Joel S. Snyder

Effects of Attention on Neuroelectric Correlates of Auditory Stream Segregation

A general assumption underlying auditory scene analysis is that the initial grouping of acoustic elements is independent of attention. The effects of attention on auditory stream segregation were investigated by recording event-related potentials (ERPs) while participants either attended to sound stimuli and indicated whether they heard one or two streams or watched a muted movie. The stimuli were pure-tone ABA-patterns that repeated for 10.8 sec with a stimulus onset asynchrony between A and B tones of 100 msec in which the A tone was fixed at 500 Hz, the B tone could be 500, 625, 750, or 1000 Hz, and was a silence. In both listening conditions, an enhancement of the auditory-evoked response (P1-N1-P2 and N1c) to the B tone varied with f and correlated with perception of streaming. The ERP from 150 to 250 msec after the beginning of the repeating ABA-patterns became more positive during the course of the trial and was diminished when participants ignored the tones, consistent with behavioral studies indicating that streaming takes several seconds to build up. The N1c enhancement and the buildup over time were larger at right than left temporal electrodes, suggesting a right-hemisphere dominance for stream segregation. Sources in Heschls gyrus accounted for the ERP modulations related to f-based segregation and buildup. These findings provide evidence for two cortical mechanisms of streaming: automatic segregation of sounds and attention-dependent buildup process that integrates successive tones within streams over several seconds.

Toward a Neurophysiological Theory of Auditory Stream Segregation.

Auditory stream segregation (or streaming) is a phenomenon in which 2 or more repeating sounds differing in at least 1 acoustic attribute are perceived as 2 or more separate sound sources (i.e., streams). This article selectively reviews psychophysical and computational studies of streaming and comprehensively reviews more recent neurophysiological studies that have provided important insights into the mechanisms of streaming. On the basis of these studies, segregation of sounds is likely to occur beginning in the auditory periphery and continuing at least to primary auditory cortex for simple cues such as pure-tone frequency but at stages as high as secondary auditory cortex for more complex cues such as periodicity pitch. Attention-dependent and perception-dependent processes are likely to take place in primary or secondary auditory cortex and may also involve higher level areas outside of auditory cortex. Topographic maps of acoustic attributes, stimulus-specific suppression, and competition between representations are among the neurophysiological mechanisms that likely contribute to streaming. A framework for future research is proposed.

Pulse and Meter as Neural Resonance

The experience of musical rhythm is a remarkable psychophysical phenomenon, in part because the perception of periodicities, namely pulse and meter, arise from stimuli that are not periodic. One possible function of such a transformation is to enable synchronization between individuals through perception of a common abstract temporal structure (e.g., during music performance). Thus, understanding the brain processes that underlie rhythm perception is fundamental to explaining musical behavior. Here, we propose that neural resonance provides an excellent account of many aspects of human rhythm perception. Our framework is consistent with recent brain‐imaging studies showing neural correlates of rhythm perception in high‐frequency oscillatory activity, and leads to the hypothesis that perception of pulse and meter result from rhythmic bursts of high‐frequency neural activity in response to musical rhythms. High‐frequency bursts of activity may enable communication between neural areas, such as auditory and motor cortices, during rhythm perception and production.

Attention, awareness, and the perception of auditory scenes.

Auditory perception and cognition entails both low-level and high-level processes, which are likely to interact with each other to create our rich conscious experience of soundscapes. Recent research that we review has revealed numerous influences of high-level factors, such as attention, intention, and prior experience, on conscious auditory perception. And recently, studies have shown that auditory scene analysis tasks can exhibit multistability in a manner very similar to ambiguous visual stimuli, presenting a unique opportunity to study neural correlates of auditory awareness and the extent to which mechanisms of perception are shared across sensory modalities. Research has also led to a growing number of techniques through which auditory perception can be manipulated and even completely suppressed. Such findings have important consequences for our understanding of the mechanisms of perception and also should allow scientists to precisely distinguish the influences of different higher-level influences.

Effects of context on auditory stream segregation.

The authors examined the effect of preceding context on auditory stream segregation. Low tones (A), high tones (B), and silences (-) were presented in an ABA- pattern. Participants indicated whether they perceived 1 or 2 streams of tones. The A tone frequency was fixed, and the B tone was the same as the A tone or had 1 of 3 higher frequencies. Perception of 2 streams in the current trial increased with greater frequency separation between the A and B tones (Delta f). Larger Delta f in previous trials modified this pattern, causing less streaming in the current trial. This occurred even when listeners were asked to bias their perception toward hearing 1 stream or 2 streams. The effect of previous Delta f was not due to response bias because simply perceiving 2 streams in the previous trial did not cause less streaming in the current trial. Finally, the effect of previous ?f was diminished, though still present, when the silent duration between trials was increased to 5.76 s. The time course of this context effect on streaming implicates the involvement of auditory sensory memory or neural adaptation.

Effects of prior stimulus and prior perception on neural correlates of auditory stream segregation

We examined whether effects of prior experience are mediated by distinct brain processes from those processing current stimulus features. We recorded event-related potentials (ERPs) during an auditory stream segregation task that presented an adaptation sequence with a small, intermediate, or large frequency separation between low and high tones (Deltaf), followed by a test sequence with intermediate Deltaf. Perception of two streams during the test was facilitated by small prior Deltaf and by prior perception of two streams and was accompanied by more positive ERPs. The scalp topography of these perception-related changes in ERPs was different from that observed for ERP modulations due to increasing the current Deltaf. These results reveal complex interactions between stimulus-driven activity and temporal-context-based processes and suggest a complex set of brain areas involved in modulating perception based on current and previous experience.

Memory for sound, with an ear toward hearing in complex auditory scenes

An area of research that has experienced recent growth is the study of memory during perception of simple and complex auditory scenes. These studies have provided important information about how well auditory objects are encoded in memory and how well listeners can notice changes in auditory scenes. These are significant developments because they present an opportunity to better understand how we hear in realistic situations, how higher-level aspects of hearing such as semantics and prior exposure affect perception, and the similarities and differences between auditory perception and perception in other modalities, such as vision and touch. The research also poses exciting challenges for behavioral and neural models of how auditory perception and memory work.

Neural encoding of sound duration persists in older adults.

Speech perception depends strongly on precise encoding of the temporal structure of sound. Although behavioural studies suggest that communication problems experienced by older adults may entail deficits in temporal acuity, much is unknown about the effects of age on the neural mechanisms underlying the encoding of sound duration. In this study, we measured neuromagnetic auditory evoked responses in young, middle-aged and older healthy participants listening to sounds of various durations. The time courses of cortical activity from bilateral sources in superior temporal planes showed specific differences related to the sound offsets indicating the neural representation of onset and offset markers as one dimension of the neural code for sound duration. Model free MEG source analysis identified brain areas specifically responding with an increase in activity to increases in sound duration in the left anterior insula, right inferior frontal, right middle temporal, and right post-central gyri in addition to bilateral supra-temporal gyri. Sound duration-related changes in cortical responses were comparable in all three age groups despite age-related changes in absolute response magnitudes. The results demonstrated that early cortical encoding of the temporal structure of sound presented in silence is little or not affected by normal aging.

Tempo dependence of middle- and long-latency auditory responses: power and phase modulation of the EEG at multiple time-scales.

OBJECTIVE We measured the influences of power and phase modulations of neuroelectric activity on auditory responses to pure-tone patterns with inter-onset intervals typical of music. METHODS Tones were presented to 8 subjects at 10 different tempos from 150 to 3125 ms and with random intervals. We quantified time-frequency (TF) power with respect to a pre-tone-onset baseline and the TF phase coherence across trials. Peak-to-peak event-related potential (ERP) amplitude values for the middle and long-latency auditory responses were obtained for comparison. RESULTS ERP amplitude, size of power modulation, and amount of phase coherence were larger at slower tempos for the long-latency response (LLR) but not for the middle-latency response (MLR). Multiple regression analysis indicated that for MLR and LLR, phase modulation was a better predictor of ERP amplitude than power modulation. CONCLUSIONS Phase modulation is a better predictor of ERP amplitude than power modulation for middle and long-latency auditory responses. SIGNIFICANCE Lack of diminution of the MLR at fast tempos indicates its usefulness for studying early cortical processing of music and speech patterns.

Enhanced sensory processing accompanies successful detection of change for real-world sounds

Change deafness is the inability of listeners to detect changes occurring in their auditory environment. It is a matter of some debate whether change deafness occurs because of a failure of auditory-specific processes or a failure of more general semantic/verbal memory. To address this issue, we measured event-related potentials (ERPs) to pairs of scenes consisting of naturalistic auditory objects while listeners made a same/different judgment for scenes presented before and after an interruption. ERPs to the post-change scene revealed an enhanced early sensory response (N1) and an enhanced late positivity (P3) for detected changes. Change detection performance was better when there was a large acoustic spread among the objects within Scenes 1 and 2, suggesting that the deficits reflected by the ERP components during change deafness are related to successfully segregating the pre- and post-change objects. We also found that a separate sensory response (P2) reflects implicit, unconscious change detection. Overall, the results provide evidence that auditory-specific sensory processing is critical for both explicit and implicit change detection in natural auditory scenes.