Reward cues readily direct monkeys' auditory performance resulting in broad auditory cortex modulation and interaction with sites along cholinergic and dopaminergic pathways.

In natural settings, the prospect of reward often influences the focus of our attention, but how cognitive and motivational systems influence sensory cortex is not well understood. Also, challenges in training nonhuman animals on cognitive tasks complicate cross-species comparisons and interpreting results on the neurobiological bases of cognition. Incentivized attention tasks could expedite training and evaluate the impact of attention on sensory cortex. Here we develop an Incentivized Attention Paradigm (IAP) and use it to show that macaque monkeys readily learn to use auditory or visual reward cues, drastically influencing their performance within a simple auditory task. Next, this paradigm was used with functional neuroimaging to measure activation modulation in the monkey auditory cortex. The results show modulation of extensive auditory cortical regions throughout primary and non-primary regions, which although a hallmark of attentional modulation in human auditory cortex, has not been studied or observed as broadly in prior data from nonhuman animals. Psycho-physiological interactions were identified between the observed auditory cortex effects and regions including basal forebrain sites along acetylcholinergic and dopaminergic pathways. The findings reveal the impact and regional interactions in the primate brain during an incentivized attention engaging auditory task.

Interaction of the effects associated with auditory-motor integration and attention-engaging listening tasks.

A number of previous studies have implicated regions in posterior auditory cortex (AC) in auditory-motor integration during speech production. Other studies, in turn, have shown that activation in AC and adjacent regions in the inferior parietal lobule (IPL) is strongly modulated during active listening and depends on task requirements. The present fMRI study investigated whether auditory-motor effects interact with those related to active listening tasks in AC and IPL. In separate task blocks, our subjects performed either auditory discrimination or 2-back memory tasks on phonemic or nonphonemic vowels. They responded to targets by either overtly repeating the last vowel of a target pair, overtly producing a given response vowel, or by pressing a response button. We hypothesized that the requirements for auditory-motor integration, and the associated activation, would be stronger during repetition than production responses and during repetition of nonphonemic than phonemic vowels. We also hypothesized that if auditory-motor effects are independent of task-dependent modulations, then the auditory-motor effects should not differ during discrimination and 2-back tasks. We found that activation in AC and IPL was significantly modulated by task (discrimination vs. 2-back), vocal-response type (repetition vs. production), and motor-response type (vocal vs. button). Motor-response and task effects interacted in IPL but not in AC. Overall, the results support the view that regions in posterior AC are important in auditory-motor integration. However, the present study shows that activation in wide AC and IPL regions is modulated by the motor requirements of active listening tasks in a more general manner. Further, the results suggest that activation modulations in AC associated with attention-engaging listening tasks and those associated with auditory-motor performance are mediated by independent mechanisms.

Intrinsic, stimulus-driven and task-dependent connectivity in human auditory cortex.

A hierarchical and modular organization is a central hypothesis in the current primate model of auditory cortex (AC) but lacks validation in humans. Here we investigated whether fMRI connectivity at rest and during active tasks is informative of the functional organization of human AC. Identical pitch-varying sounds were presented during a visual discrimination (i.e. no directed auditory attention), pitch discrimination, and two versions of pitch n-back memory tasks. Analysis based on fMRI connectivity at rest revealed a network structure consisting of six modules in supratemporal plane (STP), temporal lobe, and inferior parietal lobule (IPL) in both hemispheres. In line with the primate model, in which higher-order regions have more longer-range connections than primary regions, areas encircling the STP module showed the highest inter-modular connectivity. Multivariate pattern analysis indicated significant connectivity differences between the visual task and rest (driven by the presentation of sounds during the visual task), between auditory and visual tasks, and between pitch discrimination and pitch n-back tasks. Further analyses showed that these differences were particularly due to connectivity modulations between the STP and IPL modules. While the results are generally in line with the primate model, they highlight the important role of human IPL during the processing of both task-irrelevant and task-relevant auditory information. Importantly, the present study shows that fMRI connectivity at rest, during presentation of sounds, and during active listening provides novel information about the functional organization of human AC.

Evidence for cue-independent spatial representation in the human auditory cortex during active listening.

Few auditory functions are as important or as universal as the capacity for auditory spatial awareness (e.g., sound localization). That ability relies on sensitivity to acoustical cues-particularly interaural time and level differences (ITD and ILD)-that correlate with sound-source locations. Under nonspatial listening conditions, cortical sensitivity to ITD and ILD takes the form of broad contralaterally dominated response functions. It is unknown, however, whether that sensitivity reflects representations of the specific physical cues or a higher-order representation of auditory space (i.e., integrated cue processing), nor is it known whether responses to spatial cues are modulated by active spatial listening. To investigate, sensitivity to parametrically varied ITD or ILD cues was measured using fMRI during spatial and nonspatial listening tasks. Task type varied across blocks where targets were presented in one of three dimensions: auditory location, pitch, or visual brightness. Task effects were localized primarily to lateral posterior superior temporal gyrus (pSTG) and modulated binaural-cue response functions differently in the two hemispheres. Active spatial listening (location tasks) enhanced both contralateral and ipsilateral responses in the right hemisphere but maintained or enhanced contralateral dominance in the left hemisphere. Two observations suggest integrated processing of ITD and ILD. First, overlapping regions in medial pSTG exhibited significant sensitivity to both cues. Second, successful classification of multivoxel patterns was observed for both cue types and-critically-for cross-cue classification. Together, these results suggest a higher-order representation of auditory space in the human auditory cortex that at least partly integrates the specific underlying cues.

Functional Imaging of Audio-Visual Selective Attention in Monkeys and Humans: How do Lapses in Monkey Performance Affect Cross-Species Correspondences?

The cross-species correspondences and differences in how attention modulates brain responses in humans and animal models are poorly understood. We trained 2 monkeys to perform an audio-visual selective attention task during functional magnetic resonance imaging (fMRI), rewarding them to attend to stimuli in one modality while ignoring those in the other. Monkey fMRI identified regions strongly modulated by auditory or visual attention. Surprisingly, auditory attention-related modulations were much more restricted in monkeys than humans performing the same tasks during fMRI. Further analyses ruled out trivial explanations, suggesting that labile selective-attention performance was associated with inhomogeneous modulations in wide cortical regions in the monkeys. The findings provide initial insights into how audio-visual selective attention modulates the primate brain, identify sources for "lost" attention effects in monkeys, and carry implications for modeling the neurobiology of human cognition with nonhuman animals.

Processing of pitch and location in human auditory cortex during visual and auditory tasks.

The relationship between stimulus-dependent and task-dependent activations in human auditory cortex (AC) during pitch and location processing is not well understood. In the present functional magnetic resonance imaging study, we investigated the processing of task-irrelevant and task-relevant pitch and location during discrimination, n-back, and visual tasks. We tested three hypotheses: (1) According to prevailing auditory models, stimulus-dependent processing of pitch and location should be associated with enhanced activations in distinct areas of the anterior and posterior superior temporal gyrus (STG), respectively. (2) Based on our previous studies, task-dependent activation patterns during discrimination and n-back tasks should be similar when these tasks are performed on sounds varying in pitch or location. (3) Previous studies in humans and animals suggest that pitch and location tasks should enhance activations especially in those areas that also show activation enhancements associated with stimulus-dependent pitch and location processing, respectively. Consistent with our hypotheses, we found stimulus-dependent sensitivity to pitch and location in anterolateral STG and anterior planum temporale (PT), respectively, in line with the view that these features are processed in separate parallel pathways. Further, task-dependent activations during discrimination and n-back tasks were associated with enhanced activations in anterior/posterior STG and posterior STG/inferior parietal lobule (IPL) irrespective of stimulus features. However, direct comparisons between pitch and location tasks performed on identical sounds revealed no significant activation differences. These results suggest that activations during pitch and location tasks are not strongly affected by enhanced stimulus-dependent activations to pitch or location. We also found that activations in PT were strongly modulated by task requirements and that areas in the inferior parietal lobule (IPL) showed task-dependent activation modulations, but no systematic activations to pitch or location. Based on these results, we argue that activations during pitch and location tasks cannot be explained by enhanced stimulus-specific processing alone, but rather that activations in human AC depend in a complex manner on the requirements of the task at hand.

The effect of precision and power grips on activations in human auditory cortex.

The neuroanatomical pathways interconnecting auditory and motor cortices play a key role in current models of human auditory cortex (AC). Evidently, auditory-motor interaction is important in speech and music production, but the significance of these cortical pathways in other auditory processing is not well known. We investigated the general effects of motor responding on AC activations to sounds during auditory and visual tasks (motor regions were not imaged). During all task blocks, subjects detected targets in the designated modality, reported the relative number of targets at the end of the block, and ignored the stimuli presented in the opposite modality. In each block, they were also instructed to respond to targets either using a precision grip, power grip, or to give no overt target responses. We found that motor responding strongly modulated AC activations. First, during both visual and auditory tasks, activations in widespread regions of AC decreased when subjects made precision and power grip responses to targets. Second, activations in AC were modulated by grip type during the auditory but not during the visual task. Further, the motor effects were distinct from the present strong attention-related modulations in AC. These results are consistent with the idea that operations in AC are shaped by its connections with motor cortical regions.

Brain activations during bimodal dual tasks depend on the nature and combination of component tasks.

We used functional magnetic resonance imaging to investigate brain activations during nine different dual tasks in which the participants were required to simultaneously attend to concurrent streams of spoken syllables and written letters. They performed a phonological, spatial or "simple" (speaker-gender or font-shade) discrimination task within each modality. We expected to find activations associated specifically with dual tasking especially in the frontal and parietal cortices. However, no brain areas showed systematic dual task enhancements common for all dual tasks. Further analysis revealed that dual tasks including component tasks that were according to Baddeley's model "modality atypical," that is, the auditory spatial task or the visual phonological task, were not associated with enhanced frontal activity. In contrast, for other dual tasks, activity specifically associated with dual tasking was found in the left or bilateral frontal cortices. Enhanced activation in parietal areas, however, appeared not to be specifically associated with dual tasking per se, but rather with intermodal attention switching. We also expected effects of dual tasking in left frontal supramodal phonological processing areas when both component tasks required phonological processing and in right parietal supramodal spatial processing areas when both tasks required spatial processing. However, no such effects were found during these dual tasks compared with their component tasks performed separately. Taken together, the current results indicate that activations during dual tasks depend in a complex manner on specific demands of component tasks.

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.

Source analysis of event-related potentials during pitch discrimination and pitch memory tasks.

Our previous studies using fMRI have demonstrated that activations in human auditory cortex (AC) are strongly dependent on the characteristics of the task. The present study tested whether source estimation of scalp-recorded event-related potentials (ERPs) can be used to investigate task-dependent AC activations. Subjects were presented with frequency-varying two-part tones during pitch discrimination, pitch n-back memory, and visual tasks identical to our previous fMRI study (Rinne et al., J Neurosci 29:13338-13343, 2009). ERPs and their minimum-norm source estimates in AC were strongly modulated by task at 200-700 ms from tone onset. As in the fMRI study, the pitch discrimination and pitch memory tasks were associated with distinct AC activation patterns. In the pitch discrimination task, increased activity in the anterior AC was detected relatively late at 300-700 ms from tone onset. Therefore, this activity was probably not associated with enhanced pitch processing but rather with the actual discrimination process (comparison between the two parts of tone). Increased activity in more posterior areas associated with the pitch memory task, in turn, occurred at 200-700 ms suggesting that this activity was related to operations on pitch categories after pitch analysis was completed. Finally, decreased activity associated with the pitch memory task occurred at 150-300 ms consistent with the notion that, in the demanding pitch memory task, spectrotemporal analysis is actively halted as soon as category information has been obtained. These results demonstrate that ERP source analysis can be used to complement fMRI to investigate task-dependent activations of human AC.

Acoustical and categorical tasks differently modulate activations of human auditory cortex to vowels.

The present study compared activations to prototype, nonprototype, nonphonemic, and cross-category vowel pairs during vowel discrimination, category discrimination, 2-back, and visual tasks. Our results support previous findings that areas of the superior temporal gyrus (STG) are sensitive to the speech-level difference between prototype vs. nonprototype and phonemic vs. nonphonemic vowels. Further, consistent with previous studies, we found enhanced activations in anterior-posterior STG and inferior parietal lobule (IPL) during the vowel discrimination and 2-back tasks, respectively. Unlike the vowel discrimination task, the category discrimination task was associated with strong IPL activations. Our results provide evidence that activations in STG and IPL strongly depend on whether the task requires analysis of detailed acoustical information or operations on categorical representations. Based on previous studies investigating activations during categorical pitch and spatial tasks, we argue that this distinction is probably not specific to speech.

Processing of spatial sounds in human auditory cortex during visual, discrimination and 2-back tasks.

Previous imaging studies on the brain mechanisms of spatial hearing have mainly focused on sounds varying in the horizontal plane. In this study, we compared activations in human auditory cortex (AC) and adjacent inferior parietal lobule (IPL) to sounds varying in horizontal location, distance, or space (i.e., different rooms). In order to investigate both stimulus-dependent and task-dependent activations, these sounds were presented during visual discrimination, auditory discrimination, and auditory 2-back memory tasks. Consistent with previous studies, activations in AC were modulated by the auditory tasks. During both auditory and visual tasks, activations in AC were stronger to sounds varying in horizontal location than along other feature dimensions. However, in IPL, this enhancement was detected only during auditory tasks. Based on these results, we argue that IPL is not primarily involved in stimulus-level spatial analysis but that it may represent such information for more general processing when relevant to an active auditory task.

Stimulus-dependent activations and attention-related modulations in the auditory cortex: a meta-analysis of fMRI studies.

We meta-analyzed 115 functional magnetic resonance imaging (fMRI) studies reporting auditory-cortex (AC) coordinates for activations related to active and passive processing of pitch and spatial location of non-speech sounds, as well as to the active and passive speech and voice processing. We aimed at revealing any systematic differences between AC surface locations of these activations by statistically analyzing the activation loci using the open-source Matlab toolbox VAMCA (Visualization and Meta-analysis on Cortical Anatomy). AC activations associated with pitch processing (e.g., active or passive listening to tones with a varying vs. fixed pitch) had median loci in the middle superior temporal gyrus (STG), lateral to Heschl's gyrus. However, median loci of activations due to the processing of infrequent pitch changes in a tone stream were centered in the STG or planum temporale (PT), significantly posterior to the median loci for other types of pitch processing. Median loci of attention-related modulations due to focused attention to pitch (e.g., attending selectively to low or high tones delivered in concurrent sequences) were, in turn, centered in the STG or superior temporal sulcus (STS), posterior to median loci for passive pitch processing. Activations due to spatial processing were centered in the posterior STG or PT, significantly posterior to pitch processing loci (processing of infrequent pitch changes excluded). In the right-hemisphere AC, the median locus of spatial attention-related modulations was in the STS, significantly inferior to the median locus for passive spatial processing. Activations associated with speech processing and those associated with voice processing had indistinguishable median loci at the border of mid-STG and mid-STS. Median loci of attention-related modulations due to attention to speech were in the same mid-STG/STS region. Thus, while attention to the pitch or location of non-speech sounds seems to recruit AC areas less involved in passive pitch or location processing, focused attention to speech predominantly enhances activations in regions that already respond to human vocalizations during passive listening. This suggests that distinct attention mechanisms might be engaged by attention to speech and attention to more elemental auditory features such as tone pitch or location. This article is part of a Special Issue entitled Human Auditory Neuroimaging.

Attention effects on the processing of task-relevant and task-irrelevant speech sounds and letters.

We used event-related brain potentials (ERPs) to study effects of selective attention on the processing of attended and unattended spoken syllables and letters. Participants were presented with syllables randomly occurring in the left or right ear and spoken by different voices and with a concurrent foveal stream of consonant letters written in darker or lighter fonts. During auditory phonological (AP) and non-phonological tasks, they responded to syllables in a designated ear starting with a vowel and spoken by female voices, respectively. These syllables occurred infrequently among standard syllables starting with a consonant and spoken by male voices. During visual phonological and non-phonological tasks, they responded to consonant letters with names starting with a vowel and to letters written in dark fonts, respectively. These letters occurred infrequently among standard letters with names starting with a consonant and written in light fonts. To examine genuine effects of attention and task on ERPs not overlapped by ERPs associated with target processing or deviance detection, these effects were studied only in ERPs to auditory and visual standards. During selective listening to syllables in a designated ear, ERPs to the attended syllables were negatively displaced during both phonological and non-phonological auditory tasks. Selective attention to letters elicited an early negative displacement and a subsequent positive displacement (Pd) of ERPs to attended letters being larger during the visual phonological than non-phonological task suggesting a higher demand for attention during the visual phonological task. Active suppression of unattended speech during the AP and non-phonological tasks and during the visual phonological tasks was suggested by a rejection positivity (RP) to unattended syllables. We also found evidence for suppression of the processing of task-irrelevant visual stimuli in visual ERPs during auditory tasks involving left-ear syllables.

The functional role of the frontal cortex in pre-attentive auditory change detection.

Accounts of the functional role of the frontal cortex in pre-attentive auditory change detection include attention switching, response inhibition, contrast enhancement, and activation of a predictive model. These accounts assume different sequential activation patterns between the temporal and frontal cortices: Change detection in the auditory areas of the superior temporal cortex (STC) followed by inferior frontal cortex (IFC) activation for attention switching and response inhibition; STC preceded by IFC activation for contrast enhancement; and an IFC-STC-IFC activation sequence for the predictive model. We used the event-related optical signal (EROS), which provides a temporal resolution of milliseconds and a spatial resolution of 5 to 10mm, combined with lagged correlation path modeling to examine the response of the right frontal and temporal cortices to auditory duration deviants of varying magnitude. Event-related potentials (ERPs) were also recorded, as was the slow optical (hemodynamic) brain response. The data analyses revealed temporal-frontal, frontal-temporal-frontal, and temporal-frontal activation patterns when the deviants represented relatively large, medium, and small changes from the standard stimulus, respectively. These results indicate that the degree of deviance modulates spatio-temporal dynamics within the STC-IFC auditory change detection network.

Activations of human auditory cortex to phonemic and nonphonemic vowels during discrimination and memory tasks.

We used fMRI to investigate activations within human auditory cortex (AC) to vowels during vowel discrimination, vowel (categorical n-back) memory, and visual tasks. Based on our previous studies, we hypothesized that the vowel discrimination task would be associated with increased activations in the anterior superior temporal gyrus (STG), while the vowel memory task would enhance activations in the posterior STG and inferior parietal lobule (IPL). In particular, we tested the hypothesis that activations in the IPL during vowel memory tasks are associated with categorical processing. Namely, activations due to categorical processing should be higher during tasks performed on nonphonemic (hard to categorize) than on phonemic (easy to categorize) vowels. As expected, we found distinct activation patterns during vowel discrimination and vowel memory tasks. Further, these task-dependent activations were different during tasks performed on phonemic or nonphonemic vowels. However, activations in the IPL associated with the vowel memory task were not stronger during nonphonemic than phonemic vowel blocks. Together these results demonstrate that activations in human AC to vowels depend on both the requirements of the behavioral task and the phonemic status of the vowels.

Task-dependent activations of human auditory cortex to prototypical and nonprototypical vowels.

Research in auditory neuroscience has largely neglected the possible effects of different listening tasks on activations of auditory cortex (AC). In the present study, we used high-resolution fMRI to compare human AC activations with sounds presented during three auditory and one visual task. In all tasks, subjects were presented with pairs of Finnish vowels, noise bursts with pitch and Gabor patches. In the vowel pairs, one vowel was always either a prototypical /i/ or /ae/ (separately defined for each subject) or a nonprototype. In different task blocks, subjects were either required to discriminate (same/different) vowel pairs, to rate vowel "goodness" (first/second sound was a better exemplar of the vowel class), to discriminate pitch changes in the noise bursts, or to discriminate Gabor orientation changes. We obtained distinctly different AC activation patterns to identical sounds presented during the four task conditions. In particular, direct comparisons between the vowel tasks revealed stronger activations during vowel discrimination in the anterior and posterior superior temporal gyrus (STG), while the vowel rating task was associated with increased activations in the inferior parietal lobule (IPL). We also found that AC areas in or near Heschl's gyrus (HG) were sensitive to the speech-specific difference between a vowel prototype and nonprototype during active listening tasks. These results show that AC activations to speech sounds are strongly dependent on the listening tasks.

Brain activity during auditory and visual phonological, spatial and simple discrimination tasks.

We used functional magnetic resonance imaging to measure human brain activity during tasks demanding selective attention to auditory or visual stimuli delivered in concurrent streams. Auditory stimuli were syllables spoken by different voices and occurring in central or peripheral space. Visual stimuli were centrally or more peripherally presented letters in darker or lighter fonts. The participants performed a phonological, spatial or "simple" (speaker-gender or font-shade) discrimination task in either modality. Within each modality, we expected a clear distinction between brain activations related to nonspatial and spatial processing, as reported in previous studies. However, within each modality, different tasks activated largely overlapping areas in modality-specific (auditory and visual) cortices, as well as in the parietal and frontal brain regions. These overlaps may be due to effects of attention common for all three tasks within each modality or interaction of processing task-relevant features and varying task-irrelevant features in the attended-modality stimuli. Nevertheless, brain activations caused by auditory and visual phonological tasks overlapped in the left mid-lateral prefrontal cortex, while those caused by the auditory and visual spatial tasks overlapped in the inferior parietal cortex. These overlapping activations reveal areas of multimodal phonological and spatial processing. There was also some evidence for intermodal attention-related interaction. Most importantly, activity in the superior temporal sulcus elicited by unattended speech sounds was attenuated during the visual phonological task in comparison with the other visual tasks. This effect might be related to suppression of processing irrelevant speech presumably distracting the phonological task involving the letters.

Attention-related modulation of auditory-cortex responses to speech sounds during dichotic listening.

Event-related magnetic fields (ERFs) were measured with magnetoencephalography (MEG) in fifteen healthy right-handed participants listening to sequences of consonant-vowel syllable pairs delivered dichotically (one syllable presented to the left ear and another syllable simultaneously to the right ear). The participants were instructed to press a response button to occurrences of a particular target syllable. In a condition with no other instruction (the non-forced condition, NF), they showed the well-known right-ear advantage (REA), that is, the participants responded more often to target syllables delivered to the right ear than to targets delivered to the left ear. The same was true in the forced-right (FR) condition, where the participants were instructed to attend selectively to the right-ear syllables and respond only to targets among them. In the forced-left (FL) condition, where they were instructed to respond only to left-ear targets, they responded more often to targets in this ear than to targets in the right ear. At 300-500 ms from syllable pair onset, a sustained field (SF) in ERFs to the syllable pairs was stronger in the left auditory cortex than in the right auditory cortex in the NF and FR conditions, while the opposite was true in the FL condition. Thus selective attention during dichotic listening leads to stronger processing of speech sounds in the auditory cortex contralateral to the attended direction. Our results also suggest that the REA observed for dichotic speech may involve a bias of attention to the right side even when there is no instruction to do so. This supports Kinsbourne's (1970) model of attention bias as a general principle of laterality.

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.