“…Light level is the primary determinant of pupil size, although non-light-driven pupil fluctuations, widely assumed as an indicator of arousal through locus coeruleus activity, can be used to index brain state across species ( McGinley et al, 2015 ; Lee and Margolis, 2016 ; Reimer et al, 2016 ). Higher cognitive and emotional processes are also able to evoke tonic or phasic pupillary changes, such as attention ( Binda et al, 2013a ), memory load ( Wierda et al, 2012 ), novelty ( Angulo-Chavira et al, 2017 ; Krebs et al, 2018 ; Montes-Lourido et al, 2021 ), pain ( Connelly et al, 2014 ; Azevedo-Santos and DeSantana, 2018 ; Charier et al, 2019 ), and more general cortical sensory processing ( Binda et al, 2013b ; Lee and Margolis, 2016 ) in humans and in animal models.…”