Communications in Soil Science and Plant Analysis

ISSN: 0010-3624 (Print) 1532-2416 (Online) Journal homepage: http://www.tandfonline.com/loi/lcss20

Applications in sustainable production

G. Vllora , D. A. Moreno , J. M. Ruiz , R. Rivero , J. Hernndez & L. Romero
To cite this article: G. Vllora , D. A. Moreno , J. M. Ruiz , R. Rivero , J. Hernndez & L. Romero
(2000) Applications in sustainable production, Communications in Soil Science and Plant
Analysis, 31:11-14, 2309-2320, DOI: 10.1080/00103620009370585
To link to this article: http://dx.doi.org/10.1080/00103620009370585

Published online: 11 Nov 2008.

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COMMUN. SOIL SCI. PLANT ANAL., 31(11-14), 2309-2320 (2000)

Effect of External NK Concentrations on the Leaf Content of

Divalent Cations, Their Forms and Fractions in Capsicum
Plants
G. Vllora,a D. A. Moreno,a J. M. Ruiz,a R. Rivero,a J. Hernndez,b and L. Romeroa
a

Dpto. Biologa Vegetal, Facultad de Ciencias, Universidad de Granada, E-18071 Granada, Spain
Dpto. Produccin Vegetal. EUP Ingeniera Agrcola, Universidad de Almera, 04120-Almera,
Spain

ABSTRACT
Capsicum plants grown under controlled conditions underwent crossed fertilization with N (four
levels) and K (three levels) totalling 12 combination of NK dosages. The plants were sampled
throughout their growth cycle and the leaves were analyzed to measure total and soluble Ca and
Mg, and their fractions. Soluble Mg showed its lowest leaf level in the treatment N4K3, whereas
the highest value was obtained at N1K3. Total Mg has the maximum leaf levels at N1K2, whereas
the lowest levels occurred in the treatment N2K1. The various fractions of Mg showed different
degrees of importance and in the present study the order of importance was:
Pectate-Mg<Residual-Mg<Organic acid anion-Mg<Organic anion-Mg<Chlorophyll-Mg.
Soluble Ca showed its lowest leaf level in the treatment N3K3, whereas the highest value was
obtained at N4K1. Total Ca has the maximum leaf levels at N4K3, whereas the lowest levels
occurred in the treatment N1K2. The increasing NK supply directly or indirectly affects the
integration of the ionic forms of Ca in plant tissues as Ca was integrated in proportions which
indicated the growing conditions of the crop, with the following order:
Oxalate-Ca > Inorganic-Ca > Pectate-Ca Residual-Ca > Phosphate-Ca
INTRODUCTION
The mobility of Ca2+ from cell to cell and in the phloem is very low, and it is the only
mineral nutrient other than, possibly, B which functions mainly outside the cytoplasm in the
apoplast (Douglas, 1993). Most of its activity is related to its capacity for coordination, by which
it provides stable but reversible inter-molecular linkages, predominantly in the cell walls and at
the plasma membrane (Shroeder and Thuleau, 1991). These Ca2+-mediated linkages respond to
local changes in environmental conditions and are part of the control mechanism for growth and

2309
Copyright 2000 by Marcel Dekker, Inc.

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VILLORAETAL.

developmental processes (Hanson, 1984). Ca Uptake can be strongly depressed by other cations,
such as K+, NH 4 + (Kurvits and Kirkby, 1980), and Mn 2t (Heenan and Campbell, 1981). The
functions of Mg2* in plants are related to its mobility within the cells, its capacity to interact with
strongly nucleophilic ligands throught ionic bonding and to act as a bridging element and/or form
complexes of different stabilities. Although most bonds involving Mg2+ are ionic, some are
partially covalent, as in the chlorophyll molecule (Marschner, 1995). A high proportion of the
total Mg2+ is involved in the regulation of cellular pH and the cation-anion balance (Sigel and
Sigel, 1990). The distribution and partitioning of both divalent cations is not only species
dependent but also depends upon the growth stressing conditions such drougth, ligth, nutrient
supply and nutritional status of certain elements as the case of K, which at the "luxury range"
could affects the uptake and bioavailability of Ca and Mg. Moreover, K+ is involved in cell wall
acidification (Kochian and Lucas, 1993), cell wall extension and osmotic regulation (Lindhauer,
1989). Respect to the N, its application to the soil affects Ca and Mg plant levels depending on
the applied form. Thus, NH4+ usually reduced the Ca and Mg uptake, whereas NO3" increased Ca
uptake (Mills and Jones, 1996).
It is well stablished that the N and K applications to the soil affects the uptake and
transport of Ca and Mg, as a result of this effect, the fractions of both elements in the plant
tissues is affected by the nutritional composition of the growth media (Marschner, 1995). Thus,
N and K affects the Ca and Mg fractions, and the aim of the present study was to determine the
effect of differing NK combination on forms and fractions of the divalent cations present in the
leaf in capsicum plants.
MATERIALS AND METHODS
Experimental Design
Pepper plants (Capsicum annuum L. cv. Lamuyo) were grown in cell flats (cell size 3 x 3 x
10 cm) filled with peat-lite mixture, placed on benches under the greenhouse conditions described
below, for a period of 8 weeks; then seedlings were transplanted and grown under controlled
conditions in an experimental greenhouse at Centro de Investigacin y Desarrollo Hortcola, El
Ejido, Almera, Spain. The soil used was loamy-sand with the following characteristics: sand
(37.3%), silt (48.6%) and clay (10.1%), CaCO3 (26.82%), total N (3.5 g kg 1 ), total organic C (36.1
g kg"1), PO4"3 (890 mg kg'), K* (5.34 g kg") pH (H2O, 8.45; KC1, 8.01), electrical conductivity
(E.C.= 4.63 dS m'1). The soil surface was covered with a 7-cm layer of sand. The relative humidity
was between 60-80% and the temperature range with extremes of 20 and 30 C in the greenhouse.
The experimental design was a factorial arrangement in a randomized complete block with twelve
treatments. Container-grown capsicum plants were transplanted into 2 rows 100 cm apart and
trickle irrigated. Each treatment was replicated four times in four individual plots of 4 m x 2 m wide
(48 plots). Each plot contained 8 treated plants. The irrigation water had the following properties:
pH, 8.05; E.C. 2.03 dS rn';.Cr 483.90 mg litre1; Na+ 305.76 mg litre 1 ; K+ 10.16 mg litre 1 ; HCO
278.15 mg litre 1 .
The different treatments consisted of applying increasing rates of both N and K in the
following manner: N in the form of NH4NO3 (N,: 6 g m 2 , N 2 :12 g m"2, N 3 : 18 g m 2 and N 4 : 24 g m'
2
) and K in the form of K2SO4 (K,: 4 g m"2, K2: 8 g m"2 and K3: 12 g m 2 ). Phosphorus was applied as
H3PO4 (6 g m 2 ), calcium (11 g m 2 ) and magnesium (3 g m 2 ) were supplied as sulfates. The stated
rate of each nutrient is the total applied. It was added gradually with water irrigation over the entire
growth period of the plants. Fertilization-irrigation was complemented with the following
micronutrients: Fe: 0.5 mg litre'1; B: 0.1 mg litre"1; Mn: 0.1 mg litre"1; Zn: 0.075 mg litre"1; Cu:
0.075 mg litre"1 and Mo: 0.05 mg litre"1. The pH values of the solution oscillated between 5 to 6; Fe
was applied as FeEDDHA, B as H3BO3 and the remaining micronutrients as sulfates.

LEAF CONTENT OF DIVALENT CATIONS

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Plant Sampling
Leaf samples were standardized by using only plants with fully expanded leaves of the
same size. Leaves (without petiole) were picked from a point located one third below the plant
apex. The plant samplings were carried out every 15 days throughout the growth cycle of the crop
(from 85 to 220 days old), totalling 10 samplings. Leaves were sampled from the 4 replicates of
each treatment. All the samplings were carried out at the experimental greenhouse between 10.00 h
AM and 12.00 h PM and transported under cold conditions. The time taken between sampling and
sample preparation at the laboratory was < 2 h. Leaves were washed with 1% alkaline detergent and
three times with de-ionized water (Jones et al., 1991). The 50% of the sampled material was placed
in a forced-draught oven at 70C for 24 h, ground in a wiley mill and then placed in plastic bags for
the further analysed. Other subsample (the other 50% fresh leaf matter), was stored in a cold
chamber at 4C.
Plant Analysis
The soluble forms of Ca and Mg were extracted, using 0.5 g of ground, dry plant matter
with 20 mL of distilled water for 30' (Villora et al., 1997). A certain quantity of plant matter was
weighed and mineralized with concentrated H2SO4 and 30% H2O2, free of P, (Wolf, 1982) in
order to extract the total forms of Ca and Mg.
The Ca and Mg fractions were measured using the procedures of Carpena et al., (1977)
and Lpez-Cantarero and Romero (1993), as exemplified in Figure 1.
Statistical Analysis
All data were subjected to statistical analysis by Duncan's Multiple Range Test (DMRT),
the means of of separation within columns was represented with lowercase letters in the tables
(P< 0.05).
RESULTS
The level of nutritional requirements of a plant may be expressed by both the total form
of the nutrient or by its fraction which is soluble in an acid medium, as in the cases of Ca and Mg
(Table 1). Soluble Mg showed its lowest leaf level in the treatment N4K3, whereas the highest
value was obtained with N,K 3 . The behavior of total Mg was similar, with maximum leaf levels
being obtained in the treatment N,K 2 , whereas the lowest levels occurred in the treatment N 2 K,.
For soluble Ca (Table 1), N4K, showed the maximum concentration, whereas with N3K3 the
minimum concentration was obtained. For total Ca (Table 1), the lowest value occurred with
N,K 2 and the maximum at N4K3.
Table 2 shows the relationship between the external levels of NK and Ca fractions. The
fraction of Phosphate-Ca seems depends of external N levels as the treatments with low N levels
(N,K, to N2K3), the percentage of this fraction is low, and increases as soil N concentrations rise
(N3K, to N4K3). With the N, (combinations (N,K, to N,K 3 ), leaf levels of this fraction increases,
whereas with N4 combinations (N4K, to N4K3), foliar levels of this fraction decreased. This
suggests that there could be a synergic or antagonistic relationship between this fraction and
external K, totally depends on the N concentrations in the growth medium. Thus, external K
inhibits the presence of this fraction on the leaf.
With respect to Residual-Ca (Table 2), the highest level was obtained in the treatment
N4K3, and the lowest in the treatment N,K3. The Ca fraction which usually forms an important
part of organic tissue is Pectates-Ca, where N,K, showed the lowest proportion of this fraction
and N2K, the greatest. The most useful or physiologically active Ca fraction is Inorganic-Ca,

VLLORAETAL.

2312

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1 g dried ground material

+
25 mL distilled water

Supernatant

Inorganic-Ca

Residue

Chlorophyll-Mg

Supernatant

Residue

I
Pectate-Ca

Organic anion-Ca Supernatant

Residue

Phosphate-Ca

Organic acid anion-Mg

Supernatant
I

Oxalate-Ca
O

Pectate-Mg
O

Residue

0
Residual-Ca

Residual-Mg

O Shaking 180 min; centrifugation (3,000 g 4 CC during 5 min).

Air dried 24 h at 70 C. Dried residue + 15 mL NaNO3 2N.
Shaking 120 min; centrifugation.
Dried residue + 15 mL AcOH 10%. Shaking 240 min; centrifugation.
O Dried residue + CIH 2N. Shaking 240 min; centrifugation.
remaining material dried at 70 C
O Aliquots of the corresponding extracts, after drying at 70 C, undergoing
sulphuric digestion in order to measure the Ca and Mg by atomic
absorption spectrophotometry.
Figure 1.- Extraction procedure of the Ca and Mg fractions in plant material

LEAF CONTENT OF DIVALENT CATIONS

2313

Table 1.- The effect of NK treatments on Total and Soluble Ca and Total and Soluble
Mg foliar levels (mg/g d.w.).

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Treatments

Total Ca

Soluble Ca

Total Mg

Soluble Mg

NiKi

33.31 f

17.38 ab

11.11 bc

3.02 bc

N1K2

33.26 f

17.93 ab

11.84 a

3.33 a

N1K3

36.34 ef

17.73 ab

11.37 abc

3.39 a

N2K1

38.54 cd

17.51 ab

10.46 d

3.32 a

N2K2

37.90 cd

17.86 ab

11.20 abc

3.01 bc

N2K3

40.87 abc

17.98 ab

11.19bc

3.03 b

N3K,

38.79 bed

17.87 a

10.83 cd

2.91 bed

N3K2

39.96 abc

17.70 ab

11.00 bed

3.02 bc

N3K3

40.90 ab

17.10b

10.98 bed

2.82 bed

N4K1

42.33 a

18.70 ab

11.41 abc

2.70 d

N4K2

40.15 abc

18.59 ab

11.24 abc

2.81 cd

N4K3

42.49 a

18.57 ab

11.49 ab

2.75d

DMRT atp < 0.05 are indicated by lowercase letters in columns

whose value was highest in N,K, and lowest in N4K2. The main fraction obtained was the
Oxalate-Ca, and showed opposite behavior to inorganic Ca, since N2K, gave the lowest percent
value and N4K3 the highest. The values for N,K, to N2K3 were generally lower than those for
N 3 K,toN 4 K 3 .
The capacity for integration of Mg is conditioned by the NK levels applied to the soil
(Table 3). Thus, Pectate-Mg is the fraction with the least incidence overall, although with
differences between treatments, such that the highest value was obtained from N 2 K, and the
lowest from N4K3. For Organic acid anions-Mg fraction, the highest value was obtained at N 2 K,
and the lowest in N4K3. In our experiment, the concentration of this fraction is higher with N
while an increase of NK applied (N2 to N4 combinations) decreased significantly the foliar levels

2314

VLLORAETAL.

Table 2.- Concentrations of Ca fractions (mg/g d.w.) with respect to the NK

treatments.

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Treatments

Oxalate

Inorganic

Pectate

Residual

Phosphate

NiKi

14.24

cd

8.14

3.20

3.19

bed

1.61

NiR>

14.47

bed

7.71

abc

3.31

ab

3.18

bed

1.75

ede

N,K3

14.28

cd

7.63

abc

3.24

ab

3.09

1.76

ede

N2K1

13.99

7.43

be

3.53

3.07

cd

1.70

de

NJC2

14.26

cd

7.55

be

3.29

ab

3.20

a-d

1.75

ede

N2K3

14.95

bed

7.80

ab

3.33

ab

3.30

ab

1.77

ede

N3K,

15.30

abc

7.78

abc

3.35

ab

3.22

abc

1.93

bed

N3K2

15.14

a-d

7.69

abc

3.50

ab

3.28

ab

2.22

N3K3

15.30

abc

7.33

be

3.46

ab

3.31

ab

2.03

ab

N4K1

15.04

a-d

7.33

be

3.24

ab

3.29

ab

2.12

ab

N4K2

15.56

ab

7.20

3.32

ab

3.19

bed

1.91

bed

N4K3

16.15

7.62

abc

3.27

ab

3.34

1.93

be

DMRT at p < 0.05 are indicated by lowercase letters in columns

of Organic acid anions-Mg. Thus, given N dosage, K increasing levels (K, to K3) induced a
decrease of this fraction.
The Residual-Mg fraction is one of the forms of Mg with low incidence with respect to
total Mg, and N2K2 and N4K3 are the maximum and minimum values, respectively. When N was
low (N,K, to N,K 3 ) or relatively high (N3K, to N3K3), increases in the levels of K led to a
reduction in the leaf levels of this fraction, while when N was intermediate (N2K, to N2K3) or
very high (N4K, to N4K3), the intermediate K level (N2K2 and N4K2) had a very positive effect on
the fraction.

LEAF CONTENT OF DIVALENT CATIONS

2315

Table 3.- Concentrations of Mg fractions (mg/100 g d.w.) with respect to the NK

treatments.

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Treatments

Cloroph.

Org.An.

Org.Ac.An.

Residual

Pectate

NiKi

536

de

233

be

62 a

41

bed

53

ab

N1K2

572

bed

286

61

41

bed

25

N1K3 .

580

abc

265

abc

58

ab

39

b-e

44

bc

N2K1

616

268

ab

44

de

34

ef

61

N2K2

536

de

258

abc

51

a-d

49

34

cd

N2K3

543

cde

262

abc

46

b-e

40

b-e

30

cd

N3K,

575

a-d

278

57

abc

45 ab

34

cd

N3K2

585

abc

257

abc

49

bed

43 abc

52

ab

N3K3

582

abc

257

abc

45

cde

36

c-f

27

N4K1

611

ab

240

bc

53

a-d

35

def

24

N4K2

527 e

232

47

b-e

45

ab

24

N4K3

563

242

bc

35

31

22

cde

Clorophyll: chlorophyll Mg; Org.An.: Mg linked to organic anions; Org.Ac.An.: Mg

linked to organic acid anions; Residual: residual Mg; Pectate: Mg linked to pectate
DMRT atp < 0.05 are indicated by lowercase letters in columns

The fraction Organic anions-Mg reached the maximum at N,K 2 and showed a minimum
in N4K2. In general, this fraction was negatively affected by K levels applied, since at same
dosage, increased K dosage reduced significantly Organic anions-Mg levels.
The most common Mg fraction in plants is the Chlorophyll-Mg. We observed that the
lowest value was obtained at N4K2 and the highest at N2K,. An initial approximation indicates
that when there is a large amount of another ion in the soil, K does not have any clear effect on

2316

VILLORAETAL.

this Mg fraction, since the maximum and minimum values are obtained with different K levels
and with low and high N levels.

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DISCUSSION
The response to the N dosages N, and N2 showed a higher mean concentration of total and
soluble Mg, while N3 and N4 gave mean levels which were lower in both Mg forms, while the
opposite was true for total and soluble Ca, with N3 and N4 showing higher levels than N, and N2.
This results are surprising if we bear in mind that each N dosage contained three increasing
levels of K, and total Ca increased significantly as NK increased in the growth medium. This fact
could be explained by i) the presence of K and NH/ in the soil and aerial part of the plant, ii) leaf
level of Ca affected the leaf Mg or, iii) a combination of both. The improve of uptake and
transport of Mg and Ca could be explained by a reduction of antagonistic effect of NH4+ in N
dosage, through the increase of uptake of K from the soil. Similarly, although K has an
antagonistic effect upon Mg and Ca in the uptake process (Valenzuela and Romero, 1991;
Marschner, 1995), it enhances the process of the transport of both cations (Addiscott, 1974),
which would explain the leaf levels obtained, but not prevent the antagonism between the two
divalent cations, since the foliar increase of one leads to a decrease in the other, and vice versa.
The reason for this inverse behavior might be directly related to the N levels and the
accumulation of NH4* in the plant tissue (Mills and Jones, 1996) which leads, at cellular level, to
increased Ca concentration in the cytoplasm to counteract the toxic effect of NH4+ on the plant
tissue (Findenegg et al., 1982) and stabilize the pH of the cell tissue. This accumulation of Ca
reduce both soluble and total Mg, as a competitive mechanism due of affinity with the transporter
(Schymansky, 1981). The leaf level of total and soluble Ca in our experiment is similar to that
obtained for melon (Valenzuela et al., 1993), and tomato and cucumber (Valenzuela et al., 1992),
but much lower than that obtained for eggplant (Lpez-Cantarero and Romero, 1993) and greater
than that obtained for capsicum plants grown under fertilization conditions at lower levels than
those employed in this experiment (Ruiz, 1983). Such disparity between results may be due to
the growing conditions and species used for each of the studies. Total and soluble Mg showed a
high leaf concentration in comparison with the study by Ruiz (1983), but our levels were similar
to those obtained for melon (Valenzuela et al., 1993) and lower than those found for eggplant
(Lpez-Cantarero and Romero, 1993).
The influence of the joint action of NK on total Ca led us to wonder whether it the same
occurred with the various fractions of the cation integrated into the plant structure. The opposite
behavior of the Phosphate- and Residual-Ca fractions suggests that K dosage have a positive or
negative effect on fraction, respectively. Foliar levels of the Phosphate-Ca fraction were higher
compared to those obtained by Valenzuela et al. (1992) for tomato and cucumber plants, while
our results are much higher than those for melon (Valenzuela et al., 1993) and similar to those
reported for eggplant (Lpez-Cantarero and Romero, 1993). These differences could be due to
the growing conditions and characteristics of the species under study, e.g. the eggplant was
grown with increasing P levels, and although in our experiment P levels were constant. Our
values for Residual-Ca are high compared with those obtained for eggplant (Lpez-Cantarero
and Romero, 1993). The positive result is to be expected, since this fraction usually represents
the part of Ca which is linked to proteins and highly specialized organic structures with a strong
affinity for the cation, making it difficult to extract (Lpez-Cantarero and Romero, 1992).
Comparing the Pectate-Ca results with those for other species, they are higher than those
for melon (Valenzuela et al., 1993) but considerably lower than those for eggplant (LpezCantarero and Romero, 1993), tomato and cucumber (Valenzuela et al., 1992) and even
watermelon (Vargas etal., 1991).
With respect to the fractions, Inorganic-Ca may be involved in physiological processes
which depend on the Ca and is considered as "physiologically active Ca" (Valenzuela et al.,

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LEAF CONTENT OF DIVALENT CATIONS

2317

1991). Our Inorganic-Ca values are similar to those recorded for tomato (Valenzuela et al.,
1992), melon (Valenzuela et a l , 1993) and watermelon (Vargas et a l , 1991), but higher than
those published for eggplant (Lpez-Cantarero and Romero, 1993) and considerably higher than
for cucumber (Valenzuela et al., 1992).
Oxalate-Ca is involved in specific cellular stabilizing function when the tissue is
subjected to conditions of saline stress (Lpez-Cantarero and Romero, 1993), and the Oxalate-Ca
results suggests that the plant compensates for large amounts of NK in the soil with a high
content of this fraction. Inorganic-Ca decrease with NK increase (Mills and Jones, 1996). A
possible explanation for this negative interaction may be that an excess of Inorganic-Ca leads to
changes in the cytoplasm pH (Raven, 1977). The Pectate-Ca, showed no clear relationship with
N dosage and this could be explained since P, especially Inorganic-P, is usually involved in
numerous metabolic processes (Silva et al., 1992) and acts as an intermediary when energy is
required. There is also a significant although slight relationship between N and Oxalate-Ca. This
can be explained as one of its primary functions is as osmoregulator involved in the osmotic
stabilization processes of plant cells (Martinez and Cerda, 1989).
The supply of NK directly or indirectly affects the integration of the ionic forms in the
plant tissue, and in the present study, Ca was integrated in proportions which are indicative of
such growing conditions, with the following order:
Oxalate-Ca > Inorganic-Ca > Pectate-Ca Residual-Ca > Phosphate-Ca
with mean percentages of 49%, 24%, 11%, 10% and 6%, respectively. These proportions differ
from those of Lpez-Cantarero and Romero (1993), whoes obtained similar values for OxalateCa but higher values for Phosphate- Ca, their experiment was considerably affected by P dosage.
Our results largely agree with those of Valenzuela et al. (1992), although the proportions of
Inorganic- and Residual-Ca are higher in our results. The high proportion (nearly 50%) of
Oxalate-Ca in this experience could be due to the saline grown conditions.
Like Ca, Mg is involved as a divalent cation in numerous vital processes for the plant and
therefore plays a role in many different structures of the plant tissues. Thus, maximum
concentrations of all Mg fractions were obtained when N and K levels were low, and the lowest
leaf values when NK dosage were high or very high. This response suggests that high levels of K
and NH4+ (the N was applied as NH4NO3) lead to decreases in the leaf levels of the Mg fractions
and the combined action of K and NH4* applied to the soil may reduce the uptake and transport
of Mg to the plant tissue. The antagonistic effect is more acute for K than NH 4 \ since at same N
dosage, increasing amounts of K affect the foliar levels of Mg and the effect of the NH4+ cation is
minimized by the K. This K/Mg antagonism (Valenzuela and Romero, 1991) therefore affects the
ability of the divalent cation to participate in the metabolic system. However, this does not mean
that leaf levels of the Mg fractions cannot be high, since although the antagonistic effect does not
facilitate the transport of the divalent cation it is not totally impeded (Addiscott, 1974), so
dramatic nutritional changes do not occur and the rhythm of the plant is not greatly disturbed. In
general, the levels of the different Mg fractions were similar to those obtained for other species
(Shockey and Reid, 1984; Lpez-Cantarero and Romero, 1993), with any differences, probably
due to the growing conditions and particular nutritional requirements of each species.
We could conclude that soil NK levels affected the response of the Mg fractions in
leaves. We thus find that K has a negative relationship with the Chlorophyll-Mg fraction, since in
the excess of cation, may alter the cytoplasmic pH and thus disturb the photosynthetic apparatus.
K is involved in the system of electronegative ionic balance of the cells (Valenzuela et al., 1993),
and this fact could explain the positive relationship with Organic anions-Mg. There was a
different physiological significance between the various Mg fractions which may vary according
to the growing conditions of the crop, but the dominant fractions (Organic anions-Mg or
Chlorophyll-Mg) are the same in all species (Lpez-Cantarero and Romero, 1993). In the present
study, the order of importance was:

2318

VUXORAETAL.

Pectate-Mg < Residual-Mg < Organic acid anion-Mg < Organic anion-Mg < Chlorophyll-Mg
with percentages of 3%, 4%, 5%, 27% and 55%, respectively. These proportions are different
from those obtained for other species such as eggplant (Lpez-Cantarero and Romero, 1993),
where Organic anion-Mg and Pectate-Mg were higher, while the Residual-Mg was low.

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CONCLUSIONS
As a result of the above, we could conclude that the total and soluble forms of Ca and Mg
were affected stronger by N than K dosage, thus N, and N2 gave the higher mean concentration of
total and soluble Mg but lower total and soluble Ca, while N3 and N4 gave lower levels for both
Mg forms, while the opposite was true for total and soluble Ca, independently of the K dosage
applied. However, the distribution of the Ca and Mg fractions, was notably affected by the K
treatments within each N dosage. Moreover, the percentage distribution of the Ca and Mg
fractions with respect to the total content of the elements, differed than those obtained and
established for other crops grown under similar controlled conditions. These alterations could
have its origin in the high application of K fertilizer, thus, Pectate-Mg and Oxalate-Ca showed
higher percentage than those obtained in crops under no stressing conditions and these fractions
will be employed as a useful bioassay for the knowledge of the response of plants to the stressing
growth conditions.
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