Bilateral Control and Interhemispheric Coordination in the Avian Song Motor System

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Abstract
Birdsong is a complex learned motor behavior controlled by an interconnected network of vocal control nuclei that are present in both cerebral hemispheres. Unilateral lesions of song nuclei in the left or the right hemisphere result in different effects on song structure, suggesting that normal song output results from the activation of two parallel but functionally different motor pathways. Because each syringeal half is innervated primarily by ipsilateral motor structures and activity in both halves is tightly coordinated during singing, motor commands originating from both hemispheres must be tightly coordinated to produce the appropriate vocal output. This coordination occurs despite the absence of direct interhemispheric connections between song control nuclei. In this article, we discuss how motor commands in nucleus HVC, a key forebrain song control region, are coordinated by precisely timed inputs that act to synchronize premotor activity in both hemispheres. Synchronizing inputs are tightly linked to syllable and note onset, which suggests that bilaterally organized circuits in the midbrain or brainstem act in specifying higher-order song features, such as duration, order, and possibly even structure of individual song syllables. The challenge ahead lies in identifying the networks that generate the synchronizing timing inputs and to determine how these inputs specify the motor commands in HVC. Resolving these issues will help us gain a better understanding of how pattern-generating networks in the midbrain/brainstem interface with forebrain circuits to produce complex learned behaviors.