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Nat Commun. 2016 Nov 8;7:13289. doi: 10.1038/ncomms13289.

Pupil fluctuations track rapid changes in adrenergic and cholinergic activity in cortex.

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Department of Neuroscience, Baylor College of Medicine, One Baylor Plaza, Houston, Texas 77030, USA.
Department of Neuroscience, Yale University, 333 Cedar Street, New Haven, Connecticut 06510, USA.
Department of Biomedical Engineering, Columbia University, 1210 Amsterdam Avenue, New York, New York 10027, USA.
Department of Electrical and Computer Engineering, Rice University, 6100 Main St, Houston, Texas 77005, USA.


Rapid variations in cortical state during wakefulness have a strong influence on neural and behavioural responses and are tightly coupled to changes in pupil size across species. However, the physiological processes linking cortical state and pupil variations are largely unknown. Here we demonstrate that these rapid variations, during both quiet waking and locomotion, are highly correlated with fluctuations in the activity of corticopetal noradrenergic and cholinergic projections. Rapid dilations of the pupil are tightly associated with phasic activity in noradrenergic axons, whereas longer-lasting dilations of the pupil, such as during locomotion, are accompanied by sustained activity in cholinergic axons. Thus, the pupil can be used to sensitively track the activity in multiple neuromodulatory transmitter systems as they control the state of the waking brain.

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