Brain mechanisms implicated in the preattentive categorization of speech sounds revealed using FMRI and a short-interval habituation trial paradigm

A hallmark of categorical perception is better discrimination of stimulus tokens from 2 different categories compared with token pairs that are equally dissimilar but drawn from the same category. This effect is well studied in speech perception and represents an important characteristic of how the phonetic form of speech is processed. We investigated the brain mechanisms of categorical perception of stop consonants using functional magnetic resonance imaging and a passive short-interval habituation trial design (Zevin and McCandliss 2005). The paradigm takes advantage of neural adaptation effects to identify specific regions sensitive to an oddball stimulus presented in the context of a repeated item. These effects were compared for changes in stimulus characteristics that result in either a between-category (phonetic and acoustic) or a within-category (acoustic only) stimulus shift. Significantly greater activation for between-category than within-category stimuli was observed in left superior sulcus and middle temporal gyrus as well as in inferior parietal cortex. In contrast, only a subcortical region specifically responded to within-category changes. The data suggest that these habituation effects are due to the unattended detection of a phonetic stimulus feature.     

Functional anatomy of biological motion perception in posterior temporal cortex: an FMRI study of eye, mouth and hand movements

Passive viewing of biological motion engages extensive regions of the posterior temporal-occipital cortex in humans, particularly within and nearby the superior temporal sulcus (STS). Relatively little is known about the functional specificity of this area. Some recent studies have emphasized the perceived intentionality of the motion as a potential organizing principle, while others have suggested the existence of a somatotopy based upon the limb perceived in motion. Here we conducted an event-related functional magnetic resonance imaging experiment to compare activity elicited by movement of the eyes, mouth or hand. Each motion evoked robust activation in the right posterior temporal-occipital cortex. While there was substantial overlap of the activation maps in this region, the spatial distribution of hemodynamic response amplitudes differentiated the movements. Mouth movements elicited activity along the mid-posterior STS while eye movements elicited activity in more superior and posterior portions of the right posterior STS region. Hand movements activated more inferior and posterior portions of the STS region within the posterior continuing branch of the STS. Hand-evoked activity also extended into the inferior temporal, middle occipital and lingual gyri. This topography may, in part, reflect the role of particular body motions in different functional activities.     

Listening to narrative speech after aphasic stroke: the role of the left anterior temporal lobe

The dorsal bank of the primate superior temporal sulcus (STS) is a polysensory area with rich connections to unimodal sensory association cortices. These include auditory projections that process complex acoustic information, including conspecific vocalizations. We investigated whether an extensive left posterior temporal (Wernicke's area) lesion, which included destruction of early auditory cortex, may contribute to impaired spoken narrative comprehension as a consequence of reduced function in the anterior STS, a region not included within the boundary of infarction. Listening to narratives in normal subjects activated the posterior-anterior extent of the left STS, as far forward as the temporal pole. The presence of a Wernicke's area lesion was associated with both impaired sentence comprehension and a reduced physiological response to heard narratives in the intact anterior left STS when compared to aphasic patients without temporal lobe damage and normal controls. Thus, in addition to the loss of language function in left posterior temporal cortex as the direct result of infarction, posterior ablation that includes primary and early association auditory cortex impairs language function in the intact anterior left temporal lobe. The implication is that clinical studies of language on stroke patients have underestimated the role of left anterior temporal cortex in comprehension of narrative speech.     

Neural correlates of coherent audiovisual motion perception

Real-life moving objects are often detected by multisensory cues. We investigated the cortical activity associated with coherent visual motion perception in the presence of a stationary or moving auditory noise source using functional magnetic resonance imaging. Twelve subjects judged episodes of 5-s random-dot motion containing either no (0%) or abundant (16%) coherent direction information. Auditory noise was presented with the displayed visual motion that was moving in phase, was moving out-of-phase, or was stationary. Subjects judged whether visual coherent motion was present, and if so, whether the auditory noise source was moving in phase, was moving out-of-phase, or was not moving. Performance was greatest for a moving sound source that was in phase with the visual coherent dot motion compared with when it was in antiphase. A random-effects analysis revealed that auditory motion activated extended regions in both cerebral hemispheres in the superior temporal gyrus (STG), with a right-hemispheric preponderance. Combined audiovisual motion led to activation clusters in the STG, the supramarginal gyrus, the superior parietal lobule, and the cerebellum. The size of the activated regions was substantially larger than that evoked by either visual or auditory motion alone. The congruent audiovisual motion evoked the most extensive activation pattern, exhibiting several exclusively activated subregions.     

Hemispheric specialization for processing auditory nonspeech stimuli

The left hemisphere specialization for speech perception might arise from asymmetries at more basic levels of auditory processing. In particular, it has been suggested that differences in "temporal" and "spectral" processing exist between the hemispheres. Here we used functional magnetic resonance imaging to test this hypothesis further. Fourteen healthy volunteers listened to sequences of alternating pure tones that varied in the temporal and spectral domains. Increased temporal variation was associated with activation in Heschl's gyrus (HG) bilaterally, whereas increased spectral variation activated the superior temporal gyrus (STG) bilaterally and right posterior superior temporal sulcus (STS). Responses to increased temporal variation were lateralized to the left hemisphere; this left lateralization was greater in posteromedial HG, which is presumed to correspond to the primary auditory cortex. Responses to increased spectral variation were lateralized to the right hemisphere specifically in the anterior STG and posterior STS. These findings are consistent with the notion that the hemispheres are differentially specialized for processing auditory stimuli even in the absence of linguistic information.     

Neural substrates of real and imagined sensorimotor coordination

Much debate in the behavioral literature focuses on the relative contribution of motor and perceptual processes in mediating coordinative stability. To a large degree, such debate has proceeded independently of what is going on in the brain. Here, using blood oxygen level-dependent measures of neural activation, we compare physically executed and imagined rhythmic coordination in order to better assess the relative contribution of hypothesized neuromusculoskeletal mechanisms in modulating behavioral stability. The executed tasks were to coordinate index finger to thumb opposition movements of the right hand with an auditory metronome in either a synchronized (on the beat) or syncopated (off the beat) pattern. Imagination involved the same tasks, except without physical movement. Thus, the sensory stimulus and coordination constraints were the same in both physical and imagination tasks, but the motoric requirements were not. Results showed that neural differences between executed synchronization and syncopation found in premotor cortex, supplementary motor area, basal ganglia and lateral cerebellum persist even when the coordinative patterns were only imagined. Neural indices reflecting behavioral stability were not abolished by the absence of overt movement suggesting that coordination phenomena are not exclusively rooted in purely motoric constraints. On the other hand, activity in the superior temporal gyrus was modulated by both the presence of movement and the nature of the coordination, attesting to the intimacy between perceptual and motoric processes in coordination dynamics.     

Voice recognition and cross-modal responses to familiar speakers' voices in prosopagnosia

Recognizing the voices of people we know does not only activate "voice areas" in the temporal lobe but also extraauditory areas including the fusiform "face area" (FFA). This cross-modal effect could reflect that individual face and voice information become specifically associated when becoming acquainted with a person. Here, we addressed whether the ability to have individual face representations 1) plays a role in voice recognition and 2) is required to observe cross-modal responses to voices in face areas. We compared speaker recognition performance and neuroimaging responses during the processing of familiar and nonfamiliar speakers' voices in a developmental prosopagnosic subject (SO) with the respective findings obtained in a group of 9 control subjects. Despite scoring worse than controls on recognition of familiar speakers' voices, SO had normal cross-modal responses in the FFA and normal connectivity between FFA and the voice regions. However, she had reduced activations in areas that usually respond to familiarity with people. An indication for the malfunctioning of her FFA was reduced connectivity of the FFA to a subset of these supramodal areas. In combination these data suggest that 1) voice recognition benefits from the ability to process faces at an individual level and 2) cross-modal association of voices and faces in the brain is achieved by a sensory binding and does not depend on a top-down mechanism subsequent to successful person recognition.     

Multiple stages of auditory speech perception reflected in event-related FMRI

Speech processing in auditory cortex and beyond is a remarkable yet poorly understood faculty of the listening brain. Here we show that stop consonants, as the most transient constituents of speech, are sufficient to involve speech perception circuits in the human superior temporal cortex. Left anterolateral superior temporal cortex showed a stronger response in blood oxygenation level-dependent functional magnetic resonance imaging (fMRI) to intelligible consonantal bursts compared with incomprehensible control sounds matched for spectrotemporal complexity. Simultaneously, the left posterior superior temporal plane (including planum temporale [PT]) exhibited a noncategorical responsivity to complex stimulus acoustics across all trials, showing no preference for intelligible speech sounds. Multistage hierarchical processing of speech sounds is thus revealed with fMRI, providing evidence for a role of the PT in the fundamental stages of the acoustic analysis of complex sounds, including speech.     

Face configuration processing in the human brain: the role of symmetry

Symmetry is an important cue in face perception. We manipulated symmetry and other configurational variables to study their role in face processing in the human brain. We employed 2 types of symmetry: image symmetry (where one part of the image is defined as the mirrored transform of the other part about an axis) and object symmetry (where the spatial relationships among the image components are interpreted as parts of a symmetric 3-dimensional object). We compared blood oxygenation level dependent responses in healthy human observers for upright front-view faces with responses to different symmetry-controlled images. The cortical areas activated by the face images, relative to Fourier-matched scrambled images, were the fusiform (FFA) and occipital (OFA) face areas, the middle occipital gyri (MOG), and areas around the superior temporal and intraoccipital sulci (IOS). Contrasting faces and their image-symmetric scrambled versions showed a similar activation pattern except in the right OFA, suggesting an involvement in facial symmetry processing. The upright versus inverted faces (with the same image symmetry but unfamiliar object identity) showed robust differential activation in the FFA, OFA, MOG, IOS, and precuneus. The response to frontal-view versus 3/4-view faces (having the same object symmetry but disrupted image symmetry) showed little differential activation in the FFA or the OFA but strong responses in the MOG and IOS, suggesting that face processing in the FFA and the OFA is holistic and viewpoint invariant.     

Neural correlates of rule-based and information-integration visual category learning

An emerging theory of the neurobiology of category learning postulates that there are separate neural systems supporting the learning of categories based on verbalizeable rules (RB) or through implicit information integration (II). The medial temporal lobe (MTL) is thought to play a crucial role in successful RB categorization, whereas the posterior regions of the caudate are hypothesized to support II categorization. Functional neuroimaging was used to assess activity in these systems during category-learning tasks with category structures designed to afford either RB or II learning. Successful RB categorization was associated with relatively increased activity in the anterior MTL. Successful II categorization was associated with increased activity in the caudate body. The dissociation observed with neuroimaging is consistent with the roles of these systems in memory and dissociations reported in patient populations. Convergent evidence from these approaches consistently reinforces the idea of multiple neural systems supporting category learning.     

Brain activations during judgments of positive self-conscious emotion and positive basic emotion: pride and joy

We aimed to investigate the neural correlates associated with judgments of a positive self-conscious emotion, pride, and elucidate the difference between pride and a basic positive emotion, joy, at the neural basis level using functional magnetic resonance imaging. Study of the neural basis associated with pride might contribute to a better understanding of the pride-related behaviors observed in neuropsychiatric disorders. Sixteen healthy volunteers were studied. The participants read sentences expressing joy or pride contents during the scans. Pride conditions activated the right posterior superior temporal sulcus and left temporal pole, the regions implicated in the neural substrate of social cognition or theory of mind. However, against our prediction, we did not find brain activation in the medial prefrontal cortex, a region responsible for inferring others' intention or self-reflection. Joy condition produced activations in the ventral striatum and insula/operculum, the key nodes of processing of hedonic or appetitive stimuli. Our results support the idea that pride is a self-conscious emotion, requiring the ability to detect the intention of others. At the same time, judgment of pride might require less self-reflection compared with those of negative self-conscious emotions such as guilt or embarrassment.     

Kinesthetic working memory and action control within the dorsal stream

There is wide agreement that the "dorsal (action) stream" processes visual information for movement control. However, movements depend not only on vision but also on tactile and kinesthetic information (=haptics). Using functional magnetic resonance imaging, the present study investigates to what extent networks within the dorsal stream are also utilized for kinesthetic action control and whether they are also involved in kinesthetic working memory. Fourteen blindfolded participants performed a delayed-recognition task in which right-handed movements had to be encoded, maintained, and later recognized without any visual feedback. Encoding of hand movements activated somatosensory areas, superior parietal lobe (dorsodorsal stream), anterior intraparietal sulcus (aIPS) and adjoining areas (ventrodorsal stream), premotor cortex, and occipitotemporal cortex (ventral stream). Short-term maintenance of kinesthetic information elicited load-dependent activity in the aIPS and adjacent anterior portion of the superior parietal lobe (ventrodorsal stream) of the left hemisphere. We propose that the action representation system of the dorsodorsal and ventrodorsal stream is utilized not only for visual but also for kinesthetic action control. Moreover, the present findings demonstrate that networks within the ventrodorsal stream, in particular the left aIPS and closely adjacent areas, are also engaged in working memory maintenance of kinesthetic information.     

V3A processes contour curvature as a trackable feature for the perception of rotational motion

Contour curvature (CC) is a vital cue for the analysis of both form and motion. Using functional magnetic resonance imaging, we localized the neural correlates of CC for the processing and perception of rotational motion. We found that the blood oxygen level-dependent signal in retinotopic area V3A and possibly also lateral occipital cortex (LOC) varied parametrically with the degree of CC. Control experiments ruled out the possibility that these modulations resulted from either changes in the area of the stimuli, the velocity with which contour elements were actually translating, or perceived angular velocity. We conclude that neurons within V3A and perhaps also LOC process continuously moving CC as a trackable feature. These data are consistent with the hypothesis that V3A contains neural populations that process trackable form features such as CC, not to solve the "ventral problem" of determining object shape but in order to solve the "dorsal problem" of what is going where.     

Auditory evoked potential patterns to voiced and voiceless speech sounds in adult developmental dyslexics with persistent deficits

Auditory evoked potentials (AEPs) were recorded from eight developmental dyslexic adults with persistent reading, spelling and phonological deficits, and 10 non-dyslexic controls to voiced (/ba/) and voiceless (/pa/) consonant-vowel syllables. Consistent with previous data, non-dyslexics coded these stimuli differentially according to the temporal cues that form the basis of the voiced/voiceless contrast: AEPs had time-locked components with latencies that were determined by the temporal structure of the stimuli. Dyslexics were characterized by one of two electrophysiological patterns: AEP pattern I dyslexics demonstrated a differential coding of stimuli on the basis of some temporal cues, but with an atypically large number of components and a considerable delay in AEP termination time; AEP pattern II dyslexics demonstrated no clear differential coding of stimuli on the basis of temporal cues. These data reveal the presence of anomalies in cortical auditory processing which could underlie persistent perceptual and linguistic impairments in some developmental dyslexics. Furthermore, scalp AEP distribution maps showing the difference observed between /ba/ and /pa/ activity over time suggest that the regions implicated in the processing of crucial time-related acoustic cues were not systematically lateralized to the left hemisphere like they were for non-dyslexics. These findings may be conducive to a better understanding and treatment of perceptual dysfunctions in developmental language disorders.     

Interhemispheric transmission of visuomotor information for motor implementation

Using transcranial magnetic stimulation (TMS), we addressed the contribution of both hemispheres to the visuomotor control of each hand. The subjects had to press one of two buttons as quickly as possible after the go-signal. A precue preceding this conveyed full, partial or no advance information (hand and/or button), such that reaction time (RT) shortened with increasing amount of information. We gave TMS over each hemisphere at various time intervals (100-350 ms) after the go-signal and before the expected onset of response, and measured its effect on RT, movement time (MT) and error rate. At short intervals (100-200 ms), left hemisphere TMS delayed RT and prolonged MT of both hands, while right hemisphere TMS delayed RT only of the right hand, without affecting error rates. At long intervals (250-350 ms), TMS produced slightly more pronounced RT delays of the contralateral hand. RT was delayed more if the precues were less informative. The results suggest the importance of interhemispheric transmission of visuomotor information for motor implementation. The right hemisphere may play a role mainly in calculating target and effector information, determining RT, while the left hemisphere may play a role in elaborating the motor program and determining MT.     

Working memory specific activity in auditory cortex: potential correlates of sequential processing and maintenance

Working memory (WM) tasks involve several interrelated processes during which past information must be transiently maintained, recalled, and compared with test items according to previously instructed rules. It is not clear whether the rule-specific comparisons of perceptual with memorized items are only performed in previously identified frontal and parietal WM areas or whether these areas orchestrate such comparisons by feedback to sensory cortex. We tested the latter hypothesis by focusing on auditory cortex (AC) areas with low-noise functional magnetic resonance imaging in a 2-back WM task involving frequency-modulated (FM) tones. The control condition was a 0-back task on the same stimuli. Analysis of the group data identified an area on right planum temporale equally activated by both tasks and an area on the left planum temporale specifically involved in the 2-back task. A region of interest analysis in each individual revealed that activation on the left planum temporale in the 2-back task positively correlated with the task performance of the subjects. This strongly suggests a prominent role of the AC in 2-back WM tasks. In conjunction with previous findings on FM processing, the left lateralized effect presumably reflects the complex sequential processing demand of the 2-back matching to sample task.     

Hemispheric asymmetries for different components of global/local attention occur in distinct temporo-parietal loci

Data from brain-damaged and neurologically intact populations indicate hemispheric asymmetries in the temporo-parietal cortex for discriminating an object's global form (e.g. the overall shape of a bicycle) versus its local parts (e.g. the spokes in a bicycle tire). However, it is not yet clear whether such asymmetries reflect processes that (i) bias attention toward upcoming global versus local stimuli and/or (ii) attend/identify global versus local stimuli after they are presented. To investigate these possibilities, we asked sixteen healthy participants to perform a cued global/local attention task while their brain activity was recorded using event-related functional magnetic resonance imaging (fMRI). The results indicated a novel double dissociation. Hemispheric asymmetries for deploying attention toward expected global versus local object features were specific to the intraparietal sulcus (iPs). However, hemispheric asymmetries for identifying global versus local features after they were presented were specific to the inferior parietal lobe/superior temporal gyrus (IPL/STG). This double dissociation provides the first direct evidence that hemispheric asymmetries associated with different components of global/local attention occur in distinct temporo-parietal loci. Furthermore, it parallels an analogous dissociation reported in a recent fMRI study of spatial orienting, suggesting that global/local attention and spatial attention might rely on similar cognitive/neural mechanisms.     

Patient perception of wound photography

The objectives of this study were to provide an assessment of photographic documentation of the wound from the patients' perspective and to evaluate whether this could improve patients' understanding of and involvement in their wound care. Our results revealed that most patients visiting the wound care clinic have difficult‐to‐see wounds (86%). Only 20% of patients monitor their wounds and instead rely on clinic or nurse visits to track the healing progress. There was a significant association between patients' ability to see their wound and their subsequent memory of the wound's appearance. This was especially true for patients who had recently begun visiting the wound care clinic. This relationship was not present in patients who had visited the clinic for 3 or more years. Patients reported that the inability to see their wounds resulted in feeling a loss of autonomy. The majority of patients reported that photographing their wounds would help them to track the wound progress (81%) and would afford them more involvement in their own care (58%). This study provides a current representation of wound photography from the patients' perspective and reveals that it can motivate patients to become more involved in the management of their wounds – particularly for patients with difficult‐to‐see wounds.     

Hearing lips and seeing voices: how cortical areas supporting speech production mediate audiovisual speech perception

Observing a speaker's mouth profoundly influences speech perception. For example, listeners perceive an "illusory" "ta" when the video of a face producing /ka/ is dubbed onto an audio /pa/. Here, we show how cortical areas supporting speech production mediate this illusory percept and audiovisual (AV) speech perception more generally. Specifically, cortical activity during AV speech perception occurs in many of the same areas that are active during speech production. We find that different perceptions of the same syllable and the perception of different syllables are associated with different distributions of activity in frontal motor areas involved in speech production. Activity patterns in these frontal motor areas resulting from the illusory "ta" percept are more similar to the activity patterns evoked by AV(/ta/) than they are to patterns evoked by AV(/pa/) or AV(/ka/). In contrast to the activity in frontal motor areas, stimulus-evoked activity for the illusory "ta" in auditory and somatosensory areas and visual areas initially resembles activity evoked by AV(/pa/) and AV(/ka/), respectively. Ultimately, though, activity in these regions comes to resemble activity evoked by AV(/ta/). Together, these results suggest that AV speech elicits in the listener a motor plan for the production of the phoneme that the speaker might have been attempting to produce, and that feedback in the form of efference copy from the motor system ultimately influences the phonetic interpretation.     

Volume of left Heschl's Gyrus and linguistic pitch learning

Research on the contributions of the human nervous system to language processing and learning has generally been focused on the association regions of the brain without considering the possible contribution of primary and adjacent sensory areas. We report a study examining the relationship between the anatomy of Heschl's Gyrus (HG), which includes predominately primary auditory areas and is often found to be associated with nonlinguistic pitch processing and language learning. Unlike English, most languages of the world use pitch patterns to signal word meaning. In the present study, native English-speaking adult subjects learned to incorporate foreign pitch patterns in word identification. Subjects who were less successful in learning showed a smaller HG volume on the left (especially gray matter volume), but not on the right, relative to learners who were successful. These results suggest that HG, typically shown to be associated with the processing of acoustic cues in nonspeech processing, is also involved in speech learning. These results also suggest that primary auditory regions may be important for encoding basic acoustic cues during the course of spoken language learning.