The connection between development and evolution has become the
focus
of an increasing amount of
research in recent years, and heterochrony has long been a key concept
in
this relation. Heterochrony is
defined as evolutionary change in rates and timing of developmental
processes; the dimension of time is
therefore an essential part in studies of heterochrony. Over the past
two decades, evolutionary biologists have
used several methodological frameworks to analyse heterochrony, which
differ substantially in the way they
characterize evolutionary changes in ontogenies and in the resulting
classification, although they mostly use
the same terms. This review examines how these methods compare ancestral
and descendant ontogenies,
emphasizing their differences and the potential for contradictory
results from analyses using different
frameworks. One of the two principal methods uses a clock as a graphical
display for comparisons of size,
shape and age at a particular ontogenic stage, whereas the other
characterizes a developmental process by
its time of onset, rate, and time of cessation. The literature on
human heterochrony provides particularly
clear examples of how these differences produce apparent contradictions
when applied to the same problem.
Developmental biologists recently have extended the concept of heterochrony
to the earliest stages of
development and have applied it at the cellular and molecular scale.
This extension brought considerations
of developmental mechanisms and genetics into the study of heterochrony,
which previously was based
primarily on phenomenological characterizations of morphological change
in ontogeny. Allometry is the
pattern of covariation among several morphological traits or between
measures of size and shape; unlike
heterochrony, allometry does not deal with time explicitly. Two main
approaches to the study of allometry
are distinguished, which differ in the way they characterize organismal
form. One approach defines shape
as proportions among measurements, based on considerations of geometric
similarity, whereas the other
focuses on the covariation among measurements in ontogeny and evolution.
Both are related conceptually
and through the use of similar algebra. In addition, there are close
connections between heterochrony and
changes in allometric growth trajectories, although there is no
one-to-one correspondence. These
relationships and outline links between different analytical frameworks
are discussed.