The balanced and introspective brain

Transfers of large-scale neural activity into, within and between corticothalamic neural populations and brain hemispheres are analysed using time-integrated transfer functions and state parameters obtained from neural field theory for a variety of arousal states. It is shown that the great majority of activity results from feedbacks within the corticothalamic system, including significant transfer between hemispheres, but only a small minority arises via net input from the external world, with the brain thus in a near-critical, highly introspective state. Notably, the total excitatory and inhibitory influences on cortical neurons are balanced to within a few per cent across arousal states. Strong negative intrahemispheric feedforward exists to the cortex, and even larger interhemispheric positive feedforward, but these are modified by feedback loops to yield near-critical positive overall gain. The results underline the utility of transfer functions for the analysis of brain activity.