
Figure 1 and Figure I in Box 2). For example, developmen-
tal transformations have been shown to be controlled by
environmental signaling pathways that sense abiotic cues
such as light and nitrogen [11] and drought [12], as well as
biotic signals such as Nod factors that cause nodulation in
legumes under low nitrogen conditions [13]. For many
other environmentally induced phenotypic responses,
the mechanisms of how environmental signals are sensed
and processed are still largely unknown [e.g. 14,15].An
improved understanding of the molecular basis of environ-
mentally induced changes in plant traits will yield insight
into possible ecological and evolutionary responses in wild
species and will be useful for engineering plasticity in crop
species (Box 1, Q1).
Flowering time is a good example of a crucial trait that
has been shown to be both under genetic control and plastic
(see below). Under climate change, the temperature cues
triggering the chain of events leading to flowering might
cease to be reliable if they occur at the wrong time with
respect to the lifecycle and ecology of the species. Such
changes in cue, signal or response schemes might thereby
elicit maladaptive responses [7]. Alternatively, they can
lead to the expression of phenotypic responses that are
currently hidden [16]. Current techniques in molecular
biology and genetics allow for studies of plastic trait
responses that scale from a description of molecular mecha-
nism to the assessment of adaptive value under current or
simulated future climates [17]. Thus far the genetic basis of
plasticity has been examined in greatest depth in model and
crop species. As new tools become available, the extension of
these studies to more non-model species becomes increas-
ingly possible and will help us determine the extent to which
there are genetic homologs in other species (Box 2).
Plasticity in key plant functional traits in response to
climate change
Plasticity is a characteristic of a given trait in response to a
given environmental stimulus, rather than a characteristic
of an organism as a whole. Likewise, some responses are
examples of adaptive plasticity, providing a fitness benefit,
whereas others are inevitable responses to physical pro-
cesses or resource limitations [18,19] (Figure 2). Both
adaptive and non-adaptive plasticity will play a role in
the context of plant responses to climate change. Differen-
tiating between the two is important to our understanding
of both the current value and the evolution of plasticity
(Box 1, Q2). The consensus from the theoretical literature
is that adaptive phenotypic plasticity should evolve in
heterogeneous environments where signals of environmen-
tal conditions are reliable [19,20]. Hypotheses about what
sort of species will be most plastic also abound in the
literature [21–26], yet our ability to predict patterns of
plasticity in key traits in response to climate change
remains limited.
Given that it is not feasible to assess plastic responses to
current or future environments on all species, it is impor-
tant to identify which traits are likely to show important
plastic responses to particular changing environmental
conditions and to develop predictors to enable us to gener-
alize about the sorts of species likely to exhibit these plastic
responses [9]. Those traits can then be examined in current
or projected climate conditions to determine the extent of
plasticity and assess the extent to which the underlying
molecular and genetic pathways are shared (Box 2).
Plasticity in plant functional traits
In recent years, ecologists have categorized species accord-
ing to plant functional types and have also identified
several continuous plant functional traits that vary in
predictable ways along environmental gradients. Func-
tional types are widely used in global climate models to
group species according to their function in the ecosystem
or community (e.g. C3 or C4 grasses, herbs, shrubs, decid-
uous trees, N-fixing legumes, etc.). Functional traits are
those that help describe the ecology of species using a few,
easily quantified variables (e.g. seed size, plant height, leaf
lifespan, leaf mass per area, etc.) [27]. Functional traits are
relevant to both global climate models and mechanistic
models of plant distributions (see below). Considering their
probable importance, we advocate that plant functional
traits should have priority for the investigation of (adap-
tive) phenotypic plasticity and identification of molecular
and genetic mechanisms across species (Box 3).
Adaptive plasticity in functional traits is likely to assist
rapid adaptation to new conditions. Thus, a natural ques-
Box 1. Outstanding questions
Modern techniques and the potential for cross-disciplinary ap-
proaches mean that we are now in a position to address the
following questions effectively.
Q1: Molecular basis of plasticity:
What is the genetic control of plasticity and how is it linked to
epigenetics?
Can we identify ‘plasticity genes’?
Does identifying such plasticity genes improve our ability to
predict the longer term responses of traits and species to climate
change?
Q2: Adaptive plasticity:
What traits are likely to show adaptive plasticity?
Will species with differing ecologies (i.e. differing functional
types) exhibit adaptive plasticity in different traits?
Will the incidence of adaptive plasticity vary among types of traits
(e.g. those related to anatomy versus allocation versus physiol-
ogy)?
Q3: Functional traits:
Are the traits most commonly identified as plant functional traits
also those that show adaptive plasticity?
Is plasticity in functional traits important in determining response
to climate change under future climates, regardless of current
adaptive value?
Q4: Plasticity and evolution:
How has plasticity contributed to the diversification of lineages
and can the evidence of this contribution be found by comparing
the distribution of adaptive plasticity or relevant plasticity genes
with population or species phylogenies?
How will plasticity contribute to rapid evolution in response to
climate change?
How much variation is there for plasticity and how does it respond
to selection?
Q5: Plasticity in crop species:
Has breeding led to reductions in adaptive plasticity in contem-
porary crop varieties relative to older ones or wild ancestors?
Can we breed for plasticity in key traits in agricultural systems to
improve yield stability under climate change?
Review
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