Stephen R. Arnott

What and "where" in the human auditory system.

The extent to which sound identification and sound localization depend on specialized auditory pathways was examined by using functional magnetic resonance imaging and event-related brain potentials. Participants performed an S1–S2 match-to-sample task in which S1 differed from S2 in its pitch and/or location. In the pitch task, participants indicated whether S2 was lower, identical, or higher in pitch than S1. In the location task, participants were asked to localize S2 relative to S1 (i.e., leftward, same, or rightward). Relative to location, pitch processing generated greater activation in auditory cortex and the inferior frontal gyrus. Conversely, identifying the location of S2 relative to S1 generated greater activation in posterior temporal cortex, parietal cortex, and the superior frontal sulcus. Differential task-related effects on event-related brain potentials (ERPs) were seen in anterior and posterior brain regions beginning at 300 ms poststimulus and lasting for several hundred milliseconds. The converging evidence from two independent measurements of dissociable brain activity during identification and localization of identical stimuli provides strong support for specialized auditory streams in the human brain. These findings are analogous to the “what” and “where” segregation of visual information processing, and suggest that a similar functional organization exists for processing information from the auditory modality.

Bottom-up and top-down influences on auditory scene analysis: evidence from event-related brain potentials.

The physiological processes underlying the segregation of concurrent sounds were investigated through the use of event-related brain potentials. The stimuli were complex sounds containing multiple harmonics, one of which could be mistuned so that it was no longer an integer multiple of the fundamental. Perception of concurrent auditory objects increased with degree of mistuning and was accompanied by negative and positive waves that peaked at 180 and 400 ms poststimulus, respectively. The negative wave, referred to as object-related negativity, was present during passive listening, but the positive wave was not. These findings indicate bottom-up and top-down influences during auditory scene analysis. Brain electrical source analyses showed that distinguishing simultaneous auditory objects involved a widely distributed neural network that included auditory cortices, the medial temporal lobe, and posterior association cortices.

Neural Correlates of Natural Human Echolocation in Early and Late Blind Echolocation Experts

Background A small number of blind people are adept at echolocating silent objects simply by producing mouth clicks and listening to the returning echoes. Yet the neural architecture underlying this type of aid-free human echolocation has not been investigated. To tackle this question, we recruited echolocation experts, one early- and one late-blind, and measured functional brain activity in each of them while they listened to their own echolocation sounds. Results When we compared brain activity for sounds that contained both clicks and the returning echoes with brain activity for control sounds that did not contain the echoes, but were otherwise acoustically matched, we found activity in calcarine cortex in both individuals. Importantly, for the same comparison, we did not observe a difference in activity in auditory cortex. In the early-blind, but not the late-blind participant, we also found that the calcarine activity was greater for echoes reflected from surfaces located in contralateral space. Finally, in both individuals, we found activation in middle temporal and nearby cortical regions when they listened to echoes reflected from moving targets. Conclusions These findings suggest that processing of click-echoes recruits brain regions typically devoted to vision rather than audition in both early and late blind echolocation experts.

fMR-adaptation reveals separate processing regions for the perception of form and texture in the human ventral stream

We recently demonstrated that attending to the form of objects and attending to their surface properties activated anatomically distinct regions of occipito-temporal cortex (Cant and Goodale, Cereb Cortex 17:713–731, 2007). Specifically, attending to form activated the lateral occipital area (LO), whereas attending to texture activated the collateral sulcus (CoS). Although these regions showed preferential activation to one particular stimulus dimension (e.g. texture in CoS), they also showed activation to other, non-preferred stimulus dimensions (e.g. form in CoS). This raises the question as to whether the activation associated with attention to form in CoS, for example, represents the actual processing of object form or instead represents the obligatory processing of object texture that occurred when people attended to form. To investigate this, we conducted an fMR-adaptation experiment which allowed us to examine the response properties of regions specialized for processing form, texture, and colour when participants were not explicitly attending to a particular stimulus dimension. Participants passively viewed blocks where only one dimension varied and blocks where no dimensions varied, while fixating a cross in the centre of the display. Area LO was most sensitive to variations in form, whereas the CoS was most sensitive to variations in texture. As in our previous study, no regions were found that were most sensitive to variations in colour, but unlike the results from that study, medial regions of the ventral stream along the fusiform gyrus and CoS showed some selectivity to colour. Taken together, these results replicate the findings from our previous study and provide additional evidence for the existence of separate processing pathways for form and surface properties (particularly texture) in the ventral stream.

The Functional Organization of Auditory Working Memory as Revealed by fMRI

Spatial and nonspatial auditory tasks preferentially recruit dorsal and ventral brain areas, respectively. However, the extent to which these auditory differences reflect specific aspects of mental processing has not been directly studied. In the present functional magnetic resonance imaging experiment, participants encoded and maintained either the location or the identity of a sound for a delay period of several seconds and then subsequently compared that information with a second sound. Relative to sound localization, sound identification was associated with greater hemodynamic activity in the left rostral superior temporal gyrus. In contrast, localizing sounds recruited greater activity in the parietal cortex, posterior temporal lobe, and superior frontal sulcus. The identification differences were most prominent during the early stage of the trial, whereas the location differences were most evident during the late (i.e., comparison) stage. Accordingly, our results suggest that auditory spatial and identity dissociations as revealed by functional imaging may be dependent to some degree on the type of processing being carried out. In addition, dorsolateral prefrontal and lateral superior parietal areas showed greater activity during the comparison as opposed to the earlier stage of the trial, regardless of the type of auditory task, consistent with results from visual working memory studies.

The auditory dorsal pathway: Orienting vision

A particularly prominent model of auditory cortical function proposes that a dorsal brain pathway, emanating from the posterior auditory cortex, is primarily concerned with processing the spatial features of sounds. In the present paper, we outline some difficulties with a strict functional interpretation of this pathway, and highlight the recent trend to understand this pathway in terms of one that uses acoustic information to guide motor output towards objects of interest. In this spirit, we consider the possibility that some of the auditory spatial processing activity that has been observed in the dorsal pathway may actually be understood as a form of action processing in which the visual system may be guided to a particular location of interest. In this regard, attentional orientation may be considered a low-level form of action planning. Incorporating an auditory-guided motor aspect to the dorsal pathway not only offers a more holistic account of auditory processing, but also provides a more ecologically valid perspective on auditory processing in dorsal brain regions.

An investigation of auditory contagious yawning

Despite a widespread familiarity with the often compelling urge to yawn after perceiving someone else yawn, an understanding of the neural mechanism underlying contagious yawning remains incomplete. In the present auditory fMRI study, listeners used a 4-point scale to indicate how much they felt like yawning following the presentation of a yawn, breath, or scrambled yawn sound. Not only were yawn sounds given significantly higher ratings, a trait positively correlated with each individual’s empathy measure, but relative to control stimuli, random effects analyses revealed enhanced hemodynamic activity in the right posterior inferior frontal gyrus (pIFG) in response to hearing yawns. Moreover, pIFG activity was greatest for yawn stimuli associated with high as opposed to low yawn ratings and for control sounds associated with equally high yawn ratings. These results support a relationship between contagious yawning and empathy and provide evidence for pIFG involvement in contagious yawning. A supplemental figure for this study may be downloaded from http://cabn.psychonomic-journals.org/content/supplemental.

Crinkling and crumpling: An auditory fMRI study of material properties

Knowledge of an objects material composition (i.e., what it is made of) alters how we interact with that object. Seeing the bright glint or hearing the metallic crinkle of a foil plate for example, confers information about that object before we have even touched it. Recent research indicates that the medial aspect of the ventral visual pathway is sensitive to the surface properties of objects. In the present functional magnetic resonance imaging (fMRI) study, we investigated whether the ventral pathway is also sensitive to material properties derived from sound alone. Relative to scrambled material sounds and non-verbal human vocalizations, audio recordings of materials being manipulated (i.e., crumpled) in someones hands elicited greater BOLD activity in the right parahippocampal cortex of neurologically intact listeners, as well as a cortically blind participant. Additional left inferior parietal lobe activity was also observed in the neurologically intact group. Taken together, these results support a ventro-medial pathway that is specialized for processing the material properties of objects, and suggest that there are sub-regions within this pathway that subserve the processing of acoustically-derived information about material composition.

Shape-specific activation of occipital cortex in an early blind echolocation expert.

We have previously reported that an early-blind echolocating individual (EB) showed robust occipital activation when he identified distant, silent objects based on echoes from his tongue clicks (Thaler, Arnott, & Goodale, 2011). In the present study we investigated the extent to which echolocation activation in EBs occipital cortex reflected general echolocation processing per se versus feature-specific processing. In the first experiment, echolocation audio sessions were captured with in-ear microphones in an anechoic chamber or hallway alcove as EB produced tongue clicks in front of a concave or flat object covered in aluminum foil or a cotton towel. All eight echolocation sessions (2 shapes×2 surface materials×2 environments) were then randomly presented to him during a sparse-temporal scanning fMRI session. While fMRI contrasts of chamber versus alcove-recorded echolocation stimuli underscored the importance of auditory cortex for extracting echo information, main task comparisons demonstrated a prominent role of occipital cortex in shape-specific echo processing in a manner consistent with latent, multisensory cortical specialization. Specifically, relative to surface composition judgments, shape judgments elicited greater BOLD activity in ventrolateral occipital areas and bilateral occipital pole. A second echolocation experiment involving shape judgments of objects located 20° to the left or right of straight ahead activated more rostral areas of EBs calcarine cortex relative to location judgments of those same objects and, as we previously reported, such calcarine activity was largest when the object was located in contralateral hemispace. Interestingly, other echolocating experts (i.e., a congenitally blind individual in Experiment 1, and a late blind individual in Experiment 2) did not show the same pattern of feature-specific echo-processing calcarine activity as EB, suggesting the possible significance of early visual experience and early echolocation training. Together, our findings indicate that the echolocation activation in EBs occipital cortex is feature-specific, and that these object representations appear to be organized in a topographic manner.

Attentional set modulates visual areas: an event-related potential study of attentional capture.

The present experiment offers event-related potential evidence suggesting that modulation of neural activity in the visual cortex underlies top-down attentional capture by irrelevant cues. Participants performed a covert visual search task where they identified the unique stimulus in a brief, four-location display. Targets defined uniquely by color or onset were run in separate blocks, encouraging observers to adopt different attentional sets in each block. In Experiment 1, a brief, white, abrupt-onset cue highlighted one of the locations 100 or 200 ms prior to the target display. In Experiment 2, the cue display consisted of three white and one red cues simultaneously presented at the four locations. In both experiments, participants were informed that there was no predictive relation between the location of the cue and that of the target. Reaction times were dependent on the location of the preceding cue (i.e. attention was captured), but only in those blocks where the cue shared the uniquely relevant target feature. Evoked potentials over the right hemisphere were modulated during the attention-capturing blocks just prior to the cues appearance. Additionally, the N1 wave elicited by the cue was enhanced over occipital regions during the attention-capturing blocks. These findings support the notion that attentional capture with peripheral cues is not simply reflexive but is modulated by top-down processes.