Ralph Greenspan


Dr. Greenspan’s research includes studies of the consequences of mutations and localized genetic alterations in the nervous system, molecular identification of genes underlying naturally occurring variation in behavior, and the genetic analysis of sleep and attention in the fruit fly. His current research addresses large-scale network interactions pertaining to the action of genes and neurons. In 2011, he was one of the small team of scientists that produced the white paper for the White House Office of Science and Technology Policy that eventuated in the BRAIN Initiative, and in 2014 spearheaded the effort to establish the state-funded Cal-BRAIN program and serves as its co-Director. In addition to research papers, he has authored: Fly Pushing: The Theory and Practice of Drosophila Genetics, An Introduction to Nervous Systems, and How Genes Influence Behaviour (with Jonathan Flint and Ken Kendler).

Select Publications

  • Shaw, P.J., Cirelli, C., Greenspan, R.J. and Tononi, G. (2000) Correlates of sleep and waking in Drosophila melanogaster. Science 287: 1834-1837.
  • Toma, D.P., White, K.P., Hirsch, J. and Greenspan, R.J. (2002) Identification of genes involved in Drosophila melanogaster geotaxis, a complex behavioral trait. Nature Genetics 31: 349-353.
  • Nitz, D.A., van Swinderen, B., Tononi, G. and Greenspan, R.J. (2002) Electrophysiological correlates of rest and activity in Drosophila melanogaster. Current Biology 12: 1934-1940.
  • van Swinderen, B. and Greenspan, R.J. (2003) Salience Modulates 20-30 Hz Brain Activity in Drosophila. Nature Neuroscience 6: 579-586.
  • van Swinderen, B., Nitz, D.A. and Greenspan, R.J. (2004) Uncoupling of brain activity from movement defines arousal states in Drosophila. Current Biology 14: 81-87.
  • Andretic, R., van Swinderen, B. and Greenspan, R.J. (2005) Dopaminergic modulation of arousal in Drosophila. Current Biology 15: 1165-1175.
  • Dierick, H.A. and Greenspan, R.J. (2006) Molecular analysis of flies selected for aggressive behavior. Nature Genetics 38: 1023-1031.
  • Foltenyi, K., Greenspan, R.J., and Newport, J.W. (2007) Activation of Egf-r and ERK by Rhomboid signaling regulates the consolidation and maintenance of sleep in Drosophila. Nature Neuroscience 10: 1160 – 1167.
  • Alivisatos, A.P., Chun, M., Church, G.M., Greenspan, R.J., Roukes, M.L. and Yuste, R. (2012) The brain activity map project and the challenge of functional connectomics. Neuron 74: 970 – 974.
  • Lin, C-Y., Chuang, C-C., Hua, T-E., Chen, C-C., Dickson, B.J., Greenspan, R.J. and Chiang, A-S. (2013) A comprehensive wiring diagram of the protocerebral bridge for visual information processing in the Drosophila brain. CELL Reports 3: 1739-1753.
  • Shih, C-T, Sporns, O, Yuan, S-L, Lin, Y-J, Su, T-S, Chuang, C-C., Lo, C-C, Greenspan, RJ and Chiang, A-S. (2014) Toward the Drosophila connectome: Structural analysis of the brain network. Curr. Biol., 25: 1249-1258.
  • Grover D, Katsuki T, Greenspan RJ. (2016) Flyception: imaging brain activity in freely walking fruit flies. Nat Methods. 13: 569-572.


Dr. Ralph J. Greenspan has worked on the genetic basis of behavior and brain function in fruit flies (Drosophila melanogaster) almost since the inception of the field, studying with one of its founders, Jeffery Hall, at Brandeis University in Massachusetts, where he received his Ph.D. in biology in 1979. He is currently Director of the Center for Brain Activity Mapping of the Kavli Institute for Brain and Mind at the University of California San Diego, and Professor in the Division of BIology’s Neurobiology Section and the Dept. of Cognitive Science.