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Behav Brain Res. 2016 Jan 1;296:211-232. doi: 10.1016/j.bbr.2015.09.011. Epub 2015 Sep 11.

Electrophysiological CNS-processes related to associative learning in humans.

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Dept. of Food Science, Univ. of Copenhagen, Rolighedsvej 30, DK-1958 Frederiksberg, Denmark. Electronic address:
Dept. of Psychological Sciences, Univ. of Missouri, Columbia, USA.


The neurophysiology of human associative memory has been studied with electroencephalographic techniques since the 1930s. This research has revealed that different types of electrophysiological processes in the human brain can be modified by conditioning: sensory evoked potentials, sensory induced gamma-band activity, periods of frequency-specific waves (alpha and beta waves, the sensorimotor rhythm and the mu-rhythm) and slow cortical potentials. Conditioning of these processes has been studied in experiments that either use operant conditioning or repeated contingent pairings of conditioned and unconditioned stimuli (classical conditioning). In operant conditioning, the appearance of a specific brain process is paired with an external stimulus (neurofeedback) and the feedback enables subjects to obtain varying degrees of control of the CNS-process. Such acquired self-regulation of brain activity has found practical uses for instance in the amelioration of epileptic seizures, Autism Spectrum Disorders (ASD) and Attention Deficit Hyperactivity Disorder (ADHD). It has also provided communicative means of assistance for tetraplegic patients through the use of brain computer interfaces. Both extra and intracortically recorded signals have been coupled with contingent external feedback. It is the aim for this review to summarize essential results on all types of electromagnetic brain processes that have been modified by classical or operant conditioning. The results are organized according to type of conditioned EEG-process, type of conditioning, and sensory modalities of the conditioning stimuli.


Alpha waves; Brain–computer interface; Conditioning; EEG; Gamma waves; Human; Mu-rhythm; Sensorimotor rhythm; Sensory evoked potentials

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