population’s heritable traits: selecting for beneficial alleles and, thus, increasing their
frequency in the population, while selecting against deleterious alleles and, thereby,
decreasing their frequency [172]. This process is known as adaptive evolution and can
result in ecological specialization for particular niches [66] and may eventually result
in speciation events [181]. Natural selection can favor a particular phenotype over the
others, causing the allele frequency to shift in the direction of such phenotype
(directional selection). Selection can also act on intermediate phenotypic variants
maintaining multiple alleles in the gene pool of a population at larger frequencies
(stabilizing selection) or can increase the variance of traits favoring the extreme values
over intermediates (disruptive selection) [50, 110, 142]. Disruptive selection, also
called diversifying selection, divides a population into two distinct groups. This
process may lead to divergent evolution, which is a likely outcome for coevolving
species and for species evolving in sympatry [42, 130]. Evolutionary dynamics
between species are often driven by coadaptation process (Box 2). Consequences of
the reciprocal fitting between partners of biological associations emerged at different
levels of organization (e.g., traits or genes), and as any adaptive dynamic, their
evolutionary patterns are dissected at different time, space, and organizational scales
[22, 87]. Based on biological complexity, the most contrasting standpoints for the
study of coevolution are the phylogenetic and the genetic/genomic perspectives. Tree
thinking and gene thinking suppose different patterns that involve specific mechanisms
(Table 1). Both perspectives have influenced the study of adaptive evolution in
different ways. Whereas tree thinking has helped to understand events that emerged
at supraspecific level (e.g., speciation, diversification, and adaptive radiation), in the
last two decades, the genomic approach has helped to disentangle the molecular
mechanisms behind evolutionary change. In particular, the study of plant chemistry
has been a key to discover the mechanisms of evolutionary innovation. Several studies
on enzymatic complexes of secondary metabolisms have consistently documented the
central role for gene and genome duplication as a common mechanism to achieve novel
function of traits, during speciation or adaptive radiation events (e.g., [17, 46, 47,
140]). After a long tradition of independent field-growing of phylogenetics and
ecology, the development of genomics for the study of biotic interactions has begun
to connect intra- and interspecific approaches, unraveling the genetic mechanism
linked to macroevolutionary patterns (e.g., [89, 183]). The study of plant-herbivore
coevolution provides a good example of how macro- and microevolution begin to
approach but at the same time, of how these two angles have been very prolific when
develop independently.
Box 2 Coadaptation
Coadaptation appears as a result of interaction with others, which produces a
reciprocal adaptation.
Coadaptation generates and allows coevolution [165], which favors the
survival of the systems or individuals. The two parts obtained advantages.
(continued)
52
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