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The role of nonlinear dynamics of the syrinx in the vocalizations of a songbird


Birdsong is characterized by the modulation of sound properties over a wide range of timescales1. Understanding the mechanisms by which the brain organizes this complex temporal behaviour is a central motivation in the study of the song control and learning system2,3,4,5,6,7,8. Here we present evidence that, in addition to central neural control, a further level of temporal organization is provided by nonlinear oscillatory dynamics that are intrinsic to the avian vocal organ. A detailed temporal and spectral examination of song of the zebra finch (Taeniopygia guttata) reveals a class of rapid song modulations that are consistent with transitions in the dynamical state of the syrinx. Furthermore, in vitro experiments show that the syrinx can produce a sequence of oscillatory states that are both spectrally and temporally complex in response to the slow variation of respiratory or syringeal parameters. As a consequence, simple variations in a small number of neural signals can result in a complex acoustic sequence.

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Figure 1: Spectral and temporal characteristics of zebra-finch song.
Figure 2: Patterns of acoustic signals generated in the in vitro syrinx preparation during slow modulation of syringeal and flow parameters.
Figure 3: Stroboscopic observation of in vitro syringeal oscillations.
Figure 4: Results of a simple numerical model of airflow through a compliant constriction.


  1. Greenewalt, C. H. Brid Song: Acoustics and Physiology (Smithsonian Institution Press, Washington DC, (1968)).

    Google Scholar 

  2. Nottebohm, F., Stokes, T. M. & Leonard, C. M. Central control of song in the canary, Serinus canarius. J. Comp. Neurol. 165, 457–486 (1976).

    CAS  Article  Google Scholar 

  3. Nottebohm, F., Kelley, D. B. & Paton, J. A. Connections of vocal control nuclei in the canary telencephalon. J. Comp. Neurol. 207(1982).

  4. McCasland, J. S. Neuronal control of bird song production. J. Neurosci. 7, 23–39 (1987).

    CAS  Article  Google Scholar 

  5. Scharff, C. & Nottebohm, F. Acomparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning. J. Neurosci. 11, 2896–2913 (1991).

    CAS  Article  Google Scholar 

  6. Konishi, M. Pattern generation in birdsong. Curr. Opin. Neurobiol. 4, 827–831 (1994).

    CAS  Article  Google Scholar 

  7. Vu, E., Mazurek, M. E. & Kuo, Y.-C. Identification of a forebrain motor programming network for the learned song of zebra finches. J. Neurosci. 14, 6924–6934 (1994).

    CAS  Article  Google Scholar 

  8. Yu, A. & Margoliash, D. Temporal hierarchical control of singing in birds. Science 273, 1871–1875 (1996).

    ADS  CAS  Article  Google Scholar 

  9. Immelman, K. in Bird Vocalizations (ed. Hinde, R. A.) pp. 61–74 (Cambridge Univ. Press, (1969)).

    Google Scholar 

  10. Price, P. H. Developmental determinants of structure in zebra finch song. J. Comp. Physiol. Psychol. 93, 260–277 (1979).

    Article  Google Scholar 

  11. Fletcher, N. H. Mode locking in non-linearly excited inharmonic musical oscillators. J. Acoust. Soc. Am. 64, 1566–1569 (1978).

    ADS  Article  Google Scholar 

  12. Jackson, E. A. Perspectives of Nonlinear Dynamics (Cambridge Univ. Press, (1992)).

    Google Scholar 

  13. Hartley, R. S. & Suthers, R. A. Airflow and pressure during canary song: direct evidence for mini-breaths. J. Comp. Phys. A 165, 15–26 (1989).

    Article  Google Scholar 

  14. Goller, F. & Suthers, R. A. Implications for lateralization of bird song from unilateral gating of bilateral motor patterns. Nature 373, 63–66 (1995).

    ADS  CAS  Article  Google Scholar 

  15. Goller, F. & Larsen, O. N. Anew mechanism of sound generation in songbirds. Proc. Natl Acad. Sci. USA 94, 14787–14791 (1997).

    ADS  CAS  Article  Google Scholar 

  16. Fletcher, N. H. Bird song — a quantitative acoustic model. J. Theor. Biol. 135, 455–481 (1988).

    MathSciNet  Article  Google Scholar 

  17. Ishizaka, K. & Flanagan, J. L. Synthesis of voiced sounds from a two-mass model of the vocal cords. Bell Syst. Tech. J. 51, 1233–1268 (1972).

    Article  Google Scholar 

  18. Bertram, C. D. & Pedley, T. J. Amathematical model of unsteady collapsible tube behaviour. J. Biomech. 15, 39–50 (1982).

    CAS  Article  Google Scholar 

  19. Berry, D. A., Herzel, H., Titze, I. R. & Krischer, K. Interpretation of biomechanical simulations of normal and chaotic vocal fold oscillations with empirical eigenfunctions. J. Acoust. Soc. Am. 95, 3595–3604 (1994).

    ADS  CAS  Article  Google Scholar 

  20. Herzel, H., Berry, D. A., Titze, I. R. & Saleh, M. Analysis of vocal disorders with methods from nonlinear dynamics. J. Speech Hear. Res. 37, 1008–1019 (1994).

    CAS  Article  Google Scholar 

  21. Thompson, D. J. Spectrum estimation and harmonic analysis. Proc. IEEE 70, 1055–1096 (1982).

    ADS  Article  Google Scholar 

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We thank W. Denk, M. Konishi and S. S.-H. Wang for comments on the manuscript.

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Correspondence to Michale S. Fee.

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Fee, M., Shraiman, B., Pesaran, B. et al. The role of nonlinear dynamics of the syrinx in the vocalizations of a songbird. Nature 395, 67–71 (1998).

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