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PubMed2004

Spatial and temporal factors during processing of audiovisual speech: a PET study.

Speech perception can use not only auditory signals, but also visual information from seeing the speaker's mouth. The relative timing and relative location of auditory and visual inputs are both known to influence crossmodal integration psychologically, but previous imaging studies of audiovisual speech focused primarily on just temporal aspects. Here we used Positron Emission Tomography (PET) during audiovisual speech processing to study how temporal and spatial factors might jointly affect brain activations. In agreement with previous work, synchronous versus asynchronous audiovisual speech yielded increased activity in multisensory association areas (e.g., superior temporal sulcus [STS]), plus in some unimodal visual areas. Our orthogonal manipulation of relative stimulus position (auditory and visual stimuli presented at same location vs. opposite sides) and stimulus synchrony showed that (i) ventral occipital areas and superior temporal sulcus were unaffected by relative location; (ii) lateral and dorsal occipital areas were selectively activated for synchronous bimodal stimulation at the same external location; (iii) right inferior parietal lobule was activated for synchronous auditory and visual stimuli at different locations, that is, in the condition classically associated with the 'ventriloquism effect' (shift of perceived auditory position toward the visual location). Thus, different brain regions are involved in different aspects of audiovisual integration. While ventral areas appear more affected by audiovisual synchrony (which can influence speech identification), more dorsal areas appear to be associated with spatial multisensory interactions.

Adult↗
PubMed2004

Transient brain responses predict the temporal dynamics of sound detection in humans.

The neural events leading up to the conscious experience of stimulus events have remained elusive. Here we describe stimulation conditions under which activation in human auditory cortex can be used to predict the temporal dynamics of behavioral sound detection. Subjects were presented with auditory stimuli whose energy smoothly increased from a silent to a clearly audible level over either 1, 1.5, or 2 s. Magnetoencephalographic (MEG) recordings were carried out in the passive and active recording conditions. In the active condition, the subjects were instructed to attend to the auditory stimuli and to press a response key when these became audible. In both conditions, the stimuli elicited a prominent transient response whose emergence is unexplainable by changes in stimulus intensity alone. This transient response was larger in amplitude over the right hemisphere and in the active condition. Importantly, behavioral sound detection followed this brain activation with a constant delay of 180 ms, and further the latency variations of the brain response were directly carried over to behavioral reaction times. Thus, noninvasively measured transient events in the human auditory cortex can be used to predict accurately the temporal course of sound detection and may therefore turn out to be useful in clinical settings.

Adult↗
PubMed2005

Analysis of the spectral envelope of sounds by the human brain.

Spectral envelope is the shape of the power spectrum of sound. It is an important cue for the identification of sound sources such as voices or instruments, and particular classes of sounds such as vowels. In everyday life, sounds with similar spectral envelopes are perceived as similar: we recognize a voice or a vowel regardless of pitch and intensity variations, and we recognize the same vowel regardless of whether it is voiced (a spectral envelope applied to a harmonic series) or whispered (a spectral envelope applied to noise). In this functional magnetic resonance imaging (fMRI) experiment, we investigated the basis for analysis of spectral envelope by the human brain. Changing either the pitch or the spectral envelope of harmonic sounds produced similar activation within a bilateral network including Heschl's gyrus and adjacent cortical areas in the superior temporal lobe. Changing the spectral envelope of continuously alternating noise and harmonic sounds produced additional right-lateralized activation in superior temporal sulcus (STS). Our findings show that spectral shape is abstracted in superior temporal sulcus, suggesting that this region may have a generic role in the spectral analysis of sounds. These distinct levels of spectral analysis may represent early computational stages in a putative anteriorly directed stream for the categorization of sound.

Adult↗
PubMed2005

Listening to a walking human activates the temporal biological motion area.

A vivid perception of a moving human can be evoked when viewing a few point-lights on the joints of an invisible walker. This special visual ability for biological motion perception has been found to involve the posterior superior temporal sulcus (STSp). However, in everyday life, human motion can also be recognized using acoustic cues. In the present study, we investigated the neural substrate of human motion perception when listening to footsteps, by means of a sparse sampling functional MRI design. We first showed an auditory attentional network that shares frontal and parietal areas previously found in visual attention paradigms. Second, an activation was observed in the auditory cortex (Heschl's gyrus and planum temporale), likely to be related to low-level sound processing. Most strikingly, another activation was evidenced in a STSp region overlapping the temporal biological motion area previously reported using visual input. We thus propose that a part of the STSp region might be a supramodal area involved in human motion recognition, irrespective of the sensory modality input.

Acoustic Stimulation↗
PubMed2005

The neural basis of temporal auditory discrimination.

When two identical stimuli, such as a pair of clicks, are presented with a sufficiently long time-interval between them they are readily perceived as two separate events. However, as they are presented progressively closer together, there comes a point when the two separate stimuli are perceived as one. This phenomenon applies not only to hearing but also to other sensory modalities. Damage to the basal ganglia disturbs this type of temporal discrimination irrespective of sensory modality, suggesting a multimodal process is involved. Our aim was to study the neural substrate of auditory temporal discrimination in healthy subjects and to compare it with structures previously associated with analogous tactile temporal discrimination. During fMRI scanning, paired-clicks separated by variable inter-stimulus intervals (1-50 ms) were delivered binaurally, with different intensities delivered to each ear, yielding a lateralised auditory percept. Subjects were required (a) to report whether they heard one or two stimuli (TD: temporal discrimination); or (b) to report whether the stimuli were located on the right or left side of the head mid-line (SD: spatial discrimination); or (c) simply to detect the presence of an auditory stimulus (control task). Our results showed that both types of auditory discrimination (TD and SD) compared to simple detection activated a network of brain areas including regions of prefrontal cortex and basal ganglia. Critically, two clusters in pre-SMA and the anterior cingulate cortex were specifically activated by TD. Furthermore, these clusters overlap with regions activated for similar judgments in the tactile modality suggesting that they fulfill a multimodal function in the temporal processing of sensory events.

Acoustic Stimulation↗
PubMed2006

Auditory motion perception activates visual motion areas in early blind subjects.

We have previously shown that some visual motion areas can be specifically recruited by auditory motion processing in blindfolded sighted subjects [Poirier, C., Collignon, O., De Volder, A.G., Renier, L., Vanlierde, A., Tranduy, D., Scheiber, C., 2005. Specific activation of V5 brain area by auditory motion processing: an fMRI study. Brain Res. Cogn. Brain Res. 25, 650-658]. The present fMRI study investigated whether auditory motion processing may recruit the same brain areas in early blind subjects. The task consisted of simultaneously determining both the nature of a sound stimulus (pure tone or complex sound) and the presence or absence of its movement. When a movement was present, blind subjects had to identify its direction. Auditory motion processing, as compared to static sound processing, activated the brain network of auditory and visual motion processing classically observed in sighted subjects. Accordingly, brain areas previously considered as specific to visual motion processing could be specifically recruited in blind people by motion stimuli presented through the auditory modality. This indicates that the occipital cortex of blind people could be organized in a modular way, as in sighted people. The similarity of these results with those we previously observed in sighted subjects suggests that occipital recruitment in blind people could be mediated by the same anatomical connections as in sighted subjects.

Adolescent↗
PubMed2006

Electrical brain imaging reveals spatio-temporal dynamics of timbre perception in humans.

Timbre is a major attribute of sound perception and a key feature for the identification of sound quality. Here, we present event-related brain potentials (ERPs) obtained from sixteen healthy individuals while they discriminated complex instrumental tones (piano, trumpet, and violin) or simple sine wave tones that lack the principal features of timbre. Data analysis yielded enhanced N1 and P2 responses to instrumental tones relative to sine wave tones. Furthermore, we applied an electrical brain imaging approach using low-resolution electromagnetic tomography (LORETA) to estimate the neural sources of N1/P2 responses. Separate significance tests of instrumental vs. sine wave tones for N1 and P2 revealed distinct regions as principally governing timbre perception. In an initial stage (N1), timbre perception recruits left and right (peri-)auditory fields with an activity maximum over the right posterior Sylvian fissure (SF) and the posterior cingulate (PCC) territory. In the subsequent stage (P2), we uncovered enhanced activity in the vicinity of the entire cingulate gyrus. The involvement of extra-auditory areas in timbre perception may imply the presence of a highly associative processing level which might be generally related to musical sensations and integrates widespread medial areas of the human cortex. In summary, our results demonstrate spatio-temporally distinct stages in timbre perception which not only involve bilateral parts of the peri-auditory cortex but also medially situated regions of the human brain associated with emotional and auditory imagery functions.

Acoustic Stimulation↗
PubMed2004

Cortical motion deafness.

The extent to which the auditory system, like the visual system, processes spatial stimulus characteristics such as location and motion in separate specialized neuronal modules or in one homogeneously distributed network is unresolved. Here we present a patient with a selective deficit for the perception and discrimination of auditory motion following resection of the right anterior temporal lobe and the right posterior superior temporal gyrus (STG). Analysis of stimulus identity and location within the auditory scene remained intact. In addition, intracranial auditory evoked potentials, recorded preoperatively, revealed motion-specific responses selectively over the resected right posterior STG, and electrical cortical stimulation of this region was experienced by the patient as incoming moving sounds. Collectively, these data present a patient with cortical motion deafness, providing evidence that cortical processing of auditory motion is performed in a specialized module within the posterior STG.

Acoustic Stimulation↗
PubMed2005

Deficits in temporal-order judgments in dyslexia: evidence from diotic stimuli differing spectrally and from dichotic stimuli differing only by perceived location.

The main debate concerning dyslexia focuses on the question of whether dyslexia is a language-specific disorder or a general nervous system dysfunction manifested in deficits of temporal processing. According to the temporal-order deficit hypothesis, dyslexia manifests difficulty in discriminating the temporal order of stimuli. Evidence has usually involved testing the ability to discriminate series of phonemes or pure tones whose components are separated by very short intervals. One of the difficulties in interpreting the data is the confound of changes in the spectrum with changes in temporal order. Two experiments are reported. In the first experiment, we verified the difficulty by adult dyslexics in judging the temporal order of two tones differing in frequency and presented diotically. The second experiment was designed to isolate temporal-order judgment (TOJ) from holistic frequency-based pattern discrimination processes. We tested temporal-order judgments with 15 ms duration tones of equal frequency presented dichotically (left-right, right-left) with ISI intervals ranging from 8 to 400 ms. Dichotic temporal threshold was significantly lower for adult normal readers than for the adult dyslexics. The results support the claim that adult dyslexics have difficulty in discriminating temporal order even when no spectral changes are involved.

Acoustic Stimulation↗
PubMed2005

Loss of memory for auditory-spatial associations following unilateral medial temporal-lobe damage.

The goal of the present experiment was to determine the role of medial temporal-lobe structures in episodic memory of auditory-spatial associations. By using a two-alternative forced choice paradigm in which an association between eight different sounds and their spatial location must be recognized, learning abilities over 10 learning sessions were tested in 19 patients who had undergone a right or a left medial temporal-lobe resection for the relief of intractable seizures as well as in nine normal control participants. The data demonstrated that significant learning took place over the successive sessions for all the participants. In addition, the results showed that patients with left but not right medial temporal-lobe lesion were impaired in this learning task as compared to normal participants, suggesting the predominant implication of left medial temporal-lobe structures in auditory-spatial associative learning. The predominant role of left hemisphere structures in this memory task could be explained by a spatial categorical coding, which was enhanced by the use of eight loud-speakers. This result also suggests that the ability to store an episodic event associated with a rich spatial (or temporal) context depends on the left medial temporal-lobe structures. Thus, this finding provides an interesting parallel with data obtained in the visual modality by documenting for the first time the role of the left medial temporal-lobe in episodic learning of auditory-spatial associations.

Adult↗
PubMed2005

Processing of auditory spatial cues in human cortex: an fMRI study.

The issue of where in the human cortex coding of sound location is represented still is a matter of debate. It is unclear whether there are cortical areas that are specifically activated depending on the location of sound. Are identical or distinct cortical areas in one hemisphere involved in processing of sounds from the left and right? Also, the possibility has not been investigated so far that distinct areas have a preference for processing of central and eccentric sound locations. The present study focussed on these issues by using functional magnetic resonance imaging (fMRI). Activations evoked by left, right and central sounds were analysed separately, and contrasts were computed between these conditions. We did not find areas, which were involved in the processing of exclusively left, right or central sound positions. Large overlapping areas rather were observed for the three sound stimuli, located in the temporal, parietal and frontal cortices of both hemispheres. This result argues for the idea of a widely distributed bilateral network accessing an internal representation of the body to encode stimulus position in relation to the body median plane. However, two areas (right BA 40 and left BA 37) also were found to have preferences for sound position. In particular, BA 40 turned out to be significantly more activated by processing central positions, compared to eccentric stimuli. In line with previous findings on visual perception, the latter observation supports the assumption that the right inferior parietal cortex may be preferentially involved in the perception of central stimulus positions in relation to the body.

Acoustic Stimulation↗
PubMed2005

When space merges into language.

We present data from right brain-damaged patients, with and without spatial heminattention, which show the influence of hemispatial deficits on spoken language processing. We explored the findings of a previous study, which used an emphatic stress detection task and suggested spatial transcoding of a spoken active sentence in a 'language line'. This transcoding was impaired in its initial portion (the subject-word) when the neglect syndrome was present. By expanding the original methodology, the present study provides a deeper understanding of the level of spoken language processing involved in the heminattentional bias. To ascertain the role played by syntactic structure, active and passive sentences were compared. Sentences comprised of musical notes and of a sequence of unrelated nouns were also compared to determine whether the bias was manifest with any sequence of events (not only linguistic ones) deployed over time, and with a sequence of linguistic events not embedded in a structured syntactic frame. Results showed that heminattention exerted an influence only when a syntactically structured linguistic input (=sentence with agent of action, action and recipient of action) was processed, and that it did not interfere when a sequence of non-linguistic sounds or unrelated words was presented. Furthermore, when passing from active to passive sentences, the heminattentional bias was inverted, suggesting that heminattention primarily involves the logical subject of the sentence, which has an inverted position in passive sentences. These results strongly suggest that heminattention acts on the spatial transcoding of the deep structure of spoken language.

Adult↗
PubMed2006

Extinction of auditory stimuli in hemineglect: Space versus ear.

Unilateral extinction of auditory stimuli, a key feature of the neglect syndrome, was investigated in 15 patients with right (11), left (3) or bilateral (1) hemispheric lesions using a verbal dichotic condition, in which each ear received simultaneously one word, and a interaural-time-difference (ITD) diotic condition, in which both ears received both words lateralised by means of ITD. Additional investigations included sound localisation, visuo-spatial attention and general cognitive status. Five patients presented a significant asymmetry in the ITD diotic test, due to a decrease of left hemispace reporting but no asymmetry was found in dichotic listening. Six other patients presented a significant asymmetry in the dichotic test due to a significant decrease of left or right ear reporting, but no asymmetry in diotic listening. Ten of the above patients presented mild to severe deficits in sound localisation and eight signs of visuo-spatial neglect (three with selective asymmetry in the diotic and five in the dichotic task). Four other patients presented a significant asymmetry in both the diotic and dichotic listening tasks. Three of them presented moderate deficits in localisation and all four moderate visuo-spatial neglect. Thus, extinction for left ear and left hemispace can double dissociate, suggesting distinct underlying neural processes. Furthermore, the co-occurrence with sound localisation disturbance and with visuo-spatial hemineglect speaks in favour of the involvement of multisensory attentional representations.

Acoustic Stimulation↗
PubMed2004

Efferent connections of "posterodorsal" auditory area in the rat cortex: implications for auditory spatial processing.

We examined efferent connections of the cortical auditory field that receives thalamic afferents specifically from the suprageniculate nucleus (SG) and the dorsal division (MGD) of the medial geniculate body (MG) in the rat [Neuroscience 117 (2003) 1003]. The examined cortical region was adjacent to the caudodorsal border (4.8-7.0 mm posterior to bregma) of the primary auditory area (area Te1) and exhibited relatively late auditory response and high best frequency, compared with the caudal end of area Te1. On the basis of the location and auditory response property, the cortical region is considered identical to "posterodorsal" auditory area (PD). Injections of biocytin in PD revealed characteristic projections, which terminated in cortical areas and subcortical structures that play pivotal roles in directed attention and space processing. The most noticeable cortical terminal field appeared as dense plexuses of axons in area Oc2M, the posterior parietal cortex. Small terminal fields were scattered in area frontal cortex, area 2 that comprises the frontal eye field. The subcortical terminal fields were observed in the pontine nucleus, the nucleus of the brachium inferior colliculus, and the intermediate and deep layers of the superior colliculus. Corticostriatal projections targeted two discrete regions of the caudate putamen: the top of the middle part and the caudal end. It is noteworthy that the inferior colliculus and amygdala virtually received no projection. Corticothalamic projections terminated in the MGD, the SG, the ventral zone of the ventral division of the MG, the ventral margin of the lateral posterior nucleus (LP), and the caudodorsal part of the posterior thalamic nuclear group (Po). Large terminals were found in the MGD, SG, LP and Po besides small terminals, the major component of labeling. The results suggest that PD is an auditory area that plays an important role in spatial processing linked to directed attention and motor function. The results extend to the rat findings from nonhuman primates suggesting the existence of a posterodorsal processing stream for auditory spatial perception.

Acoustic Stimulation↗
PubMed2005

The inferior colliculus of the rat: a quantitative analysis of monaural frequency response areas.

Frequency response areas (FRAs) were measured for 237 single units in the inferior colliculus (IC) of urethane-anesthetized pigmented rats using monaural pure-tone stimulation. Based on qualitative criteria [J Neurosci 21 (2001) 7303], FRAs were classified as V-shaped in 69% of neurons, non-V-shaped in 29%, and unclassifiable in the remaining 2%. Non-V-shaped FRAs were heterogeneous, comprising a number of subtypes including narrow, closed, low- and high-tilt, multipeaked, U-shaped, mosaic and inhibitory. To complement this subjective classification, we applied quantitative measures used by others (e.g. [J Neurophysiol 84 (2000) 1012]), including the inverse slope of the upper and lower FRA borders, Q-values, and other measures of bandwidth. The results suggest that FRAs in the rat IC are best described as forming a continuous distribution among subtypes, rather than clustering into discrete categories. Moreover, there is a broad range of frequency tuning characteristics and FRA types across the entire frequency spectrum. Within this general pattern, however, there are some frequency-specific differences in FRA type distribution. The relative proportion of V-shaped FRAs was greatest at the high and low ends of the auditory range, with the highest proportion of non-V-shaped FRAs in the mid-range from 6 to 12 kHz. For most neurons with multipeaked FRAs, the peak frequencies were not harmonically related. Frequency tuning in the pigmented rat IC is generally similar to that in other species. Comparison of Q values across auditory nuclei shows little evidence that FRAs are sharpened at levels above the auditory nerve. Rather, there is a broad range of frequency tuning properties at each level.

Acoustic Stimulation↗
PubMed2005

The analysis of interaural time differences in the chick brain stem.

The brain stem auditory system of the chick has proven to be a useful model system for analyzing how the brain encodes temporal information. This paper reviews some of the work on a circuit in the brain stem that compares the timing of information coming from the two ears to determine the location of a sound source. The contralateral projection from the cochlear nucleus, nucleus magnocellularis (NM), to nucleus laminaris (NL) forms a delay line as it proceeds from medial to lateral across NL. NL neurons function like coincidence detectors in that they respond maximally when input from the two ears arrive simultaneously. This arrangement may allow NL to code sound space by the relative level of activity across the nucleus. The head anatomy of the chick allows for enhancement of the functional interaural time differences. Comparing the functional interaural time differences to the length of the neural delay line suggests that each NL can encode approximately one hemifield of sound space. Finally it is suggested that inhibitory input into the NM-NL circuit may provide a means to dynamically adjust the gain of the circuit to allow accurate coding of sound location despite changes in overall sound intensity.

Acoustic Stimulation↗
PubMed2006

Impaired verbal source monitoring in schizophrenia: an intermediate trait vulnerability marker?

Patients with schizophrenia, particularly those with positive symptoms show impaired verbal source monitoring. Specific cognitive deficits have been observed during both active and remission phases of the illness as well as in groups of unaffected first degree relatives of patients with schizophrenia. This type of schizophrenia vulnerability marker may precede the onset of frank psychotic symptoms and contribute to their developments. The aim of this study was first to determine if unaffected siblings were impaired in discriminate internal vs. external generated events when compared to their remitted schizophrenics relatives and healthy subjects. Performances of healthy subjects were then compared with results from previous studies with acute hallucinating patients, acute non-hallucinating patients and patients with resistant auditory verbal hallucinations. Compared with healthy subjects, unaffected siblings are impaired (effect size, ES=0.7), remitted or acute non-hallucinating patients are more impaired than siblings (ES=1.4); patients with verbal auditory hallucinations (acute or resistant) are even more impaired than non-hallucinating patients (ES=2.1). Our results suggest that a source monitoring deficit could be considered as an intermediate vulnerability marker of schizophrenia.

Adult↗
PubMed2005

Sounds do-able: auditory-motor transformations and the posterior temporal plane.

Accumulating evidence in humans and non-human primates implicates the posterior superior temporal plane (STP) in the processing of both auditory spatial information and vocal sounds. Such evidence is difficult to reconcile with existing accounts of the primate auditory brain. We propose that the posteromedial STP generates sequenced auditory representations by matching incoming auditory information with stored templates. These sequenced auditory representations are subsequently used to constrain motor responses. We argue for a re-assessment of the much-debated dorsal auditory pathway in terms of its generic behavioral role as an auditory "do" pathway.

Animal Communication↗