Abstract

Studies of learning mechanisms critically depend on the ability to accurately assess learning outcomes. This assessment can be impeded by the often complex, multidimensional nature of behavior. We present a novel, automated approach to evaluating imitative learning. Conceptually, our approach estimates how much of the content present in a reference behavior is absent from the learned behavior. We validate our approach through examination of songbird vocalizations, complex learned behaviors the study of which has provided many insights into sensory-motor learning in general and vocal learning in particular. Historically, learning has been holistically assessed by human inspection or through comparison of specific song features selected by experimenters (e.g. fundamental frequency, spectral entropy). In contrast, our approach uses statistical models to broadly capture the structure of each song, and then estimates the divergence between the two models. We show that our measure of song learning (the Kullback-Leibler divergence between two distributions corresponding to specific song data, or, Song DKL) is well correlated with human evaluation of song learning. We then expand the analysis beyond learning and show that Song DKL also detects the typical song deterioration that occurs following deafening. Finally, we illustrate how this measure can be extended to quantify differences in other complex behaviors such as human speech and handwriting. This approach potentially provides a framework for assessing learning across a broad range of behaviors like song that can be described as a set of discrete and repeated motor actions.

Highlights

  • Songbird vocal learning shares many parallels with speech learning and is a powerful and tractable model system for elucidating neural and behavioral mechanisms underlying vocal control and vocal learning [1,2]

  • We estimate the amount of content present in a reference behavior that is absent in the resultant learned behavior. We show that this measure provides a holistic and automated assessment of vocal learning in Estrildid finches that is consistent with human assessment

  • We show that the DKL provides a measure of the quality of song learning in Bengalese finches that is well correlated with scores provided by human experts

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Summary

Introduction

Songbird vocal learning shares many parallels with speech learning and is a powerful and tractable model system for elucidating neural and behavioral mechanisms underlying vocal control and vocal learning [1,2]. Like humans, learn vocalizations early in life through exposure to the vocalizations of an adult ‘tutor’ followed by a period of practice that eventually results in typical adult vocalizations that require auditory feedback for maintenance [1]. In the finch species examined here, a given bird’s song comprises about 5–10 categorically distinct syllable types, with these distinct types defined by their unique spectro-temporal structure (Fig 1A). An individual bird’s song can be described as a specific set of categorically distinct syllable types (that can be labeled ‘A’, ‘B’, ‘C’ and so on). Juvenile ‘tutees’ could learn to produce all distinct syllable types present in an adult ‘tutor’ song, but the spectral content of the syllables might be imperfect or noisy (Fig 1Ci-1Cii), while other tutees might completely fail to learn some syllables (Fig 1Ciii), and still others might improvise new syllables (Fig 1Civ)

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