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AgroThesis (2008); Vol.6, No.

1: 20-28
[Effect of the substrate in tomato soilless culture]
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#200803
Original Research Paper
Crop Production > Soiless cultivation
Oct 2007
Apr 2008

Deployment of shredded maize stems as an alternative substrate
medium in tomato soilless culture.
Effect on yield and fruit quality per truss.
[Effect of the substrate in tomato soilless culture]
Nikos G. Tzortzakis, Costas D. Economakis
Department of Hydroponics and Aromatic plants, Institute of Olive Tree and Subtropical Plants, National
Agricultural Research Foundation (N.AG.RE.F.), Agrokipion, 73100, Chania, Greece

Nikos G. Tzortzakis
Department of Hydroponics and Aromatic plants, Institute of Olive Tree and Subtropical Plants, National
Agricultural Research Foundation (N.AG.RE.F.), Agrokipion, 73100, Chania, Greece.
Phone: 0030 28210 83435
E-mail: ntzortzakis@Googlemail.com

Abstract
The interest in the use of mixtures of inorganic and organic materials as substrate media in soilless culture
in greenhouses is increasing in parts of the world where the mixtures have not been used in common
practice. In this study, tomato (Lycopersicum esculentum Mill.), cv. Belladona, plants were grown over a
five-month period in a closed soilless culture system on four different substrates (pumice and its mixture
with 25% or 50% shredded maize stems), as well as NFT, in an unheated glasshouse. The impact of
substrate on total yield was differentiated among the substrates and trusses, and became significant after
the 3rd truss with marketable improved yield after the addition of shredded maize stems into the inorganic
substrate (pumice). The results of this study indicated that the highest total yield was obtained in plants
grown on pumice+50%maize, followed by pumice+25%maize, maize, NFT and lastly by pumice. The
decline in yield when maize substrate employed, should be attributed to the reduction in volume of this
material due to decomposition. The fact that pumice substrate alone resulted in lower yields than maizecontaining substrates suggested better nutritional conditions in the latter. The greater number of
fruits/plant produced in the 1st truss whereas no differences observed in the other trusses. Maizecontaining inorganic substrates significantly improved mean fruit weight during the whole yielding
period. The mean fruit weight was lower in the first two trusses, comparing with the 3rd-5th trusses, and
this may be due to the greater number of fruits produced on the first two trusses. In exception, plants
grown on maize, produced higher mean fruit weight in the initial trusses, and this may attributed to the
higher substrate temperature recorded due to decomposition process. Plants grown on maize resulted on
greater fruit firmness compared with other substrates up to the 4th truss. Plants grown on
pumice+50%maize produced low values in fruit juice TSS, inevitably because they also had high number
of fruits/plant. No differences observed on the pH and the EC of the tomato fruit juice between the
treatments. Shredded maize stems can serve as a partial substitute for inorganic media while maintaining
or improving tomato fruit yield and fruit quality. The new information gained from this study could be
useful for hydroponic tomato production in the greenhouses of the Mediterranean region.
Keywords: Lycopersicum esculentum, maize, NFT, organic materials, pumice, soilless culture, tomato,
yield, truss

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[Effect of the substrate in tomato soilless culture]

Introduction
Since the late 1970s, as an alternative to expensive chemical soil disinfection methods and possible plant
residues, many growers have switched progressively to soilless culture. The soilless growing system of
protected vegetables has been developed due to a number of advantages and avoids problems associated
with decreasing fertility of natural soils, due to disease limitations and the increase in salinity (Verdonck,
1975; Olympios, 1992). In addition, growers needed to improve the efficiency and quality of the products,
by means of a better control of production through technological innovations in the nursery. Soilless
culture uses various substrates, depending on the local conditions and opportunities (Moinereau et al.,
1987), each one with its own specific physico-chemical properties.
The trend for the Mediterranean region is to use a closed system and bags filled with different substrates
to obtain the maximum advantages of each individual substrate. This type of production system also
allows growers to improve efficiently yields and product quality. However, rising costs of materials for
certain soilless systems, the disposal of some materials after use, and the lack of grower knowledge on the
use of substrates, limits the development of soilless culture in some parts of the world. The possibility of
using different materials for substrate, locally available and less costly than those imported, as well as
with no pollution limitations, but adequate physical and chemical properties (Verdonck, 1975), is a major
factor to solve these problems.
Several studies reported the favorable effect of organic media on plant growth (Hardgrave and Harriman,
1995; Ayuso et al., 1996; Tzortzakis and Economakis, 2005). These effects are direct, such as absorption
by the plants of the humic compounds that affect membrane permeability and certain enzymatic activities
(Chen and Aviad, 1990; Pinton et al., 1992), or indirect, such as stimulation of microbiological activity,
and increased cationic exchange capacity (CEC) in plants (Yu and Komada, 1999). Inorganic substrates,
such as vermiculite, sand, perlite, and pumice, are chemically inert, making it possible to supply nutrients
in a controlled manner (De Rijck and Schrevens, 1998). Methods such as composting, aging, washing of
locally available material, mixing inorganic and organic substances, and adding fertilizer are also
common practices in greenhouse production (Nichols, 1981; Yates and Rogers, 1981). These practices
increase substrate porosity and water holding capacity (Hardgrave and Harriman, 1995), reduce or
eliminate toxicity problems associated with organic or inorganic substrates and unsuitable C:N ratios, and
overcome limitations of individual materials (Nichols, 1981; Yates and Rogers, 1981). Organic substrates
with a moderate, or low biostability, will release the available nutrients, decrease porosity, increase
salinity due to mineralization and vary in their chemical properties, such as pH, electrical conductivity
(EC) and CEC as a consequence of the decomposition of the substrate’s organic matter (Lemaire, 1995,
1997). Decomposition could result in increased temperatures, retardation of nutrient release, and
alteration of the properties of the growing medium (Handreck, 1992). Organic products such as wheat
straw and flax waste, with a medium to low biostability (Bunt, 1976) cannot be used directly for making
substrates; they have to be composted, to the extent where they are biostable (Lemaire, 1997).
Worldwide, 12% of the hydroponic industry uses organic media as a substrate and/or as compost
(Donnan, 1998).
Many investigators have studied the changes in fruit content that occur during ripening. Customer tests
indicated that firmness and flavour are important criteria for high tomato quality (Vesseur, 1990). The
flavour of salad tomatoes and the value of processing tomatoes are closely related to the concentration of
total soluble solids (TSS) in fruit (Adams, 1987). Several studies have shown that the typical tomato
flavour depended on the ratio between sweet and acid tastes and that sugars have been important taste
constituents (Kader, 1986). Acid content is an important component of flavour and the differences in
sources can be explained by variation in pH and titratable acids. Moreover, an increased interest in tomato
products has been created by the fact that their consumption has been correlated to a reduced risk of some
types of cancer (Clinton et al., 1996).
Maize (Zea mays L.) is an important crop grown worldwide. The edible products (green cobs and grains)
constitute only 10-15% of the total harvest while the leftovers (stover) are generated in large quantities
annually (Akanbi and Togum, 2002). Several studies examined the beneficial effect of maize compost
applied on soil, resulted in soil enrichment with organic matter and increased yields (Hartz et al., 1996;
Akanbi and Togum, 2002) as well as in soilless culture (Tzortzakis and Economakis, 2005; 2007).
The work presented here, designed to study the use of different substrates in tomato culture, in order to
access the suitability of some local materials for soilless cultivation, comparing inorganic and organic
mixtures (pumice:maize), for tomato. Although pumice also possess some of these qualities, maize is
easier to dispose of than pumice. It might also be possible to mix other organic- or inorganic-media (i.e
perlite, sand) with maize to improve the chemical and physical properties of the substrate.

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Materials and Methods
The experiment was carried out in an unheated glasshouse with a North-South orientation at the Institute
for Subtropical Plants and Olive Trees at Chania, Greece. Two substrates, pumice, and shredded maize
(Zea mays L.) stems, and mixtures of these, were used to create four treatments which were: 1) 100%
pumice, 2) 75% pumice + 25% maize, 3) 50% pumice + 50% maize, 4) 100% maize and 5) NFT.
Physicochemical properties of the substrates were presented in previous studies (Tzortzakis and
Economakis, 2005).
Maize plants were grown for yield of corn in the same region in early spring. Plants were not treated with
synthetic fertilizer or pesticides, but animal composted manure was incorporated 4 months before sowing.
Stalks (as by-products) were obtained after fruit harvest in late summer, were chopped, dried by exposure
to ambient temperature and shredded by grinding. Shredded dried maize stems were kept for 45 days in a
tank, frequently irrigated to cause decomposition, without adding any nitrogen source, and then used
alone or as mixes as substrates in bags.
The indeterminate tomato, cv. Belladona (HAZERA, Israel), which has an extended self-life with large
fruits (180-220 g), was grown in the bags. Seedlings at the first truss production were purchased from a
commercial nursery and placed in the various substrates. Maximum, minimum and average temperature,
and relative humidity were recorded every 30 min, with a data logger placed in the middle of the
glasshouse. Photosythetically active radiation (PAR) was monitored with a Millivolt Integrator (Fig. 1).
37

300000
250000

31
28

200000

25

150000

22
19

100000

16

PAR (E m-2)

Temperature (T o C)

34

50000

13
10

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Weeks

PAR

Day ToC

Night ToC

Figure 1. Weekly day and night temperature (oC) as well as photosynthetic active radiation (PAR; E.m-2)
in the glasshouse during the cultivation period for tomato plant in soilless culture.
Each substrate was arranged in a single row on a greenhouse trough in nine 16 L capacity bags (two
plants in each bag), with 18 plants/row. Rows were 1.2 m apart and plants were separated in bags by 0.4
m. An additional 18 plants were placed in an NFT trough with a 3% slope. NFT is a system in which
plants are grown in troughs in which shallow nutrient solution (up to 1 cm) is continuously re-circulated.
Of the 18 plants in each row, nine constituted a treatment, with the 3rd through 11th plants used for data
collection. Irrigation emitters were placed at the base of each plant, and the bags had slits in the bottom
to allow drainage. Before transplanting, the bags were soaked with a full strength nutrient solution, made
using commercial fertilizers with 1.44N-0.25P-1K a.i. (v/v/v) content. Pumice was originated by Lava
(Yali, Greece). As plants grew, all lateral shoots were removed manually and the resulting single stem
was trained up a string according to the high wire system that plants were layered when they reach the
wire (2.3 m) and lower old-leaves were removed periodically, avoiding foliage self-shading. Temperature
sensors connected to a data logger using thermistors monitored temperatures in air, and in maize and
pumice substrates.
The soilless culture system was closed (50 L capacity catchment tanks). A solution (1:100 v/v) in water
containing the following concentration of nutrients: NO3-N=9.6, K=6.8, ΡΟ4-P=1.7, Ca=4.0, Mg=2.8 and
Na=1.3 mmol·L-1, respectively; and Β=27, Fe=73, Μn=17, Cu=3.6, Zn=6.6, and Μο=1.2 µmol·L-1,
respectively, was applied with a drip irrigation system, via emitters for the individual plants, by means of
pressure pumps. Nutrient solution of a similar composition was used in the NFT unit at a continuous flow

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rate of 3 L·min-1. Target pH and EC values of the nutrient solution were 6.0 and 2.15–2.40 dS·m-1
respectively according the enrichment during fruit setting with iron chelate (4 ppm Fe) and potassium
sulphate (75 ppm K). Fertigation was applied during the daytime through a timer for 5 min/hr, every 0.5
hr after fruit set, at a flow rate of 100 mL·min-1. Actual pH values of the nutrient solution collected in
catchment tanks fluctuated between 6.7 and 7.8 while EC was between 1.77-2.72 dS·m-1.
The harvesting period lasted for 60 days with up to thirteen harvests were performed. Total yield and fruit
number produced per truss recorded throughout. The quality assessment included fruit fresh weight, flesh
firmness, pH and EC of the juice, total soluble solids (TSS), titratable acidity (TA). The fruits were left on
a bench in the lab, till full colour development. Fruits, at the pink stage, were weighed immediately after
harvest. Fruit firmness was measured by a Bishop FT Oil model pressure tester (probe 7.9 mm). TSS, pH
and EC of the extracted fruit juice was measured by a hand refractometer, a pH meter and an EC meter
respectively. TA was measured by titration with 0.1 N sodium hydroxide and expressed as a citric acid
percentage.
Data were analysed using SPSS (SPSS Inc., Chicago, USA) and first tested for normality and subjected to
univariate analysis of variance on multiple factors (ANOVA) followed by analysis of mean (one-way
ANOVA), investigating significant differences between substrates.

Results and Discussion
Day and night temperatures fluctuated during the cultivation period with a sharp decrease at the end of
March (week 13), when PAR also decreased. PAR and temperature increased rapidly after week 18 till
the end of the experiment from 5X104 E·m-2 and 21oC to 28X104 E·m-2 and 26oC, respectively (Fig. 1).
Plants in all treatments grew vigorously and no deficiency symptoms were observed during the
experiment.
Effect of the substrate on total yield and fruit number produced per truss
The impact of substrate on total yield was differentiated among the substrates and different trusses, and
became significant after the 3rd truss (Fig. 2). Indeed, the addition of different ratios of shredded maize
stems into the inorganic substrate (pumice) resulted in improved yield, whereas no differences observed
for pure pumice and NFT. The fact that pumice alone gave lower yields than maize-containing substrates
(with the highest yield observed in plants grown on pumice+50%maize) suggested better nutritional
conditions in the latter being in accordance with Padem and Alan (1994) in pepper cultivation. However,
Bohme et al. (2001) reported no differences between organic (coconut-fibre) and inorganic (perlite and
rookwool) substrates on yield of cucumber plants.

Cumulative yield (Kg/plant)

6

Pumice
Pum.+25%maize
Pum.+50%maize
Maize
NFT

5
4
3
2
1
0
1

2

3

4

5

6

Truss

Figure 2. Effect of the substrate medium on the cumulative yield produced per truss during the cultivation
period in hydroponically grown tomatoes
The total number of fruits/plant harvested from each treatment varied and probably affected fruit quality
characteristics. The greater number of fruits/plant produced in the 1st truss with no great differences
observed in the other trusses (Fig. 3). However, plants grown on pumice produced significantly lower
total number of fruits/plant than in maize shredded stems which contrasts with the findings of Hardgrave

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who reported that organic substrate (peat, bark and straw) produced significant lower or (hortifibre)
insignificant number of cucumber fruits/plant compared with the inorganic substrate (rockwool;
Hardgrave and Hariman, 1995). It’s worthy to mention that maize substrate volume reduced through the
cultivation period because of the decomposition took place. This was resulting in decreased plant growth
(vegetative, flowering and fruiting) compared with the other substrates, during the last 3-4 weeks,
eventually due to the lack of porosity and/or inadequate water availability to the roots in the middle of the
day when plant wilting occurred.

Number of fruits/plant

8

Pumice
Pum.+25%maize
Pum.+50%maize
Maize
NFT

7
6
5
4
3
2
1

2

3

4

5

6

Truss
Figure 3. Effect of the substrate medium on the mean fruit number produced per truss during the
cultivation period in hydroponically grown tomatoes
Effect of substrate on fruit quality
An increase in fruit size was associated with improved climatic conditions (PAR, temperature) for the
majority of the substrates. Indeed, the mean fruit weight was lower in the first two trusses, comparing
with the 3rd-5th trusses, and this was due to the greater number of fruits on the first two trusses. In
exception, plants grown on maize, produced greater mean fruit weight in the initial trusses, and this may
attributed to the higher substrate temperature recorded; on average 1.6oC and 2oC during day and night
respectively comparing with the other substrates. The increased temperature of the organic substrate
should be due to the microbial decomposing activities in it. Maize-containing inorganic substrates
significantly improved mean fruit weight over the whole yielding period, whereas no differences
observed in the no-mixed substrates. These results are in accordance with previous studies reported
insignificant differences between pure organic and inorganic substrates on mean fruit weight (g) of
cucumber and tomatoes plants (Hardgrave and Hariman, 1995) and (Islam et. al. 2002) respectively.
Plants grown on maize resulted on greater fruit firmness compared with other substrates at the 1st till the
4th truss, whereas no differences observed for the 5th and 6th truss. TSS (oBrix) content of the fruit was
found to differ significantly between substrates. Plants grown on pumice and maize compared with
pumice+50%maize, pumice+25%maize and NFT produced a higher value of TSS. Plants grown on
pumice+50%maize produced low values in fruit juice TSS, inevitably because they also had high number
of fruits/plant (Table 3 and Fig. 2). Higher sugar and organic acid content improved tomato fruit quality
(Davies and Hobson, 1981). Islam et al. (2002) reported no differences in TSS on tomato fruit juice
between organic and inorganic substrates in accordance with the present study. The increase in fruit TSS
values from the 3rd to the 6th truss should be attributed to the gradually increasing light intensity during
the ripening period (Table 2 and Fig. 1), being in accordance to Went (1957) who stated that sugar
content is entirely a function of light intensity. The pH and the EC of the tomato fruit juice were not
significantly different on tomato cultivation with different substrates being in accordance with Islam et al.
(2002). Low pH is associated with high fruit quality (Davies and Hobson, 1981) and was recorded in the
substrates, which produced early yield (maize and pumice) as reported in previously (Tzortzakis and
Economakis, 2008). The increasing earliness of the organic substrates diminished the fruit quality for the
first tomato truss (Angelis et al., 2001).

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[Effect of the substrate in tomato soilless culture]
Shredded maize stems can serve as a partial substitute for inorganic media while maintaining or
improving tomato fruit yield and fruit quality. The increased temperature of the maize substrate due to
microbial decomposition could be beneficial for crops grown under unheated glasshouse conditions. Root
temperature is thought to be one of the major factors that directly affected plant growth and increased
yield (Moss, 1983). Several studies reported the beneficial impacts of mixtures between inorganic and
organic media (Padem and Alan, 1994; Stoffella and Graetz, 2000; Tuzel et al., 2001). The results of this
study indicated that the highest total yield was obtained on pumice+50%maize, followed by
pumice+25%maize, maize and lastly by pumice. The decline in yield on maize should be attributed to the
reduction in volume of this material due to decomposition. The fact that plants grown on pumice alone
gave lower yields than maize-containing substrates suggested better nutritional conditions in the latter.
Total number of fruits per plant was greater on plants grown on NFT followed by maize substrate. Fruit
quality characteristics were differently affected by the substrate. Adding maize shredded stems in pumice
improved its properties (providing buffering capacity, essential elements, adsorptive and absorptive
properties) as pure inorganic substrate, leading to higher yields of tomato plants and better fruit quality.
Maize itself is a low cost, worldwide available and effective organic material which should be used for
tomato growing as a substrate more efficiently in a condensed form (compressed) because of quick
decomposition under the Mediterranean conditions.
The new information gained from this study could be useful for hydroponic tomato production, in the
glasshouses of the Mediterranean region. Further investigation is needed to determine the presence of
maize in pumice could promise the prolonged reuse of the mixture, as well as the point of condensation
(compression) of maize stems to assure their successful use (biostability) as substrate and the alternative
use of maize substrate on soilless culture introducing plants with shorter life-cycle ie. lettuce and spinach.
Table 1. Effect of the substrate medium on the mean fruit weight (g) on different trusses during the
cultivation period in hydroponically grown tomatoes
Truss

Pumice

Pum+25%Maize

Pum+50%Maize

Maize

1st truss

147± 6.1 (a)

2nd truss

186± 9.1
(ab)
232± 14.1
(a)
218± 12.7
(a)
229± 20.2 (ab)

153± 5.3
(ab)
189± 10.3
(ab)
240± 13.0
(a)
243± 13.9
(a)
248± 10.7
(ab)
210± 21.1
(a)

170± 6.5
(a)
172± 11.5
(b)
240± 13.5
(a)
236± 12.8
(a)
277± 15.8
(a)
227± 16.3
(a)

161± 5.5
(ab)
213± 18.2
(a)
223± 9.5
(ab)
200± 11.6
(a)
199± 16.5
(b)
157± 10.2
(a)

3rd truss
4th truss
5th truss
6th truss

196± 21.2
(a)

NFT
135± 6.1
(b)
162± 14.8
(b)
192± 11.1
(b)
227± 13.0 (a)
253± 15.9
(ab)
230± 10.3
(a)

Table 2. Effect of the substrate medium on the fruit firmness (Kg) per truss during the cultivation period
in hydroponically grown tomatoes
Truss
st

1 truss
2nd truss
3rd truss
4th truss
5th truss
6th truss

Pumice
1.28± 0.092
(b)
1.03± 0.058
(a)
1.25± 0.079
(ab)
0.91± 0.035
(b)
1.15± 0.064
(a)
1.00± 0.104
(a)

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Pum+25%maize

Pum+50%maize

Maize

1.24± 0.084
(b)
0.98± 0.608
(a)
1.18± 0.075
(ab)
0.85± 0.060
(b)
1.17± 0.057
(a)
0.97± 0.048
(a)

1.33± 0.086
(b)
0.97± 0.062
(a)
1.23± 0.036
(a)
0.98± 0.037
(a)
1.13± 0.063
(a)
1.03± 0.025
(a)

1.65± 0.099
(a)
1.18± 0.069
(a)
1.17± 0.061
(ab)
1.28± 0.071
(a)
1.03± 0.064
(a)
1.08± 0.071
(a)

NFT
1.00± 0.086
(b)
0.93± 0.060
(c)
1.03± 0.066
(b)
1.32± 0.107
(b)
1.05± 0.068
(a)
1.09± 0.056
(a)

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Table 3. Effect of the substrate medium on the total soluble solids (TSS) of the fruit juice per truss during
the cultivation period in hydroponically grown tomatoes
Truss
st

1 truss
2nd truss
3rd truss
4th truss
5th truss
6th truss

Pumice

Pum+25%maize

Pum+50%maize

Maize

3.50± 0.100
(a)
4.14± 0.147 (a)

3.50± 0.103
(a)
3.37± 0.155
(bc)
3.50± 0.151
(bc)
3.76± 0.136
(b)
3.93± 0.062
(b)
4.28± 0.101
(a)

3.71± 0.163
(a)
3.34± 0.115
(bc)
3.66± 0.105
(b)
3.89± 0.111
(b)
4.05± 0.079
(ab)
4.66± 0.333
(a)

3.31± 0.203
(a)
3.65± 0.096
(ab)
3.84± 0.164
(ab)
4.36± 0.078
(a)
4.34± 0.182
(a)
4.57± 0.229
(a)

4.18± 0.216
(a)
4.28± 0.240
(a)
4.21± 0.101
(ab)
4.25± 0.250
(a)

NFT
3.58± 0.172
(a)
3.22± 0.173
(c)
3.10± 0.124
(c)
3.60± 0.111
(b)
3.96± 0.132
(b)
3.50± 0.129
(b)

Table 4. Effect of the substrate medium on the pH of the fruit juice per truss during the cultivation period
in hydroponically grown tomatoes
Truss
st

1 truss
2nd truss
3rd truss
4th truss
5th truss
6th truss

Pumice

Pum+25%maize

Pum+50%maize

Maize

3.86± 0.027
(a)
4.00± 0.037
(a)
4.18± 0.026
(a)
4.45± 0.117
(ab)
4.12± 0.047
(ab)
4.05± 0.050
(a)

3.80± 0.034
(a)
4.20± 0.103
(a)
4.13± 0.023
(a)
4.41± 0.067
(ab)
4.10± 0.032
(b)
4.11± 0.059
(a)

3.82± 0.026
(a)
4.21± 0.101
(ab)
4.16± 0.023
(a)
4.32± 0.027
(a)
4.19± 0.028
(ab)
4.16± 0.033
(a)

3.75± 0.024
(a)
3.90± 0.036
(b)
4.19± 0.024
(a)
4.35± 0.069
(ab)
4.16± 0.036
(ab)
4.14± 0.029
(a)

NFT
3.81± 0.012
(a)
4.00± 0.056
(ab)
4.15± 0.023
(a)
4.20± 0.053
(b)
4.21± 0.031
(a)
4.11± 0.031
(a)

Table 5. Effect of the substrate medium on the EC (mS/cm) of the fruit juice per truss during the
cultivation period in hydroponically grown tomatoes
Truss

Pumice

Pum+25%maize

Pum+50%maize

Maize

1st truss

3.16± 0.071
(a)
2.87± 0.104
(a)
3.27± 0.116
(ab)
3.66± 0.320
(a)
3.36± 0.228
(a)
2.67± 0.575
(a)

3.11± 0.088
(a)
3.09± 0.084
(a)
2.87± 0.077
(b)
3.17± 0.125
(a)
2.97± 0.091
(a)
3.26± 0.109
(a)

3.03± 0.146
(a)
3.05± 0.102
(a)
3.07± 0.103
(a)
3.45± 0.113
(a)
3.31± 0.105
(a)
3.25± 0.064
(a)

3.10± 0.143
(a)
3.10± 0.088
(a)
3.15± 0.075
(ab)
3.34± 0.135
(a)
3.17± 0.092
(a)
3.43± 0.275
(a)

2nd truss
3rd truss
4th truss
5th truss
6th truss

NFT
3.12± 0.115
(a)
2.97± 0.092
(a)
2.94± 0.091
(b)
3.14± 0.127
(a)
3.28± 0.104
(a)
3.34± 0.187
(a)

Acknowledgements
This work was supported in part by M.A.I.Ch. (Mediterranean Agronomic Institute of Chania, Greece)
and N.AG.RE.F. (National Agriculture Research Foundation of Greece, Chania). We thank Dr A. Borland
(University of Newcastle, UK) for the critical revision to the manuscript.

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