An Associational Model of Birdsong Sensorimotor Learning I. Efference Copy and the Learning of Song Syllables
Abstract
Birdsong learning provides an ideal model system for studying temporally complex motor behavior. Guided by the well-characterized functional anatomy of the song system, we have constructed a computational model of the sensorimotor phase of song learning. Our model uses simple Hebbian and reinforcement learning rules and demonstrates the plausibility of a detailed set of hypotheses concerning sensory-motor interactions during song learning. The model focuses on the motor nuclei HVc and robust nucleus of the archistriatum (RA) of zebra finches and incorporates the long-standing hypothesis that a series of song nuclei, the Anterior Forebrain Pathway (AFP), plays an important role in comparing the bird's own vocalizations with a previously memorized song, or “template.” This “AFP comparison hypothesis” is challenged by the significant delay that would be experienced by presumptive auditory feedback signals processed in the AFP. We propose that the AFP does not directly evaluate auditory feedback, but instead, receives an internally generated prediction of the feedback signal corresponding to each vocal gesture, or song “syllable.” This prediction, or “efference copy,” is learned in HVc by associating premotor activity in RA-projecting HVc neurons with the resulting auditory feedback registered within AFP-projecting HVc neurons. We also demonstrate how negative feedback “adaptation” can be used to separate sensory and motor signals within HVc. The model predicts that motor signals recorded in the AFP during singing carry sensory information and that the primary role for auditory feedback during song learning is to maintain an accurate efference copy. The simplicity of the model suggests that associational efference copy learning may be a common strategy for overcoming feedback delay during sensorimotor learning.
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The feedback loops execute a delay function by increasing the distance over which corollary discharge issued from vocal centres travels before it collides with auditory feedback.
“The feedback loops execute a delay function by increasing the distance over which CD that is issued from vocal centres travels before it collides with auditory feedback72,73.”
From Corollary discharge across the animal kingdom (2008) · cites this paper as [72]
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- Authors
- Troyer, T. W., & Doupe, A. J.
- Affiliations
- Department of Psychiatry,
- University of California, San Francisco (US)
- ORCID
- Troyer, Todd W.
- Venue
- Journal of Neurophysiology
- Publisher
- American Physiological Society
- Published
- 2000-09-01
- Volume
- 84
- Issue
- 3
- Pages
- 1204-1223
- ISSN
- 0022-3077 (ISSN-L)
- 1522-1598
- OpenAlex
- W99195914
- Cited by
- 145
- References
- 99
- Citations by year
- 2026: 3 · 2024: 2 · 2023: 3 · 2022: 3 · 2021: 1 · 2020: 7 · 2019: 5 · 2018: 1 · 2017: 8 · 2016: 2 · 2015: 6 · 2014: 8 · 2013: 8 · 2012: 1
- FWCI
- 5.04
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Journal of Neurophysiology (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (5f32mh012372-02)
- National Institute of Mental Health (MH55987, MH12372, R01MH055987)
- National Institute of Neurological Disorders and Stroke (NS 34835, P01NS034835)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neuroscience and Music Perception
- Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
- Keywords
- efference copy, sensorimotor learning, zebra finch, HVC, robust nucleus of the archistriatum, anterior forebrain pathway, auditory feedback, feedback delay, Hebbian learning, reinforcement learning, sensorimotor interaction, vocal motor control
- MeSH terms
- Algorithms, Animals, Association Learning, Computer Simulation, Feedback, Models, Neurological, Neural Pathways, Synaptic Transmission, Neuronal Plasticity, Psychomotor Performance, Reaction Time, Reinforcement, Psychology, Sound, Vocalization, Animal, Prosencephalon, Songbirds
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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