Auditory feedback in learning and maintenance of vocal behaviour

ABSTRACT Songbirds are one of the best-studied examples of vocal learners. Learning of both human speech and birdsong depends on hearing. Once learned, adult song in many species remains

unchanging, suggesting a reduced influence of sensory experience. Recent studies have revealed, however, that adult song is not always stable, extending our understanding of the mechanisms

involved in song maintenance, and their similarity to those active during song learning. Here we review some of the processes that contribute to song learning and production, with an

emphasis on the role of auditory feedback. We then consider some of the possible neural substrates involved in these processes, particularly basal ganglia circuitry. Although a thorough

treatment of human speech is beyond the scope of this article, we point out similarities between speech and song learning, and ways in which studies of these disparate behaviours complement

each other in developing an understanding of general principles that contribute to learning and maintenance of vocal behaviour. KEY POINTS * Behavioural observations indicate that similar

processes contribute to the learning and maintenance of birdsong and speech. These observations suggest that during learning, both birds and humans form internal representations of

species-specific vocalizations. They then use auditory feedback to match their developing vocal output to these internal sensory models. Once learned, vocalizations tend to remain stable.

However, alterations of auditory feedback in adulthood lead to a deterioration of both birdsong and speech, indicating that they have not become ‘hard-wired’, that is, resistant to the

influence of experience. * The mechanisms responsible for the influence of auditory feedback on the maintenance of vocalizations are unknown. It is possible that the observed deterioration

in the absence of feedback is due to passive drift in vocal control structures. Alternatively, the lack of auditory feedback could cause the mechanisms that match the feedback to an internal

model to generate an error signal that actively modifies the vocalizations. * Lesion studies in combination with manipulations of auditory feedback are consistent with the idea that the

deterioration of adult song is indeed active. These experiments also raise the possibility that a basal ganglia circuit of songbirds, the anterior forebrain pathway, participates in the

evaluation of song, and in the generation of an error signal when the bird does not receive feedback that matches its internal model. For example, the destruction of the lateral

magnocellular nucleus of the anterior neostriatum (LMAN) — the output nucleus of the anterior forebrain pathway — prevents the song deterioration that normally results from the absence of

feedback. * How the songbird brain evaluates song and matches it to the internal model is not known. Song-selective neurons — cells found throughout the songbird brain that fire more

strongly to the bird's own song than to the songs of other birds of the same species — could be involved in this process. Access through your institution Buy or subscribe This is a

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BEHAVIOUR IS LEARNED THROUGH DOPAMINE-MEDIATED REINFORCEMENT Article 12 March 2025 A MEMORY-DRIVEN AUDITORY PROGRAM ENSURES SELECTIVE AND PRECISE VOCAL IMITATION IN ZEBRA FINCHES Article

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Download references ACKNOWLEDGEMENTS We thank S. Lisberger for helpful comments on the manuscript. The work of the authors was supported by the National Institutes of Health, a Burroughs

Wellcome Fund Fellowship of the Life Sciences Research Foundation, the John Merck Fund and the EJLB foundation. AUTHOR INFORMATION AUTHORS AND AFFILIATIONS * Departments of Physiology and

Psychiatry, Keck Center for Integrative Neuroscience, University of California San Francisco, San Francisco, 94143 0444, California, USA Michael S. Brainard & Allison J. Doupe Authors *

Michael S. Brainard View author publications You can also search for this author inPubMed Google Scholar * Allison J. Doupe View author publications You can also search for this author

inPubMed Google Scholar SUPPLEMENTARY INFORMATION IN ADULT ZEBRA FINCHES, DEAFENING PRODUCES A GRADUAL DETERIORATION OF LEARNED SONG. LESIONS OF THE LATERAL MAGNOCELLULAR NUCLEUS OF THE

ANTERIOR NEOSTRIATUM (LMAN) PREVENT THE DEAFENING-INDUCED DETERIORATION. Sound file A, the normal song of an adult zebra finch. (MPG 191 kb) SOUND FILE B, DETERIORATED SONG OF THE SAME BIRD

FOLLOWING DEAFENING. (MPG 246 KB) SOUND FILE C, THE NORMAL SONG OF A DIFFERENT ZEBRA FINCH. (MPG 164 KB) SOUND FILE D, THE SONG OF THIS BIRD DID NOT CHANGE APPRECIABLY AFTER DEAFENING AND

LESION OF LMAN. (MPG 145 KB) RELATED LINKS RELATED LINKS FURTHER INFORMATION Allison Doupe's laboratory ENCYLOPEDIA OF LIFE SCIENCES Bird song: steroid hormones and plasticity GLOSSARY

* PHONEME A distinct unit of sound that distinguishes one word from another. * BASAL GANGLIA A collection of brain structures that modulate cortical output. * WERNICKE'S AREA Region of

the parietal cortex involved in speech processing. RIGHTS AND PERMISSIONS Reprints and permissions ABOUT THIS ARTICLE CITE THIS ARTICLE Brainard, M., Doupe, A. Auditory feedback in learning

and maintenance of vocal behaviour . _Nat Rev Neurosci_ 1, 31–40 (2000). https://doi.org/10.1038/35036205 Download citation * Issue Date: 01 October 2000 * DOI:

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