Robert Savage

Professor of Biology

Thompson Biology Lab Rm 203
At Williams since 1997


B.A. Bowdoin College (1987)
Ph.D. Wesleyan University, Biology (1993)

Areas of Expertise



Note: courses in gray are not offered this academic year.

Scholarship/Creative Work

Selected Publications

  1. Savage, R. and Phillips, C. R. (1989). Signals from the dorsal blastopore lip region during gastrulation bias the ectoderm toward a nonepidermal pathway of differentiation in Xenopus laevis. Developmental Biology 133, 157-168.
  2. Savage, R. M. and Danilchik, M. (1993). Dynamics of germ plasm localization and its inhibition by uv irradiation in early cleavage Xenopus embryos. Developmental Biology 157, 371-382.
  3. Savage, R. M. and Shankland, M. (1996). Identification and characterization of a hunchback orthologue, Lzf2, and its expression during leech embryogenesis. Developmental Biology 175, 205-217.
  4. Shankland, M. and Savage, R. M. (1997). Annelid, the Segmented Worms. In “Embryology: Constructing the Organism” (S.F.Gilbert and A.M.Raunio, Eds.) Sinauer Associates; Sunderland, MA.
  5. Iwasa*, J., Suver*, D., and Savage, R.M. (2000) The leech hunchback gene is expressed in epidermis and CNS but not in the segmental precursor lineages. Development, Genes and Evolution 210 (6) 277-288.
  6. Werbrock*, A.H., Meiklejohn*, D.A., Sainz, A*., Iwasa*, J.H., and Savage, R.M. (2001). A Polychaete hunchback Ortholog. Developmental Biology 235, 476-488.
  7. Shimizu, T and Savage, R.M. (2002). Expression of hunchback protein in a subset of ectodermal teloblasts of the oligochaete anneldi Tubifex. Dev Genes Evol 212;520-525.
  8. Pinnell*, J., Lindeman*, P.S., Colavito*, S., Lowe, C. and Savage, R. M. (2006). The divergent roles of the segmentation gene hunchback. Integrative and Comparative Biology 46: 519-532.
  9. Schmerer, M., Savage, RM., and Shankland, M. (2009). Paxbeta: a novel family of lophotrochozoan Pax genes. Evol Dev 11(6): 689-96.
  10. Dean, D., Himes, CM., Behrman*, E, and Savage, RM. (2009). Hunchback-like protein is expressed in cleavage blastomeres, gastrula epithelium, and ciliary structures in gastropods. Biol Bull 217(2): 189-201.
  11. Simakov O, Marletaz F, Cho SJ, Edsinger-Gonzales E, Havlak P, Hellsten U, Kuo DH, Larsson T, Lv J, Arendt D, Savage R, Osoegawa K, de Jong P, Grimwood J, Chapman JA, Shapiro H, Aerts A, Otillar RP, Terry AY, Boore JL, Grigoriev IV, Lindberg DR, Seaver EC, Weisblat DA, Putnam NH, Rokhsar DS. (2012).  Insights into bilaterian evolution from three spiralian genomes.  Nature. 2012 Dec 19. doi: 10.1038/nature11696.

*Williams College student or alumnus

Book Chapter

  • Hill, SD and Savage, RM. (2009). Book chapter entitled: “Evolution, Development and Ecology of Capitella sp. I: A Waxing Model for Polychaete Studies” in Annelids in Modern Biology edited by Daniel Shain. John Wiley and Sons, Hoboken, NJ.


Awards, Fellowships & Grants

  • National Institutes of Health: “Segmental Pattern Formation in Annelids”, GM57778-01 ($108,693 7/1/98-6/30/01)
  • National Science Foundation, IBN-Evolution of Developmental Mechanisms: “The Roles of Segmentation Gene Homologues in Annelids”, IBN-0090378, ($292,489 1/1/01-12/31/04)
  • National Institutes of Health: “Segmental Pattern Formation in Annelids”, GM071444 ($342,489 8/1/04-7/31/07)
  • National Institutes of Health: “Segmental Pattern Formation in Annelids’, GM71444 ($214,990 8/1/07-7/31/11)
  • MBL Associates Endowed Fellowship Summer 2009

Current Committees

  • Biochemistry Program
  • Committee on Academic Standing, Chair

Research Interests

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. 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 potential segmentation gene homologues obtained from an unbiased subtraction screen completed in polychaetes and leeches (see Annelid Genomics below). Much of what is known about segmentation in annelids comes from studies in the highly derived body plan of the leech. Therefore, concurrent 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

Our lab has produced seven high quality primary and subtraction cDNA libraries from which the Joint Genome Institute (JGI) sequenced ~140,000 ESTs.  In the end, our lab contributed all the ESTs for the Helobdella genome project and the  majority  ESTs for the Capitella genome project.  Microtiter plates are stored at Williams College as a resource for the scientific community.

With the subtraction libraries in hand, we designed an unbiased approach that enriched for gene products expressed during annelid segmentation.  The strategy was simple yet novel.  ESTs collected from subtraction libraries, enriched with cDNAs expressed at the time of segmentation, were compared between annelids (leeches and polychaetes) as opposed to comparing ESTs from other phyla.  The assumption was that the two annelid groups were more likely to share a common mechanism of segmental patterning than any interphyletic comparison.

Our lab has begun to characterize the spatiotemporal expression patterns of annelid homologues to developmental regulatory genes identified from our subtraction libraries. We have initiated an in situ hybridization screen in leech and focused our efforts on key stages of segment formation but prior to overt morphological segmentation for the top 100 segmentation gene candidates.  This screening strategy offers the opportunity to identify  gene products expressed in a molecular reiterated prepattern before overt segmentation is observed ~24 hr later.

In Figure 1 C and D, we have discovered a bilateral punctate mRNA expression pattern that appears shortly after germinal band coalesence (left and right sides of the trunk) and that this bilateral pattern persists until midway through the fusion of the left and rights sides of the trunk.  Once the  the segmented trunk is formed, the bilateral expression pattern is lost which is exactly time period when overt morphological segmentation is first observed.  We are continuing our screen and beginning a series of functional studies to determine the significance of these molecular prepatterns.

Figure 1.  Whole mount in situ mRNA patterns in three distinct expression domains in stage 8 leech embryos. In the unfused germinal bands domain, Epidermal Growth Factor Receptor (EGFR in A) and Fibroblast Growth Factor Receptor (FGFR in B) mRNAs are expressed in unfused germinal bands (see bracketed area) after they reach the surface and begin to move circumferentially towards the ventral midline.   EGFR and FGFR are two of three gene products expressed prior to coalescence identified from our initial screen.  During the early coalescence domain, ladybird homeobox (Lbx, C) and novel MAPK (CEK-2, D) mRNAs are expressed in a punctate bilateral pattern when the two germinal bands first coalescence.  During the mid-coalescence domain, Hamlet (E) and FYVE zinc finger (F) mRNAs are expressed in stripes in four anterior segments.  The uniform pattern of expression in younger, posterior tissue resolves into stripes in the older, anterior germinal plate. Mid-coalescence stage occurs at least 20 hours before overt morphological segmentation is observed in late stage 8 leech embryos (see Fig 3).  Anterior up, sb 50 um.

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