Cation Concentrations of Plant Tissues of Fruit-Vegetable Crops as

Affected by the EC of the External Nutrient Solution and by Humidity

C. Sonneveld G.W.H. Welles
Tolhuislaan 35, 3862 WK Nijkerk Applied Plant Research, Division
The Netherlands Glasshouse Horticulture, P. O. Box 8,
2670 AA Naaldwijk
The Netherlands†1

Abstract
In a series of experiments effects of EC values in the root environment and
differences in climatic conditions were studied in relation to the uptake of cations.
Tomato, cucumber and sweet pepper were used as test crops. Differences with EC
values were realised by the addition of various concentrations of nutrients, whereby
the mutual ratios between the nutrients were kept constant. Differences of climatic
conditions were realised by ventilation, heating, screening regimes and evaporation
of water from open water basins. For all crops a spring and a fall grown crop were
incorporated in the study. The uptake of the cations was determined by tissue
samples. Young fully grown leaves and ripe fruits were sampled and analysed
systematically. Accidentally, old leaves and tops were sampled and analysed, while
in some cases the fruits were separated in a proximal part and a distal part. The K
concentration in the plant tissues was increased and the Ca concentration was
decreased by an increasing EC in the external solution. The Mg concentration in the
plant tissues was affected less evidently by an increased EC. The overall effect of the
humid climate was a decrease of the K and Mg concentrations in the leaves and a
small increase of the Ca concentration. The K, Ca as well the Mg concentrations in
the fruits were increased by the high humidity in the greenhouse. However, these
effects were not of the same size as occurred with the EC and were not consistent for
all crops. The average Ca concentration in the distal end of the tomato fruits was
40% lower than in the proximal part; for sweet pepper this difference was only
10%. Increased nutrient concentrations in equal ratios, to enhance the EC in the
external solution, had only slight effects on the uptake of cations in comparison with
the effects of nutrient concentrations realised by different ratios in the external
solution at an equal EC value. In this way it was shown that great differences can
occur in the cation uptake by plants, while the cation concentrations in the external
solution were equal. Therefore, with the interpretation of analytical data of cation
concentrations in the external solution in substrate systems, beside the concentration
at such, especially the relation to the other nutrient cations should be seriously taken
into account. Guidelines for interpretation are discussed.

INTRODUCTION
In hydroponic and hydroponic related systems the composition of the nutrient
solution in the root environment is regularly analysed to check the availability of plant
nutrients. The small root volumes in charge and the high uptake of nutrients require
specific knowledge with interpretation of the analytical data. The traditional models used
for field grown crops are an insufficient basis for an interpretation. These models are
developed for big root volumes and for the sub-optimal to optimal domain of nutrient
supply, while in hydroponic systems small root volumes and mostly the optimal to
luxurious domain of nutrient supply is in view. Because of the differences between the
volumes, with equal analytical data huge differences occur in the quantities of nutrients
available for field and substrate grown crops (Sonneveld, 1981) and because of the
different domains of nutrient supply in view linear and curvilinear uptake models should
1
† Gerard Welles died on October 27th 2004 when this paper was ready to publish. I have lost with him a
valuable colleague and a good friend.

Proc. IS on Soilless Cult. and Hydroponics

Ed: M. Urrestarazu Gavilán
Acta Hort. 697 ISHS 2005 377
be taken into account in relation to luxurious and sub-optimal nutrient supply (Sonneveld,
1991) respectively.
With equal EC levels mostly a close relationship has been found between
concentrations in the root environment and those in plant tissues (Adams, 2002; Maruo et
al., 2004; Sonneveld and Voogt, 1985; Sonneveld and Voogt, 1986). However, this is not
the case when the EC values differ, surely not when EC values differ by addition of ions
with equal mutual ratios (Sonneveld and Voogt, 1990). Another factor that affects the
nutrient uptake is the humidity. Differences in humidity will affect the uptake of nutrients
at equal concentration in the root environment.
At the Applied Plant Research, Division Glasshouse Horticulture at Naaldwijk a
series of experiments was carried out in which effects of different EC values in the root
environment and different climatic condition on the uptake of nutrients were studied. The
results offer possibility to discuss the effects of these factors on the interpretation of
analytical data in relation to the uptake of nutrients. In this paper data of cation uptake
will be presented.

MATERIALS AND METHODS

Tomato, cucumber and sweet pepper were grown in a greenhouse in rock wool
strips placed in gutters, furnished with a drip irrigation system with which the nutrient
solution was circulated. The greenhouse consisted of different compartments covered by
glass, equipped with a movable polythene thermal screen and a heating system.
Nutrient solutions were adjusted to the crop in agreement with the
recommendations to growers as published by Sonneveld and Straver, (1994). The primary
water was rain water or desalinated water. The different EC values in the root
environment varied roughly between 1.5 and 7 dS m-1 and were realised by increasing all
nutrients with equal ratios.
The differences in climatic conditions were focussed on humidity, obtained by
differences for ventilation, heating, screening and evaporation from open water basins.
The temperature between the humidity treatments was kept constant in principle. The
equipment to realise the climatic differences was used such that the crop development
never seriously was hindered. In the present paper data of the driest and the most humid
climate will be presented. The driest climate represented more or less the climatic
conditions as usual in The Netherlands and therefore will be mentioned “standard”. The
differences in humidity varied strongly by the daily weather conditions and the growing
season of the crop. The average differences of vapour pressure deficits (vpd) realised
under standard and humid conditions were in spring 0.73 and 0.53 kPa and in autumn
0.72 and 0.54 kPa for the standard and the humid climatic conditions respectively.
Two crops per year were grown, one from December until June and the second
from June until November, called spring crop and fall crop respectively. The circulating
solution was sampled and analysed frequently to control EC, pH and nutrient
concentrations. Tissue samples of young fully grown leaves and ripe fruits were gathered
of all crops at proper times. With the fall crop of sweet pepper no fruit samples were
available. For tomato and sweet pepper sometimes fruits were split in a proximal and a
distal part, because of the great differences of nutrient concentrations, especially Ca,
expected between such parts (Adams, 1990; Ehret and Ho, 1986). With cucumber also
plant tops and old leaves were sampled, to compare the composition of those plant parts
with those of young leaves. In the present publication the concentrations of cations in the
plant tissues will be discussed in relation to the EC and the cation concentrations in the
root environment and the different climatic conditions. Cation concentrations in the root
environment are expressed as mmol l-1 and the EC as dS m-1; the cation concentrations in
the plant tissues are expressed as mmol kg-1 dry matter.

RESULTS
The results of the tissue tests are listed in the Tables 1-6 for the spring and fall
crops of tomato, cucumber and sweet pepper respectively.

378
 The sums of cations in the plant tissues (C+ = Na + K + 2Ca + 2Mg) did not
show systematic differences between different EC values and the difference in humidity.
C+ was higher for the young leaves than for the fruits; the average values were 2998 and
1372 for tomato, 2758 and 2224 for cucumber and 3114 and 955 for sweet pepper,
respectively.
The Na concentrations decreased with increasing EC on average with 3 and 5
mmol per kg dry matter per unit EC for young leaves and fruits respectively. The K
concentrations in the plant tissues increased with increasing EC on average over the crops
with 48 and 44 mmol per kg dry matter for every unit EC for leaves and fruits
respectively, while the Ca concentrations decreased with 17 and 4 mmol per kg dry matter
per unit EC, respectively. Mg did not show a consistent reaction on the EC; with
increasing EC on average the concentration in the young leaves and fruit decreased with 5
and 1 mmol per kg dry matter per unit EC, respectively.
The climatic conditions did not show a consistent effect on the concentration of
cations in the plant tissues. On average, a humid climate decreased the cation
concentrations in the young leaves, while these on average were increased in the fruits.
However, the Ca concentration in the young leaves was an exception, because this
concentration was on average a little increased by humid conditions. On average, the Na,
K, and Mg concentrations in the young leaves were lowered by high humidity with 6, 26
and 7 mmol per kg dry matter, respectively, while the Ca concentration was increased
with 2 mmol. The K, Ca and Mg concentration in the fruits under humid conditions were
higher with on average 59, 11 and 7 mmol per kg dry matter, respectively. The average
Na concentrations in the fruits were equal for both treatments.
The proximal and distal parts of fruits showed relatively greatest differences for
the Ca concentrations. These were relatively greatest for the tomato crop, where the
concentration in the distal part was about 60% of that in the proximal part. For the sweet
pepper the differences occurred with the increasing EC value. At a low EC value no
difference was found between the Ca concentrations of both fruit parts, while at the high
EC values the concentration in the distal part was about 80% of that in the proximal part.
The concentrations of cations in the old leaves and tops of cucumber followed
the same pattern as found in the young leaves. The Na and the Ca concentrations
increased by the age of the tissue, K was highest in the tops and Mg in the old leaves. The
differences were greatest for Ca, where the concentration in the old leaves was more than
five times higher than of those in the tops.

DISCUSSION
The present data shows that EC values in the root environment increased with
equal ratios of nutrients only has relatively small effects on the uptake of cations for the
crops under investigation. The most striking effects were the increased K and the reduced
Ca uptake. Such effects have been found earlier (Charbonneau et al., 1987; Sonneveld
and Voogt, 1990). Increased K concentrations as found will not affect strongly growth
and production of crops. However, the decrease of Ca in plant tissues easily aggravate
deficiency symptoms of this element, because the Ca concentration in some plant parts,
like fruits and shoot tops, is as usual low and any reduction will promote possible
disorders. It has been found that high EC values promote Ca deficiency symptoms in
different crops (Adams, 1993; Van Berkel, 1987). The decrease of 4 mmol Ca per kg dry
matter for each unit increase of EC in the root environment, as found on average for fruits
in the present experiments, easily brings the Ca concentration below the critical value,
when the EC in the root environment is increased with some units. This will be
emphasized by the unequal distribution of Ca within plant organs. Youngest growing
parts can be very low in Ca concentrations (Adams and Ho, 1989; Adams and Ho 1992).
More about this subject can be found in the data of sweet pepper fruits in Table 5, where
the Ca concentration in the distal end of the fruits decreases with almost 40% by an
increase of the EC in the root environment of about 4 units.
For fruits the effects of a reduced Ca uptake by increasing EC values were

379
aggravated by a lower humidity. Thus, high EC and low humidity are synergistic and
therefore an unfavourable combination with respect to Ca uptake and its distribution to
fruits.
Described effects offer serious problems for the interpretation of analytical data
of substrate and nutrient solution samples from the root environment to control the
nutrient supply in such systems. When the EC is increased with nutrients, the different
cation concentrations in the root environment are proportionally increased, while it only
slightly increases the potassium uptake, nearly not affect the Mg uptake and even
decreases the Ca uptake. Thus, the cation concentrations in the root environment cannot
tolerate an interpretation on basis of their absolute concentrations, but should be judged in
relation to the EC. However, under saline conditions the EC also can be strongly affected
by the availability of NaCl in the root environment and thus, the Na concentration will be
considered in the interpretation of the nutrient cations too.
The effect of NaCl salinity in relation to nutrient uptake was studied in earlier
research (Sonneveld and Van der Burg, 1991), from which the data of the nutrient uptake
was not presented, but now summarized in Table 7. These data shows that the uptake of K
is reduced by the NaCl addition, the Ca uptake was reduced by the sweet pepper crop
only and the Mg uptake was not seriously affected by any of the crops presented. Thus the
behaviour of Na is different from the other cations, for the addition of relatively high
concentrations of NaCl of 25 mmol l-1 did not affect the uptake of other cations seriously.
In the data presented only the slight reduction in the uptake of K was found as a
consistent reaction on high NaCl concentrations. This will not induce serious plant
nutrition problems, because the effects were small and in addition effects of a reduced K
uptake can be substituted by Na uptake in some plant species (Besford, 1978).
The differences in the uptake of nutrient cations mainly occur by differences
between their mutual ratios. This is demonstrated by the data shown in Table 8, where the
ratio between the lowest and highest concentrations of a nutrient cation in the root
environment are compared with this ratio of the concentrations in the plant for the data
presented in this paper in comparison with the data presented by a cation ratio experiment
of Sonneveld and Voogt (1985). The differences of cation concentrations in the root
environment of the present paper are originated by EC values and thus, having the same
mutual cation ratios, while those presented by Sonneveld and Voogt (1985) are originated
by different mutual ratios at equal total concentrations (EC). Despite the much wider
range between highest and lowest concentration in the root environment in the present
experiments than in the cation ratio experiments of Sonneveld and Voogt (1985), are the
differences in the uptake much wider in the last case. Thus, the mutual ratios much more
determine the uptake than the absolute concentration.

Interpretation of Analytical Data

Last conclusion is very important with respect to interpretation of analytical data
of samples from the root environment of substrate and hydroponically grown crops. The
guide values for the cations of the nutrient solution in the root environment, used in the
Dutch horticulture industry (Sonneveld and Straver, 1994) are shown in Table 9. These
data are based on relative high EC values as maintained under Dutch growing conditions
and on the use of low salinity water, like rainwater. The sum of nutrient cations of these
guide values in relation to the total EC offers only a small space for Na, 0-3 mmol l-1.
These data need adjustment for different factors discussed in the present paper.
Firstly, the data need adjustment for the EC value. Not the concentration at such,
but the relative value with respect to the other nutrient cations should be considered, using
following formula.

Vc(X )
RV(X) = (1)
Cn +

380
In which R = relative concentration
X = nutrient cation X, respectively K, Ca and Mg
c = cation concentration mmol l-1
V = valency of the cation
Cn+ = sum of protons of the nutrient cations (K + 2Ca + 2Mg)
Secondly, the data should be adjusted for high NaCl concentrations. High EC
values are recommended to control the lush growth under poor light conditions and with a
view to an optimal quality of the produce. However, when the EC is already high by the
use of saline primary water, there is no reason to increase the nutrient levels by addition
of nutrients higher than necessary for optimal plant nutrition. Under current growing
conditions with a flow rate of the nutrient solution in the system between 1.3 and 2.0 it
can be argued that the concentration of a well balanced nutrient solution in the root
environment should not be lower than 1.5 dS m-1 (Sonneveld, 2000). Calculation of the
guide values of Table 9 to the relative contribution to Cn+ and minimum values at EC 1.5,
using formula (1), are listed in Table 10. The total sum of cations at EC = 1.5 was
calculated with the aid of formula (2) (Sonneveld et al., 1999).
EC dS m-1 = 0.1 C+ (2)
In which
C+ = sum of protons
Thirdly, there should be an adjustment for the climatic conditions. Dry climatic
conditions will reduce the Ca transport to fruits, which will stimulate deficiency
symptoms with crops sensitive to it.
The K : Ca (mol/mol) of the guide values for the different crops varied from 0.75
till 1.23 and showed a close relationship with the sensitivity of the crop to Ca deficiency
in the fruits. The lowest value is recommended for sweet pepper and the highest for
cucumber, very sensitive and not sensitive, respectively. These values are focussed on the
relative high EC values recommended in The Netherlands. It can be argued that with
lower (total) EC values a little higher K : Ca ratio will be maintained, because of the
better Ca uptake with lower EC values. Under dry climatic conditions a some what lower
K : Ca is recommended, because of the lower Ca transport to the fruits. One thing and
another should be considered in relation to the sensitivity of the crop and the cultivar
grown to the disorder expected. A too low Ca concentration in the fruit induces blossom
en rot and a too high concentration gold speck (Voogt, 2003). The already mentioned
synergistic effect between EC and humidity should be emphasized again. High EC in the
root environment combined with low humidity can be responsible for dramatic Ca
deficiencies in fruits and asks for adjustment of the K : Ca ratio.
The relative contribution of Mg to the Cn+ of the nutrients shows a close relation
with the sensitivity of the crops to Mg deficiency. It varies from 0.214 for sweet pepper to
0.312 for eggplant, not sensitive and very sensitive to Mg deficiency, respectively.
An example for an interpretation is given in Figure 1. The results of the
calculations suggest a small decrease of both, the supply of K and Mg and an increase of
the Ca supply in relation to the current EC. The ultimate decision will be made in relation
to the factors discussed in the paper, which means in relation to the EC and humidity
expected for the approached period.
The interpretation model given is suitable for optimal and luxurious supply of
nutrients, which means a balanced nutrient solution in the root environment >1.5 dS m-1.
With a lower nutrient status the uptake of some nutrients will be in the sub-optimal range
and the relation between supply and uptake can differ strongly from the relationships on
which the interpretation presented is based.

381
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Tables
Table 1. Cation concentrations of young fully grown leaves and fruits of spring grown
tomato as affected by EC value and humidity. Data expressed as mmol kg-1 dry
matter.

EC Leaves Fruits
Na K Ca Mg Na K Ca Mg
Proximal Part
2.4 66 1026 661 212 44 1150 38 64
3.2 57 1154 662 198 38 1205 36 63
4.5 64 1238 642 194 43 1272 34 67
Distal part
2.4 38 1133 22 67
3.2 33 1211 21 68
4.5 34 1210 22 70
Climate
Proximal Part
Standard 62 1118 693 208 46 1136 34 59
Humid 62 1160 616 194 38 1282 38 70
Distal part
Standard 37 1121 19 63
Humid 33 1249 24 73

Table 2. Cation concentrations of young fully grown leaves and fruits of fall grown
tomato as affected by EC value and humidity. Data expressed as mmol kg-1 dry
matter.
EC Leaves Fruits
Na K Ca Mg Na K Ca Mg
2.6 172 961 847 145 75 1049 34 63
5.0 119 1032 842 123 49 1107 27 65
Climate
Standard 161 1012 859 131 63 1043 33 60
Humid 137 927 834 128 59 1063 32 69

383
Table 3. Cation concentrations of young fully grown leaves and fruits of spring grown
cucumber as affected by EC value and humidity. Data expressed as mmol kg-1 dry
matter.

EC Leaves Fruits
Na K Ca Mg Na K Ca Mg
1.6 32 801 768 236 96 1338 143 132
3.0 19 882 746 215 70 1650 124 121
4.4 18 1007 635 200 65 1696 125 115
Climate
Standard 28 937 695 233 73 1504 128 122
Humid 18 865 741 200 80 1618 133 123

Table 4. Cation concentrations of leaves, fruits and tops of fall grown cucumber as
affected by EC value and humidity. Data expressed as mmol kg-1 dry matter.

EC Na K Ca Mg Na K Ca Mg
Young leaves Fruits
2.0 23 977 555 262 39 1615 181 150
4.0 19 1007 547 307 33 1674 153 163
5.7 17 1048 529 311 28 1621 152 153
Climate
Standard 20 1057 535 304 33 1610 136 147
Humid 20 1028 546 299 34 1677 182 161
EC Old leaves Tops
2.0 27 1146 1115 430 14 1414 191 275
4.0 22 1155 985 487 12 1464 193 312
5.7 22 1197 955 452 11 1461 177 318
Climate
Standard 26 1181 1097 486 13 1438 201 316
Humid 22 1162 996 424 12 1474 180 299

Table 5. Cation concentrations of young fully grown leaves and fruits of spring grown
sweet pepper as affected by EC value and humidity. Data expressed as mmol kg-1 dry
matter.
EC Leaves Fruits
Na K Ca Mg Na K Ca Mg
Proximal Part
1.6 4 1359 493 279 5 699 28 72
2.9 3 1424 452 277 5 756 27 67
4.2 3 1455 426 264 3 777 28 70
5.7 3 1496 407 241 4 758 24 68
Distal part
1.6 4 720 30 75
2.9 3 761 24 74
4.2 4 800 23 76
5.7 4 784 19 74
Climate
Proximal Part
Standard 3 1410 445 263 4 776 26 69
Humid 3 1452 444 267 4 719 27 69
Distal part
Standard 3 779 23 74
Humid 3 754 25 75

384
Table 6. Cation concentrations of young fully grown leaves of fall grown sweet pepper as
affected by EC value and humidity. Data expressed as mmol kg-1 dry matter.

EC Leaves
Na K Ca Mg
1.6 5 1406 707 282
3.3 6 1514 651 260
4.6 5 1501 669 259
6.7 4 1564 650 268
Climate
Standard 5 1528 642 262
Humid 5 1465 697 272

Table 7. Cation concentrations in young leaves (mmol kg-1 dry matter) as affected by EC
level (dS m-1) and NaCl concentration (mmol l-1) in the root environment. (After Van
der Burg, 1989, 1989a and 1990).

Treatments Tomato Cucumber Sweet pepper

EC/NaCl K Ca Mg K Ca Mg K Ca Mg K Ca Mg
2.5/5 5.5 5.4 2.3 924 886 158 823 1166 388 1430 712 323
3.7/12.5 6.5 5.5 2.4 906 903 171 796 1166 410 1337 672 317
5.2/25 8.5 6.0 2.6 911 839 169 758 1258 408 1360 647 327
3.7/5 9.7 8.1 3.3 978 829 154 832 1190 408 1428 656 333
5.2/5 15.7 11.0 4.4 1079 823 140 856 1212 382 1434 646 349

Table 8. Ratios of highest and lowest K, Ca and Mg concentrations in the root

environment (mmol l-1) and in the young fully grown leaves (mmol kg-1 dry matter) of
cucumber and sweet pepper crops. Results of an experiment with an equal EC, but
different cation ratios (Sonneveld and Voogt, 1985) and as found in present
experiments with different EC values, but equal cation ratios.

Highest/lowest Cucumber Sweet pepper

concentration K Ca Mg K Ca Mg
Experiment with different cation ratios
Ratios in slabs 2.98 1.90 3.36 2.21 1.50 2.22
Ratios in leaves 1.50 1.06 2.08 1.17 1.10 2.22
Present experiment with different EC values
Ratios in slabs (spring crop) 6.15 3.53 3.42 8.19 4.61 5.00
Ratios in leaves 1.26 0.83 0.85 1.10 0.83 0.86
Ratios in slabs (fall crop) 4.31 3.17 2.94 6.48 3.31 3.47
Ratios in leaves 1.07 0.95 1.19 1.11 0.92 0.95

385
Table 9. Guide values for EC (dS m-1), K, Ca and Mg concentrations (mmol l-1) in the
root environment as given by Sonneveld and Straver, (1994) and the sum of protons
(Cn+) originated by K, Ca, and Mg for fruit vegetable crops.

Parameter Eggplant Cucumber Sweet pepper Tomato

EC 3.0 3.0 3.0 4.0
K 6.25 8.0 6.0 8.0
Ca 6.25 6.5 8.0 10.0
Mg 4.25 3.0 3.0 4.5
Cn+ 27.25 27.0 28.0 37.0

Table 10. Relative contribution to Cn+ of the guide values from Table 9 and minimum
cation concentrations (mmol l-1) at EC 1.5 as calculated by formula (1).

Crop Relative contribution to Cn+ Minimum values mmol l-1

EC K Ca Mg EC K Ca Mg
Eggplant 3.0 0.229 0.459 0.312 1.5 3.4 3.4 2.4
Cucumber 3.0 0.296 0.481 0.222 1.5 4.4 3.6 1.6
Sweet pepper 3.0 0.214 0.571 0.214 1.5 3.2 4.3 1.6
Tomato 4.0 0.216 0.541 0.243 1.5 3.2 4.0 1.8

Figures
Analytical data of a sample of a nutrient solution from rock wool slabs with tomato
growing.

EC Na K Ca Mg
4.5 10 8 8 5
Comparison with standard values
Calculations formula (1)
from Table 10
K = 0.235 K = 0.216
Ca = 0.471 Ca = 0.541
Mg = 0.294 Mg = 0.243

Conclusion: Standards recommended at

K and Mg relatively high existing EC and Na concentration
and Ca low K = 7.3
Ca = 9.2
Mg = 4.1

Adjustments in the nutrient

solution added, see text

Fig. 1. Example of a scheme for interpretation.

386