Chemoecology (2017) 27:1–24
DOI 10.1007/s00049-016-0225-x
REVIEW
Allelopathy in agro-ecosystems: a critical review of wheat
allelopathy-concepts and implications
Farhena Aslam1 • Abdul Khaliq1 • Amar Matloob2 • Asif Tanveer1
Saddam Hussain3 • Zahir Ahmad Zahir4
Received: 5 December 2015 / Accepted: 11 October 2016 / Published online: 19 October 2016
 Springer International Publishing 2016
Abstract Allelopathy is an important mechanism of plant
interference mediated by the release of plant-produced
secondary metabolites or decomposition products of
microbes to the aerial or soil environment. It plays a key
role in natural as well as cultivated ecosystems. Allelo-
chemicals are released into the soil rhizosphere by a variety
of mechanisms, including volatilization, decomposition of
residues, and root exudation. Along with inhibitory/stim-
ulatory effects of allelochemicals, several other ecological
roles of these chemicals, including plant defense, nutrient
chelation, and regulation of soil biota, have been reported.
Wheat is extensively studied and used as an allelopathic
crop, and numerous chemicals are reported to be released
from the wheat living plants and decomposing residues. In
this review, we presented a contemporary synthesis of the
existing data that how wheat allelopathy can be exploited:
(a) to biologically control the insects, pests, and diseases,
(b) to enhance the soil quality by adding nutrients for crop
plants during decomposition from residues and ameliorate
soil environment for microbes, (c) to increase the crop
diversification by rotation while reducing the weeds and
Handling Editor: Liliane Ruess.
& Amar Matloob
amar.matloob@mnsuam.edu.pk
1
Department of Agronomy, University of Agriculture,
Faisalabad 38040, Pakistan
2
Department of Agronomy, Muhammad Nawaz Shareef
University of Agriculture, Multan, Pakistan
3
College of Resources and Environment, Huazhong
Agricultural University, Wuhan 430070, China
4
Institute of Soil and Environmetal Sciences, University of
Agriculture, Faisalabad 38040, Pakistan
CHEMOECOLOGY
•
pests infestation, (d) to develop the low-cost biological
pesticides with a novel mode of action from crop plants,
and (e) to confer tolerance against abiotic stresses. Based
on our hypothetical concepts and previous evidences, we
briefly discussed the mode of action of allelochemicals, and
extent and rate of their production based on crop growth
stage. We also addressed the interaction of root exudates
and allelochemicals with soil biotic and abiotic compo-
nents to explore the role of allelopathy in rhizosphere
ecology.
Keywords Allelochemicals  Ecosystem 
Microorganisms  Rhizosphere  Weeds
Introduction
Allelopathy—the conceptual framework
Plants naturally produce secondary metabolites which are
important for biological and ecological functions and
interactions among neighboring plants and environment
(Duke 2015; Cheng and Cheng 2015; Sangeetha and Baskar
2015). These chemical interactions are complex, diverse,
and provide many advantages to the donor plant (Reigosa
et al. 1999; Reigosa et al. 2013; Cheng and Cheng 2015).
Sometimes, allelochemicals at low concentration also
behave as growth regulators and show stimulatory effects
on receiver plants (Inderjit and Keating 1999; Duke 2006).
The term ‘allelopathy’ was introduced by an Austrian plant
physiologist Molisch (1937) to refer to biochemical inter-
actions between all plants, including microorganisms.
Molisch referred to the both inhibitory and stimulatory
biochemical interactions. Later, it was defined as any direct
or indirect, beneficial or destructive effect by one plant on
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2 F. Aslam et al.

another through production of chemical compounds (alle-
lochemicals/phytotoxins) that escape into the environment
(Rice 1984) from plant parts by leaching, stem flow, root
exudation, volatilization, residue decomposition, and other
processes in both natural and agricultural systems (Fig. 1;
Birkett et al. 2001; Ferguson et al. 2003). Khaliq et al.
(2011) defined allelopathy as the mechanism explaining
inhibitory and/or stimulatory interactions in soil–plant
interface through bioactive products produced by bio-
chemical pathways. An important point regarding
allelopathy is that its effects depend upon chemical com-
pounds released to the environment from living plant or
decaying plant parts (Dayan et al. 2000; Einhellig 2004). It
is thus distinct from competition that involves the removal
or reduction of some factors from the environment that is
required by some other plant sharing the habitat (Tukey
1969). Allelopathy reflects inter-disciplinary, challenging,
and complex studies to unravel myths and facts and also
invites the attention of global agronomist, botanist, soil
scientists, microbiologists, phytochemists, foresters, and
ecologists (Reigosa et al. 2013). Allelopathic studies to
understand various ecological problems, such as replanting
problems, in stubbles of several crops, poor crop stand, and
low crop yields in rotation, detrimental effects of living
plants, or their decomposition products on plant growth and
crop yields, autotoxicity, and direct interference by certain
weeds and crops form an important area of continued
research.
In this review, we discussed the present state of
knowledge on wheat allelopathy, prospects, and challenges
regarding exploitation of wheat allelopathy in agro-
ecosystems, physiological and ecological mechanisms
underlying wheat allelopathy, effect of wheat allelopathy
on soil microorganisms, implications for rhizosphere
ecology, and genetic bases of wheat allelopathy. Efforts are
undertaken to reveal that how this eco-physiological phe-
nomenon facilitates ecological weed management, nutrient
acquisition and recycling, and crop growth stimulation, and
confers biotic and abiotic stresses tolerance in the back
drop of sustainable agriculture. Rather than being exhaus-
tive, this review paper is aimed to provide a critical update
of the current state of wheat allelopathy in the domains of
plant protection, environmental safety, and resistance
breeding along with future research thrusts.

Fig. 1 Possible mechanisms of

allelochemicals release and Plant aqueous
transformation From living Plant debris/
residues extracts
plants

Directly through Decomposition of plant material and Absorbtion or

root exudation release of allelochemicals in to the soil adsorbtion in soil

Action of soil microbes

Degradation Transformation
products, retention in /degradation
soil

Plant exposure
(Receiver species)

Stimulation/Inhibition

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Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
Role of allelopathy in agro-ecosystems
Allelopathy plays an important role in natural ecosystems,
and has the potential to become an important tool in cul-
tivated ecosystems too (Olofsdotter 1998; Tesio and
Ferrero 2011; De Albuquerque et al. 2011). Xuan et al.
(2004) proposed that if crop allelopathy is properly
exploited for agricultural practices and production, agro-
nomic significance could be applied to: (a) biologically
control of insects, pests, and diseases, (b) enhancement of
soil quality by adding nutrients for crop plants during
decomposition from residues and ameliorate soil environ-
ment for microbes, (c) increasing crop diversity by rotation
while reducing the weeds and pests infestation, and if crops
are appropriately rotated improvement of soil quality,
(d) development of low-cost biological pesticides with
novel modes of action from crop plants for sustainable pest
management, and (e) development of tolerance against
abiotic stresses. Biochemical interactions take place when
allelopathic compounds produced by one plant are released
into the environment and influence the growth and devel-
opment of another plant. Thus, allelopathy can be defined
by interaction among plants and microorganisms through
chemical pathways. These interactions may be stimulatory
or inhibitory. In agricultural practices, suppressive effects
may be exploited for pest and weed control (Khanh et al.
2005; Lithourgidis et al. 2011).
Weed management
Indiscriminate use of herbicides has resulted in serious
ecological implications, such as resistance and shifts, in
weed populations, minor weeds becoming dominant,
greater environmental pollution, and health hazards (Ber-
tholdsson 2012; Khaliq et al. 2012a). Research attention is
now focused on finding alternative strategies for weed
management. Allelopathy is a tool that can be manipulated
to manage weeds in agro-ecosystems (Tesio and Ferrero
2011). Use of allelopathy for weed management could be
either through direct natural allelopathic interactions, crop
plants, or through the use of allelochemicals as natural
phytotoxic compounds (Singh et al. 2003; Bhowmik and
Inderjit 2003; Jabran et al. 2015). A number of crop plants
with allelopathic potential can be used as cover, green
manure, and smother crops for handling weeds by making
anticipated manipulation in the cultural cropping patterns
(Weston 1996; Singh et al. 2003; Khanh et al. 2005). These
can be suitably rotated and intercropped with main crops to
manage the target weeds selectively. The use of crop
aqueous extracts, residues/mulches can also give desirable
benefits. These approaches for weed management reduce
the deleterious effect of current agriculture practices and
input costs in agro-ecosystem (Singh et al. 2003; Tesio and
3
Ferrero 2011; De Albuquerque et al. 2011; Jabran et al.
2015). Not only terrestrial weeds, even allelopathy can be
suitably manipulated for the management of aquatic weeds.
Allelochemical present in plants as well in microbes can be
directly used for weed management on the pattern of her-
bicides (Kennedy 1999; Reigosa et al. 2013; Jabran et al.
2015). Allelochemical effect on weed species can be
selective (Xuan et al. 2005). Allelopathic interactions can
be used in diverse cropping patterns often for weed man-
agement and suppression (Weston 1996; Weston and Duke
2003; Tesio and Ferrero 2011; Jabran et al. 2015; San-
geetha and Baskar 2015; Cheng and Cheng 2015; Duke
2015). Allelopathic potential for weed management under
field conditions has been utilized in various ways, i.e.,
surface mulch (Khaliq et al. 2015), incorporation into the
soil (Matloob et al. 2010), aqueous extracts (Khaliq et al.
2013), rotation (Narwal 2000), smothering (Narwal and
Sarmah 1996; Singh et al. 2003), mix cropping/intercrop-
ping (Hatcher and Melander 2003; Iqbal and Cheema
2007), or combination of allelopathic plant extracts with
reduced dose of synthetic herbicides (Khaliq et al. 2012a).
Allelopathic research can be applied to many current weed
problems in agricultural systems (Narwal 1999). Utiliza-
tion of allelopathic properties of native plant/crop species
offers promising opportunities for this purpose. Allelo-
pathic potential of crops, such as sorghum (Sorghum
bicolor L.) (Weston 1996; Cheema et al. 2003), sunflower
(Helianthus annuus L.) (Macias et al. 2002; Weston and
Duke 2003; Anjum and Bajwa 2005), rice (Oryza sativa L.)
(Kong et al. 2006), and wheat (Triticum aestivum L.) (Wu
et al. 2001a; Labbafi et al. 2010; Khaliq et al. 2011, 2012b)
against weeds of economic significance has been reported.
Insect pest and disease management
Estimates indicate that on a global scale, insect pests and
disease causing pathogens cause a yield reduction of 18
and 15 %, respectively, in major food and cash crops
across the world (Oerke and Dehne 2004). Natural
compounds of plant origin manifesting allelopathic
activity have been suggested as novel weapons to deter
insect pests and pathogen of field crops (Bertin et al.
2003). Contrary to synthetic pesticides that result in
ecological and health issues beside pest control, natural
compounds are on the other side of the window, as these
have a relatively short life, are biodegradable and water
soluble, lack halogen, and hence are comparatively safer
in the perspective of environmental toxicology (Duke
et al. 2002). Over 300 plants exhibit suppressive ability
against a wide spectrum of insects (Devakumar and
Parmar 1993). Ding et al. (2000) observed resistance in
some wheat cultivars against wheat midge (Sitodiplosis
mosellana), possibly owing to the production of p-
123
4 F. Aslam et al.
coumaric acid and ferulic acid. Ferulic acid treatment
killed the hatched larvae when the concentration excee-
ded 0.35 mg g-1 of fresh plant tissues. Valenzuela and
Smith (2002) reported inhibition of wireworm larvae
(Conoderus rudis) and consequently reduced population
in the field previously cultivated with buckwheat. Wheat,
rye, maize, and barley contain high concentration of
hydroxamic acid or tramine which are useful for the
control of higher population of aphid. Neem (Azadir-
achta indica) and Eucalyptus (Eucalyptus globulus) are
potent higher plants with arsenal of secondary metabo-
lites as a mean of natural defense. Their insecticidal
potential is manifested in the form of antifeedant prop-
erties, inhibition of post-embryonic development, and
larval emergence against a number of agricultural and
household pests (Farooq et al. 2011).
Roots are complicated morphologically and physiolog-
ically, and their metabolites are often released in large
quantities into the soil rhizosphere from living root hairs or
fibrous root systems (Bertin et al. 2003). Root exudates
containing root-specific metabolites have critical ecologi-
cal impacts on soil macro- and microbiota as well as on the
whole plant (Bertin et al. 2003). Phytotoxic compounds
released from decomposing residue or root exudation of
preceding crops play an important role in inhibiting soil
born plant pathogens (Batish et al. 2001; Bertin et al.
2003). Brassica spp. (B. nigra, B. napus, B. campestris, B.
juncea, and B. hirta) and Lepidium sativum are useful for
green manuring, weed and insect management (Mayton
et al. 1996). Potato dry rot (Fusarium sambucinum) was
controlled by the higher concentration of allyl-isothio-
cyanate (ranged 0.78–0.288 mg g-1 leaf tissue) present in
the Brassica species (Mayton et al. 1996). The addition of
these crops in rotation of those areas infested with aphid
can be helpful for the development of natural herbicides
(Rizvi and Rizvi 1992). Javed et al. (2007) reported 38 %
reduction in root-knot nematode (Meloidogyne javanica)
females per root, and 64 % in egg mass per root in response
to neem allelopathy (3 % w/w neem cake). Aiyaz et al.
(2015) evaluated the efficacy of 100 extracts of 20 different
plant species for their antimicrobial potential against tox-
igenic Fusarium verticillioides and Aspergillus flavus.
Among these, extracts of Toddalia asiatica, Decalepis
hamiltonii, Vitex negundo, and Psoralea corylifolia
reduced A. flavus and F. verticillioides incidence signifi-
cantly suggesting that hydropriming of maize seeds with
some plant extracts can be a potent alternative strategy to
control seed-borne toxigenic fungi.
Thus, pests and diseases can be control with natural
chemicals produced by allelopathic crops in the cropping
systems, antibiotic allelochemicals of plant origin, insect
pest resistant crop varieties, and their rotation as well as
intercropping.
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Plant growth regulation/hormetic effect
An et al. (1993a, b) postulated the phenomenon of biphasic
dose–response as an important universal biological char-
acteristic of allelochemicals. Stimulatory effect (not
necessarily beneficial) in response to low dose of a toxic
substance is termed hormesis (Duke et al. 2006). This term
was first used by Southam and Erlich (1943) after they
noticed that an oak bark compound was inhibitory to fungal
growth at higher concentration but stimulated the same at
lower concentrations. They coined the term using Greek
base ‘‘hormo’’ meaning to excite, the same root as used in
word hormone. Even Molisch, who coined the term ‘‘al-
lelopathy’’, was well-versed with the phenomenon of
allelopathic hormesis. He stated that the often observed
rule is confirmed that poisons and irritating compounds are
harmful at higher concentrations, but stimulatory in diluted
form (Chobot and Hadacek 2009). Likewise, Paracelus
proposed that poison is in the dose (all things are poison
and are not poison; only dose controls the process of tox-
icity). Most allelopathic research describes implications of
inhibitory allelopathic effects for weed management in
agro-ecosystems (Weston and Duke 2003; Weston 2005),
and have chosen to neglect hormesis, as it was a non-target
objective (Streibig 1980). Hence, hormesis in plants has
not been studied in detail until recently (Calabrese 2005;
Calabrese and Blain 2005). Studies document about
20–30 % stimulation due to hormesis effect in plants
(Cedergreen et al. 2005). Nevertheless, hormetic effect
observed in response to herbicides, such as glyphosate,
could be a potential factor in resistance development (Belz
2014) by increasing the competitiveness, reproductive
potential, and resistance evolution to a second weed control
measure rather than causing direct selection pressure (Belz
et al. 2011).
Allelochemicals are well known to induce hormetic
effects (Liu et al. 2011). Previous studies have revealed
the growth stimulatory potential of secondary metabolites
and compounds of natural origin (Ambika 2012; Maq-
bool 2010; Abbas et al. 2013). Allelopathic compounds
can confer abiotic stress tolerance and help maintain
metabolic events conducive to better plant growth (Ein-
hellig 1996). Allelopathic compounds are reported to act
as a first line of defense against biotic and abiotic
stresses (Jimanez et al. 2003; Asao et al. 2004). Identi-
fication of allelopathic donor plants exhibiting hormesis,
and threshold concentration, at which growth promotion
or stress tolerance is imparted, has formed an important
area of research in allelopathy nowadays. Stimulatory
potential of allelopathic crops due to hormetic effect can
be realized by treating seeds with aqueous extracts, foliar
or root application (Maqbool et al. 2012). Pre-sowing
seed treatment with growth-promoting substances has
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
been reported as an effective remedy to improve ger-
mination, growth, and yield under stressful environments
(Ashraf et al. 2008). Work done in Yugoslavia concluded
that the application of agrostemin at 100 g ha-1 either as
seed treatment of foliar application improved yield of
crops, such as wheat, maize, and sunflower by 10-15 %.
Besides yield increment, this strategy was equally good
in promoting oil yield of sunflower by 4 % (Biprodukt
1984). Aqueous allelopathic crop water extracts can also
be utilized as foliar treatment to modulate plant growth.
Basra et al. (2011) found that moringa leaf extracts were
more effective in promoting plant growth under ambient
and stressful environments than artificial cytokinin source
(benzyl amino purine; BAP). Chon et al. (2003a, b)
documented an increase in alfalfa root length to the tune
of 13–33 % by aqueous extracts of several plants of
compoistae family when applied at low concentration.
The plants producing flavonoids often exhibited auto-
toxicity (Weston ad Mathesius 2013). Sorghum is one of
the most potent and extensively studied allelopathic
crops (Weston and Duke 2003). Munir (2011), from her
wire house study concluded that application of 5 and
10 % aqueous extract of sorghum at vegetative growth
stage of wheat crop stabilized membrane integrity, and
enhanced yield and yield components under drought.
At low concentration, these may act as growth regu-
lators (Narwal 1994). Growth regulators are exogenous
non-nutritional compounds that influence the growth and
composition and development of plants and functions
with the endogenous phytohormones groups. Their
actions include growth regulation and retardation, flower
initiation, leaf senescence, and stimulation of biomass
production (Narwal 2006). Plant allelochemicals provide
a number of growth-promoting hormones, such as cyto-
kinins from rice root exudates and moringa leaf sap
(Soejima et al. 1992; Makkar and Becker 1996),
brassinolides from Brassica spp. (Maugh 1981) and
brassinosteroids (Andrzej and Hayat 2009). Bio-regula-
tors have shown increase plant growth and yields of
major crops. However, difficulties in their commercial-
izing have been encountered, thus limiting their
commercial use. This suggests the need for further
investigations to optimize the hormetic allelopathic
effects with scientific evidence. Studies determining the
optimum concentration of allelopathic product as well as
threshold concentration of the secondary metabolites at
which hormetic effect is observed are inevitable for
large-scale utilization of such an approach for growth
promotion in field crops under abiotic stresses. Need is
also felt to explore the mechanisms responsible for
growth modulation by allelopathic products at lower
rates.
5
Crop nutrition
Soil moisture availability and nutrient uptake is most
important factors for regulating crop growth. Soil is an
important medium for providing most nutrients to plants
with some exceptions. Plants take up nutrients from the soil
solution through their roots. A relationship between the
plant-released allelochemicals and nutrient uptake has been
observed. Allelochemical activities are well-defined by
form, quantity, and balance of nutrients present in soil
(Karmarkar and Tabatabai 1991). In some cases, allelo-
chemicals are used as to improve the nutrient uptake.
Oxalic acid and citric acid improved the uptake of K, Mg,
P, and Fe under deficient conditions and consequently
improved the root and shoot growth of plant under stress
conditions (Gent et al. 2005). Under low phosphorus con-
ditions, plants produce phosphatase which improves the
phosphorus availability through hydrolysis process (Gilbert
et al. 1999). Allelochemicals help to bind the nutrient
radicals in chelate form, and improve their stay in rhizo-
sphere to reduce losses. Beans acquire nutrients through
this process by exuding potential chelators, such as citrate
and oxalate, etc. (Delhaize et al. 1993). It is helpful to
avoid toxicity of metal ions and released the require ones
(Jabran et al. 2012). However, some time allelochemicals
have adverse effect on the uptake of nutrients. For exam-
ple, ferulic acid and cinnamic acid decrease the availability
of Fe and phosphorus and other nutrients. This reduction is
attributed to altered cell membrane permeability due to the
presence of these allelochemicals (Franche et al. 2009).
Recently, the role of allelopathic compounds in biological
nitrification inhibition has gained attention as a mean to
improve nitrogen use efficiency (Jabran et al. 2012). The
phenolics released from plant roots can suppress the oxi-
dation of NH4? to NO3- by inhibiting the activity of
nitrifying bacteria (Rice 1984). A recent breakthrough in
the biological nitrification inhibition (BNI) by allelopathic
mechanism comes from the work of Subbarao et al. (2013).
These authors’ explored quantification and characterization
of the BNI function of sorghum that covered inhibitors
production, functionality, and factors regulating their
release as well their chemical identity. Results revealed
that sorghum releases two types of nitrification inhibitors,
i.e., hydrophobic and hydrophilic. Release rates of these
BNIs varied from 10 to 25 ATU (allylthiourea units) g-1
root dry weight day-1. The addition of hydrophilic BNIs at
10 ATU g-1 soil inhibited nitrification by 40 % over a
30 day incubation period. Two BNI compounds isolated
were sorgoleone (13.0 lM, from hydrophobic fraction) and
sakuranetin (0.6 lM, from hydrophilic fraction). Adding
further, Tesfamariam et al. (2014) revealed genotypic dif-
ferences for nitrification inhibition among sorghum
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6 F. Aslam et al.
cultivars owing to variation in sorgoleone production and
release. These authors also established that purified sor-
goleone inhibited nitrosomonas activity and hence soil
nitrification. Thus, allelochemicals have potential of
nutrient regulation and cycling in agro-ecosystem, and
offer an eco-friendly and sustainable way to manage the
crop nutrient requirements and nutrient related problems.
Allelopathy and rhizosphere ecology
The chemicals exuded from living or actively growing
roots into the soil are broadly termed root exudates. A wide
array of these exudates regulate the microbial community
in rhizosphere, encourage beneficial symbiosis, change the
physcio-chemical characteristics of the soil, and inhibit the
growth of competing plant species (Nardi et al. 2000). The
ability of plant roots to exude biochemically diverse
compounds in the rhizosphere is one of the most interesting
features of plant roots (Marschner 1995). Metabolites
exuded through roots include carbohydrates, amino acids,
proteins, vitamins, and organic compounds. It is estimated
that through plant root exudation, 5–21 % photosyntheti-
cally fixed carbon is transferred to the rhizosphere
(Marschner 1995; Pinton et al. 2001). In the rhizosphere,
root exudates act as an important source of nutrients and
carbon for microorganisms and help in the early colo-
nization of rhizospheric bacteria (Bacilio-Jimenez et al.
2003). Microorganisms in rhizosphere have continuous
access to the flow of high- and low-molecular weight
substrates (sugars, amino acids, organic compounds, and
allelochemicals) which are exuded by roots. This flow,
with specific biological, physical, and chemical factors, can
markedly affect the microbial activity and community
structure of the rhizospheric soil (Brimecombe et al. 2001).
The low-molecular weight compounds of root exudates act
as allelochemicals that participate in various interactions of
rhizosphere between plant and microorganisms (Bais et al.
2006). The phenolic compounds, such as flavonoid in
legume roots, are responsible for the activity of Rhizobium
meliloti gene which activates the nodulation process
(Peters et al. 1986; Weston and Mathesius 2013).
Wheat allelopathy
Wheat is an important cereals crop that is extensively
studied and is also used as an allelopathic cover crop. It has
the ability to produce and exude allelochemicals under field
conditions which adversely affect weed growth (Wu et al.
2001b; Bertin et al. 2003). Numerous chemicals, such as
hydroxamic and phenolic acid and short-chain fatty acids,
are released from the wheat living plants and decomposing
residues which are responsible for wheat allelopathy.
Several studies have been conducted for the identification
123
of these wheat allelochemicals (Wu et al.
2000b, 2001a, b, c), their mode of release and degradation
(Fomsgaard et al. 2004), and screening of allelopathic
varieties and further improving the allelopathic potential at
genetic level (Bertholdsson 2005; Villagrasa et al. 2006;
Mahmood et al. 2013). In the late 1960s, Guenzi et al.
(1967) and Kimber (1967) reported that wheat residue
allelopathy varied among cultivars. Further studies found
that allelochemicals production varied among plant parts,
including root, shoot, and leaves (Villagrasa et al. 2006)
and infested rhizospheric soil (Khaliq et al. 2011). Wheat
allelopathy needs to be further studied to elaborate new
potent wheat varieties, effect of wheat on weeds, crops,
disease, and pests, isolation and identification of allelo-
pathic compounds, and their synergistic and additive
effects on physiological processes of plants. It is also
important to study wheat allelopathic activity in rhizo-
sphere and its implications for soil ecology and wheat
residue management in cropping systems.
Wheat allelochemicals
Allelochemicals are non-nutritional secondary metabolites
or waste products of the main metabolic pathways pro-
duced by living organisms (plants, microorganisms, fungi,
and viruses) and are not important for primary metabolism
but necessary for plant survival (Swain 1977; Schoonhoven
et al. 2005; Gordon-Weeks and Pickett 2009). Allelo-
chemicals produced as plant secondary metabolites or
decomposition products of microbes are the active media
of allelopathy (Cheng and Cheng 2015). Within this con-
text, the secondary metabolites involved in allelopathic
interactions have been designated as allelochemicals.
Allelochemicals are important for the growth, behavior, or
population biology and health of other organisms (Reese
1979). These compounds play a role in plant–soil, plant–
plant, plant–insect, plant predator, and plant–disease
interactions (Yaduraj and Ahuja 1996). From plants and
microorganisms, about 40,000 allelochemicals have been
identified and isolated (Rice 1984; Worsham 1989). These
chemicals are water soluble and can be released through
root exudation, leaching from plants by rain or decompo-
sition of residues (Fig. 1; Rice 1984). The allelopathic
activity of wheat has been attributed to hydroxamic acids
and its derivative compounds, phenolic acids (Blum et al.
1991; Bertin et al. 2003) and short-chain fatty acids (Lynch
1978; Tang and Waiss 1978).
Phenolic compounds
Wheat cultivars with strong allelopathic potential possess
higher quantity of phenolic compounds (Wu et al. 1998).
Guenzi and McCalla (1966) reported that wheat straw
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
contain p-cumaric, ferulic, vanillic, syringic, and p-hy-
droxybenzoic acids. Tsuzuki (2001) also reported that
buckwheat contains phenolic (chlorogenic, caffeic, and
ferulic) and fatty acids (palmitic, stearic, behenic, and
arachidic). Wu et al. (2000b) identified and quantified
phenolic compounds from wheat seedlings which were p-
hydroxybenzoic acid, cis-p-coumaric, trans-p-coumaric,
vanillic, syringic, cis, and trans-ferulic acids. Further
research results showed that wheat accessions significantly
varied in the production of these phenolic compounds in
the roots and shoots (Wu et al. 2000c, 2001a). These
compounds are reported as natural herbicidal source for
weed control and suggest that allelopathic wheat cultivars
can be used for natural weed suppression (Prasanta et al.
2003). Regarding wheat allelopathy, more concerned phe-
nolic compounds are p-hydroxybenzoic acid, p-cumaric
acid, vanillic acid, and ferulic acid (Guenzi and McCalla
1966; Koch et al. 1992; Lodhi et al. 1987). Higher con-
centrations of these phenolic compounds are present in
wheat straw (Koch et al. 1992). Studies establish the mode
of action of these phenolic compounds on nutrient uptake
(Baziramakenga et al. 1994), respiration (Penuelas et al.
1996), photosynthesis (Mersie and Singh 1993; Bazira-
makenga et al. 1994), protein synthesis (Mersie and Singh
1993), enzyme activity (Devi and Prasad 1992), membrane
permeability (Baziramakenga et al. 1995), hormonal bal-
ance (Holappa and Blum 1991), and plant water potential
(Einhellig 1986). Allelochemicals have both additive and
synergistic inhibitory effects (Jia et al. 2006). Wheat resi-
due allelopathy was highly associated with the quantity of
total phenolic contents. Wheat seedling exuded allelo-
chemicals through roots in the agar medium that were
identified and quantified (Wu et al. 1998). Total phenolic
acids varied among 58 wheat accessions ranging from 93.2
to 453.8 mg kg-1. Wheat accessions with higher phenolic
acids in their shoot inhibited the growth of annual ryegrass
(Lolium rigidum) (Wu et al. 2001a).
Hydroxamic acid and benzoxazinones
The hydroxamic acids most abundantly found in wheat are
DIMBOA (2,4-dihydroxy-7-methoxy-2H-1,4-benzoxazin-
3(4)-one) and DIBOA (2,4-dihydroxy-1,4(2H)-benzoxasin-
3-one) (Fig. 2), and studies report that their contents vary
among wheat genotypes (Niemeyer 1988; Wu et al. 2001a).
These are novel class of secondary products and considered
as natural defense agent against weeds, diseases, insects,
and pests (Niemeyer et al. 1989; Niemeyer and Perez
1995). These compounds are produced throughout the plant
life cycle and remain biologically active (Perez 1990;
Niemeyer 1988). The specific benzoxazinoids produced in
wheat are: ‘‘DIMBOA’’ [(2,4-dihydroxy-7-methoxy-2H-
1,4-benzoxazin-3(4)-one (benzoxazilinone)], ‘‘MBOA’’
7
6-methoxy-benzoxazolin-2(3H)-one (hydroxamic acid),
and ‘‘HMBOA’’ [2-hydroxy-7-methoxy-1,4-benzoxazin-3-
one (lactam)] and ‘‘DIMBOA glycoside’’ [2-(2,4-dihy-
droxy-7methoxy-2H-1,4-benzoxazin-3(4H)-one)-b-D-glu-
copyranoside (benzoxazilinone)] (Macias et al. 2006;
Morant et al. 2008). Wu et al. (2001d) found higher
DIMBOA content in root extracts as compared to shoots of
young seedlings and root exudates, while others reported
higher DIMBOA content in stem and leaves than roots
(Villagrasa et al. 2006). Givovich and Niemeyer (1995)
reported that five cereal aphid species (Rhopalosiphum
padi, Sitobion avenae, Schizaphis graminum, Me-
topolophium dirhodum, and Rhopalosiphum maidis) were
significantly affected when feed on DIMBOA and DIM-
BOA-glucoside at concentration 6–8 mM. They reported
that these species are also sensitive to the hydroxamic acid
present in wheat seedling that ranged from 2.5 to
3.0 mmol kg-1 FW in concentration.
Hydroxamic acid as DIMBOA glycoside accumulates in
vacuole of cereal plants, such as wheat, rye, and maize.
Upon damage, disrupted cell releases b-glucosidase,
cleaving the glucoside, and subsequently produces more
toxic aglycones compounds, DIMBOA in wheat and maize,
and DIBOA in rye (Fomsgaard et al. 2004). When DIM-
BOA or DIBOA are liberated by glucosidase, they are
generally degraded to their respective benzoxazilinones,
MBOA, and BOA (2(3H)-benzoxazolinone) (Nikus and
Jonsson 1999) and subsequently transformed to some other
compounds belonging to the acetamides, malonamic acids,
and aminophenoxazicones (Fomsgaard et al. 2004). The
transformation of the aglycone to the benzoxazilinone is
the result of non-enzymatic hydrolysis (Virtanen and Hie-
tala 1955). In aqueous solution, DIMBOA and DIBOA
have a half-life of less than 24 h (Bredenberg et al. 1962).
Both phenolic and hydroxamic acids were simultaneously
produced from wheat seedlings in growth medium (Wu
et al. 2000b; Huang et al. 2003). Several studies have been
conducted to ascertain the allelopathic effect of mixtures of
allelochemicals from wheat seedlings, but contradicting
results were found. Studies of Reigosa et al. (1999) showed
additive effects of phenolic compounds, leading to grater
activity. On the other hand, Inderjit et al. (2002) found the
antagonistic interactions among phenolic acids. The mix-
ture of DIMBOA, ferulic acid and vanillic acid had
antagonistic effects against Lolium perenne and Myosotis
arvensis (Jia et al. 2006).
Fatty acids
These are recognized as a third important category of
allelochemicals involved in the wheat allelopathy (Wu
et al. 2008). Insoluble polysaccharides represent a major
fraction of wheat straw and anaerobic fermentation of these
123
8 F. Aslam et al.
Fig. 2 Some allelochemicals
found in Poaceae family
(Brooks 2008)
BENZOXAZILINONES
BOA (Rye), MBOA
(Wheat, Maize)
compounds results in the formation of short-chain fatty
acids with allelopathic activity (Lynch 1978; Tang and
Waiss 1978). Previous studies document significant con-
centration of acetic acid in wheat straw (Lynch et al. 1980;
Hairston et al. 1987). Extract from fermented suspension of
wheat straw was found to contain salts of acetic, propionic
and butyric acids. Pentanoic, isopentanoic, and isobutyric
were present in traceable amounts (Tang and Waiss 1978).
Propionic acid at a concentration of above 1.0 mM sig-
nificantly inhibited the root growth of herbicide resistant
and sensitive biotypes of annual rye grass, with complete
suppression being observed at concentrations above
5.0 mM. Compared with p-coumaric acid, this short-chain
fatty acid was inhibitorier to germination and seedling
growth of both the biotypes of rye grass (Wu et al. 2003a).
Long-chained carboxylic acids, such as stearic, oleic, and
linoleic acids, have also been identified from wheat (Dong
et al. 2005). These fatty acids significantly inhibited the
germination of weeds, such as Leptochloa chinensis, but
had no effects on its root growth.
Sterols and ketosteroids have also been reported as
wheat allelochemicals (Gaspar et al. 1999a, 1999b).
Evidence of allelochemical activity
In nature, a number of bioactive compounds exist with
unexplored characteristics. Some of these compounds may
have potential for use against pests and weeds (Vyvyan
2002). The knowledge of chemical relationship of these
compounds on sensitive plants may aid in developing new
herbicides (Macias et al. 2001; Vyvyan 2002). Macias et al.
(2002) isolated about 125 toxic compounds from sunflower
cultivars which showed phytotoxicity against several
weeds. Haig et al. (2005) screened 130 plant species to
explore their toxic compounds against annual ryegrass.
Amongst agricultural crops, wheat and rye are important
cereal crops that produce significant quantities of hydrox-
amic acids and phenolic compounds. Isolation of these
allelochemicals from plants and subsequent test in bioas-
says has provided substantial proof for their toxicity. The
123
BENZOXAZINOIDS
LACTAMS HYDROXAMIC ACID
BOA (Rye), MBOA DIBOA and DIBOA (Rye),
(Wheat, Maize) DIMBOA (Wheat, Maize)
activity of allelochemicals, such as MBOA and DIMBOA,
produced primarily by wheat and maize, has been studied
in bioassays (Perez 1990). The root growth of Avena fatua
was reduced by 50 % by the application of MBOA and
DIMBOA at 0.5 and 0.7 mM concentration (Perez 1990).
Another study reported greater phytotoxicity of MBOA
than DIMBOA (Blum et al. 1992). Burgos et al. (1999) also
found that high concentration of BOA and DIBOA were
correlated with greater inhibition of goosegrass (Galium
aparine). The activity of BOA and DIBOA was investi-
gated against weed species, such as barnyard grass
(Echinochloa crus galli (L.) P. Beauv.), garden cress (L.
sativum), and crop species cucumber (Cucumis sativa L.)
and snap bean (Phaseolus vulgaris L.). Results indicated
that the shoot and root length suppression of barnyard
grass, garden cress, and cucumber was pronounced at
higher concentrations of BOA and DIBOA (Chase et al.
1991).
Chum et al. (2010) evaluated five benzoxazinoides
BOA, H-BOA, Me-BOA, M-BOA, and Hy-BOA for their
phytotoxic potential against three weed species (Phalaris
minor Retz.), (E. crus galli), and (Senna occidentalis L.).
The results indicated the phytotoxicity of benzoxazinoids
on weed germination and growth were concentration
dependent. In all test species, maximum effect was
observed with the M-BOA and Me-BOA, followed by
H-BOA, while BOA was least phytotoxic compound.
Seeds of P. minor and E. crus-galli were more sensitive to
all derivatives of benzoxazinoids than S. occidentalis.
Effect of growth stages on allelochemical production
Wu et al. (2001d) screened 58 wheat accessions to study
the production and accumulation of hydroxamic acids in
17-day-old roots and shoots. Fifty accessions had higher
levels of hydroxamic acids in shoot and root with an
average range of 439.4 and 643.0 mg kg-1, respectively.
Contrary to these findings, Mogensen et al. (2006) reported
that winter wheat cultivars had lower concentration of
DIMBOA in roots than leaf tissues at the early growth
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
stages. Copaja et al. (1999) studied the production of
DIBOA and DIMBOA in three wheat cultivars and found
that the production of these compounds was higher in first
week of germination but began to decline after seven days.
They also reported that DIMBOA level was higher than
DIBOA in the leaves of three cultivars, and were lower in
roots of two cultivars out of three. The concentration of
these compounds and their derivatives remain relatively
constant in plant roots while concentration decline gradu-
ally in the shoot with plant age (Mogensen et al. 2006).
Several studies reported the detection of hydroxamic acid
and their derivatives in ungerminated wheat seed (Copaja
et al. 1999; Villagrasa et al. 2006).
Mode of action of allelochemicals
Allelochemicals cause a generalized cytotoxicity through
number of physiological effects (Einhellig 2004). A num-
ber of physiological effects induced in response to
allelochemicals include reduction in overall plant growth
(Weir et al. 2004; Mohamadi and Rajaie 2009), ion uptake,
water absorption and mineral nutrients (Akemo et al.
2000), leaf water potential, osmotic potential, shoot turgor
pressure (Sheteawi and Tawfik 2007), leaf area expansion,
dry matter production, photosynthesis (Batish et al. 2001)
and stomatal aperture size and diffusive conductance
(Gniazdowska and Bogatek 2005), protein synthesis, and
enzyme activities (***Muscolo et al. 2001; Bertin et al.
2007;). These physiological processes may be inhibited or
stimulated by allelochemicals in different ways. By the
action of phenolic compounds, cell membrane becomes
non-specifically permeable, disturbing the ion flux and
hydraulic conductivity in root. These changes in plant
membrane may upset the plant–water relationship, rate of
photosynthesis, respiration, stomata opening and closure,
and ion balance (Einhellig et al. 1985; Gerald et al. 1992).
Blum et al. (1999) also indicated that the depolarization of
cell membrane and ion fluxes was affected by phenolic
acids. Baziramakenga et al. (1997) concluded that cin-
namic and benzoic acid produced by quack grass have
negative allelopathic effect on soybean, inhibit the germi-
nation and seedling growth and biomass by altering
membrane permeability, transport of ions and reduction of
chlorophyll contents by damaging thylakoid membrane.
Barkosky and Einhellig (1993) reported that p-coumaric,
ferullic, salicylic, and caffeic acids cause water stress in
treated seedlings of Glycine max and Sorghum bicolor.
Lettuce plant treated with BOA resulted in 10 % reduction
of relative water content which was simultaneously corre-
lated with the decrease in leaf water potential (Table 1:
Sànchez-Moreiras and Reigosa 2005). Gulzar et al. (2014)
indicated that the reduction in relative water content of leaf
initially caused stomatal closure, and reduced the CO2
9
supply to mesophyll cells consequently reducing photo-
synthesis. Mersie and Singh (1993) found that phenolic
compounds, e.g., vanillic, p-coumaric, and ferulic acid,
inhibited protein synthesis and photosynthesis in velvetleaf
(Table 1). Muscolo et al. (2004) observed seed germination
inhibition in Pinus laricio grown in the soils nearby Fagus
sylvatica and P. laricio trees. These authors concluded that
the mechanism of germination inhibition involved the
disruption of mitochondrial or dark respiration, and this
disruption may involve metabolic enzymes of oxidative
pentose phosphate pathway and glycolysis. Several phe-
nolic acids reduce protein synthesis by reducing the
assimilation of certain amino acids into proteins (Hussain
et al. 2010). Different studies reported the inhibition of
germination owing to allelochemical-induced oxidative
stress (Gniazdowska et al. 2007; Javed 2011). The appli-
cation of BOA enhanced the production of reactive oxygen
species in shoot and root of mung bean which caused
oxidative stress and increased the lipid peroxidation, and
proline accumulation (Batish et al. 2006). It is also
hypothesized that abnormalities regarding the nutrient
uptake through plasma membrane decrease the root length
(Friebe et al. 1998). Chou and Chiou (1979) indicated that
rice straw incorporation reduced the available soil nitrogen
and cations Zn?2, Ca?2, Mn?2, Cu?2, and Na?. Autotoxins
that produce in cucumber also reduce ion uptake, PO43-,
Mg2?, NO3-, K?, and Ca2? (Wang and Wang 2005). Yang
et al. (2004a, b) reported that absorption of IAA oxidase
and some essential macro- and micro-nutrients was
reduced by various allelochemicals which inhibited the
root dry weight and root water content in germinating
mustard seedling.
Root growth is categorized by a high metabolic rate in
root tissues so it is affected to much greater extent by
the environmental stresses, such as production of alle-
lochemicals in the soil (Cruz-Ortega et al. 1998).
Baerson et al. (2005) found that BOA at 540 lM con-
centration inhibited root system expansion, resulting in
decreased root length and a complete failure of root hairs
and lateral root formation in 10-day-old seedling of
Arabidopsis thaliana. Similarly, Sànchez-Moreiras et al.
(2004) found that mitotic interference occured due to
hydroxamic acid in Lactuca sativa L. which inhibited the
seedling growth. Friedman and Waller (1983) found
negative effect of caffeine on tea and coffee hypocotyl
and radical elongation. This negative affect of caffeine
may be due to the restriction of cell division by
inhibiting cell plate formation or incorporation into
nucleic acid chain. It also causes sequence of abnor-
malities in root cells of maize (Anaya et al. 2002).
Phenolic acids (p-hydroxy phenyl acetic acid, p-cou-
maric, ferulic acids) may affect the photosynthesis by the
disruption of chlorophyll synthesis pathway.
123
10 F. Aslam et al.

Table 1 Mode of action of some allelopathic compounds

Compounds Mechanisms Receiver species References

Ferulic, p-coumaric, Stimulation of chlorophyll degradation mechanism Oryza sativa Yang et al. (2002)
o-hydroxyphenyl
acetic acid
P-hydroxybenzoic acid Inhibits seedling growth, induce water stress, stomatal closure Glycine max Barkosky and Einhellig
(2003)
Hydroxyamic acid Mitotic interference, inhibits seedling growth Lactuca sativa Sànchez-Moreiras et al.
(2004)
Caffeine Inhibits cell division, abnormal root growth Zea mays Anaya et al. (2002)
Caffiec acid Inhibits seed germination, plant growth, disruption of plant–water Euporbia esula Barkosky et al. (2000)
relationship, reduce chlorophyll contents
2-Benzoxazolinone Inhibits plasma membrane bound H?-ATPase in roots Avena fatua Friebe et al.(1997)
(BOA) Inhibits germination, seedling growth, induces oxidative stress Phaseolus Singh et al. (2005); Batish
aureus et al. (2006)
Disruption of plant–water relationship, adverse effect on transpiration Lactuca sativa Sànchez-Moreiras et al.
and photosynthesis (2005)
Inhibits seedling growth Raphanus Chiapusio et al. (2004);
sativa, Kato-Noguchi and Ino
Lactuca (2005)
sativa
Caffeic, p-coumaric, Induces water stress Glycine max, Barkosky and Einhellig
ferulic, salicylic acids Sorghum (1993)
bicolor
Benzoic acid and Disruption of membrane or alter membrane permeability, efflux of Lactuca sativa, Baziramakenga et al.
cinnamic acid ions, reduce chlorophyll content by damage of thylakoid membrane Glycine max (1995), (1997)
Phenolic compounds Reduction in hydraulic conductivity, net nutrient uptake Glycine max Baziramakenga et al.
(1995)
DIMBOA, MBOA Inhibits seed germination Avena fatua Perez (1990)
p-coumaric, vanillic, Inhibit photosynthesis and protein synthesis Abutilon Mersie and Singh (1993)
ferulic acids theophrasti

Allelopathic compound has also been reported as
intermediate in some plant hormones synthesis and
degradation, such as phenolic compound ferulic acid acti-
vates the synthesis of abscisic acid (Holappa and Blum
1991). The dihydroflavonone naringenin stimulates indole-
3-acetic acid oxidase which degrades the indole-3-acetic
acid (Stenlid 1970). Komai et al. (1981) and Watanabe
et al. (1982) reported that the sesquiterpene lactone
argrophylline A and b–selinene and sesquiterpene farnesol
act as anti-gibberellin and anti-auxin.
Wheat allelopathy and weed management
Allelopathic potential of wheat for weed and pest man-
agement has been well documented (Table 2; An et al.
1998; Wu et al. 1999). The allelopathic effect of alle-
lochemicals depends upon species and tissue
concentration (Belz and Hurle 2005), movement, per-
sistence and fate of such chemicals in soil (Inderjit
2001). Wheat allelopathy also varies with plant age and
more weed suppression was observed at vegetative and
post-harvest stages (Wu et al. 2000d). At vegetative
stage, allelopathy can be exploited to suppress weeds
through the exudation of allelochemicals into the growth
medium. These allelochemicals suppress the growth of
weeds and also decrease the need for application of
synthetic chemicals at the early growth stages. At har-
vest stage, concentration of allelochemicals increases in
plants root and shoot, which upon leaching into the soil
affect the emergence and seedling growth of weeds (Wu
et al. 2000d).
Wheat aqueous extracts
The aqueous wheat straw extract adversely affected the
emergence and seedling growth, and antioxidant system of
Trianthema portulacastrum (Khaliq et al. 2012b). Mah-
mood et al. (2013) screened the 35 wheat genotypes to
check their allelopathic potential and found that wheat
aqueous extract, residue incorporation, and rhizosphere soil

123
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and… 11

Table 2 Allelopathic influence of wheat on selected weed species

Source Weed species Effect References

Aqueous extract Trianthema portulcastrum Reduce germination by 64 %, shoot and root length by 43 Khaliq et al.
and 55 %, respectively (2012a, b)
Avena fatua Reduction in shoot (76 %) and root dry biomass (89 %) Mahmood et al.
(2013)
Amranthus retroflexus, Echinochloa Germination inhibition (30 and 12 %, respectively) and dry Zwain (1996)
crus-galli biomass reduction (126 and 58 %)
Eclipta prostrate L., Echinochloa Diminished root growth by 27 % and 25 %, respectively Chon and Kim
crus-galli L. (2004)
Amaranthus retroflexus, Stellaria Inhibition of germination and seedling growth Ma (2005)
media and Digitaria ciliaris
Lolium rigidum Inhibit germination and root growth Wu et al. (2000b,
2003)
Residue Trianthema portulcastrum Suppressed germination by 36 % and seedling dry biomass Khaliq et al.
incorporation/mulch by 65 %, reduce antioxidant activities (2011)
Avena fatua Reduction in shoot (72 %) and root dry biomass (86 %) Mahmood et al.
(2013)
Ivy-leaf morning glory, red rot Inhibition of emergence Lehman and
pigweed, Amranthus retroflexus, Blum (1997)
Ipomoea hederacea
Infested-rhizosphere Trianthema portulcastrum Germination inhibition (43 %), root and shoot length were Khaliq et al.
soil reduced (38 and 25 %), seedling dry biomass suppression (2011)
(49 %), also reduced activities of enzymatic antioxidant
Under field conditions Alopecurus myosuroides Huds. Root growth (-60 %) and seedling dry biomass (-50 %) Bertholdsson
(2012)
Rabi season weeds present in wheat Reduction in weed density (76 to 95 %) Mahmood (2013)
field
Morning glory, Ipomoea lacunose Reduced density Liebl and
Worsham
(1983)
Ipomoea spp. Seedling dry biomass was reduced (97 %) Shilling et al.
(1985)
Root exudates Secale cereal, Lolium rigidum Inhibited root elongation (25 %) Labbafy et al.
(2009); Wu
et al. (2000d)
Sinapis alba L. Inhibit root growth Belz and Hurle
(2004)
Different winter weeds in field trials Reduce the weed density, and seedling growth (75 %) Rizvi et al.
(2004)

significantly affected the germination and seedling growth
of A. fatua. A series of greenhouse experiments conducted
by Zwain (1996) aimed to evaluate the allelopathic
potential of wheat aqueous extract and residue incorpora-
tion against E. crus-galli and Amaranthus retroflexus. He
found that residue aqueous extract at 2 and 4 % concen-
tration inhibited the germination and seedling growth of
test species. The extract was less inhibitory on A. retro-
flexus than E. crus-galli. In other subsequent experiments,
the wheat residue incorporation was surprisingly more
suppressive to A. retroflexus than E. crus-galli and the
phytotoxicity of decomposed residue persisted for 6 weeks
and decreased afterwards. Rambakudzibga (1991) reported
that aqueous extract of wheat residues has phytotoxic
effects on the germination of shoo-fly plant (Nicandra
physalodes), pigweed (Portulaca oleracea), itchgrass
(Rottboellia cochinchinesis), and Amaranthus spp. hybrids.
Wheat aqueous extracts were also allelopathic to common
ragweed (Ambrosia artemisiifolia L.), pitted morning glory
(Ipomoea lacunosa L.), and to herbicide-susceptible and
resistant biotype of annual ryegrass (Wu et al. 2000a).
Wheat extracts also adversely affected the seed emergence
and seedling growth of Stellaria media, Digitaria ciliaris,
and A. retroflexus (Ma 2005).

123
12
Wheat root exudates
Phytotoxins produced by wheat seedlings can be released
via root exudation and can inhibit the germination and
seedling growth of several weed species (Wu et al. 2000d;
Belz and Hurle 2004; Bertholdsson 2005). Wu et al.
(2000d) conducted ‘‘equal compartment agar method’’
(ECAM) bioassays to study wheat seedling allelopathy
against annual ryegrass. This method successfully sepa-
rated the allelopathic effect of both competitive species
(wheat and ryegrass) and allowed study of the release and
accumulation of phtotoxins from wheat seedlings into the
agar medium (growth medium). These accumulated phy-
totoxins in the agar medium affected the growth of annual
ryegrass. This method was successfully employed for the
screening of 453 wheat accessions, and seedling allelopa-
thy differed among wheat accessions which inhibited the
growth of ryegrass roots ranging from 10 to 91 % (Wu
et al. 2000d). Bertholdsson (2004) similar screening
bioassays for 1104 wheat accessions (from South America,
Soviet Union, Africa, Asia, Sweden, America, and some
other European countries) were accomplished based on
agar medium. He concluded that certain wheat genotypes
produced phytotoxins as root exudates which inhibited the
root growth of perennial ryegrass by 50–60 %. Similarly,
some hydroponic culture bioassays were designed to assess
the allelopathic potential of root exudates and found that
wheat accessions varied in their allelopathic potential
against white mustard (Sinapis alba L.) root growth (Belz
and Hurle 2004). Wheat root exudate allelopathy has also
been verified in field trials and pot screening bioassays. In a
field trial, 200 wheat accessions were screened for their
allelopathic potential and most of the accessions inhibited
the seedling growth of weeds up to 75 % which was similar
to that achieved by hand weeding (Rizvi et al. 2004).
Wheat straw mulch/cover crop
The negative impact of wheat straw/residue has been
studied on agriculture production systems (Alsaadawi
2001; Wu et al. 2001c). It is acknowledged that wheat
straw can help manage pests, weeds, and diseases (Mu-
minovic 1991). Phytotoxic compounds are released from
the straw during decomposition, or produced by the
microorganisms which utilize the straw as carbon and
nutrient source which help to control the weeds (Chou
1992). Khaliq et al. (2011) found that wheat straw-amen-
ded and infested-rhizosphere soil had allelopathic effect on
T. portulacastrum and significantly reduced its germina-
tion, seedling growth, and biochemical attributes. Wang
et al. (2004) compared the effect of wheat straw as mulch
and chemical weed control on rice weeds. They found that
weeds Leptochloa chinensis L., E. crus-galli, and
123
F. Aslam et al.
Commelina diffusa were significantly decreased with
increasing rate of wheat straw mulch as compared to
chemical control alone.
Allelopathic cover and smother/rotational crops con-
siderably reduce the weeds infestation in crop lands.
Appropriate manipulation of these crops in agricultural
systems may provide a way towards non-chemical weed
management (Weston 1996; Chou 1999; Anaya 1999). The
cover crop residues, such as wheat, rye, barley reduced the
germination, and seedling growth of many weeds, espe-
cially annual broad leaf weeds (Barnes and Putnam 1983).
Phytotoxins, such as b-hydroxybutyric and b-phenyl lactic
acid, were identified in wheat and rye mulch (Shilling et al.
1985). Rye residues produce DIBOA which along with its
byproduct BOA reduced the emergence and growth of
weeds (Barnes et al. 1987).
The presence of phenolic acid and triterpenoids in wheat
straw residue makes it a suitable cover crop in no until
cropping system (Gaspar et al. 1999a, b). Wheat straw
mulch reduced the germination, seedling growth, and
seedling dry weight of several weed species (Muminovic
1991). Shilling et al. (1985) also reported that some broad-
leaved weed species were suppressed by allelopathic
activity of wheat mulch. Narwal (1998) evaluated that
residues of several cover crops including wheat had
allelopathic effect on the biomass and density of weeds.
They found that wheat straw inflicted 16.8 % reduction of
broad-leaved weeds but was least effective against grassy
weeds. Both above- and below-ground wheat residues were
allelopathic against broad-leaved weeds, such as redroot
pigweed, prickly sida, and ivy-leaf morning glory (Blum
et al. 2002). Contrary to these findings, biomass of both
grassy and broad leaf weeds was significantly reduced by
wheat straw (Alsaadawi 2008). According to Bertholdsson
(2012), if allelochemicals released from living wheat crop
are able to suppress the growth of herbicide resistant
weeds, such as black grass (Alopecurus myosuroides
Huds.), this will make wheat cultivation possible in areas
infested with this resistant weed.
Allelopathic wheat cultivars
The ability of a cultivar to suppress weeds under natural
settings is an important criterion for its commercial
acceptance (Worthington and Reberg-Horton 2013), and is
related to the allelopathic potential (Jabran et al. 2015)
besides several morpho-physiological attributes. Genetic
variability for the allelopathic trait is quite common (Tesio
and Ferrero 2011) in crop species (especially in cereals),
and offers a genetic pool for selection (Wu et al. 1999).
Table 3 summarizes the promising allelopathic wheat
cultivars from various parts of the world. For example,
Mahmood et al. (2013) while investigating the allelopathic
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and… 13

Table 3 Allelopathic wheat cultivars around the world

Cultivar Weed/test species suppressed Country References

Tasman Lolium rigidum China (Wu et al.

2003a,b)
Vinjett Lolium perenne Sweden (Bertholdsson
2005)
Astron Apera spica-Venti L. Denmark (Mathiassen
Ritmo Chenopodium album L. et al. 2006)
Stakado Stellaria media L.
No 6 Lankao Lolium rigidum China (Zuo and Ma
No 22 Xiaoyan 2007)
Shiraz Convolvulus arvensis Iran (Labbafi et al.
Niknejad 2008)
Roshan Secale cereal L. Iran (Labbafy et al.
2009)
Mohan 73 Lolium perenne Tunis (Bertholdsson
2010)
Rasool, Mahdavi, Navid, Karaj1, Adl1, Gaspard, Gascogen, Amaranthus retroflexus, Hordeum spontaneum and Iran (Miri 2011)
Bam, Arvand, Baiat and Azadi Avena fatua
Rohtas 90, V6007, Pak 81, AS 2000, V7189, Bhakkar 2002, Avena fatua Pakistan (Mahmood
V6111, Chanab 2000, V6034, Uqab 2000 and V4611 et al. 2013)
MACS 6272, PBW 550 and RAJ 4077 Avena ludoviciana Durieu., Celosia argentea L. India (Rajput and
and Echinochloa crus-galli (L.) P. Beauv. Rao 2013)
Xiaoyan Descurainia sophia (L.) Webb ex Prantl China (Zuo et al.,
Sin-Altheeb 2014)
Coupon
Jing411 28 weed species of wheat fields (narrow and broad- China (Zhang et al.
leaved) 2016)
AARI-2011 and Lasani-2008 Phalaris minor Retz. Pakistan (Aslam et al.
2016)

activity of 35 Pakistani wheat cultivars against A. fatua
found a pronounced genetic divergence regarding allelo-
pathic activity among wheat cultivars, as 11 out of 35
cultivars had a high allelopathic activity. Thus, identifica-
tion of wheat cultivars with strong allelopathic potential
can contribute directly to weed suppression by their
inclusion into crop rotation, or their use in breeding pro-
grams to incorporate allelopathic trait into future genotypes
making them compete as well as suppress weeds more
effectively (Mahmood et al. 2013). Allelopathic crop cul-
tivars can produce and release their own ‘herbicides’, i.e.,
allelochemicals to interfere with weeds (Zhang et al. 2016).
The identification of cultivars with greater allelopathic
potential and transfer of this characteristic to improved
cultivars could help restore a property that was inadver-
tently lost while breeding for fast growth rate and higher
yield (Wu et al. 1999) without incurring extra cost for weed
control. Understanding the genetics of crop allelopathic
activity remains a germane issue to be researched (Cheng
and Cheng 2015) and successful genetic manipulation of
this trait has been demonstrated (Olofsdotter 1998). Con-
certed research efforts have been devoted to unveil the
genetics of this trait, since the production and exudation of
the allelochemicals are regulated by the action of several
genes. Allelopathic crop cultivars can be developed using
the conventional breeding or transgenic approaches (Fra-
gasso et al. 2013; Cheng and Cheng 2015) for inexpensive,
and environment benign weed control. Promising cultivars
would be those that integrate weed suppressing ability with
other traits of interest, such as high yield, early maturity,
disease resistance, and quality traits (Gealy and Yan 2012).
Bertholdsson (2010) bred spring wheat for improved
allelopathic potential by conventional breeding. Based on
preliminary screening, a Swedish cultivar (Zebra) with low
potential allelopathic activity and a Tunisian cultivar
(Mohan 73) with high-potential allelopathic activity was
crossed and material originating from this cross was used
in the evaluation of a potential breeding program for high
allelopathic activity. In the F6 and F7 generations, three
breeding lines with an average improved allelopathy of

123
14
20 % and one line with an unimproved allelopathy activity,
but with the same phenotype as the high allelopathic lines,
were tested together with the low allelopathic Swedish
parent. An important finding of the field study was a 19 %
average reduction in weed biomass accompanied by a 9 %
reduction grain yield in the high allelopathic lines. The
biomass accumulation by high allelopathic lines was less
than the control. Allelopathic potential of crops due to its
polygenetic nature is weakly correlated with the yield and
other traits (Olofsdotter et al. 1999).
Weed suppressive ability of wheat is lower than that of
the barley and rye (Bertholdsson 2005, 2011). Bertholdsson
(2005) postulated a 60 % improvement in the weed com-
pletive ability of wheat if its allelopathic potential could be
raised to the level of the highly allelopathic barley culti-
vars. In the case of wheat, we can know the position of the
genes coding for production of DIMBOA, an allelopathic
compound in wheat, and modern genetic techniques or
conventional breeding can help to transfer such genes
between cultivars of the same species or between species.
Benzoxazinoids are the allelochemicals present in wheat as
well in rye. In wheat, their biosynthesis is maximum at
7 days after germination, while the same is achieved after
60 days in rye. Hence, transfer and incorporation of genes
from rye could extend the period of allelopathic activity in
wheat (Bertholdsson et al. 2012). The polyploidy level of
common wheat makes it tolerant to changes in its chro-
mosomes. However, part of the wheat genome that
disappears, to some extent, can be substituted with genome
of a related species indicating the potential of gene transfer
from other species. Using quantitative trait loci identifica-
tion, putative genes regarding weed suppressive ability
have been located on chromosomes 1A, 2B, and 5D which
can be useful in breeding wheat with higher allelopathic
potential (Zuo et al. 2012). However, contrary to rice, a
successful allelopathic cultivar has not been developed so
far for wheat (Cheng and Cheng 2015). Coordinated
breeding programs focusing on the location of genes
involved in the production of allelochemicals, control of
allelopathic activity, and mapping the populations between
allelopathic and non-allelopathic accessions can be crucial
in this regard (Fragasso et al. 2013). Due attention should
also be given to autotoxicity and detoxification potential
while breeding allelopathic cultivars (Cheng and Cheng
2015).
Wheat allelopathy and rhizospheric microorganisms
The free movement of allelochemicals within soil envi-
ronment is influenced by a wide array of crop and soil-
related management processes, such as plant growth,
weed control, crop compatibility within rotations, residue
management, nitrogen transfer, soil fertility, and
123
F. Aslam et al.
management of diseases caused by soilborne pathogens
(Narwal 2000; Einhellig 1995). Allelochemicals in the
rhizosphere are produced directly or indirectly from plant
roots, from precursor compounds released into the root
zone and subsequently transformed through biochemical
reactions (enzyme mediated) or abiotic (oxidation)
through the action of microbes or higher organisms (Tang
et al. 1989). Allelochemicals enter the soil environment
through the release of water soluble leachates from
underground and aerial plant parts (Tukey 1966; Guenzi
et al. 1967), root exudation (Wu et al. 2000b, b), break-
down product from the microbial decomposition of crop
residues (Harpers and Lynch 1981a, b; Chou and Patrick
1976), and the production of allelochemicals by root-as-
sociated microbial communities (Gurusiddaiah et al. 1986;
Thomashow and Weller 1988; Sarwar and Kremer 1995).
When one allelochemical molecule enters the soil envi-
ronment through above-mentioned processes, it may be
activated or detoxified by soil microorganisms or serve as
carbon source for the production of new toxins by
microorganisms in the soil (Liebl and Worsham 1983;
Blum and Shafer 1988). Blum (1998) reported that phe-
nolic compounds are readily transformed by the activity
of soil microorganisms from one form to another, such as
ferulic acid into vanillic acid and p-coumaric acid into p-
hydroxybenzoic acid. Understrup and his co-workers
found that plant-released BOA, H-BOA, and MBOA that
are transformed into three major groups (aminophenox-
azinones, malnomic acids, and acetamides) in the
presence of G. graminis var. graminis and G. graminis
var. tritici (Understrup et al. 2005). Fomsgaard et al.
(2004) also reported that BOA was transformed in the
presence of G. graminis var. tritici (fungal pathogen), but
such transformation was not observed in sterile soil.
Microorganisms also produce secondary metabolites
either directly or indirectly by decomposition of organic
residues (Ren et al. 2006). Microbial metabolism and co-
metabolism with phenolic compounds are helpful for the
production of phytotoxic compounds by deletion or addi-
tion of side groups (Martin and Haider 1976; Blum 1998),
polymerization of soil organic matter, or incorporation of
phenolic carbon into microbial biomass (Martin and Haider
1976; Haider et al. 1977). Secondary metabolites can have
phytotoxic effect on other plants (Kohli et al. 2001). Sec-
ondary metabolites when exposed to various chemical and
biological processes may prove toxic to other plants or
serve as carbon source for microorganisms (Blum et al.
1999). The inhibitory effects of plant residues were col-
lectively due to the toxins both from microorganisms and
plant residues (Norstadt and McCalla 1963). The avail-
ability, persistence, and biological activity of
allelochemicals is also regulated by microbial communities
(Inderjit 2001).
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
Preferential utilization of carbon source in soil affects
the soil microbial system and expression of allelopathic
phenomenon (Singh et al. 2001). The phenolic compounds
have higher energy content than simple sugars, so carbon
limited soil microorganisms would rapidly degrade phe-
nolic compounds to form carbon rich soils (Schmaidt and
Ley 1999). Rhizospheric microbes can bring about quali-
tative and quantitative changes in allelopathic compounds,
having implications for allelopathy (Inderjit 2005). In
many situations, the transformed products of microorgan-
isms are important for allelopathic growth inhibition
(Inderjit 2005). Several studies report on the significance of
microbial metabolites (Blum 1998). Blum et al. (1999)
reported that microbial activity in soil alters phenolic
chemistry and modified phytotoxicity. For example, in soil,
DIM2BOA and DIMBOA exuded from roots of corn and
wheat, respectively, were catabolized by soil microorgan-
isms to 2-amino-4,6,7-trimethoxy-3H-phenoxazin-3-one
and 2-amino-7-methoxy-3H-phenoxazin-3-one, respec-
tively. (Kumar et al. 1993). Weidenhamer and Romeo
(2004) reported that soil microorganisms transformed the
allelopathic compound hydroquinone glycoside arbutin
into hydroquinone and then into benzoquinone. Nair et al.
(1990) reported that regarding rye (Secale cereale)
allelopathy, microbial degradation of benzoxazinones is
very important. Gram-negative bacteria, Actinetobacter
calcoaceticus, isolated from field soil converted BOA to
2,20 -oxo-l,l0 -azobenzene (AZOB) (Chase et al. 1991).
Comparison of both revealed that AZOB was more inhi-
bitory to garden cress and barnyard grass than BOA (Nair
et al. 1990). In non-sterile soil, transformation of BOA into
2-amino-3H-phenoxazin-3-one (APO) inhibited radical
growth of barnyard grass. This suggested the greater phy-
totoxic potential of degraded byproduct 2-amino-3H-
phenoxazin-3-one for barnyard grass than its precursor
(Gagliardo and Chilton 1992). Another study also con-
cluded that MBOA was more suppressive than its precursor
DIMBOA to the germination and seedling growth of Tri-
folium incarnatum and Ipomoea hereracea (Blum et al.
1992).
Wachowska et al. (2006) conducted an experiment to
study the fungi and bacteria population in rhizosphere soil
of winter wheat cultivars, i.e., Roma and Sakwa. They
found that fungi of genus Penicillium species were more
dominant than others, which was 22.3 and 44 % in soils of
Roma and Sakwa, respectively. Regarding bacteria, higher
counts of genus pseudomonas were isolated. These authors
concluded that the population of fungi and bacteria is
dependent on species and developmental stage of wheat.
The species of these microorganisms particularly at the late
developmental stage may increase the grain yield of cereal
crops (Singh and Kapoor 1999). Grayston et al. (1998) also
suggested that the plant species affect the population of
15
microorganisms in rhizosphere. Population of bacteria of
genus Actinomycetales increase at flowering and milking
stage and Pseudomonas at milk and ear formation stage of
winter wheat (Wachowska et al. 2006). The microbial
counts were higher in rhizosphere of winter wheat than in
the fallow soil (Bergsma-Vlami et al. 2005).
Genetic architecture of wheat allelopathy
Substantial research efforts now have been undertaken to
improve the understanding of genetics of crop allelopathy
which once remained as the least studied area in the overall
allelopathy discipline (Wu et al. 1999). Successful genetic
manipulation of allelopathy has been demonstrated for
crops, such as rice (Olofsdotter et al. 1999, 2002; Amb and
Ahluwalia 2016) and sorghum (Weston et al. 2013); nev-
ertheless, more research is needed to have through insight
into the genetic control of allelopathic activity in wheat.
Wu et al. (2000a) screened 453 wheat accessions for their
allelopathic activity against rye grass. Allelopathic activity
was normally distributed within the collection from 50
countries suggesting the polygenic control of allelopathic
trait. Later on, these authors concluded that wheat
allelopathy is genetically controlled and a multigenic
model was proposed (Wu et al. 2001b) because of
involvement of complex biochemical pathways and distinct
categories of allelochemicals. Allelopathic effects showed
high heritability (55–95 %) during the whole life cycle of
the wheat (Zuo and Ma 2007). Nevertheless, heritability
was highest at the tillering stage and minimum at the grain
filling stage. Moreover, allelopathic potential was variable
and discontinuous throughout the wheat life cycle.
Research has concluded that wheat allelopathy is quanti-
tatively inherited (Wu et al. 2000b). This notion was
further supported by the finding of the studies that used
near isogenic wheat lines of Hartog (weakly allelo-
pathic) 9 Janz (strongly allelopathic). The BC2-Hartog
lines (back cross to Hartog) showed a weak alleopathic
activity as that of the Hartog; whereas, the allelopathic
activity of Janz lines was strong similar to that of Janz.
Niemeyer and Jerez (1997) studied the location of genes
responsible for the production of hydroxamic acids in
wheat. Euploids and aneuploids of variety Chinese Spring
(high in DIMBOA) were studied with the substitution lines
of a cross between variety Chinese Spring and variety
Cheyenne (low in DIMBOA). These authors proposed a
multigenic model for the accumulation of hydroxamic
acids in wheat. It was found that genes involved in the
transformation of DIBOA into DIMBOA were located on
chromosomes 4A and 4B, while the chromosome 5B may
contain genes for the transformation of methoxylated lac-
tam into DIMBOA. Moreover, there might be an inhibitory
gene on chromosome 4D. With the advent of modern
123
16
molecular techniques, the genetic background of allelopa-
thy, and underlying genetic and molecular mechanisms is
becoming more apparently known than ever before (Duke
et al. 2001).
Molecular techniques have been employed to study
genetic markers associated with wheat allelopathy (Wu
et al. 2003b). These authors studied the genetic control of
wheat allelopathy using a double haploid population of
wheat derived from cv. Tasman (strongly allelopathic) and
cv. Sunco (weakly allelopathic). Two major quantitative
trait loci (QTLs) associated with wheat allelopathy were
identified on chromosome 2B through restriction fragment
length polymorphism, amplified fragment length poly-
morphism, and microsatellite markers. Simple interval
mapping-based regression revealed that one QTL located
on chromosome 2B was responsible for about 29 % phe-
notypic variation in wheat seedling allelopathy. Zuo et al.
(2005) postulated that allelopathic activity of wheat was
enhanced with the evolution as the genome changed from
2n to 4n and to 6n. The identification of plant Cytochrome
P450s encoding the biosynthesis of benzoxazinones has
been acknowledged as a major breakthrough in wheat
allelopathy (Wu et al. 2008). Frey et al. (2003) demon-
strated that two genes Bx6 and Bx7 are involved in the
transformation of DIBOA into DIMBOA. Bxs genes (Bx1–
Bx8) regulating the entire biosynthesis of DIMBOA have
been reported to cluster on short arm of chromosome 4,
while Bx9 is located on chromosome 1 (Frey et al. 2003;
Rad et al. 2001). The cDNAs of five P450 s conducive to
DIBOA biosynthesis have been isolated (Nomura et al.
2002).
New insights to improve the genetic architecture of wheat
allelopathy
Both the conventional breeding methods and those devel-
oped using transgenic technology can be applied in the
breeding of allelopathic cultivars. Genetic variability for
allelopathic trait is widespread within wheat cultivars and
offers promising opportunity to select cultivars with the
greatest allelopathic ability. The quantity of allelochemi-
cals produced by a wheat cultivar can be an intrinsic
determinant of allelopathic potential, since the allelopathic
potential of the wheat cultivars is positively correlated with
the total phenolic contents. Direct screening for allelo-
chemicals will help complement the results of biological
screening and can be used to elucidate the genetic control
of this trait among wheat cultivars. Attempts to enhance
allelopathic ability of crop plants using the traditional
breeding have not met with fair degree of success; thus,
biotechnology is expected to play a major role (Tranel and
Horvath 2009).
123
F. Aslam et al.
The molecular biology and transgenic approaches have
been successfully exploited to enhance production and
secretion of sorgoleone in sorghum (Duke et al. 2001;
Weston et al. 2013). The use of restriction fragment length
polymorphism has markedly increased the efficiency of
mapping QTLs as compared to isozymes and morpholog-
ical markers (Wu et al. 1999). This technique scores greater
number of markers in a single population than other tech-
niques. The availability of suitable mapping populations
has been regarded as one of the major limiting factor in
studies dealing with the genetic control of allelopathy
(Worthington and Reberg-Horton 2013). Marker-assisted
backcrossing works best in situation in which fewer than
five QTLs control allelopathy (Courtosis and Olofdotter,
1998). Involvement of more number of QTLs makes it
difficult to recover important agronomic parameters from a
non-allelopathic recurrent parent and to simultaneously
retain all the QTLs responsible for allelopathic activity
from donor plant. The linkage analysis of the genetic
markers and QTLs is supposed to complement allelopathic
activity through marker-assisted selection for breeding
wheat with strong allelopathic potential (Wu et al. 1999).
With the increased availability of single nucleotide poly-
morphism and diversity arrays technology markers,
association mapping has emerged as a potential tool for
allelopathic researchers (Worthington and Reberg-Horton
2013). Effective molecular markers can be worked out and
used in marker-assisted selection for improved allelopathic
potential of wheat if additional fine resolution QTLs con-
trolling allelopathy are discovered. However, such marker-
assisted selection may be hindered by the large number of
minor effects QTLs (Worthington and Reberg-Horton
2013). Contrary, while the QTL mapping is crucial for
marker-assisted selection, it seldom results on gene dis-
covery, although hundreds of candidate genes may be
located within a single QTL spanning only 5–10 cM
(Bertin et al. 2008).
Wu et al. (2008) proposed that overexpression of P450 s
genes as a tool to modify the flux across biosynthesis
pathways that regulate the production and accumulation of
allelochemicals. Alternatively, the possibility of introduc-
ing foreign P450 s genes into intended crop, such as wheat,
may lead towards engineering novel biochemical pathways
and synthesis of bioactive allelochemicals. Identification of
genes controlling the biosynthesis of allelopathic com-
pounds can achieved through activation tagging (Hayashi
et al. 1992), use of gene knockout libraries (Krysan et al.
1999), and discovery, isolation, and purification of
enzymes and related metabolites (Yang et al.
2004a, 2004b). Genes specific to benzoxazionoides
biosynthesis and activity have been implicated (Frey et al.
2009). Transgenic approaches facilitating the isolation,
Allelopathy in agro-ecosystems: a critical review of wheat allelopathy-concepts and…
cloning, and introduction in the sense orientation of PAL2
(phenylalanine ammonia lyase) genes will greatly increase
the overexpression of PAL activity which is believed to be
positively correlated with the biosynthesis of most phenolic
compounds. However, the feasibility of these transgenic
approaches is challenged by complex genetic nature of
allelopathy. Introduction of multigene expression system
and its regulation to produce and localize enough quantity
of desired allelochemicals in the proper plant tissue/organ
is a difficult task (Bertin et al. 2008). Furthermore, besides
these genetic barriers, the social and political opposition to
transgenic crops is also crucial aspect while using trans-
genic approach (Worthington and Reberg-Horton 2013).
Recently, metabolomics (non-targeted qualitative or
quantitative analysis of all low-molecular weight metabo-
lites in a biological sample; Kim et al. 2011) had emerged
as an important aid to the biochemical profiling with ref-
erence to allelopathy (D’Abrosca et al. 2013). Contrary to
protemics, transcriptomic and genomic approaches which
aim to predict that what will happen to a biological system,
metabolomics intends to unravel what is actually happen-
ing (Aliferis and Jabaji 2011). ***Asaduzzaman et al.
(2014) demonstrated the usefulness of metabolomics
analysis in differentiating allelopathic and non-allelopathic
genotypes of canola. Similar studies on wheat are entirely
lacking and same needs to be explored for wheat as well.
Conclusions
Allelopathy can be a tool for a more bio-rational man-
agement of natural resources in agro-ecosystems or in
natural ecosystems. It has a key role in regulating plant
biodiversity and in sustainable agriculture. In the present
review, we mainly focused on the role of wheat allelopathy
in sustainability of agro-ecosystem. Allelopathy has been
suggested as an innovative approach to tackle multi-facet
issues of contemporary agriculture. We found a volumi-
nous body of knowledge suggesting that secondary
metabolites present in living wheat plants and dead resi-
dues are involved in many biochemical interactions in the
biosphere. Wheat allelopathy can be implicated for the
biological control the insects, pests, weeds, and diseases.
Allelochemicals in wheat may impact the availability of
nutrients in the soil through their effects on the microbial
diversity and functioning. It offers crop diversification by
rotation while reducing the pest infestation. Many different
types of useful products, such as low-cost biological pes-
ticides and drugs, can arise from allelopathy studies.
Wheat allelopathy can exert profound influence on the
establishment of other plants in the vicinity and alter
community structure and functional diversity of soil
microbiota and enzymes-the two important domains of soil
17
ecology. Thus, it seems imperative to study these aspects
under the influence of wheat allelopathy to unravel wheat
allelopathic phenomenon in the context of soil ecology for
a better understanding of subject matter and its utilization
for weed management in agro-ecosystems.
Although a lot of studies have been carried out on wheat
allelopathy in the past, still a lot of work needs to be done.
Future studies should be carried out: (a) to establish the
effective, economic and feasible techniques of using wheat
allelochemicals under natural field settings, (b) to elucidate
the biochemical pathways responsible for allelochemicals
synthesis, release, and transformation, (c) the uptake
mechanisms and mode of action of allelochemicals in the
receptor organisms, (d) to explore the advanced biotech-
nological tools to enhance allelopathic potential of wheat
for biological weed control, (e) to screen the diverse wheat
genotypes with greater weed/pest suppressive ability and to
incorporate them into the cropping systems, and (f) to
identify the enzymes responsible for the biosynthesis of
wheat allelochemicals (having herbicidal and allelopathic
properties) and genes encoding them.
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