The neural basis of stream segregation in the primary auditory cortex.
Christophe Micheyl (1), Biao Tian (2), Robert P. Carlyon (3), and Joseph P. Rauschecker (2)
1- UMR CNRS 5020, Lyon, France
2- Georgetown University, Washington DC, USA
3- MRC - Cognition & Brain Sciences Unit, Cambridge, UK
This study was funded in part by an Engineering and Physical Sciences Research Council Life Sciences Network grant GR/M90146 to Prof. C.J. Darwin, Exptl. Psychol., University of Sussex.
The sounds reaching our ears often contain components arising from multiple acoustic sources. The task of our brain is to organize this complex raw sensory material into meaningful perceptual entities. One of the fundamental perceptual processes of auditory scene analysis is known as "stream segregation". Auditory stream segregation refers to the perceptual teasing apart, and linking over time of frequency components in sound sequences, which results in the formation of auditory "streams". In everyday life, a stream may be the voice of a speaker in a cocktail-party background, or the sound of a violin among other instruments in an orchestral symphony. In the laboratory, a now classic demonstration of the stream segregation phenomenon involves the presentation of tones alternating in frequency in an ABAB pattern: when the tempo is slow and/or the frequency difference between the A and B tones is small, listeners report hearing a single stream of tones alternating in frequency; when the tempo is fast and/or the frequency difference is large, listeners report hearing two distinct, monotonic streams. Although stream segregation has been largely studied over the past fifty years, the brain mechanisms behind it still remain unclear to date. In the present study, we investigated further the hypothesis suggested by earlier results from Fishman et al. (2000), that perceptual auditory stream segregation is reflected in the primary auditory cortex by the differential suppression of neural responses to off-BF (best frequency) tones by preceding on-BF tones. We recorded the responses of single units in the primary auditory cortex (A1) of an awake monkey to 10-sec ABA_ sequences of tones - where A and B represent 125-ms tone pips of a different frequency, and _ stands for a 125-ms silent gap. The A-tone frequency was always close to the unit's BF, whilst the B-tone frequency was set to 1, 3, 6, or 9 semitones (STs) above A. The results revealed that for A-B separations of 3, 6, and 9 ST, but not 1 ST, the amplitude of the neural response to the B-tones decreased over part or all of the sequence duration. Conversely, a trend for A tone responses to increase over time was observed. As a result, the ratio of B-tone to A-tone responses increased over time. These electrophysiological results are consistent with psychoacoustical data in humans, which indicate that with the type of sequences used here, auditory stream segregation "builds up" over roughly the first 5-10 secs of stimulation. They reveal that the differential suppression of responses to off-BF tones by preceding on-BF tones, which Fishman et al. (2000) have proposed to be the neural basis of stream segregation at the cortical level, is a dynamic phenomenon. The decrease in amplitude of the B-tone responses was not observed when these tones were presented alone - i.e. with the surrounding A tones removed -, suggesting that the effect was not simply due to long-term adaptation of off-BF responses.