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 biolog-
ical 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
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 expres-
sion within species, (2) community level variation in
allelopathy across species, and (3) variation in the
impacts of allelopathy on associated species. Address-
ing 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

C.-H. Kong
Department of Ecology, College of Resources and
Environmental Sciences, China Agricultural University,
Beijing 100193, China
Introduction
L. M. Ladwig
Department of Biology, University of New Mexico, Allelopathy is broadly defined as any chemical-
Albuquerque, NM 87131, USA
mediated interaction among plants, though it is
N. L. Pisula typically thought of as a mechanism of inhibition
Gewalt Hamilton Associates, Inc., 850 Forest Edge Drive, (Rice 1974). Within ecological research, allelopathy
Vernon Hills, IL 60061, USA has rapidly become a favored mechanism of non-
native species impacts and success (Lawrence et al.
K. A. Lang
Department of Biology, Bradley University, Peoria, 1991; Heisey 1996; McCarthy and Hanson 1998;
IL 61625, USA Roberts and Anderson 2005; Abhilasha et al. 2008;

123

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 evolu-
tionary 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 allelo-
chemicals 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 allelo-
pathic interaction is therefore mediated by soil chem-
ical 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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Plant Ecol
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 selec-
tion 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 advan-
tageous. 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
Genetic Genetic
Environmental Environmental
Inducible Inducible
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
Lankau (2008) found genetic variation in the produc-
tion 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 quantifica-
tion 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, geno-
types 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.
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
Species 3 Species n
Genetic Genetic Variation
Environmental Environmental within species
Inducible Inducible
diverse range of interactions and contingencies into its
conceptual framework. This approach will allow the develop-
ment of a more complete theoretical basis for allelopathy and its
role in 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 respon-
siveness 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 com-
petitive mechanism, it is unknown whether it functions
to alleviate competition specific to water, light or
mineral nutrients or competition in general. Compe-
tition 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.
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,
123

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 com-
petitors (Kong et al. 2002). For example, allocation
of carbon and mineral nutrients towards growth and
reproduction instead of allelochemicals when com-
petition 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 link-
age between competitive environment and allelopa-
thy 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 physiolog-
ical 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. Toler-
ance characteristics may be similar to those proposed
in the herbivory literature. Tolerant plants may
respond to competitive pressures by rapid root prolif-
eration (a form of scramble competition), compensa-
tory 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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Plant Ecol
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. Var-
iation in defense is set by the constraints imposed by
differences in allocation tradeoffs, life history and
evolutionary context across species. Allelopathy stud-
ies 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 inter-
actions 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 non-
allelopathic. 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 strate-
gies 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 strate-
gies (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 inter-
actions, 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, produc-
tion 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 alle-
lopathy. 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 intro-
duced 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 allelo-
pathic, 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 suscep-
tible 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 commu-
nities 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 spe-
cies 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 herbi-
vores. 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 consis-
tently present in a community may generate stronger
selective pressures for allelopathy and select for
chemicals that have a strong specificity. For example,
123

herbaceous dicots in grasslands would more consis-
tently compete with grasses than with other herba-
ceous 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 effective-
ness 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 consis-
tent 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 produc-
tion 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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Plant Ecol
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.
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