By Dorothy Lehman Hoerr
What do a human being and a butterfly have in common?
You might expect them to share some genes. But only recently
are we discovering common links in the development of
various organisms, often in unexpected ways. For example, one
of the genes used to make appendages in many organisms -
such as the legs of both humans and insects - is also involved
in the development of eyespots on a butterfly's wings. 
Discoveries like this fall within the purview of evolutionary
developmental biology, known as EvoDevo, a field of science
that has developed rapidly in the past 30 years. Although the
relationship between evolution and development has attracted
interest ever since Darwin, the more recent resurgence in
EvoDevo has been enabled by the advent of molecular biology
as well as a desire to provide a more complete picture of how
evolution works.
Assistant Professor Gregory Davis uses the example of the
acquisition of bipedalism in the lineage leading to humans
from a more ape-like ancestor."To have a complete
understanding of that evolutionary transition, you would like
to know not only what genetic changes were responsible, but
exactly how those genetic changes altered development to
produce all the skeletal and musculature changes associated
with walking upright. This is the type of information we'd like
to know to obtain a more complete picture."
Over the last 30 years,Davis says, the search for that complete
picture has branched out in multiple directions, some
rather different from the field's origins.One of the major goals
of the senior seminar in EvoDevo is to introduce students to
what Davis calls in the syllabus the many "flavors of
evolutionary developmental biology."
EvoDevo has even gone beyond its initial boundaries of
filling the gap between evolution and development."The more
radical version is that there are ways in which the
developmental process of an organism actually influences the
direction that organisms can evolve," Davis explains."So once
you've got this complicated developmental process, you can
only tinker with it in some ways. It's this idea of developmental
constraint,which is really different. It's not just saying that to
have a complete picture, you need development. But it's saying
that development actually controls or influences the direction,
the evolutionary possibilities, that are out there. In amphibian
species that have evolved loss of digits, for example, the digits
that are lost tend to be those formed last during limb
development in species that possess all of their digits."
For the first half of the course, students in the class select
articles for discussion in areas that might include "classic"
EvoDevo (the broader comparisons of the original field),micro
or genetical EvoDevo (the study of small changes in more
closely related species),molecular developmental evolution,
polyphenism, or other miscellaneous topics such as
environmental sex determination or evolutionary loss."Because
the field has really matured and broken and splintered into all
these flavors,"Davis says,"I try to provide a sort of a la carte
menu of different case studies and topics and let students
choose which ones they want to pursue."
In each class period, two students present articles on their
topics and solicit feedback from the group to help them
identify a problem on which to focus. In a recent seminar, Jean
Chung chose to study the symbiotic relationship of the
Hawaiian bobtail squid with the bioluminescent bacterium
V. fisheri. Her question was how the squid, as it develops, is able
to select that particular bacterium. Caroline Feldman's thesis
focused on the sources of morphological variation in
vertebrates. She used dogs as the basis of her study because, due
to selective breeding, dog morphology is highly variable and yet
the genetic changes that cause these variations are relatively
easy to pinpoint. Similarly, Benjamin Pyenson chose to study
turtles in which sex determination is the result of
environmental factors like temperature and acidity."You could
really tease apart these bi-partite influences," he explained in his
presentation.
Another major goal of the course,Davis says, is for students
to "effectively communicate a scientific problem and how
they're going to approach it."
To this end, students give two
presentations of their thesis.One is a 10-minute talk to the
entire biology department; the other is an hour-long defense
before Davis and another faculty member who has also read
the student's paper.
In the second half of the course, students choose their own
topic to be the focus of a 20 to 25-page thesis.While the
seminar students don't do any actual research in the lab, the
papers are written in the style of a grant proposal, identifying a
problem in the chosen field and designing an experiment to
address it. The goal,Davis says, is for students "to become more
adept at weighing evidence and identifying holes in arguments, and then designing experiments to fill those holes, even if they
don't actually pursue the experiments."
This is one of the ways in which students grow most
in the course, Davis says.He describes how, in early
conversations with students about their topic, they may be
reluctant to share their ideas, but over the term, he watches
their confidence and ownership of the project grow."By the
time they get to their defense, there's less need to have
qualifiers on the ends of their sentences. Based on the
intonation, there's less hesitancy."One of the greatest rewards,
he says, is when the other faculty member questions some
aspect of the student's work and the student is able to respond
to the criticism confidently and convincingly. 
Davis stresses to students that the defense is a unique
opportunity to discuss their project with two other experts in
the field. But a lot of preparation is necessary to take full
advantage of that opportunity. Over the term, the students'
theses go through numerous revisions, receiving feedback from
Davis and a student partner. Students also practice their
presentations in front of the class and work together to
improve them.
Nicolette Belletier '09 found the presentations the most
rewarding part of the course."The most challenging aspect of
the presentation," she says,"was to explain my project in a
nutshell while providing enough detail for everyone in the
audience to understand the topic and appreciate why it is
interesting." She feels the assignment gave her valuable
experience in making a formal presentation.
Belletier says she was nervous before the thesis defense but
quickly became comfortable.My defense was with Professor
Davis and another professor I felt comfortable with and we just
talked through what I had written." In the end, she says,"The
whole experience was kind of fun and made me less
intimidated about receiving criticism of my writing."
For a reasonably good survey of different topics in the field:
For recent discussions on the role of developmental plasticity in evolution and medicine:

"Development actually controls
or influences the direction, the
evolutionary possibilities, that
are out there."

The advantage of large horn size for
dung beetles in terms of mating
success may come at the cost of
diminished copulatory organs. The
size of other features, such as eyes,
wings, or antennae may also be
affected in different species by the
location of the horns.

Seminar students' papers are
written in the style of a grant
proposal, identifying a problem
in the chosen field and
designing an experiment to
address it.

Professor Davis says one of the
greatest rewards of teaching the
course is when the other faculty
member questions some aspect of
the student's work and the student
is able to respond to the criticism
confidently and convincingly.

"The senior seminar in
developmental evolutionary
biology introduces students to
the many ‘flavors' of this field."

Possible genetic changes underlie changes in the skulls of
Bull Terriers favored by breeders over the past 80 years.