Developmental Biology 427 (2017) 177–178
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Developmental Biology
journal homepage: www.elsevier.com/locate/developmentalbiology
Editorial
EvoDevo: Changes in developmental controls underlying the evolution
of animal body plans
Some have two feet
And some have four.
Some have six feet
And some have more.
Where do they come from? I can’t say.
But I bet they have come a long, long way.
 One Fish, Two Fish, Red Fish, Blue Fish 
By Dr. Seuss
When our son was little, we used to read Dr. Seuss’ One Fish,
Two Fish, Red Fish, Blue Fish to him every night. The most exciting
part for him was to count the eleven fingers on an imaginary an-
imal that is illustrated in the book. This hairy animal has seven
fingers on the right hand and four on the left. It was a fun exercise
to wonder how developmental programs could have possibly
produced asymmetric hands and whether natural selection would
favor this trait. However, if one goes to the real world and looks
carefully, it becomes apparent that in the history of life, the great
diversity of animal body plans and adaptive modifications within
each animal lineage are even more spectacular than illustrations in
children's books and easily exceed one's imagination.
More than a half a billion years ago, in the early Cambrian,
representative body plans of most metazoan animal phyla had
emerged (Valentine et al., 1999). The phylogenetic relationships
between these animal phyla, as resolved by DNA sequences, have
provided a glimpse into the evolutionary history of animal di-
versity (Dunn et al., 2014). It is now known that non-bilaterians,
including ctenophores, sponges, placozoa and cnidarians, evolved
from the common ancestor of all multicellular animals and bran-
ched out early during metazoan evolution. Among the non-bila-
terians, cnidarians are considered to be the closest group to the
bilaterians. All of the bilaterians, except possibly early branching
acoel flatworms, are divided into two superphyla – the Proto-
stomia and the Deuterostomia. The Protostomia consist of more
than twenty phyla that fall into two diverse clades, the Lopho-
trochozoa (sometimes called Spiralia) and the Ecdysozoa. The
Deuterostomia superphylum contains Echinodermata, Hemi-
chordata and Chordata. Over the past few decades, developmental
genetic studies on model organisms have revealed the basic
principles behind constructing animal body plans. Since the dis-
covery that developmental genes are conserved in distantly re-
lated animal phyla [for examples see (Francois and Bier, 1995;
Halder et al., 1995; Holley et al., 1995; McGinnis et al., 1984)], the
field of Evolutionary Developmental Biology (EvoDevo) has laid
down a generalized approach for reconstructing possible genetic
toolkits that may build ancestral animal forms. Further, comparing
http://dx.doi.org/10.1016/j.ydbio.2017.05.023
0012-1606/& 2017 Published by Elsevier Inc.
differences between developmental mechanisms in extant animals
provides a framework for understanding Charles Darwin's evolu-
tionary theory of “descent with modification”. The morphological
characters of diverse animal forms are controlled by develop-
mental toolkit genes, which are wired into gene regulatory net-
works that are directed by information stored in the genome
(Davidson and Erwin, 2006). Changes in genomic sequences thus
alter network architecture and result in different morphologies
that are then selected by the environment. In recent years, with
the advent of high-throughput sequencing technologies and re-
vived interest in using diverse non-traditional model organisms
for developmental research, EvoDevo studies have provided many
new insights into the evolution of animal forms.
In this special issue, nine review articles, covering re-
presentative and emerging topics in the field of EvoDevo, are in-
cluded. With an increasing number of animal genomes having
been sequenced, Simakov and Kawashima take a comparative
perspective and discuss evolutionary changes of various genomic
characters (Simakov and Kawashima, 2017). Among these char-
acters, non-coding sequences are intimately related to the dy-
namic control of gene expression and transcript levels that are
essential for building a multicellular animal. Gaiti and colleagues
review recent insights gained from analyses of non-coding RNAs
and regulatory DNA in bilaterian, non-bilaterian and non-animal
holozoans (Gaiti et al., 2017). Linkages between regulatory factors
and non-coding sequences constitute gene regulatory networks
(GRNs) that direct animals to develop into their specific forms.
Cary and Hinman discuss GRN evolution in the phylum of Echi-
nodermata based on the framework of the well-resolved sea
urchin GRN (Cary and Hinman, 2017). Minokawa provides a review
focusing on the evolution of the network circuits that control
skeletogenesis in sea urchins (Minokawa, 2017). Modern verte-
brate head skeletons are derived mainly from cranial neural crest
cells that represent one of the great innovations in the vertebrate
lineage. Square and colleagues review current understandings
about the origin and diversification of the vertebrate head skeleton
(Square et al., 2017). During evolution, how an ancestral devel-
opmental mode may have been transformed into a drastically
different one has been a puzzle. In annelids, direct development in
clitellates, such as leech, is considered to be a derived trait, which
evolved from an indirect developing polychaete-like ancestor. Kuo
proposes that the polychaete-to-clitellate transition was likely
triggered by habitat changes from marine to freshwater and ter-
restrial environments (Kuo, 2017). Similar to animals shaped by
natural environments, domesticated species display dramatic
178 Editorial / Developmental Biology 427 (2017) 177–178
variations owing to human preferences. Two articles review recent
discoveries on the molecular basis of artificially selected traits in
pigeons and goldfish (Abe and Ota, 2017; Domyan and Shapiro,
2017). The last article in this special issue discusses the develop-
ment and evolution of a specific cell type, germ cells. Lochab and
Extavour provide a comprehensive review concluding that BMP
signaling regulates multiple reproductive processes in a wide
range of animal models (Lochab and Extavour, 2017).
We learn lessons from history. The history of animal life on this
planet provides an amazing showcase of numerous experimental
trials, producing a wide variety of morphological characters and
body forms. Since human beings began to dominate the earth, the
environment has changed with unprecedented speed, creating a
new epoch that has been called the Anthropocene. Will the evo-
lution of developmental programs in modern animals be able to
keep pace with selection pressures imposed by this changing
world? Understanding the evolutionary history of extant animals
may perhaps allow us to better predict evolution in the future.
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Yi-Hsien Su, Jr-Kai Yu
Institute of Cellular and Organismic Biology,
Academia Sinica, Taipei 11529, Taiwan