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Allelopathic Interactions in Agroforestry Systems

S. J. H. Rizvi; M. Tahir; V. Rizvi; R. K. Kohli; A. Ansari
Online publication date: 24 June 2010
To cite this Article Rizvi, S. J. H. , Tahir, M. , Rizvi, V. , Kohli, R. K. and Ansari, A.(1999) 'Allelopathic Interactions in
Agroforestry Systems', Critical Reviews in Plant Sciences, 18: 6, 773 — 796
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Critical Reviews in Plant Sciences, 18(6):773–796 (1999)
Allelopathic Interactions in Agroforestry Systems

S. J. H. Rizvi,1,2* M. Tahir,3 V. Rizvi,1,2 R. K. Kohli,4 and A. Ansari 1
1Plant Pests and Diseases Research Institute, P.O. Box 19395–1454, Evin, Tabnak Av., Tehran, Iran;
2Permanent address: Faculty of Basic Sciences, Rajendra Agricultural University, Pusa-848125, Samastipur,
India; 3International Center for Agricultural Research in Dry Areas (ICARDA) Regional Office, Agricultural
Research, Education and Extension Organization, P.O. Box 19835–111, Tehran, Iran; 4Department of Botany,
Panjab University, Chandigarh, India
* Corresponding author.
ABSTRACT: Agroforestry is a modern tool to develop sustainable land use and to increase food production by
growing woody species (trees, shrubs, palms, bamboos, etc.) with agricultural crops and/or animals in some form
of spatial arrangement or temporal sequence. Because these species co-exist with the agricultural crops, their
allelopathic compatibility may be crucial to determine the success of an agroforestry system. A survey of the
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available information reveals that most of the agroforestry species (AF species) have negative allelopathic effects
on food and fodder crops. Therefore, it is desirable to do further research in this direction so that AF species with
no or positive allelopathic effects on the companion crops may be promoted for agroforestry programs. As AF
species remain a part of the agroecosystem for a longer period, and most of them produce a large amount of leaves
and litter, their allelochemicals may play an important role in developing an eco-friendly pest management
strategy. Besides these generally studied aspects of allelopathy, some comparatively newer aspects of research
have been identified, such as evaluation of qualitative yield of agroforestry systems, selective behavior of the
allelochemicals, effect on soil quality, and the role of tree allelochemicals in animal and human nutrition. If given
due consideration, allelopathy could play a pivotal role in conservation of the highly threatened environment,
biodiversity, natural resource base, and making agriculture more sustainable through broadening the scope of
agroforestry.
KEY WORDS: agroforestry, allelochemicals, allelopathy, eco-friendly pest control, qualitative yield.
I. INTRODUCTION tude and the rate of the world population growth,

hunger, and worldwide environmental degrada-
Agroforestry is a sustainable land manage- tion. Unfortunately, by the end of nineteenth cen-
ment system that increases the yield of the land, tury the establishment of forest plantations had
combines the production of crops (including tree become the dominant objective of agroforestry.
crops) and forest plants, and/or animals simulta- Landless laborers were employed with a provi-
neously or sequentially on the same unit of land, sion that they can cultivate the land left between
and applies management practices that are com- the rows of forest trees and use the agricultural
patible with the cultural practices of the local products. However, the priority for forest trees
population (Bene et al., 1977; King and Chandler, always controlled the agricultural operations in a
1978). It is evident that the pioneers in way that the interests of forestry should not be
“agroforestry” considered that the ultimate objec- compromised (King, 1968). This philosophy cre-
tive of the system was not tree production but ated a serious problem especially in the less-de-
food production in totality. Further, development veloped countries because development of forest
of the agroforestry system was considered a means and forest industry debarred the local people from
of sustainable land use to cope with the magni- basic forest products. Thus, the most important
Copyright© 1999, CRC Press LLC — Files may be downloaded for personal use only. Reproduction of this material without
the consent of the publisher is prohibited.
773
role of forestry, that is, to support agriculture and the system (spatial/temporal), relative importance
rural welfare was almost totally ignored (Westoby, and role of various components, production aims,
1975). and social/economic features. On the basis of
Such realization coupled with other socio- components, it is mainly divided into agriculture
economic pressures resulted in reassessment and (crops-trees-shrubs-vines), silvopastoral (pasture-
reexamination of developmental policies by agen- animals-trees), and agrisilviculture (crops-pasture-
cies like the World Bank and Food and Agricul- animals-aquaculture-sericulture-apiculture-trees).
ture Organization (FAO). This led the Interna- Based on the space arrangement, it can be divided
tional Development Research Center (IDRC) to into mixed dense, mixed sparse, support tree, etc.
commission a project under the leadership of John When classified from the viewpoint of protection,
Bene to identify the priorities for tropical forestry agroforestry is divided into soil conservation,
research. FAO focused its attention toward the moisture conservation, shelterbelts, and wind-
rural poor and emphasized the importance of for- breaks. However, when the emphasis is on pro-
estry for rural development. During 1976 to 1978 duction, it is divided into food, fodder, timber,
a series of seminars and workshops was held and etc. Recently, attention has been paid to the pos-
as a result agroforestry was identified as a system sible allelopathic interactions between different
to provide food, fuel, fodder, and other useful components of the agroforestry system to make it
products in a sustainable manner. However, the more productive and sustainable (Rizvi et al.,
formal rediscovery of agroforestry by the global 1992). We intend to discuss here the problems
scientific community can be credited to Bene and and prospects of such interactions.
his group and the IDRC. Bene and his associates
recommended that the priority should be given to
combining production systems, which would in- II. ALLELOPATHY IN AGROFORESTRY
tegrate forestry, agriculture, and/or animal hus- SPECIES
bandry in order to optimize land use (Bene et al.,
1977). Finally, with the establishment of the In- Owing to a worldwide awareness about the
ternational Center for Research in Agroforestry benefits and potential of agroforestry, extensive
(ICRAF), one of the oldest integrated traditional research has been conducted in this area in a
land use systems was transformed into a scientific relatively short period. Some of the best known
discipline called agroforestry. Presently, one of positive points about agroforestry are an overall
its most widely accepted scientific definition is, increased productivity (Avery et al., 1991; Burch
and Parker, 1992), enrichment of soil with or-
Agroforestry is a collective name for land-use sys-
tems and technologies where woody perennials
ganic matter and nitrogen (Agboole and Fayemi,
(trees, shrubs, palms, bamboos, etc.) are deliber- 1975; Ta and Farris, 1988), transport of nutrients
ately used on the same land management units as from lower to the upper layer of soil (Yamoah
agricultural crops and/or animals, in some form of et al., 1986), conservation of environment
spatial arrangement or temporal sequence. In the (Turnbull, 1984; Baumer, 1990), improved mi-
agroforestry system there are both ecological and
croclimate (Harris and Natarajan, 1987), and ru-
economical interactions between the different com-
ponents ral development through employment generation
(Lundgren and Raintree, 1982) (Chambers, 1983).
To optimize the gains of agroforestry, several
Agroforestry is being practiced in a variety of standard management practices have been evolved
climatic conditions to achieve one or more goals, and selection of suitable agroforestry species (AF
and thus needs a classification. Some of the com- species*) is one of those. Any such selection is
monly employed criteria for classification are based on a number of important characters of AF
based on arrangement of different components of species, such as fast growth rate, thorough pas-
* The term agroforestry species (AF species) covers all kinds of woody perennial trees, shrubs, palms, and bamboos grown with annual crops
in agroforestry systems.
774
sage for sunlight to the ground, rooting pattern, neem tree than in open plots. Similarly, the reduc-
and multipurposeness of AF species. It is surpris- tion in crop yields due to neem trees varies from
ing that allelopathic properties of AF species so 7 to 33% in sorghum (Sorghum bicolor), and 3 to
far have not been paid due attention. Research in 16% in safflower (Carthamus sp.) (Srivastava
this area began in the 1980s (Kuo et al., 1983; and Rammohanrao, 1989). Further studies, how-
Melkania, 1984; Rizvi and Rizvi, 1987), but the ever, suggest a selective effect of neem tree. Puri
surface has only been scratched. Detailed infor- and Bangawa (1992) have found that neem tree
mation about the allelopathic effects of AF spe- has no adverse effect on the yield of wheat (Triti-
cies on other components (annual plants) of cum aestivum) if grown 5 m apart from the main
agroforestry systems is limited. If available, such stem. Some studies suggest a direct role of neem
information would prove useful to identify allelochemicals in this effect on crop plants.
‘allelopathically compatible’ AF species (having Melkania (1984) found inhibition of germination
either beneficial or at least no adverse effect on of seeds of barnyard grass (Echinochloa crus-
companion crop) or ‘noncompatible’ ones with galli), buckwheat (Fagopyrum sagittatum), soy-
inhibitory effects. This kind of knowledge would bean (Glycine max), and turnip (Brassica rapa)
greatly facilitate formulation of agroforestry sys- by leachates of leaf, wood, and leaf litter. Maize
tems with higher yields by avoiding harmful al- (Zea mays), mustard (Brassica campestris), pea
lelopathic interactions and through exploitation (Pisum sativum), and wheat germination was also
of beneficial effects of particular AF species. In- inhibited by litter extract (Joshi and Prakash, 1992).
formation available on allelopathic activities of The extracts were particularly inhibitory to the
some of the important AF species is summarized root growth (Alam, 1990). Although a number of
in Table 1. All the AF species (except Moringa biologically active chemicals have been isolated
oleifera) tested for allelopathic activity have shown and identified, a correlation is yet to be estab-
an inhibitory effect on crop plants. lished between their presence and allelopathic
properties of neem.
III. TREE ALLELOPATHY: SOME

EXAMPLES B. Eucalyptus spp.
A. Azadirachta indica Eucalyptus has been promoted on a large scale

in various parts of the world. This is because of its
Azadirachta indica (commonly known as fast growth, adaptability to various edaphic and
neem) is a tree that has been used for centuries for climatic conditions, lesser soil coverage, least post-
medicinal and pesticidal properties. The tree is plantation care, and above all the industrial value.
indigenous to the Indian subcontinent, but it has However, its indiscriminate promotion without
been introduced and promoted in other regions of giving any significance to edaphic and ecological
the world. With the onset of agroforestry research, stability has evoked concern among environmen-
new frontiers for its exploitation have been dis- talists. In fact, during the last decade it has suf-
covered besides its traditional uses in pest con- fered from a dramatic fall in popularity because
trol, toiletries, pharmaceutical, cosmetics, plant of its antiphytosocial nature (Kohli, 1987) and ill
and animal nutrition, industry, and energy gen- effects on the ecology (Poore and Fries, 1985). Its
eration. Owing to its enormous uses, the U.S. monoculture plantations are reported to support
National Academy of Sciences has recognized it either very little or almost negligible understorey
as a tree for solving global problems (Anon., vegetation (del Moral and Muller, 1969; del Moral
1992). However, results of some agroforestry tri- et al., 1978; Bhaskar and Dasappa, 1986; Singh
als indicate its possible allelopathic effects on et al., 1993). The species diversity index is also
companion crops. Hazra and Tripathi (1989) have highly reduced under eucalypt monoculture plan-
reported that under semi-arid conditions, forage tations when compared with the other native plan-
yield of oats (Avena sativa) was 26% less under tations. Allelopathy has often been considered as
775
TABLE 1
Allelopathic Activity of Some Agroforestry Species
Agroforestry Target Plant parts/ Uses of agroforestry
species species allelochemicals species Ref.
Abies alba Abies alba Natural leachate, aqueous Fuel wood Becker and Drapier, 1984 ,85
Lepidium sativum extracts of fresh needles Georgiev, 1983
Picea abies and seeds
Pinus sylvestris
Acacia arabia Triticum aestivum Canopy effect, extracts of Building material/fuel wood/ Prakash et al., 1989
Pyricularia oryzae various parts farm timber/pest control Sheikh and Haq, 1978
A. auriculiformis Cicer arietinum Aqueous leaf leachate Aforestation Jadhav and Gaynar, 1992
Oryza sativa Rao et al., 1994
Triticum aestivum
A. confusa Bidens bipinnata Aqueous leachates and Aforestation Kuo et al., 1989
Brassica chinensis extracts of fresh and dried
Lactuca sativa leaves, leaf litter, seed/pod
extract
A. cyclops Eriocephalus racemosus Aqueous extract of Aforestation Rutherford and Powrie, 1993
and several other non-crop of phyllode
plants
A. dealbata Lolium perenne Aqueous extracts of Aforestation Casal et al., 1985
Trifolium pratense leaves, flowers, and soil
T. repens
A. excelsa Casuarina equisetifolia Aqueous extract of mature Shade Balasubramanian and

leaves Ravichandran, 1996
A. leucopholea Cajanus cajan Bark leachate Multipurpose tree Swaminathan, 1996
Sesamum indicum (MPT)
Zea mays
A. mangium Shorea leprosula Soil Timber Anwar, 1992
S. stenoptera
A. melanoxylon Lactuca sativa Phyllode extract Aforestation Gonzalez et al., 1995
A. nilotica Casuarina equisetifolia Aqueous extract of mature Beverage/fuel wood/MPT Balasubramanian and
Cajanus cajan leaves, bark leachate Ravichandran, 1996
Sesamum indicum Duhan et al., 1994
Rhizobium sp. Swaminathan, 1996
Zea mays
A. tortilis Cicer arietinum Aqueous extracts of fresh Fodder/fuel wood Saxena and Sharma, 1996
Gossypium hirsutum leaves and roots, soil from
Trifolium alexandrum under canopy
Triticum aestivum
A. xanthopholea A. xanthopholea Leaf and bark litter Fodder/fuel wood Nsolomo et al., 1995
Albizia lebbek leachates
Vigna radiata
Zea mays
Adhathoda vasica Brassica campestris Aqueous extract, rain Live fences/soil conservation Ayaz et al., 1989
Triticum vulgare leachate, litter, soil
Zea mays
Aegle marmelos Sclerotonia sclerotirum Leachates of roots, seed, Fruit/timber/medicinal use/ Ram, 1989
and bark pathogen control
Albizia lebbek Oryza sativa Canopy effect Fuel/fodder Bhatt et al., 1997
Parthenium hysterophorus Aqueous extract of Dhawan and Dhawan, 1995
leaves
A. stipulata Eleusine coracana Aqueous extracts of green Fuel/fodder/land Uniyal and Nautiyal, 1996
Echinochloa colonum leaves, leaf litter, and bark rehabilitation/nitrogen
Phaseolus radiata fixation
Lens esculentus
Annona squamosa Amaranthus spinosus Ethanolic extracts of Food/fuel wood Rizvi et al., 1980b
leaves and seeds
Azadirachta indica Fagopyrum sp., Glycine max, Leaf, wood and leaf litter Timber/lumber/manure/oil/ Joshi and Prakash, 1992
Oryza sativa, Pisum sativum leachates, litter, and fuel/food/pest control Melkania, 1984
Triticum aestivum, Zea mays, mature leaf extracts Rao et al., 1994
a number of microorganisms Schmutterer, 1995a
Bambusa arundinacea Arachis hypogea Aqueous leaf extract Building material/domestic Eyini et al., 1989
uses/fence
B. indica Costus speciosus Leaf leachate Building material/domestic Konar, 1996
uses/fence
Bauhinia variegata Vigna unguiculata, Leachates of leaves and Food/fuel wood Kaletha et al., 1996
Zea mays naturally flaked bark
776
TABLE 1 (continued)
Camellia sinensis C. sinensis Purine alkaloids in seeds Beverages Suzuki and Waller, 1987
Carica papaya Amaranthus spinosus Ethanolic extracts of Food/fruit/shade Rizvi et al., 1980b
leaves and seeds
Casuarina equisetefolia Cajanus cajan Leaf mulch, top soil, bark Charcoal/fuelwood/timber Suresh and Vianaya Rai, 1987
Helianthus annuus leachate Swaminathan, 1996
Sorghum bicolor
Zea mays
Celtis australis Brassica campestris Soil, dry leaf mulch, Food/fuelwood/timber Bhatt and Todaria, 1990
Glycine max aqueous leaf extract Melkania, 1992
Hordeum vulgare and leachate
Lepidium sativum
Citrus aurantium Amaranthus retroflexus Aqueous leaf extract, Essential oils/food Al-Saadawi and
Avena sativa decaying material Al-Rubeaa, 1985
Chenopodium album Al-Saadawi et al., 1985
Citrus aurantium Hassan et al.,1989
Cynodon dactylon
Citrus sinensis Citrus sinensis Soil from old orchards Food/shade/firewood/fruit Hassan et al., 1989
Coffea arabica Amaranthus spinosus Aqueous/ethanolic Beverage/fuel wood/food Chou and Waller, 1980
Coffea arabica extract, caffeine, Evenari, 1949
Lactuca sativa paraxanthine, scopoletin, Friedman and Waller, 1983
Secale cereale theobromine, theophylline, Rizvi and Rizvi, 1984, 1992b,

and phenolic acids Rizvi et al., 1987
Dalbergia sissoo Cicer arietinum Soil, exudates, mature leaf Shade/timber poles Puri and Bangawa, 1992
Oryza sativa extract Rao et al., 1994
Triticum aestivum
Emblica officinalis Pyricularia oryzae Extracts of various parts Fruit/pest control Prakash et al., 1989
Eucalyptus sp. Acacia saligna Shelterbelt effect Lumber/firewood/stake/ May and Ash, 1990
Lemna minor timber/essential oil/pole Onyewotu, 1985
Lolium perenne
E. alba Shorea palembanica Fresh leaves, leaf litter, Lumber/firewood/stake/ Anwar, 1991a,b
Zea mays root, and stem extracts timber/essential oil/pole
E. baxteri Casuarina pusilla Soil, topsoil extract, foliar, Lumber/firewood/stake/ del Moral et al., 1978
Leptospermum viminalis and litter leachates timber/essential oil/pole
Triticum aestivum
E. blakelyi Lemna minor Soil-decomposing litter Lumber/firewood/stake/ May and Ash, 1990
timber/essential oil/pole
E. camaldulensis Abelmoschus escutentus Leachates and aqueous Lumber/firewood/stake/ del Moral and Muller, 1970
Amaranthus caudatus extracts of dried/fresh timber/essential oil/pole Igboanugo, 1986
Avena fatua leaves, bark, leaf litter, Jensen, 1983
Bromus mollis volatiles, soil, 1,8- Lisanework and
Bromus rigidus cineole, α-pinene, Michelsen, 1993
Cicer arietinum α-phellandrene, phenolic, Mizutani, 1989
Vigna radiata gallic, and ferulic acids
Zea mays
E. citriodora Avena sativa Aqueous extract of fresh Lumber/firewood/stake/ Igboanugo, 1986, 1988a,b
Capsicum annuum leaf litter, leaf leachate, timber/essential oil/pole Kohli and Singh, 1991
Helianthus annuus crude volatile oils, oil Singh et al., 1991
Hordeum vulgare adsorbed soil, canopy Vicherková and Polová,
Lens esculentum effect 1986
Lycopersicon esculentum
Zea mays
E. delegatensis E. delegatensis Aqueous extracts of roots Lumber/firewood/stake/ Bowman and Kirpatrick,
and leaves, leachate of timber/essential oil/pole 1986
chopped leaves
E. deglupta Shorea palembanica Fresh leaves, leaf litter, Lumber/firewood/stake/ Anwar, 1991a,b
Zea mays stem flow timber/essential oil/pole
E. elata Lemna minor Decomposing litter, soil Lumber/firewood/stake/ May and Ash, 1990
E. globulus Cicer arietinum Leaf extract and leachate, Lumber/firewood/stake/ Baker, 1966
Cucumis sativus essential oils, oils, soil timber/essential oil/pole del Moral and Muller, 1969
Glaucium flavum percolate, canopy effect Kohli and Singh, 1991
Phaseolus aureus Molina et al., 1991
Lactuca sativa Singh and Bawa, 1982
Singh et al., 1991
E. grandis Abelmoschus esculentus Decomposing plant parts Lumber/firewood/stake/ Igboanugo, 1987
Amaranthus caudatus timber/essential oil/pole
777
TABLE 1 (continued)

E. macrorrhyncha Lemna minor Leaf and bark litter Lumber/firewood/stake/ May and Ash, 1990
Lolium perenne leachates, stem flow timber/essential oil/pole Nayyar et al., 1994
Raphanus sativus
Triticum aestivum
E. maculata Lemna minor Leaf and bark litter Lumber/firewood/stake/ May and Ash, 1990
leachates timber/essential oil/pole Nayyar et al., 1994
E. mannifera Lemna minor Decomposing litter, soil Lumber/firewood/stake/ May and Ash, 1990
timber/essential oil/pole Nayyar et al., 1994
E. melliodora Lemna minor Decomposing litter, soil Lumber/firewood/stake/ May and Ash, 1990
E. microtheca Herbaceous species Phytotoxins Lumber/firewood/stake/ Al-Mousawi and
timber/essential oil/pole Al-Naib, 1975, 1976
E. polyanthemosa Lemna minor Decomposing litter, soil Lumber/firewood/stake/ May and Ash, 1990
E. pulverulenta Lipidium sativum Grandinol Lumber/firewood/stake/ Bolte et al., 1984
E. radiata Lemna minor Volatile substances Lumber/firewood/stake/ May and Ash, 1990
E. robusta Zea mays Fresh leaves, leaf litter, Lumber/firewood/stake/ Anwar, 1991a
stem flow timber/essential oil/pole
E. rossii Lemna minor Stem flow, bark and leaf Lumber/firewood/stake/ May and Ash, 1990
Lolium perenne litter leachates timber/essential oil/pole
E. rubida Acacia saligna Leaf and bark litter Lumber/firewood/stake/ May and Ash, 1990
Eucalyptus globulus leachates, decomposing timber/essential oil/pole
Lemna minor litter, soil
Lolium perenne
E. saligna Cicer arietinum Aqueous leaf extract Lumber/firewood/stake/ Lisanework and
Eragrostis sp. timber/essential oil/pole Michelsen, 1993
Pisum sativum
Zea mays
E. tereticornis Brassica spp. Leaf and flower extracts, Lumber/firewood/stake/ Ahmad et al., 1984
Cicer arietinum leaf litter and mulch, timber/essential oil/pole Bhaskar and Dassappa, 1986
Helianthus annuus canopy effect, shelterbelt Sidhu and Hans, 1988
Lens esculentum effect Singh and Kohli, 1992
Nigella sativa Suresh and Vinaya Rai,
Phaseolus mungo 1987, 1988
Setaria italica
Sorghum vulgare
Triticum aestivum
Ficus bengalensis Brassica campestris Aqueous leaf extract, Charcoal/food/fuelwood/ Akram et al., 1990
Lens culinaris litter, soil beverage/shade/soil
Phaseolus vulgaris conservation
Raphanus sativus
Triticum aestivum
Zea mays
F. palmata Trifolium alexandrum Aqueous leaf extract, Charcoal/food/fuelwood/ Akram et al., 1990
Zea mays litter, soil beverage/shade/soil
conservation
F. racemosa Brassica campestris Aqueous leaf extract, Charcoal/food/fuelwood/ Akram et al., 1990
Lens culinaris litter, soil beverage/shade/soil
Raphanus sativus
Triticum aestivum
Zea mays
Aqueous leaf extract, Charcoal/food/fuelwood/ Akram et al., 1990
litter, soil beverage/shade/soil
conservation
F. religiosa Brassica campestris Aqueous leaf extract, Charcoal/food/fuelwood/ Akram et al., 1990
Lens culiraris litter, soil beverage/shade/soil
Raphanus sativus
Triticum aestivum
Zea mays
F. roxburghii Vigna unguiculata Leaf, naturally flaked bark Charcoal/food/fuelwood/ Kaletha et al.,1996
Zea mays beverage/shade/soil
conservation
778
TABLE 1 (continued)

Fraxinus micrantha Brassica campestris Aqueous leaf extract, Timber Joshi et al., 1996
Raphanus sativus coumarins
Triticum aestivum
Gliricidia maculata Helianthus annuus Leaf aqueous extract Shade/soil fertility Patil, 1994
Oryza sativa
Phaseolus mungo
Vigna radiata
G. sepium Bidens pilosa Leaf and bark (root) Crop shade/food/fuel wood/ Inostrosa and Founier,
Lycopersicon esculentum extracts, protocatechuic live fence/poles/shade 1982
acid, canavanine
Gmelina arborea Zea mays Leaf Firewood/lumber/shade/ Hauser, 1993
food/building material
Grewia optiva Brassica campestris Leaf leachate Food/shade/wind break/soil Melkania, 1984
Fagopyrum esculentum conservation
Glycine soja
Lepidium sativum
Setaria italica
Grevillea robusta Grevillea robusta Aqueous root extract Food/fuelwood/tools, and Webb et al., 1967
utensils
Inga edulis Oryza sativa Soil, exudate Crop shade/food/soil Salazar et al., 1993
conservation
Leucaena leucocephala Acacia confusa Aqueous leachate/ Fuelwood/pole/timber/food/ Chaturvedi and Jha, 1992
A. nilotica extracts of leaves, litter, soil conservation/fodder Chou and Kuo, 1986
Ageratum conyzoides soil, dry leaf mulch, Kuo et al., 1983
Alnus formosana topsoil Prasad and Subhashini, 1994
Lactuca sativa Mimosine, p-hydroxy- Rizvi et al., 1990a,b
Oryza sativa phenylacetic acid, cis- and
Sorghum bicolor trans-p-hydroxycinnamic
Vigna radiata acid, p-hydroxybenzoic
V. mungo acid, sinapic acid, vanillic
acid, caffeic acid,
p-coumaric acid, quercetin
Melia azadirach Brassica campestris Leaf leachate Crop shade/fuel wood/ Melkania, 1984
Lepidium sativum timber/lumber
Raphanus sativus
Setaria italica
A number of
microorganisms
Moringa oleifera Oryza sativa Leachate of intact and Fruit/vegetable/medicinal Rizvi and Rizvi, 1996
Vigna mungo chopped leaves, soil value
mixing of leaves
Pinus radiata P. radiata Volatile substances, Pulp/timber mulch Lill and McWha, 1976
Rhizopogon sp. soil, root extract
Secale cereale
Trifolium repense
Populus deltoides Saccharum officinarium Soil, leachate Timber Ralhan et al., 1992
Triticum aestivum Sheikh and Haq, 1986
Singh et al., 1993
Prosopis cineraria Triticum aestivum Soil, leachate Shade/soil conservation/ Puri and Bangawa, 1992
wind break
P. juliflora P. juliflora Leaf, stem, litter, and Fodder/fire wood/shade Chellahmuthu et al., 1997
Sorghum bicolor soil leachates Sundarammorthy et al., 1995
Vigna mungo Warrag, 1995
Prunus cerasoides Eleusine coracana Soil, dry leaf mulch, Food/fuel wood/timber Bhatt and Todaria, 1990
Glycine max aqueous leaf extract
Hordeum vulgare
P. jacquemontii Raphanus sativus Root extract Food/fuel wood/timber Joshi et al., 1997
Psidium guajava Lactuca sativa Living root exudate, Fruit/fuel wood/tools Brown et al., 1983
Setaria verticillata alcohol extract of
dried roots
Ricinus communis Meloidogyne incognita Oil cake Pest control/medicinal value Alam and Khan, 1974
M. javanica Singh, 1969
779
TABLE 1 (continued)

Tamarindus indica Amaranthus spinosus Ethanolic extracts of Beverage/fruit/fuel wood/ Rizvi et al., 1980b
leaves and seeds food/shade/tools/rituals
Tectona grandis Arachis hypogea Extract of fallen leaves, Firewood/lumber/shade/ Chaturvedi and Sharma, 1997
Cajanus cajan bark leachate timber Jeyakumar et al., 1987
Sesamum indicum Swaminatham, 1996
Zea mays
Terminalia arjuna Cicer arietinum Mature leaf extract Building material/ Rao et al., 1994
Oryza sativa timber/fuel
Triticum aestivum
T. tomentosa Oryza sativa Leaf leachate Building material/timber/ Gaynar and Jadhav, 1992
Vigna unguiculata fuel wood
Vitex negundo Andropogon nodosus Dried leaf extract, Weed control/medicinal Chou and Yao, 1983
Brassica chinensis p-hydroxybenzoic, value Kuo et al., 1989
Digitaria decumbens p-coumaric, ferulic,
Mimosa pudica vanillic, syringic acids,
Secale cereale flavonoids
A number of insects
a possible reason for the species depletion (Suresh fect growth of the associated vegetation (Kohli,
and Vinaya Rai, 1987; Kohli et al., 1992). 1990). Various volatile terpenes like limonene,
Eucalypt species have been used as a favor- cineole, citronellal, citronellol, α-pinene, and
able AF species. These are usually planted on the grandinol, etc. identified from the crude oil are
field boundaries as windbreaks, shelterbelts, or highly toxic and affect the germination and growth
simply scattered in the fields. Studies have shown of native vegetation (Baker, 1966; del Moral and
that the shelterbelts of eucalypts are very harmful Muller, 1970; Al-Mousawi and Al-Naib 1975,
to the crops growing in the adjoining area (Jensen, 1976; Bolte et al., 1984; Kohli et al., 1992). Un-
1983; Onyewotu, 1985; Igbuanugo, 1988a,b; Kohli der natural conditions, volatile oils are released
1990; Malik and Sharma, 1990; Puri and Bangawa, from the leaves through diffusion and being
1992; Singh and Kohli, 1992). Kohli and his as- heavier than air, travel downward, get adsorbed to
sociates have reported significant reduction in the the surface of soil particles, and thus affect the
density, root and shoot length, biomass, and eco- vegetation supported by this soil. The content of
nomic yield of crops up to 11 m from the the oil in leaves varies with species, climatic con-
shelterbelts of Eucalyptus. They suggested that ditions, and because of seasonal changes.
the performance of the crops can be assessed by The germination, seedling vigor, and seed-
a mathematical formula (% Performance = 100 + ling length of the four crops, namely, Phaseolus
m (x-24)/2, where x represents distance from the aureus, Hordeum vulgare, and Avena sativa were
tree and m represents value of regression slope). significantly reduced when placed in chambers
Later, based on the evaluation of bioefficacy of flushed with eucalypt oil (Kohli and Singh, 1991).
phytotoxins (extracted from the soil collected at Further, when seeds of P. aureus were placed in
different distances from the tree, and at various Petri dishes having soil adsorbed with eucalypt
depth from the soil surface), the poor crop perfor- oils, the germination was greatly reduced (Singh
mance was attributed to the allelopathic property et al., 1991). The volatile allelochemicals have
of Eucalyptus (Kohli et al., 1990; Singh and Kohli, also been found to inhibit respiration (Vicherková
1992). It is desirable to conduct similar studies and Polová, 1986), reduce the chlorophyll con-
with other AF species. tent, and cause wilting (Kohli and Singh, 1991).
Eucalypts are reported to release a number of In addition, the leachates and extracts from the
volatile and nonvolatile allelochemicals that af- eucalypt leaves, litter, bark, flowers, and leaf mulch
780
have been reported to reduce the germination and mination, root length, and dry matter production
initial growth of a number of plant species (Singh were reduced. Several other workers have also
and Bawa, 1982; Ahmed et al., 1984; Igboanugo, reported allelopathic effect of aqueous extracts
1986; Sidhu and Hans, 1988; Kohli, 1990; May of leaves, litter, soil, leaf leachate, seed exudate,
and Ash, 1990; Lisanework and Michelson, 1993). dry leaf mulch, topsoil, and its allelochem-
A number of agricultural crops were tested for icals. Abelmoschus esculentus, Acacia confusa,
their susceptibility or resistance to the aqueous A. nilotica, Ageratum conyzoides, Alnus for-
leachates of Eucalyptus. Interestingly, the level of mosana, Bidens pilosa, Brassica chinensis,
resistance was determined by the ratio of seed B. juncea, Cajanus cajan, Casuarina equisetifolia,
coat thickness and seed volume. The critical val- C. glauca, Cicer arietinum, Helianthus annuus,
ues of this ratio were determined mathematically, Lactuca sativa, Liquidambar formosana, Mimosa
and it was found that seeds with a ratio of seed pudica, Miscanthus floridulus, Phaseolus vulgaris,
coat thickness to seed volume greater than 0.79 Pinus taiwanensis, Stachytarpheta jamaicensis,
were resistant or the vice versa (Kohli, 1994). Sorghum bicolor, and Vigna radiata are some of
Studies conducted by Kohli (1990) and Singh the common species that are negatively sensitive
(1991) demonstrate that both volatile as well as to allelochemicals of Leucaena (Kuo et al., 1983;
nonvolatile allelochemicals are continuously be- Chou and Kuo, 1986, Rizvi and Rizvi, 1987; Kuo
ing added to the soil system beneath the planta- et al., 1989; Chaturvedi and Jha, 1992; Rizvi et al.,
tions. The soil collected from the floor of these 1990a,b, 1994; Narwal, 1996; Sinha, 1996).
plantations was found to be rich in phenolic com- There are reports on the allelopathic effects
pounds. Their content varied with distance as well of various plants on nitrogen fixation (Rice, 1984).
as depth. Later, a number of phenolic acids like Therefore, Rizvi and his associates selected nitro-
gallic acid, gentisic acid, syringic acid, vanillic genase, the enzyme for nitrogen fixation as a test
acid, caffeic acid, p-coumaric acid, ferulic acid, parameter to measure the allelopathic activity of
and cinnamic acid were identified in the soil as Leucaena leaves. They grew V. radiata in soil
well as in the leaves (Kohli, 1990). having Leucaena leaves, and in soil collected from
under the canopy of 5-year-old Leucaena plants.
They found that nitrogenase (N-ase) activity of
C. Leucaena leucocephala plants grown with Leucaena leaves (8 g/kg) was
34% more than in the control plants, but this
Leucaena is a widely recommended tree spe- increase was reduced by 29% when the plants
cies for agroforestry because of its fast growth were grown with higher amount of leaves
rate, fodder, fuel and wood value, ability to fix (16 g/kg). The N-ase activity was found to be
nitrogen, and to improve the overall productivity reduced further when plants were grown in
of land (Chou and Waller, 1989; Nair, 1989; Leucaena-canopy soil, causing an inhibition of
Lantican and Taylor, 1991). However, the pres- 28% when compared with the control (Rizvi, 1996;
ence of a non-protein amino acid along with some Sinha 1996).
phenolic compounds in its leaves and seeds is a According to a report of the International
cause of concern to allelopaths and ecologists. Institute for tropical Agriculture (Anon., 1980),
Chou and his associates after working with the yield of maize (Zea mays) and rice (Oryza
different species of Leucaena for several years sativa) was increased when grown in association
have concluded that exclusion of understorey with Leucaena. Rachie (1983) also found an in-
vegetation by Leucaena is at least partly mediated crease in the yield of maize intercropped with
by allelopathy (Kuo et al., 1983; Chou and Kuo, Leucaena. Studies conducted by Jeyaraman (1991)
1986; Chou, 1993). Suresh and Vinaya Rai (1987, and Salazar et al. (1993) further supported a posi-
1988) tested the allelopathic influence of Leucaena tive effect of Leucaena green leaf mulch on sev-
on sorghum and sunflower using topsoil collected eral growth and yield contributing parameters of
from the field either mulched with dry leaves rice, resulting in a higher yield. Cajanus cajan,
or irrigated with aqueous leaf extract. Seed ger- Sesamum indicum, Ricinus communis, and
781
Sorghum vulgare are some of the other plants that IV. AGROFORESTRY AND PEST
are positively affected by Leucaena (Singh, 1983). CONTROL
However, there are contradictory reports about its
effect on maize. Karim and co-workers (1991) Plants are known to synthesize allelochemicals
have reported an inhibition of growth and yield of that affect germination, growth, metabolism, de-
maize grown in association with Leucaena hedges. velopment, distribution, behavior, and reproduc-
The allelopathic effects of Leucaena are at- tion of other organisms (Inderjit et al., 1994; Rice,
tributed to the presence of a number of phenolic 1995; Narwal et al., 1997). The presence of these
compounds and mimosine (Table 1). Various allelochemicals often imparts plant resistance to
concentrations of mimosine have been found to pathogens, insects, nematodes, and reduces infes-
be inhibitory when applied to different plants. tation of weeds (Rice, 1984, 1995; Green and
Radicle growth of lettuce (Lactuca sativa), rice, Hedin, 1986; Chou and Waller, 1989; Rizvi and
radish (Raphanus sativus), and turnip (Brassica Rizvi, 1992a; Copping, 1996).
rapa) was inhibited by 10 to 20 ppm of mimosine Agroforestry systems provide an excellent
(Kuo et al., 1983; Tawata and Hongo, 1987). opportunity to explore the pest controlling prop-
Germination, radicle, and plumule length of erties of AF species. Most of the AF species
Abelmoschus esculentus, Brassica campestris, produce a good amount of leaf, litter and debris
Phaseolus aureus, Raphanus sativus, Triticum that are rich in allelochemical content. These
aestivum, and Vigna mungo have been found to allelochemicals in turn provide various kinds of
be inhibited by 1 mM mimosine. Phaseolus aureus pest controlling properties to AF species. Thus,

and V. mungo were affected the most and showed their allelopathic materials can be used as mulch,
an 83 and 86% inhibition of radicle growth, re- and their leachates and purified compounds may
spectively (Rizvi et al., 1990a,b). be eco-friendly alternatives to synthetic pesticides.
Despite the reports on the allelopathic effects
of mimosine, not much information on the mode
of action is available. Rizvi and his associates A. Agroforestry in Weed Control
have found that mimosine inhibited a large num-
ber of physiological and biochemical parameters Trees can regulate the germination and growth
in V. mungo and P. aureus. They found that and development of weeds, through allelopathy.
mimosine inhibited seedling vigor, food mobili- Scopolin and Scopoletin isolated from Celtis
zation efficiency, solubilization of starch, break- laevigata are reported to suppress Amaranthus
down of proteins, and activity of amylase. The palmeri (Lodhi and Rice, 1971). Ethanolic ex-
reduced amylase activity was at synthetic as well tracts of seeds of Annona squamosa, Carica pa-
as catalytic level, and it was mediated by gibber- paya, Coffea arabica, and Tamarindus indica were
ellic acid. They further reported that mimosine found to inhibit germination of Amaranthus
altered the hormonal balance of the seedlings lead- spinosus by 13, 58, 100, and 36%, respectively
ing to an inhibition in their growth. When (Rizvi et al., 1980 b). They isolated and identified
V. mungo plants were grown in the soil having an allelochemical as 1, 3, 7-trimethylxanthine
different amounts of Leucaena leaves, nitroge- (1,3,7-T) from the seeds of C. arabica. When
nase activity of root nodules was inhibited (Rizvi tested for herbicidal potential, it completely in-
et al., 1990a,b, 1994; Rizvi and Rizvi, 1998). hibited the seed germination of A. spinosus at
A report by Prasad and Subhashini (1994) 1200 ppm. The compound also suppressed the
also confirms that the inhibitory effects of germination of Avena fatua (40.1%), Echinochloa
mimosine on germination and seedling growth of colonum (100%), and E. crusgalli (91.2%) at a
rice is mediated through its effect on nitrate re- concentration of 2000 ppm. Rizvi and his associ-
ductase, catalase, IAA-oxidase, peroxidase, and ates further studied the selective behavior of
its isozymes. Further studies on the mode of ac- 1,3,7-T and its mode of action on A. spinosus.
tion of allelochemicals produced by AF species While completely inhibiting the weed, it did not
would help in understanding the mechanism of affect the germination and growth of Vigna mungo
tree-crop interaction in agroforestry system. in which the weed is a problem (Rizvi et al.,
782
1980c, 1987; Rizvi and Rizvi, 1983, 1984). The herbicidal properties of AF species ap-
Allelochemi-cals like mimosine, 3,4-dihydroxy pear promising. Some of the examples of
pyridine, and phenolics present in Leucaena have allelochemicals that have demonstrated weed-sup-
also been found to totally exclude weeds like pressing ability are given in Table 2.
Ageratum conyzoides and Mimosa pudica (Chou
and Kuo, 1986; Chou, 1993).
Seed germination and seedling growth of B. Agroforestry in Pathogen Control
Amaranthus retroflexus, Avena sativa, Chenopo-
dium album, and Cynodon dactylon were found to One strategy to exploit allelopathy is the
be inhibited by aqueous extracts, decaying mate- use of allelochemicals for the control of patho-
rials, and volatile compounds of senescent and gens (Rice, 1995). The plant allelochemicals
nonsenescent leaves of Citrus aurantium (Al- can be used either in the purified form, in the form
Saadawi and Al-Rubeaa, 1985; Al-Saadawi et al., of crude plant extracts or as volatile extracts.
1985). Dried mango (Mangifera indica) leaf pow- However, very few tree-based allelochemicals
der @ 250 g/10 kg mixed with soil in pot culture have been exploited for this purpose. Neem
completely inhibited the germination of Cyperus (Azadirachta indica) is one tree, which possesses
rotundus up to 21 days, and its application to potential to kill pathogens (Ghwande, 1989;
seedling considerably inhibited (60%) the tillering Schmutterer, 1995a). Its seed cake, seed and fruit
(Mohanty et al., 1994). Eucalyptus spp., because extracts, seed kernel powder, and seed oil have
of high allelopathic activity, are expected to con- been reported to control a wide spectrum of fun-
trol weeds (Kohli et al., 1998a). Eucalyptus leaf gal pathogens (Gunasekaran et al., 1986; Jeyarajan
leachate and oil showed differential effects on the et al., 1987; Srivastava et al., 1997). The biologi-
growth of two weeds. A 20% leaf leachate sup- cal activity of neem against pathogens is attrib-
pressed the biomass production of Cynodon uted to the presence of sulfurous compounds in its
dactylon by about 50%, whereas 1% oil caused a seed oil. Moreover, neem products also act as
68% reduction. Application of 1% oil signifi- deterrents to pathogen-carrying insects, thereby
cantly inhibited shoot and root length, leaf chlo- decreasing the disease incidence (Saxena et al.,
rophyll, and total biomass production of Cyperus 1985; Eppler, 1995). Besides neem, some other
rotundus (Babu et al., 1996). Aqueous extracts of AF species have also been evaluated for their
its bark, leaves, and oil inhibited Parthenium effect on pathogens. Some of the examples can be
hysterophorus (Kohli et al., 1998b). Based on the seen in Table 3.
chemistry of cineole (a component of Eucalyptus
oil), a commercially used bioherbicide — cin- C. Agroforestry in Nematode Control
methylene — has been developed (Duke, 1986).
Ailanthone, another chemical (isolated from Ail- Neem (Azadirachta indica) is a plant species
anthus alissisma), has been reported to possess a that has been studied for the nematode control.
post-emergence herbicidal property similar to Various parts of neem or their extracts have been
glyphosate and paraquat (Heisey, 1996). found nematicidal against Meloidogyne incognita.
TABLE 2
Allelochemicals Isolated from Agroforestry Species with Weed-Suppressing
Properties
Allelochemical Chemical nature Natural source Ref.
Ailanthone Quassinoid Ailanthus altissima Heisey, 1996

Caffeine Alkaloid Coffea arabica Rizvi et al., 1980c
Cineole Terpenoid Eucalyptus globulus Kohli et al., 1998a
Citronellal Terpenoid Eucalyptus citriodora Kohli et al., 1998a
Mimosine Non-protein amino acid Leucaena leucocephala Rizvi, 1998
Azadirachtin Sesquiterpene lactone Azadirachta indica Koul et al., 1990
783
TABLE 3
Agroforestry Species with Potential to Control some Crop Pathogens
AF species Plant part/chemicals Pathogen Reference
Acacia arabica Roots, seeds, and bark Pyricularia oryzae Prakash et al., 1989
extracts
Aegle marmelos Leaves Sclerotonia sclerotirum Ram, 1989
Azadirachta indica Leaf extract, oil, Phaeoisariopsis personata Ghwande, 1989
essential oils Mycobacterium tuberculosis Singh and Dwivedi, 1990
Sclerotum rolfsii Ganapathy and
Staphylococcus aureus, Narayansamy, 1990
TOSWV
Neem cake Fusarium solani Jeyarajan et al., 1987

Ganoderma lucidium
Macrophomina phaseolina
Phytophthora capsici
Rhizoctania solani
Aqueous bark extract, TMV Murty, 1982
Nimbidin PVX Verma, 1974
Commercial products TMV, ZYMV, vectors Eppler, 1995
Seed kernel extract Xanthomonas campestris Eswaramurthy et al., 1993

Callistemon lanceolatus Essential oil Pythiun aphanidermatum Kishore and Dwivedi, 1991
Citrus limone Fresh leaves Cockliobolus miyabeanus Tewari and Nayak, 1991
Pyricularia oryzae
Rhizoctonia solani
Essential oil Pythium aphanidermatum Kishore and Dwivedi, 1991
Coffea arabica 1,3,7-Trimethylxanthine Drechslera maydis Rizvi et al., 1980a
Eucalyptus spp. Volatile and nonvolatile Sclerotum rolfsii Singh and Dwivedi, 1990
fractions
E. rostrata Essential oil, leaf powder Sclerotum capivorum Salama et al., 1988
Emblica officinalis Extracts of different parts Pyricularia oryzae Prakash et al., 1989
Juniperus communis Essential oil Pythiun aphanidermatum Kishore and Dwivedi, 1991
Lawsonia inermis Leaf extract Phaeoisariopsis personata Ghwande, 1989
Lonchocarpus castilloi Flavonoids of hardwood Lenzites trabea Gomez-Garibay et al., 1990
Pinus spp. Essential oils Pythiun aphanidermatum Kishore and Dwivedi, 1991
P. taeda Soil amended with bark Pythium aphanidermatum Huang and Kuhlman, 1991
Fusarium sp.
Rhizoctonia sp.
Pongamia pinnata Leaf extract Phaeoisariopsis personata Ghwande, 1989
Saraca indica Leaf extract Pythium debaryanum Kumar and Tripathi, 1991
Fusarium oxysporum
Terminalia arjuna Extracts of different parts Pyricularia oryzae Prakash et al.,1989
T. belerica Extracts of different parts Pyricularia oryzae Prakash et al.,1989
Vitex negundo Extracts of different parts Pyricularia oryzae Prakash et al.,1989
Leaves (Singh and Sitaramaiah, 1967, 1969; Sitaramaiah, 1967; Rossher and Zebitz, 1987);
Vijayalskshmi et al., 1979); flower, bark, and gum chickpea (Cicer arietinum) (Gupta and Ram,
(Siddiqui and Alam, 1985); seeds (Mishra 1981); mungbean (Vigna radiata), pulses and
et al., 1989) and seed coat kernel/cake (Mojumder vegetable crops (Mojumder, 1997). Reddy et al.
and Mishra, 1991a,b) are some of the most (1997) listed more than 20 nematode species that
studied parts. Nematode populations and root are susceptible to neem or chinaberry (Melia
galling have been inhibited by extracts of neem in azadirach) derivatives.
Abelmoschus esculentus, Lycopersicon escu- Castor (Ricinus communis) is another AF
lentum, and Solanum tuberosum (Singh and species known to have nematicidal activity. Its oil
784
cake has been found to reduce galling on okra products. Some of the neem allelochemicals tested
(Abelmoschus esculentus) roots caused by for their efficacy against various insect pests are
Meloidogyne javanica (Singh, 1969). In spinach azadirachtins, azadirones, nimo- and nim-
(Spinacea oleracea), soil population of M. incog- bocinolides, salannins, vilasinines, nimbenene and
nita was suppressed and root galling was reduced 6-deacetylnimbinene, margosinolides, meliantriol,
by the treatment of its oil cakes (Alam and Khan, alkanes, and several sulfur-containing compounds
1974). Khan and his associates have found that (Koul, 1992). Neem products or their formula-
the soil population of M. incognita in tomato tions have been found to be effective against in-
fields was reduced when treated by castor oil sect pests in corn (Hellpap, 1995); oil crops
cakes (Khan et al., 1969, 1973). A reduction in (Zebitz, 1995a); vegetables and grain legumes
the number of M. incognita and Meloidogyne sp., (Ostermann and Dreyer, 1995); fruit trees, root,
root galling, and an increase in tomato yield when and tuber crops (Zebitz, 1995b,c); forest, orna-
cultivated in the presence of castor cake have mental trees, and shrubs (Schmutterer, 1995b);
been reported by Hasan (1992). Some of the other blood sucking and other parasites of man and
tree species, which are reported to suppress nema- domestic animals (Schmutterer, 1995c); and pests
todes, are Acacia auriculiformis (Sinhababu et al., of stored products (Saxena, 1995). Readers inter-
1992), Coffea sp. (Tronocon et al., 1986), and ested in details may refer to Schmutterer and
L. leucocephala (Jain and Hasan, 1985). Ascher (1984, 1987), Schmutterer (1995a), and
Apart from using plant parts or their extracts, Narwal et al., (1997).
purified allelochemicals could also be used for Besides neem, only a few plant species (known
nematode control. However, only a few allelo- for agroforestry uses) have been evaluated for
chemicals have been isolated and identified that their insect-controlling property. Rizvi and his
are active against nematodes. Alkaloids like associates isolated a compound 1,3,7-trimethyl-
nimbidin and thionemone isolated from neem have xanthine from seeds of coffee (Coffea arabica)
been found toxic to a number of phytoparasitic and tested it as a chemosterilent against stored
nematodes (Khan et al., 1974a,b). Further studies grain pest-Callosobruchus chinensis. They found
are needed to demonstrate the benefits of that at 1.5% concentration the compound signifi-
agroforestry in nematode control. cantly inhibited the oviposition (Rizvi et al.,
1980d). Oil extracted from seeds of Annona squa-
mosa showed a significant reduction in survival
D. Agroforestry and Insect Control of rice leafhopper (Nephotettix virescens) and
thereby transmission of rice tungro virus (Prakash
The insect repellent property of neem et al., 1989). They further reported that leaves of
(Azadirachta indica) was first discovered in India Vitex negundo are effective against Rhizopertha
in 1928, and the neem leaves were used in pro- dominica, Sitotroga cerealella, and Tribolium
tecting stored rice from insect infestation. How- castanem. Insects like Achaea janta, Bruchus
ever, significant advancement in the insect con- chinensis, Diacrisia obliqua, Euproctis fraterna,
trolling potential of neem started in the 1960s. Sitotroga cerealella, Spodoptera litura, and
H. Schmutterer rediscovered the properties of Scirpophaga sp. were also inhibited by V. negundo.
neem tree that control insects while working in Shin-Foon (1987) reported that seed oil of Melia
Sudan during a locust invasion in 1959. The next azadirach is effective against citrus red mite
3 decades witnessed an unprecedented increase in (Panonychus citri) and orange spiny white fly
research activities throughout the world. (Aleurocanthus spiniferus). He further reported
Neem products, ranging from simple leaf and that toosendanin, a triterpenoid isolated from the
seed kernel powders, their extracts, oil, cake, ac- bark of M. toosendan, possesses antifeedant and
tive compounds, and several commercial prod- growth-disturbing properties against cabbage
ucts, have been tested against 450 to 500 species worm — Pieris rapae. When methanolic extract,
of insects. Schmutterer (1995a) has listed 413 stem, and bark of Grewia microcos were tested
species or subspecies that are susceptible to neem for their anti-insect properties against Aedes
785
aegytii, Plutella xylostella, and Callosobruchus and Sinha (1996) have demonstrated that
chinensis, the LC-50 of methanolic extract against allelochemicals released from AF species also
second-instar mosquito larvae was found to be affect the quality of food grains. When Vigna
47.5 ppm (Permaratne et al., 1996). radiata was grown in monoculture and in alley
Indiscriminate use of synthetic pesticides is a cropping with Leucaena, iron content of grains
serious global problem. Several of them have been obtained from the latter was reduced by 72%.
found to be hazardous due to their long persis- Thus, even if the grain yield potential is high, the
tence, health-related problems, environmental possibility of producing grains with low nutritive
pollution, and non-target toxicity (Rizvi et al., value cannot be ruled out. On the contrary, when
1998, Rizvi and Rizvi, 1992 b). The organophos- plants of V. radiata were grown in the presence of
phorus compounds are especially dangerous be- leaves of Moringa oleifera, the level of zinc and
cause they are known to attack the nervous sys- iron content in grains was increased by 28 and
tem (Vighi and Funari, 1995; Van Emden and 45%, respectively (Rizvi and Rizvi, unpublished
Peakall, 1996). By contrast, the botanicals data). Thus, it is extremely important to conduct
(allelochemicals), with some exceptions, are con- similar studies with other AF species to ensure
sidered to be less toxic to non-target species and the supply of food with at least normal nutritive
environmentally safer owing to their biodegrad- value or if possible, with an improved quality.
able nature (Copping, 1996). Thus, plants may Agroforestry systems designed on the basis of the
prove a neverending reservoir of eco-friendly above considerations could improve the quality
allelochemicals because they are likely to be re- of life of the poor and malnourished.
cycled through nature. Agroforestry species re-
main a component of agroecosystem for com-
paratively longer period; therefore, their possible B. Soil Quality and Replant Problem
ability to control pests could be of great value. with Agroforestry Species
In agroecosystems, nurseries, and agroforestry

V. APPROACHES FOR FUTURE systems where the same crops are grown year
RESEARCH after year, the Replant Syndrome often appears,
which pertains to the injurious effect on the suc-
A. Qualitative Yield of Agroforestry ceeding crops. It often makes the land unfit for
Systems further use. To rejuvenate such soils and reestab-
lish the crops in these is difficult. Data suggest
Agroforestry, besides being an important tool that in most of these problems allelopathy plays
for developing sustainable land use, is also con- an important role either directly, or through me-
sidered a system to increase food production, es- diation of microorganisms, nematodes, and other
pecially for rural people. In less-developed and soil biota. The replant problem is at least partly
developing countries, the latter goal of agroforestry due to the presence of allelochemicals released in
becomes more crucial because the majority of the the soil from the roots of the intact plants, fallen
rural population depends on food from their small litter, or the post-harvest residues left in the soil.
land holdings. Thus, any recommended agro-for- In some cases, the allelochemicals responsible for
estry system should ensure production of suffi- the same have been identified, such as amygdalin
cient food (especially the grains obtained from in peach (Prunus persica) and phlorizin and phlo-
the crop component of agroforestry), without com- retin in apple (Malus baccata) (Rice, 1984).
promising the nutritional value of the product. The problem is prevalent especially in the
Unfortunately, while evaluating the grain yield of orchards, nurseries, and agricultural fields where
agroforestry systems, a gross increase in the quan- the same cropping pattern is repeated year after
tity of grains (if any) invariably has been consid- year. As early as in the beginning of this century,
ered a measure of the increase in food production. Schreiner and Reed (1908) reported the replant
However, studies conducted by Rizvi et al. (1990b) problem of crops and attributed this to deteriora-
786
tion of soil caused by the toxins released from the ass and density of its understorey vegetation are
previous crops. Their study concluded that the relatively lower than in adjacent pastures. Field
removal of the toxins from the soil overcomes results showed that the natural leachates of
this problem. Although continuous cropping of V. negundo significantly retarded the growth of
the same species is successful in many parts of the Digitaria decumbens but stimulated the growth of
world, replant problem has been documented from Andropogon nodosus when compared with the
some of the countries. A number of crop plants rainfall control. The growth of D. decumbens,
like coffee (Friedman and Waller, 1983), alfalfa grown in pots under green house conditions, was
(Medicago sativa) (Miller, 1983), wheat (Thorne significantly retarded by watering with a 1% aque-
et al., 1990), rye (Secale cereale) (Wojcik- ous extract of V. negundo, but the growth of
Wojtkowiak, 1990), maize (Yakle and Cruse, A. nodosus and Mimosa pudica was inhibited. Kil
1983), and rice (Chou and Lin, 1976) in addition (1992) reported a selective pattern in understorey
to orchard crops are known to suffer with this vegetation of pine stands when compared with
problem. Recently, Duhan et al. (1994) found that the adjoining area. Species such as Aster tataricus,
soil of Acacia nilotica rhizosphere inhibits the Cymbopogon tortilis, Themeda triandra, and Plan-
nitrogenase (N-ase) activity in Rhizobium sp. Rizvi tago asiatica were found to grow well inside the
(1996) has also found a 28% inhibition in the N- pine forest, where species such as Boehmeria
ase activity of root nodules of Vigna radiata plants plantanifolia, Cassia tora, Chenopodium album,
when grown in Leucaena-canopy soil. These find- and Digitaria sanguinalis were growing only
ings indicate toward the possibility of soil health outside the forest. This selectivity was attributed
problem in agroforestry systems. to the presence of a number of phenolic
allelochemicals in pine leaves and canopy soil.
The yield of wheat, green gram (Vigna ra-
C. Selective Behavior of Tree diata), and turmeric (Curcuma longa) was re-
Allelochemicals duced when grown in alley cropping with
Leucaena, but the yield of maize and rice was
It has been reported that AF species behave increased (Anon., 1991). Research conducted at
differentially with companion crops (Anon., 1991). the International Institute for Tropical Agricul-
Kuo and his associates reported that only a few ture (Anon., 1980), and by Rachie (1983) further
understorey species grow under the Leucaena supported a positive effect of Leucaena on the
canopy, but a substantial number of its own seed- yield of maize and rice. Studies conducted by
lings were able to grow. Experimental data on Jeyaraman (1991) and Salazar et al. (1993) also
light, soil moisture, and nutrients, etc. revealed demonstrate a positive effect of Leucaena green
that competition for these factors was not the leaves on several growth and yield parameters in
major cause for the phenomenon of exclusion of rice. They found that the rice plant was not only
understorey vegetation (Kuo et al., 1983; Chou tolerant to negative allelopathic effects, but it gave
and Kuo, 1986; Chou, 1993). Rizvi and his asso- a higher yield with Leucaena mulch. Such obser-
ciates evaluated the effect of Leucaena leaf vations on the selective allelopathic behavior of
leachate on Abelmoschus esculentus, Brassica AF species for companion crops could be of prac-
juncea, Cajanus cajan, Cicer arietinum, Oryza tical value in designing agroforestry systems.
sativa, Phaseolus vulgaris, Vigna radiata, and Zea Selective action of a chemical can be attrib-
mays. Except for O. sativa, all other crops were uted to several factors viz. some detoxifying
significantly inhibited by Leucaena leachate mechanism in the receptor plant; concentration of
(Rizvi, 1996; Sinha, 1996). Similar effects were allelochemical in the producer plant, soil, and site
observed with pure mimosine (Rizvi and Rizvi, of action at a particular time; activity of microbes
1987; Rizvi et al., 1990a,b). Vitex negundo, a and environmental conditions, etc. However, re-
dominant component of coastal vegetation, is search is lacking in this area. Smith and Fowden
widely distributed in the southern parts of Tai- (1966) have demonstrated that Leucaena seed-
wan. Chou and Yao (1983) found that the biom- lings detoxify mimosine into 3,4-dihydroxypyri-
787
dine, which is further broken down into nontoxic animals. Most conspicuous of these adverse ef-
metabolites. Rizvi and his associates found varia- fects are the loss of hair in nonruminant animals
tion in allelochemical concentration in Leucaena and the reduction of egg production in poultry.
leaves over the year. While studying the effect of Cattle fed excessively on Leucaena meal suffer
leachates (prepared from the leaves collected ev- from hair loss, poor growth, and excessive saliva-
ery month) on V. mungo, they found the radicle tion (Ritchie, 1974). Mimosine can also cause the
growth inhibited by 10 to 62%. Thus, the adverse formation of goiter, deterioration in the quality of
effect on the companion crops grown during the wool, retarded growth, fertility problems, and
high allelochemical concentration period is obvi- general adverse effects on health, which may
ous (Rizvi et al., 1994; Sinha, 1996). Similar stud- eventually lead to mortality. Mimosine is known
ies with other AF species are needed to make to inhibit a number of biochemical reactions. It
suitable selections of companion crops. Further, acts as a tyrosine antagonist and competes with
such studies may also prove to be useful from the tyrosine to inhibit the activity of tyrosinase, re-
viewpoint of animal nutrition. duces synthesis of high-tyrosine proteins and
DNA, and leads to reduction of blood thyroxin
(Ries et al., 1975; Hegarthi et al., 1976; ter-Meulen
D. Allelochemicals: Role in Animal and and El-Harith, 1985; Bray, 1986). It has also been
Human Nutrition reported to inhibit tyrosine decarboxylase, aspar-
tate glutamate transaminase, and synthesis of RNA
One of the major goals of agroforestry is to and proteins (Bell, 1972; Kuo et al., 1983). Ear-
provide fodder for farm animals. This is even lier, the toxicity of Leucaena was ascribed to its
more important for the farmers of developing ability to accumulate selenium from soil (Arnold,
countries. The populations of most of these coun- 1944). However, Yoshida (1994) working with
tries have only small land holdings, and they rats has shown that the loss of hair and other toxic
almost completely depend on farm for their food, symptoms that developed when rats were fed on
fodder, and other needs. In these circumstances Leucaena meal were not due to selenium accu-
farmers have a high expectation from their mulation but rather to mimosine.
agroforestry endeavors. This includes a regular The expression of mimosine toxicity depends
supply of suitable, nutritious, and less-expensive on the rumen microbial ecology, and the level of
fodder. Being completely dependent on their farm toxicity is related to the extent and rate of bacte-
animals for agricultural and economic purposes, rial breakdown of mimosine. During degradation
the health of livestock is always a concern. There- of mimosine, two isomers viz. 3-hydroxy-4 (1H)-
fore, the agroforesters must guarantee that the pyridone (3,4-DHP) and 2,3-DHP are produced.
fodder available through agroforestry is not only Some of the rumen bacteria are capable of detoxi-
cheaper and nutritious, but safer as well. Unfortu- fying both forms of DHP, but the possibility ex-
nately, the roles of allelochemicals in this regard ists that considerable quantities of mimosine and
have almost been totally ignored. Rizvi and Rizvi 3,4-DHP may escape such degradation (Bray,
(1992b) have emphasized that it is desirable to 1986; D’Mello, 1994). The nature of bacteria in
evaluate the tree component of agroforestry sys- ruminants varies in different geographical regions.
tems from this viewpoint. Only limited informa- In those regions where Leucaena is indigenous
tion is available with the exception of allelo- (Central America) or is naturalized (Hawaii and
chemicals such as mimosine and canavanine pro- Indonesia), ruminants possess the bacteria required
duced by Leucaena leucocephala and Gliricidia for breakdown of mimosine and its degradation
sepium, respectively. products. This leads to the absence of Leucaena
The presence of a high amount of crude pro- toxicity. However, in other regions like Australia,
tein, potassium, calcium, phosphorus, carotenes, Kenya, and the U.S.A., ruminants lack the bacte-
vitamin K, and riboflavin in Leucaena leaves make ria that are required to detoxify mimosine and its
it desirable as an animal feed. However, research- isomers. Therefore, in these regions it is common
ers have found that Leucaena can be toxic to to observe mimosine toxicity in ruminants fed on
788
Leucaena meal for a long period. If ruminants are and productivity besides conservation of the envi-
inoculated with DHP-degrading bacteria, the prob- ronment, biodiversity, and natural resource base.
lem of mimosine toxicity can be solved. Such
uses have been demonstrated successfully in
Australia (Brewbaker, 1989). Another AF spe- ACKNOWLEDGMENTS
cies, G. sepium, is known to contain a highly
toxic allelochemical — canavanine. Besides be- The authors thank the International Center
ing allelopathic (Bass et al., 1995), it is known to for Research in Agroforestry, Nairobi, for provid-
enhance arginine catabolism; reduce the synthe- ing access to its database, and the Plant Pests and
sis of polyamine, creatine, and DNA; compete Diseases Research Institute, Tehran, for provid-
with lysine and arginine for transport and induce ing necessary facilities. Generous support extended
synthesis of aberrant proteins (D’Mello, 1994). by Profs. P. K. R. Nair, University of Florida;
Thus, the presence of harmful allelochemicals Estella D. Elakovich, University of Southern
in fodder may totally jeopardize one of the impor- Mississippi and Manuel J. Reigosa, University of
tant goals of agroforestry. The reports that Vigo, Spain, is gratefully acknowledged. The
mimosine has also caused toxic effects in humans cooperation of Dr. Daizy Batish, Panjab Univer-
are even more disturbing (Brewbaker, 1989). Toxic sity, Chandigarh, India, deserves a thankful men-
allelochemicals or their degradation products may tion.
enter the human body in several ways: (1) through
direct consumption of allelochemical containing

material, (2) as a residue in crop plants grown in
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Allelopathic Interactions in Agroforestry Systems

I. INTRODUCTION tude and the rate of the world population growth,

III. TREE ALLELOPATHY: SOME

A. Azadirachta indica Eucalyptus has been promoted on a large scale

A. excelsa Casuarina equisetifolia Aqueous extract of mature Shade Balasubramanian and

Secale cereale theobromine, theophylline, Rizvi and Rizvi, 1984, 1992b,

Agroforestry Target Plant parts/ Uses of agroforestry

Agroforestry Target Plant parts/ Uses of agroforestry

Agroforestry Target Plant parts/ Uses of agroforestry

be inhibited by 1 mM mimosine. Phaseolus aureus pest controlling properties to AF species. Thus,

Allelochemical Chemical nature Natural source Ref.

Ailanthone Quassinoid Ailanthus altissima Heisey, 1996

AF species Plant part/chemicals Pathogen Reference

Neem cake Fusarium solani Jeyarajan et al., 1987

Seed kernel extract Xanthomonas campestris Eswaramurthy et al., 1993

In agroecosystems, nurseries, and agroforestry

direct consumption of allelochemical containing

of Acacia tortilis in agroforestry systems of arid re- Ecol. 9: 21–28.

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