Abstract

Bird songs range in form from the simple notes of a Chipping Sparrow to the rich performance of the nightingale. Non-adjacent correlations can be found in the syntax of some birdsongs, indicating that the choice of what to sing next is determined not only by the current syllable, but also by previous syllables sung. Here we examine the song of the domesticated canary, a complex singer whose song consists of syllables, grouped into phrases that are arranged in flexible sequences. Phrases are defined by a fundamental time-scale that is independent of the underlying syllable duration. We show that the ordering of phrases is governed by long-range rules: the choice of what phrase to sing next in a given context depends on the history of the song, and for some syllables, highly specific rules produce correlations in song over timescales of up to ten seconds. The neural basis of these long-range correlations may provide insight into how complex behaviors are assembled from more elementary, stereotyped modules.

Highlights

  • Brains build complex behaviors from simple modules [1,2].The ultimate example is speech where sequences of phonemes form words that in turn are rearranged to form sentences

  • Birdsong has often been described in terms of first-order transition statistics, e.g. between adjacent syllables [13] in the zebra finch or syllable chunks [9,14] in the nightingale and Bengalese finch

  • Analysis of the Bengalese finch song reveals non-adjacent dependencies where transition probabilities between syllables depend on the current active syllable, and one or more prior syllables sung [15,16,17]. This implies that song syntax must be modeled with a second-order or higher order Markov chain

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Summary

Introduction

Brains build complex behaviors from simple modules [1,2].The ultimate example is speech where sequences of phonemes form words that in turn are rearranged to form sentences. The complex performances of a musician or swordfighter are composed of discrete motor gestures that may be composed of more elementary motor modules or muscle synergies [3] Songbirds, in their own ways, build complex vocal forms from elementary units known as syllables. Analysis of the Bengalese finch song reveals non-adjacent dependencies where transition probabilities between syllables depend on the current active syllable, and one or more prior syllables sung [15,16,17]. This implies that song syntax must be modeled with a second-order or higher order Markov chain. Statistically complex sequences will require a large number of hidden states, relative to the number of observed syllables

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