An introduction to Forest Genetics

Gsta Eriksson Inger Ekberg David Clapham 2006

http://www.slu.se/Forest-Genetics-online

Evolution
In this chapter we mainly focus on the principles of evolution and we will present a
limited number of empirical data. Population differentiation observed for many traits
are shown in the next chapter. First we give a short presentation of natural selection,
random genetic drift, mutations, and gene flow. The phenotypic plasticity of a trait and
its role in evolution is also discussed. Later on we present in more detail the
evolutionary factors. We raise the question whether or not any perfect form could
be reached in nature.

If we look upon adaptation from an analytical perspective
we can distinguish two steps. During the fi rst step genetic
variation is created and recombination of alleles takes
place. This is mainly a random process. Natural selection
constitutes the second step, during which the allele frequencies
of populations are changed.
It is stressed that natural selection is one of several factors
that infl uence genetic variation within and among populations.
Natural selection is a change of gene frequencies
and it reduces the genetic variation within populations.
There are three types of natural selection. Stabilizing
selection means that phenotypes close to the population
mean are favoured. In directional selection individuals
in one tail of the distribution are favoured. Finally, disruptive
selection favours individuals in both tails of the
distribution. Stabilizing selection is common within stationary
populations. A stabilizing selection within a series
of populations growing along an environmental gradient
will be experienced as disruptive selection among populations.
Natural selection improves the adaptedness but
other evolutionary factors participate in the evolution.
Therefore, perfect adaptedness will never be observed in
nature.
Genetic drift is a random process that leads to allele fi xation
independent of the fi tness contribution of the fi xed
allele; this reduces the within-population genetic variation.
By chance different alleles will be fi xed in different
populations, contributing to among-population variation.
The effect of genetic drift increases exponentially with
decreasing effective population size.
Mutations occur at a low frequency and increase the genetic
variation within populations. Since the mutation rate
per locus and generation is so low, the probability for the
same mutation to arise in two populations is infi nitesimal.
Therefore, mutations will give rise to a small difference among populations.
Gene fl ow is a strong constraint to among-population differentiation.
At the population level it is a strong contributor
to increased within-population variation. Data on
gene fl ow and outcrossing suggest that a large gene fl ow
is not restricted to wind-pollinated species with a wide
and continuous distribution but also occur in scattered
and insect pollinated tree species.
The role of phenotypic plasticity is ambiguous. On the
one hand it can confer fi tness to its carrier and thus is
favoured by natural selection. On the other hand it may
be regarded as a disguise of the genotype. In this way
natural selection becomes less effi cient in the presence of
pronounced phenotypic plasticity.
The relationship between evolution, natural selection,
and genetic drift is illustrated in Figure 6-23. All three
require that there is genetic variation available. Differences
in fi tness are not a prerequisite for evolution but it is
facilitated if it exists. Difference in fi tness is what characterises
natural selection and separates natural selection
from the other two. Evolution means that genetic change
has taken place.
Dependending on the ecological characteristics of a
species they show ecoclinal or ecotypic differentiation.
Ecotypic differentiation occurs if gene fl ow among populations
is much restricted. Ecoclinal variation occurs in
widespread species with a large gene fl ow among populations.
Ecoclinal variation means a continuous variation
along environmental gradients while ecotypic variation
occurs stepwise.
From an evolutionary point of view differentiation of populations
and speciation are related. The difference is that
reproductive isolation exist at the species level whereas
some gene fl ow might occur among populations within
a species. Small populations with no or restricted gene
fl ow are a good basis for rapid speciation. Speciation
is facilitated by geographical isolation. Doubling of the
chromosomal number in species hybrids has been shown
to have occurred frequently in plant speciation and is an
outstanding example of speciation without geographical
isolation.
Evolution in the past has created various patterns of population
structure in different tree species. Some of the
types of population structure are illustrated schematically
in next chapter, Box 7-1. Examples of species having the
various patterns are also given. These patterns are discussed
in the next chapter while variation within populations
is discussed in Chapter 8.