Soil Science and Plant Nutrition

ISSN: 0038-0768 (Print) 1747-0765 (Online) Journal homepage: http://www.tandfonline.com/loi/tssp20

Effect of the addition of activated charcoal to

the nutrient solution on the growth of tomato in
hydroponic culture

Jing Quan Yu , Kwang Seek Lee & Yoshihisa Matsui

To cite this article: Jing Quan Yu , Kwang Seek Lee & Yoshihisa Matsui (1993) Effect of the
addition of activated charcoal to the nutrient solution on the growth of tomato in hydroponic culture,
Soil Science and Plant Nutrition, 39:1, 13-22, DOI: 10.1080/00380768.1993.10416970

To link to this article: http://dx.doi.org/10.1080/00380768.1993.10416970

Published online: 04 Jan 2012.

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 Soil Sci. Plant Nutr., 39 (I), 13-22, 1993 13

Effect of the Addition of A c t i v a t e d Charcoal to the

N u t r i e n t Solution on the G r o w t h of T o m a t o in
Hydroponic Culture

Jing Quan Yu, K w a n g Seek Lee, and Y o s h i h i s a Matsui

Facultj: of Agriculture, Shhnane University, Matsue, 690 Japan

Received September 24, t991

The addition of activated charcoal to a n u t r i e n t solution for t h e hydroponic

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culture of t o m a t o resulted in both a considerable decrease in the carbon

c o n c e n t r a t i o n in the solution and significant increases in the dry w e i g h t of
plant and in f r u i t yield. No appreciable changes were b r o u g h t about in the
c o n c e n t r a t i o n s of m a j o r and t r a c e elements b o t h in the solution and in the
plant by the addition of activated charcoal. The n u t r i e n t solution collected
a f t e r the cultivation without activated charcoal was toxic to the g r o w t h of
t o m a t o seedlings and to the g e r m i n a t i o n of t o m a t o seeds. No toxic effect was
observed for the residual n u t r i e n t solution t r e a t e d with activated charcoal.
These facts indicate t h a t the g r o w t h of t o m a t o in hydroponic culture is inhib-
ited by organic substances which arise f r o m the r o o t exudates and are removed
f r o m the n u t r i e n t solution by adsorption on activated charcoal.
K e y Words: activated charcoal, hydroponics, p h y t o t o x i c substance, r o o t
exudate, t o m a t o .

Hydroponics has lately attracted a great deal of attention, since it enables to avoid the
growth injury associated with consecutive cropping, a notorious problem in greenhouse crop
production, and to control easily rhizosphere factors such as temperature, aeration, pH,
water supply, and nutrient supply. However, reduction of plant growth has often been
observed, especially in hydroponic culture with a re-circulating nutrient solution system,
when the plant was bearing a large amount of fruits (Tucker 1981; Rahman and Newton
1984). This phenomenon was ascribed to the accumulation of phytotoxic substances and ion
imbalance in the nutrient solution. Periodic renewal of the nutrient solution was recom-
mended to solve this problem (Shimada 1989). However, this practice is associated with the
loss of a large amount of fertilizer and requires a great deal of labor. As early as in the 1950s,
Takijima and Hayashi (1959) and Hirayoshi et al. (1959) reported that the residual nutrient
solution after cultivation of tomato was toxic to the plant itself and its relatives rather than
to other species. Recently, several investigators have reported that some plants exuded
substances which were toxic to the plants themselves (autotoxicity) (Young 1984) or to other
species (allelopathy) (Tang and Young 1982; Pope et al. 1985; Ito et al. 1986; Yasuda et al.
1991). However, limited information is available about the chemical structure and bio-
activity of phytotoxic substances.
The present investigation was undertaken to confirm the accumulation of phytotoxic
substances in the nutrient solution for hydroponic culture of tomato, as the initial step to
 14 J.Q. YU, K.S. LEE, and Y. MATSUI

identify the substances. For this purpose, we examined the effect of the addition of activated
charcoal to a nutrient solution on the growth of tomato and on the chemical composition
of the nutrient solution and plant lamina. It is known that activated charcoal (Rahman and
Newton 1984), together with Amberlite X A D - 4 resin (Tang and Young 1982; Yasuda et al.
1991), enables to remove phytotoxic substances from a nutrient solution to improve the
growth of plants. We also examined the phytotoxic effect of the nutrient solutions collected
during cultivation on the germination of seeds and the growth of seedlings of tomato.

MATERIALS AND METHODS

C u l t i v a t i o n of t o m a t o . Tomato plants (Lycopersicum esculentum Mill, cv. TVR-2)

were cultivated three times, i.e., in the fall of 1989 and in the spring and fall of 1990, using
hydroponic systems in a greenhouse. Each system was composed of a culture bed (94 cm •
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94 cm • 5 cm), a nutrient solution tank ( 150 L), a pump with a time switch, an inlet with an
air-sucker and a heater with a temperature controller (Fig. 1). The nutrient solution in the
tank was pumped up and introduced to a bed through an air-sucker for 15 rain every hour.
The excess amount of the solution in the bed overflowed from the outlet and returned to the
tank by gravitation. The volume of the nutrient solution in the bed was adjusted to 30 L at
the beginning of cultivation and decreased with increasing volume of plant roots.
The outline of the cultivation practices is shown in Table 1. Tomato seeds were sown
in rockwool and raised in a nutrient solution prepared based on the ENSHI formula by use
of solid fertilizers for hydroponic culture (Ohtsuka Chemicals Co.). The full strength
ENSHI solution contains Ca(NO3)2, 4.0; KNOa, 8.0; NH4H2PO4, 1.3; MgSO4, 2.0 in mmol
L -1 and Fe, 2.9; Mn, 0.5; B, 0.5; Cu, 0.02; Zn, 0.04; Mo, 0.01 in mg k -1. The pH and EC
values at 25~ of the solution were 6.8 and 2.5 dS m -~, respectively. Six (1989) or eight (1990)
uniform seedlings with the third to the fourth leaves were transplanted to each hydroponic
bed and cultivated for ca. four or six months with an E N S H I solution. Plants were topped
at the level of two leaves above the third (fall cropping) or fifth (spring cropping) truss.
T r e a t m e n t s in c u l t i v a t i o n e x p e r i m e n t s . The following two kinds of treatments
were applied for the cultivation experiments in the fall of 1989: TI) Cultivation without the
renewal of the nutrient solution but with the replenishment of the water and nutrients
consumed by the plants. T2) Cultivation similar to TI, except that 600g of granular
activated charcoal (Takeda Chemical Industry Co., Shirasagi W2C, 4-8 mesh) was added to

Fig. 1. Schematic diagram of the hydroponic system used. B,

culture bed; H, heater; I, inlet with air-sucker; O, outlet; P, pump;
S, nutrient solution; T, tank.
 Effect of Charcoal on the Growth of Tomato in Hydroponics 15

Table 1. Outline of cultivation practices in the three experiments.

Expt. 1 Expt. 2 Expt. 3
Year 1989 1990 1990
Sowing Aug. 14 Feb. 21 Aug. 22
Planting Sep. 13 Mar. 17 Sep. 22
Addition of charcoal Sep. 16 Apr. 11 Oct. 22
Topping Nov. 4 May 20 Nov. 31
End of harvesting Dec. 25 Jul. 20 Feb. 18, 1991
No. of trusses 3 5 3
No. of plants 12 8 8

the nutrient solution (150 L) in the tank. Each treatment was duplicated by use of two
hydroponic systems for each treatment. Experiments in the spring of 1990 were carried out
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with two additional treatments: T3) Cultivation similar to TI, except that the initial nutrient
solution was prepared by mixing 75 L of the residual nutrient solution (TI) in the fall
cropping of 1989 and 75 L of a freshly prepared nutrient solution. T4) Cultivation similar
to T3, except that 600 g of granular activated charcoal was added to the nutrient solution in
the tank. Experiments in the fall of 1990 were carried out with four treatments, in which the
amounts of granular activated charcoal added to freshly prepared nutrient solutions in the
tanks were 0, 300, 600 and 1,200 g.
In order to keep the nutrient composition of the nutrient solutions as constant as
possible during cultivation, the nutrient solutions were collected every week and the
concentrations of major nutrient elements were determined. The amounts of nutrients
consumed by the plants were compensated by the addition of fertilizers to the nutrient
solutions within a week after sampling time. Water consumed was replenished at each
sampling time. The pH was adjusted to 5.5 to 6.8 by the addition of an aqueous H=SO4 or
NaOH solution.
Chemical analysis. At the end of cultivation, plants were harvested and separated
into root, stem, petiole, lamina, fruit and fruit stalk. Each part was dried at 80~ weighed
and ground with a vibrating mixer mill. Digestion was initiated with concentrated HNOa
and completed with HCIO4 in a test tube on a hot block bath. Concentrations of P, K, Ca,
Mg, S, Fe, Mn, Cu and Zn in either the nutrient solution or the digestion solution of the
plant samples were determined with an inductively coupled plasma emission spectrometer
(Shimadzu ICPS-2000) (Lee et al. 1991a). The concentrations of NOa-N and NH4-N in the
nutrient solution were determined by the methods described previously (Lee et al. 1991b).
The concentration of N in the plant was determined with a NC analyzer (Sumitomo
Chemical, NC-80). The concentration of C in the nutrient solution was also determined with
the NC analyzer after the nutrient solution was freeze-dried.
P h y t o t o x i e i t y o f n u t r i e n t solution. The nutrient solutions used for the assay were
collected three times during the fall cropping of 1989, i.e., on October 14 (vegetative stage),
November 24 (fruit-enlarging stage), and December 24 (fruit-maturing stage). The collected
solutions were stored at 4~ until use. Prior to the assay, the solutions were adjusted to pH
6 with aqueous NaOH or H2SO4 and to EC 2.2 dS m -1 with deionized water. The mineral
composition of the test solutions was also adjusted to become virtually equal to that of a
freshly prepared nutrient solution by the addition of mineral salts as needed. For the seed
germination assay, fifteen randomly selected tomato seeds were placed on two pieces of filter
paper in a Petri dish (90 mm in diameter) containing 12.5 mL of a test solution. The seeds
 16 ,I.Q. YU, K.S. LEE, and Y. MATSUI

were incubated in darkness at 200C for 7 d. Each treatment was replicated 4 times.
For the growth test of the seedlings, a 7-d-old tomato seedling with a root length of ca.
5 cm was immersed in 30 mL of either a residual oi" freshly prepared nutrient solution in a
18X 180 mm test tube. The seedling was allowed to grow in a growth chamber under a 12
h light regime at 280C and a 12 h dark regime at 180C. The light intensity was 8,500 lx at the
level of the seedling. The length of the root was determined as an index of seedling growth
after cultivation for 14 d. Each treatment consisted of about 20 seedlings. The TTC-reducing
activity of the seedling root was determined according to the method of Yoshida (1966). The
thermostability of phytotoxicity was examined by heating the residual nutrient solution in
the spring cropping of 1990 at 1210C under a pressure of 0.12 MPa for 2 0 m i n in an
autoclave. The autoclaved solution was tested by the assay described above.

RESULTS
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E f f e c t of a c t i v a t e d c h a r c o a l on t h e g r o w t h o f t o m a t o p l a n t
Table 2 shows the dry weight of various organs and fruit yield at the end of the fall
cropping in 1989. The addition of activated charcoal to the nutrient solution resulted in the
increase of the dry weight of most organs, which was particularly significant in the case of
the lamina and whole plant. The dry weight of the lamina, fruits, and whole plant, together
with the fruit yield, for the cultivation with activated charcoal was about 1.2 times that in
the case of cultivation without activated charcoal.
Table 3 shows the results at the end of the spring cropping in 1990. As in the previous
cropping, the dry weight of the lamina and fruits, together with the fruit yield, increased by
a factor of about 1.2 by the addition of activated charcoal. The dry weight and fruit yield
were not significantly different regardless of the presence of the residual solution from the
previous cropping in the initial nutrient solutions.
Table 4 shows the effect of the amount of activated charcoal added to the nutrient
solution on the growth of tomato for the fall cropping in 1990. The growth of the tomato
plants in this season was less satisfactory than that in the other seasons, due to unsuitable
weather conditions. However, as in the other seasons, the dry weight of the lamina and fruits
increased by the addition of activated charcoal. Moreover, the increase in the amount of
activated charcoal resulted in the increase of the dry weight.

Table 2. Efl'ect of addition of charcoal to the nutrient solution

on dry matter and fruit production of tomato (fall of 1989).
Organ Control Charcoal
Dry weight (g plant -1)
Root 29.0 30.1
Stem 33.9 34.8
Petiole 38.3 43.3
Lamina 73.6 86.5*
Fruit Stalk 8.6 7.7
Fruit 147.0 174.9
Total 330.4 377.3**
Fresh weight (kg plant -1)
Fruit 3.1 3.7
* 5% and ** 1% levels of significance (n=2) by t-test.
 Effect of Charcoal on the Growth of Tomato in Hydroponics

Table 3. Effect of addition of charcoal to the nutrient solution on dry matter and
fruit production of tomato (spring of 1990).
Treatmenta
Organ
T1 T2 T3 T4
Dry weight (g plant-:)
Root 28.4 33.4 31.2 35.2
Stem 46.0 47.3 45.5 46.8
Petiole 35.7 43.7 34.3 38.4
Lamina 76.0 98.7 79.4 92.3
Fruit Stalk 9.6 11.6 9.2 9.0
Fruit 218.1 258.4 207.3 247.3
Total 413.8 493.1 407.0 469.0
Fresh weight (kg plant 1)
Fruit 4.4 5.2 4.0 4.8
" TI, freshly prepared nutrient solution; T2, T1 +activated charcoal; T3, nutrient solution containing residual
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nutrient solution; T4, T3+aetivated charcoal.

Table 4. Effect of addition of charcoal to the nutrient solution on dry matter and
fruit production of tomato (fall of 1990).
Amount of charcol added (kg (150L) :)
Organ
0 0.3 0.6 1.2
Dry weight (g plant-:)
Root 9.8 10.7 11.7 11.7
Stem 66. l 66.1 67.7 74.8
Petiole 42.6 50.1 53.0 71.8
Lamina 93. t 103.3 114.6 143.3
Fruit Stalk 6.8 8. I 9.9 13.7
Fruit 76.6 90.4 90.3 103.4
Total 295.0 328.7 347.2 418.7
Fresh weight (kg plant -1)
Fruit 1.5 1.7 1.8 2.0

Carbon c o n c e n t r a t i o n in n u t r i e n t s o l u t i o n s
Figure 2 shows the changes in the c a r b o n c o n c e n t r a t i o n of the n u t r i e n t s o l u t i o n s d u r i n g
the fall c r o p p i n g in 1989 a n d spring c r o p p i n g in 1990. T h e i n i t i a l c a r b o n c o n c e n t r a t i o n in
a freshly prepared n u t r i e n t s o l u t i o n was ca. 3 mg L :. It was assumed that the m a i n c a r b o n
source was E D T A ( e t h y l e n e d i a m i n e t e t r a a c e t i c acid) and the b i c a r b o n a t e ion. E D T A was
present in the fertilizer as the c h e l a t i n g agent of Fe. The c a r b o n c o n c e n t r a t i o n in the
s o l u t i o n s collected d u r i n g c u l t i v a t i o n was always higher t h a n that in a freshly prepared
solution. T h e c o n c e n t r a t i o n of the b i c a r b o n a t e ion in the s o l u t i o n r e m a i n e d v i r t u a l l y
constant (1.7• m g L -~) d u r i n g the c u l t i v a t i o n . Thus, the increments of the c a r b o n
c o n c e n t r a t i o n s were attributed to organic exudates from the t o m a t o root. It is i m p o r t a n t to
note that the c a r b o n c o n c e n t r a t i o n in the n u t r i e n t s o l u t i o n s c o n t a i n i n g activated charcoal
was always c o n s i d e r a b l y lower than that in the s o l u t i o n l a c k i n g charcoal. Activated
charcoal adsorbed organic substances on its particle surface and reduced the c a r b o n c o n c e n -
t r a t i o n in the s o l u t i o n . In the fall c r o p p i n g in 1990, the mean c a r b o n c o n c e n t r a t i o n in the
n u t r i e n t s o l u t i o n s collected biweekly was 17.5, 5.2, 4.9, and 2.7 mg L -~ for the treatments in
which the a m o u n t s of activated charcoal added to 150 L of n u t r i e n t s o l u t i o n s were 0, 300,
600 and 1,200 g, respectively. Thus, the increase in the a m o u n t of activated charcoal resulted
 18 J.Q. YU, K.S. LEE, and Y. M A T S U I

40 I I I I I

i I

32 A S
E

~ 24

~-
,,-I--'
C
16- /
O
/
=
n
8
(D
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(D
0 1 I I I I I I I
0 30 60 90 120 50 0 30 60 90 120 150

Days after s0~ing

Fig. 2. Changes in carbon concentrations in the nutrient solutions during cultivation in the fall of 1989
(A) and the spring of 1990 (B). Activated charcoal was added 33 (A) and 48 (B) d after sowing. O, cultivation
without activated charcoal (T I); e, cultivation with activated charcoal (T2); A, cultivation with the residual
nutrient solution which did not contain activated charcoal (T3); A T3 + activated charcoal (T4).

in the decrease in the carbon concentration in the nutrient solution.

C o n c e n t r a t i o n s o f m a j o r a n d t r a c e e l e m e n t s in n u t r i e n t s o l u t i o n s
Table 5 shows the concentrations of major and trace elements, as well as the values of
the pH and EC, in the initial nutrient solution and nutrient solutions collected every week
during the fall cropping in 1989. Apparently, the addition of activated charcoal to the
nutrient solution did not affect the concentrations of all the elements, except for Fe, in the
solution. The observed values of pH, EC, and the concentration of NO3-N were virtually
equal to the corresponding initial values. On the other hand, the concentrations of P, K, and
NH4-N were apparently lower than the initial values. The tomato plant absorbed these
elements more actively than water at the stages of fruit enlargement and ripening (Lee et al.
1991b), and the complete control of the concentrations was difficult. The concentrations of
Ca, Mg, and S were significantly higher than the initial values. These elements were absorbed
by the plant less actively than water and tended to accumulate in the nutrient solution (Lee
et al. 1991b).

C o n c e n t r a t i o n s o f n u t r i e n t e l e m e n t s in l a m i n a
Table 6 shows the concentrations of major and trace elements in the tomato lamina
which were collected at the end of the fall cropping in 1989. The concentrations of all the
elements, except for P and Fe, were not significantly differentregardless of whether the
cultivation was carried out with or without activated charcoal. The same results were
obtained in the experiments in the spring of 1990 (data not shown). These facts indicate that
the addition of charcoal to a culture solution or the reduction of the concentration of
organic compounds in the solution virtually did not affect the concentrations of the majority
 Effect of Charcoal on the Growth of Tomato in Hydroponics 19

Table 5. Effect of addition of charcoal on means and standard deviations (in parenthesis)
for pH, EC, and the concentrations of nutrient elements in solutions (fall of 1989).
Item Unit Control Charcoal Initial solution
pH 6.6 (0.7) 6.6 (0.6) 6.5
EC dS m -1 2.5 (0.2) 2.4 (0.2) 2.5
NOa-N mg L -~ 224 (28) 212 (26) 224
NH4-N mg L -1 2.5 (5.2) 2.3 (5.1) 18
P m g L -1 19 (13) 20 (14) 41
K m g L -1 287 (44) 273 (33) 312
Ca mg L -1 200 (23) 196 (20) 160
Mg mg L a 92 (32) 94 (37) 48
S r a g e -a 102 (23) 110 (27) 64
Fe mg L -~ 1.5 (0.9) 0.6 (0.6)** 2.9
Mn mg L -~ 0.02 (0.07) 0.04 (0.11) 0.5
B mg L -~ 0.69 (0.21) 0.78 (0.30) 0.5
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** 1% level of significance ( n = 2 6 ) by t-test.

Table 6. Effect of addition of charcoal to the nutrient solution on the concentrations of

nutrient elements in tomato lamina collected at the end of harvesting (fall of 1989).
Element Control Charcoal
(g kg -1 dry weight)
N 38 39
P 8.2 9.3*
K 60 53
Ca 45 47
Mg 5.2 5.2
S 20 23
C 350 350
Fe 0.087 0.064*
Mn 0.236 0.229
Zn 0.088 0.042
Cu 0.157 0.139
* 5N level of significance ( n = 12) by t-test.

Table 7. Phytotoxicity of nutrient solutions collected during cultivation in the

fall of 1989 with (T2) and without (TI) activated charcoal.
Seed germination Root length TTC reducing activity
Treatment Sampling time
(%) (cm) (g kg-* h -~)
Fresh solution 95 19.3 0.22
TI Oct. 14 93 17.4 0.16"
TI Nov. 24 90 14.6"** 0.13"
TI Dec. 24 87* 14.1"** 0.12"
T2 Oct. 14 93 2[.4 0.27
T2 Nov. 24 92 17.6 0.17
T2 Dec. 24 95 20.2 0.22
* 5% and *** 0. I% levels of significance by t-test.

o f t h e n u t r i e n t s in t h e t o m a t o p l a n t .
 20 J.Q. YU, K.S. LEE, and Y. MATSUI

P h y t o t o x i e i t y of nutrient solution
Table 7 shows the effect of nutrient solutions collected during cultivation on the seed
germination and on the length and TTC-reducing activity of the seedling root. The nutrient
solution collected at the end of cultivation in the absence of activated charcoal inhibited the
germination of seed, the growth of root, and the TTC-reducing activity. On the other hand,
the nutrient solution treated with activated charcoal did not show such an inhibitory effect.
No clear inhibitory effect was observed in nutrient solutions collected at the vegetative stage,
even in the absence of treatment with activated charcoal. Autoclaving did not alleviate the
phytotoxicity of the residual nutrient solution in the cultivation without activated charcoal
(data not shown).

DISCUSSION
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All the data on the hydroponic culture of tomato (Tables 2-4) showed that the addition
of activated charcoal was effective in improving the growth of the plant, especially that of
the lamina and fi-uits. When activated charcoal was absent in the solution, the carbon
concentration increased in general with the cultivation time (Fig. 2). On the other hand, the
concentration of the bicarbonate ion in the solution remained virtually constant during the
cultivation. Hence, the increase of the carbon concentration in the solution was considered
to be associated with the accumulation of organic exudates fi-om the roots or their secondary
products by microbes. The observed decrease in the carbon concentration with the addition
of activated charcoal was attributed to the adsorption of the organic substances on charcoal.
In contrast, the addition of activated charcoal to the nutrient solution did not affect the
concentrations of the majority of the elements in the solution (Table 5). Only the concentra-
tion of Fe decreased by the addition of activated charcoal. Although charcoal adsorbs
E D T A , a chelating agent of Fe, it is unlikely that the decrease in the Fe concentration was
responsible for the promotion of the growth of tomato by the treatment of activated
charcoal.
The addition of activated charcoal also did not affect the concentrations of major and
trace elements, except for P and Fe, in the tomato lamina (Table 6). The content of P in the
lamina increased by the charcoal treatment. Glass (1973) and Mersie and Singh (1988)
reported that the uptake of P by the roots of barley and tomato, respectively, w a s inhibited
by phenolic acids, which are known to be phytotoxic substances. Activated charcoal may
reduce the concentration of phenolic acids in the solution, and promote the absorption of
P by the tomato plant. Further studies are necessary to elucidate the mechanism. The
decrease of the Fe content in the lamina by charcoal may be due to the decrease of the Fe
concentration in the culture solution.
The length and TTC-reducing activity of the root o f t o m a t o seedlings, together with the
percentage of germination of the tomato seeds, decreased by the use of the culture solution
collected at the end of cultivation without activated charcoal (Table 7). Autoclaving of the
solution did not alleviate the phytotoxicity, indicating that heat-stable substances that
accumulated in the solution and not microbes were responsible for the phytotoxicity. The
toxic effects which were scarcely observed in the culture solution collected at the vegetative
stage became evident at the fruit enlarging and maturing stages. On the other hand, no toxic
effect was detected in the culture solution treated with charcoal. In these experiments, the
pH, EC, and the mineral composition of the test solutions were so carefully regulated that
they were not significantly different from one another. These results indicate that the
 Effect of Charcoal on the Growth of Tomato in Hydroponics 21

p h y t o t o x i c substances which a c c u m u l a t e d d u r i n g the h y d r o p o n i c c u l t u r e o f t o m a t o were

r e m o v e d f i o m the s o l u t i o n by t r e a t m e n t with activated c h a r c o a l .
T h e T T C - r e d u c i n g activity o f the roots is c o n s i d e r e d to be an i n d e x o f the delay-
d r o g e n a s e activity which is i n v o l v e d in energy p r o d u c t i o n . It was suggested that the decrease
o f the T T C - r e d u c i n g activity caused a decrease in energy p r o d u c t i o n , f o l l o w e d by a decrease
in n u t r i e n t u p t a k e ( W a k i m o t o and Y a m a d a 1985). R e d u c e d p l a n t g r o w t h a s s o c i a t e d with
the decrease o f the r o o t T T C - r e d u c i n g activity was also o b s e r v e d in p e a n u t s ( T a k a h a s h i
1968).
Recently, F u j i i et al. (1991) r e p o r t e d that the t o m a t o p l a n t does not e x h i b i t an
a l l e l o p a t h i c effect on the g r o w t h o f c u c u m b e r plant. H o w e v e r , the c u l t i v a t i o n p e r i o d in t h e i r
e x p e r i m e n t was o n l y 3 0 d , a n d the dry weight o f the t o m a t o s h o o t s was less t h a n 4 g at the
end o f the experiment. It is p o s s i b l e that the c o n c e n t r a t i o n o f the p h y t o t o x i c substances in
the n u t r i e n t s o l u t i o n was t o o low to affect p l a n t growth. In the present e x p e r i m e n t s , p l a n t s
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were c u l t i v a t e d for 120 d or more, d u r i n g which toxic substances a c c u m u l a t e d in sufficient

amounts.
U p to now, several p h y t o t o x i c substances have been i s o l a t e d from p l a n t tissues and from
the rhizosphere, most o f t h e m being o r g a n i c acids a n d p h e n o l i c substances. T h e critical
c o n c e n t r a t i o n for p h y t o t o x i c i t y varies with the k i n d s o f i n h i b i t o r s , m e t h o d s for bioassay,
p l a n t species e x a m i n e d , r a n g i n g from m m o l L -1 to ,umol L -I (Rice 1984; T s u c h i y a 1990). In
the present experiment, the c a r b o n c o n c e n t r a t i o n o f the residual n u t r i e n t s o l u t i o n ranged
from 5 to 40 mg L -1. Since the s o l u t i o n m a y c o n t a i n c o n s i d e r a b l e a m o u n t s o f b i c a r b o n a t e s ,
sugars, a m i n o acids, etc., the c o n c e n t r a t i o n s o f p h y t o t o x i c substances m a y be mtlch l o w e r
than those o f c a r b o n . It is essential to i s o l a t e and identify the p h y t o t o x i c substances, w h i c h
are u n d e r investigation.

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