Task-dependent activations of human auditory cortex during spatial discrimination and spatial memory tasks.

In the present study, we applied high-resolution functional magnetic resonance imaging (fMRI) of the human auditory cortex (AC) and adjacent areas to compare activations during spatial discrimination and spatial n-back memory tasks that were varied parametrically in difficulty. We found that activations in the anterior superior temporal gyrus (STG) were stronger during spatial discrimination than during spatial memory, while spatial memory was associated with stronger activations in the inferior parietal lobule (IPL). We also found that wide AC areas were strongly deactivated during the spatial memory tasks. The present AC activation patterns associated with spatial discrimination and spatial memory tasks were highly similar to those obtained in our previous study comparing AC activations during pitch discrimination and pitch memory (Rinne et al., 2009). Together our previous and present results indicate that discrimination and memory tasks activate anterior and posterior AC areas differently and that this anterior-posterior division is present both when these tasks are performed on spatially invariant (pitch discrimination vs. memory) or spatially varying (spatial discrimination vs. memory) sounds. These results also further strengthen the view that activations of human AC cannot be explained only by stimulus-level parameters (e.g., spatial vs. nonspatial stimuli) but that the activations observed with fMRI are strongly dependent on the characteristics of the behavioral task. Thus, our results suggest that in order to understand the functional structure of AC a more systematic investigation of task-related factors affecting AC activations is needed.

Task-dependent activations of human auditory cortex during pitch discrimination and pitch memory tasks.

The functional organization of auditory cortex (AC) is still poorly understood. Previous studies suggest segregation of auditory processing streams for spatial and nonspatial information located in the posterior and anterior AC, respectively (Rauschecker and Tian, 2000; Arnott et al., 2004; Lomber and Malhotra, 2008). Furthermore, previous studies have shown that active listening tasks strongly modulate AC activations (Petkov et al., 2004; Fritz et al., 2005; Polley et al., 2006). However, the task dependence of AC activations has not been systematically investigated. In the present study, we applied high-resolution functional magnetic resonance imaging of the AC and adjacent areas to compare activations during pitch discrimination and n-back pitch memory tasks that were varied parametrically in difficulty. We found that anterior AC activations were increased during discrimination but not during memory tasks, while activations in the inferior parietal lobule posterior to the AC were enhanced during memory tasks but not during discrimination. We also found that wide areas of the anterior AC and anterior insula were strongly deactivated during the pitch memory tasks. While these results are consistent with the proposition that the anterior and posterior AC belong to functionally separate auditory processing streams, our results show that this division is present also between tasks using spatially invariant sounds. Together, our results indicate that activations of human AC are strongly dependent on the characteristics of the behavioral task.

Top-down controlled and bottom-up triggered orienting of auditory attention to pitch activate overlapping brain networks.

A number of previous studies have suggested segregated networks of brain areas for top-down controlled and bottom-up triggered orienting of visual attention. However, the corresponding networks involved in auditory attention remain less studied. Our participants attended selectively to a tone stream with either a lower pitch or higher pitch in order to respond to infrequent changes in duration of attended tones. The participants were also required to shift their attention from one stream to the other when guided by a visual arrow cue. In addition to these top-down controlled cued attention shifts, infrequent task-irrelevant louder tones occurred in both streams to trigger attention in a bottom-up manner. Both cued shifts and louder tones were associated with enhanced activity in the superior temporal gyrus and sulcus, temporo-parietal junction, superior parietal lobule, inferior and middle frontal gyri, frontal eye field, supplementary motor area, and anterior cingulate gyrus. Thus, the present findings suggest that in the auditory modality, unlike in vision, top-down controlled and bottom-up triggered attention activate largely the same cortical networks. Comparison of the present results with our previous results from a similar experiment on spatial auditory attention suggests that fronto-parietal networks of attention to location or pitch overlap substantially. However, the auditory areas in the anterior superior temporal cortex might have a more important role in attention to the pitch than location of sounds. This article is part of a Special Issue entitled SI: Prediction and Attention.

Effects of significance of auditory location changes on event related brain potentials and pitch discrimination performance.

We examined effects of significance of task irrelevant changes in the location of tones on the mismatch negativity (MMN) and P3a event related brain potentials. The participants were to discriminate between two frequency modulated tones differing from each other in the direction of frequency glide. Each tone was delivered through one of five loudspeakers in front of the participant. On most trials, a tone was presented from the same location as the preceding tone, but occasionally the location changed. In the Varying Location Condition, these changes, although irrelevant with regard to pitch discrimination, were still significant for performance as the following tones were presented from the new location where attention had to be therefore shifted. In the Fixed Location Condition, the location changes were less significant as the tones following a location change were presented from the original location. In both conditions, the location changes were associated with decreased hit rates and increased reaction times in the pitch discrimination task. However, the hit rate decrease was larger in the Fixed Location Condition suggesting that in this condition the location changes were just distractors. MMN and P3a responses were elicited by location changes in both conditions. In the Fixed Location Condition, a P3a was also elicited by the first tone following a location change at the original location while the MMN was not. Thus, the P3a appeared to be related to shifting of attention in space and was not tightly coupled with MMN elicitation.

Brain networks of bottom-up triggered and top-down controlled shifting of auditory attention.

During functional magnetic resonance imaging (fMRI), our participants selectively attended to tone streams at the left or right, and occasionally shifted their attention from one stream to another as guided by a centrally presented visual cue. Duration changes in the to-be-attended stream served as targets. Loudness deviating tones (LDTs) occurred infrequently in both streams to catch attention in a bottom-up manner, as indicated by their effects on reaction times to targets. LDTs activated the right temporo-parietal junction (TPJ), posterior parts of the left inferior/middle frontal gyrus (IFG/MFG), ventromedial parts of the superior parietal lobule (SPL), and left frontal eye field/premotor cortex (FEF/PMC). In addition, LDTs in the to-be-ignored sound stream were associated with enhanced activity in the ventromedial prefrontal cortex (VMPFC) possibly related to evaluation of the distracting event. Top-down controlled cue-guided attention shifts (CASs) activated bilateral areas in the SPL, intraparietal sulcus (IPS), FEF/PMC, TPJ, IFG/MFG, and cingulate/medial frontal gyrus, and crus I/II of the cerebellum. Thus, our results suggest that in audition top-down controlled and bottom-up triggered shifting of attention activate largely overlapping temporo-parietal, superior parietal and frontal areas. As the IPS, superior parts of the SPL, and crus I/II were activated specifically by top-down controlled attention shifts, and the VMPFC was specifically activated by bottom-up triggered attention shifts, our results also suggest some differences between auditory top-down controlled and bottom-up triggered shifting of attention.

Auditory selective attention modulates activation of human inferior colliculus.

Selective auditory attention powerfully modulates neural activity in the human auditory cortex (AC). In contrast, the role of attention in subcortical auditory processing is not well established. Here, we used functional MRI (fMRI) to examine activation of the human inferior colliculus (IC) during strictly controlled auditory attention tasks. The IC is an obligatory midbrain nucleus of the ascending auditory pathway with diverse internal and external connections. The IC also receives a massive descending projection from the AC, suggesting that cortical processes affect IC operations. In this study, 21 subjects selectively attended to left-ear or right-ear sounds and ignored sounds delivered to the other ear. IC activations depended on the direction of attention, indicating that auditory processing in the human IC is not only determined by acoustic input but also by the current behavioral goals.