IJBPAS, September, 2015, 4(9): 5915-5932
ISSN: 2277–4998
EFFECT OF MAGNESIUM ON GROWTH, FRUIT QUALITY AND SUGAR CONTENT IN CUCUMBER UNDER VARIOUS LIGHT INTENSITIES RASOUL AZARMI1*, SEYED JALAL TABATABAEI2, NADER CHAPARZADEH3 1
Department of Horticultural Sciences, Faculty of Agriculture, Tabriz University, Tabriz, Iran 2 3
Faculty of Agriculture, Shahed University, Tehran, Iran
Department of Biology, Faculty of Science, University of Azarbaijan Shahid Madani, Tabriz, Iran *Corresponding Author: E Mail:
[email protected]; Fax: +98 45 32715417 ABSTRACT
The effect of various Mg concentrations (0, 1, 2, 3 and 4 mM) in the nutrient solution on plant growth, fruit quality and sugars content in hydroponicallygrown cucumber (Cucumis sativuscv. Nagen 792) under optimum(100%)and low (50%) light intensities was evaluated.The results showed that the decreases in the dry matter, SPAD index and soluble protein and accumulation of soluble sugar and starch content in the leaves Mg deficiency (0 mM Mg)are suggestive of decreased growth, and the decrease induced by Mg deficiency was bigger under low light intensity than under optimum light intensity. Plant growth was improved at 3 mM Mg, but it was reduced when the Mg concentration increased (4 mM Mg). Concentration of Mg in the leaf and fruit increased drastically with increasing Mg in the nutrient solution. This became steadily more pronounced under low light intensity. Mg deficiency plants (0 mM Mg) developed visible symptoms- interveinal necrosis in middle leaves, especially optimum light intensity.Fruit quality traits such as fruit dry matter percentage and total soluble solid (TSS) and fruit Fv/Fm were increased at higher concentration of Mg (3 mM) in the solution, especially under optimum light intensity. But, fruit firmness was improved at lower concentration of Mg (1 mM) in the solution especially in optimum light intensity In conclusion, Mg requirement of cucumber plants likely increases with light intensity. Thus, higher concentration of Mg (3 mM) in the
5915 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
nutrient solution was the most favorable for cucumber plant growth and fruit quality grown in hydroponics. Keywords: Mg, Growth, Quality, Cucumber, Light intensity INTRODUCTION Mg is an essential element for plant
supply on Mg-deficiency plants caused to
growth and development and fruit quality.
increase fruittotal soluble solid (TSS), dry
Apart from being a central atom of the
matter and juice acidity, [21]. Excessive
chlorophyll molecule, Mg also acts as
supply of Mg to fruit has negative effects
activator or regulator of many key
on fruit firmness, texture and storability
enzymes in plant physiological processes
that
[38,
antagonistic
50].Both
Mg
deficiency
and
are
mainly
determined
by
relationship
with
oversupply have detrimental effects on
Ca[36].Despite
plant photosynthesis [49], consequently
fundamental
roles
resulting in abnormal or restricted growth
metabolism,
there
of plants [50]). Mgplays a fundamental
information on interactions between Mg
roles in phloem export of photosynthates
and light intensity on fruit quality of
so that a deficiency of Mg restricts the
cucumber.
partitioning of dry matter between roots
The responses of plants to different Mg
and shoots to result in excessive sugar,
concentrations are not only affected by
starch and amino acid accumulation in
Mg availability in the root zone, but also
leaves
chlorophyll
depend on light intensity, temperature and
breakdown, an over- reduction in the
species[10, 27]. The roles of Mg in plant
photosynthetic electron transport chain
metabolism
and the generation of highly reactive
conditions are well known [9]. The
oxygen
of
authors indicated that the Mg requirement
CO2
is increased under high-light conditions.
(source
species
impairment
in
tissues),
(ROS)
because
photosynthetic
the
its
well-known
of Mg is
particularly
in
very
under
plant limited
stress
fixation [9,25].
The higher requirement of Mg under high
Mg had a greater effect on quality
light might be reduced to the fact that
parameters, when Mg was supplies to
under suboptimal Mg supply and high
low-Mg plants. However, there were no
light processes are induced which finally
additional benefits when Mg was supplied
lead to accumulation of reactive oxygen
to plants already grown under adequate
species (ROS) and thus plant damage. As
Mg supply conditions [8]. Increasing Mg
plants are subjected to various light
5916 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
intensities at different seasons, this may
variations. The greenhouse was under
alter the ability of plants to take up and
natural
translocate
the
spring and summer and air temperature
adjustment of Mg concentration in the
was set to 27 ± 2 ˚C and 18 ± 2 ˚C in the
nutrient solution according to the light
day and at night, respectively. The
intensity should be crucial. The objective
experiment was a split-plot design with
of this experiment was to determine the
light intensity as the main plot and various
effects of Mg and light intensity on
Mg concentrations as subplot with three
cucumber growth and qualitytraits in
replications in each treatment. Each plot
hydroponics.
contained three plants. The plants were
MATERIALS AND METHODS
treated with five Mg concentrations (0, 1,
Plant materials and growth conditions
2, 3 and 4 mM) as MgSO4.7H2O.
The experiment was carried out at the
Treatments were labelled Mg 0, Mg 1, Mg
Department of Horticultural
Science,
2, Mg 3 and Mg 4. The plants were
University of Tabriz, Iran. Cucumber
subjected to two light intensity treatments
(Cucumis sativus L. cv. Nagen 792) seeds
[optimum
were sown in cells plug trays filled with
intensities] using green shade netting
vermiculite, after emergence of two true
suspended above the box frame with the
leaves, seedlings were transplanted to a
size of 1.5 m × 8 m × 4 m. the box frames
14l growth bags (70, 20,10 cm) filled with
were randomly placed in the greenhouse.
a mixture of perlite and vermiculite (1:1
Everyday light intensity at the canopy
v/v). The nutrient solution was prepared
height under the shaded netting and in the
based on full strength of Hoagland's
glasshouse was monitored using a light-
solution [26]containing: 5.6 mM Ca
meter (Skye Instrument. Powys. UK). The
(NO3)2, 4 mM KNO3, 1 mM KH2PO4. The
average of light intensity under shaded
solution pH was maintained close to 6.5
netting and in the glasshouse (unshaded)
by
over entire period of experimentation is
adding
Mg.
It
H2SO4.
seems
The
that
electrical
photoperiod
(100%)
condition
low
(50%)
during
light
conductivity (EC) of the nutrient solution
shown in Fig.1.
was within the range 2.2 - 2.4 dS m- 1.in
Data collection and chemical analysis
order to keep the anionic-cationic balance
At the end of the experiment, two plants
and a similar electrical conductivity for
from each treatment harvested and the
the five solutions, mineral concentrations
internode diameter were recorded. The
were adjusted leading to only slight
plant organs divided into leaf and stem,
5917 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
weighed and then all plant parts dried at
Japan). Titratable acidity was measured
80 ˚C in an air-forced oven for 48 h for
by titrating with 0.1 M NaOH to the
determination of leaf and stem dry matter.
neutralization point. Before chlorophyll
The percentage leaf and stem dry weight
fluorescence measurements, fruits were
was then calculated as below: [(Dry
dark-adapted for 30 min using a dark
weight/ Fresh weight) ×100]. Chlorophyll
towel. Measurements were taken in the
index value of fully expanded young
middle and near the neck position
leaves was determined using a portable
positions of each fruit at the same location
SPAD-502 meter (Minolta, Tokyo, Japan)
in the fruit surface and then averaged. The
during the period of the plant's growth.
maximal
Fruit
a
photochemistry (Fv/Fm) was measured
representative sample collected at the
using a plant efficiency analyzer, Handy
same position from plants in each
PEA (Hansatech Instruments). Ten fruits
treatment. The samples were taken from
per treatment per replicate were used for
fruits with the same size. Each fruit was
the determination of fruit dry weight.
cut in to pieces and homogenized in a
Each individual fruit from each treatment
conventional blender in order to obtain the
was placed in a sampling bag and dried in
fruit juice. Thereafter, the fruit juice was
the oven at a temperature of 80°C for 48 h
filtered using a Whitman No. 4 filter
until a constant weight was obtained. The
paper and the filtrate was used to
percentage dry weight was calculated as
determine the pH, EC and TSS. The TSS
below: [(Dry weight/ Fresh weight)
content of the fruit was determined by
×100].
using a digital refractometer (Atago Co.,
Soluble sugars were extracted using the
Tokyo, Japan). The juice pH and EC was
method
measured by pH meter and EC meter,
About 0.5 g of dried leaf samples were
respectively.
extracted three times in 5ml of hot 80 %
quality
was
The
measured
measurement
in
fruit
quantum
described
yield
by
of
PS
II
Sheligl(1986).
a
ethanol (80 ˚C)[51]. The supernatants
penetrometer (Model: ST 977. Italy). A
from each extraction were combined and
thin layer of the middle of fruit skin (0.5
made to a convenient volume. 1 ml 5 %
mm) was removed by a sharp razor and
(w/v) phenol and 5ml concentrated H2SO4
fruit color or the extent of greening was
were added to 2 ml the plant extract and
measured
chlorophyll-meter
mixed thoroughly. The reaction mixture
(SPAD-502, Konica, Minolta, Osaka,
was allowed to stand for 30 min before
firmness
was
using
determined
a
using
5918 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
the absorbance was recorded at 485 using
Statistical Analysis
a spectrophotometer (Motic, CL-45240-
A statistical analysis was made using
00, Hong Kong, china). Total sugar
analysis of variance the SPSS 21 software
content of the sample was calculated
and the means were separated by the
based on calibration curve from a glucose
Duncan test at a significance level of 0.05.
working standard.Starch
The graphs were drawn using Excel
content
was
extracted from the residual plant material
software.
from
RESULTS AND DISCUSSION
the
soluble
sugar
extraction
described above. This was done by
Vegetative growth
incubating the dry pellet with 2 ml HCl
The results showed that the highest leaf
(4.68M) in boiling water bath for 15 min.
dry matter percentage was obtained in 3
the soluble products were assayed by the
and 4 mM Mg treatments. The leaf dry
same phenol-sulphuric method described
matter percent in concentration of 0 mM
above.
was
Mg was slightly higher than concentration
determined in according to Bradford
of 1 mM Mg. Low light intensity largely
(1976) using bovine serum albumin as
decreased
standard[5]. To measure the Mg, Leaves
compared to optimum light intensity
and fruits washed with distilled water
(Table
were oven-dried at 80 ˚C for 48 h and
agreement with the finding of Lasa et al
weighed. The dry samples were ground to
(2000) who showed that concentration of
pass through a 0.5-mm screen. 1 g dry
0 mM Mg decreased 40 – 50% of shoot
samples of leaves and fruits a were soaked
biomass compared with Mg sufficient
in 10mL nitric acid (HNO3) for 24 h then
plants in sunflower plants[32]. Low light
digested in digestion systems in a fume
intensity
hood, heated to 110 ˚C for 3 h. The
photosynthates from vegetative organs to
extracted solution was transferred to 100
the fruits. The reason for increase in leaf
mL volumetric flasks, and then diluted to
dry matter at 0 mM Mg may be due to
100mL with deionized water for Mg
impaired export of carbohydrates from
assays [35]. The Mg concentration in the
source to sink sites and accumulation of
leaf and fruit were measured at a
soluble sugars in source leaves. Stem dry
wavelength
matter was not affected by various Mg
Soluble
absorption
protein content
285.2
nm
by
spectrophotometry
Elmer, Model 110, and USA).
atomic (Perkin-
1).
leaf
dry
This
observation
reduces
concentrations,
matter
but
the
percent
is
export
optimum
in
of
light
intensity significantly increased stem dry
5919 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
matter than low light intensity (Table 1).
Soluble
In the present experiment, there was a
soluble protein
significant difference in leaf SPAD value
The results showed that under both light
between low Mg concentrations and
intensities, concentration of 0 mM Mg in
sufficient Mg concentrations (Table 1).
the solution had higher leaf soluble sugar
Significant
chlorophyll
and starch content compared to other
concentration in Mg deficiency leaves has
treatments. However, leaf soluble sugar
been widely reported[23, 55]. A reason
and starch content in optimum light
for higher chlorophyll content under
intensity was higher than in low light
adequate Mg supply could be an enhanced
intensity (Table 1). In almost all higher
production of chlorophyll and chlorophyll
plants, the principle end products of leaf
associated proteins. It is well documented
photosynthates are sucrose and starch.
that chlorotic and necrotic symptoms
However, partitioning of sucrose and
appearance in Mg deficiency leaves is
starch and their effect on dry matter
associated with chlorophyll destruction
distribution is influenced by several
due to photo-oxidation and accumulation
environmental
of soluble sugar and starch in source
temperature, drought and essential mineral
leaves [7].
In both light intensities,
nutrients [28, 56]. Mineral nutritional
internode diameter was increased with
status of plants has a considerable impact
increasing
the
on partitioning of carbohydrates and dry
solution. However, internode diameter
matter between shoots and roots [15,
was higher under optimum light intensity
34,38]. Under Mg deficiency, starch
compared to low light intensity (Table 1).
concentrations
Light quantity and quality are major
leaves[18] and low in sink organs such as
determinants
cereal grains and fruits [2]. This may
decrease
Mg
in
concentration
of
internode
in
growth.
and
insoluble
factors,
are
sugars
such
high
as
low
source
Reductions in both photosynthetically
demonstrate
active radiation (PAR), and red: far-red
transport from source leaves to sink
ratio (R: FR) result in similar shade
organs. Hence, in Mg-deficient plants
avoidance responses, such as increased
higher shoot/root
internode
thicker
compared with Mg-sufficient plants[4, 10,
internode. In plant communities, the R:
16]. The translocation of amino acids and
FR ratio seems to act as an early
sugars from sink to source might be
competition signal [20, 31].
inhibited under magnesium deficiency
elongation
and
impaired
in
and
photosynthate
ratios were found
5920 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
because of the effect of Mg on the H+-
appeared only in 0 mM Mg concentration
ATPase [9]. The results clearly showed
and under both light intensities. However
that by increasing Mg concentration in the
the symptoms severity became more
solution, increased soluble protein content
pronounced
in the cucumber leaves (Table 3).
intensity. These symptoms observed after
Andrews et al (1999) reported that Mg
35 days of treatment initiation and in
deficiency induced an increase in the
middle leaves as necrotic lesion. Whereas,
protein content of Pisun sativum and
no visual symptoms of Mg deficiency
Phaseolus vulgaris[1]. The reduction of
were found in leaves of any other
protein in Mg deficiency plants could be
treatments in the range of 1 to 4 mM Mg
attributed to
a decrease in protein
concentrations, under both light intensity.
synthesis due to the participation of Mg in
The incidence of Mg deficiency was
the aggregation of ribosome subunits and
attenuated by the initial amount Mg
its requirement for RNA polymerases
present within the plant. Because the
[12]. Protein biosynthesis also is strongly
cucumber seedlings had been grown in
reduced under Mg deficiency leading to
one
increased concentrations of the precursor
(containing 0.3 mM Mg) for four weeks
amino acids [19,40].
prior to treatment initiation, the initial
Leaf and fruit Mg concentration
accumulated Mg and its internal recycling
The results indicated that with increased
in the seedling attenuated the visible signs
Mg concentration in the nutrient solution
of Mg deficiency. The Mg concentration
led to a significant increase in Mg content
sufficient for optimal growth varied with
in leaves and fruits under both light
species. Kirkby and Mengel, (1979)
intensity.
reported that 0.35- 0.8% in the dry weight
But,
Mg concentration
in
third
of
full
nutrient
solution
light
However, the results obtained in this
higher
than
in
well
with
cucumber[29].
cucumber leaves under optimum light
study
intensity (Fig. 2). Greater concentration of
threshold line for the occurrence of Mg
Mg was observed with increasing Mg
deficiency determined by Kirkby and
levels in the nutrient solution. However,
Mengel, (1979)[29]. The Mg- deficiency
Mg concentration of leaf in shaded plant
visible symptoms observed partially only
was higher than in unshaded plants.
on the full developed middle leaves [6, 7,
Visual
19, 43]. In cucumber, deficiency visible
symptoms of Mg deficiency
agree
for
light
to
was
sufficient
optimum
cucumber leaves and fruits under low intensity
be
under
the
general
5921 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
symptoms observed initially as interveinal
that the firmest fruit under optimum light
chlorosis
interveinal
intensity in 1 and 2 mM Mg and in low
The occurrence of
light intensity at 1 mM Mg were obtained.
and
finally,
necrosis on leaves.
as
Mg- deficiency on the middle leaves
Underoptimum
could
the
conditions,the fruits were firmer than
photoassimilate production and supply to
under low light intensity conditions. This
other parts of plants. Both shading and
is consistent with findingby Marcelle,
Mg levels in the nutrient solution altered
(1995) who showed that an optimal Mg
Mg concentration in the leaves. This is
concentration ‘has to be relatively low’
consistent with findings by Zhao and
for good storage properties[38]. The
Oosterhuis (1998) and Sonneveld (1987)
reduced firmness in plant grown with low
who indicated that high light intensity will
Mg content
decrease the ability of plants to absorb and
concentration of fruit Ca. Fruit Firmness
translocate magnesium, since transpiration
is an extremely important quality attribute
is reduced and the translocation of
of cucumber and consumer prefer a firm
magnesium is driven by transpiration
and crisp product. The Mg: Ca ratio
rates[57, 54].In general, the breakdown of
mainly determines service and stability
chlorophyll under magnesium deficiency
aspects as components of the total food
is associated with the accumulation of
quality, such as product firmness, texture
sugars and starch in the cells of deficient
and storability that are mainly determined
leaves [24, 9]. This causes an over-
by the role of Ca in stabilizing cell
reduction of the photosynthetic electron
walls[21]. Since Mg is capable of
transport chain, which leads to the
replacing
formation of reactive oxygen species.
imbalance Mg: Ca ratios in the tissue
Fruit quality traits
often negatively affect product quality.
Despite the well-known roles of Mg in
The fruit TSS increased with increasing
plant
limited
Mg concentration up to 3mM Mg and
information there have been concerning
then decreased with increasing Mg (4
the significance of Mg for the quality of
mM). The fruit TSS under optimum light
agriculture and horticulture produce, as
intensity was higher than under low light
compared to other major nutrients.A fruit
intensity. The increase of fruit juice TSS
quality trait like fruit firmness was
with increasing Mg concentration in the
significantly affected by treatments. So
solution reported by Quaggio et al (1992)
significantly
metabolism,
affect
very
Ca
light
may
be
from
intensity
due
to
binding
high
sites,
5922 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
who indicated that juice pH, Soluble Solid
application
and titratable acidity of fruit orange
allocation to the fruit, whereas the
increased with increasing Mg[45]. Mg:K
allocation
ratio
influence primarily
pointing to the decisive role of Mg in
quality properties through the role of both
carbohydrate partitioning[22].The highest
mobile cations in metabolite formation
value in fruit Fv/Fm was obtained in 2 and
and translocation to fruits [46]. The
3 mM Mg treatments. Fruit Fv/Fm was
highest marketable, high quality yield was
higher under low light intensity than that
observed when K and Mg were supplied
under optimum light intensity (Table 2).
in the highest amounts at a ratio of 5:1.
Chlorophyll fluorescence is an indirect
This clearly points to the facts that only
measurement of the physiological status
balanced crop nutrition can result in
of green tissues [48, 41], being used in
optimal quality [3].Fruit juice pH and EC
both green leaves and several chlorophyll-
were not affected by the treatments (Table
containing fruits [52, 53, 13]. Concerning
3).Fruit
with
the evaluation of changes in fruit tissues,
the
chlorophyll fluorescence measurement has
solution. Low light intensity decreased
the advantage of detecting cellular injury
fruit dry matter (Table 2). Dry matter
due
content
environmental stresses in advance to the
appears to
dry
increasing
matter
Mg
is
concentration
an
criterion.Marcelis
increased
important
quality
the
natural
leaves
biomass
decreased,
senescence
or
development of visible symptoms [14, 52,
positive relationship between the dry
47].Fruit chlorophyll was significantly
matter distribution of the fruit and
higher under optimum light intensity than
irradiance[37].Increases of dry matter
under low light intensity (Table 2). Fruit
percent can be due to the importance of
color is one of the few practical criteria
Mg for photosynthesis and assimilate
for assessing cucumber quality after
translocation. This finding was underlined
harvest at present. A dark green cucumber
by Feltran et al (2004) and Poberezny and
is expected to have a longer shelf-life than
Wszelaczynska (2001) who showed that
a light green cucumber [33]. This result is
increasing
consistently
agree with observation of Klieber et al
potato[17,
(1993) who showed that high light
and
intensity is necessary for high chlorophyll
Papadopoulos(2004)indicated that at a
content in cucumber[30]. They also
given Ca supply increasing the Mg
confirmed that high chlorophyll content
increased 44].Also,
dry
showed
to
to
the
a
Mg
(1993)
in
enhanced
supply matter Hao
in
5923 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
was positively correlated with a high
[4] Bouma D, Dowling EJ, Wahjoedi
percentage of PPFD reaching the fruit
H.
surface.
potassium and magnesium on the
CONCLOSION
growth of subterranean clover
It can be concluded that Mg requirement
(Trifolium subterraneum). Annals
of cucumber plants likely increases with
Botany 43529–538.
light
intensity.
The
moderate
1979.
Some
sensitive
solution
quantitation
the
most desired
for
of
[5] Bradford MM. 1976. A rapid and
concentration of Mg (2 mM) in the was
effects
method
for
of
the
microgram
cucumber fruit quality. While, higher
quantities of protein utilizing the
concentration of Mg (3 mM) in the
principle of protein dye- binding.
solution was the most favorite for
Analytical Biochemistry72, 248–
cucumber growth in hydroponics system.
254. [6] Broschat
REFERENCES [1] Andrews M, Sprent JI, Raven J A,
TK.
1997.
distribution,
Nutrient
dynamics,
and
Eady P E. (1999). Relationships
sampling in coconut and Canary
between shoot to root ratio, growth
Island date plants. Journal of the
and
American Society for Horticultural
leaf
soluble
protein
concentration of Pisum sativum, Phaseolus vulgaris and Triticum
Science 122, 884-890. [7] Cakmak I. 1994. Activity of
aestivum under different nutrient
ascorbate-dependent
deficiencies.
scavenging
Plant
Cell
and
Environment22, 949–958.
enzymes
chlorosis
[2] Beringer H, Forster H. 1981.
H2O2-
are
magnesium-
and
leaf
enhanced
in
and
Einfluss variierter Mg-ernahrung
deficient
auf tausendkorngewicht und P-
phosphorus–deficient
fraktionen des gerstenkorns. Z
Journal of Experimental Botany
Pflanzenernahr Bodenk 144: 8–15.
278, 1259-1266.
[3] Borkowski J, Szwonek E. 1986. Effect
of
potassium
and
magnesium on the quality of tomato fruits. Acta Horticulture 191: 133–139.
leaves,
Potassiumbut
not
in
leaves.
[8] Cakmak I. 2013. Magnesium in crop production, food quality and human health. Plant Soil 368: 1-4. [9] Cakmak I, Kirkby EA. 2008. Role of
magnesium
in
carbon
5924 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
partitioning
and
alleviating
photooxidative
damage.
firmness. In: Proceedings of the International
Conference
Physiologia Plantarum 133: 692–
Sensors
704.
Testing: Measuring the Quality
[10] Cakmak I, Marschner H. 1992. Magnesium- deficiency and high light intensity enhance activities of
superoxide
ascorbate
dismutase,
peroxidase
and
for
on
Nondestructive
of Fresh Fruits and Vegetables, p. 67–73. [15] Druege U, Zerche S, Kadner R, Ernst M. 2000. Relation between nitrogen
status,
carbohydrate
glutathione reductase in bean
distribution
and
leaves. Plant Physiology 98:
rooting
1222–1227.
cuttings as affected by pre-
of
subsequent
chrysanthemum
[11] Cakmak I, Yazici AM. 2010.
harvest nitrogen supply and cold-
Magnesium: a forgotten element
storage. Annals of Botany 85(5):
in crop production. Better Crops
687-701.
94: 23-25.
[16] Ericsson T. 1995. Growth and
[12] Cammarano P, Felsani A, Gentile
shoot root ratio of seedlings in
M, Gualerzi C, Romeo C, Wolf
relation to nutrient availability.
G. 1972. Formation of active
Plant and Soil 168/169: 205-14.
hybrid
80-S
from
[17] Feltran JC, Lemos LB, Vieites
subunits of pea seedlings and
RL. 2004. Technological quality
mammalian Biochemistry
particles
liver
ribosomes.
and utilization of potato tubers.
and
Biophysica
Scientia Agricola 61: 593–597.
Acta281: 625–642.
[18] Fischer ES, Bremer E.
[13] DeEll JR, Toivonen PMA. 2000.
Influence
of
magnesium
Chlorophyll fluorescence as a
deficiency
nondestructive
expansion, starch and sucrose
indicator
of
on
1993.
rates
of
broccoli quality during storage in
accumulation,
modified-atmosphere packaging.
assimilation
Hort Science 35: 256–259.
vulgaris. Physiologia Plantarum
[14] DeEll JR, Prange RK, Murr DP.
and
leaf
in
net
Phaseolus
89: 271-276.
1997. Chlorophyll fluorescence
[19] Fischer ES, Lohaus G, Heineke
as an indicator of apple fruit
D, Heldt HW. 1998. Magnesium
5925 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
deficiency
results
in
[25] Hermans C, Bourgis F, Faucher
accumulation of carbohydrates
M, Delrot
and amino acids in source and
Verbruggen N. 2005. Magnesium
sink
spinach.
deficiency in sugar beet alters
Physiologia Plantarum 102: 16-
sugar partitioning and phloem
20.
loading in young mature leaves.
leaves
of
[20] Franklin KA, Whitlam GC. 2005. Phytochromes
and
S, Strasser RJ,
Planta 220: 441-449.
shade-
[26] Hoagland DR, Arnon DS. 1950.
avoidance responses in plants.
The water culture method for
Annals of Botany 96: 169–175.
growing plants
[21] Gerendas J, Fuhrs H. 2013. The
without
California
soil.
Agricultural
significance of magnesium for
ExperimentStation Circular 374:
crop quality. Plant Soil 368: 101-
1–32.
128.
[27] Huang JW, Welch RM, Grunes
[22] Hao X, Papadopoulos AP. 1999.
DL. 1990. Magnesium, nitrogen
Effects of supplemental lighting
form,
and cover materials on growth,
effects on grass tetany potential
photosynthesis,
of
partitioning,
biomass
early
yield
and
quality of greenhouse cucumber. Scientia Horticulture 80: 1-18. [23] Hariadi Y, Shabala S. 2004.
and
wheat
root
forage.
temperature
Agronomy
Journal 82(3): 581–587. [28] Huber SC. 1989. Biochemical Mechanism for Regulation of Sucrose Accumulation in Leaves
Screening broad beans (Vicia
during
faba) for magnesium deficiency.
physiology 91(2): 656- 662.
ll. Photosynthetic performance
[29] Kirkby EA, Mengel K. 1979. The
and leaf bioelectrical responses.
role of magnesium in plant
Functional Plant Biology 31:
nutrition. - Z. Pflanzenernahr.
539-549.
Bodenkd 2: 209-222
[24] Hermans C, Verbruggen N. 2005. Physiological
Photosynthesis.
Plant
[30] Klieber A, Lin WC, Joliffe PA,
characterization
Hall JW. 1993. Training methods
of Mg deficiency in Arabidopsis
affect canopy light exposure and
thaliana. Journal of Experimental
shelf
Botany 56: 2153-2161.
cucumber. Journal of American
life
of
long
English
5926 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
Society Horticulture Science 118, 786–790.
[36] Malundo TMM, Shewfelt RL, Scott JW. 1995. Flavor quality of
[31] Kurepin LV, Emery RJN, Pharis
fresh
tomato
(Lycopersicon
RP, Reid DM. 2007. Uncoupling
esculentum Mill.) as affected by
light quality from light irradiance
sugar and acid levels. Post
effects
Biology Technology 6:103–110.
in
Helianthus
annus
shoots: putative roles for plant
[37] Marcelis
LFM.
1993.
Fruit
hormones in leaf and internode
growth and biomass allocation to
growth. Journal of Experimental
the fruits in cucumber. 1. Effect
Botany 58: 2145–2157.
of fruit load and temperature.
[32] Lasa B, Frechilla S, Aleu M, Gonzales-Moro B, Lamsfus C, Aparicio- Tejo PM. 2000. Effects
Scientia Horticulturae 54: 107– 121. [38] Marcelle RD. 1995.
Mineral
of low and high levels of
nutrition and fruit quality. Acta
magnesium on the response of
Horticulture 383:219–226.
sunflower plants grown with
[39] Marschner H. 1995. Mineral
ammonium and nitrate. Plant and
nutrition of higher plants. 2nd
soil 225:167-174.
end.
[33] Lin
WC,
Ehret
DL.
1991.
Nutrient concentration and fruit
(Academic
Press.
San
Diego, CA) [40] Marschner
P.
(ed)
2012.
thinning affect shelf life of long
Marschner mineral nutrition of
English cucumber. Hortscience
higher plants (Third Edition).
26 (10): 1299–1300.
Elsevier Ltd.
[34] López-Bucio J, Cruz-Ramirez A,
[41] Maxwell K, Johnson GN. 2000.
Herrera-Estrella L. 2003. The
Chlorophyll
role of nutrient availability in
practical
regulating
Experimental Botany 51: 659–
root
architecture.
Current Opinion in Plant Biology 6(3): 280-287.
fluorescence
guide.
Journal
a of
668. [42] Mengel K, Kirkby EA. 1987.
[35] MAFF. 1985. The analysis of
Principles of plant nutrition, 4th
agricultural materials. Ministry
Edition,
of Agriculture
Institute, Switzer-land.
Fisheries
and
International
Potash
Food, London, UK.
5927 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
[43] Papenbrock J, Pfundel E, Mock HP, Grimm B. 2000. Decreased and increased expression if the
Caricapapaya. Canadian Journal Botany 70: 364–373. [48] Schreiber U, Bilger W. 1987.
subunit CHL1 diminishes Mg
Rapid
assessment
chelatase activity and reduces
effects
on
chlorophyll
chlorophyll
synthesis
in
plant
stress
leaves
by
fluorescence
transgenic tobacco plants. The
measurements.In:
Plant Journal 22: 155-164
J.D.,
[44] Poberezny J, Wszelaczynska E.
of
Catarino,
Tenhunen, F.M.,
Lange,
O.L., Oechel, W.C. (Eds.), Plant
2011. Effect of bioelements (N,
response
to
stress-functional
K, Mg) and long-term storage of
analysis
in
Mediterranean
potato tubers on quantitative and
ecosystems.
qualitative
II.
science institute Series. Springer-
Content of dry matter and starch.
Verlag Berlin Heidelberg New
Journal of Elementology 16:
York p. 27–53.
losses
Part
237–246. [45] Quaggio
NATO
advanced
[49] Shabala S, Hariadi Y. 2005. JA,
Sobrinho
JT,
Effects of magnesium availability
Dechen AR. 1992. Magnesium
on
the
activity
of
influences on fruit yield and
membrane ion transporters and
quality
of
orange
on
‘Valencia’
sweet
light-induced
responses
Rangpur
lime.
broad
Leaf
Proceedings International Society Citriculture 2: 633–637.
bean
plasma
from
mesophyll.
Planta 221(1): 56-65. [50] Shaul
O.
2002.
Magnesium
[46] Romheld V, Kirkby EA. 2007.
transport and function in plants;
Magnesium functions in crop
the tip of the iceberg. Biology of
nutrition and yield. Proceedings
Metals 15: 309- 323.
of a Conference in Cambridge (7th Dec. 2007), 151–171. [47] Sanxter SS, Yamamoto
HQ.
1986.
Die
verwertung orgngischer souren HY,
Fisher DG, Chan HT. 1992. Development
and decline of
chloroplasts
in
exocarp
[51] Sheligl
of
durch chlorella lincht. Planta Journal 47-51. [52] Smillie RM, Hetherington SE, Nott R, Chaplin GR, Wade NL. 1987. Application of chlorophyll
5928 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
fluorescence to the postharvest
Bertic B. 2009. Relationship
physiology and storage of mango
among
and banana fruit and the chilling
concentration
tolerance of mango cultivars.
meter readings in soybean under
Asian Food Journal 39: 55–59.
influence of foliar magnesium
[53] Song F, Deng W, Beaudry RM.
chloroplast and
application.
pigments chlorophyll
Communicationsin
1997. Changes in chlorophyll
Soil Scienceand Plant Analysis
fluorescence of apple fruit during
40: 706-725.
maturation,
ripening,
and
[56] Wardlaw A. 1990. The control of
senescence.
HortScience
32:
carbon partitioning in plants.
891–896.
New Phytologist 116(3): 341-
[54] Sonneveld C. 1987. Magnesium
381.
deficiency in rockwool-grown
[1] Zhao D, Oosterhuis DM. 1998.
tomatoes as affected by climatic
Influence of shade on mineral
conditions and plant nutrition.
nutrient
Journal of Plant Nutrition 10:
cotton. Journal of Plant Nutrition
1591–1604.
21(8): 1681–1695,
status
of
field-grown
[55] Teklic T, Vrataric M, Sudaric A, Kovacevic V, Vukadinovic V, Table 1: The effect of Mg and light intensity on cucumber vegetative growth Light intensity
Mg (mM) 0 1 2 3 4
Leaf Dwt (%) 14.16 13.55 14.44 16.63 15.36
Stem Dwt (%) 7.60 7.85 7.80 8.02 7.75
Internode diagonal (cm) 3.85 4.46 4.54 4.79 4.11
Leaf SPAD index 55.30ef 58.70c 59.50bc 61.30ab 62.63a
Soluble sugar (mg g-1 DW) 35.82 31.41 25.42 23.73 24.39
0 1 2 3 4
11.23 11.20 11.74 13.17 12.13
6.29 6.66 7.00 6.94 6.60
3.83 4.05 4.22 4.00 3.83
53.76f 54.33ef 55.06ef 56.23de 57.43cd
28.18 24.73 20.35 19.48 18.74
Optimum Light Low Light
14.83 11.89
7.80 6.70
4.35 3.98
59.48 55.36
28.15 22.29
Light intensity Mg Light intensity×Mg
0.001 0.01 ns
0.001 ns ns
0.001 0.01 ns
0.001 0.001 0.05
0.001 0.001 ns
Optimum Light
Low Light
(ns) non significance; (0.05) significance at 0.05 probability level; (0.01) significance at 0.01 probability level; (0.001) significance at 0.001 probability level. Within each column, same letters indicate no significant difference between treatments (P < 0.01
5929 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
Table 2: The effect of Mg and light intensity on cucumber fruit quality traits Light intensity
Mg (mM) 0 1 2 3 4
Firmness (Kg) 3.66bc 3.73ab 4.20a 3.30bc 3.26bc
Fruit dry matter (%) 3.92 4.83 4.35 4.90 4.71
Fruit Fv/Fm value 0.737 0.744 0.753 0.760 0.741
Fruit color (SPAD UNIT) 53.83 56.40 57.20 56.96 55.94
0 1 2 3 4
3.10bc 3.66a 3.63bc 3.23bc 3.33bc
4.21 4.31 4.48 4.46 4.67
0.749 0.751 0.763 0.764 0.751
51.10 53.96 53.23 53.03 50.76
Optimum Light Low Light
3.63 3.39
4.85 4.42
0.747 0.756
56.07 52.42
Light intensity Mg Light intensity×Mg
0.01 0.001 0.05
ns 0.001 ns
0.01 0.01 ns
0.01 ns ns
Optimum Light
Low Light
(ns) non significance; (0.05) significance at 0.05 probability level; (0.01) significance at 0.01 probability level; (0.001) significance at 0.001 probability level. Within each column, same letters indicate no significant difference between treatments (P < 0.01)
Table 3: The effect of Mg and light intensity on cucumber fruit quality traits Light intensity
Mg (mM)
Fruit juice pH
Optimum Light
0 1 2 3 4
Low Light
0 1 2 3 4
Optimum Light Low Light
Acidity (%)
Soluble protein (mg g-1 FW)
Fruit Mg (mgg-1 DW)
5.93 5.86 5.93 5.74 5.83
Fruit juice EC (dS m-1) 0.50 0.56 0.56 0.56 0.56
1.993 2.213 2.327 2.417 2.650
1.00 1.22 1.40 1.57 1.61
0.720 1.253 2.133 2.727 3.047
5.72 5.64 5.69 5.55 5.63
0.46 0.56 0.56 0.55 0.56
1.703 1.960 2.027 2.193 2.387
0.83 0.91 1.24 1.45 1.43
0.880 1.480 2.687 2.753 3.213
5.86 5.65
0.55 0.54
2.320 2.054
1.36 1.17
1.97 2.16
Light intensity 0.05 ns ns 0.01 0.01 Mg ns ns ns 0.001 0.001 Light intensity×Mg ns ns ns ns ns (ns) non significance; (0.05) significance at 0.05 probability level; (0.01) significance at 0.01 probability level; (0.001) significance at 0.001 probability level. Within each column, same letters indicate no significant difference between treatments (P < 0.01)
5930 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
Unshaded
1200
Shaded
Light intensity (µmol m-1s-1)
1000 800 600 400 200 0 May
Jun
July
Agaust
September
October
November
Months Fig. 1: Light intensity at the optimum (100%) and low (50%) light intensity during the entire of period of experimentation
0M
1M
2M
3M
4M
Leaf Mg concentration (mg g-1 DW)
6 5 4 3 2 1 0 Optimum light
Low light
Fig. 2: The effect of Mg and light intensity on the leaf Mg concentration in cucumber plants (error bars on the columns represent standard error)
5931 IJBPAS, September, 2015, 4(9)
Rasoul Azarmi et al
Research Article
0M
1M
2M
3M
4M
6
Fruit TSS (%)
5 4 3 2 1 0 Optimum light
Low light
Fig. 3: The effect of Mg and light intensity on fruit TSS in cucumber plants (error bars on the columns represent standard error)
Starch content (mg g-1 DW)
0M
1M
2M
3M
4M
90 80 70 60 50 40 30 20 10 0 Optimum light
Low light
Fig. 4: The effect of Mg and light intensity on starch content in cucumber plants (error bars on the columns represent standard error) A
B
Fig. 5: Visual symptoms of Mg sufficient leaves (A) and Mg deficiency leave (B)
5932 IJBPAS, September, 2015, 4(9)