Marc Schmidt

Dr. Marc Schmidt

Professor of Biology

Co-Director, Undergraduate Neuroscience Program


Research Interests

217 Leidy Laboratory


B.A., Swarthmore College, Biology, 1986

PhD, Colorado State University, Neuroscience, 1993

Postdoc, California Institute of Technology, Neuroethology, 1993-1999


Research Interests

Our laboratory is interested in the neural mechanisms that underlie courtship display. Much of our current work is focused on studying vocal and non-vocal displays in the brown-headed cowbird (Molothrus ater), a highly gregarious species whose complex social structure is necessary for group cohesion, mating and reproductive success. The work in the laboratory ranges from neural circuits level experiments to computational-based approaches for tracking group behavior. Specifically, we have built a "smart aviary" equipped with an array of computer vision cameras and microphones that allow us to track moment-to-moment behavior of an entire group of songbirds throughout the entire breeding season (see link for story on our aviary). By its nature, our work is highly interdisciplinary and we have close collaborations with several laboratories both. These include the Daniilidis lab (Penn Engineering) where we implement novel computer vision approaches for tracking birds in our “smart aviary”, the Balasubramanian lab (Penn Physics) where we develop novel techniques for quantifying social network dynamics and the Aflatouni lab (Penn Electrical Engineering) where we are developing novel non-invasive wireless neural recording devices for long term tracking of brain activity within our aviary. We also continue our productive collaboration with the White lab (Wilfried Laurier University, Ontario, CA) where we can study the effect of brain circuit manipulations on cowbird social behavior and reproductive success across many years.

Courses Taught


BIOL251/BBB251: Cellular and Molecular Neurobiology (Fall)

BIOL231: Evolution of Animal Behavior (Fall)

Selected Publications

Perkes A, Pfrommer B, Daniilidis K, White DJ, and M. F. Schmidt (2022). Variation in female songbird state determines signal strength needed to evoke copulation.  eLIFE (under revision); bioRxiv preprint doi:

Perkes A, Anderson H, Gros-Louis J, Schmidt M.F. and D. White (2022) Cohesion in male singing behavior predicts group reproductive output in a social songbird. (submitted); bioRxiv preprint doi:

Badger M., Xiao S., Wang Y., Perkes A., Schmidt M.F. and Kostas Daniilidis (2022) Multi-view Tracking, ReID, and Social Network Analysis of a Flock of Visually Similar Birds in an Outdoor Aviary. International Journal of Computer Vision (submitted)

Schwark R. W., Fuxjager M. J. and M. F Schmidt (2022) Proposing a neural framework for the evolution of elaborate courtship displays. eLIFE

Anderson H, Perkes A, Gottfried J, Davies H, White D and M. F. Schmidt (2021) Female signal jamming in a socially monogamous brood parasite. Animal Behaviour. 2021 February; 172:155-169. Available from: DOI: 10.1016/j.anbehav.2020.10.011.

Badger M, Wang Y, Modh A, Perkes A, Kolotouros N, Pfrommer B, Schmidt M and Daniilidis K. (2020) Lecture Notes in Computer Science. Cham: Springer International Publishing; 2020. Chapter 1, 3D Bird Reconstruction: A Dataset, Model, and Shape Recovery from a Single View. 1-17p. Available from: DOI: 10.1007/978-3- 030-58523-5_1

Sheldon*, Z., Castelino C. E., Glaze, C., Yau, E., Bibu S. and M.F. Schmidt (2020) Regulation of vocal precision by noradrenergic modulation of a motor nucleus. Journal of Neurophysiology 124: 458–470, 2020. doi:10.1152/jn.00154.2020. (*chosen as APS select paper of the month)

Perkes, A., White. D. J., J. M. Wild and M. F. Schmidt (2019) Female songbirds: the unsung drivers of social behavior. Behavioral Processes 163: 60 – 70

Schmidt M. F. and F. Goller (2016) Breathtaking songs: Coordinating the neural circuits for breathing and singing. Physiology. 31: 442-451

Albersheim-Carter, J., A. Blubaum, I. H. Ballagh, K. Missaghi, E. R. Siuda, G. Mc Murray, A. H. Bass, R. Dubuc, D. B. Kelley, M. F. Schmidt, R. J.A. Wilson, and P. A. Gray (2016) Testing the evolutionary conservation of specialized vocal motoneurons in vertebrates. Respiratory Physiology and Neurobiology. 224: 2 – 10.

Schmidt M.F. and J. M. Wild (2014) The respiratory-vocal system of songbirds: Anatomy, physiology and neural control. Progress in Brain Research, Vol 212: 297 - 335. []

Maguire S., M. F.  Schmidt and D. J. White (2013) Social brains in context: Lesions to the song control system in female cowbirds affect their social network. PLoS ONE 8(5): e63239. [Maguire et al. (PLOS ONE 2013)]

Lewandowski B.C, Alexei A., Hahnloser R. and M.F.Schmidt (2013) At the interface between the auditory and vocal motor system: NIf and its role in vocal processing, production and learning J. Physiology (Paris) 107:178-192 [Lewandowski et al. (J. Physiol Paris 2013)]

Mclean J., S. Bricault and M. F. Schmidt (2013) Characterization of respiratory neurons in the rostral ventrolateral medulla, an area critical for vocal production in songbirds. J. Neurophysiology 109: 948-957. [McLean et al (JNP 2013)]

Raksin J. N., C. Glaze, S. Smith and M. F. Schmidt (2012) Linear and Nonlinear Auditory Response Properties of Interneurons in a High Order Avian Vocal Motor Nucleus During Wakefulness. J. Neurophysiology 107:2185-2201 [Raksin et al. (JNP 2012)]

Schmidt, M. F., J. Mc Lean and F. Goller (2011) Breathing and Vocal Control: The Respiratory System as both a Driver and Target of Telencephalic Vocal Motor Circuits in Songbirds. J. Exp. Physiology 97 (4) 455-461 [Schmidt et al. (Exp Physiol-2012)]

 Lewandowski B. C. and M. F. Schmidt (2011) Short bouts of vocalization induce long lasting fast gamma oscillations in a sensorimotor nucleus. J. Neuroscience 31(39): 13936-13948; doi: 10.1523/ JNEUROSCI.6809-10.2011 [Lewandowski and Schmidt (JN 2011)]

Margoliash, D. and M.F. Schmidt (2010) Sleep, off-line processing, and vocal learning. Brain and Language 115: 45 – 58. [Margoliash ad Schmidt (Brain and Language 2009)]

Castelino, C. B. and M. F. Schmidt (2010) What birdsong can teach us about the central noradrenergic system. J. Chem. Neuroanatomy 39: 96 – 111 [Castelino and Schmidt (J. Chem. Neuroanat. 2010)]

Schmidt, M. F. (2008) Using Both Sides of Your Brain: The Case for Rapid Interhemispheric Switching. PLoS Biology 6: 2089 – 2093 [PLoS Primer (2008)]

Ashmore R. C., J. A. Renk and M. F. Schmidt (2008) Bottom-up Activation of Forebrain Vocal Motor Structures by the Respiratory Brainstem. J. Neuroscience 28: 2613 – 2623. [Ashmore, Renk and Schmidt (JN 2008)]

Schmidt, M. F. and R. C. Ashmore (2008) Integrating breathing and singing: Forebrain and brainstem mechanisms in Neuroscience of Birdsong (ed. Zeigler, H. P. and P. Marler) Cambridge University Press. [Schmidt and Ashmore (Book Chapter 2008)]

Ashmore R. C., M. Bourjaily and M. F. Schmidt (2008) Hemispheric coordination is necessary for song production in adult birds: Implications for a dual role for forebrain nuclei in vocal motor control. J. Neurophysiology 99: 373–385. [Ashmore, Bourjaily and Schmidt (JNP 2008)]

Nealen P. M. and M. F. Schmidt (2006) Distributed and selective auditory representation of song repertoires in the avian song system. J. Neurophysiology 96: 3433-3447 [Nealen and Schmidt (JNP 2006)]

Ashmore R. C., J. M. Wild and M. F. Schmidt (2005) Brainstem and forebrain contributions to the generation of learned motor behaviors for song J. Neuroscience 25: 8543-8554. [Ashmore, Wild and Schmidt (JN 2005)]

Cardin J.A., Raksin J. N. and M.F. Schmidt (2005) The sensorimotor nucleus NIf is necessary for auditory processing but not vocal motor output in the avian song system. J. Neurophysiology 93: 2157-2166. [Cardin, Raksin and Schmidt (JNP 2005)]

Cardin J.A. and M.F. Schmidt (2004) Noradrenergic inputs mediate state dependence of auditory responses in the avian song system. J. Neuroscience 24: 7745-7753. [Cardin_and_Schmidt (JN 2004)]

Cardin, J.A. and M. F. Schmidt (2003) Song system auditory responses are stable and highly tuned during sedation, rapidly modulated and unselective during wakefulness, and suppressed by arousal. J. Neurophysiology 90: 2884-2899. [Cardin_and_Schmidt (JNP 2004)]

Schmidt M. F. (2003) Pattern of interhemispheric synchronous premotor activity in HVc correlates with key transitions in the song pattern. J. Neurophysiology 90: 3931-3949. [Schmidt (JNP 2003)]

Schmidt M. F. and M. Konishi (1998) Gating of auditory responses in the song control system of awake songbirds. Nature Neuroscience 1: 513-518. [Schmidt and Konishi (Nat. Neurosci. 1998)]

  • Society for Neuroscience
  • Animal Behavior Society
  • American Physiological Society



CV (file)