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Role of Allelopathy in vegetables crops production
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Allelopathy Journal 25 (2): 275-312 (2010) International Allelopathy Foundation 2010
Tables: 7, Fig : -
Role of Allelopathy in vegetables crops production

JACOB JOHN*, J. SHIRMILA, S. SARADA and S. ANU
Cropping Systems Research Centre, P.O. Karamana

Kerala Agricultural University, Thiruvananthapuram, Kerala 695 002
E. Mail: jjkau@rediffmail.com
(Received in revised form: January 21, 2010)
CONTENTS
1. INTRODUCTION
2. SOIL SICKNESS
2.1 Tomato
2.2 Cucumber
2.3 Eggplant
3. AUTOTOXICITY
3.1 Cucumber
3.2 Tomato
3.3 Pea
3.4 Watermelon
3.5 Red pepper
3.6 Brinjal
4. ALLELOPATHIC INTERACTIONS
4.1 Vegetables and other crops interactions
4.1.1 Vegetables on other crops
4.1.2 Other crops on vegetables
4.1.3 Vegetables on vegetables
4.2 Vegetables and weeds interactions
4.2.1 Summer season vegetables
4.2.2 Cool season vegetables
4.3 Effects of vegetables on weeds
4.4 Allelopathic effects of trees on vegetables (Agroforestry)
5. PESTICIDAL POTENTIAL OF VEGETABLE CROPS
5.1 Fungitoxic/antibiotic effects
5.2 Nematicidal effects
6. ALLELOCHEMICAL STUDIES IN VEGETABLES
6.1. Cucumber
6.2. Other vegetables
7. FUTURE RESEARCH PROSPECTS
8. REFERENCES
*
Correspondence author
276 John et al
ABSTRACT
Vegetables are important worldwide, but their production faces problems
of yield decline due to soil sickness and autotoxicity, when grown continuously for
several years. Besides, allelopathic effects of other crops, weeds and trees also reduces
yields. Thus the allelochemical interactions and their effects on vegetables are
important in vegetable production. Although, research on various aspects of
allelopathy in vegetable crops has been done but not compiled. Soil sickness is
complex phenomenon due to several factors involved and autoxicity is major one. The
autotoxic potential of certain vegetables has been discussed. In multi-storey cropping
systems, where numerous crops and trees are grown together, vegetables are essential
components and allelopathic interactions arise. Several vegetables possess
antimicrobial principles and hence, allelopathically inhibit phytopathogenic fungi and
bacteria. Certain vegetables possess nematicidal principles and therefore, offer
immense potential for nematode control in their cultivation. Several studies have been
done to elucidate the role of allelochemicals in vegetables across the world. The
allelopathic interactions between the vegetables and other crops/weeds/trees and the
potential of vegetables for pathogen and nematode management/control are reviewed
in this paper. Future allelopathic research in vegetables should focus on (i) separating
the allelopathic interference from competition in vegetable fields and vegetables based
cropping systems, (ii) screening the germplasm/varieties of vegetable crops for
allelopathic potential and later on do genetic manipulations to breed new varieties, (iii)
exploiting the allelopathic potential of vegetable crops for weed control and plant
protection, (iv) determining the critical concentrations of allelochemicals in each
vegetable crop to express their inhibitory/stimulatory influences, (v) identifying the
compatible and beneficial associations of vegetable crops with other crops and trees
and (vi) determine the harmful and beneficial effects of allelopathy in vegetable crops
in pot culture and field studies.
Keywords: Agroforestry, allelopathy, allelochemicals, autotoxicity, crop, nematode,

pathogen, soil sickness, tree, vegetable crops, weeds
1. INTRODUCTION
The term allelopathy includes both harmful and beneficial biochemical

interactions between all types of plants including microorganisms, through the release of
chemicals from plant parts by leaching, root exudation, volatilization, residue
decomposition and other processes in both natural and agricultural systems (119).
Allelopathy can affect many aspects of plant ecology including occurrence, growth, and
plant succession, the structure of plant communities, dominance, diversity and plant
productivity (150). Allelopathy is a universal phenomenon and is not the domain of few
selected plants. Some vegetables with strong allelopathic effects have been investigated
(201). The allelopathic interactions between the vegetables and other crops/weeds/trees
and the potential of vegetables for pathogen and nematode management/control are
reviewed in this paper. Allelopathic interactions are complex and can involve different
classes of chemicals. Allelochemicals are present in all plant tissues, several chemicals can
be released together and may exert additive or synergistic effects (145). The vegetables
production faces many problems (i). Yield decline due to autotoxicity and soil sickness
(when vegetables are grown continuosly for several years) and (ii). Low yield of
vegetables in monoculture and cropping systems due to allelopathic effects of crops, weeds
Role of Allelopathy in vegetables crops production 277
and trees (72,80,146,201). Researchers study only the competition for nutrients and
moisture and not the allelopathic effects (145).
The allelochemical interactions and their effects on vegetables are important
aspects to be considered in vegetables production. Although, research on allelopathy in
vegetable crops has been done worldwide, however, it has not been compiled. Hence this
review on allelopathy in vegetables production has been prepared. It deals with (i).
problems of soil sickness and autotoxicity, (ii). use of vegetables allelopathy for weed
management, pest management and (iii). identify ideal combinations (companion, rotations
and agroforestry systems) of vegetable crops.
2. SOIL SICKNESS
The soil sickness has been observed in many vegetables (Table 1) and has been
attributed to several factors, viz. depletion of soil nutrients, accumulation of phytotoxic
compounds, and accumulation of soil-borne pathogens due to improper cultural practices,
such as continuous monoculture and improper management of soil-borne diseases (136).
Continuous cultivation of vegetables in same field reduces their growth, yield and quality
(50,72). However, the magnitude of yield decline varies with crops. Legumes, cucurbits
and solanaceae crops are most vulnerable to soil sickness (91).
Table 1. Vegetable crops affected by soil sickness and autotoxicity
Crop References
Soil Sickness
Bean (Phaseolus vulgaris L.) 75
Pea (Pisum sativum L.) 58, 122, 183
Potato (Solanum tuberosum L.) 13, 64
Asparagus (Asparagus officinalis L.) 17, 49, 64, 119, 139, 140, 197, 198, 157
Cabbage (Brassica oleracea L.) 12
Celery (Apium graveolens L.) 19
Cucurbits (Cucumis spp. and Cucurbita spp.) 196, 200, 210, 207
Eggplant (Solanum melongena L.) 210
Tomato (Lycopersicon esculentum Mill.) 133, 210
Autotoxicity
Cucumber (Cucumis sativus L.) 107, 109, 201, 205, 207
Tomato (Lycopersicon esculentum Mill.) 57, 88, 108, 167, 175
Pea (Pisum sativum L.) 33, 54, 91, 176, 189
Watermelon (Citrullus lanatus (Thunb.) Mansf.) 53, 124
Red pepper (Capsicum annuum L.) 184
Egg plant (Solanum melongena L.) 54, 175
2.1. Tomato (Lycopersicon esculentum Mill.)

Tomato (Lycopersicon esculentum Mill.), when grown in monoculture for long
periods leads to soil sickness (212). Autotoxicity and soil problems in tomato production
have been reported long ago (111,213). Continuous cropping reduces the root vitality and
fruit quality of tomatoes (196). Zhou et al. (214) reported the presence of di-iso-octyl
phthalate, di-iso-butyl phthalate, tannic acid and salicylic acids in tomato plants.
278 John et al
Singh et al. (164) studied the autotoxic effect of tomato on germination, seedling
growth, biophysical and biochemical parameters of plants grown in hydroponic culture.
The root, stem and leaf leachates of tomato had varying effects on various parameters. The
germination was stimulated by higher concentration of root and leaf leachates. The
moderate concentration of root leachates inhibited the germination than lower and higher
concentrations. The higher concentration of leachates stimulated the radicle growth, while
lower concentration inhibited it. However, the reverse was true for plumule growth.
Chlorophyll a, chlorophyll b and total chlorophyll content decreased with increasing
concentration of leachates. The inhibition of chlorophyll synthesis was concentration
dependent with maximum inhibition in leaf leachate. The root and stem leachates
decreased the protein content to minimum level. Superoxide dismutase and peroxidase
activity was inhibited but catalase activity was stimulated. The leachates adversely affect
the biophysical and biochemical parameters resulting in arrested growth and low yield of
tomato. The influence of leachates on growth followed the order roots > stem > leaf. The
root leachates drastically decreased the tomato fruit growth than control, while leaf
leachates completely prevented fruit formation.
Soil amendment with organic matter (56,69), or some compounds (172) controls
the soil sickness due to soil-borne plant pathogens (Table 3). In Brassica spp., allyl-
isothiocyanate was the main toxin to control the soil-borne pathogens (30,31,115). Kauri
Paasuke (86) reported that amendment of soil with milled peat, or green manuring of
weeds [50% couchgrass (Agropyron repens (L.) Beauv.)] suppressed the soil sickness in
apple tree nursery, but led to the buildup of nematophagous fungi and tardigrade beetles.
The soil amendment with formulated organic compounds [S-H mixture (a composted
mixture of bagasse, rice husks, oyster shell powder, urea, potassium nitrate, calcium
superphosphate and mineral ash (172)], controls the Fusarium wilt of radish (Raphanus
sativus L.) and watermelon (Citrullus lanatus (Thunb.) Mansf. and clubroot of crucifers
(66). Use of PerlkaTM (granulated calcium cyanamide product, SKW Trostberg
Aktiengesellschaft, Germany), as nitrogen fertilizer controls the plant pathogens
[Cercosporella herpotrichoides Fron., Plasmodiophora brassicae (152) and Sclerotinia
sclerotiorum (Lib.) de Bary (74,97,114,152)]. The mechanism of pathogen suppression
may be due to the release of toxic ammonia gas from urea fertilizer (56,70,71,73).
Table 2. Effects of crucifers biomass added as soil amendments to control soil sickness
Soil amendment Effects on test plant References

Crucifers Reduced severity of damping-off of cabbage caused by 30, 31
Rhizoctonia solani Kuhn
Crucifers Reduced severity of damping-off of watermelon caused by 32
Pythium sp.
Crucifers Reduced severity of potato dry rot caused by Fusarium 115
sambucinum Fuckel
Crucifers Reduced severity of root rot of pea caused by Aphanomyces 120
euteiches Drechsler
2.2. Cucumber (Cucumis sativus L.)

The effects of Fusarium oxysporium f. sp. cucumerinum [the pathogen causing
Fusarium wilt in cucumber (Cucumis sativus L.)] and cinnamic acid, a principal autotoxic
component in the root exudates of cucumber were studied, on the plant growth,
photosynthesis and incidence of Fusarium wilt in cucumber, to elucidate the interaction of
autotoxins and soil borne pathogens in the soil sickness. F. oxysporum f. sp. cucumerinum
(FO) and cinnamic acid (CA) at 0.1 or 0.25 mM significantly decreased net photosynthetic
rate, stomatal conductance and yield of Photosystem II photochemistry (Phi PS II).
Thereafter the plant biomass production was reduced, but did not induce photoinhibition.
Pretreatment with CA before inoculation with FO increased the effectiveness of FO,
together with a slight photoinhibition. CA pretreatment significantly increased the
percentage of plants affected by Fusarium wilt, browning index of vascular bundle and
Fusarium population in the nutrient solution. All these results indicate that CA enhanced
the Fusarium wilt by predisposing cucumber roots to infection by FO through a direct
biochemical and physiological effect. It is likely that soil sickness results from an
interaction of many factors (198). Introduction of antagonistic bacteria and beneficial
microbes could overcome the problems caused by pathogenic microbes and autotoxic
compounds (212) that led to soil sickness in tomato, cucumber, melon (Cucumis sp.) and
eggplant or brinjal (Solanum melongena L.).
2.3. Eggplant (Solanum melongena L.)

The possibility of repeated use of substrate in the growing of eggplant was
studied (138). It was found that peat repeatedly used showed a negative effect on the
growth and yield of eggplant due to the decrease in water in substrate and accumulation of
phenolic compounds. On the other hand in the mixture of bark and peat, a positive effect
of repeated use of substrate was observed. In this type of substrate, the amount of water
increased and phenolic compound concentration was low. Response of eggplant depended
on the type of substrate and on the time of its use. The longer the peat substrate was used,
the lower was the dry matter of eggplant leaves and the fresh weight of fruits. An inverse
dependence was found, when eggplant was grown in bark with peat substrate. In
comparison to first eggplant growing, in the third substrate use, the dry weight of leaves of
plants grown in peat was 30% lower and the fresh weight of fruits was 16% lower. On the
other hand, in bark with peat substrate, the dry weight of leaves increased by 150% and
fresh weight of fruits by 21%. In both types of substrates, the dry weight of stems was
similar and did not change depending on the time of substrate use. Comparison of the
effects of repeated use of substrates indicates that symptoms of substrate fatigue that
occurred in peat substrate were not observed in the bark+peat substrate. It is postulated that
this finding reflects differences in the physico-chemical properties of the substrates
(mainly the water content and C/N ratio). These differences were responsible for the
differences in the speed of release of phenolic compounds from the decaying crop residues
of eggplant and their subsequent degradation (138).
3. AUTOTOXICITY
Soil sickness is complex phenomenon and of the several factors involved,

autoxicity is major one. Autotoxicity has been reported in several plants grown in
monocultures (162,206,208,209) and also in vegetables crops (Table 1). Mostly, these
plants release autotoxins through leachings from the living and dead plant parts, root
280 John et al
exudates, biomass decomposition and volatilization (18,148). Many autotoxins [simple

organic acids, straight chain alcohols, aldehydes or ketones, unsaturated lactones, fatty
acids, naphthoquinones, complex terpenoids, polypeptides, alkaloids, glucosinolates etc.]
have been isolated from plants and they affect many biochemical and physiological
processes (126,162,192). The autotoxic potential of certain vegetables are discussed.
3.1. Cucumber (Cucumis sativus L.)

The poor growth due to successive cropping of cucumber has been observed.
Gaidmark (44) grew several crops of tomato and cucumber in gravel culture and detected
some unidentified toxins in the nutrient solution. Testa of cucumber var. PI 169391 seed
contains inhibitor (s) that suppressed the seedling growth of cucumber (107). Seed
fermentation and/or leaching with distilled water reduced the inhibition, while activated
charcoal eliminated the toxicity. Inhibitory effects of cucumber seed extract on cucumber
decreased with maturity of the cucumber fruit. Extracts of seeds and fruits of cucumber
were non-toxic, however, the fruit juice extracts from PI 169391 suppressed the growth of
cucumber (107).
Root exudates: The cinnamic and hydroxybenzoic acids present in the cucumber root
exudates drastically reduced the activities of dehydrogenase, root ATPase, nitrate
reductase and superoxide dismutase and inhibited the uptake of K+, NO3- and H2PO4- ions
(109). Yu and Matsui (204,206) investigated the autotoxic potential of cucumber root
exudates and reported that addition of activated charcoal to nutrient solution for the culture
of cucumber did not affect the inorganic composition in nutrient. But, it significantly
improved the growth of cucumber. Root exudates at reproductive stage showed higher
phytotoxicity than the vegetative stage. Nine compounds including benzoic acids,
cinnamic acids and p-thiocyanatephenol were identified as phytotoxic substances (205).
These compounds enhanced ion leakage from cucumber roots and inhibited ion uptake
(112, 205, 207). Compounds such as cinnamic acid showed great activity even at a
concentration less than 0.01 mM, indicating the possibility of autotoxicity under field
conditions (207). Yu and Matsui (205) examined the effects of root exudates of cucumber,
aromatic carboxylic acids in root exudates and their analogues upon the uptake of NO3 ,
H2PO4, SO42-, K+, Ca2+, Mg2+, and Fe2+ by intact cucumber seedlings. Root exudates
inhibited the uptake of all ions analyzed except H2PO4. Inhibition of ion uptake by
cinnamic acid, a main component of root exudates, was both concentration and pH
dependent. Inhibitory effect on ion uptake increased with decreasing pH. In continuous
cropping, allelochemicals from root exudates of cucumber roots are inhibitory to its own
growth (201).
Cucumber plants were grown hydroponically with or without addition of
activated charcoal (AC) to the nutrient solution at different temperatures and photoperiods.
Significant growth differences were observed between charcoaled and non-charcoaled
plants. Plants without AC were severely retarded in growth than those with AC. A number
of simple phenolic and aliphatic acids were identified in the cucumber root exudates along
with a chlorinated compound, 2,4-dichlorobenzoic acid (2,4-DCBA). All the identified
compounds inhibited the growth of test plants in a concentration dependent manner and
2,4-DCBA was most inhibitory. To overcome autotoxicity, cucumber plants were grown in
the nutrient solution added with 2,4-DCBA and also with or without addition of a
microbial strain or AC. It was revealed that vegetative growth of cucumber plants grown
with 2,4-DCBA and with a microbial strain recovered significantly. Susceptible cucumber
plants grafted on tolerant cucumber plants decreases the autotoxicity (143).
3.2. Tomato (Lycopersicon esculentum Mill.)

Hirano (57) and Soil Microbiological Laboratory of Japanese Agriculture
Technology Institute (167) confirmed the release of inhibitors during the decomposition of
tomato plant residues. The phytotoxicity of the residual nutrient solution after cultivation
of tomato was greater on itself than on other plant species (175). The phytotoxicity was
mainly due to the acidic substances in the solution (176,203). Mizutani (118) isolated
vanillic, p-hydroxybenzoic and gentisic acids as allelochemicals from tomato roots.
Rahman and Newton (148) reported that the addition of activated charcoal to the nutrient
solution during the growth of tomato significantly improved its growth. The aqueous
extract of tomato leaves inhibited the seedlings growth and biomass accumulation in
tomato (88). Yu et al. (208) confirmed this effect and reported that only the nutrient
solution at the reproductive stage of tomato showed phytotoxicity. Phthalic acid (PA) and
phloroglucinol dihydrate (PD) was isolated and identified in the root exudates of tomato
(90). Yu and Matsui (203) identified benzoic, palmitic and sinapic acids from the residual
culture solution and benzoic, phenylacetic, 2-hydroxy-3-phenylpropanoic, p-
hydroxybenzoic, vanillic, palmitic, ferulic, caffeic and sinapic acids from the used
activated charcoal.
The detrimental effects of autotoxicity decreases the plant population,
regeneration failure and subsequent yield decline in tomato (88). Potential autotoxicity
also exists in root exudates of tomato growing in hydroponics (108). The autotoxic
substances viz., phthalic acid and phloroglucinol dehydrate caused root oxidative damage
of tomato seedlings. Potted tomato seedlings were cultured in perlite and treated with
phthalic acid (PA) and phloroglucinol dihydrate (PD) as exogenous autotoxins at 1 mM, 5
mM and 10 mM concentrations. The application of both PD and PA and especially PD
increased the MDA (malondialdehyde) contents. The activities of SOD (superoxide
dismutase), CAT (catalase) and POD (peroxidase) depended on autotoxins (PA or PD),
their time of action and concentration. The enzyme activities increased with application of
PA on 5th day and decreased on 10th day except at 10 mM PA. On 20th day, the activities of
all enzymes decreased except SOD at 1 mM. Similar trend of enzyme changes was present
in the treatments of PD, except POD activity that kept growing on the 10th day. Results
indicated the adverse effects of exogenous PA and PD on enzymes of antioxidant defence
system, resulting in lipid peroxidation in roots of tomato seedlings (211).
3.3. Pea (Pisum sativum L.)

In pea grown continuously, nearly 50% yield decreases in the subsequent crop
(91). Soil sickness of pea could be overcome by leaching soil with water or ethanol,
suggesting that phytotoxins played an important role in this problem (33). The addition of
sap of living or decaying root of pea to nutrient solution stunted the growth of pea (54).
The residual nutrient solution after culture of pea not only inhibited the growth of pea, but
also showed significant phytotoxicity to other crops such as carrot (Daucus carota L.),
eggplant, bean (Phaseolus vulgaris L.) and Chinese cabbage (Brassica oleracea cv.
Chinensis) (59,60,61). Hatsuda et al. (52) isolated vanillic acid and p-coumaric acid from
282 John et al
the water extract of pea root. Takijima and Hayashi (176) fractionated the residual solution
after hydroponic culture of pea and found that acidic and neutral substances accumulated
in the nutrient solution mainly caused the phytotoxicity. Hirayoshi et al. (61) reported that
phytotoxicity of root exudates was mainly due to acidic substances and at least two
phenolic acids were present in the exudates. Vaughan and Ord (189) identified ferulic,
vanillic, p-coumaric and p-hydroxybenzoic acids in the root exudates of pea seedlings.
3.4. Watermelon (Citrullus lanatus (Thunb.) Mansf.)

In commercial production of watermelon, an interval of 5 to 7 years is
recommended, in areas where soil sickness is a serious problem. Hatsuda et al. (53)
isolated o-hydroxybenzoic acid from water extracts of watermelon root as a phytotoxic
substance. Nishimura (124) observed that o-hydroxybenzoic acid was a precursor chemical
causing the browning of vascular bundles in Fusarium wilt. When seedling was incubated
in o-hydroxybenzoic acid solution, it exhibited symptoms similar to that caused by
Fusarium (181). Studies indicated that among the tested phenolic acids, o-hydroxybenzoic
acid had the strongest ability to inhibit ion uptake by cucumber (207). Confirmatory
studies on the role of o-hydroxybenzoic acid in the soil sickness problem of watermelon
are needed.
3.5. Red pepper (Capsicum annuum L.)

In Korea, the yield of red pepper decreases due to continuous cropping. Although
this decline had been mainly attributed to the occurrence of diseases, it was suggested that
allelopathy may partly account for the decline. Tsuchiya et al. (184) assayed the growth
inhibition of red pepper by a germination test using water or organic solvent extracts of its
leaf, stem and root. Methanol extracts of stem and root of red pepper strongly inhibited the
radicle growth of red pepper. Methanol extracts of leaf and root and water extract of root
inhibited the hypocotyl growth. The methanol and water extracts of red pepper root
contained several phenolic acids such as p-hydroxy benzoic acid and vanillic acid. It was
inferred that the red pepper decline under continuous cropping was due to allelopathy.
3.6. Brinjal (Solanum melongena L.)

Hayashi and Takijima (54) reported that amending soil with dry roots of brinjal
inhibited the growth of brinjal itself. Takijima and Hayashi (175) investigated the
phytotoxicity of root exudates and found that the exudates of brinjal showed great
phytotoxicity to brinjal itself and tomato, whereas, it improved the growth of wheat and
upland rice. However, no inhibitors have been identified from roots or root exudates.
4. ALLELOPATHIC INTERACTIONS
4.1. Vegetable crops and their interactions

Several workers have investigated the allelopathic interaction of vegetables with
other crops and between vegetables.
4.1.1. Effects of Vegetable crops on other crops

Bhatt and Dhawan (16) reported that addition of crushed seeds of carrots and
coriander (Coriandrum sativum L.) gave significantly higher grain yield of wheat. Carrot
at all levels and coriander at 4 % resulted in reduction of Heterodera avenae cyst
population.
The aqueous extracts of fresh and dried roots of Chinese cabbage reduced its own
radicle growth, fresh and dry weight and the growth of mustard (Brassica campestris L.).
Extracts of dried roots were more inhibitory than fresh roots. Inhibition of growth was
more in mustard than Chinese cabbage (2).
Peterson and Harrison (133) screened yellow squash (Cucurbita pepo L.) and two
cucumber (PI 165046 and PI 169391) accessions for the presence of constitutive chemical
factors which interfere with the germination and growth of proso millet (Panicum
milliaceum L.). Hexane, ethylacetate and water soluble components were all inhibitory,
especially those obtained from leaves, stems or roots. Extracts obtained from these plant
organs inhibited germination between 61 and 97 % at 50 mg DW equivalent per ml.
Extracts from seed parts inhibited germination by 18-64 % at the same extract
concentrations (50 mg DW equivalent per ml). The combined hexane and ethanol extracts
of leaves strongly inhibited germination. Aqueous leaf homogenates of the three cucurbits,
incorporated into the potting media, inhibited growth of millet. Squash leaf material
inhibited shoot length of millet by 26%. The cucumber accessions had no effect. Shoots
dry weight was inhibited by 49, 32 and 35% and root dry weight by 73, 56 and 67% by
homogenates of squash, cucumber 165046 and cucumber 169391 leaves, respectively.
Allelopathic effects of potato (Solanum tuberosum L.) tuber shoots on
germination and seedling growth of bean, soyabean (Glycine max (L.) Merr), lentil (Lense
ulinaris Medik.), maize (Zea mays L.) and wheat were studied (131). Fresh shoots were
allowed to decay for 72, 120, 168, 240 and 336 h in normal water in the ratio of 1:10 w/v.
These water extracts were applied to the crop seeds. All extracts reduced seed germination
and seedling growth. For lentil and maize, the lowest negative effect on germination was
obtained with the 120 h extract, for bean and wheat with the 168 h extract and for
soyabean with the 240 h extract. Maize had least negative effect from 72 and 120 h
extracts, bean and lentil from the 120 h extract, wheat from the 120 and 168 h extracts and
soyabean from the 240 h extract (132).
4.1.2. Effects of other crops on vegetable crops

Volatile emissions from residues of the winter cover legumes, Berseem clover
(Trifolium alexandrinum L.), hairy vetch (Vicia hirsute (L.) SF.Gray), and crimson clover
(Trifolium incarnatum L.), inhibited germination and seedling development of onion
(Allium cepa L.), carrot and tomato. Hydrocarbons, alcohols, aldehydes, ketones, esters,
furans, and monoterpenes were identified in these residue emission mixtures (20).
Growth chamber tests demonstrated that alfalfa residue is toxic to cucumber seed
germination and seedling growth. Ground alfalfa roots (0.5% w/w, dry weight) inhibited
germination when added to the growing medium. Alfalfa roots (0.5% w/w, dry weight)
were also toxic to pre-germinated cucumber seed. However, cucumber seedlings grew
normally if the medium in which ground roots were added was irrigated so as to leach
down chemicals and kept for >1 day before planting (39).
284 John et al
Water leachates from guayule (Parthenium argentatum) processing residues, or

residues mixed with peat moss or soil significantly delayed onset of germination and
decreased percentage germination of broccoli (Brassica oleraceae var. italica cv.
Emperor) cantaloupe (Cucumis melo cv.Topmark), cauliflower (Brassica oleraceae var.
botrytis cv. Snow crown), lettuce cv. Empire, pepper (Capsicum annuum cv. NM 6-4) and
tomato cv.GS 12. These effects can be ascribed to p-anisic acid present in leaf resin and a
variety of substituted benzoic acid, cinnamic acids present as carboxylate salts in leaves
and bagasse (193).
Stoimenova (170) reported that cucumbers grown after soyabean had shorter
roots than the controls or those grown after Amaranthus retroflexus or the mixtures.
Cucumber hypocotyl length was slightly greater after soyabean and/or A. retroflexus than
in the control.
The effect of sesbanimide (an alkaloid from Sesbania seeds) and extracts from the
ground powders of whole Sesbania bispinosa seeds, and the isolated embryos, testae and
intact seeds of S. punicea (a noxious weed), on the germination and growth of vegetable
(including cucumbers cv. Stono, lettuce cv. Queen and radish Sparkler) was investigated
(188). An inhibitory effect was observed for all treatments, being most noticeable
regarding seedling growth than germination. In all cases, roots were more sensitive and
hence, its growth more severely retarded than shoots. This inhibitory effect was also
detected if cucumber seeds were soaked in extracts prior to sowing. The embryo and testa
extracts of S. punicea produced an inhibitory response in the test species, the former being
the most inhibitory (188).
Seeds of cucumber, green beans (Phaseolus vulgaris L.), tomatoes, redroot
pigweed (Amaranthus retroflexus) and annual Italian ryegrass (Lolium multiflorum) were
exposed to 0, 16.7, 33.3, and 66.7 g L-1 of kenaf (Hibiscus cannabinus L.) extract (154).
Distilled water and three concentrations of polyethylene glycol (PEG) were included as
controls. Germination in tomato was reduced by 30% when exposed to the highest
concentration of unweathered kenaf (154).
Rye seed densities of 50 and 100 kg ha-1 inhibited lettuce seedling growth and
vigour. The visual symptoms were similar to atrazine and benzoxazolinone (BOA) effects.
Leaf yellowing and vigour were dependent on rye density, atrazine and BOA
concentrations, where, leaf yellowing increased while vigour decreased with an increase in
density and concentration (168).
Nearly 239 medicinal plants were evaluated for allelopathic activity on lettuce cv.
Great Lakes using the sandwich method (43). Around 223 species inhibited, whereas 17
species promoted radicle growth in lettuce. Radicle growth inhibition of >80, 60-79, 40-
59, 20-39, and 0.30-19.0% was attributed to 19, 16, 43, 72 and 73 species, respectively.
Artabotrys odoratissimus (A. hexapetalus) inhibited lettuce growth the most. Strong
inhibitory activity was also recorded for Annona cherimola, Dialium guianense,
Tamarindus indica, Emblica pectinata, Hevea brasiliensis, Garcinia oblongifolia,
Elaeocarpus serratus, Schleichera oleosa, Paeonia lactiflora and Sandoricum koetjape
(43).
Aqueous leaf extracts of Aloe vera extracts did not significantly affect seed
germination but promoted root length and seedling height of lettuce, radish and turnip
(Brassica rapa L.). The extracts also increased the number of roots in lettuce and turnip,
but not in radish (103).
Pot culture experiments revealed that aqueous extracts of cotton (Gossypium sp.)
significantly inhibited the seed germination and seedling growth of tomato (104).
4.1.3. Effects of Vegetable crops on vegetable crops

Kim and Kil (89) reported that the aqueous extract of leaves, stem and root of
tomato heavily inhibited the seed germination and growth of lettuce and tomato. In
general, the phytotoxicity of the tomato extracts increased in the following order: time for
extracting 25 h < 48 h < 72 h. Tomato exhibited auto inhibition. Four phytotoxic
phenolics, salicylic acid, gallic acid, protocatechuic acid and vanillic acid were obtained
from the aqueous extracts of tomato leaves by paper chromatography. The aqueous
leachates of tomato significantly inhibited growth of certain crops in pot culture.
Root exudates from cucumber, squash [Cucurbita moschata (Duch.) Poir] and
melon seedlings inhibited tomato seed germination. Squash exudates had the greatest
effect, reducing germination from 91.4% (control) to 76.8%. Squash root exudates reduced
the germination rate index from 0.46 (controls) to 0.43, whereas root exudates from
cucumbers and melons increased it to 0.50 and 0.48, respectively. The effects of root
exudates on 30 or 45 day old seedlings were mixed. All the exudates reduced seedling DW
when applied to 30 day old plants but had no significant effect on seedling DW when
applied to 45 day old plants. Squash and melon root exudates significantly increased root
and seedling FW when applied to 30 and 45 day old plants (129).
Aqueous extracts of asparagus roots inhibited seed germination in tomato and
lettuce (Lactuca sativa L.), but not in cucumber. The extracts reduced hypocotyl growth in
lettuce, shoot growth in asparagus, and inhibited radicle elongation in lettuce and
asparagus. Seedling growth in tomato was not affected. Inhibition was concentration
dependent. Radicle growth in 'Grand Rapids' lettuce was sensitive to an extract
concentration as low as 0.05 g dry root tissue/100 ml H2O. Asparagus radicles were more
sensitive than asparagus shoots. Aqueous root extracts of A. racemosis also inhibited
germination and radicle growth in 'Grand Rapids' lettuce (55).
Water extracts of soils from pea and taro fields with continuous cropping
inhibited the radicle growth of lettuce. Since the same phenomenon was observed even
when the extracts were autoclaved, it is considered that the inhibition may be caused by
some allelochemicals rather than by the effect of harmful soil microorganisms (182). Leaf
extracts of vegetables such as asparagus, watermelon, pea, tomato and cucumber inhibited
the radicle growth of lettuce, regardless of the low EC (electrical conductivity) value of the
extracts. In contrast, root extracts of these vegetables enhanced the inhibition of the root
growth of lettuce as the EC value and the contents of total polyphenols increased. The root
extract of asparagus strongly inhibited the germination of lettuce in spite of low EC value.
It is considered that the root of asparagus may contain some allelochemicals (182).
Hydrophobic root exudates of cucumber collected at different growth stages with
Amberlite XAD 4 resin were bioassayed with lettuce seedlings (205). The exudates at the
reproductive stage were more phytotoxic than those at the vegetative stage. The exudates
contained organic acids such as benzoic, p-hydroxybenzoic, 2,5 dihydroxybenzoic, 3
phenylpropionic, cinnamic, p-hydroxycinnamic, myristic, palmitic and stearic acids, as
well as p-thiocyanatophenol and 2 hydroxybenzothiazole. All chemicals, except 2
hydroxybenzothiazole, were toxic to the growth of lettuce (205).
286 John et al
Pot trials were carried out to investigate the effects of plant incorporation into
sand and of plant extracts and isolated chemicals on the growth of several vegetables and
on lettuce seed germination (183). Pea growth was inhibited up to 30% by the
incorporation of pea tops (shoots) into sand (4 g/1.7 kg sand). The growth of pea tops was
inhibited by incorporation of taro, especially the roots. The incorporation of taro roots also
inhibited taro root growth. Cucumber growth was inhibited by incorporation of taro roots
or tops or cucumber tops. Pea growth was inhibited 10-15% by water extracts of pea or
taro roots. Ethyl acetate extracts of pea roots inhibited pea growth and lettuce seed
germination. Several phenolic acids, considered to act as allelochemicals in the roots of
vegetables, inhibited lettuce seed germination in the order salicylic acid > p-
hydroxybenzoic acid > vanillic acid > p-coumaric acid > gentisic acid. Pea growth was
inhibited 20% by vanillic acid and p-coumaric acid at 300 ppm. Tomato growth was
inhibited by vanillic acid, gentisic acid and p-hydroxybenzoic acid (in descending order).
Watermelon growth was inhibited by 10 and 70% with 10 and 500 ppm salicylic acid,
respectively. Concentrations >500 ppm induced stem rot like symptoms, leading to the
conclusion that salicylic acid is an indirect cause of soil sickness in watermelons (183).
Allelopathy properties of solvent extracts from broccoli harvested before the head
formation were evaluated through biological assays on germination and root elongation of
seedlings of broccoli, lettuce and tomato. Chloroform extracts of broccoli drymatter as
well as its chromotographic fractions showed high allelopathic activity on both seed
germination and root elongation of broccoli, lettuce and tomato. The allelopathic activity
was exhibited mainly as inhibition in root elongation rather than in the seed germination.
Among the five chromatographic fractions, three of them (I, II, III) were very effective,
while subfractions obtained from the fraction II of chloroform extract showed non-
significant allelopathic effects. As the allelopathic activity decreased with fractionation, it
may be attributed to a complex of non-polar compounds (35).
The aqueous extract of tomato significantly inhibited the growth of cucumber,
radish, lettuce, Chinese cabbage (Brassica pekinensis (Lour.) Rupr.) and broccoli
seedlings. But, tomato volatiles had no significant effect on growth of green gram (Vigna
radiata (L.) Welczek), chinese cabbage, lettuce or tomato seedlings (213). Forty days after
transplanting, tomato seedling root exudates significantly inhibited cucumber, but not
lettuce, growth. It is suggested that tomato should be avoided as an intercrop with
cucumbers in solution culture or in solar energy greenhouse (213).
4.2. Vegetables and weeds interactions
4.2.1. Summer season vegetable crops

Weeds, besides competing with vegetables for nutrients and water, also exert
allelopathic effects on vegetables. Tripati and Srivastava (180) studied that effect of
aqueous extracts of certain weeds on the seed germination of brinjal. The plant extracts
exerted varied effects on seed germination. The extracts of Holarrhena antidysentrica,
Cannabis sativa, Crinum asiaticum, Nelumbo nucifera, Sansevieria roxburghiana and
Salvinia natans (leaf extract) inhibited germination. Extracts of Spirodela polyrrhiza,
Marsilea minuta, Salvinia natans (shoot and root extracts), Ceratoplyllum demersum and
Eichhornia crassipes (extracts of shoot, root and flowers) had stimulatory effects.
Almodovar et al. (7, 8) determined the allelopathic effects of root exudates of

seven weed species viz., Parthenium hysterophorus, Echinochloa colonum, Amaranthus
dubius, Sorghum halepense, Rottoelia exalata, and Trianthema portulacastrum on
pumpkin (Cucurbita maxima (Duch.) Poir) and egg plant. The treatments included (0):
root exudates not added; (3): root exudates applied in 3 alternate days for 6 weeks and (5):
root exudates applied during 5 consecutive days for 6 weeks. Root exudates of all weed
species evaluated caused adverse allelopathic effects on egg plant seedlings through
decreased stem length and dryweight. Besides, the effect on vine length and dry weight, a
decrease of more than 50 % in pumpkin yields was recorded when weeds grew freely in
association with the crop.
Laboratory and polyhouse experiments were conducted to determine the
allelopathic effects of aqueous extracts of leaves of common weeds (Cynodon dactylon,
Cyperus rotundus, Eupatorium odoratum, Imperata cylindrica, Ipomoea sepiaria, Mikania
cordata and Parthenium hysterophorus) on tomato (157). The results revealed that all
weeds inhibited germination of tomato seeds than control (distilled water). The aqueous
leaf extracts of I. sepiaria and E. odoratum were most inhibitory to germination of tomato
seeds, even 15 days after sowing. Leaf extracts of these weeds, obtained 240 h after
decaying, recorded more adverse allelopathic effect in seed germination upto 7 DAS (days
after sowing) and upto 15 DAS in pots than weed extracts of 120 h after decaying. Shoot
length of tomato was shorter in presence of C. dactylon, I. sepiaria and E. odoratum
extracts in both decaying periods of 120 and 240 h, while plants were taller in extracts of
other weeds than control. These extracts also decreased the root length of tomato over the
control and C. dactylon, I. sepiaria and E. odoratum caused maximum reduction. Water
soluble allelochemicals extracted from Dittrichia viscosa inhibited the germination rate,
final germination (%) and radicle growth of Lycopersicon esculentum (169).
Chauhan et al. (24) investigated the effect of pollen of weed species on in-vitro
pollen germination and pollen tube growth of okra/lady’s finger (Abelmoschus esculentus
L.) and brinjal. The pollen of Cannabis sativa and Ricinus communis completely inhibited
the pollen germination (100% inhibition) in brinjal. The inhibition in pollen germination
and tube length of this crop in the presence of the pollens of weeds was very drastic. In
okra, neither Cannabis nor Ricinus weed pollen imposed any significant inhibitory
allelopathic effect on the pollen germination or pollen tube growth and time taken for
germination.
The results of in-vivo studies are similar to in-vitro results (24). The pollens of
both Cannabis sativa and Ricinus communis completely inhibited the germination of
brinjal. The pollen grains of brinjal failed to germinate on their own stigma, when their
stigma surface was artificially dusted with pollens and the pollen of any of the two weeds,
Cannabis sativa or Ricinus communis. The flowers of brinjal pollinated with the mixture
of the pollens of the weeds and its own pollen withered and fell-off within a week and
thus, no fruit-set was observed. Neither Cannabis sativa nor Ricinus communis showed
any pollen allelopathic effect on the pollen germination and tube length of the okra.
Alsaadawi and Salih (9) reported that root exudates of Cyperus rotundus
significantly reduced the root and shoot growth of tomato and cucumber plants, while, its
residues incorporated at 3 and 6 g per kg soil inhibited the seedling growth of cowpea
(Vigna unguiculata (L.) Walp.). The reduction increased with the increased rates of
residues. In soil, the toxicity of C. rotundus residues started 2 weeks after incorporation,
288 John et al
continued up to 8 weeks and then declined. Allelopathy was the causative factor for the
reduction in growth of test crops.
4.2.2. Cool season vegetable crops

In laboratory experiments, tomato, onion and cucumber were treated with 0.1, 1
and 10% solutions of ethanolic root extracts of Parthenium hysterophorus before
flowering (101). No effect was evident on germination, but radicle length was generally
reduced in all plants by the 10% extract. The 1% extract increased the radicle length of
tomato.
The effects of aqueous extracts from Mimosa bimucronata spring, summer and
autumn leaves and from green and mature fruits on seed germination and radicle growth
were tested using lettuce, carrot, cucumber, cabbage (Brassica oleracea L. var. capitata)
and tomato (81). Extracts from green or mature fruits did not inhibit germination but those
from green fruits inhibited radicle growth. Extracts from autumn (dry) leaves inhibited
germination in lettuce, carrots and tomatoes, and radicle growth was inhibited in all
species. The effects increased proportionally with the extract concentration.
Laboratory, greenhouse and field studies were conducted to determine the
allelopathic potential of Lantana camara and Cromolaena odorata on the germination and
growth of spinach (Spinacia oleracea L.), chinese cabbage, cucumber and chilly
(Capsicum frutescens) (155). Emergence and the dry weight were affected when L.
camara or C. odorata debris was present on the soil surface or incorporated into the soil.
Emergence and the dry weight of crops, except the emergence of spinach, were not
affected when crops were grown in soil samples collected from beneath field grown L.
camara. The dry weight of Chinese cabbage and Capsicum frutescens was reduced when
these crops were grown in C. odorata contaminated soil. The germination of Chinese
cabbage and Capsicum frutescens decreased progressively when exposed to increasing
concentration of aqueous L. camara extract. However, the L. camara extract at full
strength (66.7 g L-1) did not reduce the germination of spinach and cucumber seeds. C.
odorata extract, when applied at full strength to seeds of spinach, Chinese cabbages and
Capsicum frutescens, reduced germination by 10, 12, 21 and 19% of controls, respectively.
Full strength extracts of L. camara and C. odorata decreased seedling length and the FW
of all crops.
Dried shoot extracts of redroot pigweed significantly reduced germination of
cabbage seeds grown in Petri dishes. Extracts of nettle leaved goosefoot (Chenopodium
murale) inhibited germination of cabbage, carrot and pepper seeds (146). Shoot dry
weights of all crops were reduced with extracts of both weed species compared with the
control. The inhibitory effect was more pronounced at higher extract concentrations and
the crops differed in their sensitivity to weed extracts. Incorporation of dried shoot
residues of either weed species in soil mixtures severely reduced germination of cabbage,
carrot and cauliflower and aubergine (brinjal) seeds, and delayed germination of pepper
and tomato. Residues of C. murale delayed, while those of Amaranthus retroflexus
strongly inhibited germination of the cucumber. Cauliflower and tomato were the most
sensitive crop species to weed residues. Plants from direct sown seeds or transplants were
greatly affected by weed residues. The inhibitory effect was residue rate dependent.
Decayed residues of C. murale delayed germination of cauliflower and tomato but
increased their shoot dry weights, while those of A. retroflexus retained their toxicity on
tomato and significantly lowered shoot dry weight and leaf surface area of this crop.
Results showed that the inhibitory effects of weed residues on tomato could be overcome
by supplying more nutrients to the growth medium.
Ismail and Kumar (79) conducted laboratory and greenhouse experiments to
determine the effects of aqueous extracts and residues of Mikania micrantha decomposed
at different periods on the germination and growth of cucumber and Chinese cabbage.
They found that germination, radicle length and fresh weight of the 2 crops decreased
progressively when plants were exposed to increasing concentrations (12.5, 25 and 50
g/litre) of aqueous leaf or root extract of M. micrantha. The phytotoxic effect of root
extract on the germination and growth of both test crops was greater than that of the leaf
extract. The fresh weight and the rate of emergence of the 2 bioassay species decreased
with an increase of M. micrantha residues (leaf or root residues). The emergence of both
species was greatly reduced when M. micrantha debris (root or leaf) was incorporated into
the soil after 2 weeks of decomposition. The fresh weight of cucumber seedlings was
considerably reduced by exposure to root debris at both 1 and 2 weeks of decomposition,
whereas that of Chinese cabbage was reduced only when exposed to debris after 2 weeks
of decomposition.
Root exudates of Cymbopogon citratus, Ageratum conyzoides and Bidens pilosa
with or without treatment with XAD 4 resin were applied to seeds of radish and cucumber
(210). The exudate decreased germination rate, root length of seedlings and seedling
height of both the crops. Treatment with resin reduced the effect of the exudates. Exudates
of B. pilosa had the greatest effect.
The allelopathic potential of volatiles, foliage leachates, root exudates, and shoot
dried residues of white top (Cardaria draba) and Syrian sage (Salvia syriaca) on cabbage
cv. Pronzwik, carrot cv. Natus, cucumber cv. Beithalpha, squash cv. Byrouti, onion cv.
Texas Early Grana, pepper cv. Red Common and tomato cv. Special Back) was assessed
through different laboratory and glasshouse experiments (145). Volatiles from Syrian sage
fresh shoots reduced germination and inhibited seedling growth of most crops. Foliage
leachates or root exudates of both weeds were toxic to different crops under laboratory
conditions, with tomato and cabbage being most affected. In pot experiments, surface
placed shoot residues of both weeds significantly delayed seed germination and reduced
seedling growth of all crops with carrot, onion, and tomato being the most affected.
Decayed residues of white top were also toxic, but lesser than when fresh materials were
used. Foliage leachates or root exudates of both weed species added or released into the
soil mixture reduced seedling growth of cabbage and tomato.
The effect of different concentrations of underground organ and foliage extracts
of Cyperus rotundus were tested on germination of cucumber seeds (38). The underground
organ and foliage extracts (5%) of C. rotundus inhibited cucumber germination, reduced
radicle and plumule lengths, fresh and dry weights, contents of endogenous hormones
(auxins, gibberellins and cytokinins), altered the protein pattern of cucumber seedlings,
and increased the contents of endogenous inhibitors (ABA and phenols). The foliage
extract tended to be more potent. Chromatography fractionations and bioassays of growth
regulating substances lead to the isolation of many phenolic compounds (38).
290 John et al
Table 3. Inhibitory allelopathic effects of Ageratum conyzoides on vegetable crops in bioassays and
pot culture
Inhibitory effects on recipient

Allelochemical Recipient plant spp. References
spp.
Laboratory Bioassay
Volatile oil Cucumis sativus L. Fresh weight, root length and 93, 94
shoot height
Lycopersicon esculentum Mill. Fresh weight, root length and 93, 94
shoot height
Raphanus sativus L. Fresh weight, root length, shoot 68, 93, 94,
height and chlorophyll content 95
Aqueous Allium sativum L. Germination, fresh weight, root 67
extracts length and shoot height
Cucumis sativus L. Germination, fresh weight, root 67
length and shoot height
Lactuca sativa L. Germination, root length and 67
shoot height
Purified allelo- Cucumis sativus L. Germination, fresh weight, root 93, 95
chemicals Lycopersicon esculentum Mill. length and shoot height
Raphanus sativus L. Root length and shoot height of 93, 95,
seedling 110
Pot culture
Volatile oil Cucumis sativus L. Fresh weight, content of 92, 96
chlorophyll
Allelopathic activity of weeds (Amaranthus gracilis Desf., Convolvulus arvensis

L., Lactuca serriola L. and Portulaca oleracea L.) was studied on cabbage, carrot,
cucumber, onion, pepper, squash and tomato under glasshouse conditions (124). The root
exudates of weeds released into the soil had variable effects on different test crops. L.
serriola root exudates were most toxic to growth of different crops. Soil-incorporated dried
shoot residues, reduced the seedlings growth of the test crops and the degree of inhibition
was residue rate dependent. Soil-surface placed weed residues delayed seedling
emergence, but their effects on seedling growth was lower than soil-incorporated residues.
Decayed residues of different weeds inhibited the seedlings growth but their effect was
less as compared to un-decayed residues. In all experiments, roots were more sensitive
than shoots to allelopathic effects and both positive and negative effects were observed. C.
arvensis proved most harmful to all test crops, while cabbage, onion and tomato were most
affected than other crops.
The leaf leachates of Cassia uniflora weed stimulated the seed germination of
radish at low concentrations (2.5-5%), but higher concentrations (15-20%) inhibited both
seed germination and seedling growth (47). Application of either fresh or dry material of
Sicyos deppei weed decreased the relative growth rate, leaf area and dry weight of tomato
seedlings (130). Variation in growth was caused by the decreased net assimilation rate and
the physiological traits. These results linked with an imbalance of antioxidant enzyme
activity, increase in hydrogen peroxide and NADPH oxidase activity and an increase in
lipid peroxidation in leaves of tomato plants.
Laboratory and greenhouse assays revealed that aqueous whole plant extract from
Conyza canadesis inhibited germination, germination speed and seedling growth of
cucumber and radish (45). Radish was more sensitive to the extracts. The aqueous extract
decreased the photosynthesis activity but increased the malondialdehyde contents of
cucumber and radish.
4.3. Effects of vegetables on weeds

Some vegetables allelopathically inhibits the weeds. Cucumber accessions (256
nos.) from 41 countries were grown in pots with Brassica hirta and proso millet.
Cucumber variety 'PI211728' inhibited growth of the former species by 59 % and the latter
by 87 %, while 25 accessions inhibited growth by 50 % or more. The effect of leachates
collected from separate pots containing a nontoxic accession and two toxic accessions on
the growth of seedlings the weed species suggested that inhibition was due to allelopathy.
The possibility of incorporation of the allelopathic trait into plants can be considered to
enhance their competitive ability against weed species (144).
Lockerman and Putnam (105, 106) investigated the allelopathic potential of a
number of cucumber accessions on weeds. Cucumber wild accession PI 169391
suppressed adjacent weeds much more effectively than ‘Pioneer’ in the field. PI 169391
suppressed fresh weight and population of proso millet (Panicum miliaceum) two times
more than ‘Pioneer’. They suggested that introducing the allelopathic characters from wild
accessions to cultivated crops may be a useful approach to control weeds.
The volatiles from the youngest fully expanded leaves of 35 days old, field-grown
plants of turnip and kale (Brassica oleracea var. acephala L.) slightly inhibited the
germination of barnyard grass [Echinochloa crusgalli (L.) Beuv.] (128).
Radish has the potential to control summer weeds like Sorghum halepense,
Cynodon dactylon, Amaranthus retroflexus, Portulaca oleraceae and Xanthium.
strumarium that cause yield losses in maize and cotton fields (187). Radish successfully
controls Sorghum halepense in cotton fields, when grown in rotation with cotton (186).
When watermelon seeds were cultured in a Petri dish together with amaranthus
(Amaranthus retroflexus), barnyard grass, cockscomb (Amaranthus caudatus), lettuce or
tomato seeds, the shoot growth of amaranth and cockscomb was markedly promoted,
whereas the shoot growth of lettuces and tomatoes was inhibited (100). The shoot growth
of barnyard grass was not affected. The results suggest that plant selective allelopathic
substance(s) affecting the shoot growth of other plant seedlings were exuded from
watermelon seeds. An allelopathic substance was isolated from the exudates of
germinating watermelon seeds and identified as vanillic acid. Vanillic acid was found to
promote the shoot growth of cockscomb at concentrations of 10 to 300 ppm and that of
amaranth at concentrations of 3-30 ppm, although the shoot growth of amaranth was
inhibited by 300 ppm of vanillic acid. The shoot growth of lettuces and tomatoes was
inhibited at concentrations higher than 30 mg/litre by vanillic acid. However, the shoot
growth of barnyard grass was not affected at the concentrations used. All these results
suggest that vanillic may be the allelopathic substance in exudates of germinating
watermelon seeds (100).
Chilli root, shoot and rhizosphere soil (fresh) leachates (10, 20 and 30 %)
decreased the germination of Amaranths viridis L. and Trianthema portulacastrum L
(161). As the concentration of the leachate increased, the magnitude of inhibition of
292 John et al
germination also increased. The lower concentration (10%) did not affect germination. The
leachates also reduced the seedling growth of the weeds. The inhibition was more
pronounced at higher concentrations.
Silva et al. (160) reported a concentration dependent inhibition of Portulaca
oleracea germination by the juice of ground roots of radish. The aqueous extracts of radish
(33, 50, 60, 100 %) reduced the germination and rhizome development of Sorghum
halepense, Cynodon dactylon, Amaranthus retroflexus, Portulaca oleraceae, Xanthium
strumarium (36).
An allelopathic vegetable can potentially be used to control weeds by planting a
variety with allelopathic qualities, either as a smother crop, in a rotational sequence, or
when left as a residue or mulch, especially in low-till systems, to control subsequent weed
growth. Alternatively, application of allelopathic vegetable residues before, along with, or
after synthetic herbicides could increase the overall effect of both materials, thereby
reducing application rates of synthetic herbicides.
4.4. Allelopathic effects of trees on vegetable crops (Agroforestry)

In multi-storey cropping systems, where numerous crops and trees are grown
together, vegetables are essential components. The multipurpose trees comprise a virtually
untapped reservoir of allelochemicals. The allelopathic effects of several trees on crops has
been reported (Table 4).
Jacob et al. (80) conducted laboratory bioassays and pot culture experiments to
assess the allelopathic influence of leaf leachates of multipurpose trees (Artocarpus
heterophyllus, Mangifera indica, Ailanthus triphysa, Anacardium occidentale, Tamarindus
indica, Tectona grandis, Thespesia populnea, Casuarina equisetifolia, Gliricidia sepium,
Strychnos nux-vomica) planted in the multi-storey home gardens of Kerala, on cowpea,
bitter gourd (Momordica charantia L.) and brinjal. The nature and degree of allelopathic
effects of trees varied with crop species (Table 5). Tamarind, teak, casuarina and strychnos
caused severe allelopathic inhibition of cowpea, hence were incompatible. Gliricidia,
portia, tamarind, teak and cashew were incompatible with brinjal. Ailanthus, cashew,
strychnos, mango, portia, tamarind and teak were detrimental to bitter gourd. In general,
the inhibitory effects were most prominent in brinjal than cowpea and bitter gourd and
cannot be overcome with dilution. The findings will help in differential selection of
vegetables, for combining with trees in agroforestry systems.
A green house pot experiment assessed the allelopathic effects of Acacia nilotica
leaves on the growth and metabolic activities of 45-day-old pea plants (190). The lower
doses of Acacia leaf residue (0.25 and 0.5% w/w) stimulated the growth of pea shoot and
root, but the higher doses (0.75, 1.0, 1.5 and 2%, w/w) inhibited seedling growth and the
effect was concentration dependent. The total phenolic content of pea shoots (particularly
phenolic glycosides), increased at lower doses of Acacia residue and decreased with higher
ones. Chlorophylls a, b and carotenoids accumulated in pea shoot at lower doses of Acacia
leaf residues, accompanied by accumulation of total sugar, mainly the insoluble fraction.
On the other hand, the inhibition in the contents of photosynthetic pigments at higher doses
of Acacia residues was paralleled by significant reduction in all sugar fractions.
294 John et al
Table 5. Effect of fresh leaf leachate (1:10 w/v) of trees on vegetable crops (Pot culture)
Treatment Plant No. of DW per plant (g).

height leaves Pod Seed Root
(cm) MAP MAP
COWPEA (Vigna unguiculata L.)
Ailanthus triphysa (Dennst)Alston 65.97 21.93 12.33 3.78 1.28
Anacardium occidentale L. 60.17 23.67 11.75 3.11 1.38
Casuarina equisettifolia J.R. 35.53 16.00 9.30 2.11 1.30
Gliricidia sepium Jacq Walp. 57.60 21.67 7.71 3.11 0.84
Artocarpus heterophyllus Lamk. 57.53 20.27 8.66 2.56 1.64
Strychnos nuxvomica L. 36.27 23.00 8.01 2.33 1.51
Mangifera indica L. 58.00 20.33 11.25 2.89 1.56
Thespesia populnea (L.) Soland 52.50 22.93 8.56 2.56 1.63
Tamarindus indicus L. 0.00 0 0 0 0
Tectona grandis L.f. 0.00 0 0 0 0
Control 66.97 24.00 12.29 3.56 1.70
CD (0.05) 19.445 4.238 2.959 1.003 0.313
Treatment Plant height (cm) No. of leaves Weight (g) per

plant
1MAP 2 MAP 1MAP 2 MAP Fruit Root
FW DW
BITTER GOURD (Momordica charantia L.)
Ailanthus triphysa (Dennst)Alston 40.10 148.00 11.20 30.67 453.67 3.78
Anacardium occidentale L. 39.63 168.67 12.83 30.00 522.10 3.71
Casuarina equisettifolia J.R. 30.30 200.33 10.17 36.00 624.33 4.10
Gliricidia sepium Jacq Walp. 40.40 194.67 9.87 37.00 628.4 3.93
Artocarpus heterophyllus Lamk. 39.43 186.67 11.63 31.00 613.97 3.40
Strychnos nuxvomica L. 32.53 138.33 8.53 32.33 432.57 1.79
Mangifera indica L. 43.73 186.33 11.53 36.00 563.33 2.73
Thespesia populnea (L.) Soland 48.50 216.00 12.43 40.67 543.70 1.41
Tamarindus indicus L. 25.83 157.33 7.97 38.67 472.63 3.10
Tectona grandis L.f. 31.07 150.33 10.70 31.67 453.67 3.37
Control 49.83 226.33 11.87 43.33 625.33 3.81
CD (0.05) 11.160 43.889 2.517 6.831 41.036 0.487
EGG PLANT (Solanum melongena L.)
Ailanthus triphysa (Dennst)Alston 15.00 41.33 7.00 21.33 317.64 3.03
Anacardium occidentale L. 11.00 38.33 7.00 15.33 260.39 1.94
Casuarina equisettifolia J.R. 13.67 42.00 8.33 21.33 329.98 3.72
Gliricidia sepium Jacq Walp. 0 0 0 0 0 0
Artocarpus heterophyllus Lamk. 13.00 35.67 6.33 23.33 303.77 3.41
Strychnos nuxvomica L. 15.67 35.33 6.33 24.00 291.37 2.10
Mangifera indica L. 14.67 32.33 6.67 23.00 314.05 3.89
Thespesia populnea (L.) Soland 0 0 0 0 0 0
Tamarindus indicus L. 10.33 27.00 6.33 16.67 248.43 1.93
Tectona grandis L.f. 10.67 37.00 6.67 19.00 255.90 1.67
Control 15.07 47.67 7.33 24.33 308.29 4.82
CD (0.05) 2.498 8.005 NS 4.715 46.433 1.193
MAP: Months after planting; DW: Dry weight; FW Fresh weight.
296 John et al
Eucalyptus: Eucalyptus tereticornis leaf extracts significantly decreased the germination,

shoot and root length of okra (Abelmoschus esculentus L), than bark and root extracts. The
inhibitory effects of extracts were concentration dependent. Leaf extract of Eucalyptus
proved most detrimental to seedling vigour of okra than the bark and root (6). The aqueous
extract of dry leaves of Eucalyptus camaldulensis Dehn. inhibited the germination, vigour
and growth of seedlings of cucumber and radish than green leaves (1). Pina et al. (135)
studied the influence of leaf extracts of Eucalyptus dysentrica on sesame (Sesamum
indicum L.) and radish. Their study also evaluated the effects of fresh and dry leaf extracts
and the extract pH (4.7-7.0) on target species. The extract did not influence the
germination but drastically reduced the seedling growth. The roots were more affected by
the extracts than shoots. Dry-leaf extracts were more inhibitory to sesame and radish
seedlings than fresh leaf extracts. The extract pH did not influence the allelopathic
response of target species. The extracts resulted in the abnormal growth of seedlings viz.,
shorter roots, root tissue darkening, early lateral root development, less number of root
hairs and lateral roots and altered gravitropic response. The inhibitory effects of leaves
extracts on seedling growth were more intense, when soil was substrate. The aqueous leaf
extracts of E. dysenterica at concentration as low as 1% inhibited the initial growth, root
hair and lateral root differentiation and impaired gravitropic responses of sesame and
radish seedlings.
Gatti et al. (46) reported the allelopathic potential of aqueous extracts of the leaf,
stem and root of Ocotea odorifera on the germination and growth of lettuce and radish.
The leaf extract of Ocotea odorifera drastically reduced the seed germination in lettuce
and radish, whereas, the stem and root extracts delayed their germination. The roots of
vegetables were the most sensitive to allelochemicals.
Aqueous extracts of fresh leaves of five mangrove tree spp. (Avicennia marina
(Forsk.) Vierh., Aegiceras corniculata (L.) Blanco, Kandelia candel (L.) Druce,
Rhizophora stylosa Griff. and Bruguiera gymnorrhiza (L.) Lam.) reduced the germination
rate and seedling growth of cabbage (25). The allelopathic intensity of aqueous extracts on
cabbage increased with the development of mangrove succession.
5. PESTICIDAL POTENTIAL OF VEGETABLE CROPS
5.1. Fungitoxic/antibiotic effects

Several vegetables possess antimicrobial principles and hence, allelopathically
inhibit phytopathogenic fungi and bacteria. Extraneous application of plants or its
constituents controls several fungal pathogens (Table 6).
The phytochemical α-tomatine is a steroidal glycoalkaloid found in tissues of
members of the genus Lycopersicon and has been shown to exhibit antibiotic activity
against a wide range of organisms. Studies have revealed that this secondary plant
compound is toxic to many microorganisms (14,78,158). Tomatine and tomatidine (0.3
mM) when incorporated into growth media (agar), inhibited the growth of three fungal
bioherbicidal phytopathogens. Alternaria cassiae, the most sensitive pathogen was
inhibited 70% by both compounds (42). Tomatine inhibited the growth of Colletotrichum
truncatum and Fusarium subglutinasns (produces fumonisin, phytotoxic to many plants)
Table 6. Vegetable crops with fungicidal activity
Vegetable Pathogens affected Product Reference

tested
Daucus carota L. Aspergillus sp, Curvularia sp., Penicillium sp. Oil 51
Momordica Dreschlera oryzae, Pyricularia oryzae, SAE, EE 177
charantia L. Rhizoctonia solani
Allium sativum L. Dreschlera oryzae, Pyricularia oryzae, SAE, EE 177
Rhizoctonia solani
Allium cepa L. Alternaria alternata, Asperigillus niger, Botrytis Oil 82, 113,
allii, Chrysoporium tropicum, Claviceps 161
purpurea, Kerratinophyton terreum, Malbranchea
pulchella, Microsporum gypseum, Phytophthora
infestans, Verticillium tenuipes
Capsicum annum L. Aspergillus niger Oil 22
SAE: Steamed aqueous extract, EE: Ethanolic extract
by 63% and 50%, respectively; while tomatidine inhibited the growth of these latter two
pathogens by 50% and 15%, respectively. These natural plant products have broad range
phytotoxicity and fungitoxicity, which may be important in plant defence mechanisms. α-
Tomatine is a saponin (steroidal glycol-alkaloid) produced by tomato and some other
Solanum species (153). α-Tomatine consists of a branched tetrasaccharide (β-D-
glycopyranosyl-(1→2)- (β-D- xylopyranosyl- (1→3)–β-D-glucopyranosyl- (1→4)-D-
galactose), attached to 0-3 of the steroidal aglycone, tomatidine. This tetrasaccharide,
called lycotetraose, occurs in solanaceous species: tomatoes and potatoes etc.(42).
Tomatine accumulates in plant stems, leaves, and roots and is fungitoxic to some plant
pathogens (63).
Ajoene: Ajoene, a compound derived from the garlic (Allium sativum L.) and produced by
chemical synthesis, was investigated in vitro and in vivo for its activity against 20
phytopathogenic and epiphytic fungi and bacteria (149). The main interest was focussed on
the fungi which belonged to the group of obligate biotrophic parasites (powdery mildews),
other leaf pathogens, soil-borne pathogens, vascular wilt fungi, and yeasts. The minimum
inhibitory concentrations (in vitro tests), ranged from 2 to 200 mg L-1 depending on the
organism, method, and nutrient medium used. The phytopathogenic fungi Cladosporium
fulvum and Verticillium dabliae and the phytopathogenic bacterium Erwinia amylovora
were the most sensitive species. In greenhouse experiments, the inhibiting action of ajoene
against Cladosporium fulvum could be confirmed with tomato plants after protective
treatment. However, only powdery mildew of tomatoes (Oidium lycopersicum) and roses
(Sphaerotheca pannosa var. rosae) could be inhibited completely.
Aqueous and ethanolic extracts of seeds of radish, cauliflower and cress were
tested for allelopathic effects on Rhizoctonia solani in vitro. The tested extracts were
inhibitory to R. solani. The application of powdered seeds of radish to the soil infested
with R. solani reduced the damping off in cotton seedlings (40).
Two compounds, separated chromatographically from extracts of potato peel and
from extracts of pulp tissue, were found fungistatic to Helminthosporium carbonum and
reduced its growth (98). These compounds were identical to chlorogenic and caffeic acid
298 John et al
in physical and chemical properties. It appears these acids are associated with the
immunity of white potatoes to attack of H. carbonum.
5.2. Nematicidal effects

Certain vegetables possess nematicidal principles and therefore, offer immense
potential for nematode control in vegetables cultivation (Table 7). Tomato contains α-
tomatine effective against Panagrellus redivivus (4). Potato contains £-chaconine effective
against Meloidogyne incognita (5). Asparagus contains glucoside asparagusic acid
effective against Trichodorus christie (131).
Table 7. Vegetables with nematicidal activity
Name of plant Nematicidal Nematode test species Reference

plant part
Allium fistulosum L. Bulb Meloidogyne incognita 172
A. sativum L. Bulb Meloidogyne incognita 169
Amaranthus gracilis Desf. expoir Leaf Meloidogyne javanica 121
Asparagus sp. Root Trichodorus christie 130
Capsicum annum L. Pod Meloidogyne incognita 169
Momordica charantia L. Seed Meloidogyne incognita 77
Raphanus sativus L. Cover crop Pratylenchus 62
Tylenchorhynchus
Trichosanthes anguina L. Seed Meloidogyne incognita 77
α-tomatine and α-solanine are major glycoalkaloids biosynthesized in plants of

Solanaceae family, for plant defence against phytopathogens. They are also plasma
cholinesterase inhibitors and teratogens. Using liquid chromatography, their levels in 15
tomato cultivars were estimated in response to root-knot nematode (Meloidogyne
incognita) attack at an infestation level of 2 juveniles. α-tomatine content was found much
higher in roots and shoots of resistant over the susceptible cultivars (116). In roots, the
concentration of α-tomatine was maximum (1773 mg kg-1) in PAU-8 among the resistant
cultivars versus 356 mg kg-1 in susceptible cultivar Hybrid-2. In shoots, the α-tomatine
level ranged from 710 mg kg-1 in Hybrid -1 (susceptible) to 5530 mg kg-1 in Rutgers
(resistant) cultivars. The levels of both α-tomatine and α-solanine decreased with time.
However at fruiting, the level increased in green fruit to maximum of 877 mg kg-1 in
cultivar PAU-9 as against 276 mg kg-1 in Hybrid -2 and declined rapidly in yellow and red
fruits. α-tomatine content in roots could be correlated to reduction in population of M.
incognita in soil and the hazard indices measured in terms of galling of tomato roots. The
studies revealed that α-tomatine contents in roots of tomatoes may prove an excellent
marker for resistance against M. incognita.
The application of wheat flour at 5% rhizospheric soil drench to tomato cv. Pusa
Ruby and okra cv. Purbani Kranti plants (24 h after inoculation of roots with Meloidogyne
incognita juveniles), reduced the infestation (99). Wheat flour, α-amylase and α-amylase
inhibitor did not kill the nematodes in in vitro test, but decreased the α-amylase activity in
inoculated roots of tomato and okra. α-amylase inhibitor applied on tomato and okra leaves
(inoculated with M. incognita), ameliorated the root-knot. Both α- amylase and α-amylase
inhibitors enhanced the enzyme activity in inoculated roots than in non-inoculated roots of
tomato and okra plants.
6. ALLELOCHEMICAL STUDIES IN VEGETABLES
6.1. Cucumber
Holappa and Blum (65) assessed the relative sensitivity of cucumber to
exogenously applied concentrations of ferulic acid. Ferulic acid inhibited leaf growth and
water utilization of cucumber. Increased endogenous abscisic levels were found in
cucumber following ferulic acid treatment.
p-Thiocyanatophenol was extracted from the root exudates of hydroponically
grown cucumber in a column filled with Amberlite XAD-4 resin, through which the
nutrient solution was constantly recycled. In bioassays, p-thiocyanatophenol markedly
retarded the root elongation of lettuce seedlings and the nutrient ion uptake of cucumber
seedlings (204).
Lehman et al. (102) reported that ferulic acid was more inhibitory to cucumber
leaf expansion than p-coumaric acid. If present together, the effects of ferulic and p-
coumaric acids on leaf expansion are additive.
Activated charcoal added to the nutrient solution of growing cucumber cv. PI
169319 seedlings when desorbed was found to contain four compounds viz., benzoic, p-
hydroxybenzoic, 2,4 dichlorobenzoic and phthallic acids (15). These compounds were
used in a cucumber bioassay at 0, 2, 10 and 20 µl L-1. Only 2,4 dichlorobenzoic acid at 20
µl L-1 inhibited seedling growth, while phthallic acid stimulated its growth at 2 µl L-1.
Hence, 2,4 dichlorobenzoic acid may be responsible for growth reduction of cucumbers in
hydroponic solution. In another bioassay, 2,4 dichlorobenzoic acid at 20 µl L-1 was added
in mixtures with all other compounds at 10 µl L-1, and the growth inhibition of cucumber
seedlings was found to be ameliorated.
Terzi et al. (178) investigated the effects of juglone on the growth of cucumber
cv. Beith Alpha) seedlings with respect to physiological and anatomical parameters.
Growth parameters (seedling elongation, and fresh and dry weights) were reduced by 1
mM juglone. Juglone also reduced chlorophyll a and b contents, and some anatomical
tissues (xylum vessel and bundle radius of stem, stomatal length, and number of stomata
on cotyledons). The anatomical changes in stem and cotyledon were related to the growth-
inhibiting effect of juglone. On the other hand, catecholase and tyrosinase (catechol
oxidase) activities also increased due to juglone application.
The effects of exogenous cinnamic acids were studied on cucumber growth and
physiological characteristics at seedling stage (195). The soils used were from a
greenhouse, where cucumbers were continuously planted for 18 years and an open field
where no cucumbers were planted, respectively. Seedling growth, contents of
photosynthetic pigment, root activities and H+-ATPase activities of root membrane were
inhibited by cinnamic acids. Inhibitions on growth and root activities of seedlings were
significantly different between the greenhouse soil treated with 100 mg kg-1 and open field
soil treated with 200 mg kg-1.
The dynamics of cinnamic acid uptake in cucumber plants and its residue in soil
were studied. The cucumber ‘Changchun Mici’ was grown in pots and cinnamic acid was
300 John et al
applied at 0, 25, 50, 100 and 200 mg kg-1 soil. The application of cinnamic acid increased
its content in cucumber plants and in soil with its increasing dose. The retention rate (in
cucumber and in soil) decreased with high doses. The rate of allelochemicals addition and
soil retention probably controls the allelochemicals content in soil. The allelochemicals
content in soil was the result of interactions among the plant and soil microbes (197).
Cucumber seedlings were subjected to allelochemical stress by treating their roots
with 0.5 mM solutions of ferulic and p-coumaric acids (139). Production of ethylene by
cucumber seedlings, the increase in phenylalanine ammonia-lyase activity and lignin
synthesis as well as the reduction in growth indices of cucumber roots were observed. The
results indicated that ethylene participates in the retardation of cucumber root growth by
these phenolic compounds.
Cucumber seeds were treated with solutions of ferulic and p-coumaric acids. The
phenolic acids changed the activity of hydrolytic enzymes and impeded the seeds
germination. Their effects depended on the plant species, enzyme type, concentration of
applied phenolic acid and stage of germination (137).
6.2. Other vegetable crops

Alsaadawi et al. (10) conducted a study to test whether interference of
chlorophyll metabolism and ion uptake may be mechanisms through which some phenolic
acids inhibit the growth of cowpea seedlings. Three concentrations (10-4 M, 5 x 10-4 M and
10-3 M) of each of syringic, caffeic, and protocatechuic acids were used in sand-culture
medium. It was found that seedling growth, chlorophyll a, total chlorophyll, chlorophyll
a/b ratio, and the uptake of N, P, K, Fe and Mo were significantly reduced by most of the
test concentrations of the phenolic acids. However, chlorophyll b content and the Mg
uptake were not significantly affected by all the phenolic acid concentrations. Calcium
uptake was significantly inhibited by 5 x 10-4 M and 10-3 M of caffeic acid and 5 x 10-4 M
of protocatechuic acid. In most cases, the reduction in dry weight was parallel to the
reduction in chlorophyll content and ion uptake, and the reduction in chlorophyll was also
parallel to the reduction in ion uptake.
Phenolic compounds such as gallic acid, ferulic acid, p-hydroxybenzoic acid,
vanillic acid, salicylic acid, tannic acid and hydroquinone were identified from the aqueous
extracts and volatile substances of tomato plant (90). The seed germination and seedling
growth of the lettuce and egg plant was severely inhibited by 5 x 10-3 Molar of phenolic
reagents identical to those identified from tomato plant. Germination and growth rate of 5
x 10-4 M and 5 x10-5 M were higher than that of 5 x 10-3 M of phenolic compounds would
be assumed to be threshold concentration for allelopathic effects.
The inhibitory effect of the glycoalkaloids chaconine, solanine, tomatine,
solasonine and solamargine and of the aglycones solanidine, solasodine and tomatidine
(200 and 400 µg ml-1) on lettuce seed radicle elongation were determined. Solasodine and
tomatidine had negligible effects on the elongation, while solanidine produced over 50 %
of the total measured inhibition (48).
A methanol:water extract of tomato apices increased the growth of the alga
Chlamydomonas reinhardtii. The active substance from the dried shoot apices was purified
by C18 flash column and high performance liquid chromatography. The purified extract
enhanced the growth of tomato, corn and rice seedlings at concentrations less than 1.0
ppm. Nuclear magnetic resonance and mass spectroscopy indicated that the purified
fraction was a mixture of compounds having sugar moieties. Analysis by thin layer
chromatography showed that the fraction was ninhydrin positive and more polar than the
known plant hormone studied (191).
Holappa and Blum (65) assessed the sensitivities of tomato and bean to
exogenously applied concentrations of ferulic acid. Ferulic acid inhibited leaf growth and
water utilization of wild type tomato and flacca tomato, but not of bean. Increased
endogenous abscisic levels were found in wild type tomato and flacca tomato subsequent
to ferulic acid treatment.
Allelochemicals alpha-pinene, borneol, chlorogenic-acid, coumarin, scopoletin,
limonene, terpinolene in roots of carrot; α-pinene, chlorogenic-acid, cinnamaldehyde, o-
cresol, quercetin, rutin, salicylaldehyde in fruit of tomato; and alpha-pinene, chlorogenic-
acid, limonene, scopoletin, terpinolene (fruit) in chilly have been identified (37).
Thirteen natural and synthetic phenylpropanoids as well as coumarin were tested
for their biological activity on radish germination and subsequent root growth in light and
darkness (3). Coumarin was the most potent inhibitor. Coumarin was formed
spontaneously by photooxidation of 2-hydroxycinnamic acid. Microscopic observation of
root treated with coumarin suggest that this substance inhibits the elongation of cells of the
differentiating zone of the root.
Sterile root exudates from carrot seedlings stimulate the hyphal development of
Gigaspora margarita, a vesicular-arbuscular mycorrhizal fungus, in the presence of
optimal CO2 enrichment (142). Three flavonols (quercetin, kaempferol, rutin or quercetin
3-rutinoside) and two flavones (apigenin, luteolin) were identified in carrot root exudates.
Flavonols like quercetin and kaempferol are known to have stimulatory effects on hyphal
growth of G.margarita.
The influence of carrot seed oil and its major components (caryophyllene and
carotol) on the growth and germination of cress and carrot was studied in two independent
tests, viz., (i) the action of vapours of volatile substances and (ii) the action if applied
uniformly dispersed in water (83). The oil and its components exhibited phytotoxic
properties regardless of the mode of application and inhibited the germination of test plants
at the highest concentrations. Their post emergence influence on plant growth was even
more pronounced than that observed in seed germination.
Capsaicin (from chilli) inhibited germination, roots and shoots of alfalfa, cress,
lettuce, crabgrass (Digitaria sanguinalis), timothy (Phleum pratense) and ryegrass (85).
Increasing the dose of capsaicin increased the inhibition. The concentrations for 50 %
inhibition of the root growth were 2.7, 0.32, 2.1, 0.27, 0.29 and 0.57 mM for alfalfa, cress,
lettuce, crabgrass, timothy and ryegrass, respectively. The concentrations for 50 %
inhibition of shoot growth were 17, 0.87, 6.7, 2.3, 1.4 and 6.2 mM for alfalfa, cress,
lettuce, crabgrass, timothy and ryegrass, respectively. Germination was inhibited by 50 %
at 82, 88, 68. 4.8, 22 and 11 mM concentrations of alfalfa, cress, lettuce, crabgrass,
timothy and ryegrass, respectively. Thus, effectiveness of capsaicin on the plant growth
differed with species and targets, and suggests that capsaicin may act as an allelochemical
to other plants.
The volatile extracts of essential oils of cinnamon (Cinnamomum zeylanicum
Blume), alecrim-pimenta, capim-citronella and alfavaca-cravo revealed allelopathic
potentialities on lettuce seed germination and radicle growth. The effect varied according
302 John et al
to the oil concentration (11). The volatile extract of essential oil stimulated radicle growth
and did not inhibit lettuce seeds germination.
The phytotoxic activity of the compounds isolated from sugarcane straw was
evaluated on seedling growth of lettuce (156). Three compounds were identified: ferulic
(FA), syringic (SA) and vanillic (VA) acids. VA drastically inhibited the root elongation,
followed by FA and SA. These phytotoxins increased the root membrane permeability and
depressed the root metabolic activity in lettuce. VA and FA inhibited the mitotic index,
while, FA and SA stimulated the proliferation of secondary root.
Pea seeds were treated with solutions of ferulic and p-coumaric acids. The
phenolic acids changed the activity of hydrolytic enzymes and impeded the seeds
germination. Their effects depended on the plant species, enzyme type, concentration of
applied phenolic acid and stage of germination (137).
Hoagland (63) conducted studies to examine the phytotoxic and antibiotic effects
of tomatine and tomatidine in a variety of plant species (including weeds). α-tomatine was
not highly phytotoxic and slightly inhibited the stem elongation (7 to 13%) when applied
as a spray to etiolated 4-d-old seedlings of sesbania (Sesbania exaltata (Raf) Rybd),
sicklepod (Senna obtusifolia L.), mungbean, wheat and sorghum (Sorghum vulgare L.).
Tomatidine had less effect on stem elongation than tomatine and reduced the elongation (5
to 10%). Tomatine was more effective than tomatidine in reducing chlorophyll content in
excised etiolated tissues of sesbania, sicklepod, mungbean, wheat and sorghum. Inhibition
of chlorophyll accumulation by tomatine ranged from 16 to 89% of control values,
whereas inhibition by tomatidine ranged from 0 to 30 % of control in these species. Both
tomatine and tomatidine increased electrolyte leakage of corn, kudzu (Pueraria lobata
(Willd.) Ohwi), palmleaf morning glory (Ipomea wrightii Gray) and wild senna (Cassia
marilandica L) leaf disks at 24 to 72 h after exposure to the compounds (0.5 mM) and
light (100 µE m2 s-1). Tomatidine caused greater electrolyte leakage than tomatine in
tissues of these species.
7. FUTURE RESEARCH PROSPECTS
The thrust areas for future allelopathic research are as under:

i. Separate the allelopathic interference from competition in vegetable fields and
vegetables based cropping systems.
ii. Screen the germplasm/varieties of vegetables for allelopathic potential and later on do
genetic manipulations to breed new varities.
iii. Exploit the vegetables allelopathy for weed control and plant protection.
iv. Determine the critical concentrations of allelochemicals of each vegetable needed to
express their inhibitory/ stimulatory influences.
v. Identify the compatible and beneficial associations of vegetables with other crops,
vegetables and trees.
vi. Determine the harmful and beneficial effects of allelopathy through detailed pot
culture and field studies.
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Role of Allelopathy in vegetables crops production

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Role of Allelopathy in vegetables crops production

Cropping Systems Research Centre, P.O. Karamana

(Received in revised form: January 21, 2010)

Keywords: Agroforestry, allelopathy, allelochemicals, autotoxicity, crop, nematode,

The term allelopathy includes both harmful and beneficial biochemical

Table 1. Vegetable crops affected by soil sickness and autotoxicity

2.1. Tomato (Lycopersicon esculentum Mill.)

Soil amendment Effects on test plant References

2.2. Cucumber (Cucumis sativus L.)

2.3. Eggplant (Solanum melongena L.)

Soil sickness is complex phenomenon and of the several factors involved,

exudates, biomass decomposition and volatilization (18,148). Many autotoxins [simple

3.1. Cucumber (Cucumis sativus L.)

3.2. Tomato (Lycopersicon esculentum Mill.)

3.3. Pea (Pisum sativum L.)

3.4. Watermelon (Citrullus lanatus (Thunb.) Mansf.)

3.5. Red pepper (Capsicum annuum L.)

3.6. Brinjal (Solanum melongena L.)

4.1. Vegetable crops and their interactions

4.1.1. Effects of Vegetable crops on other crops

4.1.2. Effects of other crops on vegetable crops

Water leachates from guayule (Parthenium argentatum) processing residues, or

4.1.3. Effects of Vegetable crops on vegetable crops

4.2. Vegetables and weeds interactions

4.2.1. Summer season vegetable crops

Almodovar et al. (7, 8) determined the allelopathic effects of root exudates of

4.2.2. Cool season vegetable crops

Inhibitory effects on recipient

Allelopathic activity of weeds (Amaranthus gracilis Desf., Convolvulus arvensis

4.3. Effects of vegetables on weeds

4.4. Allelopathic effects of trees on vegetable crops (Agroforestry)

Treatment Plant No. of DW per plant (g).

Treatment Plant height (cm) No. of leaves Weight (g) per

Eucalyptus: Eucalyptus tereticornis leaf extracts significantly decreased the germination,

5. PESTICIDAL POTENTIAL OF VEGETABLE CROPS

5.1. Fungitoxic/antibiotic effects

Table 6. Vegetable crops with fungicidal activity

Vegetable Pathogens affected Product Reference

5.2. Nematicidal effects

Table 7. Vegetables with nematicidal activity

Name of plant Nematicidal Nematode test species Reference

α-tomatine and α-solanine are major glycoalkaloids biosynthesized in plants of

6. ALLELOCHEMICAL STUDIES IN VEGETABLES

6.2. Other vegetable crops

7. FUTURE RESEARCH PROSPECTS

The thrust areas for future allelopathic research are as under:

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