It’s just a stage. A phylotypic stage. Part I.

by Stephen F. Matheson
Originally posted on Panda's Thumb, December 2010.

Disputes and controversies in science are always a good thing. They're fun to read about (and to write
about), and they're bellwethers of the health of the enterprise. Moreover, they tend to stimulate thought
and experimentation. Whether scientists are bickering about evo-devo, or about stem cells in cancer, or
about prebiotic chemistry, and whether or not the climate is genial or hostile, the result is valuable.
Now of course, some controversies are invented by demagogues for political purposes. The dispute in
such cases is far less interesting and clearly less profitable, even if participation by scientists is
necessary.
In December 2010, two papers in Nature weighed in on a major scientific controversy that has its roots
in pre-Darwin embryology, fueled by some gigantic scientific personalities and even tinged with what
some would call fraud. This intense scientific dispute spawned a sort of doppelganger, a manufactured
controversy that is just one more invention of anti-evolution propagandists. The Nature cover story
gives us a great opportunity to look into the controversies, real and imagined, and to learn a lot about
evolution and development and the things we're still trying to understand about both.
The scientific dispute is an old one, dating to when scientists first began to study embryonic
development in earnest. Embryologists like the great Karl Ernst von Baer noticed that the embryos of
very different animals often looked so similar that they could hardly be distinguished from each other. A
chicken embryo, at some point, looks an awful lot like a human embryo. What does this mean? Two
schools of thought (roughly speaking) entered into competition, with evolution as the major subtext.
One set of ideas envisioned development as recapitulation: development was a sort of re-play of
evolution, with the organism recapitulating its evolutionary history as it took shape. Recapitulation
theory was the brainchild of Ernst Haeckel, whose view of development was codified as his Biogenetic
Law and sloganeered as "Ontogeny recapitulates phylogeny." Against recapitulation were the views of
von Baer and others; von Baer formulated his own set of laws, the third of which repudiates
recapitulation rather directly. Everyone agreed that embryos of different animals often looked quite
alike; the dispute was about what this meant. And it seems that those who opposed evolutionary
explanations (like von Baer) were eager to point to difference and divergence during development, while
those who championed evolutionary views wanted to emphasize the shocking similarities between, say,
chickens and mammals when compared at key embryological junctures.
Haeckel, famously, went on to point to those similarities as evidence for common ancestry and,
infamously, to create a certain illustration of that evidence. His picture, thought by even some
embryologists to be partly fraudulent (more accurately, "doctored"), is now a staple of anti-evolution
propaganda. You can read all about that elsewhere; suffice it to say that Haeckel's drawings have long
since been "corrected" without creating any problems for evolutionary theory. (For a much more
detailed treatment of this saga, see Richardson and Keuck, Biological Reviews, 2002.)
But interestingly, the debates about recapitulation
morphed (wink) over the years into a distinct but
related disagreement about whether animal
development passes through a stage that is common
to – or typical of – the lineage of the organism.
Because although Haeckel's recapitulation idea
didn't survive, it remained clear that development
seemed to reflect evolutionary commonalities.
Consider the photos on the right. (The figure was
created by Michael K. Richardson, who led the
research group that critiqued Haeckel's drawings in

1
1998.) While the various embryos shown all end up looking quite different – looking like the adult form,
in other words – they seem to "start" at a place that's notably similar. (Compare the embryos in the first
row.) That starting point is not the beginning of development, and in fact those different kinds of
embryos got to that starting point via rather different beginnings. In other words, it seems that animal
embryos pass through roughly three phases of development: an early phase that can vary from group to
group (say, between birds and mammals), a late phase in which group-specific forms are established,
and a middle phase that is eerily similar among groups. That middle phase has come to be known as the
"phylotypic stage" of development, meaning that it is a stage at which the embryo looks like a typical
example of its evolutionary group. For insects, this is thought to be the "extended germband" stage; for
vertebrates, it's roughly the tailbud stage. The point is that there is a middle phase of development
during which animal embryos of varying morphological destinies look very similar, even if their earlier
stages seemed very distinct. This model of developmental trajectories, compared across groups, is
known as the "hourglass model," nicely depicted by Richardson and colleagues in the cartoon below.
Why all the controversy? Well, the disputes all seem to be
related to the fact that the model is mostly descriptive. And
so, one criticism is that the model is based on what embryos
look like, and not strongly anchored in carefully-defined and
-measured characters. Moreover, some critics have noted
that the comparisons were often restricted to popular
laboratory species, such that when the analysis was
expanded to include a broader set of species, the similarities
in the waist of the hourglass become less striking. In other
words, the dispute centered on the basis of the model.
Critics were disputing the very existence of the phylotypic
stage.
Oh, and while this interesting scientific debate was ongoing,
some propagandists were shadowboxing with Haeckel's
ghost, shrieking about fraud while creating in the minds of
their dupes the illusion of a different debate: one about
whether development and evolution are conceptually linked.
Along the way, these busy demagogues suggested that the
phylotypic stage is an illusion, cherry-picking their data
more shamelessly than Haeckel ever did. In any case, these
folks were exploiting the real scientific dispute: whether the
phylotypic stage can be defined more rigorously, in a way that links the similarities (whatever they are)
to common ancestry.
And that brings us to the cover story in the 9 December 2010 issue of
Nature. The cover image depicts a version of Haeckel's infamous
illustration. The issue includes two reports, very different in their approach
and in the animals they examined. Both reports provide striking support for
the hourglass model, by showing that the phylotypic stage is indeed
characterized by distinctive and fascinating patterns of gene expression.
Parts II and III will explore those two papers.

Image credits: 1) embryo images from Gilbert, Developmental Biology, 6th

Edition, online at PubMed; 2) cartoon fromRichardson et al. 1998.