Autotoxicity in Vegetables and Ornamentals and Its Control: Toshiki Asao and Md. Asaduzzaman

4
Autotoxicity in Vegetables
and Ornamentals and Its Control
Toshiki Asao and Md. Asaduzzaman
Department of Agriculture,
Faculty of Life and Environmental Science,
Shimane University, Kamihonjo, Matsue, Shimane
Japan
1. Introduction
Allelopathy comes from the Latin words allelon ‘of each other’ and pathos ‘to suffer’ refers
to the chemical inhibition of one species by another. The ‘inhibitory’ chemical is released
into the environment where it affects the development and growth of neighboring plants.
Allelopathic chemicals can be present in any parts of an allelopathic plant. They can be
found in leaves, flowers, roots, fruits, or stems and also in the surrounding soil. Around 300
BC, the Greek botanist Theophrastus was possibly the first person to recognize the
allelopathic properties of plants when he observed and recorded that chickpea plants
exhausted the soil and destroyed weeds. Later, Pliny the Elder, a Roman scholar and
naturalist, noted that walnut trees were toxic to other plants, and that both chickpea and
barley ruined crop lands for maize. The term allelopathy was first introduced by a German
scientist Molisch in 1937 to include both harmful and beneficial biochemical interactions
between all types of plants including microorganisms. Rice (1984) reinforced this definition
in the first monograph on allelopathy. Research on the recognition and understanding of
allelopathy has been well documented over the past few decades (Rice, 1984; Rizvi & Rizvi,
1992). These include the symptoms and severity of adverse effects of living plants or their
residues upon growth of higher plants and crop yields, interactions among organisms,
ecological significance of allelopathy in plant communities, replanting problems, problems
with crop rotations, autotoxicity, and the production, isolation and identification of
allelochemicals in agro ecosystem.
Autotoxicity is a phenomenon of intraspecific allelopathy that occur when a plant species
releases chemical substances which inhibit or delay germination and growth of the same
plant species (Putnam, 1985; Singh et al., 1999). It been reported to occur in a number of crop
plants in agro ecosystem causing serious problems such as growth reduction, yield decline
and replant failures (Singh et al., 1999; Pramanik et al., 2000; Asao et al., 2003). Plants when
experiences autotoxicity it releases chemicals to its rhizosphere (Singh et al., 1999) through
various mechanisms such as leachation (Overland, 1966), volatilization (Petrova, 1977), root
exudation (Tang & Young, 1982), and crop residue decomposition (Rice, 1984). Autotoxicity
was found to be pronounced if the plants were cultivated consecutively for years on the
same land or grown by hydroponic culture without renewal of nutrient solution. One of the
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68 Hydroponics – A Standard Methodology for Plant Biological Researches
principal causes of this autotoxic growth inhibition in the successive culture of plants has
been attributed to the effect of exuded chemicals from plant roots. Root extracts and
exudations are the common sources of allelochemicals with potent biological activity and
are produced by numerous plant species, with great variation in chemical components
(Inderjit & Weston, 2003). It represents one of the largest direct inputs of plant chemicals
into the rhizosphere environment. The synthesis and exudation of allelochemicals, along
with increased overall production of root exudates, is typically enhanced by stress
conditions that the plant encounters such as extreme temperature, drought and UV
exposure (Inderjit & Weston, 2003; Pramanik et al., 2000). Accumulations of these
allelochemicals are immense in reused nutrient solution during hydroponic culture.
Vegetable and ornamental plants generally cultured through hydroponics in Japan and
other developed countries and recently closed type hydroponic systems gained popularity
for the production of these crops on a commercial basis. However, this managed and viable
technique has the autotoxicity constraint. Therefore, we have studied autotoxicity
phenomenon in several vegetables crops such as cucumber (Cucumis sativus), taro (Colocasia
esculenta), strawberry (Fragaria × ananassa Duch.), some leafy vegetables, and many
ornamentals at the glasshouse of Experimental Research Center for Biological Resources
Science, Shimane University, Matsue, Japan using hydroponic culture. We have also
investigated the isolation, identification, phytotoxicity evaluation of the allelochemicals and
means to recover growth inhibition. In this chapter we illustrate autotoxicity of the above
crops in hydroponics, its occurrence, autotoxic substances isolation and phytotoxicity
evaluation, and control methods.
2. Materials and methods

2.1 Plant materials
In our laboratory we have investigated the autotoxicity from root exudates of several
vegetable crops such as cucumber, taro, strawberry, some leafy vegetables, and many
ornamentals following hydroponic systems. Uniform seedlings or plantlets of similar
growth stage produced through seeds or tissue culture means were used as the test plant
materials.
2.2 Plant cultivation in hydroponics

Plant materials under investigation were planted into plastic containers (34 cm × 54 cm × 20
cm) and three containers were used for each treatment (plants with or without Activated
charcoal, AC). The containers were filled with ‘Enshi’ nutrient solution (Table 1) for each
crop (Hori, 1966). The nutrient solution concentration employed for each crop was 75% for
cucumber, taro and strawberry, and 50% for several ornamentals. Nutrient solution in the
container was continuously aerated (3.8 liter/min.) using air pumps with two small air
filters each packed with 100 g of AC (Type Y-4P, 4-8 mesh, Ajinomoto Fine Techno Co.,
Kawasaki, Japan). The same aeration system was maintained for the nutrient solution
without AC. The AC was used to trap the chemicals exuded from the plants and was
replaced by fresh AC at 2-week intervals until the end of the experiment for efficient
adsorption of the chemicals. The used AC was either immediately extracted with alkaline
methanol or stored at 4 ºC for later extraction. FeSO4.7H2O (0.75 g) was added to each
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Autotoxicity in Vegetables and Ornamentals and Its Control 69
Chemicals Concentration (µM/l)

Ca(NO3) 2・4H2O 4.03
KNO3 8.02
MgSO4・7H2O 2.03
NH4H2PO4 1.35
H3BO3 0.05
ZnSO4・7 H2O 7.64×10-4
MnSO4・5 H2O 8.30×10-3
CuSO4・5 H2O 2.00×10-4
Na2MoO4 9.71×10-5
NaFe-EDTA 0.06
Table 1. Mineral nutrient concentrations in full strength ‘Enshi` nutrient solution (Hori, 1966).
solution container at 2-day intervals since the AC that absorbed Fe-EDTA and Fe2+ was
rapidly oxidized to Fe3+ and less available for the plants (Yu et al., 1993). During cultivation,
the water level of the solution containers was kept constant by regularly adding tap water.
Nutrient concentrations (NO3-, PO42-, K+, Ca2+, Mg2+, and Fe3+) in the solution were adjusted
as close as possible to the initial concentration at 2-week intervals on the basis of chemical
analyses with an atomic absorption spectrometer (AA-630, Shimadzu Co., Kyoto, Japan), a
spectrophotometer (UVmini-1240, Shimadzu Co., Kyoto, Japan), and an ion meter (D-23,
Horiba, Kyoto, Japan). At the end of the experiment growth parameters, yield and yield
components were compared with untreated control.
2.3 Gas chromatography-mass spectrometry (GC-MS) analysis of root exudates

The AC used to trap the exudates (organics) were desorbed three-times using 200 ml 1:1
(v/v) methanol (100 ml):0.4 M aqueous NaOH (100 ml) (Pramanik et al., 2001). Each batch of
AC (200 g) was gently shaken with the mixture for 12 h at room temperature (25 ºC) with an
electric shaker (20 rpm). The three extracts (600 ml) were combined and filtered through
Whatman (No. 6) filter paper. The filtrates were neutralized with 6 M HCl and concentrated
to 25 ml in a rotary vacuum evaporator at 40 ºC. Organic compounds in the concentrate
were then extracted according to Yu & Matsui (1993). The concentrated AC-extract was
adjusted to pH 2.0 with 4 M HCl, extracted three times with 35 ml of refined diethyl ether
(DE), and a further three times with 35 ml of ethyl acetate (EA). DE2 and EA2 were the
pooled DE and EA extract fractions (105 ml), respectively at pH 2.0. DE2 and EA2 fractions
were dried over anhydrous CaSO4 and concentrated to 5 ml each in a rotary evaporator at 40
ºC. Both concentrated fractions (DE2 and EA2) extracted from the AC were analyzed using a
gas chromatograph coupled to a mass spectrometer (GC-MS, Hitachi M-80B, Hitachi, Tokyo,
Japan) before or after methylation with diazomethane from N-methyl-N-nitoso-p-toluene
sulfonamide. An aliquot of each concentrated fraction (1 or 2 ml) was diluted in 50 ml ether,
treated with diazomethane and concentrated to 5 µl in a rotary evaporator then in a N2
stream in a water bath at 35 ºC. One microliter of the concentrated sample was injected into
a GC-MS with a capillary column (0.25 mm × 60 m) of TC-5 (GL Science, Tokyo, Japan).
Helium was used as the carrier gas at a pressure of 78.4 kN/m2. The column was held
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initially at 100 ºC for 2 minutes and then raised at 5 ºC/min. to a final temperature of 260 ºC
for 10 minutes. The injector temperature was held at 270 ºC. The ionization voltage and
temperature in the electron impact (EI) mode were 70 eV and 250 ºC, respectively.
2.4 Bioassay with the identified chemicals

The bioassays with identified chemicals were carried out according to Asao et al., (1998b).
Inhibitions of the test solution were assayed by their effects on seedling growth of the source
plant species. Aqueous solutions of the identified compounds at several concentrations
between 0 (control) and 400 µM were prepared with nutrient solutions for each crops
studied. These test solutions were added to glass flasks (approx. 420 ml) wrapped in black
polythene to exclude light from the roots (Asao et al., 1999a). The selected plants were
transplanted into each flask with urethane foam as a support. The planted flasks were then
placed in a growth chamber at 25ºC with a light intensity of 74-81 µM/m2/s and required
photoperiod under fluorescence lights. To minimize the effects of aeration and microbial
degradation of the organic acids (Sundin & Waechter-Kristensen, 1994) on the bioassay, the
test solutions in the flasks were renewed every 3-4 d. The plantlets were grown for 3 weeks
then growth parameters were measured in terms of fresh weight (FW) and dry weight (DW)
of the shoots, DW of roots, and the longest root per plant.
2.5 Statistical analysis

A randomized complete block design with three replicates was used for growth chamber
bioassay and hydroponic culture in the greenhouse as described above. These experiments
were not repeated over time due to the consistent result. Multiple-comparison test were
performed by SPSS 11.0J for Windows (SPSS Japan Inc., Tokyo, Japan) as Tukey’s test at a
level of significance of P=0.05. Regression analyses were performed by Statcel2 (Add-in soft
for Microsoft Excel, OMS Publishing Inc., Saitama, Japan) at a level of significance of P=0.05.
3. Autotoxicity from the root exudates of vegetables and ornamentals in

hydroponics
Successive culture of the same crop on the same land for years cause soil sickness or
replanting injuries (Bonner & Galson, 1944; Davis, 1928; Hirano, 1940; Rice, 1984; Tsuchiya,
1990) resulting reduction in both crop yield and quality. This phenomenon is evidenced in
agricultural production especially in the production of horticultural crops (Grodzinsky,
1992; Tsuchiya, 1990; Young, 1984). It leads to resurgence of disease pest, exhaustion of soil
fertility, and developing chemical interference in the rhizosphere referring allelopathy
(Hegde & Miller, 1990; Komada, 1988; Takahashi, 1984; Young, 1984). Allelopathic effects
from crop residues and root exudates have extensively studied in vegetable crops such as in
alfalfa (Miller, 1983; Nakahisa et al., 1993, 1994; Chon et al., 2002, Chung et al., 2011),
asparagus (Young, 1984; Young & Chou, 1985; Hartung et al., 1990), cucumber (Yu &
Matsui, 1994, 1997), watermelon (Kushima et al., 1998; Hao et al., 2007), taro (Asao et al.,
2003), strawberry (Kitazawa et al., 2005), tomato (Yu & Matsui, 1993b), lettuce (Lee et al.,
2006) and so on. Therefore, growth of these plants found to be inhibited by the released
allelochemicals. So far several methods has been found to be effective in removing or
degradating the phytotoxic substances released from plant roots during autotoxicity such as
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adsorption by activated charcoal (Asao et al., 1998a), degradation by microbial strain (Asao
et al., 2004a) or auxin (2,4-D and NAA) supplementation (Kitazawa et al., 2007), electro-
degradation of root exudates (Asao et al., 2008) and TiO2 photocatalysis (Sunada et al., 2008).
Similar to successive culture, in closed hydroponics accumulation of phytotoxic chemical
leads to the occurrence of autotoxicity. In our lab we have investigated the autotoxic
potentials of several vegetables and ornamentals and suggested means to overcome it. Our
research history started with the selection of cucumber cultivars suitable for a closed
hydroponics system using bioassay with cucumber seedlings (Asao at al., 1998b).
Experiments were conducted to clarify why fruit yield decrease during the late growing
period of cucumber cultured in non-renewed hydroponic nutrient solution and results were
suggested that root exudates had induced the decrease in fruit yield, especially by affecting
young fruits, the decrease was reversible through removal of root exudates by AC (Asao et
al., 1998a). We found extended harvesting period in a closed nutrient flow system by
grafting 'Shogoin-aonaga-fushinari' on 'Hokushin' or 'Aodai' (Asao et al., 1999b) and
increased number of harvested fruits by adding AC in the nutrient solution (Asao et al.,
2000). Growth inhibiting substances of unknown origin found in the growing nutrient
solution of cucumber plants were isolated and identified. The growth inhibitors were
adsorbed on AC and extracted by organic solvent (Asao et al., 1999a). We developed a
bioassay technique to evaluate toxicity of aromatic acids to cucumber seedlings and to select
cucumber cultivars that release little or no 2,4-dichlorobenzoic acid (an autotoxic chemical
found in cucumber root exudates), thereby avoiding cucumber autotoxicity in the closed
hydroponics system (Asao et al., 1999a). Root exudates, which are detrimental to vegetative
growth and yield of cucumber plants, were adsorbed by the AC irrespective of dissolved
oxygen levels (Asao et al., 1999d).
The number of organic acids and their exudation rates were higher in high temperatures
and long photoperiods than that in low temperature with short photoperiod condition and
caused higher cucumber autotoxicity in the former conditions (Pramanik et al., 2000).
Species differences in the susceptibility to autotoxicity among leaf vegetables were also
investigated in hydroponics (Asao et al., 2001a). Autotoxicity of root exudates from taro
were showed benzoic acid as the potent growth inhibitor (Asao et al., 2003). A number of
aromatic organic acids were identified in several leaf vegetables (Asao et al., 2004b). 2,4-
dichlorobenzoic acid (DCBA)-degrading microbial strains, may degrade DCBA including
other growth inhibitors exuded from cucumber roots and avoid autotoxicity in cucumber
resulting increased fruit yield (Asao et al., 2004a). In strawberry we found vegetative and
reproductive growth inhibition due to autotoxicity developed in non-renewed nutrient
solution through accumulation of autotoxic root exudates, and the most potent inhibitor was
benzoic acid (Kitazawa et al., 2005). Foliar application of auxin such as 1-naphthaleneacetic
acid (NAA) avoided the growth reduction of strawberry caused by autotoxicity. NAA at 5.4
µM found to be the most effective for alleviating autotoxicity of strawberry and increasing
the yield (Kitazawa, et al., 2007). Autotoxicity in some ornamentals were investigated in
hydroponics with or without the addition of AC to the nutrient solution and several organic
compounds were detected (Asao et al., 2007). Benzoic acid being the strongest growth
inhibitor, its removal from the nutrient solution is imperative for sustainable production of
taro and strawberry or other crops exudates containing it. Therefore, electro-degradation
method have been tried to degrade benzoic acid and it was found to be recovered
strawberry yield up to 71% from non-renewed nutrient solution (Asao et al., 2008).
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3.1 Autotoxicity in cucumber (Cucumis sativus)

In closed hydroponic culture without renew of the nutrient solution, we found that the fruit
yield of cucumber plants decreased significantly in the late reproductive stage (2 weeks
ahead of final harvest) and the growth was recovered by the biweekly renewal of nutrients
or supplementation of AC to the nutrient solution (Asao et al., 1998a). Shrunken fruits were
harvested from the plant grown in non-renewed culture solution (Fig. 1). This inhibition has
been attributed due to the autotoxicity from root exudates (Yu et al., 1994). The autotoxicity
of cucumber also differs among cultivars (Asao et al., 1998b). Fruit harvesting of a
susceptible cucumber cultivar grown in a closed nutrient flow system was prolonged by
grafting onto a non-autotoxic cultivar (Asao et al., 1999b). Thus, cucumber root exudates
from a closed hydroponic system were analyzed and among a number of growth inhibitors
detected (Asao et al., 1999c; Pramanik et al., 2000), 2,4-dichlorobenzoic acid (DCBA) was the
strongest inhibitor. Microorganisms can degrade chemical substances in soil and water
(Markus et al., 1984; Nanbu, 1990; Sundin & Waechter-Kristensen, 1994). Van den Tweel et
al. (1987) reported that 2,4-dichlorobenzoate was degraded through reductive
dechlorination by microorganisms. Recently, we found that the inhibitory effect of DCBA on
cucumber seedlings could be reversed using strains of microorganisms (Asao et al., 2001b).
However, the effects of such strains on cucumber reproductive growth in the presence or
absence of DCBA have yet to be elucidated. Therefore, in this study we investigated the
effects of microbial strains on the autotoxicity of cucumber plants grown with or without
DCBA in the nutrient solution.
A B
Fig. 1. Fruits of cucumber cv. ‘Shogin-aonaga-fushinari’, white arrow indicating (A) a

normal developing fruit at ten days after anthesis; (B) shrunken fruit.
3.1.1 Cultivation of cucumber plants with or without microbial strain in presence of

DCBA
The DCBA-degrading microorganism (microbial strain) was isolated and screened from soil
in Aichi prefecture (Asao et al., 2001b). Nutrient solutions with DCBA (10 mg/l) and sucrose
(1 g/l) were prepared and sterilized by autoclave. A 200 ml volume of sterile nutrient
solution was inoculated with the DCBA-degrading microorganism and shaken continuously
by machine at 25 ºC for 9 days to have stock microbial suspension. Cucumber (Cucumis
sativus L. var. Shougoin-aonaga-fushinari) plants were grown in a greenhouse by
hydroponics at different concentrations of DCBA with or without addition of DCBA-
degrading microorganisms to the nutrient solution. One-week-old cucumber seedlings
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raised in vermiculite were transplanted into plastic containers containing 50 l of

continuously aerated (3.8 l/min) 75% Enshi nutrient solution having an electrical
conductivity (EC) of 2.0 dS/m. Three seedlings with four leaves were transplanted to each
container with three replications. The solutions were prepared at concentrations of 0
(control), 2 or 10 µM/l of DCBA with or without bacterial suspension in the nutrient
solution. The solutions were renewed biweekly. Three plants were planted in each container
with three repetitions. At the 15-leaf stage, the apical buds of cucumber plants were plucked
to maintain 15 leaves on the main stem. The terminal buds of all the developing primary
and secondary branches were removed keeping only one node in each branch. The mean air
and water temperature during the experiment ranged from 23.0 to 25.5 ºC and from 23.3 to
34.4 ºC, respectively. At the end of the experiment, data were recorded on plant growth,
dates of anthesis in male and female flowers, number of healthy female flowers, and
harvested fruit number.
3.1.2 Cucumber cultivation with microbial suspension in absence of DCBA

Similar cultivation procedure was followed without addition of DCBA to the nutrient
solution in another set of experiments. Three cucumber seedlings with four leaves were
transplanted to each container containing the nutrient solution without DCBA and three
containers were used for each treatment. During culture, the water level of containers was
kept constant by regularly adding tap water. Nutrient contents of the solutions were
adjusted to the initial concentrations following procedures described earlier. In all the
treatments, the EC and pH in the nutrient solution ranged from 1.4 to 2.8 dS/m and 6.4-7.9,
respectively. The microbial suspension was supplied to the nutrient solution added (a) at
planting, at the plucking of apical buds and at 2 weeks after initial harvest, (b) at the
plucking of apical buds and at 2 weeks after initial harvest, and (c) at 2 weeks after initial
harvest. No DCBA was added. An additional cultivation with biweekly renewal of the
nutrient solution in the absence of the microbial suspension or DCBA was set up to serve as
a control. At the 15-leaf stage, the apical buds of cucumber plants were removed to maintain
15 leaves on the main stem. The terminal buds of the branches were removed keeping only
one node on each branch. The mean air and water temperature during the experiment
ranged from 24.9 to 31.3 ºC and from 25.8 to 32.1 ºC, respectively. Data were recorded as
mentioned for the preceding experiment.
3.1.3 Effects of DCBA with or without microbial suspension on the growth and yield of
cucumber
Cucumber plants were grown in hydroponics using different concentrations of DCBA with
or without the addition of microorganisms to the nutrient solution. Results reveal that the
length of the main stem and primary branches decreased with the increase in DCBA
concentration (Table 2). The dry weight of stem, leaf, root and primary branches of plants
grown with DCBA (10 µM/l) was also decreased by about 60, 30, 26 and 32% of that without
DCBA, respectively. This growth inhibition was significantly recovered by the addition of
soil microorganisms to the nutrient solution. This indicates that the microbes efficiently
degraded the added DCBA in the nutrient solution and thus restored the inhibitory effect of
DCBA on the plants. The date of male flower anthesis was unaffected by the addition of
DCBA and the microbial suspension. However, the presence of DCBA at a concentration of
10 µM/l shifted the date of female flower anthesis and harvesting time by about 5 and 16
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z stem and leaf; ysignificant at the 5 % level (*), 1 % level(**), and not significant (NS) by T-test;
x significant at the 1 % level (**), and not significant (NS) by regression analysis of the concentrations.
Table 2. Effects of DCBA with or without microbial suspension on the growth and yield of
cucumber plants grown in hydroponics.
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days, respectively. Addition of the microbial suspension to the nutrient solution enhanced
early flowering and fruit setting in the cucumber plants treated with DCBA. The number of
healthy female flowers and the harvested fruit number per plant also decreased as the
DCBA concentration increased, and this decrease was significantly compensated by the
microbial suspension.
3.1.4 Effects of microbial suspension on cucumber growth in the absence of DCBA

The suspension of DCBA-degrading microorganisms was added once at 2 weeks after the
initial harvest, twice upon plucking the apical buds and at 2 weeks after the initial harvest,
and three times at the beginning of the culture, on plucking the apical buds and at 2 weeks
after the initial harvest. There was no significant difference in the growth of cucumber
except in the dry weight of roots and fruit number (Table 3). Root dry weight increased by
about 43% with the addition of the suspension of DCBA-degrading microorganisms once at
2 weeks after the initial harvest. The treatments did not affect the dates of anthesis in male
and female flower, the beginning of harvest, or the number of flowering female flowers per
plant. The harvested fruit number per plant was the lowest (14.2 per plant) in non-renewed
nutrient solution. The number of fruits recovered increased from 14.2 to 17.4 on addition of
the suspension to the nutrient solution once at 2 weeks after the initial harvest.
DCBA is one of the growth inhibitors found in cucumber root exudates (Pramanik et al.,
2000) and we found it as the most effective inhibitor of the growth of cucumber plants (Asao
et al., 1999c). In these experiments we also found that DCBA strongly retarded the growth of
cucumber plants (Table 2). However, this inhibition was significantly recovered by the
addition of DCBA-degrading microbes (Asao et al., 2001b) into the nutrient solution. This
result reveals that the microbial strain appreciably deactivated the inhibitory action of
DCBA including the other inhibitors in cucumber root exudates in the nutrient solution and
thus, the cucumber plant growth was enhanced. The recovery of growth, especially the dry
weight of roots and branches, in the plants grown with the microbial suspension, was about
three times higher than that of cucumber grown with DCBA alone at a concentration of 10
µM/l. Consequently, dates of male and female flower anthesis, and initial harvest were
several days earlier. The number of healthy female flowers as well as fruits also significantly
increased on the addition of the microbial suspension.
Experiments to clarify the influence of root exudates and microbes on cucumber plant
growth were conducted with or without biweekly renew of the nutrient solutions (Table 3).
Results revealed that the root growth and fruit number of the cucumber plants grown with
biweekly renewed nutrient solution were significantly increased than those grown without
renew of nutrient solution. The addition of the microbes to the nutrient solutions also
increased the growth of cucumber plants compared to the non-renew nutrient solution.
However, the microbial suspension added to the nutrient solutions in vegetative stage (at
the start of the culture or the plucking of apical buds) did not make significant yield
difference from non-renewed solution culture. Addition of DCBA-degrading microbial
suspension applied once at 2 weeks after the initial harvest was effective enough to recover
yield reduction of cucumber from autotoxicity.
DCBA causing autotoxicity in the cucumber was detected in their root exudates only in the
reproductive stage (Pramanik et al., 2000). Apparently it indicates that the growth inhibitors
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z(＋)= total renewal of the nutrient solution every other week; (−)= only supplement of the nutrient
solution which decreased during culture; yat the plucking of apical buds; xat two weeks after the initial
harvest; wstem and leaf; vdifferent letters within a column indicate significance at the 5 % level and not
significant (NS) by the Tukey test.
Table 3. Effects of addition of the microbial suspension at different growth phases on the
growth and yield of cucumber plants grown in the absence of DCBA.
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including DCBA would have sufficiently accumulated in the nutrient solution through
cucumber root exudation at the reproductive stage. The degrader (applied two or three
times) probably became a source of nutrients for other microorganisms. In this case, the
degrader did not dominated more than other microorganisms. However, when supplied
once at 2 weeks after the start of harvest, the degrader did not become a nutrient source for
other microorganisms and degraded the DCBA exuded from cucumber. This was why the
DCBA exuded from cucumber sustained the microbial activity. In conclusion, DCBA-
degrading microorganisms, if added to the nutrient solution, may degrade DCBA including
other growth inhibitors exuded from cucumber roots and avoid autotoxicity in cucumber
resulting increase the fruit yield. Addition of the microbial suspension in the reproductive
stage of cucumber plants appears to degrade the growth inhibitors efficiently. However, the
timing of degrader addition to the nutrient solution for efficient mitigation of cucumber
autotoxicity needs further study.
3.2 Autotoxicity in strawberry (Fragaria ananassa Duch.)

Closed hydroponics is a system used for plant cultivation in environmentally sensitive areas
(Van Os, 1995) where the nutrient solution is not released into the surrounding environment,
but recycled (Ruijs, 1994). However, in a closed hydroponic system, plants can suffer
autotoxicity, due to the accumulation of toxic exudates from the roots themselves in the
nutrient solution (Yu et al., 1993). Recently, closed hydroponics has been considered for
strawberry cultivation (Takeuchi, 2000; Oka, 2002; Koshikawa & Yasuda, 2003). However, it
was reported that a yield reduction, caused by unknown factors, occurred in closed
hydroponic system for strawberry (Oka, 2002). Koda et al., (1977, 1980) reported that a growth
reduction in mitsuba (Cryptotaenia japonica Hassk.) in hydroponic system was caused by root
exudates such as organic acids. Some aromatic acids also accumulated in the nutrient solution
during hydroponic cultivation of tomato and had inhibitory activity on growth (Yu & Matsui,
1993). Asao et al., (1998a, 1999c) demonstrated that a reduction in fruit yield in the late
reproductive stage of cucumber was induced by root exudates and that the most potent
inhibitor was 2,4-dichlorobenzoic acid. In a closed hydroponic culture of rose plantlets, root
and shoot growth were reduced by root exudates (Sato, 2004). However, little is known about
similar effects in strawberry culture. It was therefore felt necessary to examine the effects of
root exudates on the growth of strawberry. Root exudates can be removed by adding activated
charcoal (+AC) to the nutrient solution (Koda et al., 1977; Asao et al., 1998a, 1999c; Sato, 2004).
In this study, we investigated the effects of non-renewal of the nutrient solution, and of adding
AC on vegetative and reproductive growth of strawberry.
3.2.1 Cultivation of strawberry plants in hydroponics

Strawberry (Fragaria ananassa Duch. cv. ‘Toyonoka’) was used for these experiments
following the cultivation method explain above (Section 2.2 & Fig. 2). Pollination was aided
by vibrating the anthers over stigma with a soft brush at 2 d intervals. The fruits were
collected when ripe. At harvest growth parameters were measured among the treatments.
During cultivation, the number of flower clusters, flowers and fruits per plant were
recorded. The allelochemicals in the root exudates of strawberry were identified by GC-MS
(section 2.3) and their phytotoxicity were evaluated in growth chamber bioassay (section
2.4) using strawberry plantlets.
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Fig. 2. Hydroponic system used for strawberry cultivation at the greenhouse of Shimane
University, Matsue, Japan.
3.2.2 Effects of non-renewed nutrient solution on the growth and yield of strawberry
In the treatment '-AC', the number of leaves, FW of shoots, DW of shoots and roots per plant,
and the root length decreased by 75%, 59%, 50%, 81% and 45% of control values,
respectively. In the '+AC' treatment, each value was 103%, 83%, 75%, 102% and 98% of
control values (Table 4). Although the number of flower clusters per plant was not
significant by different between treatments, values in '-AC' and '+AC' treatment were 85%
and 89% of control values, respectively (Table 4). In the '-AC' treatment, the number of
flowers per plant decreased significantly to about 74% of control value. In the '+AC'
treatment, each value was approx. 102% of the control value and there was no significant
difference between the '+AC' treatment and control. In the '-AC' treatment, the number of
harvested fruit per plant decreased significantly by approx. 49% of control values. In the
'+AC' treatment, this value was about 107% of the control value, and there was no
Nutrient AC No. of FW of DW of DW of Root No. of No. of No. of

solutionz supplement leaves shoots/ shoots/ root/ length flower flower/ fruits/
/plant plant (g) plant (g) plant (g) (cm) cluster/ plant plant
plant
＋－AC 23.2ay 71.8a 15.5a 4.7ab 43.8a 4.6ab 21.0a 12.3a
－－AC 17.5b 42.6b 7.8b 3.8b 19.8b 3.9a 15.6b 6.0b
－＋AC 24.0a 59.8a 11.6a 4.8a 42.8a 4.1a 21.4a 13.1a
z(+)= complete renewal of the nutrient solution every second week, and (-)= only supplement of the
nutrient solution which decreased during culture; yvalues in column followed by a different letter differ
significantly by Tukey's test (P = 0.05; n = 9).
Table 4. Effects of non-renewed nutrient solution and activated charcoal on the growth,
yield components and yield of strawberry in hydroponics.
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significant difference between the '+AC' treatment and control (Table 4). In the ‘+AC'
treatment with non-renewed nutrient solution, the growth and yield of strawberry plants
were eventually equivalent to control plants.
3.2.3 Phytotoxicity of the identified chemicals in root exudates of strawberry

GC-MS analysis of strawberry root exudates (Fraction 'DE2') showed more than 20 peaks,
whereas the 'EA2' fraction had only a few detectable peaks (Fig. 3). Based on a comparison
of retention times and the mass spectra of standard samples, five peaks were identified as
methyl esters of lactic acid, benzoic acid, succinic acid, adipic acid and p-hydroxybenzoic
acid. The autotoxic effects by the five exudates compounds identified were evaluated by
using micro-propagated the plantlet of the same species from which they originated. Lactic,
succinic, adipic and p-hydroxybenzoic acid did not significantly reduce the FWs of shoot, or
the DWs of shoots and roots (Table 5). No correlation was found between these growth
parameters and increasing concentrations of these four exuded compounds. Benzoic acid,
however, significantly reduced both FW and DW of shoots, even at 50 μM and, these growth
parameters decreased further with increasing concentration. Benzoic acid also significantly
reduced the DWs of roots at all concentrations. Root length was reduced by increasing
concentrations of all five compounds, although lactic, succinic and adipic acids reduced root
length significantly at concentrations of 100 μM or above, while benzoic and p-
hydroxybenzoic acids significantly reduced root length even at 50 μM (Table 5).
The effects of non-renewed hydroponic nutrient solution and of adding AC ('+AC') on the
vegetative and reproductive growth of strawberry were investigated. Non-renewed nutrient
solution resulted in a significant decrease in the growth of strawberry plantlets compared to
growth when the nutrient solution was renewed. The number of flower clusters, flowers and
fruit harvested all decreased in non-renewed nutrient solution (Table 4). Growth and/or yield
reductions in plants caused by non-renewed nutrient solution have been reported by many
researchers (Koda et al., 1980; Yu & Matsui, 1993). Asao et al., (1998a) reported that fruit yields
in cucumber plants decreased significantly at the late reproductive stage (2 weeks before final
harvest) when nutrient solution was not renewed and that the reduction was caused by root
exudates. In a closed hydroponic system for rose plantlets, the root and shoot growth were
reduced by the root exudates (Sato, 2004). Thus, it was thought that vegetative and
reproductive growth in strawberry was inhibited by root exudates during non-renewal of the
nutrient solution. Growth and/or yield inhibition in cucumber (Asao et al., 1998a), mitsuba
(Koda et al., 1980) and rose (Sato, 2004), caused their root exudates could be avoided by
adding AC to the nutrient solution. AC added to the nutrient solution adsorbs the organic
compounds exuded from plant roots and thereby removes the inhibitory effects of exudates.
In this study, there was no significant difference in vegetative and reproductive growth of
strawberry between addition of AC to non-renewed nutrient solution, and renewed the
nutrient solution (Tables 4). Thus, our results suggest that the chemicals exuded from
strawberry roots inhibit vegetative and reproductive growth, which can be avoided by
adsorption of root exudates using AC. The substances adsorbed on the AC were extracted,
analyzed and some identified as phenolic and aliphatic acids (Fig. 3). Growth and/or yield
reduction in cucumber plant by root exudates had been confirmed to be due to (Pramanik et
al., 2001) and alipathic acids (Yu & Matsui, 1997). In cucumber, 2,4-dichlorobenzoic acid was
the most potent inhibitor of growth and yield (Asao et al., 1999c).
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Compoundz (Concentration, FW of shoot/ DW of shoot/ DW of Root length/

μM) plant (g) plant (g) root/ plant plant (cm)
(g)
Control (No
0 0.98ab 0.17a 0.07a 17.0a
compound)y
Lactic acid 50 0.95a 0.16a 0.06a 16.2a
100 0.88a 0.15a 0.06a 13.7b
200 0.88a 0.16a 0.07a 11.9bc
400 0.85a 0.15a 0.06a 11.4c
Benzoic acid 50 0.48b 0.09b 0.03b 12.1b
100 0.42b 0.09b 0.03b 10.5b
200 0.42b 0.08b 0.03b 10.3b
400 0.36b 0.08b 0.03b 9.4c
Succinic acid 50 0.80a 0.14a 0.06a 16.2ab
100 0.74a 0.13a 0.04a 13.6bc
200 0.69a 0.19a 0.04a 12.5c
400 0.87a 0.16a 0.05a 11.7c
Adipic acid 50 0.70a 0.12a 0.03a 14.1ab
100 0.88a 0.15a 0.05a 12.0bc
200 0.74a 0.13a 0.05a 10.6c
400 0.76a 0.15a 0.05a 9.4c
p-
hydroxybenzoic 50 0.77a 0.14a 0.04a 11.9b
acid
100 0.67a 0.13a 0.03a 10.8bc
200 0.77a 0.15a 0.05a 9.1c
400 0.65a 0.12a 0.04a 10.4bc
zcompounds identified in the root exudates adsorbed onto the activated charcoal; yvalues in column
followed by a different letter differ significantly by Tukey's test (P = 0.05; n = 9).

Table 5. Effects of purified exudate compounds at different concentrations on the growth of
strawberry plantlets.
Benzoic acid was also the main inhibitor of growth and yield in taro plants (Asao et al.,
2003). The phenolic and aliphatic acids identified in strawberry root exudates may have the
potential to inhibit growth. The inhibitory potential of root exudates has been tested using
germination test of lettuce seeds (Tsuchiya & Ohno, 1992). However, Asao et al. (2001a)
suggested that lettuce plants suffer autotoxicity from their own root exudates. Vanillic acid
was the most inhibitor of the growth of lettuce (Asao et al., 2004b). Thus, the potential of
each compound identified in our study was evaluated using plantlets of the species which
they originated. Five compounds including phenolic and aliphatic acid were found to affect
plant growth but only benzoic acid significantly inhibited increases in the FW of shoots,
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Fig. 3. Gas Chromatograms of all components from root exudate of strawberry plants
adsorbed and released from AC. Methyl esters of lactic acid (peak 1), benzoic acid (peak 2),
succinic acid (peak 3), adipic acid (peak 4) and p-hydroxy benzoic acid (peak 5) are
identified based on known standard retention times.
DWs of shoots and roots, and root length at all concentrations (Table 5). Therefore, it was
thought that benzoic acid was the strongest inhibitor of vegetative and reproductive growth
in strawberry. In conclusion, inhibition of the vegetative and reproductive growth in
strawberry caused by non-renewed nutrient solution may occur through autotoxic root
exudates, and the most potent inhibitor was benzoic acid. Reduction in the yields of
strawberry grown closed hydroponic systems would, therefore, appear to be related to the
allelochemicals exuded by the strawberry plant itself. Finally, for strawberry cultivation in a
closed hydroponic system, AC should be added to the nutrient solution to relieve the
autotoxicity caused by root exudates.
3.3 Autotoxicity in taro (Colocasia esculenta Schott.)

Taro plants do not grow well if cultivated consecutively for years on the same land (Takahashi,
1984). Rotation with other crops for at least three years (Miyoshi et al., 1971a), in combination
with organic matter and soil disinfectants (Murota et al. 1984), has been suggested to improve
the yield of taro. However, even in a fixed crop rotation system, there was a great difference in
the growth and yields of taro plants. This depends upon the kinds of crops in rotation, and the
order in which they were rotated. In combination with burdock, the yield of taro was equal to
or more than that of taro in the first-year planting, and the extent of corm injury were slight
(Murota et al., 1984). Harmful microbes (Atumi, 1956, 1957; Atumi & Nakamura, 1959; Nagae
et al., 1971) and nematodes (Miyoshi et al., 1971b; Oashi, 1973; Matsumoto et al., 1973, 1974) in
the soil are the main causes of damage in the successive culture of taro. Takahashi (1984)
suggested that unknown factors were also involved whereas; Miyaji et al. (1979) found that
taro residues in soils after harvest were inhibitory to its growth. Methanol extracts of taro
residues alone or of soils with taro residues were found to strongly inhibit the elongation of
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hypocotyls and radicle growth of turnip. The foregoing results reveal that growth inhibitors
from taro were connected with replanting problems. Our laboratory has established used
hydroponic culture system to assess autotoxicity in crop plants (Asao et al., 1999c; Pramanik et
al., 2000). Thus, an attempt was made to identify the chemicals exuded by taro roots and to
evaluate the allelopathic effects of these exudates on the growth and yield of taro through this
established hydroponic culture system.
3.3.1 Cultivation of taro plants in hydroponics

Taro cv. Aichi-Wase was used for this experiment. Corms were planted in plastic tray (32
cm × 47 cm × 7 cm) containing vermiculite in the green house. At the third leaf stage, taro
plantlets were transplanted into plastic containers following the cultivation method
described above in section 2.2 (Fig. 4). At the end of this experiment, measurements were
made on the longest leaf stalk, maximum leaf length and width, leaf number per plant,
shoot dry weight and corm yield. Phytotoxic chemicals adsorbed in the ACs were extracted
and identified by a Gas-chromatograph coupled with mass-spectrometer as mentioned
earlier. Bioassay of the identified acids at concentrations of 0 (control) or 400 μM/l were
prepared with a 75 % Enshi nutrient solution (EC 2.0 dS/m) in growth chamber condition.
The taro plantlets were grown for 26 days and then the fresh and dry weights of shoots,
number of leaves, longest root length and root dry weight were measured. Each treatment
was replicated 15 times. We carried out further bioassays following the same procedure
with benzoic and adipic acids at concentrations of 0 (control), 25, 50, 100 200 and 400 μM/l.
In this case, the taro plantlets were grown for 20 days.
A B
Fig. 4. Hydroponic system used for taro cultivation (A) taro plants in plastic containers, (B)
sketch of taro hydroponics showing different components.
3.3.2 Effects of non-renewed nutrient solution on the growth and yield of taro
Results revealed that plants grown without AC had experienced significant shoot growth
retardation compared to those grown with AC. The leaf numbers and shoot dry weights of
the plants grown without AC decreased to about 90% and 67% of those grown with AC,
respectively (Table 6). Addition of AC to the nutrient solution also improved yield
significantly. The total yield per plant without AC decreased to about 34% compared to that
on the addition of AC (Table 6). Larger corms were harvested from the nutrient solution
with AC.
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Charcoal No. of DW of Total yield/ Yield/plant by corm sizez (g)

supplement leaf/plant shoot/plant (g) plant (g) 2L L M S 2S 3S
− 14.6 10.9 429 72 38 123 74 86 109
+ 16.1 16.1 649 154 80 225 97 111 103
* y * ** x ** ** ** NS NS NS
z2L (>60 g), L (35-59 g), M (20-34 g), S (15-19 g); 2S (10-14 g) and 3S (<10 g); ysignificant at the 5 % level
(*); xsignificant at the 1 % level (**), and not significant (NS) by T-test.
Table 6. Influence of nutrient solutions in the absence and presence of activated charcoal on
the vegetative growth, yield and yield components of taro plants grown by hydroponic
culture.
3.3.3 Phytotoxins in root exudates of taro and their phytotoxicity

Analysis of the extracted taro root exudates with GC-MS gave more than thirty peaks (Fig. 5).
Based on the comparison of retention times and mass spectra with those of authentic
samples, seven peaks were assigned as methyl esters of lactic acid, benzoic acid, m-
hydroxybenzoic acid, p-hydroxybenzoic acid, vanillic acid, succinic acid, and adipic acid.
Fig. 5. Gas chromatograms of all components from root exudate of strawberry plants
adsorbed and released from AC. Methyl esters of lactic acid (peak 1), benzoic acid (peak 2),
succinic acid (peak 3), adipic acid (peak 4), m-hydroxybenzoic acid (peak 5), p-
hydroxybenzoic acid (peak 6) and vanillic acid (peak 7) are identified based on known
standard retention times.
the test material. Benzoic acid at 400 M/l induced severe growth inhibition of shoots and
The allelopathic potential of the identified compounds was evaluated using taro plantlets as
roots, while adipic acid at the same concentration reduced only dry weight of roots
acid at concentrations ranged from 0 to 400 M/l using taro plantlets. Both acids
(Table 7). Thus, we further evaluated growth inhibition potential of benzoic acid and adipic
retardation even at 50 M/l and growth decreased with increasing concentration of the
significantly inhibited the growth of plantlets (Table 8). Benzoic acid induced growth
acid. Benzoic acid at the highest concentration of 400 μM/l reduced fresh weight, shoot dry
weight, root length and root dry weight in taro plants to 54 %, 53 %, 54 %, and 75 % of
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Allelochemicalz FW of shoot/ DW of No. of Root length DW root/plant

plant (g) shoot/plant (g) leaf/plant (cm) (g)
None(control) 26.58ay 2.05a 4.3 17.0a 0.52a
Lactic acid 35.40a 2.11a 4.1 18.8a 0.51a
Benzoic acid 20.79b 1.30b 3.6 14.5b 0.29b
m-hydroxybenzoic
27.23a 1.80a 4.0 16.5a 0.38a
acid
p-hydroxybenzoic
32.87a 2.56a 4.2 16.7a 0.54a
acid
Vanillic acid 27.37a 1.84a 3.5 17.7a 0.45a
Succinic acid 27.17a 1.84a 3.9 19.5a 0.36a
Adipic acid 23.54a 1.61a 4.3 16.7a 0.27b
NS
z400 μM/l; ydifferent letters within a column indicate significance at the 5 % level and not significant
(NS) by the Tukey test.

Table 7. Inhibition potentials of chemicals identified in taro root exudates (400 μM/l) on the
growth of taro plantlets.
Allelochemical Concentrations FW of shoot/ DW of No. of Root length DW

(μM/l) plant (g) shoot/plant leaf/plant (cm) root/plant
(g) (g)
None(control) 0 3.76az 0.235a 4.0 16.0a 24a
Benzoic acid 25 3.46a 0.200a 4.6 13.5a 24a
50 3.14b 0.167b 4.2 14.2a 19a
100 2.81b 0.159b 4.0 11.8a 20a
200 2.16c 0.131c 3.0 10.5b 18b
400 2.03c 0.125c 4.0 8.6c 18b
Significance
Linear *y * NS * *
Quadratic * * NS * *
Adipic acid 25 3.91a 0.251a 4.2 16.6a 31a
50 3.35a 0.020a 4.0 14.6a 23a
100 3.43a 0.021a 3.7 12.3a 23a
200 3.33a 0.195a 4.3 12.0a 27a
400 3.18b 0.191a 4.3 10.8b 26a
Significance
Linear * NS NS * NS
Quadratic * NS NS * NS
zDifferent letters within a column indicate significant at the 5 % level and not significant (NS) by the
Tukey test; ysignificant at the 5 % level (*) and non-significant (NS) by regression analysis of the
concentrations.
Table 8. Effects of benzoic acid and adipic acid at different concentrations on the growth of
taro plantlets.
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control values, respectively. Adipic acid only at 400 μM/l reduced fresh weight of shoots
and root length. Lower concentrations of this acid did not affect shoot or root growth.
As the nutrient concentrations and growth environment in the hydroponic cultures of taro
plants were apparently identical, the significant growth differences between the plants
grown with and without AC could be attributed to the variation in the chemical
composition of the nutrient solution. These chemicals would have exuded from taro roots.
Tsuchiya and Ohno (1992) indicated that water extracts from soils used consecutively for
taro cultivation over a period of years inhibited the growth of lettuce. Since the same
phenomenon was observed even when the extracts were autoclaved, it was considered that
the inhibition was caused by allelochemicals rather than by harmful soil
microorganisms. There have been many reports that taro residues exhibited an allelopathic
effect on plant growth (Miyaji et al., 1979; Tsuzuki et al., 1995; Pardales & Dingal, 1988). It
was made clear here that the vegetative growth and corm yield of taro plants were
decreased in the non-renewed culture solution and the loss was recovered by adding AC to
the nutrient solution. This result suggests that the chemicals exuded from taro roots had
induced the inhibition of growth and reduced yield. This inhibition was prevented by the
adsorption of the exuded allelochemicals in AC.
The substances adsorbed on the AC were extracted, analyzed and some of them identified
as phenolic and aliphatic acids although many compounds in the root exudates are yet to be
detected. The allelopathic potential of each identified compound was evaluated and found
induced significant growth inhibition in taro plantlets even at concentration of 50 M/l.

that almost all the compounds inhibited the growth of taro plantlets (Table 7 & 8). Benzoic
Inhibitory effects of phenolic acids (Pramanik et al., 2001) and aliphatic acids (Yu & Matsui,
reduction of absolute leaf expansion brought about at 0.23 M of phenolic acid per gram
1997) to plant growth have been well recognized. In a bioassay, Blum (1996) found that 30 %
soil, while it required only 0.05 M in the presence of 0.06, 0.17, and 0.04 M of p-coumaric,
p- hydroxybenzoic, and vannilic acids per gram of soil, respectively. Thus, mixture of
allelochemicals can below their inhibitory levels. This indicates that taro plants exude a
number of compounds (Fig. 5) into its surroundings and those inhibit the growth taro plants
by synergistic or additive actions. In conclusion, taro roots exude a number of
allelochemicals including aromatic acids such as benzoic acid and aliphatic acids such as
adipic acid which inhibit the growth of taro plants by additive or synergistic actions.
Benzoic acid induced strongest inhibition. Thus, the decline in yield on the successive
culture of taro would appear to be related to the allelochemicals exuded from the taro plant
itself.
3.4 Autotoxicity in some ornamental plants

Plants synthesize, store, and exude various kinds of organic compounds in their
surroundings as exudates, volatiles, or residues of decomposition (Hale & Orcutt, 1987).
Some of the released compounds (allelochemicals) inhibit the growth of the source plants
(autotoxicity) or the other species grown in the vicinity of source plants (heterotoxicity). This
autotoxicity or heterotoxicity can be treated as allelopathy and the autotoxicity was found to
be increased if the plants were cultivated consecutively for years on the same land (Rice,
1984) or grown by hydroponic culture without renew of nutrient solution (Asao et al., 1998a,
2001a). One of the principal causes of this growth inhibition in the successive culture of
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plants has been attributed to the effect of exuded chemicals from plants (Pramanik et al.,
2000). Growth of some vegetables such as asparagus, taro, cucumber, and tomato was
inhibited by allelochemicals found in their root exudates (Asao et al., 1998a, 2003, 2004b;
Shafer & Garrison, 1986; Yang, 1982; Yu & Matsui, 1993). Inhibition in growth of apple,
peach, rice, strawberry, and sugarcane has been documented for the autotoxicity (Kitazawa
et al., 2005; Mizutani et al., 1988; Rice, 1984). This autotoxicity in tomato (Yu et al., 1993) and
cucumber (Asao et al., 1998a; Pramanik et al., 2000) has been recovered by addition of AC to
the nutrient solution, because the added AC adsorbed the phytotoxic root exudates and thus
favored plant growth. However, research on autotoxicity in ornamentals is limited. Tukey
(1969) showed that when chrysanthemum was grown repeatedly in the same place for
several years, growth was reduced owing to accumulation of toxic substances in the soil.
Kaul (2000) reported on autotoxicity in African marigold, but did not identify the
allelochemicals involved. Therefore, we attempted to investigate autotoxicity, if any, in
selected ornamentals along with a possible remedial measure to overcome the growth
inhibition from autotoxicity.
3.4.1 Cultivation of ornamental plants in hydroponics

Thirty-seven different ornamentals belonging to 16 different families were chosen for this
experiment (Table 9). Plant cultivation was carried out according to Pramanik et al. (2000) as
described above. Seedlings, scions, germinated bulbs, and corms of the plants under study
were transplanted to plastic containers (34 cm × 54 cm × 20 cm) in the greenhouse. At the
end of the experiment, plant length, number of leaves per plant, maximum root length, flesh
and dry weight of shoot and dry weight of root, and number of flowers per plant were
recorded.
3.4.2 Bioassay of the identified autotoxic chemicals in nutrient solution

Gas chromatography-mass spectroscopy analysis of root exudates adsorbed in activated
charcoal identified the responsible autotoxic chemicals. Bioassay of aqueous solutions of the
identified compounds was carried out according to Asao et al. (1998b) at concentrations of 0
(control), 50, 100, 200, and 400 µM with 50% Enshi nutrient solution (EC 1.3 dS/m) having
ten replications. The plants were grown for 2 weeks and then the fresh and dry weights of
shoots were measured.
3.4.3 Bioassay in soils amended with activated charcoal

Bioassay in soils amended with activated charcoal has also been carried out. Soils were
collected from a field successively cultivated with prairie gentian [Eustoma grandiflorum (Raf.)
Shinn.] for over 10 years in Nagano prefecture, Japan, and was used as medium of growth
for the bioassay. Three kilograms of the soil was pulverized and placed in each plastic
container (17 cm × 29 cm × 9.5 cm) after amending with AC corresponding to the rate of 0
(control), 30, 60, 120, 240, and 480 kg/10a. Soil collected outside the prairie gentian field was
also used as a reference to compare the growth performance of the test plants growth with
or without AC (control). The physical and chemical properties of the reference soil were
essentially similar to the soil in the prairie gentian field (data not shown). Ten prairie
gentian seedlings were planted into the treated containers. Irrigation (500 ml water) was
applied to each container at 2-week intervals and 500 ml Enshi nutrient solution (50%) with
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Family Ornamental Scientific name Cultivar

Compositae Pot marigold Callendula officinalis L. ‘Gold-star’
Cornflower Centaurea cyanus L. ‘Echo-sultan’
Chrysanthemum Chrysanthemum morifolium Ramat. ‘Shuhou-no-chikara’
Cosmos Cosmos bipinnatus Cav. ‘Dearboro’
Zinnia Zinnia elegans Jacq. ‘Sunbow-orange’
Thistle Cirsium japanicum DC. ‘Rakuonzi-Azami’
Sunflower Helianthus annuns L. ‘Big-smile’
Safflower Carthamus tinctorius L. −z
African marigold Tagetes erecta L. ‘Orange-isis’
China aster Callistephus chinensis Nees ‘Kurenai’
Coneflower Rudbeckia hirta L. ‘Gloriosa-daisy’
Liliaceae Tulip Tulipa gesneriana L. ‘Blue-champion’
Thunberg lily Lilium × elegans Thunb. ‘Iberu-flora’
Toritelia Tritelelia laxa Benth ‘Bridgesii’
Lily Lilium × formolongi Hort. ‘Hananomai’
Labiatae Rocket larkspur Delphinium ajacis L. ‘Lilac’
Love-in-a-mist Nigella damascena L. ‘Transformer’
Scarlet sage Salvia splendens Ker. ‘Lavender’
Fan columbine Aquilegia flabellate Sieb. et Zucc. ‘Macana-giant’
Caryophyllaceae Corn cockl Agrostemma githago L. ‘Purple queen’
Gypsophilla Gypsophila elegans M.B ‘Covent-garden’
Carnation Dianthus caryophyllus L. ‘Feminist’
Leguminosae Sweet pea Lathyrus odoratus L. ‘Rolay-lavender’
Lupine Lupine luteus L. ‘Lassell’
Cruciferae Rape blossoms Brassica rapa L. ‘Wase-fushimi-kanzaki’
Stock Matthiola incana R.Br. ‘Love-me rose’
Onagraceae Farewell-to-spring Godetia amoena G.Don ‘Kyokuhai’
Umbelliferae Bishop's weed Ammi majus L. −z
Scrophulariaceae Snapdragon Antirrhinum majus L. ‘F1-butterfly-bronze’
Papaveraceae Corn poppy Papaver rhoeas L. ‘Red-sales’
Amaryllidaceae Narcissus Narcissus tazetta L. ‘Fernandesii’
Amaranthaceae Feather cockscomb Celosia argentea L. ‘Red-cupid’
Globe amaranth Gomphrena globosa L. ‘Strawberryfields’
Gentianaceae Prairie gentian Eustoma grandiflorum (Raf.) Shinn. ‘Blue-line I’
Campanulaceae Balloon flower Platycodon grandiflorum A. DC. ‘Samidare-murasaki’
Plumbaginaceae Statice Limonium sinuatum Mill. ‘Marine-blue’
Solanaceae Chinese-lantern plant Physalis alkekengi L. var. franchetii ‘Tanba houzuki’ .
z Unknown
Table 9. Planting materials used for investigating autotoxicity from their root exudates in
hydroponics.
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EC of 1.3 dS/m was applied to each container at 2-week intervals. The cultivation was
continued for 8 weeks. At the end of the experiment plant length, number of leaves per
plant, maximum root length, shoot dry weight and root dry weight, and number of flowers
per plant were recorded.
3.4.4 Growth performances of the ornamental plants grown in hydroponics

Thirty-seven ornamentals were grown through hydroponic culture with or without addition
of AC in the nutrient solution. Plant growth was significantly affected by the added AC.
Performances of the plants were evaluated as percent comparing the growth of the plants
grown without AC (control) with those grown with AC. Different plants responded
differently to the addition of AC (Table 10). Growth in lily was the most severely retarded.
Plant length, number of leaves and flowers per plant, root length, and plant dry weight
almost all declined significantly in most of the plants grown without AC compared with
those grown with AC. However, root growth was found to be more responsive to AC than
the other studied parameters possibly for being the roots in direct contract with the exuded
chemicals (Pramanik et al., 2000). Root dry weight of lily and rocket larkspur was reduced to
approx. 85% and 74%, respectively, followed by prairie gentian with growth reduced to
55%. Root length of lily was reduced to approx. 58%, whereas that in prairie-gentian was
reduced to approx. 49%. It appears that lily, prairie gentian, corn poppy, pot marigold,
toritelia, and farewell-to-spring were the most sensitive to autotoxicity. Autotoxicity in
plants from their own exuded chemicals is also observed in natural ecosystems (Rice, 1984)
and was well documented in many crops (Asao et al., 1998a; Kitazawa et al., 2005; Mizutani
et al., 1988; Pramanik et al., 2000; Yu et al., 1993). Asao et al. (2001a) detected autotoxicity in
some species of Umbellifeae, Compositiae, and Cruciferae. So, autotoxicity in the
ornamentals might be incited by the exuded chemicals from their roots. Stimulated growth
was observed in the plants such as African marigold, love-in-a-mist, and rape blossoms
grown in non-renewed nutrient solution. The exact reasons for this growth stimulation in
the latter plants were not discovered. It is well known that a chemical at low concentration
acts as a growth stimulant to a plant and the same chemical at high concentration becomes
toxic or growth-retardant to the same plant (Rizvi & Rizvi, 1992). Functional activity of an
allelochemical depends on its concentration and time exposure to the test plants. So, it is
possible that the quality and quantity of root exudates in the nutrient solution in absence of
AC might not be sufficient to inhibit growth in the latter ornamental plants, but rather their
growth was stimulated.
3.4.5 Growth and yield of prairie gentian plants when grown its replant soil
Performances of prairie gentian were very poor when successively grown for years in the
same land. Significant growth inhibition was noticed in the plants grown in soils from a
prairie gentian field without AC compared with those grown in reference soil (Table 11). It
suggests that soil from a prairie gentian field has some growth inhibitors. In hydroponic
culture, we also detected some growth inhibitors in the root exudates of the test plant
(Tables 12 and 13). Those inhibitors should have been adsorbed when the soil was amended
with AC. Thus, the growth of the test plants was increased with an increase in amount of
AC from 30 to 60 kg/10a followed by a gradual decline at the highest dose of AC (480
kg/10a). This high dose of AC might have affected other chemical properties in soil. Results
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Family Ornamental Plant No. of Root FW of DW of DW of No. of

length leaves length shoot shoot root Flowers/
plant
Compositae Pot marigold 89.9*y 95.8NS 101.9NS 55.9** 79.9* 70.4** －
Cornflower 102.9NS 115.5** 102.1NS － 111.3NS 86.8NS －
Chrysanthemum 103.8NS －－ 99.9NS 98.9NS 126.6** －
Cosmos －x －－ 119.9NS 120.1NS 111.2NS －
Zinnia 93.7NS －－ 88.6NS 91.7NS 96.8NS －
Thistle 114.8NS － 114.6* 99.9NS 118.1NS 120.8NS 142.9NS
Sunflower 106.1NS 96.8NS 84.4NS 113.3NS － 95.8NS 100.0NS
Safflower 104.8NS 89.7** 79.4** 91.6NS 100.2NS 84.6NS 100.0NS
African marigold 146.1** 95.5NS － 146.7** 176.2** － 100.0NS
China aster 103.2NS 97.3NS 79.1** 80.7* 82.4* 70.6** 68.4*
Coneflower 93.7NS 87.2NS 102.8NS 79.2* 84.2* 119.4NS 80.3NS
Liliaceae Tulip 110.6NS 102.6NS 86.2NS 104.4NS 110.5NS 69.7NS 100.0NS
Thunberg lily 88.2* 96.0NS 118.2NS 107.3NS 97.1NS 155.3NS －
Toritelia 93.1* 100.0NS 55.9** 77.2** 80.2** 74.8** 71.5**
Lily 37.2** 64.6** 42.1** 13.5** 13.2** 15.6** －
Labiatae Rocket larkspur 71.5** 93.8NS 51.4** 25.5** 38.1** 26.3** 88.3NS
Love-in-a-mist 181.4** 110.3NS 122.7NS 151.6** 127.1* 162.5** 100.0NS
Scarlet sage 99.5NS 101.0NS 91.6NS 103.6NS 106.1NS 112.5NS －
Fan columbine 104.4NS － 68.1** 74.6* 74.2* 80.3NS －
Caryophyllaceae Corn cockl 74.1** 85.4** 62.1** 27.9 33.1** 83.7NS －
Gypsophilla 105.3NS 102.6NS 83.9** 99.9NS 118.1NS 121.8NS 100.0NS
Carnation 42.4** 75.0** 61.2** 34.6** 46.5** 58.5** －
Leguminosae Sweet pea 85.1* 105.8NS － 78.5* 82.2* 79.8NS －
Lupine 98.1NS 106.5NS － 120.3NS 107.2NS 96.3NS 71.9NS
Cruciferae Rape blossoms 106.1* 100.0NS 95.6NS 121.2** 113.3* 50.2* －
Stock 60.3* 89.9NS 101.5NS 62.9** 78.3** 100.0NS 95.3NS
Onagraceae Farewell-to-spring 78.4** 92.1* 75.1** 44.7** 51.4** 28.3** 56.3**
Umbelliferae Bishop's weed 91.3* 97.5NS － 66.3** 69.4* － 91.1NS
Scrophulariaceae Snapdragon 72.8** 96.7NS 100.7NS 46.1** 56.3** 79.5NS 73.1*
Papaveraceae Corn poppy 50.4* 75.3NS 98.1NS 32.1** 52.5* 52.6* －
Amaryllidaceae Narcissus 97.1NS 102.0NS 78.8** 96.3NS 89.2NS 97.7NS 100.0NS
Amaranthaceae Feather cockscomb 92.9NS 80.7* 85.7NS 100.5NS －－ 100.0NS
Globe amaranth 102.8NS 100.0NS 102.8NS 84.5** 83.2** 100.0NS 82.7**
Gentianaceae Prairie gentian 83.8** 107.9* 51.1** 50.8** 60.2** 45.4** 62.2**
Campanulaceae Balloon flower 117.5* 102.5NS 78.8** 95.7NS 89.4NS 112.5NS 113.2NS
Plumbaginaceae Statice 109.2NS 94.2NS 98.1NS 94.7NS 97.8NS 68.5* 114.2NS
Chinese-lantern
105.3NS 104.7NS － 67.6** 64.8** 74.8** 114.7NS
Solanaceae plant
zGrowth performance (%) = Growth in absence of AC/Growth in presence of AC × 100; ysignificant at
5% level (*), 1% level (**) and not significant (NS) by t-test (n=36); xno data.
Table 10. Growth performances of some ornamental plants grown in hydroponics in the
presence or absence of activated charcoal (AC) in the nutrient solution (%)z.
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Soil Addition of Plant length No. of DW of Root length DW of root No. of

AC (kg/10a) (cm) leaves shoot (g) (cm) (g) flowers/plant
New (control) − 50.6az 11.4b 2.06a 19.2a 0.18b 6.7a
Successive − 39.9c 11.1bc 1.29c 15.6b 0.25a 5.6b
Successive 30 40.8c 11.7b 1.31c 14.6bc 0.18b 5.2c
Successive 60 48.4a 12.2a 1.85a 18.1a 0.19b 6.8a
Successive 120 44.0b 11.4b 1.60b 16.5b 0.19b 6.7a
Successive 240 42.2bc 11.2bc 1.54b 14.5bc 0.20ab 5.8b
Successive 480 40.3c 10.9c 1.35c 10.1c 0.11c 5.4c
z Values in a column followed by a different letter differ significant by Tukey's test (P=0.05; n=10)
Table 11. Effects of activated charcoal (AC) on the growth of prairie gentian, an ornamental
plant, grown on the soil of prairie gentian field amended with different amount of the AC.
Allelochemicals Pot Toritelia Lily Rocket Sweet Stock Farewell Bishop's Snap- Prairie
marigold larkspur pea -to- week dragon gentian
spring
Lactic acid +z + − + − + − + − −
Valeric acid − + − − − − − − − −
Malonic acid − − − − + + − − − +
Fumaric acid − + − − − − − − − −
Maleic acid − + − − − − − − − +
n-Caproic acid − + + − − − − − + +
Succinic acid + + − + − + − − − −
Benzoic acid + − + − + − − − − +
Malic acid − + − − − − − − − +
m-Hydroxybenzoic
− − − − − − + − − +
acid
p-Hydroxybenzoic
− − + − + − − − − +
acid
Adipic acid − + + − − − − − − −
o-
Hydroxyphenylacetic − − − + − − − − − −
acid
p-
Hydroxyphenylacetic − + − − − − − − − −
acid
Vanillin − − + − − − − − − −
3,4-Dihydroxybenzoic
− + − − − − − − − −
acid
Vanillic acid − − − + + − − − − −
n-Capric acid − + − − − − − − − −
z Detected (+) and not detected (-).
Table 12. The compounds identified in the exudates of some ornamentals adsorbed on
activated charcoal (AC) added in the nutrient solution.
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Allelochemicals Conc. Pot marigold Lily Rocket Sweet pea Stock Prairie
(μM) larkspur gentian
FW DW FW DW FW DW FW DW FW DW FW DW
shoot root shoot root shoot root shoot root shoot root shoot root
None(control) 0 530bz 4.7b 1640a 63a 140a 4.4a 1210a 18a 130a 1.1a 560a 23a
Lactic acid 50 420c 3.1c − − 130a 3.5a − − 150a 1.1a − −
100 420c 3.3c − − 140a 3.7a − − 120a 0.9a − −
200 430c 3.4c − − 160a 4.5a − − 120a 1.1a − −
400 420c 3.4c − − 150a 4.6a − − 110a 0.8a − −
Malonic acid 50 − − − − − − 1340a 23a 110a 1.1a 530a 24a
100 − − − − − − 1320a 32a 140a 1.2a 510a 24a
200 − − − − − − 1330a 21a 140a 1.1a 510a 25a
400 − − − − − − 1070b 17b 110a 1.5a 490b 29a
Maleic acid 50 − − − − − − − − − − 490b 17b
100 − − − − − − − − − − 460b 17b
200 − − − − − − − − − − 420b 18b
400 − − − − − − − − − − 390c 18b
n-Caproic acid 50 − − 1410a 43b − − − − − − 530a 22a
100 − − 1380a 41b − − − − − − 580a 25a
200 − − 940b 34b − − − − − − 550a 23a
400 − − 850b 35b − − − − − − 530a 25a
Succinic acid 50 510b 4.7b − − 120a 4.3a − − 130a 1.1a − −
100 490b 4.3b − − 160a 4.7a − − 120a 1.7a − −
200 510b 3.7b − − 140a 3.9a − − 120a 1.2a − −
400 490b 4.1b − − 140a 3.9a − − 110a 1.3a − −
Benzoic acid 50 470b 4.2b 810b 34b − − 1150b 18a − − 460b 19b
100 750a 6.2a 810b 34b − − 1110b 18a − − 470b 18b
200 530b 4.3b 800b 34b − − 1090b 21a − − 480b 17b
400 440c 3.1c 890b 42b − − 1110b 21a − − 470b 16b
Malic acid 50 − − − − − − − − − − 510a 22a
100 − − − − − − − − − − 480b 22a
200 − − − − − − − − − − 380c 22a
400 − − − − − − − − − − 390c 22a
m-Hydroxybenzoic
50 − − − − − − − − − − 520a 19b
acid
100 − − − − − − − − − − 510a 18b
200 − − − − − − − − − − 510a 18b
400 − − − − − − − − − − 420b 16b
p-Hydroxybenzoic
50 − − 990b 28b − − 1330a 21a − − 510a 22a
acid
100 − − 1170b 35b − − 1310a 34a − − 550a 23a
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Allelochemicals Conc. Pot marigold Lily Rocket Sweet pea Stock Prairie
(μM) larkspur gentian
FW DW FW DW FW DW FW DW FW DW FW DW
shoot root shoot root shoot root shoot root shoot root shoot root
200 − − 1010b 31b − − 980b 15b − − 610a 26a
400 − − 1010b 35b − − 870b 16b − − 470b 25a
Adipic acid 50 − − 1370a 36b − − − − − − − −
100 − − 1210a 28b − − − − − − − −
200 − − 910b 29b − − − − − − − −
400 − − 970b 22c − − − − − − − −
o-
Hydroxyphenylacetic 50 − − − − 110a 3.0b − − − − − −
acid
100 − − − − 110a 2.8b − − − − − −
200 − − − − 110a 2.4b − − − − − −
400 − − − − 60b 2.2b − − − − − −
Vanillin 50 − − 1340a 38b − − − − − − − −
100 − − 1310a 34b − − − − − − − −
200 − − 1030b 27b − − − − − − − −
400 − − 1010b 26b − − − − − − − −
Vanillic acid 50 − − − − 140a 4.6a 1230a 19a − − − −
100 − − − − 140a 4.6a 1190a 19a − − − −
200 − − − − 120a 2.8b 1010b 23a − − − −
400 − − − − 110a 2.4b 1110b 21a − − − −
z Values in a column followed by a different letter differ significant by Tukey's test (P=0.05; n=10)
Table 13. Effects of the identified compounds at different concentrations on the fresh and
dry weights (mg) of shoot and root of some ornamental plants.
revealed that the test plant length was increased by 96% over control as a result of the
addition of AC (60 kg/10a). Shoot dry weight and root length were increased by 90% and
94%, respectively, over control for the same concentration (60 kg/10a). Flower setting was
also increased at 60 kg AC/10a. This indicated that the reduced growth of prairie gentian
after prolonged cultivation in a field could be corrected by amending the soil with AC at the
rate of 60 kg/10a. In conclusion, of the ornamentals experiencing autotoxicity owing to the
chemicals exuded from their roots being more specific, this autotoxicity could be reduced, at
least to some extent, using AC in the root media.
3.4.6 Phytotoxicity of the identified allelochemicals

Root exudates from the ornamentals were analyzed and some compounds were detected.
The identified chemicals were mainly some small chain aliphatic acids and some simple
phenolic acids or phenolic compounds and those varied from extract to extract in the
ornamentals that experienced autotoxicity. Eleven organic compounds were detected in the
root exudates of toritelia roots and seven in prairie gentian (Table 12). Many compounds in
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the root exudates of the plants are yet to be identified. However, at least one aliphatic acid
or phenolic compound has been detected in the root exudates of the studied plants. A
bioassay was carried out to evaluate the inhibition potential of some identified compounds.
Different test concentrations were made with the compounds and a bioassay was furnished
with some test plants. Almost all the compounds inhibited the growth of tested plants in a
concentration dependent manner. Lactic acid significantly reduced fresh shoot weight and
concentration (50 µM) (Table 13). Benzoic and p-hydroxybenzoic acid in lily, even at 50 M,
root dry weight in pot marigold to 79% and 66% of control, respectively, even at low
significantly reduced fresh weight to 49% and 60% of over control, n-caproic, benzoic,
57%, and 60% of control, respectively. o-Hydroxyphenylacetic acid at 50 M reduced root

phydroxybenzoic, and adipic acid and vanillin decreased root dry weight to 68%, 54%, 44%,
dry weight in rocket larkspur to 68% of control (Table 13). Quantity and quality of exuded
allelochemicals varied from plants to plants (Inderjit, 1996) and in cucumber plants, root
exudation rate of different chemicals was found to range from 0.20 to 4.17 mg/d per plant
(Pramanik et al., 2000). This low concentration is apparently not enough to cause
autotoxicity in cucumber plants, but those cucumber plants experienced autotoxicity when
grown in absence of AC in the nutrient solution plant (Pramanik et al., 2000). Actually, in
natural conditions, occurrence of a chemical at high concentrations (100 μM or more) is rare
or absent. However, under field conditions or hydroponic culture, the exuded compounds
affect plant growth by additive or synergistic means (Inderjit, 1996) and thus, the
compounds even at low concentrations could induce significant growth inhibition in plants,
although their threshold inhibition at the individual level is quite high (Rice, 1984). Identical
results were found in the experiment (Table 13). So, it appears that the identified
compounds would be toxic enough to affect growth of the ornamental plants by additive or
synergistic effects.
We found a number of ornamental plants with autotoxic potential due to their root
exudation. Growth and yield of the ornamentals under investigation were found to be
improved in culture solution with AC supplementation where it used to trap the exudates.
Therefore, cultivation through hydroponics enables us to isolate the responsible
allelochemicals. Most of the autotoxic ornamental plants released mainly aliphatic acids or
phenolic compounds to the nutrient solution. Phytotoxicity was evaluated in terms of fresh
hydroxybenzoic acid showed phytotoxicity even at lower concentration (50 M). The
weight of shoot and dry weight of root and some compounds like lactic, benzoic acid and p-
aforesaid results would be useful for sustainable production of ornament plants. We have
tried AC to remove the allelochemicals from culture solution; however, more practical
measures should be investigated to manage the rhizosphere free of inhibitory exudates.
4. Conclusion
Intensive and continuous culture of vegetable crops on the same land for several years
causes replanting injuries like outbreak of disease and insect pest, exhaustion of soil fertility,
development of chemical interference (allelopathy) leading to growth and yield reduction.
Similarly in closed hydroponic system for the commercial cultivation of vegetables,
autotoxicity is also evidenced. Hydroponic culture solution accumulates root exudates
which in terns hamper water and mineral uptake due to root injuries. This managed culture
technique has the facility of trapping and isolating the chemicals released through plant
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roots. Therefore, autotoxicity phenomenon can clearly be investigated through hydroponics.

Our lab aims to investigate autotoxicity in a number of vegetables and ornamental crops, to
identify potential allelochemicals, their phytotoxicity through bioassay and suggest the
control measures. We cultured the plants in hydroponics using plastic containers in growth
chamber or in the greenhouse. Mineral nutrients were supplied as Enshi nutrient solution
and nutrient concentration were adjusted throughout the culture period. Activated
charcoals were supplemented in the air filters for trapping allelochemicals. Root exudates
were extracted and analyzed through GC-MS. Phytotoxicity of the identified chemicals was
assayed at several concentrations and potential growth inhibitors were identified for each
crop. We have showed species differences in the susceptibility to autotoxicity among leaf
vegetables in hydroponics. Therefore, knowledge regarding autotoxicity in vegetable crops,
autotoxic chemicals, and their phytotoxicity with control measures are useful for sustainable
crop production.
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Hydroponics - A Standard Methodology for Plant Biological
Researches
Edited by Dr. Toshiki Asao
ISBN 978-953-51-0386-8
Hard cover, 244 pages
Publisher InTech
Published online 23, March, 2012
Published in print edition March, 2012
Hydroponics-A standard methodology for plant biological researches provides useful information on the
requirements and techniques needs to be considered in order to grow crops successfully in hydroponics. The
main focuses of this book are preparation of hydroponic nutrient solution, use of this technique for studying
biological aspects and environmental controls, and production of vegetables and ornamentals hydroponically.
The first chapter of this book takes a general description of nutrient solution used for hydroponics followed by
an outline of in vitro hydroponic culture system for vegetables. Detailed descriptions on use of hydroponics in
the context of scientific research into plants responses and tolerance to abiotic stresses and on the problems
associated with the reuse of culture solution and means to overcome it are included. Some chapters provides
information on the role of hydroponic technique in studying plant-microbe-environment interaction and in
various aspects of plant biological research, and also understanding of root uptake of nutrients and thereof
role of hydroponics in environmental clean-up of toxic and polluting agents. The last two chapters outlined the
hydroponic production of cactus and fruit tree seedlings. Leading research works from around the world are
brought together in this book to produce a valuable source of reference for teachers, researcher, and
advanced students of biological science and crop production.
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4

2. Materials and methods

2.2 Plant cultivation in hydroponics

Chemicals Concentration (µM/l)

2.3 Gas chromatography-mass spectrometry (GC-MS) analysis of root exudates

2.4 Bioassay with the identified chemicals

2.5 Statistical analysis

3. Autotoxicity from the root exudates of vegetables and ornamentals in

3.1 Autotoxicity in cucumber (Cucumis sativus)

Fig. 1. Fruits of cucumber cv. ‘Shogin-aonaga-fushinari’, white arrow indicating (A) a

3.1.1 Cultivation of cucumber plants with or without microbial strain in presence of

raised in vermiculite were transplanted into plastic containers containing 50 l of

3.1.2 Cucumber cultivation with microbial suspension in absence of DCBA

3.1.4 Effects of microbial suspension on cucumber growth in the absence of DCBA

3.2 Autotoxicity in strawberry (Fragaria ananassa Duch.)

3.2.1 Cultivation of strawberry plants in hydroponics

Nutrient AC No. of FW of DW of DW of Root No. of No. of No. of

3.2.3 Phytotoxicity of the identified chemicals in root exudates of strawberry

Compoundz (Concentration, FW of shoot/ DW of shoot/ DW of Root length/

followed by a different letter differ significantly by Tukey's test (P = 0.05; n = 9).

3.3 Autotoxicity in taro (Colocasia esculenta Schott.)

3.3.1 Cultivation of taro plants in hydroponics

Charcoal No. of DW of Total yield/ Yield/plant by corm sizez (g)

3.3.3 Phytotoxins in root exudates of taro and their phytotoxicity

Allelochemicalz FW of shoot/ DW of No. of Root length DW root/plant

(NS) by the Tukey test.

Allelochemical Concentrations FW of shoot/ DW of No. of Root length DW

induced significant growth inhibition in taro plantlets even at concentration of 50 M/l.

3.4 Autotoxicity in some ornamental plants

3.4.1 Cultivation of ornamental plants in hydroponics

3.4.2 Bioassay of the identified autotoxic chemicals in nutrient solution

3.4.3 Bioassay in soils amended with activated charcoal

Family Ornamental Scientific name Cultivar

3.4.4 Growth performances of the ornamental plants grown in hydroponics

Family Ornamental Plant No. of Root FW of DW of DW of No. of

zGrowth performance (%) = Growth in absence of AC/Growth in presence of AC × 100; ysignificant at

Soil Addition of Plant length No. of DW of Root length DW of root No. of

3.4.6 Phytotoxicity of the identified allelochemicals

57%, and 60% of control, respectively. o-Hydroxyphenylacetic acid at 50 M reduced root

roots. Therefore, autotoxicity phenomenon can clearly be investigated through hydroponics.

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