Plant Ecol

DOI 10.1007/s11258-012-0078-5

Developing an ecological context for allelopathy

Scott J. Meiners Chui-Hua Kong
Laura M. Ladwig Nikki L. Pisula
Kimberly A. Lang

Received: 28 March 2012 / Accepted: 7 June 2012

Springer Science+Business Media B.V. 2012

Abstract There has been a renewed interest in

allelopathy as a plantplant interaction as more plant
ecologists have become involved in studying biological invasions. This resurgence highlights a major
deficiency in our understanding of allelopathythe
lack of a well-developed ecological context for the
interaction. In contrast to allelopathy, the plantplant
interaction of competition has a strong theoretical
foundation as well as a large body of supporting
empirical studies. We suggest that the plant-herbivore
defense literature provides a mature and well-devel-

S. J. Meiners (&)
Department of Biological Sciences, Eastern Illinois
University, Charleston, IL 61920, USA
e-mail: sjmeiners@eiu.edu
C.-H. Kong
Department of Ecology, College of Resources and
Environmental Sciences, China Agricultural University,
Beijing 100193, China
L. M. Ladwig
Department of Biology, University of New Mexico,
Albuquerque, NM 87131, USA
N. L. Pisula
Gewalt Hamilton Associates, Inc., 850 Forest Edge Drive,
Vernon Hills, IL 60061, USA
K. A. Lang
Department of Biology, Bradley University, Peoria,
IL 61625, USA

oped framework from which a broader ecological

context for allelopathy can be developed. Here, we
discuss three broad classes of questions, drawn from the
herbivore defense literature, which may help to develop
an appropriate ecological context for allelopathy. These
questions focus on (1) variation in allelopathic expression within species, (2) community level variation in
allelopathy across species, and (3) variation in the
impacts of allelopathy on associated species. Addressing such broad population and community level themes
in a variety of systems will be necessary to fully develop
an ecological context for allelopathy and provide a
theoretical basis for understanding its role in plant
communities.
Keywords Allelopathy
Allocation
Competition

Cost of defense
Impacts
Variation

Introduction
Allelopathy is broadly defined as any chemicalmediated interaction among plants, though it is
typically thought of as a mechanism of inhibition
(Rice 1974). Within ecological research, allelopathy
has rapidly become a favored mechanism of nonnative species impacts and success (Lawrence et al.
1991; Heisey 1996; McCarthy and Hanson 1998;
Roberts and Anderson 2005; Abhilasha et al. 2008;

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Gomez-Aparicio and Canham 2008; Inderjit et al.

2008; Lankau et al. 2009). While there has been some
conceptual development in the context of invasion
(e.g., Callaway and Aschehoug 2000; Thorpe et al.
2009), allelopathy as an ecological phenomenon has
evolved relatively slowly since its inception therefore
its importance to communities and populations
remains unclear (Wardle et al. 1998).
In contrast to allelopathy, there is a very strong
conceptual basis for the understanding of herbivore
defenses (e.g. Atsaat and ODowd 1976; Coley et al.
1985; Rosenthal and Kotanen 1994; de Jong 1995).
This rich body of theory allows us to predict the types of
species and tissues that may be defended, the flexibility
of defense (i.e., inducible or constituent), the evolutionary constraints of allocation to defenses and shared
evolutionary history with herbivores. As allelopathy
may be thought of as a chemical defense against
competitive suppression, we may expect to see similar
ecological patterns to the interactions between plants
and their herbivores. Herbivore defenses and allelochemicals are similar in that they require allocation of
energy and nutrients and should incur costs to the
organism that must be balanced with the benefits of
protection. However, defenses against herbivores differ
from allelopathy in the directness of the interactions
involved. While herbivore activity is a direct impact on
the affected plant, allelopathic interactions rely on
chemicals released into the environment. The allelopathic interaction is therefore mediated by soil chemical properties and soil microbial communities, and
may directly influence plant growth or indirectly
through altered microbial associations. Despite the
more direct nature of plantherbivore interactions, they
represent a reasonable starting point to guide our
understanding of allelopathy.
Drawing heavily from the conceptual foundation of
plant antiherbivore defenses, we briefly outline three
broad research questions that should be addressed to
build an appropriate ecological context for allelopathy
(Fig. 1). Our goal is to simply point out parallels
between the two areas of research and illustrate
productive avenues for allelopathic research. These
questions are not the only challenges to the study of
allelopathy as there are many methodological issues
that must still be resolved. However, we believe that
by addressing these broad questions plant ecologists
will be more able to understand the importance of
allelopathy in natural systems.

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What are the sources of variation in allelopathy

within species?
The vast majority of allelopathic research focuses on
whether a species is allelopathic and what the
causative agent may be. The conclusion of many of
these allelopathic studies is that the test species is
determined to be either allelopathic, or not. While
such studies are beneficial, they neglect to address the
potential for intraspecific variation in allelopathy.
From the herbivore defense literature, we know that
within a species there may be considerable variation in
defensive chemistry based on genetic variation in
chemical production, the environmental conditions
that an individual is growing in, the exposure of the
individual to herbivores, or the developmental stage of
the individual (Coley et al. 1985; Siemens et al. 2002;
Agrawal 2004). Allelopathic studies need to include
sources of intraspecific variation to fully understand
the ecological context of allelopathy at the population
level. Central to the population ecology of allelopathy
is the idea of energetic costs. Plants that allocate
resources to allelochemicals are expected to incur
costs to growth and reproduction that natural selection should balance with the benefits of reduced
competition.
Genetic variation
As ecologists, we know remarkably little about the
variation in allelopathy that would serve as the base
variance upon which selection would function. In
contrast, genetic controls on herbivore defenses are
commonly examined (e.g. Gols et al. 2008). As with
any trait, we would expect genetic variation among
individuals even when allelopathy is strongly advantageous. Genetic variation may allow local adaptation
which would favor increased allocation to allelopathy
in competitive environments and a decrease when the
costs exceed the benefits. Some evidence for genetic
variation in allelopathy comes from the agricultural
literature that evaluates crop varieties based on
allelopathic control of weed species (e.g. Weston
1996). Selection imposed from crop breeding has
generated variation in allelopathy among varieties,
though this was not the original selection goal
(Bertholdsson 2004). We may expect similar variation
in plant populations, where allelopathy may be
beneficial and therefore selected for. For example,

Plant Ecol

Variation among species

Prevalence of allelopathy
Plant strategies
Invasion

Species 1

Species 2

Species 3

Species n

Genetic
Environmental
Inducible

Genetic
Environmental
Inducible

Genetic
Environmental
Inducible

Genetic
Environmental
Inducible

Variation
within species

Variation in impacts
Specificity
Evolutionary responses

Fig. 1 Conceptual diagram that highlights the primary sources

of variation in allelopathy within plant communities. Though
dealt with separately in this article, there is clearly potential for
interactions among these sources of variation. The study of
allelopathy must incorporate a broader view that includes the

diverse range of interactions and contingencies into its

conceptual framework. This approach will allow the development of a more complete theoretical basis for allelopathy and its
role in plant communities

Lankau (2008) found genetic variation in the production of allelopathic chemicals in Brassica. In this
study, allocation towards chemical defense resulted in
greater interspecific competitive ability, but reduced
intraspecific competition from allocation costs. These
types of studies suggest that understanding genetic
variation and selective pressures may yield important
insights into allelopathy as an ecological process
(Lankau et al. 2009). Identifying genotypes of varying
allocation to allelopathy will allow for the quantification of the cost of allelopathy by comparing growth
rates in the presence and absence of competition and
will provide opportunities to understand the underlying
tradeoffs that constrain allelopathy. Furthermore, genotypes with varying degrees of allelopathy can provide
experimental opportunities to test for the direct benefits
of allelopathy as well as its impact on experimental
plant communities.

chemicals, may be primarily carbon molecules or may

contain nitrogen (Duke and Dayan 2006) and may be
favored by different growing conditions. The responsiveness of plants to the environment would result in
phenotypic plasticity across landscapes that vary in
resource availability and competitive interactions.
Understanding the basis for environmentally-driven
variation will allow the prediction of conditions under
which we would expect allocation to allelochemicals
and potentially strong effects of allelopathy. An
environment-allelopathy linkage would also suggest
a specific triggering mechanism for allelopathy.
Though allelopathy is commonly considered a competitive mechanism, it is unknown whether it functions
to alleviate competition specific to water, light or
mineral nutrients or competition in general. Competition has been found to both increase and decrease the
abundance of allelochemicals (Kong et al. 2002;
Rivoal et al. 2011), leaving its role as a constraint on or
trigger for allelopathy unclear.

Environmental variation
The ability of plants to produce defensive chemicals is
often constrained by the environment in which the
plant is growing (Coley et al. 1985). Environments
that are limiting in resources needed to construct
particular allelochemicals or are physiologically
stressful may alter the production of allelochemicals
(Kong et al. 2002; Kong et al. 2004; Rivoal et al.
2011). Allelochemicals, just as herbivore defense

Inducible versus constituent allelopathy

Many species maintain a base level of antiherbivore
chemicals, increasing their production only if the plant
is damaged by an herbivore (e.g. Harvell and Tollrian
1999; Siemens et al. 2002). Flexibility in defense
effectively minimizes the cost of producing and
maintaining defenses in the absence of herbivores,

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though an energetic cost is incurred from the initial

damage. As there are likely costs to allelopathy
(Lankau 2008), it may be selectively advantageous
for plants to similarly vary allelochemical production
based on the presence/absence of interspecific competitors (Kong et al. 2002). For example, allocation
of carbon and mineral nutrients towards growth and
reproduction instead of allelochemicals when competition is minimal would maximize fitness in that
environment. Interestingly, allelochemicals may
function as both an inhibitor of competing plant
species and defense against herbivores (Wardle et al.
1998; Cipollini 2004; Thelen et al. 2005; Cipollini
et al. 2008), leading to more a complex suite of
potential regulators. Establishing a causal linkage between competitive environment and allelopathy would strengthen its role as a competitive
mechanism.

Tolerance versus resistance to competition

Another major concept used to explain variation in
defense in the plant-herbivore literature is the trade-off
between resistance (defensive) and tolerance traits.
Plant species may allocate resources to defenses
against herbivore damage or may develop physiological responses such as compensatory growth, resource
use efficiency or utilization of stored carbohydrates to
minimize the impact of herbivore damage on plant
fitness (Strauss and Agrawal 1999; Stamp 2003).
While the presence of a trade-off is not always found
between defensive and tolerance traits (Mauricio et al.
1997; Leimu and Koricheva 2006), similar processes
may also occur with competition. Allelopathy in this
context would be the resistance end to the strategy
gradient where the release of chemicals would inhibit
the competitive ability of neighboring plants. Tolerance characteristics may be similar to those proposed
in the herbivory literature. Tolerant plants may
respond to competitive pressures by rapid root proliferation (a form of scramble competition), compensatory allocation to root or shoot tissues to increase
resource uptake, increased efficiency of resource
utilization or phenological shifts in resource uptake
with storage. While the potential for such alternate
strategies has not been explored in the context of
allelopathy, it would help to explain variation in
allelopathic strength among populations and species.

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What are the sources of variation in allelopathy

among species?
Plant species are expected to exhibit a range in levels
of antiherbivore defenses with some species very
poorly defended, others heavily defended and many
species intermediate between the two extremes. Variation in defense is set by the constraints imposed by
differences in allocation tradeoffs, life history and
evolutionary context across species. Allelopathy studies typically focus on a single target species that is
a priori thought to be allelopathic. This scenario
generates challenges for understanding variation in
allelopathy among species as the pool of studies is
likely biased towards more allelopathic species. As
case studies often determine whether a species is
allelopathic or not (a binary response), it makes
comparisons among species more difficult. To place
allelopathy in an accurate ecological context, we need
to address the continuous range of allelopathic interactions across communities (Pisula and Meiners
2010).
Prevalence of allelopathy in communities
Allelopathy is typically treated as a somewhat rare
plant characteristic. When experiments are conducted,
one species is designated as allelopathic and the
other(s)the target speciesare assumed to be nonallelopathic. In contrast to allelopathy, the herbivore
defense literature embraces the diversity of defensive
modes and levels within plant communities. As
competition is as ubiquitous, if not more so than
herbivory, it is possible that allelopathy is similarly
widespread in plant communities. To address the
community level aspects of allelopathy, we will need
surveys of large numbers of species (e.g. Moral and
Cates 1971) or meta-analyses of individual studies.
However, surveys, like those of any plant trait, must
include a range of species to avoid biases based on the
identity of the species selected for study. As larger and
more detailed surveys are developed, phylogenetic
patterns will need to be addressed, particularly with
regard to the types of chemicals produced. Ultimately,
knowing the prevalence of allelopathy may help us to
understand its importance in plant communities. This
information may also help us to understand why
documenting the community impacts of allelopathy
are so difficult (Wardle et al. 1998).

Plant Ecol

Ecological correlates of allelopathy

Within the herbivore defense literature is the concept
of whether species should grow or defend (Herms and
Mattson 1992). Selection may favor different strategies based on an individuals potential life span and
other life history characteristics. If we were to place
allelopathy into Grimes C-S-R view of plant strategies (Grime 1977), we would expect allelopathy, as a
competitive mechanism, to be predominately found
with the C-strategists. In this set of tradeoffs, ruderals
would typically not be exposed to competitive interactions, and therefore would be less likely to be
allelopathic. Stress tolerant species may also be
allelopathic to capitalize on limited resources, but this
may be more context-specific. For example, production of allelochemicals by light limited understory
species would be unlikely to increase light availability
to the forest floor and may detrimentally reduce
allocation to growth. However, if species in the
understory were limited by soil resources, production
of allelochemicals may be beneficial. While Grimes
suite of plant strategies may be a good starting place,
allelopathy may not cleanly fit into the scheme as
annuals (ruderals) are often found to be allelopathic
(e.g. Bertholdsson 2004; Wang et al. 2005; Lankau
2008). Broad scale analyses may also be able to find
linkages between environmental variables and allelopathy. The benefit of looking for either life history or
environmental correlates of allelopathy is that it will
help us to predict the type of species and communities
where allelopathy may be important. Building this
context will also suggest a clear ecological function
for allelopathy within plant communities.
Linkage with invasion
Allelopathy is an often-cited mechanism of the
success and impacts of non-native species in introduced habitats (Callaway and Aschehoug 2000;
Abhilasha et al. 2008; Inderjit et al. 2008) and is at
least partially responsible for the resurgence of interest
in allelopathy. Though part of this effect may be the
sensitivity of the native species to the invaders
chemicals (see below), allelopathy as a plant trait
may be important in determining which non-native
species become successful. To effectively determine
whether invasion success is linked with allelopathy we
would need to appropriately compare invaders with

native species (van Kleunen et al. 2010). A strong

argument for allelopathy as a key trait in invasion
would be if dominant non-native species have a
greater incidence of allelopathy than dominant native
species. If non-native and native species have similar
rates of allelopathy, then allelopathy may be a general
mechanism of dominance rather than one specific to
the invasion process. Alternatively, it may be that
non-native invaders are not more likely to be allelopathic, just more likely to be tested than native species.
This scenario would not suggest that allelopathy is
unimportant in any individual invasion, just that it is
not a mechanism of dominance unique to non-native
species. Allelochemical production may still be
important to an invaders success if the environmental
constraints of the new habitat result in increased
production or if the new community is more susceptible to those chemicals (Inderjit et al. 2011).

What are the sources of variation in the impacts

of allelopathy?
Ultimately we are interested in understanding what
allelopathy does in plant communities, just as we are
interested in how herbivores structure plant communities and populations. Allelopathy impacts need to be
examined at two temporal scales, ecological and
evolutionary, to fully understand their importance.
Again, the focus here is to generate a broader view of
allelopathy by looking at larger ecological patterns
rather than individual species as case studies.
Specificity of effect
Studies often identify differences among target species in their response to an allelopathic plant species
(e.g. Abhilasha et al. 2008). Similar to antiherbivore
defenses, we may expect to see selection on the
efficacy of allelochemicals analogous to the selective
pressures imposed by generalist and specialist herbivores. Rare or unpredictable herbivores are unlikely to
impose large selective pressures on plant populations,
whereas herbivores that consistently damage a species
should shape the chemical defenses towards their
activity. Similarly, strong competitors that are consistently present in a community may generate stronger
selective pressures for allelopathy and select for
chemicals that have a strong specificity. For example,

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herbaceous dicots in grasslands would more consistently compete with grasses than with other herbaceous species. An effective allelochemical in this
situation may be much more effective on grasses than
other life forms. Therefore, we can postulate that
plants with a more unpredictable pool of competitors
may evolve allelochemicals with broader effectiveness the equivalent of generalist herbivore defenses.
Identifying the rules that govern species responses to
allelopathy would address some of the difficulty in
showing community level impacts of allelopathy
(Wardle et al. 1998). As competitive interactions are
not equivalent across a community, likewise we
should not expect allelopathy to have uniform effects.
Selection in response to allelopathy
Not only should we expect selection to shape the
specificity of allelopathy, but we should also expect
selection to shape species response to allelopathy. If
the production of allelochemicals generates a consistent selective pressure on resident species, then
genotypes that are less affected by the allelochemicals
should be favored in the community (Callaway et al.
2005). Over time, this trend may lead to suites of
allelopathic species and their primary competitors that
are adapted to the chemicals produced. Populations
without exposure to allelochemicals may therefore
remain more susceptible to their effects (Callaway
et al. 2005). In the context of invasion, this is called the
novel weapons hypothesis, where species nave to an
allelochemical may be more affected than those which
have coevolved with the allelopathic competitor
(Thorpe et al. 2009; Kim and Lee 2011). If the
selective pressure maintaining allelochemical production is alleviated, this change may lead to shifts in
allocation from allelochemicals towards growth and
reproduction as has been seen in herbivore defenses
(Blossey and Notzold 1995; Inderjit et al. 2011).

Conclusions
The general themes proposed here are not the only
challenges to understanding the complexity and
importance of the ecology of allelopathy, nor is this
an exhaustive list of potential avenues for research.
There are still methodological issues associated with
examining large suites of species as well as in

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separating out competitive effects from allelopathic

ones (e.g., Lau et al. 2008). Other important classes
of questions not dealt with here involve the role of
allelopathy in ecosystems and the interaction of
allelochemicals with soil processes (Wardle et al.
1998; Inderjit et al. 2011). From a developmental
perspective, the study of allelopathy is still in its
ecological infancy and needs to mature conceptually
before its importance to populations and communities
can be fully assessed. While the plant defensive
literature provides a reasonable starting point to
develop a broader context to allelopathy, we must
also remember the long history and controversies that
characterized its development (Stamp 2003). It is
important that we not only learn from the outcomes of
those debates, but also from the process so that we may
avoid similar pitfalls. By critically examining the
conceptual foundation of plant defenses, we may
wisely select ideas that may be appropriate to the field
of allelopathy and move forward.
The time is ripe for ecologists to develop a broader
conceptual view of allelopathy. Many studies focus on
determining whether a species is allelopathicthe
equivalent of asking whether a species is defended from
herbivores. While this approach is important to those
interested in the autecology of that species, it does not
directly move the broader field forward. The case study
approach is an important first step to determine whether
allelopathy can be important. As the answer to this query
is clearly yes, it is now time to move forward, build upon
the rich foundation of case studies, and develop the
ecological context of allelopathy.

References
Abhilasha D, Quintana N, Vivanco J, Joshi J (2008) Do allelopathic compounds in invasive Solidago canadensis s.l.
restrain the native European flora? J Ecol 96:9931001
Agrawal AA (2004) Resistance and susceptibility of milkweed:
competition, root herbivory, and plant genetic variation.
Ecology 85:21182133
Atsaat PR, ODowd DJ (1976) Plant defense guilds. Science
193:2429
Bertholdsson NO (2004) Variation in allelopathic activity over
100 years of barley selection and breeding. Weed Res
44:7886
Blossey B, Notzold R (1995) Evolution of increased competitive
ability in invasive nonindigenous plantsa hypothesis.
J Ecol 83:887889

Plant Ecol
Callaway RM, Aschehoug ET (2000) Invasive plants versus
their new and old neighbors: a mechanism for exotic
invasion. Science 290:521523
Callaway RM, Ridenour WM, Laboski T, Weir T, Vivanco JM
(2005) Natural selection for resistance to the allelopathic
effects of invasive plants. J Ecol 93:576583
Cipollini D (2004) Stretching the limits of plasticity: Can a plant
defend against both competitors and herbivores? Ecology
85:2837
Cipollini D, Stevenson R, Enright S, Eyles A, Bonello P
(2008) Phenolic metabolites in leaves of the invasive shrub,
Lonicera maackii, and their potential phytotoxic and antiherbivore effects. J Chem Ecol 34:144152
Coley PD, Bryant JP, Chapin FS (1985) Resource availability
and plant antiherbivore defense. Science 230:895899
de Jong TJ (1995) Why fast-growing plants do not bother about
defense. Oikos 74:545548
Duke SO, Dayan FE (2006) Modes of action of phytotoxins
from plants. In: Pedrol N, Gonzalez L, Reigosa MJ (eds)
Allelopathy: a physiological process with ecological
implications. Springer, Dordrecht, pp 511536
Gols R, Wagenaar R, Bukovinszky T, van Dam NM, Dicke M,
Bullock JM, Harvey JA (2008) Genetic variation in defense
chemistry in wild cabbages affects herbivores and their
endoparasitoids. Ecology 89:16161626
Gomez-Aparicio L, Canham CD (2008) Neighbourhood analyses of the allelopathic effects of the invasive tree Ailanthus altissima in temperate forests. J Ecol 96:447458
Grime JP (1977) Evidence for the existence of three primary
strategies in plans and its relevance to ecological and
evolutionary theory. Am Nat 111:11691194
Harvell CD, Tollrian R (1999) Why inducible defenses? In:
Tollrian R, Harvell CD (eds) The ecology and evolution of
inducible defenses. Princeton University Press, Princeton
Heisey RM (1996) Identification of an allelopathic compound
from Ailanthus altissima (Simaroubaceae) and characterization of its herbicidal activity. Am J Bot 83:192200
Herms DA, Mattson WJ (1992) The dilemma of plants: to grow
or defend. Q Rev Biol 67:283335
Inderjit I, Seastedt T, Callaway R, Pollock J, Kaur J (2008)
Allelopathy and plant invasions: traditional, congeneric, and
bio-geographical approaches. Biol Invasions 10:875890
Inderjit, Inderjit DA, Karban R, Callaway RM (2011) The
ecosystem and evolutionary contexts of allelopathy.
Trends Ecol Evol 26:655662
Kim Y, Lee E (2011) Comparison of phenolic compounds and
the effects of invasive and native species in East Asia:
support for the novel weapons hypothesis. Ecol Res
26:8794
Kong CH, Hu F, Xu XH (2002) Allelopathic potential and
chemical constituents of volatiles from Ageratum conyzoides under stress. J Chem Ecol 28:11731182
Kong CH, Hu F, Liang WJ, Wang PW, Jiang Y (2004) Allelopathic potential of Ageratum conyzoides at various growth
stages in different habitats. Allelopath J 13:233240
Lankau R (2008) A chemical trait creates a genetic trade-off
between intra- and interspecific competitive ability. Ecology 89:11811187
Lankau RA, Nuzzo V, Spyreas G, Davis AS (2009) Evolutionary limits ameliorate the negative impact of an invasive
plant. Proc Natl Acad Sci USA 106:1536215367

Lau JA, Puliafico KP, Kopshever JA, Steltzer H, Jarvis EP,

Schwarzlander M, Strauss SY, Hufbauer RA (2008)
Inference of allelopathy is complicated by effects of activated carbon on plant growth. New Phytol 178:412423
Lawrence JG, Colwell A, Sexton OJ (1991) The ecological
impact of allelopathy in Ailanthus altissima (Simaroubaceae). Am J Bot 78:948958
Leimu R, Koricheva J (2006) A meta-analysis of tradeoffs
between plant tolerance and resistance to herbivores:
combining the evidence from ecological and agricultural
studies. Oikos 112:19
Mauricio R, Rausher MD, Burdick DS (1997) Variation in the
defense strategies of plants: are resistance and tolerance
mutually exclusive? Ecology 78:13011311
McCarthy BC, Hanson SL (1998) An assessment of the allelopathic potential of the invasive weed Alliaria petiolata
(Brassicaceae). Castanea 63:6873
Moral RD, Cates RG (1971) Allelopathic potential of the
dominant vegetation of western Washington. Ecology
52:10301037
Pisula NL, Meiners SJ (2010) Relative allelopathic potential of
invasive plant species in a young disturbed woodland.
J Torrey Bot Soc 137:8187
Rice EL (1974) Allelopathy. Academic Press, New York
Rivoal A, Fernandez C, Greff S, Montes N, Vila B (2011) Does
competition stress decrease allelopathic potential? Biochem Syst Ecol 39:401407
Roberts KJ, Anderson RC (2005) Effect of garlic mustard
[Alliaria petiolata (Beib, Cavarra & Grande)] extracts on
plants and arbuscular mycorrhizal (AM) fungi. Am Midl
Nat 146:146152
Rosenthal JP, Kotanen PM (1994) Terrestrial plant tolerance to
herbivory. Trends Ecol Evol 9:145148
Siemens DH, Garner SH, Mitchell-Olds T, Callaway RM (2002)
Cost of defense in the context of plant competition: Brassica rapa may grow and defend. Ecology 83:505517
Stamp N (2003) Out of the quagmire of plant defense hypotheses. Q Rev Biol 78:2355
Strauss SY, Agrawal AA (1999) The ecology and evolution of
plant tolerance to herbivory. Trends Ecol Evol 14:179185
Thelen GC, Vivanco JM, Newingham B, Good W, Bais HP,
Landres P, Caesar A, Callaway RM (2005) Insect herbivory stimulates allelopathic exudation by an invasive plant
and the suppression of natives. Ecol Lett 8:209217
Thorpe AS, Thelen GC, Diaconu A, Callaway RM (2009) Root
exudate is allelopathic in invaded community but not in
native community: field evidence for the novel weapons
hypothesis. J Ecol 97:641645
van Kleunen M, Dawson W, Schlaepfer D, Jeschke JM, Fischer
M (2010) Are invaders different? A conceptual framework
of comparative approaches for assessing determinants of
invasiveness. Ecol Lett 13:947958
Wang P, Liang WJ, Kong CH, Jiang Y (2005) Allelopathic
potential of volatile allelochemicals of Ambrosia trifida L.
on other plants. Allelopath J 15:131136
Wardle DA, Nilsson M-C, Gallet C, Zackrisson O (1998) An
ecosystem-level perspective of allelopathy. Biol Rev
73:305319
Weston LA (1996) Utilization of allelopathy for weed
management in agroecosystems. Agron J 88:860866

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