Kristi A. Wharton Professor of Biology

A.B. Biology 1978
Cornell University, Ithaca, NY

Ph.D. Biology  1986
Yale University, New Haven, CT


Postodocatoral Fellow  1986-87                             Biology Department, Yale University, New Haven, CT     

Postodocatoral Fellow  1987
Institute of Molecular Biology and Biotechnology  University of Crete Greece

Postodocatoral Fellow  1988-91
Department of Cell and Debelopmental Biology   Harvard University, Cambridge, MA  

Assisstant Professor (Research)  1991-95
Department of Molecular Biology, Cell Biology and Biochemistry                                          Brown University, Providence, RI               

Assisstant Professor of Medical Sciences  1995-2001
Department of Molecular Biology, Cell Biology and Biochemistry                                          Brown University, Providence, RI

Manning Assisstant Professor  2000-2001
Department of Molecular Biology, Cell Biology and Biochemistry                                          Brown University, Providence, RI          

Associate Professor of Medical Sciences  2001-2012
Department of Molecular Biology, Cell Biology and Biochemistry                                          Brown University, Providence, RI

Visiting Scientist  2003
Department of Cell Biology                                           Biozentrum, University of Basel, Switzerland

Visiting Scientist  2010
Departement Genetique et Developpement                     Institut de Genetique Humaine, CNRS           Montpellier, France

Professor of Medical Sciences  2012-present
Department of Molecular Biology, Cell Biology and Biochemistry                                          Brown University, Providence, RI

Brown Affiliations

scholarly work

James, R.E., Hoover, K.M., Bulgari, D., McLaughlin, C.N., Wilson, C.G., Wharton, K.A., Levitan, E.S., Broihier, H.T. (2014) Crimpy enables discrimination of presynaptic and postsynaptic pools of a BMP at the Drosophila neuromuscular junction. Developmental Cell 31, 586-598. PMCID: PMC4283799

Wharton, K.A. and Serpe, M. (2013) Fine tuned shuttles for bone morphogenetic proteins.  Curr Opin Genet Dev 23, 374-384. PMCID: PMC3760970 (invited review)

Boulanger, A., Farge, M., Ramanoudjame, C., Wharton, K., Dura, J-M. (2012) Drosophila motor neuron retraction during metamorphosis is mediated by inputs from TGF-b/BMP signaling  and orphan nuclear receptors. PLoS ONE 7(7): e40255. doi:10.1371/journal.pone.0040255. PMCID: PMC339-346

Akiyama, T., Marqués, G. and Wharton, K.A. (2012) Conserved unconventional proconvertase processing site produces a large bioactive BMP ligand with signaling properties distinct from the conventional C-terminally derived ligand. Science Signalling 5, ra28.  PMCID: PMC3699424

Le, V.Q. and Wharton, K.A. (2012) Hyperactive BMP signaling induced by ALK2R206H requires type II receptor function in a Drosophila model for classic Fibrodysplasia Ossificans Progressiva. Developmental Dynamics 241, 200-214. (Special Issue: Drosophila as a model for understanding development and disease) PMCID: PMC3454517

Ballard, S., Jarolimova, J. and Wharton, K.A. (2010) Gbb/BMP signaling is required to maintain energy homeostasis in Drosophila.  Developmental Biology 337, 375-85. PMCID: PMC2838617

Twombly, V., Bangi, E., Le, V.Q., Malnic, B., Singer, M. and Wharton, K.A. (2009).  Functional analysis of saxophone, the Drosophila BMP type I receptor ortholog of human ALK1/ ACVRL1 and ACVR1/ALK2. Genetics 183, 563-79. PMCID: PMC2766317

Wharton, K.A. and Derynck, R. (2009) TGF-β family signaling: Novel insights in development and disease. Development 136, 3691-97. PMID: 19855012 

Prabhat, Forsberg, A., Katzourin, M., Wharton, K.A. and Slater, M (2007).  A Comparative Study of Desktop, Fishtank and Cave Systems for the Exploration of Volume Rendered Confocal Data Sets. IEEE Transactions on Visualization and Computer Graphics, epub. 06 Aug 2007.

Bangi, E. and Wharton, K. A. (2006). Dual function of the Drosophila ALK1/ALK2 orthologue, Saxophone, influences the BMP activity gradient critical for wing patterning. Development 133, 3295-3303.  PMID: 16887821

Bangi, E. and Wharton, K.A. (2006) Dpp and Gbb exhibit different effective ranges in the establishment of the BMP activity gradient critical for Drosophila wing patterning. Developmental Biology 295, 178-193.  PMID: 16643887

Sinenko, S.A., Kim, E.K., Wynn, R., Manfruelli, P., Ando, I., Wharton, K.A., Perrimon, N. and Mathey-Prevot, B. (2004). Yantar is a novel and evolutionary conserved gene involved in Drosophila hemocyte differentiation. Developmental Biology 273: 48-62. 

Ray, R.P. and Wharton, K.A. (2001) Twisted perspective: New insights into extracellular modulation of BMP signaling during development. Cell 104, 801-804. 

Ray, R. and Wharton, K.A. (2001) Context-dependent relationships between the BMPs gbb and dpp during development of the Drosophila wing imaginal disc. Development 128, 3913-3925.  PMID: 11641216

Wharton, K. A., Cook, J., Torres-Schumann, S., de Castro, K., Borod, E., Phillips, D. A., (1999). Genetic analysis of the BMP-related gene, gbb, identifies multiple requirements during Drosophila development. Genetics 152: 629-40. PMID: 10353905

Haerry, T. E., Khalsa, O., O'Connor, M. B., Wharton, K. A., (1998). Synergistic signaling by two BMP ligands through the SAX and TKV receptors controls wing growth and patterning in Drosophila. Development 125: 3977-87. PMID: 9735359

Khalsa, O., Yoon, J-W., Torres-Schumann, S., Wharton, K.A., (1998). TGF-β/BMP superfamily members, Gbb-60A and Dpp,cooperate to provide pattern information and establish cell identity in the Drosophila wing. Development 125: 2723-34. PMID: 9636086

Wharton, K.A., Ray, R.P., Findley, S.D., Duncan, H.E. and Gelbart, W.M. (1996) Molecular lesions associated with alleles of decapentaplegic identify residues necessary for TGF-β/ BMP cell signaling in Drosophila melanogaster. Genetics 142, 493-505. PMID: 8852848

Raftery, L.A., Twombly, V., Wharton, K.A. and Gelbart, W.M. (1995) Genetic screens to identify elements of the decapentaplegic signaling pathway in Drosophila. Genetics 139, 241-54. PMID: 7705627

Wharton, K. A. (1995) How many receptors does it take? Bioessays 17, 13-16.

Wharton, K.A., Ray, R. and Gelbart, W.M. (1993) An activity gradient of decapentaplegic is required for dorsal-ventral patterning in the Drosophila embryo. Development 117, 807-22. PMID: 8330541

Wharton, K.A., Thompsen, G.and Gelbart, W.M. (1991) Drosophila 60A gene, a TGF-β family member, is closely related to human bone morphogenetic proteins. PNAS 88, 2914-18. PMID: 1924384

Ramos, R.G.P., Grimwade, B.G., Wharton, K.A., Scottgale, T.N. and Artavanis-Tsakonas, S. (1989) Physical and functional definition of Drosophila Notch locus by P -element transformation. Genetics 123, 337-48. PMID: 2555253

Yedvobnick, B., Muskavitch, M.A.T., Wharton, K.A., Halpern, M.E., Paul, E., Grimwade, B.G., and Artavanis-Tsakonas, S. (1985) Molecular genetics of Drosophila neurogenesis. Cold Spring Harbor Sympos.Quant.Biology 50:841-854.

Artavanis-Tsakonas, S., Grimwade, B., Halpern, M.E., Johansen, K. Markopoulou, K., Ramos, R.G.P. Schlesinger-Bryant, R., Wharton, K.A. and Yedvobnick, B. (1985) Molecular biology of genes involved in the neurogenesis of Drosophila. Society for Developmental Biology 44th Annual Symposium. ed. Joseph Gall

Wharton, K.A., Johansen, K.M., Xu, T. and Artavanis-Tsakonas, S. (1985) Nucleotide sequence from the neurogenic locus Notch implies a gene product which shares homology with proteins containing EGF-like repeats. Cell 43, 567-81. PMID: 3935325

Wharton, K.A., Yedvobnick,B., Finnerty, V.G. and Artavanis-Tsakonas, S. (1985) opa: A novel family of transcribed repeats shared by the Notch locus and other developmentally regulated loci in Drosophila melanogaster. Cell 40: 55-62. PMID: 2981631

Artavanis-Tsakonas, S., Grimwade, B.G., Harrison, R.G., Markopoulou, K.,Muskavitch, M.A.T., Schlesinger-Bryant, R., Wharton, K.A. and Yedvobnick, B. (1984) The Notch locus of Drosophila melanogaster: A molecular analysis. Developmental Genetics 4: 233-254.

Mooseker, M.S., Bonder, E.M., Grimwade, B.G., Howe, C.L., Keller, T.C.S., Wasserman,R.H. and Wharton, K.A. (1982) Regulation of contractility, cytoskeletal structure and filament assembly in brush borders of the intestinal epithelial cells. Cold Spring Harbor Sympos.Quant.Biology 46: 855-870.

Mooseker, M.S., Pollard, T.D. and Wharton, K.A. (1982) Nucleated polymerization of actin from the two ends of microvillar filaments in the intestinal brush border. J.Cell Biol. 95:223-233.

research overview

Cell-cell communication is essential for the proper manifestation of many events that take place during the development of multi-cellular organisms, including fate specification, cell shape change and migration, growth, and differentiation. We are interested in the underlying molecular mechanisms that mediate cell signaling during these developmental events, as well as the developmental consequences associated with signaling dysfunction.  We are particularly interested in the regulation of signaling output by the BMP pathway. Our research in the Drosophila model system has led to new discoveries with regard to regulating ligand production and differential receptor contribution in the signaling complex.  We are currently applying these findings to studies of ALS, cleft lip and/or palate and fibrodysplasia ossificans progressiva (FOP). 

research statement

The goal of the research in our laboratory is to understand how TGF-B signaling molecules facilitate the communication between cells. Cellular communication is essential to the proper manifestation of cell movements, growth, and differentiation during the development of multi-cellular organisms. Secreted proteins, such as those which belong to the transforming growth factor-B (TGF-B ) superfamily, are thought to mediate intercellular communication through their binding and/or association with integral membrane proteins. The transmission of such a signal culminates in the activation or repression of gene expression in the responding cell, resulting in alterations in the cell's phenotype, its position in the cell cycle, the types or extent of proteins expressed on the cell's surface, and/or in further transcription of new sets of genes. TGF-B -related proteins have been identified in both vertebrates and invertebrates as signaling factors which are instrumental in the control of cellular growth and differentiation, as well as in inductive events that specify cell fates. In order to understand the control of cellular growth and differentiation central to normal development, it is important to elucidate the regulation and molecular and cellular mechanisms of TGF-B signaling. Our approach to elucidating these signaling mechanisms is to exploit the genetic, molecular, and cell biological techniques of Drosophila to analyze the genes and their products, which act in TGF-B-type signaling pathways, and to investigate the function and regulation of these molecules in vivo.

To initiate this research program and as a means toward understanding the specificity of TGF-B -related signaling molecules, we have identified a new Drosophila member of the superfamily, the 60A gene. As a first step toward understanding 60A function, we have generated mutations in the locus. The phenotype of 60A mutants coupled with its wild type expression pattern suggest that 60A is important in multiple developmental processes. We are currently analysing the role of 60A in these processes, as well as beginning to identify other components of the signaling pathway which mediates the 60A signal. It will be of particular interest to compare the action and signaling pathway of 60A with that of decapentaplegic, dpp, which is probably the TGF-B family member best understood in terms of in vivo function.

funded research


Biogen idec MA Inc

Title: SOD1-based ALS: Identification of biomarkers and therapeutics

Role in project: PI

Award dates: January 1, 2015 – Dec 31, 2017

Collaborator: Robert Reenan


The Judith and Jean Pape Adams Foundation

Title: Elucidating cellular dysfunction in ALS and restoring function

Role in project: PI

Award dates: Dec 31, 2014 – Dec 31, 2015


Emerging Areas of New Science DEANS Award

Title: Importance of BMP processing in craniofacial development

Role in project: PI

Co-PIs: Helena Taylor, Asst. Prof. Surgery and Pediatrics, Stephen Sullivan, Asst. Prof. Surgery and Pediatrics, Warren Alpert Medical School, Brown University

Award dates: July 1, 2014 – June 30, 2015 


National Institutes of Health - 2R01 GM068118-05

Title of project: Differential regulation of BMP signals in vivo

Role in project: PI

Project dates: 07/01/2011 – 06/30/2015



Cali Center Development Grant

Center for FOP Research and Related Disorders - University of Pennsylvania

Total project period: 02/01/2010 - 01/31/2011

Role in project: PI

Title of project: FOP in Drosophila: Playing (the) Sax and ACVR1 to decipher the tune

Center for FOP Research and Related Disorders - University of Pennsylvania

Total project period: 06/01/2007 – 10/31/2009

Role in project: PI

Title of project: FOP in Drosophila: Playing (the) Sax and ACVR1 to decipher the tune

Richard B. Salomon Faculty Research Award
Brown University
Office of the Vice President for Research
Total project period: 02/01/2004-–07/01/2005
Role in project: PI
Title of project: "Establishing assays to investigate BMP ligand type and receptor choice"

NIH/NCRR - 1 P20 RR15578-01
Center for Genetics and Genomics (Sedivy, J.-PI)
Total project period: 09/30/2000–-08/31/2005
Role in project: Investigator (K. Wharton)
Title of project: "Mechanism of BMP signaling"

American Cancer Society - RPG DDC-98790
Total project period: 07/01/1999-06/30/2002
Title of Project: "Functional analysis of signaling by multiple TGF-/BMP ligands"

National Science Foundation - IBN-9604769
Total project period: 03/15/1997-08/31/1999
Title of Project: "Analysis of TGF-/BMP-mediated Signaling"

American Heart Association - Established Investigatorship 94002580
Total project period: 07/01/1994-06/30/1999
Title of project: "Functional Analysis of TGF--mediated Cellular Signaling"

Salomon Faculty Research Award
Total project period: 02/01/1996-01/31/1997
Title of project: "TGF-β-mediated signaling during oogenesis"

National Science Foundation - IBN-9205808
Total project period: 09/01/1992-08/31/1996
Title of project: "Cellular communication during Drosophila development"

American Cancer Society - DB-32
Total project period: 01/01/1993-12/31/1995
Title of project: "TGF- cell signaling during development"

NECUSE –The New England Consortium for Undergraduate Science Education/
The Pew Charitable Trusts
Support for two Brown undergraduates to attend Annual Drosophila Conference
Awarded: 12/13/1993

American Cancer Society - Institutional Award
Total project period: 08/01/1991-06/30/1993
Title of project: "Functional analysis of a TGF--like gene during development"