Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

How sleep affects the developmental learning of bird song


Sleep affects learning and development in humans and other animals, but the role of sleep in developmental learning has never been examined. Here we show the effects of night-sleep on song development in the zebra finch by recording and analysing the entire song ontogeny. During periods of rapid learning we observed a pronounced deterioration in song structure after night-sleep. The song regained structure after intense morning singing. Daily improvement in similarity to the tutored song occurred during the late phase of this morning recovery; little further improvement occurred thereafter. Furthermore, birds that showed stronger post-sleep deterioration during development achieved a better final imitation. The effect diminished with age. Our experiments showed that these oscillations were not a result of sleep inertia or lack of practice, indicating the possible involvement of an active process, perhaps neural song-replay during sleep. We suggest that these oscillations correspond to competing demands of plasticity and consolidation during learning, creating repeated opportunities to reshape previously learned motor skills.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Tracing vocal changes.
Figure 2: Vocal changes during night-sleep.
Figure 3: Recovery of syllable structure during the morning.
Figure 4: Progression of song learning.
Figure 5: Vocal changes after song prevention and sleep manipulation.
Figure 6: Comparison of post-sleep deterioration and post-deafening deterioration.


  1. Karni, A., Tanne, D., Rubenstein, B. S., Askenasy, J. J. & Sagi, D. Dependence on REM sleep of overnight improvement of a perceptual skill. Science 265, 679–682 (1994)

    Article  ADS  CAS  Google Scholar 

  2. Wilson, M. A. & Mc Naughton, B. L. Reactivation of hippocampal ensemble memories during sleep. Science 265, 676–679 (1994)

    Article  ADS  CAS  Google Scholar 

  3. Maquet, P. The role of sleep in learning and memory. Science 294, 1048–1052 (2001)

    Article  ADS  CAS  Google Scholar 

  4. Peigneux, P., Laureys, S., Delbeuck, X. & Maquet, P. Sleeping brain, learning brain. The role of sleep for memory systems. Neuroreport 12, 111–124 (2001)

    Article  Google Scholar 

  5. Stickgold, R., Hobson, J. A., Fosse, R. & Fosse, M. Sleep, learning, and dreams: off-line memory reprocessing. Science 294, 1052–1057 (2001)

    Article  ADS  CAS  Google Scholar 

  6. Fenn, K. M., Nusbaum, H. C. & Margoliash, D. Consolidation during sleep of perceptual learning of spoken language. Nature 425, 614–616 (2003)

    Article  ADS  CAS  Google Scholar 

  7. Walker, M. P., Brakefield, T., Hobson, A. & Stickgold, R. Dissociable stages of human memory consolidation and reconsolidation. Nature 425, 616–620 (2003)

    Article  ADS  CAS  Google Scholar 

  8. Wagner, U., Gals, S.,, Halder, H., Verleger, R. & Born, J. Sleep inspires insight. Nature 427, 352–355 (2004)

    Article  ADS  CAS  Google Scholar 

  9. Walker, M. P. & Stickgold, R. Sleep-dependent learning and memory consolidation. Neuron 44, 121–133 (2004)

    Article  CAS  Google Scholar 

  10. Huber, R., Ghilardi, M. F., Massimini, M. & Tononi, G. Local sleep and learning. Nature 430, 78–81 (2004)

    Article  ADS  CAS  Google Scholar 

  11. Peigneux, P. et al. Are spatial memories strengthened in the human hippocampus during slow wave sleep? Neuron 44, 535–545 (2004)

    Article  CAS  Google Scholar 

  12. Frank, M. G., Issa, N. P. & Stryker, M. P. Sleep enhances plasticity in the developing visual cortex. Neuron 30, 275–287 (2001)

    Article  CAS  Google Scholar 

  13. Zuckerman, B., Stevenson, J. & Bailey, V. Sleep problems in early childhood: continuities, predictive factors, and behavioral correlates. Pediatrics 80, 664–671 (1987)

    CAS  PubMed  Google Scholar 

  14. Doupe, A. S. & Kuhl, P. K. Birdsong and human speech: common themes and mechanisms. Annu. Rev. Neurosci. 22, 567–631 (1999)

    Article  CAS  Google Scholar 

  15. Brainard, M. S. & Doupe, A. J. What songbirds teach us about learning. Nature 417, 351–358 (2002)

    Article  ADS  CAS  Google Scholar 

  16. Immelmann, K. in Bird Vocalizations (ed. Hinde, R. A.) 61–74 (Cambridge Univ. Press, Cambridge, 1969)

    Google Scholar 

  17. Tchernichovski, O., Mitra, P. P., Lints, T. & Nottebohm, F. Dynamics of the vocal imitation process: how a zebra finch learns its song. Science 291, 2564–2569 (2001)

    Article  ADS  CAS  Google Scholar 

  18. Tchernichovski, O. & Mitra, P. P. Towards quantification of vocal imitation in the zebra finch. J. Comp. Physiol. [A] 188, 867–878 (2002)

    Article  CAS  Google Scholar 

  19. Konishi, M. The role of auditory feedback in the control of vocalization in the white-crowned sparrow. Z. Tierpsychol. 22, 770–783 (1965)

    CAS  PubMed  Google Scholar 

  20. Doupe, A. J. Song- and order-selective neurons in the songbird anterior forebrain and their emergence during vocal development. J. Neurosci. 17, 1147–1167 (1997)

    Article  CAS  Google Scholar 

  21. Bottjer, S. W. & Johnson, F. Circuits, hormones, and learning: vocal behavior in songbirds. J. Neurobiol. 33, 602–618 (1997)

    Article  CAS  Google Scholar 

  22. Schmidt, M. F. & Konishi, M. Gating of auditory responses in the vocal control system of awake songbirds. Nature Neurosci. 1, 513–518 (2001)

    Article  Google Scholar 

  23. Nick, T. A. & Konishi, M. Dynamic control of auditory activity during sleep: correlation between song response and EEG. Proc. Natl Acad. Sci. USA 98, 14012–14016 (2001)

    Article  ADS  CAS  Google Scholar 

  24. Rauske, P. L., Shea, S. D. & Margoliash, D. State and neuronal class-dependent reconfiguration in the avian song system. J. Neurophysiol. 89, 1688–1701 (2003)

    Article  Google Scholar 

  25. Nick, T. A. & Konishi, M. Neural song preference during vocal learning in the zebra finch depends on age and state. J. Neurobiol. 62, 231–242 (2005)

    Article  Google Scholar 

  26. Dave, A. S. & Margoliash, D. Song replay during sleep and computational rules for sensorimotor vocal learning. Science 290, 812–816 (2000)

    Article  ADS  CAS  Google Scholar 

  27. Hahnloser, R. H., Kozhevnikov, A. A. & Fee, M. S. An ultra-sparse code underlies the generation of neural sequences in a songbird. Nature 419, 65–70 (2002)

    Article  ADS  CAS  Google Scholar 

  28. Margoliash, D. Evaluating theories of bird song learning: implications for future directions. J. Comp. Physiol. A 188, 851–866 (2002)

    Article  CAS  Google Scholar 

  29. Tchernichovski, O., Nottebohm, F., Ho, C. E., Pesaran, B. & Mitra, P. P. A procedure for an automated measurement of song similarity. Anim. Behav. 59, 1167–1176 (2000)

    Article  CAS  Google Scholar 

  30. Tchernichovski, O., Lints, T., Derégnaucourt, S., Cimenser, A. & Mitra, P. P. Studying the song development process: rationale and methods. Ann. NY Acad. Sci. 1016, 348–363 (2004)

    Article  ADS  CAS  Google Scholar 

  31. Tchernichovski, O. & Mitra, P. P. Sound analysis Pro user manual. (2004).

  32. Lombardino, A. J. & Nottebohm, F. Age at deafening affects the stability of learned song in adult male zebra finches. J. Neurosci. 20, 5054–5064 (2000)

    Article  CAS  Google Scholar 

  33. Nordeen, K. W. & Nordeen, E. J. Auditory feedback is necessary for the maintenance of stereotyped song in adult zebra finches. Behav. Neural Biol. 57, 58–66 (1992)

    Article  CAS  Google Scholar 

  34. Tassi, P. & Muzet, A. Sleep inertia. Sleep Med. Rev. 4, 341–353 (2000)

    Article  Google Scholar 

  35. Jarvis, E. D. & Nottebohm, F. Motor-driven gene expression. Proc. Natl Acad. Sci. USA 94, 4097–4102 (1997)

    Article  ADS  CAS  Google Scholar 

  36. Nordeen, K. W. & Nordeen, E. J. Projection neurons within a vocal motor pathway are born during song learning in zebra finches. Nature 334, 149–151 (1988)

    Article  ADS  CAS  Google Scholar 

  37. Wang, N., Hurley, P., Pytte, C. & Kirn, J. R. Vocal control neuron incorporation decreases with age in the adult zebra finch. J. Neurosci. 22, 10864–10870 (2002)

    Article  CAS  Google Scholar 

  38. Brainard, M. S. & Doupe, A. J. Postlearning consolidation of birdsong: stabilizing effects of age and anterior forebrain lesions. J. Neurosci. 21, 2501–2517 (2001)

    Article  CAS  Google Scholar 

  39. Leonardo, A. & Konishi, M. Decrystallization of adult birdsong by perturbation of auditory feedback. Nature 399, 466–470 (1999)

    Article  ADS  CAS  Google Scholar 

  40. Keasar, M., Motro, U., Shur, Y. & Schmida, A. Overnight memory retention of foraging skills by bumblebees is imperfect. Anim. Behav. 52, 95–104 (1996)

    Article  Google Scholar 

  41. Golani, I. A mobility gradient in the organization of vertebrate movement: the perception of movement through symbolic language. Behav. Brain Sci. 15, 249–308 (1992)

    Article  Google Scholar 

  42. Tchernichovski, O., Benjamini, Y. & Golani, I. The dynamics of long-term exploration in the rat. Part I. A phase-plane analysis of the relationship between location and velocity. Biol. Cybern. 78, 423–432 (1998)

    Article  CAS  Google Scholar 

  43. Marinari, E. & Parisi, G. Simulated tempering: a new Monte Carlo scheme. Europhys. Lett. 19, 451–458 (1992)

    Article  ADS  CAS  Google Scholar 

  44. Kirkpatrick, S., Gelatt, C. D. & Vecchi, M. P. Optimization by simulated annealing. Science 220, 671–680 (1983)

    Article  ADS  MathSciNet  CAS  Google Scholar 

  45. Ho, C. E., Pesaran, B., Fee, M. S. & Mitra, P. P. in Proc. Joint Symposium on Neural Computation Vol. 5, 76–83 (Univ. California Press, San Diego, 1998)

    Google Scholar 

  46. Loader, C. Local Regression and Likelihood (Springer, New York, 1999)

    MATH  Google Scholar 

Download references


We thank D. Swigger for programming; J. Wallman for comments; P. Andrews for help with graphics; and T. Lints, K. Maul, A. Alexander and L. Dayani for help in analysing data. This work was supported by grants from the NIH to O.T. and P.P.M., by an NIH RCMI (National Institutes of Health Research Centers in Minority Institutions) grant to the City College of New York, by internal support from the Cold Spring Harbor Laboratory and the Robertson foundation to P.P.M., and from the Fondation Fyssen to S.D.

Author information

Authors and Affiliations


Corresponding author

Correspondence to Sébastien Derégnaucourt.

Ethics declarations

Competing interests

The authors declare that they have no competing financial interests.

Supplementary information

Supplementary Methods

1. Source separation in the case of live tutoring; 2. Observations on sleep; 3. Segmentation into syllables; 4. Computing syllable features; 5. Smoothing of duration histograms; 6. Clustering syllables and tracing the evolution of clusters; 7. Estimating vocal changes across timescales; 8. Similarity measurements. (PDF 97 kb)

Supplementary Data

1. Analysis of vocal changes triggered in adult birds; 2. Observations on sleep; 3. Analysis of sleep effect on syllable duration; 4. Comparison between birds trained from day 43 versus day 60; 5. Analysis of the causality chain between post-sleep deterioration and eventual similarity to the song model. (PDF 73 kb)

Supplementary Table 1

Relative contribution of syllable features to post-sleep deterioration. (PDF 77 kb)

Supplementary Figure 1

Developmental change in the structure of a syllable produced by a bird trained from day 43 (a) and a bird trained from day 90 (b) as captured by Wiener Entropy variance (EV). (PDF 37 kb)

Supplementary Figure 2

Cross-correlation analysis between post-sleep deterioration curves of birds trained from day 43 (n=12) and birds trained from day 60 (n=6). (PDF 58 kb)

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Derégnaucourt, S., Mitra, P., Fehér, O. et al. How sleep affects the developmental learning of bird song. Nature 433, 710–716 (2005).

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI:

This article is cited by


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing