and the plant water consumption were
decreased only for higher salt concentrations. This is possibly
due to the fact that at lower salt concentrations water stress may
affect photosynthesis by interfering with the Rubisco effec-
tiveness (Vu and Yelenonsky, 1991). In addition, lowering the
water potential in the protoplast is altering the integrity of
photosynthetic apparatus via photophosphoryliosis and elec-
tron transfer (Karambourniotis, 2003), and only at higher salt
concentrations, water stress may be responsible for stomatal
closure. Moreover, Munns (1993) proposed that the reduction
in leaf cellular turgor is not the main cause for the reduction in
stomatal conductance, net assimilation rate and limited leaf
expansion in saline environments.
It is believed (Ho, 2003) that salinity reduces plant growth by
water stress in the root zone, or salt toxicity in the plants tissue. In
our experiment, salinity influenced the growth of raspberry plants
similarly to its effects on many other species (Parida and Das,
2005). The plants responded to salinity by decreased relative
growth rate, reduced cane length and diameter and smaller
leaves. The number of leaves was reduced only at the highest salt
concentration but the dead leaves (%) were increased with
287
salinity as a mean of protecting the young growing leaves to toxic
levels of the salts as well as off-loading the plants of excess salts
(Wahome, 2001). Additionally, the leaf expansion rate was also
reduced with salinity with accordance to similar results found in
Rosa spp. (Wahome, 2000), Vaccinium ashei (Wright et al.,
1995), Pisum sativum (Hernandez et al., 1995) and azalea (Risse
and Schenk, 1990). In the case of strawberries, the number of
leaves was unaffected by salinity (Saied et al., 2003) and this was
true in our experiment too. In this experiment, the plant’s fresh
and dry weight was reduced by salinity and this could be because
of the decreased photosynthetic capacity (Morsy, 2003).
Furthermore, the total dry weight reduction may attribute to
partial stomatal closure caused by turgor loss in guard cells or
hormonal signal from the roots (Giorgio et al., 1996). Regarding
the root to shoot ratio, it was significantly increased only at the
highest salt concentration and this reduction is generally
recognized to be a general morphological adaptation to
decreased water potentials (Cuartero and Fernandez-Munoz,
1999; Romero-Aranda et al., 2002). Finally, the fresh to dry
weight ratio did not change with salinity and this could be an
evidence that the observed reduction of growth was only partially
depended in the limited water accessibility in plant tissues.
The accumulation of Cl and Na ions in raspberry plants
increased linearly with increased NaCl application in accor-
dance with the results found in Citrus spp. leaves by Banuls and
Primo-Millo (1995). As it was shown earlier (Brunold et al.,
1996), despite the selective nutrient ion uptake, increased salt
concentration in growing medium leads to toxic accumulation
in both halophytes and glycophytes.
The Cl ions concentration increased in all plant parts with
increased NaCl application. The highest Cl ions accumulation
was observed in new and older lower leaves. The transport of Cl
ions occurs mainly in the transpiration stream, which explains
the high concentrations of Cl ions in leaves (El-Sidding and
Luedders, 1994). From the literature, it is revealed that Cl ions
accumulate in higher amounts in lower older leaves and this
explains the occurrence of leaf injury on the lower leaves
(Wahome, 2003) and this occurred in our experiment as well.
Chloride accumulation is probably the main factor for reduced
growth and yield in citrus (Bar et al., 1998), avocado (Wiesman,
1995), kiwi (Chartzulakis, 1991) and grapevines (Walker et al.,
1981), and the results obtained in this study support these
findings.
The Na concentration increased in all plant parts with
salinity. The highest Na accumulation was observed in roots
and lower part of the stems. Investigations on salt-sensitive
plants like Pumica granatum (Doering and Luedders, 1987)
revealed that Na accumulation in shoots could be inhibited by
accumulation of these ions in the lower part of the shoot and
roots. This Na retention in the roots plays an important role in
the adaptation of the glycophytes under salt conditions
(Greenway and Munns, 1980), even though the avoidance of
Na accumulation in shoots of glycophytes is normally limited
especially under long term conditions. This particular pattern
may be suitable in our case as well.
Even though there is no study to report critical leaf Cl and Na
concentrations range in red raspberry, in this study the Na
288 D. Neocleous, M. Vasilakakis / Scientia Horticulturae 112 (2007) 282–289
concentration remained at low, non-toxic levels in leaves and
especially in the younger growing leaves according to Bernstein
and Hayward (1958), who reported that for most crops the salts-
toxic levels are similar. This points out that the salt injury effect,
observed in the present study, could be attributed to Cl toxicity
and Na could be a factor affecting water relation in the roots.
In conclusion, the results obtained in the present study
indicate that the reduction in photosynthesis was mostly linked
to non-stomatal factors, such as the accumulation of toxic ions,
primarily chloride and water imbalance. Particularly, this
reduction may be due to damages in the photosynthetic
apparatus rather than from factors affecting stomatal closure. In
addition, a pronounced toxic effect of Cl was clearly
demonstrated and this points toward the fact that the salt
injury effect could be attributed to Cl toxicity (uptake and
translocation). As a consequence, the limited supply of
metabolic energy and salt toxicity in the plant tissues restricted
plant growth under saline conditions.
Acknowledgements
This study is a portion of the thesis, in partial fulfillment of
the requirements for the degree of Ph.D. Research supported by
Onassis Foundation, Aisxinou 4, 105 58 Athens, Hellas.
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