Selective attention to sound location or pitch studied with event-related brain potentials and magnetic fields.

Event-related brain potentials (ERPs) and magnetic fields (ERFs) were used to compare brain activity associated with selective attention to sound location or pitch in humans. Sixteen healthy adults participated in the ERP experiment, and 11 adults in the ERF experiment. In different conditions, the participants focused their attention on a designated sound location or pitch, or pictures presented on a screen, in order to detect target sounds or pictures among the attended stimuli. In the Attend Location condition, the location of sounds varied randomly (left or right), while their pitch (high or low) was kept constant. In the Attend Pitch condition, sounds of varying pitch (high or low) were presented at a constant location (left or right). Consistent with previous ERP results, selective attention to either sound feature produced a negative difference (Nd) between ERPs to attended and unattended sounds. In addition, ERPs showed a more posterior scalp distribution for the location-related Nd than for the pitch-related Nd, suggesting partially different generators for these Nds. The ERF source analyses found no source distribution differences between the pitch-related Ndm (the magnetic counterpart of the Nd) and location-related Ndm in the superior temporal cortex (STC), where the main sources of the Ndm effects are thought to be located. Thus, the ERP scalp distribution differences between the location-related and pitch-related Nd effects may have been caused by activity of areas outside the STC, perhaps in the inferior parietal regions.

Orienting and maintenance of spatial attention in audition and vision: multimodal and modality-specific brain activations.

We studied orienting and maintenance of spatial attention in audition and vision. Functional magnetic resonance imaging (fMRI) in nine healthy subjects revealed activations in the same superior and inferior parietal, and posterior prefrontal areas in the auditory and visual orienting tasks when these tasks were compared with the corresponding maintenance tasks. Attention-related activations in the thalamus and cerebellum were observed during the auditory orienting and maintenance tasks and during the visual orienting task. In addition to the supratemporal auditory cortices, auditory orienting, and maintenance produced stronger activity than the respective visual tasks in the inferior parietal and prefrontal cortices, whereas only the occipital visual cortex and the superior parietal cortex showed stronger activity during the visual tasks than during the auditory tasks. Differences between the brain networks involved in auditory and visual spatial attention could be, for example, due to different encoding of auditory and visual spatial information or differences in stimulus-driven (bottom-up triggered) and voluntary (top-down controlled) attention between the auditory and visual modalities, or both.

Orienting and maintenance of spatial attention in audition and vision: an event-related brain potential study.

We examined the effects of orienting and maintenance of attention on performance and event-related brain potentials (ERPs) in audition and vision. Our subjects selectively attended to sounds or pictures in one location (Maintenance of attention) or alternated the focus of their auditory or visual attention between left and right locations (Orienting of attention) in order to detect and press a response button to infrequent targets among the attended stimuli. Reaction times were longer in the Auditory Orienting condition and hit rates were lower and false alarm rates higher in the Visual Orienting condition than in the corresponding Maintenance conditions. Comparison of ERPs to the attended and unattended stimuli in the Auditory and Visual Orienting and Maintenance conditions revealed attention-related modulations of ERPs. In each modality, ERPs to attended stimuli were negatively displaced in relation to unattended stimuli at 100-250 ms from stimulus onset. These negative differences (Nds) showed modality-specific distributions and they were larger over the hemisphere contralateral to the attended sounds and pictures than over the ipsilateral hemisphere. Moreover, the Nd was larger in the Auditory Orienting condition than in the Auditory Maintenance condition, while no such difference was observed in the visual modality. In addition to the Nd, attended visual stimuli elicited a late positive response (LPR) in both Orienting and Maintenance conditions. In contrast to our recent functional magnetic resonance imaging (fMRI) study employing the same experimental paradigm and indicating orienting-related activity in the frontal and parietal cortices, no ERP responses specifically related to orienting were found in either modality.

Distributed cortical networks for focused auditory attention and distraction.

We used behavioral measures and functional magnetic resonance imaging (fMRI) to study the effects of parametrically varied task-irrelevant pitch changes in attended sounds on loudness-discrimination performance and brain activity in cortical surface maps. Ten subjects discriminated tone loudness in sequences that also included infrequent task-irrelevant pitch changes. Consistent with results of previous studies, the task-irrelevant pitch changes impaired performance in the loudness discrimination task. Auditory stimulation, attention-enhanced processing of sounds and motor responding during the loudness discrimination task activated supratemporal (auditory cortex) and inferior parietal areas bilaterally and left-hemisphere (contralateral to the hand used for responding) motor areas. Large pitch changes were associated with right hemisphere supratemporal activations as well as widespread bilateral activations in the frontal lobe and along the intraparietal sulcus. Loudness discrimination and distracting pitch changes activated common areas in the right supratemporal gyrus, left medial frontal cortex, left precentral gyrus, and left inferior parietal cortex.

Human brain activity associated with audiovisual perception and attention.

Coherent perception of objects in our environment often requires perceptual integration of auditory and visual information. Recent behavioral data suggest that audiovisual integration depends on attention. The current study investigated the neural basis of audiovisual integration using 3-Tesla functional magnetic resonance imaging (fMRI) in 12 healthy volunteers during attention to auditory or visual features, or audiovisual feature combinations of abstract stimuli (simultaneous harmonic sounds and colored circles). Audiovisual attention was found to modulate activity in the same frontal, temporal, parietal and occipital cortical regions as auditory and visual attention. In addition, attention to audiovisual feature combinations produced stronger activity in the superior temporal cortices than attention to only auditory or visual features. These modality-specific areas might be involved in attention-dependent perceptual binding of synchronous auditory and visual events into coherent audiovisual objects. Furthermore, the modality-specific temporal auditory and occipital visual cortical areas showed attention-related modulations during both auditory and visual attention tasks. This result supports the proposal that attention to stimuli in one modality can spread to encompass synchronously presented stimuli in another modality.

Selective attention to sound location or pitch studied with fMRI.

We used 3-T functional magnetic resonance imaging to compare the brain mechanisms underlying selective attention to sound location and pitch. In different tasks, the subjects (N = 10) attended to a designated sound location or pitch or to pictures presented on the screen. In the Attend Location conditions, the sound location varied randomly (left or right), while the pitch was kept constant (high or low). In the Attend Pitch conditions, sounds of randomly varying pitch (high or low) were presented at a constant location (left or right). Both attention to location and attention to pitch produced enhanced activity (in comparison with activation caused by the same sounds when attention was focused on the pictures) in widespread areas of the superior temporal cortex. Attention to either sound feature also activated prefrontal and inferior parietal cortical regions. These activations were stronger during attention to location than during attention to pitch. Attention to location but not to pitch produced a significant increase of activation in the premotor/supplementary motor cortices of both hemispheres and in the right prefrontal cortex, while no area showed activity specifically related to attention to pitch. The present results suggest some differences in the attentional selection of sounds on the basis of their location and pitch consistent with the suggested auditory "what" and "where" processing streams.

Two separate mechanisms underlie auditory change detection and involuntary control of attention.

We used behavioral and event-related potential (ERP) measures to study the neural mechanisms of involuntary attention switching to changes in unattended sounds. Our subjects discriminated two equiprobable sounds differing in frequency (fundamental frequency 186 or 196 Hz) while task-irrelevant intensity decrements or increments (-3, -6, -9, +3, +6, or +9 dB, standard intensity 60 dB HL) infrequently occurred in the same sounds. In line with the results of previous studies, discrimination performance deteriorated with increasing magnitude of the task-irrelevant intensity change. However, these distraction effects were dissimilar for intensity increments and decrements: while there were no differences in reaction time (RT) between intensity decrements and increments, hit rates (HR) were lower for large intensity increments than for large decrements. ERPs to task-irrelevant intensity increments and decrements were also distinctly different: the response to intensity increments consisted of an N1 enhancement, mismatch negativity (MMN), and P3a, while the response to intensity decrements consisted only of MMN. These results are consistent with the assumption that two separate mechanisms (indexed by N1 and MMN) underlie auditory change detection. However, the finding that distinct distraction effects were obtained for both intensity decrements and increments but that the P3a is elicited only by the intensity increments seems to suggest that P3a may not be regarded as a general index of attentional shift but rather it is only generated in conditions in which an enhanced N1 is elicited, too.

Superior temporal and inferior frontal cortices are activated by infrequent sound duration decrements: an fMRI study.

Functional magnetic resonance imaging (fMRI) was used to examine the processing of infrequent changes occurring in an unattended sound sequence. In event-related brain potentials (ERPs), such sound changes typically elicit several responses, including an enhanced N1, the mismatch negativity (MMN), and the P3a. In the present study, subjects were presented with a repeating sound of 75 ms in duration, which was occasionally replaced, in separate blocks, by a 15-ms, 25-ms, or 35-ms sound (large, medium, and small change, respectively). In the baseline block, only the frequent 75-ms sound was presented. During the scanning, the subjects were instructed to ignore the sounds while watching a silent wildlife documentary. We assumed that in this condition, the MMN mechanism would contribute more to the observed activation than the other change-related processes. We expected sound changes to elicit fMRI activation bilaterally in the supratemporal cortices, where the electric MMN is mainly generated, and that the magnitude of this activation would increase with the magnitude of sound duration change. Unexpectedly, however, we found that only blocks with medium duration changes (25 ms) showed significant activation in the supratemporal cortex. In addition, as reported in some previous EEG and fMRI studies, contrasts between different levels of sound duration change revealed additional activation in the inferior frontal cortex bilaterally. This activation tended to be greater for the small and medium changes than for the large ones.

Modulation of auditory cortex activation by sound presentation rate and attention.

We studied the effects of sound presentation rate and attention on auditory supratemporal cortex (STC) activation in 12 healthy adults using functional magnetic resonance imaging (fMRI) at 3 T. The sounds (200 ms in duration) were presented at steady rates of 0.5, 1, 1.5, 2.5, or 4 Hz while subjects either had to focus their attention to the sounds or ignore the sounds and attend to visual stimuli presented with a mean rate of 1 Hz. Consistent with previous observations, we found that both increase in stimulation rate and attention to sounds enhanced activity in STC bilaterally. Further, we observed larger attention effects with higher stimulation rates. This interaction of attention and presentation rate has not been reported previously. In conclusion, our results show both rate-dependent and attention-related modulations of STC indicating that both factors should be controlled, or at least addressed, in fMRI studies of auditory processing.

Spectral and temporal stimulus characteristics in the processing of abstract auditory features.

The processing of abstract stimulus features in the human brain was studied by presenting the subjects with frequent standard tone pairs and infrequent deviant tone pairs. Both pairs varied randomly over a wide frequency and/or intensity range, there being no physically constant standard stimulus. The common feature of the standard pairs was the direction of change within the pair, e.g. the second tone was louder in intensity and/or higher in frequency than the first tone. Deviant pairs, having opposite feature-change direction, elicited the mismatch-negativity (MMN) event-related potential component. MMN was similar to deviations in the direction of frequency and intensity changes and showed no additivity for simultaneous changes in both feature directions. Moreover, MMN was elicited even when the within-pair interval exceeded the 200 ms limit of auditory temporal integration. Results demonstrate that extraction of abstract features is not limited to frequency-based rules, nor is it dependent on temporal integration mechanisms. The lack of MMN additivity between violations of multiple abstract rules suggests that the processing of higher-order invariances differs from that of simple physical features.