EVOLUTION & DEVELOPMENT

5:4, 327330 (2003)

Knowing your ancestors: themes in the history of evo-devo

Alan C. Love* and Rudolf A. Raff
Department of Biology, Indiana University, Bloomington, IN 47405, USA
*Author for correspondence (email: aclove@indiana.edu)

In his magnum opus on the structure of evolutionary theory,

Stephen Jay Gould oers the following justication for his
extensive attention to the history of evolutionary ideas: I
regard such analysis not as an antiquarian indulgence, but as
an optimal path to [a] proper understanding of our current
commitments, and the underlying reasons for our decisions
about them (Gould 2002). Evolutionary developmental
biology (evo-devo) has become a legitimate research
discipline in contemporary biology, and it is time to more
adequately trace its origins. A number of articles and
conferences have begun an attempt at addressing the
historical background to the emergence of evo-devo. The
Dibner Institute for the History of Science and Technology
has held two workshops entitled From Embryology to EvoDevo (summer 2001 at Marine Biological Laboratories in
Woods Hole, MA; fall 2002 at the Dibner Institute in
Cambridge, MA), with biologists, historians, and philosophers attempting to seek key contours for a history of evodevo. Some participants have attempted to highlight the
relevance of particular researchers, whereas others have tried
to isolate key institutional changes that fostered evo-devo as
we now see it. Similar work is also emerging in refereed
journals (Guralnick 2002).
Our goal here is to suggest that a dierent sorting of the
intellectual themes that constitute a history of evo-devo is
necessary. This requires an examination of assumptions
underlying recent discussions. As preliminary evidence for the
necessity of this reective step, it is striking that 20th century
work in comparative embryology, which clearly ts into an
early history of evo-devo, has yet to be fully explored. For
example, in the above-mentioned conferences, most of the
presentations were devoted to researchers within the tradition
of experimental embryology. Identifying some assumptions found in recent historical work reveals that current
histories of evo-devo have overlooked a major intellectual
stream. Exposure of this richer history is particularly
relevant for biologists involved in evo-devo because it serves
to set a perspective for ongoing research eorts in evolution
and development.
Much of the attention in discussions of an early history of
evo-devo have centered on the 19th century work of Ernst
Haeckel, and rightly so (Sander 2002; Hofeld and Olsson,
& BLACKWELL PUBLISHING, INC.

in press). Haeckel produced a revolutionary blending of

evolutionary (phylogenetic) and developmental themes, most
familiar of which is the biogenetic law. Various similar threads
of research in comparative embryology and morphology were
developed under dierent individuals (e.g., Gegenbaur,
Brooks, and Balfour) with the primary aim of discerning
the phylogenetic relationships among various taxa. The
emergence of a more experimental approach to organismal
development in the 1880s and 1890s by Roux, Driesch, and
others (Entwicklungsmechanik) gained a large portion of its
impetus from dissatisfaction with the stalemate over alternative explanations of these phylogenetic relationships. The
blossoming of experimental embryology, with its emphasis on
mechanical manipulation as opposed to mere description,
was connected with an increasingly vocalized expression that
comparative embryology and morphology were dead end
research programs, poignantly represented in the inability to
decide between competing explanations for particular phylogenetic sequences (Nyhart 1995, 2002; Bowler 1996). William
Batesons (1894 [1992], vi) trenchant account of these
problems captures the rationale of those turning to developmental mechanics or studies of variation at the time:
Out of the same facts of anatomy and development men of equal
ability and repute have brought the most opposite conclusions.
. . . Facts of the same kind will take us no further. The issue turns
not on the facts but on the assumptions. Surely we can do better
than this. Need we waste more eort in these vain and sophistical
disputes?

Many of the contemporary tools for exploring the

evolution of development have arisen by descent through
the lineage of an experimental approach to embryology, most
notably those from developmental genetics. It is because
of the dominance of genetic techniques in contemporary
developmental biology that this stream is seen as so crucial to
the history of evo-devo. But if we shift our attention away
from the tools of investigation to the agenda of problems
within contemporary evo-devo, experimental embryology
is not the appropriate intellectual ancestor for most of the
20th century. Although overshadowed by the empirical success
of experimental embryology and its descendent disciplines,
many key issues such as phylogenetic relationships, the
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origin of evolutionary innovations, and signicance of

developmental constraints are the problems of comparative
evolutionary embryology. These core agenda items are very
much in the tradition of Kowalevsky, Haeckel, Gegenbaur,
Balfour, and others that continued long after developmental
mechanics constituted a thriving and distinct research
program. Gavin de Beer, the most visible and inuential
theoretician in this tradition during the mid-20th century, has
received some attention (Hall 2000). His work primarily
emphasized heterochrony, but probably more important was
his ability to synthesize empirical studies to make general
theoretical points (de Beer 1958 [1st edition, 1930]).
Vigorous programs in comparative embryology were
carried out through the 20th century. For example, N. J.
Berrills studies on tunicates led him to explain the origin of
vertebrates via a mechanism of size increase by slippage in the
number of cell division cycles relative to gastrulation to get a
larger basic vertebrate embryo from a tiny ascidian (Berrill
1935, 1955). These studies had a far more integrative role with
respect to evolutionary problems at the time than did highly
visible and important contemporary studies in experimental
embryology, such as Viktor Hamburgers manipulations of
the chick embryo (Hamburger 1934) or Spemanns work on
the amphibian organizer (Spemann 1938). It is also the case
that research programs utilizing comparative developmental
data to draw phylogenetic inferences continued, despite a lack
of mainstream visibility. For example, D. T. Anderson
synthesized results from the descriptive embryology literature
to prepare presumptive fate maps for a number of taxa, which
then provided the raw material for detailed phylogenetic
comparisons among annelids and arthropods (Anderson
1973). Again, this approach was ancestral to modern
evo-devo in terms of its agenda of research problems. Though
the perceived impact of individuals in 20th century comparative evolutionary embryology may appear marginal when
compared with the visibility of Haeckelian inuences, there
are at least two reasons for reconsideration. First, the
empirical work of researchers like Berrill was being cited
in inuential presentations of the relationship between
evolution and development (e.g., de Beer 1958, pp. 3839).
Second, the restatement of key problems, such as the origin of
the vertebrates, at the beginning of the renewed interest in
rejoining evolution and development of the early 1980s
explicitly acknowledged these sources (Gans and Northcutt
1983; Northcutt and Gans 1983). (Notably, a recent review of
the same topic emphasizing results from developmental
genetics did not [Holland and Chen 2001].) Succinctly, we
should not confuse the success of a non-evolutionary
developmental genetic research program and the tools it
provided with an enduring investigative agenda that keeps the
intersection of evolution and development as its focus.
The overemphasis on experimental embryology is not
merely a function of the present success of its manipulative

tools. Such a viewpoint has also been fostered by the common

perspective that the decisive historical schism relevant for
understanding contemporary evo-devo occurred between
genetics and embryology before the Modern Synthesis. This
has encouraged some contemporary scientists to subscribe to
a characterization of evo-devo in terms of developmental
genetics as the bridge between genetic accounts of evolution
and a molecularized embryology (Arthur 2002; Wilkins 2002;
cf. Love, in press). Importantly, our target is not the oftdiscussed issue of whether embryology was excluded from the
Modern Synthesis. Rather we want to call attention to
research questions in embryology during and after the period
of the Modern Synthesis that provided theoretical and
experimental inspiration to those architects of contemporary
evo-devo who were all trained before the emergence of any
recognizable research program on developmental genetics.
Our criticism and positive proposal can be distilled into two
pictorial representations. Figure 1 represents a common view,
observable to dierent degrees and with varying emphasis in
the discussions of Wallace Arthur, Sean Carroll, Scott Gilbert,
and Adam Wilkins (among others), which sees evo-devo as a
merging of themes from the Modern Synthesis with the
developmental genetics that emerged from the tradition of

Fig. 1. Common view of the relevant developmental research

component (experimental embryology now transformed into
developmental genetics, highlighted by thickened dotted arrows)
that feeds into contemporary evo-devo.

Love and Ra
experimental embryology. The historically important ssure
occurs when genetics breaks with embryology and joins the
evolutionary intellectual lineage. The investigation of developmental biology, now a multifaceted experimental discipline,
remains separate from the evolutionary-genetic lineage.
Twentieth century comparative embryology is simply not
noticed. Figure 2, our proposed revision, highlights comparative embryology as a strand of research not captured by the
common view. Although less popular after the rst quarter of
the 20th century, it continues throughout (also largely separate
from the Modern Synthesis) and includes gures like de Beer,
Berrill, and Anderson (among others). The emphasis on the
problem of reconstructing phylogenetic relations is probably
most distinctive of this lineage, but there was also a strong
interest in uncovering developmental mechanisms underlying
evolution. The major proposed cause was heterochrony, which
was seen as a universal mechanism through much of the 20th
century (de Beer 1958; Gould 1977; McKinney and McNamara 1991; Ra 1996; McNamara 1997).
The revised pictorial time line diagrams the existence of
two separate tracks of developmental thought and practice in

Fig. 2. Revised view of relevant developmental research components that feed into contemporary evo-devo, highlighting the
lineage of comparative evolutionary embryology (indicated by
thickened dotted arrows) as a key source of the current agenda of
research problems that demand an integration of evolution and
development.

Themes in the history of evo-devo

329

the 20th century. It reconstructs part of the renewed

theoretical interest in the intersection between evolution and
development that began before the developmental genetic
discoveries of the early 1980s (Gould 1977; Alberch et al.
1979; Bonner 1982; Goodwin et al. 1983; Ra and Kaufman
1983). Finally, it illustrates that an evo-devo synthesis arose
out of the joining of each developmental stream in the wake
of major empirical advances within developmental genetics
(e.g., the discovery of the Hox genes).
Drawing attention to the dierence between the research
problems of comparative evolutionary embryology and the
investigative questions and techniques of experimental
embryology generates a more accurate vantage point on the
roots of the present incarnation of evo-devo. The dramatic
success of developmental genetics was important to the evodevo synthesis in three ways. First, the discovery of widely
conserved developmental regulatory genes caught the eye of
mainstream developmental biologists. Second, developmental
genetics provided powerful experimental methods for a
preexisting set of biological questions that had interested
many others long before. Finally, developmental genetics
oers new mechanistic explanations relevant to the evolution
of developmental processes. This aspect was visible early, for
example, in Lewis discussion of Ultrabithorax in Drosophila
(Lewis 1978). Incidentally, the evo-devo investigators of the
1980s also reected the distinctly dierent trajectories of
developmental thought identied here. Some were very much
a part of the comparative embryological theme in culture and
training. Others were trained in molecular developmental
biology and genetics but drew their inspiration and problems
from comparative evolutionary embryology.
Once we recognize the centrality of particular research
questions connected to the relation between evolution and
development, we can more readily recognize the input of other
themes. Thus, morphological and paleontological researchers
(e.g., D. Dwight Davis or William K. Gregory) who have
shared much of the problem agenda with comparative evolutionary embryology are part of the history of evo-devo, as well
as its present construction (Love, in press, and submitted).
More eort should be expended to see historical developments
in systematics during the 1970s1980s, eventuating in the
predominance of cladistics, as a key component of the present
structure of evo-devo. It is only a multifaceted history such as
this that will match multidisciplinary descriptions of the
present evo-devo research program.
EDB strives to forge a unication of genomic, developmental,
organismal, population, and natural selection approaches to
evolutionary change. It draws from development, evolution,
paleaeontology, molecular and systematic biology, but has its
own set of questions, approaches and methods (Hall 1999, xv).

The heterogeneity of disciplinary threads that compose

evo-devo, past and present, suggests that sleuthing the

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relevant history is going to be complex and demands

consideration of a wider swathe of disciplinary contributions.
(Our proposed revision in Fig. 2 would thus require the
addition of these components to begin moving toward a more
adequate representation of intellectual and disciplinary
lineages important to contemporary evo-devo.) Recognition
of this historical disciplinary heterogeneity is critical to
present eorts at forging a multidisciplinary evo-devo capable
of oering adequate answers to these research questions.
We conclude that the title of the recent Dibner Institute
workshops should be rephrased as a question: Did
embryology contribute to evo-devo? Instead of providing a
direct answer we have rephrased the query: What kind of
embryology contributed to the emergence of evo-devo, and
how? If we consider the laboratory tools most commonly
deployed today, our attention is drawn to experimental
embryology in its most prominent current manifestationFdevelopmental genetics. But if we are interested in a broad set of
research problems that evo-devo proponents claim have been
ignored within the framework of neo-Darwinism (Arthur
2000; Ra 2000; Wagner et al. 2000), then comparative
evolutionary embryology (along with fellow travelers such as
morphology and paleontology) is the appropriate place to
look. Highlighting this inherited problem agenda from
comparative embryology and morphology is a reminder that
the recent success of developmental genetics should not be
confused with robust answers to the perennial questions of
phylogeny, innovation, and constraints in the history of life
that provide a continuing impetus to discern the relationship
between evolution and development.
Acknowledgment
We thank Wallace Arthur and Alec Panchen for critical comments
on an earlier version of this article.

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