Evolutionary invasion analysis, also known as adaptive dynamics, is a set of mathematical modeling

techniques that use differential equations to study the long-term evolution of traits in asexually
reproducing populations. It rests on the following four assumptions about mutation and natural selection in the
population under study:[1]

1. Individuals reproduce clonally.

2. Mutations are infrequent, and natural selection acts quickly. The population can be assumed to be
at equilibrium when a new mutant arises.
3. The number of individuals with the mutant trait is initially negligible in the large, established resident
population.
4. Phenotypic mutations occur in small but not infinitesimal steps.
Evolutionary invasion analysis makes it possible to identify conditions on model parameters for which the
mutant population dies out, replaces the resident population, and/or coexists with the resident population.
Long-term coexistence (on the evolutionary timescale) is known as evolutionary branching. When branching
occurs, the mutant establishes itself as a second resident in the environment.
Central to evolutionary invasion analysis is the mutant's invasion fitness. This is a mathematical expression for
the mutant's long-term exponential growth rate when it is introduced into the resident population in small
numbers. If the invasion fitness is positive, the mutant population can grow in the environment set by the
resident organism. If the invasion fitness is negative, the mutant population swiftly goes extinct. [1]

Contents

 1Introduction and background

 2Fundamental ideas
 3Monomorphic evolution
o 3.1Invasion exponent and selection gradient
o 3.2Pairwise-invasibility plots
o 3.3Evolutionarily singular strategies
 4Polymorphic evolution
o 4.1Invasion exponent and selection gradients in polymorphic populations
o 4.2Evolutionary branching
o 4.3Trait evolution plots
 5Other uses
 6References
 7External links

Introduction and background[edit]

The basic principle of evolution via natural selection was outlined by Charles Darwin in his 1859 book, On the
Origin of Species. Though controversial at the time, the central ideas remain largely unchanged to this date,
even though much more is now known about the biological basis of inheritance. Darwin expressed his
arguments verbally, but many attempts have since then been made to formalise the theory of evolution. The
best known are population genetics which models inheritance at the expense of ecological detail, quantitative
genetics which incorporates quantitative traits influenced by genes at many loci, and evolutionary game
theory which ignores genetic detail but incorporates a high degree of ecological realism, in particular that the
success of any given strategy depends on the frequency at which strategies are played in the population, a
concept known as frequency dependence.
Adaptive dynamics is a set of techniques developed during the 1990s for understanding the long-term
consequences of small mutations in the traits expressing the phenotype. They link population
dynamics to evolutionary dynamics and incorporate and generalise the fundamental idea of frequency-
dependent selection from game theory.
Fundamental ideas[edit]
Two fundamental ideas of adaptive dynamics are that the resident population is in a dynamical equilibrium
when new mutants appear, and that the eventual fate of such mutants can be inferred from their initial growth
rate when rare in the environment consisting of the resident. This rate is known as the invasion exponent when
measured as the initial exponential growth rate of mutants, and as the basic reproductive number when it
measures the expected total number of offspring that a mutant individual produces in a lifetime. It is sometimes
called the invasion fitness of mutants.
To make use of these ideas, a mathematical model must explicitly incorporate the traits undergoing
evolutionary change. The model should describe both the environment and the population dynamics given the
environment, even if the variable part of the environment consists only of the demography of the current
population. The invasion exponent can then be determined. This can be difficult, but once determined, the
adaptive dynamics techniques can be applied independent of the model structure.

Monomorphic evolution[edit]
A population consisting of individuals with the same trait is called monomorphic. If not explicitly stated
otherwise, the trait is assumed to be a real number, and r and m are the trait value of the monomorphic
resident population and that of an invading mutant, respectively.

Invasion exponent and selection gradient[edit]

The invasion exponent is defined as the expected growth rate of an initially rare mutant in the
environment set by the resident (r), which means the frequency of each phenotype (trait value) whenever this
suffices to infer all other aspects of the equilibrium environment, such as the demographic composition and the
availability of resources. For each r, the invasion exponent can be thought of as the fitness landscape
experienced by an initially rare mutant. The landscape changes with each successful invasion, as is the case in
evolutionary game theory, but in contrast with the classical view of evolution as an optimisation process
towards ever higher fitness.

We will always assume that the resident is at its demographic attractor, and as a consequence for all r,
as otherwise the population would grow indefinitely.

The selection gradient is defined as the slope of the invasion exponent at , . If the sign of the
selection gradient is positive (negative) mutants with slightly higher (lower) trait values may successfully
invade. This follows from the linear approximation

which holds whenever .

Pairwise-invasibility plots[edit]
The invasion exponent represents the fitness landscape as experienced by a rare mutant. In a large

(infinite) population only mutants with trait values for which is positive are able to successfully
invade. The generic outcome of an invasion is that the mutant replaces the resident, and the fitness
landscape as experienced by a rare mutant changes. To determine the outcome of the resulting series of

invasions pairwise-invasibility plots (PIPs) are often used. These show for each resident trait value

all mutant trait values for which is positive. Note that is zero at the diagonal . In
PIPs the fitness landscapes as experienced by a rare mutant correspond to the vertical lines where the

resident trait value is constant.

Evolutionarily singular strategies[edit]

The selection gradient determines the direction of evolutionary change. If it is positive (negative) a
mutant with a slightly higher (lower) trait-value will generically invade and replace the resident. But what

will happen if vanishes? Seemingly evolution should come to a halt at such a point. While this is a

possible outcome, the general situation is more complex. Traits or strategies for which , are
known as evolutionarily singular strategies. Near such points the fitness landscape as experienced by a
rare mutant is locally `flat'. There are three qualitatively different ways in which this can occur. First, a
degenerate case similar to the saddle point of a qubic function where finite evolutionary steps would lead
past the local 'flatness'. Second, a fitness maximum which is known as an evolutionarily stable
strategy (ESS) and which, once established, cannot be invaded by nearby mutants. Third, a fitness
minimum where disruptive selection will occur and the population branch into two morphs. This process is
known as evolutionary branching. In a pairwise invasibility plot the singular strategies are found where the
boundary of the region of positive invasion fitness intersects the diagonal.
Singular strategies can be located and classified once the selection gradient is known. To locate singular

strategies, it is sufficient to find the points for which the selection gradient vanishes, i.e. to find such

that . These can be classified then using the second derivative test from basic calculus. If the second

derivative evaluated at is negative (positive) the strategy represents a local fitness maximum

(minimum). Hence, for an evolutionarily stable strategy we have

If this does not hold the strategy is evolutionarily unstable and, provided that it is also convergence

stable, evolutionary branching will eventually occur. For a singular strategy to be convergence
stable monomorphic populations with slightly lower or slightly higher trait values must be invadable by

mutants with trait values closer to . That this can happen the selection gradient in a

neighbourhood of must be positive for and negative for . This means that the slope

of as a function of at is negative, or equivalently

The criterion for convergence stability given above can also be expressed using second
derivatives of the invasion exponent, and the classification can be refined to span more than the
simple cases considered here.