Biology at Williams

Faculty and Staff

Steven Swoap
Rob Savage
Robert Savage
Professor at Williams since 1997

Office: 017 TBL
Phone: (413) 597-3314
Area of Interest: Development

Courses taught 2009-2010
Honors and Summer Students 2009–2010

Research in the Savage lab


Research Interests

Metazoan pattern formation

Annelids. The long-term objective of my research is to investigate the cellular and molecular processes that control annelid development with an emphasis on understanding the evolution of segmental pattern formation. The characterization of developmental regulatory genes in species outside traditional model systems addresses fundamental questions regarding their role in divergent body plans. In fact, the emphasis of my research program addresses to what extant important regulatory genes have changed their function in the course of annelid evolution. Currently our lab is characterizing homologues to Drosophila segmentation genes, hunchback and caudal in two classes of annelids, polychaetes and leeches. Much of what is known about segmentation in annelids comes from studies in highly derived body plan of the leech. However, studies in the basal polychaete annelids will help to distinguish general developmental mechanisms from phylum-specific ones, and may show how these processes are modified to account for morphological diversity.

Annelid Genomics

The polychaete Capitella sp I and the leech Helobdella robusta are the two annelid genomes that have been sequenced (8x coverage for both) by JGI. The combination of available genomic data with over 300,000 ESTs has provided the opportunity to conduct genome-level comparisons within and outside the Lophotrochozoans. Our lab maintains cultures of each annelid and we currently house all the ESTs generated from the two sequencing projects. We are currently pursuing a number of wet lab and computer lab projects based on the available genomic information.



NIH Developmental Clusters Panel Member

NSF Developmental Mechanisms Panel Member

ad hoc reviewer for NSF, NIH and scientific journals

Director of MBL-Williams Summer Research Program

There are two goals to this undergraduate research experience. The first goal is to have students participate fully in an original research project.  The research question reflects my interests in experimental developmental biology and evolution and development known as “Evo-Devo".  During the eight-week program, the students characterized the spatiotemporal expression pattern of the zinc finger transcription factor protein called Hunchback in mollusk, nemertean, and annelid embryos using a cross-species antibody developed in my lab.  Using their data, the students compared hunchback-like expression patterns between phyla and afterwards they were able to infer evolutionary roles that this protein plays in lophotrochozoans.

An important second goal of the program is to expose students to a broad array of opportunities in biological sciences offered at the MBL each summer. This exposure is accomplished by having the students attend graduate-level courses, general lectures, and meet with resident scientists.

Williams College Student Participants


John Vu '09 , Fati Sammy '09, Jessica Diehl '09, Taylor Wilson-Hill '09, Rob Savage, Tynisha Smalls '06, and Emily Behrmen '09


Based on our preliminary data, we focused our efforts on characterizing the hunchback-like expression patterns in two gastropod embryos, Crepidula fornicata and Ilyanassa obsoleta. We also explored new animal systems such as nemerteans and echiurid worms.  These two latter lophotrochozoan phyla posed exciting challenges for us.  In the end, the students worked extensively with animals from four different lophotrochozoan phyla and generated an outstanding body of preliminary data (see below).

Molluscs           Annelids           Nemerteans                  Echiurids

Gastropods           Polychaetes       Cerebratulus lacteus     Urechis caupo

Ilyanassa             Chaetopterus

Crepidula              Nereis


In all the images below, hunchback-like reactive nuclie are labeled by the cross-species polyclonal antibody generated in my lab.

Crepidula fornicata: 32 cell embryo

Ilyanassa obsoleta: 16 cell embryo

Nereis succinea: 24 hr gastrula embryo// The night-lighting group!

Urechis caupo: 24 hr embryo

Our data suggest that the cross-species antibody recognizes hunchback-like protein in three lophotrochozoan phyla: annelids, molluscs and echiurans. To date, tthe spatiotemporal expression patterns of hunchback-like protein in embryogenesis in the three phyla are highly conserved.


As part of our finale of our summer research experience, I made a wonderful seafood soup packed with representatives from five marine phyla. Can you name the five??

From left to right: Jessica Diehl '09, Maia Hirsch and Fatti Sammy '09 loved the soup. Children

were not exempt from the phyletic experience!

Selected Publications

Book Chapter

*Williams College student or alumnus