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Epigenetic Mechanisms
Stuart A Newman, New York Medical College, Valhalla, New York, USA
Gerd B Muller, University of Vienna, Vienna, Austria
Keynote article
Article Contents
. Introduction
. Physics of Multicellularity and the Origin of Body Plans
. Interplay of Generic and Programmatic Mechanisms of
Development
. Epigenetics of Advanced Development
. Origin of Morphological Homology
Introduction
Materials of the nonliving world take on forms dictated by
external forces to which they are susceptible by virtue of
their inherent physical properties. Water, for example,
forms waves and vortices if it is mechanically agitated,
whereas clay bears the record of its most recent physical
impressions long after they have been exerted. Living
metazoa multicellular animals seem to obey dierent
rules: their forms appear to be expressions of intrinsic
developmental programmes. Although organisms must, of
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ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
See also: Evolutionary Developmental Biology: Developmental and Genetic Mechanisms of Evolutionary Change;
Evolutionary Developmental Biology: Gene Duplication,
Divergence and Co-option; Genetic Redundancy
Once basic body plans were established, selection for biochemical integration, promoting physiological homeostasis and developmental reliability stabilized the
relationship between genotype and phenotype. The dominant role of genetic control in these more advanced
developmental systems is undisputed, and its proximate
workings in modern species represent the primary research
focus of modern developmental biology. But even in highly
controlled forms of development the realization of
morphology, particularly at the level of organogenesis,
continues to depend on nonprogrammatic, epigenetic
mechanisms. Among these are physicochemical, topological and biomechanical factors, as well as generic, stochastic and self-organizational properties of developing
tissues, and the complex dynamics of interactions between
these tissues (Salazar-Ciudad et al., 2003). Although there
is ample empirical evidence for the participation of these
factors in individual ontogenies, their inuence in setting
trajectories of morphological evolution is only recently
coming to be incorporated into the framework of evolutionary theory (Muller, 2007; Salazar-Ciudad, 2006).
See also: Morphology and Disparity through Time
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ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
level will contribute to a further locking in of new characters and will gain increasing organizational importance
as increasingly elaborate design features are incorporated.
The result is an ever closer mapping between genotype and
phenotype (Figure 2b).
But the evolution of homology does not stop at this
point. Once new building elements have become integrated
into the body design of a taxon, they can gain independence
from the mechanisms responsible for their initial establishment. This is suggested by those cases in which dierent
ontogenetic pathways are employed for the realization of
the same structures in dierent species. Skeletogenesis in
sea urchins, for example, involves the use of dierent progenitor lineages in direct developing species than it does in
those that pass through a larval stage (Wray and Ra,
1989). The orbitosphenoid, a component of the skull,
develops as membrane bone in worm lizards but as
replacement bone in other vertebrates (Bellairs and Gans,
1983). Meckels cartilage of the mandibular arch is induced
by the endoderm in amphibians but by the ectoderm in
higher vertebrates (Hall, 1983). Segmental development
in long germ band insects such as the fruity diers
considerably from this process in short germ band insects
like beetles, whereas the resulting structures are clearly
homologous. In other cases the expressed genes and eventual molecular make-up of embryonic structures have
changed during evolution of a lineage (Kiontke et al.,
2007). See also: Evolutionary Developmental Biology:
Homologous Regulatory Genes and Processes
These examples demonstrate that the same phenotypic
end-point can be reached by alternative developmental
modes and pathways. In other words, morphological
homology persists while its molecular, genetic and developmental components become free to drift (True and
Haag, 2001), a process that has been termed autonomization (Muller and Newman, 1999; Figure 2c). But since
the homologues themselves are maintained as constructional units of the phenotype, they assume a role as
independent organizers of body design and of the evolving
gene regulatory hierarchy (the Organizational Homology
Concept; Muller, 2003b). In this account, phenotypic
organization and the evolution of morphological form
becomes strongly determined by the specic set of homologues that a phylogenetic lineage has acquired.
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Further Reading
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