Perceptual demand and distraction interactions mediated by task-control networks.

Modulation of selective attention appears to be under the guidance of a cluster of distinct task-control networks, the frontroparietal (FPN) and cingulo-opercular (CON). Yet, their role in mediating the relationship between task perceptual load and presence/absence of distraction in the auditory modality is unclear. Here, we examined this interaction using functional magnetic resonance imaging (fMRI) and an auditory signal detection task. The auditory stimulus signal-to-noise ratio (SNR) was parametrically manipulated, by varying the amplitude of the Tone while holding the Noise constant, to create four perceptual load conditions presented in combination with or without acoustic distraction. Regions of the FPN (e.g., dorsolateral prefrontal cortex, inferior parietal lobule) and CON (e.g., dorsal anterior cingulate cortex/medial superior frontal cortex, anterior prefrontal cortex, anterior insula/frontal operculum) were modulated by perceptual load and distraction, such that lower loads induced a pattern of increased activity when there was no distraction. On the other hand, a trend of augmented activity was found in higher loads during distraction. These findings suggest a role for the FPN and CON in mediating the allocation of attentional resources to competing auditory information under varying degrees of perceptual demand.

The functional organization of the left STS: a large scale meta-analysis of PET and fMRI studies of healthy adults.

The superior temporal sulcus (STS) in the left hemisphere is functionally diverse, with sub-areas implicated in both linguistic and non-linguistic functions. However, the number and boundaries of distinct functional regions remain to be determined. Here, we present new evidence, from meta-analysis of a large number of positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) studies, of different functional specificity in the left STS supporting a division of its middle to terminal extent into at least three functional areas. The middle portion of the left STS stem (fmSTS) is highly specialized for speech perception and the processing of language material. The posterior portion of the left STS stem (fpSTS) is highly versatile and involved in multiple functions supporting semantic memory and associative thinking. The fpSTS responds to both language and non-language stimuli but the sensitivity to non-language material is greater. The horizontal portion of the left STS stem and terminal ascending branches (ftSTS) display intermediate functional specificity, with the anterior-dorsal ascending branch (fatSTS) supporting executive functions and motor planning and showing greater sensitivity to language material, and the horizontal stem and posterior-ventral ascending branch (fptSTS) supporting primarily semantic processing and displaying greater sensitivity to non-language material. We suggest that the high functional specificity of the left fmSTS for speech is an important means by which the human brain achieves exquisite affinity and efficiency for native speech perception. In contrast, the extreme multi-functionality of the left fpSTS reflects the role of this area as a cortical hub for semantic processing and the extraction of meaning from multiple sources of information. Finally, in the left ftSTS, further functional differentiation between the dorsal and ventral aspect is warranted.

Neural effects of cognitive control load on auditory selective attention.

Whether and how working memory disrupts or alters auditory selective attention is unclear. We compared simultaneous event-related potentials (ERP) and functional magnetic resonance imaging (fMRI) responses associated with task-irrelevant sounds across high and low working memory load in a dichotic-listening paradigm. Participants performed n-back tasks (1-back, 2-back) in one ear (Attend ear) while ignoring task-irrelevant speech sounds in the other ear (Ignore ear). The effects of working memory load on selective attention were observed at 130-210ms, with higher load resulting in greater irrelevant syllable-related activation in localizer-defined regions in auditory cortex. The interaction between memory load and presence of irrelevant information revealed stronger activations primarily in frontal and parietal areas due to presence of irrelevant information in the higher memory load. Joint independent component analysis of ERP and fMRI data revealed that the ERP component in the N1 time-range is associated with activity in superior temporal gyrus and medial prefrontal cortex. These results demonstrate a dynamic relationship between working memory load and auditory selective attention, in agreement with the load model of attention and the idea of common neural resources for memory and attention.

Hierarchical organization of speech perception in human auditory cortex.

Human speech consists of a variety of articulated sounds that vary dynamically in spectral composition. We investigated the neural activity associated with the perception of two types of speech segments: (a) the period of rapid spectral transition occurring at the beginning of a stop-consonant vowel (CV) syllable and (b) the subsequent spectral steady-state period occurring during the vowel segment of the syllable. Functional magnetic resonance imaging (fMRI) was recorded while subjects listened to series of synthesized CV syllables and non-phonemic control sounds. Adaptation to specific sound features was measured by varying either the transition or steady-state periods of the synthesized sounds. Two spatially distinct brain areas in the superior temporal cortex were found that were sensitive to either the type of adaptation or the type of stimulus. In a relatively large section of the bilateral dorsal superior temporal gyrus (STG), activity varied as a function of adaptation type regardless of whether the stimuli were phonemic or non-phonemic. Immediately adjacent to this region in a more limited area of the ventral STG, increased activity was observed for phonemic trials compared to non-phonemic trials, however, no adaptation effects were found. In addition, a third area in the bilateral medial superior temporal plane showed increased activity to non-phonemic compared to phonemic sounds. The results suggest a multi-stage hierarchical stream for speech sound processing extending ventrolaterally from the superior temporal plane to the superior temporal sulcus. At successive stages in this hierarchy, neurons code for increasingly more complex spectrotemporal features. At the same time, these representations become more abstracted from the original acoustic form of the sound.

Neural dynamics of phonological processing in the dorsal auditory stream.

Neuroanatomical models hypothesize a role for the dorsal auditory pathway in phonological processing as a feedforward efferent system (Davis and Johnsrude, 2007; Rauschecker and Scott, 2009; Hickok et al., 2011). But the functional organization of the pathway, in terms of time course of interactions between auditory, somatosensory, and motor regions, and the hemispheric lateralization pattern is largely unknown. Here, ambiguous duplex syllables, with elements presented dichotically at varying interaural asynchronies, were used to parametrically modulate phonological processing and associated neural activity in the human dorsal auditory stream. Subjects performed syllable and chirp identification tasks, while event-related potentials and functional magnetic resonance images were concurrently collected. Joint independent component analysis was applied to fuse the neuroimaging data and study the neural dynamics of brain regions involved in phonological processing with high spatiotemporal resolution. Results revealed a highly interactive neural network associated with phonological processing, composed of functional fields in posterior temporal gyrus (pSTG), inferior parietal lobule (IPL), and ventral central sulcus (vCS) that were engaged early and almost simultaneously (at 80-100 ms), consistent with a direct influence of articulatory somatomotor areas on phonemic perception. Left hemispheric lateralization was observed 250 ms earlier in IPL and vCS than pSTG, suggesting that functional specialization of somatomotor (and not auditory) areas determined lateralization in the dorsal auditory pathway. The temporal dynamics of the dorsal auditory pathway described here offer a new understanding of its functional organization and demonstrate that temporal information is essential to resolve neural circuits underlying complex behaviors.

Perceptual demand modulates activation of human auditory cortex in response to task-irrelevant sounds.

In the visual modality, perceptual demand on a goal-directed task has been shown to modulate the extent to which irrelevant information can be disregarded at a sensory-perceptual stage of processing. In the auditory modality, the effect of perceptual demand on neural representations of task-irrelevant sounds is unclear. We compared simultaneous ERPs and fMRI responses associated with task-irrelevant sounds across parametrically modulated perceptual task demands in a dichotic-listening paradigm. Participants performed a signal detection task in one ear (Attend ear) while ignoring task-irrelevant syllable sounds in the other ear (Ignore ear). Results revealed modulation of syllable processing by auditory perceptual demand in an ROI in middle left superior temporal gyrus and in negative ERP activity 130-230 msec post stimulus onset. Increasing the perceptual demand in the Attend ear was associated with a reduced neural response in both fMRI and ERP to task-irrelevant sounds. These findings are in support of a selection model whereby ongoing perceptual demands modulate task-irrelevant sound processing in auditory cortex.

Neural events leading to and associated with detection of sounds under high processing load.

The neural events that lead to successful or failed detection of suprathreshold sounds are not well established. In this experiment, event-related potentials (ERPs) and functional magnetic resonance imaging (fMRI) were recorded while participants performed two tasks: a primary difficult duration judgment task on a sequence of tones presented to one ear, and a secondary target detection task on an auditory oddball stream presented to the other ear. The paradigm was designed to elicit competition and variability in detection of auditory targets despite identical input. Successful detection of auditory targets was associated mainly with greater fMRI activity in superior parietal cortex and thalamus. In the ERPs, successful detection was linked with a larger fronto-central negativity at 200-400 ms, and a later centro-posterior positivity. Failure to detect targets was associated with greater fMRI signal in the default mode network, a significantly smaller electrical fronto-central negativity and no late positivity. These findings demonstrate that variability in auditory detection is related to modulation of activity in multimodal parietal and frontal networks active ∼ 200 ms after target onset. Results are consistent with a limited capacity and late selection view of attention.

Attentional and linguistic interactions in speech perception.

The role of attention in speech comprehension is not well understood. We used fMRI to study the neural correlates of auditory word, pseudoword, and nonspeech (spectrally rotated speech) perception during a bimodal (auditory, visual) selective attention task. In three conditions, Attend Auditory (ignore visual), Ignore Auditory (attend visual), and Visual (no auditory stimulation), 28 subjects performed a one-back matching task in the assigned attended modality. The visual task, attending to rapidly presented Japanese characters, was designed to be highly demanding in order to prevent attention to the simultaneously presented auditory stimuli. Regardless of stimulus type, attention to the auditory channel enhanced activation by the auditory stimuli (Attend Auditory>Ignore Auditory) in bilateral posterior superior temporal regions and left inferior frontal cortex. Across attentional conditions, there were main effects of speech processing (word+pseudoword>rotated speech) in left orbitofrontal cortex and several posterior right hemisphere regions, though these areas also showed strong interactions with attention (larger speech effects in the Attend Auditory than in the Ignore Auditory condition) and no significant speech effects in the Ignore Auditory condition. Several other regions, including the postcentral gyri, left supramarginal gyrus, and temporal lobes bilaterally, showed similar interactions due to the presence of speech effects only in the Attend Auditory condition. Main effects of lexicality (word>pseudoword) were isolated to a small region of the left lateral prefrontal cortex. Examination of this region showed significant word>pseudoword activation only in the Attend Auditory condition. Several other brain regions, including left ventromedial frontal lobe, left dorsal prefrontal cortex, and left middle temporal gyrus, showed Attention x Lexicality interactions due to the presence of lexical activation only in the Attend Auditory condition. These results support a model in which neutral speech presented in an unattended sensory channel undergoes relatively little processing beyond the early perceptual level. Specifically, processing of phonetic and lexical-semantic information appears to be very limited in such circumstances, consistent with prior behavioral studies.

Involvement of the left anterior insula and frontopolar gyrus in odor discrimination.

Discriminating between successively presented odors requires brief storage of the first odor's perceptual trace, which then needs to be subsequently compared to the second odor in the pair. This study explores the cortical areas involved in odor discrimination and compares them with findings from studies of working-memory, traditionally investigated with n-back paradigms. Sixteen right-handed subjects underwent H(2) (15)O positron emission tomography during counterbalanced conditions of odorless sniffing, repeated single odor detection, multiple odor detection, and conscious successive discrimination between odor pairs. Eight odorants were delivered using a computer-controlled olfactometer through a birhinal nasal cannula. Conscious successive odor discrimination evoked significantly greater activity in the left anterior insula and frontopolar gyrus when compared to reported sensory detection of the identical odors. Additional activation was found in the left lateral orbital/inferior frontal and middle frontal gyri when discrimination was compared to the odorless condition. The left anterior insula is likely involved in the evaluation of odor properties. Consistent with other studies, frontopolar and middle frontal gyrus activation is more likely related to working memory during odor discrimination.

Is the failure to detect stimulus deviance during sleep due to a rapid fading of sensory memory or a degradation of stimulus encoding?

The mismatch negativity (MMN) is thought to reflect the outcome of a system responsible for the detection of change in an otherwise repetitive, homogenous acoustic environment. This process depends on the storage and maintenance of a sensory representation of the frequently presented stimulus to which the deviant stimulus is compared. Few studies have been able to record the MMN in non-rapid eye movement (NREM) sleep. This pattern of results might be explained by either a rapid fading of sensory memory or an inhibition of stimulus input prior to entry into the cortical MMN generator site. The present study used a very rapid rate of presentation in an attempt to capture mismatch-related negativity prior to the fading of sensory memory. Auditory event-related potentials were recorded from 12 subjects during a single sleep period. A 1000 Hz standard stimulus was presented every 150 ms. At random, on 6.6% of the trials, the standard was changed to either a large 2000 Hz or a small 1100 Hz deviant. In wakefulness, the large deviant elicited an extended negativity that was reduced in amplitude following the presentation of the small deviant. This negativity was also apparent during REM sleep following the presentation of the large deviant. These deviant-related negativities (DRNs) were probably a composite of N1 and MMN activity. During NREM sleep (stage 2 and slow-wave sleep), only the large deviant continued to elicit a DRN. However this DRN might be overlapped by the initial activity of a component that is unique to sleep, the N350. There was little evidence of the DRN or the MMN during sleep following the presentation of the small deviant. A rapid rate of presentation, therefore, does not preserve the MMN following small deviance within sleep. It is possible that inhibition of sensory input occurs before entry into the MMN generating system in the temporal cortex.

Neural correlates of olfactory change detection.

Detecting changes in a stream of sensory information is vital to animals and humans. While there have been several studies of automatic change detection in various sensory modalities, olfactory change detection is largely unstudied. We investigated brain regions responsive to both passive and active detection of olfactory change using fMRI. Nine right-handed healthy, normosmic subjects (five men) were scanned in two conditions while breathing in synchrony with a metronome. In one condition, subjects mentally counted infrequent odors (Attend condition), whereas in the other condition, subjects' attention was directed elsewhere as they counted auditory tones (Ignore condition). Odors were delivered via a nasal cannula using a computer-controlled air-dilution olfactometer. Infrequently occurring olfactory stimuli evoked significant (P < .05, corrected) activity in the subgenual cingulate and in central posterior orbitofrontal cortex, but only in the Ignore condition, as confirmed by direct comparison of the Ignore session with the Attend session (P < .05, corrected). Subgenual cingulate and posterior orbital cortex may therefore play a role in detecting discrepant olfactory events while attention is otherwise engaged in another sensory modality.

Alcohol-related olfactory cues activate the nucleus accumbens and ventral tegmental area in high-risk drinkers: preliminary findings.

BACKGROUND: The mesocorticolimbic dopamine system is implicated in motivation and reward and may be involved in the development of alcoholism. METHODS: We used functional magnetic resonance imaging to study the blood oxygen level-dependent (BOLD) response to alcohol-related olfactory stimuli (AROS; odors of beer and whiskey) and non-alcohol-related olfactory stimuli (NAROS; odors of grass and leather) in 10 high-risk (HR) drinkers (average drinks per week, 19.99; SD, 6.99; all with > or = 2 first- or second-degree alcoholic relatives) and 5 low-risk (LR) social drinking controls (drinks per week, 2.82; SD, 2.87; 1 subject had 1 second-degree alcoholic relative). Data were analyzed with SPM99 and random effects analysis by using regions of interest and corrected cluster statistics (p < 0.05) to focus on the nucleus accumbens (NAc) and ventral tegmental area (VTA). RESULTS: In HR subjects, there was a greater BOLD signal increase in the NAc during AROS than during clean air. BOLD signal increases during AROS were also greater in the NAc than the signal increases induced by NAROS. The AROS signal was significantly greater than the NAROS signal in a small number of voxels in the VTA. Finally, the AROS/NAROS difference signal was larger in HR drinkers in both the NAc and VTA. CONCLUSIONS: Alcoholic olfactory cues may invoke the dopaminergic mesocorticolimbic system to a greater degree than nonalcoholic odors and could be effective tools in exploring the role of the dopamine system in susceptibility to alcoholism.

Olfactory system activation from sniffing: effects in piriform and orbitofrontal cortex.

Neuroimaging studies suggest that piriform cortex is activated at least in part by sniffing. We used H(2)(15)O positron emission tomography (PET) to study 15 healthy volunteers while they participated in four conditions, two of which were sniffing odorants and odorless air. The remaining two conditions involved a constant, very low flow of either odorized or odorless air during velopharyngeal closure (VPC), a technique that prevents subject-induced airflow through the nasal passages. Contrary to expectation, sniffing under odorless conditions did not induce significant piriform and surrounding cortical (PC+) activity when compared to odorless VPC, even at a liberal statistical threshold. However, a small correlation emerged in PC+ between the difference signal of [odorless sniffing - odorless VPC] and peak rate of nasal pressure change. PC+ activity was, however, strongly evoked by odorant exposure during sniffing and VPC, with neither technique showing greater activation. Activity in orbitofrontal (olfactory association) cortex was absent during odorant stimulation (OS) with VPC, but present during odorant sniffing. Sniffing may therefore play an important role in facilitating the higher-order analysis of odors. A right orbitofrontal region was also activated with odorless sniffing, which suggests a possible orbitofrontal role in guided olfactory exploration.

Neural correlates of auditory sensory memory and automatic change detection.

An auditory event-related potential component, the mismatch negativity (MMN), reflects automatic change detection and its prerequisite, sensory memory. This study examined the neural correlates of automatic change detection using BOLD fMRI and two rates of presentation previously shown to induce either a large or no MMN. A boxcar block design was employed in two functional scans, each performed twice. A block consisting of 1000-Hz standards (S) alternated with one consisting of 1000-Hz standards and 2000-Hz infrequent deviants (S + D). Presentation rate was either 150 or 2400 ms. Fourteen participants were instructed to ignore all auditory stimulation and concentrate on a film (no audio) by reading subtitles. Data analysis used SPM99 and random effects approach. Cluster statistics (P < 0.05, corrected) were employed at a height threshold of P < 0.001. At the short ISI, there was a significant BOLD response in the right superior temporal gyrus (STG), the left insula, and the left STG (including parts of primary auditory cortex). There were no suprathreshold clusters at the long rate, with S + D blocks inducing no greater activity than S blocks. These results support the hypothesis that the automatic detection of auditory change occurs in the STG bilaterally and relies on the maintenance of sensory memory traces.

Effects of sleep onset on the mismatch negativity (MMN) to frequency deviants using a rapid rate of presentation.

This study examined the effects of sleep onset-the transition from a waking, conscious state to one of sleep and unconsciousness-on the mismatch negativity (MMN) following frequency deviants when a rapid rate of stimulus presentation is employed. The MMN is thought to reflect a brief-lasting sensory memory. Rapid rates of stimulus presentation should guard the sensory memory from fading. A 1,000 Hz standard stimulus was presented every 150 ms. At random, on 6.6% of the trials, the standard was changed to either a large 2,000 or a small 1,100 Hz deviant. During alert wakefulness (when subject ignored the stimuli and read a book), the large deviant elicited a larger deviant related negativity (DRN) than did the small deviant. This negativity may be a composite of both N1 and MMN activity while that following the small deviant is probably a 'true' MMN. The large deviant continued to elicit a DRN in relaxed wakefulness (eyes closed) and Stages 1 and 2 of sleep, although it was much reduced in amplitude. A significant MMN was recorded for the small deviant only in alert wakefulness. The failure to observe an MMN to small deviance and the attenuation of the DRN to large deviance at sleep onset therefore is probably not due to a decay of sensory memory. It is more likely that cortical encoding of both the standard and deviant is weakened during sleep onset because of prior thalamic inhibition of sensory input.

The effects of digital filtering on mismatch negativity in wakefulness and slow-wave sleep.

The mismatch negativity (MMN) is a response to a deviant auditory stimulus that occurs infrequently in a sequence of otherwise repetitive, homogeneous standard auditory stimuli. The MMN is presumed automatic and independent of conscious awareness. Recording of the MMN during unconscious states may be problematic. The frequency content of the long-lasting MMN may overlap and summate with other event-related slow potentials and low-frequency background electroencephalogram (EEG) activity. The purpose of this study is to determine the optimal filter settings for recording the MMN during unconscious states. Auditory event-related potentials (ERPs) were recorded from eight subjects in an oddball paradigm during wakefulness and Stages 3 and 4 of sleep [slow-wave sleep (SWS)] using a 0.16-35 Hz analogue bandpass. Deviant probability was 0.033. Stimulus-onset asynchrony was 150 ms. The EEG data were subsequently digitally filtered in the frequency domain. The low-pass filter was set at either 24, 12 or 6 Hz, and the high-pass filter at either 1, 2, 3 or 4 Hz. Applying a low-pass filter down to 12 Hz had a minimal impact on the waking or sleeping MMN amplitude. On the other hand, increasing the high-pass setting from 2 to 3 Hz permitted the visualization of the MMN recorded during sleep. The 4 Hz filter showed a similar trend but also markedly attenuated the amplitude of the waking MMN. A high-pass setting of 3 Hz provides a reasonable compromise. It has only a slight effect on the MMN when the subject is conscious, but still attenuates most of the unwanted slow potential activity when the subject enters SWS.