Darcy B. Kelley, PhD

Academic Appointments

  • Harold Weintraub Professor of Biological Sciences

Current Administrative Positions

  • Executive Committee, Frontiers of Science
  • Co-Director, Doctoral Program in Neurobiology and Behavior
Darcy B. Kelley, PhD

The research in my laboratory focuses on the neurobiology of vocal communication using an NIH model organism, Xenopus laevis, as an experimental model system. In their dark, crowded ponds, sound is the primary mode of social communication and is essential for finding and attracting a mate.  The importance of vocalizations has shaped a nervous system specialized for detecting acoustic cues that can be used to respond appropriately to potential mates and to rivals.   

Xenopus cannot breathe while calling underwater and they produce sounds simply by contracting their vocal muscles. We used motor nerve stimulation in the isolated larynx to determine how neuromuscular activity and vocal organ tuning control acoustic properties of calls. Because activity on the laryngeal nerve precisely matches actual calls, components of the hindbrain vocal pattern generator have been identified in the isolated brain ex vivo using fictive calling patterns recorded from this nerve. 

Tracing auditory projections from the nerve from the hindbrain to the forebrain led to the Xenopus amygdala (CeA).  The amygdala both receives input from the auditory midbrain and projects to the component of the hindbrain vocal pattern generator responsible for determining call patterns and rhythms. Damage to the amygdala interferes with a male's ability to recognize acoustic cues conveying sex and reproductive state. In collaboration with Christensen-Dalsgaard at SDU, we devised a method to effectively simulate the temporal and spectral features of underwater sounds while recording from auditory neurons. We have also developed a set of behavioral assays to determine the importance of spectral and temporal components of vocal signals for recognition of sex and state.  Our combined behavioral and neurophysiological experience in this system provides unique expertise for determining the subset of spectrotemporal acoustic parameters essential for successful social communication.  In collaboration withe members of the Sociogenomics Initiative, we will be using a casette of highly conserved, socially responsive genes to drive neural activity in the amygdala and determine effects on fictive call patterns in transgenic frogs.

In each Xenopus species, males produce a unique advertisement call distinguished by its temporal pattern and spectral properties.  A cross-species analysis of these calls suggests that call types evolved from an ancestral burst pattern, becoming either simpler or more complex.  Each advertisement call type occurs in each of the major branches of the phylogenetic tree and probably serves to advertise species identity.  Some spectral features, however, are highly conserved within groups of closely related species and may reflect conserved morphological features of their vocal organs.

Departments and Divisions

  • Department of Biological Sciences (Columbia University)

Languages Spoken

  • French

Education and Training

  • BS, 1970 Biopsychology, Barnard College - Columbia University
  • PhD, 1975 Neuroscience, Rockefeller University

Lab Locations

  • ColumbiaDoctors - 1244 Amsterdam Avenue

    1244 Amsterdam Avenue
    913 Fairchild
    New York, NY 10027
    (212) 854-4939
    (212) 531-0425

Past Positions

  • 1970 - 1975
    Graduate Fellow, D. Pfaff, Rockefeller University
  • 1975 - 1977 
     Post-doctoral Fellow, F. Nottebohm, Rockefeller University
  • 1977 - 1978
    Assistant Professor, The Rockefeller University
  • 1978 - 1981        
    Assistant Professor, Department of Psychology, Princeton University
  • 1982 - 1984
    Instructor, Neural Systems and Behavior, Marine Biological Laboratory
  • 1982 - 1987
    Associate Professor, Dept. of Biological Sciences, Columbia University
  • 1985 - 1989
    Course Director, Neural Systems and Behavior, Marine Biological Laboratory
  • 1987 -  Present
    Professor, Department of Biological Sciences, Columbia University
  • 1995 -  Present
    Director, Doctoral Program in Neurobiology and Behavior, Columbia University


Recent Seminars and Invited Lectureships, 2010 – 2013 (see also: Invited Symposia, Plenary and Keynote Lectures)

  • USC, Neuroscience Program
  • Dalhousie University, Neuroscience Program
  • Retreat, Champalimaud Neuroscience Programme, Instituto Gulbenkian de Ciencia
  • UCLA, Sawyer lecture
  • Fowler Symposium, the Florida State University
  • Columbia University, Department of Neuroscience
  • Columbia University, Department of Ecology, Evolution and Environmental Biology
  • 2007 - 2009
  • University of Montana, Bozeman
  • Wake Forest University
  • Davidson College
  • Rochester Institute of Technology
  • University of Illinois at Champaign-Urbana

2003 - 2006

  • Washington University
  • University of British Columbia
  • Cornell University
  • Stanford
  • UCSF
  • UC Berkeley
  • UCSD
  • University of Missouri, Columbia University
  • Columbia University, Department of Physiology and Cellular Biophysics
  • University of California, Irvine
  • Forbes lectures, Marine Biological Laboratory (Grass Foundation)
  • North Carolina State University, Keck Center for Behavioral Biology
  • University of Chicago, Committee on Neurobiology
  • Queens College

2000 - 2002

  • Indiana University, Systems Neuroscience and Neuroethology
  • Harvard University, Neurobiology Department
  • Arbus lecture, University of Arizona
  • University of Pennsylvania, Program in Neuroscience
  • Skirball Institute, NYU Medical School
  • Georgetown University, Neuroscience Program
  • NYU, Department of Biological Sciences
  • Emory University, Department of Neuroscience, Graduate Student Invitee
  • NYU, Seminar on developmental cognitive neuroscience
  • Columbia, Department of Physiology
  • Purdue University, Neuroscience
  • UC Davis, Biology
  • Columbia University, Center for Neurobiology and Behavior
  • Fordham University, Biology
  • Marine Biological Laboratory, Neurobiology and Behavior Course
  • Marine Biological Laboratory, Computational Neuroscience Course
  • City University of New York, Biology
  • University of British Columbia, Neuroscience
  • Bowdoin College, Biology
  • Weil Medical College of Cornell University, Endocrine Grand Rounds

1997 - 1999

  • University of Arizona, Distinguished Lecturer in Developmental Neurobiology
  • University of Maryland, College Park
  • Smithsonian Tropical Research Station, Visiting Scholar
  • Marine Biological Laboratory, Computational Neuroscience Course
  • University of California at Davis, Biology Department
  • University of Oregon, Neuroscience Department
  • Oregon State University, Zoology Department
  • Swarthmore College, lecture series on "Communication in Biological Systems"
  • Columbia Univ., Graduate School of Arts and Sciences, Dean's Distinguished Lecture
  • The Rockefeller University, Neurobiology of Learning and Memory Course
  • Marine Biological Laboratory, Computational Neuroscience Course
  • University of Virginia, Biological Sciences
  • Case Western Reserve University, Depts of Neuroscience, Biological Sciences
  • New York University, Center for Neural Science
  • Smith College, Neuroscience and Behavior
  • Farleigh Dickenson University, Department of Biology
  • Lehman College, Department of Biology
  • University of Maryland School of Medicine, Department of Neurobiology
  • Phi Beta Kappa Visiting Scholar University of Iowa
  • Phi Beta Kappa Visiting Scholar Coe College

1995 - 1996

  • Purdue University, Keynote speaker, Neuroscience retreat
  • The Rockefeller University
  • Phi Beta Kappa Visiting Scholar University of Delaware and Villanova University
  • University of Washington, Bloedel Center
  • Society for Neuroscience, Special Lecture
  • Salk Institute for Biological Studies, Sloan Center for Theoretical Neurobiology
  • University of California at San Diego, Dept. of Biology
  • Mt. Sinai School of Medicine, Brookdale Institute for Molecular Biology
  • Emory University, Dept. Anatomy and Cell Biology
  • Dartmouth College, Department of Physiology
  • Center for Biomedical Research, Population Council
  • USC, Distinguished Lecturer in Neuroscience

1993 - 1994

  • Cornell University
  • University of Pittsburgh, Neuroscience retreat speaker
  • University of California at San Francisco
  • Northeastern University
  • Stanford University, Neuroscience retreat speaker
  • Marine Biological Laboratory, Neural Systems and Behavior Scholar-in-Residence
  • City University of New York
  • University of Florida at Talahasee
  • State University of New York at Albany
  • University of Texas at Austin
  • State University of New York at Buffalo
  • New York State Psychiatric Institute
  • University of Colorado, Health Sciences Center, Neuroscience retreat speaker
  • University of Massachusetts at Amherst
  • Indiana University

1990 - 1992

  • Oberlin College
  • Duke University
  • Rockefeller University
  • NIH
  • Columbia University, Department of Psychology
  • Princeton University
  • Rutgers University
  • New York Academy of Sciences
  • Mt. Sinai Medical School
  • New York Psychoanalytical Society
  • New York University
  • University of British Columbia
  • Pennsylvania State University
  • Williams College
  • University of Washington, Seattle
  • Mayo Clinic
  • Cornell University Medical School
  • University of Madrid

1988 - 1990

  • University of California, San Francisco
  • Cornell University
  • University of California, San Diego
  • Yale University
  • Wesleyan University
  • Bell Laboratories
  • University of Southern California
  • Hunter College
  • University of Orgeon, Eugene
  • Wellesley College
  • University of California, Riverside
  • Brown University
  • University of Pittsburgh
  • University of Minnesota
  • Roche Institute for Molecular Biology
  • Vassar College
  • Cornell University Medical College
  • Hunter College
  • The Rockefeller University
  • University of Pennsylvania
  • College of Physicians and Surgeons, Columbia University

1986 - 1987

  • Rockefeller University
  • University of Virginia
  • State University of New York, Stony Brook
  • Rutgers, Newark
  • University of California, Los Angeles
  • Vanderbilt University
  • American Museum of Natural History
  • Rutgers, New Brunswick
  • Michigan State University
  • Dalhousie University
  • University of Colorado
  • University of Chicago
  • Princeton University

1983 - 1985

  • State University of New York, Downstate
  • Rockefeller University
  • State University of New York at Buffalo
  • Cornell University
  • University of Connecticut at Storrs
  • Harvard University
  • University of Pittsburgh
  • College of Physicians and Surgeons
  • Cornell Univ. Med. College
  • Washington University
  • University of California, Los Angeles

Invited Symposia, Plenary and Keynote Lectures, 2003 – 2013

  • Hormonal control of circuits for complex behavior, HHMI Janelia Farm Research Campus Conference, 2013
  • International Society for Neuroethology, Roots Symposium, 2012
  • CABI Symposium, CUNY, 2012
  • Discussant, Gordon Conference on Genes and Behavior, 2012
  • Evolution of Neural Circuits, Cold Spring Harbor Laboratory, 2011
  • Keynote address, Champalimaud Neuroscience Programme, 2011
  • UCLA, the Sawyer Lecture, 2011
  • The Florida State University, Fowler Symposium, 2011
  • Conference on sexual differentiation, brain and behavior, HHMI, Janelia Farm, 2010
  • NMNH Smithsonian Institution, Senate of Scientists Speaker 2009
  • Indiana University Animal Behavior Conference, Plenary Speaker, 2008
  • Maine Biomedical Symposium, Plenary Speaker, 2008
  • SNRP meeting, New York City, Keynote address, 2008
  • AACU meeting on science in the liberal arts curriculum, Keynote address, 2008
  • Union College, Keynote Address, General Education Meeting, 2004
  • University of Chicago, Neural Coding Workshop, 2003
  • Gordon Conference, Hormones and Development ,2003
  • Associacion Estudiantes Biologia, University of Puerto Rico, 2003


  • International Society for Neuroethology, Plenary Lecture, Bonn Congress August, 2001
  • Gordon Conference on Metamorphosis, June 2001
  • The Rockefeller University, Alumni Reunion speaker, May 2001
  • Rand Symposium, International Society of Herpetologists, 1998

1994 - 1997

  • No-Coast-Nerve-Net-Meeting, Model Systems in Neuroethology, 1997
  • Winter Animal Behavior Conference, 1996
  • Yale University Center for Neurological Sciences, Neurobiology Symposium, 1995
  • Sexual selection and the nervous system, Int. Congress of Neuroethology, 1995
  • Breckenridge Conference on Steroid Hormones and Trophic Factors, 1995
  • Microscopy Society of America, 1994
  • Keystone Conference on Muscle Development, 1994

1990 - 1993

  • Conference on Steroid Actions on Excitable Cells, 1993
  • The Development of Brain Function, NYU, 1993
  • Dahlem Conference, Flexibility and Constraint in Behavioral Systems 1993
  • Biology of Xenopus , University of London, 1992
  • Conference on Vertebrate Reproductive Endocrinology, US Fisheries Commission, 1992            
  • Society for Developmental Psychobiology, 1992
  • Meeting in Neural Development, 1991
  • University of Toronto, Neuroscience Symposium,1991
  • National Dyslexia Research Foundation, Scientific Research Conference,1990

1986 -1989

  • Int. Cong. on Prostaglandins and Related Compound,   Winter Conference on Brain Research
  • Soc. Neurosci:Neural Control of Communication Conf. on Developmental Psychobiology
  • 1st International Cong. Neuroethology, Winter Conference on Animal Behavior           
  • 2nd International Congress of Neuroethology  Soc. Neurosci. - Presidential Symposium                               

Teaching Responsibilities

Current Graduate

  • Experimental Appeoches in the Neurosciences
  • Developmental and Systems Neuroscience

Current Undergraduate

  • Developmental and Systems Neuroscience


Committees / Societies / Memberships

Federal Government Public Advisory Panels; Current Membership (14 total)

  • NSF Activation 1 Preproposal review panel, 2012          
  • NICHHD T32 review panel, 2012, 2013  


  • Society for Neuroscience
  • Society for Experimental Biology

Current Boards

  • Wenner-Gren Foundation
  • American Association of Colleges and Universities

Honors and Awards

  • Sawyer lecture, UCLA , 2011
  • Harold Weintraub Chair in Biological Sciences,  2010
  • Distinguished lecture, Champalimaud Neuroscience Programme, 2010
  • Lenfest award, Columbia University,  2009
  • Kravitz lecture, Neurobiology, Marine Biological Laboratory,  2008
  • HHMI Professor 2002 – 2006; 2006 - 2010
  • Forbes lectureship, the Grass Foundation and the MBL,  2003
  • Arbas lectureship, University of Arizona,  2002
  • Society for Neuroscience, Special Lecturer,  2001
  • Society for Neuroethology, Plenary Lecturer,   2001
  • University of Arizona, Distinguished Lecturer in Developmental Neuroscience,  1999
  • Javits Neuroscience Investigator Award, NIH,   1995 - 2002
  • Phi Beta Kappa Visiting Scholar,   1996-97
  • Society for Neuroscience, Special Lecturer,   1995
  • University of Southern California, Distinguished Lecturer in Neuroscience,   1995
  • University of Florida at Talahasee, the Rushton Lectures,   1993
  • Society for Developmental Psychobiology, Wiley Distinguished Lecturer,   1992
  • Fellow, American Association for the Advancement of Science,   1989
  • Society for Neuroscience, Presidential Symposium,   1988
  • Wellesley College, Mayer Lectureship,  1988
  • Dalhousie University, Soc. Neuroscience Chapter Lecturer,  1987
  • Javits Neuroscience Investigator Award, NIH,   1988 - 1995
  • Vanderbilt University, Kennedy Lectureship,   1982
  • Research Career Development Award, NIH,   1981 - 1986
  • Alfred P. Sloan Foundation Research Fellowship in Neuroscience,   1978 - 1981
  • NIH Post-doctoral Fellowship,   1975 - 1977
  • National Science Foundation Graduate Fellowship,   1970 - 1973

Research Interests

  • Cell Specification and Differentiation
  • Cognitive/Systems Neuroscience
  • Motor Systems
  • Sensory Physiology
  • Synapses and Circuits

Lab Members

  • Ian Hall, Revson Post-doctoral Fellow
  • Taffeta Elliott, Columbia Science Fellow
  • Charlotte Barkan, Neurobiology and Behavior Graduate Fellow
  • Martha L. Tobias, Senior Research Scientist


Sarah Woolley  Department of Psychology, Columbia University  Spatiotemporal Receptive Fields of Neurons in the Xenopus Central Amygdala

Jakob Christensen-Dalsgaard Un


Albersheim-Carter, J., Blubaum, A., Ballagh, I., Missaghi, K., Siuda, E.R., McMurray, G., Bass, A.H., Dubuc, R., Kelley, D.B., Schmidt, M.F., Wilson, R. J.A., and Gray, P.A. 2015 Testing the evolutionary conservation of vocal motoneurons in vertebrates. Respiratory Physiology and Neurobiology. DOI: http://dx.doi.org/doi:10.1016/j.resp.2015.06.010

Leininger, E.C. and Kelley, D.B. 2015. Evolution of courtship songs in Xenopus; vocal pattern generation and sound production. Cytogenetic and Genome Research. DOI:10.1159/000433483

Leininger, E.C., Kitayama, K. and Kelley, D.B. 2015. Species-specific loss of sexual dimorphism in vocal effectors accompanies vocal simplification in African clawed frogs (Xenopus). Journal of Experimental Biology, 218, 849 - 857.

Tobias, M.L., Korsh, J. and Kelley, D.B. 2014. Evolution of male and female release calls in Xenopus. Behaviour, 148, 519 - 549.

Sweeney, L.S. and Kelley, D.B. 2014. Harnessing vocal patterns for social communication. Current Opinion in Neurobiology 28: 34 - 41.

Hall, Ian C., Irene H. Ballagh, and Darcy B. Kelley. 2013 The Xenopus amygdala mediates socially appropriate vocal communication signals.The Journal of Neuroscience 33.36, 14534-14548.

Leininger, Elizabeth C., and Darcy B. Kelley. 2013 Distinct neural and neuromuscular strategies underlie independent evolution of simplified advertisement calls. Proceedings of the Royal Society B: Biological Sciences 280.1756.

Nasipak, B.T. and Kelley, D.B. 2012. Developing laryngeal muscle of Xenopus laevisas a model system; androgen-driven myogenesis controls fiber type transformation. Devel. Neurobiol. 72, 664 - 675.

Tobias. M.T., Kelley, D.B. and Evans, B.J. 2011. Evolution of advertisement calls in African clawed frogs. Behaviour, 148, 519 - 549.

Elliott, T.M., Christensen-Dalsgaard, J., and Kelley, D.B. 2011. Temporally selective processing of communication signals by auditory midbrain neurons, J. Neurophysiol., 105, 1620 - 1632.

Zornik, E. and Kelley, D.B. 2011 Neuroendocrine basis for the hierarchical control of frog courtship vocalizations. Frontiers in Neuroendocrinology. 32, 353-366

Kelley, D.B. and Bass, A.H. 2010. Neurobiology of vocal communication: mechanisms for sensorimotor integration and vocal patterning. Current Opinion in Neurobiology, 20, 748 - 753.

Tobias, M.L., Kelley, D.B., Yin, D. Korsch, J. and Corke, A. 2010. Vocal competition in male Xenopus laevis. Behav. Ecol. and Sociobiol., 65: 1791 -1803.

Yang, E.-J. and Kelley, D.B. 2009. Hormones and the regulation of vocal patterns in amphibians: Xenopus laevis vocalizations as a model system. In Hormones, Brain and Behavior, D. Pfaff, A. Arnold, A. Etgen, S. Fahrbach and R. Rubin (Eds), Academic Press, Vol. 2, 693 - 705.

Baur, L., Nasipak, B.T. and Kelley, D.B. 2008. Sexually differentiated, androgen-regulated, larynx-specific myosin heavy chain isoforms in Xenopus tropicalis andXenopus laevis. Development, Genes and Evolution, 218, 371 - 179/

Nasipak, B.T. and Kelley, D.B. 2008. The genome of the diploid anuran Xenopus tropicalis contains a novel array of sarcoplasmic Myosin Heavy Chain genes expressed in larval muscle and larynx. Development, Genes and Evolution, 218, 389 - 397.

Zornik, E. and Kelley, D.B. 2008. Regulation of respiratory and vocal motor pools in the isolated brain of Xenopus laevis. J. Neurosci., 28, 612 - 621.

Elliott, T.M., Christensen-Dahlsgaard, J. and Kelley, D.B. 2007. Tone and call responses of units in the auditory nerve and dorsal medullary nucleus of Xenopus laevis. J. Comp. Physiol. 193, 1243 - 1257.

Yang, E-J., Nasipak, B.Y. and Kelley, D.B. 2007. Direct action of gonadotropin in brain integrates behavioral and reproductive functions. PNAS, 104, 2477 - 2482.

Vignal, C. and Kelley, D. 2007. Significance of temporal and spectral acoustic cues for sexual recognition in Xenopus laevis. Proceedings of the Royal Society B, 274, 479 - 488.

Elliott, T.M. and Kelley, D.B. 2007. Male discrimination of receptive and unreceptive female calls by temporal features. J Exp Biol. 210, 2836-42.

Zornik, E. and Kelley, D.B. 2007. Breathing and calling: neuronal networks in theXenopus laevis hindbrain. J. Comp. Neurol., 501, 303 - 315.