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PubMed2000

Cross-modal reorganization of human cortical functions.

Recent technological development has opened fascinating opportunities in research on cognitive functions of the human brain. For example, cortical representations of sensory functions and their reorganization, which have been studied thoroughly in animals, are far better understood in humans now than they were only a decade ago. Hemodynamic and electromagnetic studies have demonstrated that a modality-specific brain area that is totally deprived of its normal sensory input becomes responsive to stimulation of other modalities. The functional significance of this cross-modal activation was recently indicated by, for example, studies showing that the occipital cortex of the blind is activated by sound changes, when the task is to detect these changes. Moreover, trans-cranial magnetic stimulation applied to the occipital cortex of blind individuals results in distortions and omissions of letters in Braille text being read by the subject. Contrary to prevailing views, cross-modal neural reorganization might, as shown by recent results, take place even in the mature human brain.

Adult↗
PubMed2001

Spatio-temporal constraints for auditory--visual integration.

The perceptual coherence of auditory and visual information is achieved by integrative brain processes. Specialized single neurons with spatial and temporal interactions of auditory and visual stimuli have been demonstrated by several neurophysiological studies. The present, psychophysical, study investigates possible perceptual correlates of these neuronal features. Subjects had to indicate the point of subjective spatial alignment (PSSA) for a horizontally moving visual stimulus that crossed the position of a stationary sound source. Auditory and visual stimuli consisted of periodic pulses that were systematically varied in their phase relationship or repetition rate. PSSAs obtained for continuous visual stimuli served as a reference. When sound and light pulses were coincident in phase at a repetition rate of 2 Hz, PSSAs were shifted by approximately 3 degrees in a direction opposite to the movement of the visual stimulus (with respect to the reference condition). This shift markedly decreased when the temporal disparity exceeded approximately 100 ms and disappeared near phase opposition (250 ms disparity). With 4 Hz repetition rate (temporal disparity < or =125 ms), there was no significant effect of phase relationship on PSSAs, but still an approximately constant shift with respect to the reference value. Variation of the repetition rate resulted in almost constant shifts in PSSA of approximately 3 degrees between 1 and 4 Hz and a linear decrease (slope 0.27 degrees /Hz) with higher repetition rates. These results suggest a spatio-temporal 'window' for auditory-visual integration, that extends over approximately 100 ms and approximately 3 degrees : when auditory and visual stimuli are within this window, they are always perceived as spatially coincident. These psychophysical findings may be related to properties of bimodal neurons such as have been demonstrated by neurophysiological recordings in midbrain and cortex.

Adult↗
PubMed2003

Roles of the auditory midbrain and thalamus in selective phonotaxis in female gray treefrogs (Hyla versicolor).

Diencephalic and midbrain auditory nuclei are involved in the processing of auditory communication signals in anurans [Comparative Hearing: Fish and Amphibians, Springer-Verlag, New York, 1999, p. 218], but their exact roles in acoustically guided behavior, such as female phonotaxis, are unclear. To address this question, behavioral experiments were combined with lesions of dorsal thalamic nuclei and the midbrain torus semicircularis. Females were tested in two-alternative-forced-choice phonotactic experiments before and after a defined brain area was lesioned. During phonotactic tests, females had to choose between a "standard" synthetic call and one of three different variants, each of which had a single acoustic property (pulse rate, pulse rise-time, sound spectrum) that differed from the standard synthetic call. Results showed that dorsomedial thalamus lesions produced little or no effect on phonotaxis. In contrast, superficial and deep thalamus lesions, as well as lesions of the torus semicircularis, significantly decreased the number of phonotactic responses and increased the response time. Superficial thalamus lesions also abolished or reversed preferences for the standard call in the rise-time and sound spectrum tests. This effect is likely to have been caused by an imbalance in the stimulation of the thalamus by the low- and high-frequency pathways because these preferences were not affected in animals with more extensive lesions that included the superficial thalamus. Our data suggest that the torus semicircularis, but not the dorsal thalamus is crucial for phonotaxis in gravid, reproductively active females. Although dorsal thalamic nuclei seem to play a role in spectral sensitivity, they may additionally have motivational or attentional functions that contribute to achieving a state of phonotactic readiness.

Acoustic Stimulation↗
PubMed1989

Conditioned orienting (alpha) and delayed behavioral and evoked neural responses during classical conditioning.

A differentiation of short-latency (alpha) and long-latency (delayed) classically conditioned behavioral and evoked neural (hippocampal) responses was attempted. Further, facilitation and retardation of these responses were studied in an experimental design in which 10 paired conditioning sessions either preceded (CC-CO group) or followed (CO-CC group) 10 randomly unpaired presentations of conditioned stimuli (CS) and unconditioned stimuli (UCS). A 2024-ms tone (1000 Hz) was delivered directly through a miniature earphone to the left ear, eliciting an orienting head movement ('alpha' response) to the left. The unconditioned stimulus (UCS) was a direct 1024-ms stimulation of the lateral hypothalamic area overlapping the CS (delayed paradigm) so that both stimuli terminated simultaneously. The UCS elicited approach behavior and a specific head movement in each animal. The latency and the direction of the head movement were used as criteria for a differentiation of the short-latency and long-latency conditioned responses (CR). All cats showed conditioned short-latency responses. Pairing specific long-latency head movements were observed in 10 of 13 cats and 6 of them showed a long-latency CR which was a head movement to the right, while the short-latency CR on the same trials was a head movement to the left. Hippocampal (subiculum, dentate fascia and CA1) evoked responses also showed pairing specific CRs appearing as increased negativity (short-latency CR), or increased positivity (long-latency CR). Additional reversed stimulus order (backward) sessions supported an assumption of the different nature of the short-latency and long-latency CRs: the long-latency CRs showed extinction while the short-latency CRs remained. The unpaired pre-exposure to the CSs and UCSs in the CO-CC group resulted in the retarded acquisition of the behavioral responses during the subsequent paired sessions.

Animals↗
PubMed1998

Categorical discrimination of direction in frequency-modulated tones by Mongolian gerbils.

Discrimination of the direction of linearly frequency-modulated tones (FMs) was investigated in adult Mongolian gerbils (Meriones unguiculatus) using a footshock motivated shuttle box avoidance go/no go procedure. Symmetric pairs of FMs with frequency linearly increasing with time (ascending FMs) and with frequency linearly decreasing with time (descending FMs) were used as conditioned stimuli, CS+ and CS-, respectively. Stimuli were presented in randomized order in daily sessions over a period of several months. After a number of sessions, the set of conditioned stimuli was changed with respect to frequency range, steepness of modulation and duration. In experiment 1, we observed that gerbils could discriminate between the ascending 2-4 kHz CS+ and the descending 4-2 kHz CS- after a training period of 10-15 days. In experiment 2, we used FM pairs of six other frequency ranges in successive sessions (6-13; 1-2; 13-25; 0.5-1; 3 6; 0.25 0.5 kHz). We found that in the final session the last FM pair (0.25-0.5 kHz) was discriminated already after 3-4 days. Experiment 3 showed that the animals were able to discriminate five of the FM pairs learned in the separate sessions of experiment 2 (i.e. 10 different stimuli) when they were given in randomized order during one training session. In experiment 4, novel FM pairs (not heard before) and familiar FM pairs (trained in experiments 1-3) were presented within one session. It was found that, except for FMs of very short duration and small frequency range, novel FMs were discriminated according to their modulation direction. These results show that Mongolian gerbils are able to discriminate FM tones by modulation direction and, after familiarization with a number of different FM pairs, transfer the ascending-descending concept to stimuli not heard before.

Acoustic Stimulation↗
PubMed1998

Role of acoustic striae in hearing: reflexive responses to elevated sound-sources.

This report is the fourth in a series describing the results of ablation-behavior experiments directed to the ascending output of the cochlear nuclei as it is conducted centrally within the acoustic striae. This fourth report focuses on the unique physiology of the fusiform or 'output' cells of the dorsal cochlear nucleus whose axons course through the dorsal acoustic stria (DAS). Because electrophysiological studies have shown that the cues for sensing the elevation of a sound source would seem to be best analyzed by the dorsal cochlear nucleus and projected centrally via its DAS, we tested normal cats and cats deprived of DAS for their ability to orient to elevated sources of broad-band noise. For behavioral testing, we made use of reflexive or unconditioned orienting responses to elevated sound sources using a similar method to one we have used previously for azimuth testing (Thompson GC, Masterton RB. Brainstem auditory pathways involved in reflexive head orientation to sound. J Neurophysiol 1978;41:1183-1202). The results show that cats deprived of their DAS do indeed have a marked deficit in their ability to orient to an elevated sound source. Further behavioral testing indicated that this deficit is not the secondary result of an attentional or peripheral motor deficit. Although the present results do not prove that the reflexive deficit is strictly auditory in nature, the deficit is notable in that it is the only one yet known to result from a lesion of the dorsal cochlear nucleus or its central projections.

Acoustic Stimulation↗
PubMed2003

Temporal ventriloquism: crossmodal interaction on the time dimension. 1. Evidence from auditory-visual temporal order judgment.

In the well-known visual bias of auditory location (alias the ventriloquist effect), auditory and visual events presented in separate locations appear closer together, provided the presentations are synchronized. Here, we consider the possibility of the converse phenomenon: crossmodal attraction on the time dimension conditional on spatial proximity. Participants judged the order of occurrence of sound bursts and light flashes, respectively, separated in time by varying stimulus onset asynchronies (SOAs) and delivered either in the same or in different locations. Presentation was organized using randomly mixed psychophysical staircases, by which the SOA was reduced progressively until a point of uncertainty was reached. This point was reached at longer SOAs with the sounds in the same frontal location as the flashes than in different places, showing that apparent temporal separation is effectively longer in the first condition. Together with a similar one obtained recently in a case of tactile-visual discrepancy, this result supports a view in which timing and spatial layout of the inputs play to some extent inter-changeable roles in the pairing operation at the base of crossmodal interaction.

Acoustic Stimulation↗
PubMed2002

Encoding of sound motion by binaural brainstem units in a Horseshoe bat.

In order to study how and if single brainstem units respond to moving compared with stationary sounds, radially moving sound sources were presented to the bat, Rhinolophus ferrumequinum. This time-variant binaural stimulation was simulated dichotically through earphones (closed-acoustic-field for the virtual azimuth range of +/-40 degrees from the midline). Neurophysiologically recorded responses primarily showed a function of interaural intensity difference (IID) which is considered a direct correlate of the sound source's azimuth angle. However, this is only true for the stationary case. Unit's response did not remain unaffected by the dynamic stimulus cues of sound source movement (velocity and direction). Maximal discharge rate became a function of motion velocity as well as the slopes of the response profiles. Hence, coding of IID became ambiguous as, depending on the unit, the response profiles and therefore a unit's receptive field, became spatially shifted with respect to one another when the direction of the sound source movement was reversed. Shifts within the movement direction (hysteresis) as well as against it (termed here 'advance') were observed: hysteresis is typical for units with non-monotonic, stationary rate/intensity functions, whereas those units with monotonic functions predominantly show advances. Further dynamic response features in form of transient peaks and troughs, superimposed on the response profiles, were registered. It appears that the ongoing firing rate no longer represents azimuth position alone, but vigorously reproduces the dynamic cues (velocity and movement direction), too. With respect to the neural mechanisms leading to dynamic response features, it is proposed that, as long excitation and inhibition act with similar short time constants, neural activity can rapidly and faithfully follow changing IIDs. Different time constants for excitation, inhibition, facilitation, and depression may be responsible for the dynamic 'features' such as transient responses and hysteresis/advance. They may provide biologically relevant information for nocturnally hunting bats to efficiently guide their flight maneuvers.

Animals↗
PubMed2000

Human auditory-cortex mechanisms of preattentive sound discrimination.

Intracranial event-related potentials (ERPs) were recorded in neurological patients to infrequent higher-pitch 'deviant' tones and to frequent 'standard' tones when they occurred, in random order in a mixed sequence of standard and deviant tones and when they occurred in separate sequences, that is, infrequent tones alone with intervals similar to inter-deviant intervals of the mixed sequence and frequent tones alone with intervals similar to those between the standard tones of the mixed sequence. When the tones were ignored, ERPs showed three types of responses revealing three different processes involved in stimulus discrimination in the superior temporal cortex: (1) a pitch-dependent response in the primary auditory cortex; (2) an interstimulus-interval dependent response in the secondary auditory cortex; and (3) a change-detection ('mismatch') response in the auditory association cortex. When the tones were attended, ERPs to deviant and standard tones showed differences also in the basal ganglia-thalamic circuits and in the hippocampus, indicating their involvement in attentive processing of auditory stimulus changes.

Acoustic Stimulation↗
PubMed2001

Dorsolateral prefrontal cortex activation during automatic auditory duration-mismatch processing in humans: a positron emission tomography study.

This study aimed to identify the neural networks underlying automatic and active auditory deviant detection in six healthy subjects using positron emission tomography. Eight alternating blocks of standard and standard plus duration-deviant tones were presented while subjects performed a visual discrimination task. In an additional four blocks, the subjects then performed an auditory discrimination task on the deviant tones. Actively attending the deviant tones increased regional cerebral blood flow (rCBF) in the superior temporal and inferior frontal gyrus as well as in the superior and medio-frontal gyrus. When performing the visual task and presented with deviant tones, significant increase of rCBF was detected in the caudate nucleus, cerebellum, posterior cingulate, inferior frontal and pre-central gyrus thus indicating automatic extra-pyramidal processing of auditory duration deviants.

Acoustic Stimulation↗
PubMed2001

Location changes enhance hemispheric asymmetry of magnetic fields evoked by lateralized sounds in humans.

Auditory mismatch negativity, the brain's change-detection response, has been shown to be more sensitive than other early auditory cortex responses to the hemispheric specialization of speech processing. The present study used magnetoencephalography to assess hemispheric differences in cortical evoked responses during auditory spatial processing. We compared N1m to lateralized vowels presented with equal probabilities with mismatch fields (MMNm) to rare lateralized noises interspersed in a sequence of frequent midline sounds. Both N1m and MMNm dipole amplitudes were higher in the hemisphere contralaterally to the side of sound lateralization, but this effect was about four times bigger in the mismatch paradigm. Moreover, only MMNm dipoles showed shorter latencies in the hemisphere contralaterally to stimulation. Apparently stimulus changes activate specialized auditory networks more strongly than non-deviant events.

Acoustic Stimulation↗
PubMed2003

Spatial asymmetries of auditory event-synthesis in humans.

We used the mismatch negativity event-related potential to examine how spatial location and feature variation affect the capacity of the auditory system to automatically respond to pairs of rapid (180 ms apart) acoustic changes within a single tone. When a tone first deviated from a standard tone in source location and then in its duration, we found independent responses to both deviations for right but not left field stimuli. In contrast, when the first deviation was in pitch and the second in duration, only the first deviation elicited a response, regardless of presentation side. These results suggest that information from either side of space is asymmetrically processed even in a free-field, and that the extent of the temporal window of integration is not a fixed property of the auditory system.

Acoustic Stimulation↗
PubMed2003

Tracking of multiple sound sources defined by interaural time differences: brain potential evidence in humans.

Among other cues, interaural time differences (ITD) can be used to compute the location of a sound. To investigate whether ITD can be used for the preattentive detection of sounds coming from a different location than standard sounds (P=0.7, no ITD), left and right 'far' (900 micros ITD, 80 degrees excentricity) and 'near ' (300 micros ITD, 30 degrees ) deviant stimuli (each deviant P=0.075) were presented via headphones. While 15 young healthy subjects passively listened to these sequences, the mismatch negativity (MMN) of the event-related potential was obtained for the deviant sounds. MMN was reliable for all four deviants but larger for the far-lateralized sounds. Moreover, MMN was larger contralateral to the location of the deviant. This indicates a role of preattentive mechanisms in the spatial analysis of auditory scenes.

Adult↗
PubMed1997

Reduced activity in the extrastriate visual cortex of individuals with strabismic amblyopia.

In order to examine the relationship between reduced visual acuity in human strabismic amblyopia and the cortical activation pattern, we studied, by use of positron emission tomography (PET) and the H2(15)O bolus technique, changes in the regional cerebral blood flow (rCBF) induced by monocular visual stimulation of 8 individuals with this disorder. Individual amblyopic thresholds for monocular detection of the checkerboard pattern were employed as stimuli for both eyes during PET scans. Statistical analysis of subtracted images showed significant increases in rCBF (P < 0.05) by the stimulation of the sound eye localized bilaterally to Brodmann's areas (BAs) 17-19. The cortical response evoked by the amblyopic eye was significantly reduced (P < 0.05) in the ipsilateral BAs 18, 19. These results suggest that the reduction in contrast sensitivity (pattern vision) in amblyopia is coupled with deactivation in identifiable regions of occipital visual areas, including ipsilateral BAs 18,19.

Adult↗
PubMed1997

Fast preattentive processing of location: a functional basis for selective listening in humans.

Spatial separation of sound sources provides a primary cue for selecting relevant from irrelevant acoustic input. This competence for selective listening is important in every-day life, when several concurrent sound sources are simultaneously active. The present study demonstrated a temporal advantage in the preattentive processing of location information relative to frequency information. This was indicated by shorter latency of the mismatch negativity (MMN), generated by the brain's automatic detection of a sound change, to a location change than to a frequency change. Results suggest that the superior role of spatial location for selective listening may be due to faster automatic encoding of spatial information into the neural representations underlying attentional selection.

Adult↗
PubMed1998

Preattentive processing of auditory spatial information in humans.

Auditory event-related potentials were recorded from reading subjects to frequent and infrequent tones. Frequent tones presented by a loudspeaker in front of the subject were interspersed with infrequent tones delivered either by one of the symmetrically-placed lateral loudspeakers, or by both lateral loudspeakers simultaneously. This latter sound was perceived as originating from a spacious source in the direction of the central loudspeaker. A sizable mismatch negativity (MMN) and P3a were elicited by all three infrequent stimuli, suggesting that infrequent changes in the direction or perceived spaciousness of the sound source were preattentively detected. In addition, a dissociation between the MMN and P3a amplitudes was found: whereas lateral deviants elicited a larger P3a than the simultaneous left + right deviant, the MMN amplitude was approximately equal for all three deviants.

Acoustic Stimulation↗
PubMed1986

The effect of aging on the P3 component in different auditory paradigms.

Seventy-two healthy volunteers aged 24-75 years were submitted to different auditory "oddball" ERP (Event Related Potential) paradigms which included an intensity discrimination and a right/left discrimination task. In both conditions, a late positive component (P3) of the vertex potential appeared. However its latency was about 30 msec greater and its amplitude was smaller in the intensity discrimination paradigm. Moreover, P3 latency increased with age in a strictly linear fashion in the intensity discrimination paradigm, whereas in the right/left discrimination paradigm an increase of latency with age was confined to the older age-groups, which is expressed by a quadratic latency/age function. A relation was found between the subjective feeling of difficulty in performing the tasks and P3 latency in the different paradigms.

Adult↗
PubMed2000

Sound lateralization and interaural discrimination. Effects of brainstem infarcts and multiple sclerosis lesions.

Subjects with brainstem lesions due to either an infarct or multiple sclerosis (MS) underwent two types of binaural testing (lateralization testing and interaural discrimination) for three types of sounds (clicks and high and low frequency narrow-band noise) with two kinds of interaural differences (level and time). Two major types of abnormalities were revealed in the lateralization performances: perception of all stimuli, regardless of interaural differences (time and/or level) in the center of the head (center-oriented), or lateralization of all stimuli to one side or the other of the head (side-oriented). Similar patterns of abnormal lateralization (center-oriented and side-oriented) occurred for MS and stroke patients. A subject's pattern of abnormal lateralization testing was the same regardless of the type of stimulus or type of interaural disparity. Lateralization testing was a more sensitive test than interaural discrimination testing for both types of subjects. Magnetic resonance image (MRI) scanning in three orthogonal planes of the brainstem was used to detect lesions. A semi-automated algorithm superimposed the auditory pathway onto each MRI section. Whenever a lesion overlapped the auditory pathway, some binaural performance was abnormal and vice versa. Given a lateralization test abnormality, whether the pattern was center-oriented or side-oriented was mainly determined by lesion site. Center-oriented performance was principally associated with caudal pontine lesions and side-oriented performance with lesions rostral to the superior olivary complex. For lesions restricted to the lateral lemniscus and/or inferior colliculus, whether unilateral or bilateral, just noticeable differences (JNDs) were nearly always abnormal, but for caudal pontine lesions JNDs could be normal or abnormal. MS subjects were more sensitive to interaural time delays than interaural level differences particularly for caudal pontine lesions, while stroke patients showed no differential sensitivity to the two kinds of interaural differences. These results suggest that neural processing of binaural stimuli is multilevel and begins with independent interaural time and level analyzers in the caudal pons.

Adult↗