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Abstract

Functional MR imaging (fMRI) is being used increasingly to explore the human central auditory system. The considerable background noise produced by echo-planar imaging (EPI) and other fMRI sequences, however, interferes in an unpredictable way with the experimental stimuli. Several approaches exist to overcome this problem. Each has its advantages and disadvantages. These different approaches allow researchers to tailor the experimental designs to specific research questions. Recent studies have yielded significant information about human auditory function. Compared with other sensory systems such as the visual system, the auditory database still is relatively small. It is expected that novel methodologic approaches will stimulate scientific exploration of auditory processing and eventually lead to clinically meaningful applications of auditory fMRI.

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... Because real sound sources (loudspeakers) can not be used inside the headcoil of an fMRI scanner, virtual sound sources were presented over headphones. Normal (electrodynamic ⁄ electrostatic, circum ⁄ supra-aural) headphones may not be used with magnetic brain imaging techniques [fMRI, magnetoencephalography (MEG)], as they all create spurious electric and magnetic fields causing artefacts to the neuroimaging data (Hytönen, 2005). In the case of electrodynamic headphones the sound stimuli themselves, including the embodied subtle spectral localization cues, would be distorted due to the magnetic fields in fMRI scanner ( Baumgart et al., 1999). ...
... The present experiment applied custom-made high-quality inserttype headphones (UD ADU2a, Unides Design, Helsinki, Finland), with plastic tubes conveying the sound stimulus over air into the subject's ears, thus avoiding any magnetic or electric disturbance to the fMRI scanner (empirically verified in Hytönen, 2005). The complex acoustics and perception involved with insert-type headphones is investigated in detail in Riederer & Niska (2002) and Riederer (2005). ...
... Piezoelectric insert headphones, which are commonly used in MEG experiments, yield a very limited and uneven frequency response (about 0.1–3 kHz) with notable harmonic distortions non-suitable for realistic sound reproduction. The present experiment applied custom-made high-quality inserttype headphones (UD ADU2a, Unides Design, Helsinki, Finland), with plastic tubes conveying the sound stimulus over air into the subject's ears, thus avoiding any magnetic or electric disturbance to the fMRI scanner (empirically verified in Hytönen, 2005). The complex acoustics and perception involved with insert-type headphones is investigated in detail in Riederer & Niska (2002) and Riederer (2005). ...
Article
This functional magnetic resonance imaging study was focused on the neural substrates underlying human auditory space perception. In order to present natural-like sound locations to the subjects, acoustic stimuli convolved with individual head-related transfer functions were used. Activation foci, as revealed by analyses of contrasts and interactions between sound locations, formed a complex network, including anterior and posterior regions of temporal lobe, posterior parietal cortex, dorsolateral prefrontal cortex and inferior frontal cortex. The distinct topography of this network was the result of different patterns of activation and deactivation, depending on sound location, in the respective voxels. These patterns suggested different levels of complexity in processing of auditory spatial information, starting with simple left/right discrimination in the regions surrounding the primary auditory cortex, while the integration of information on hemispace and eccentricity of sound may take place at later stages. Activations were identified as being located in regions assigned to both the dorsal and ventral auditory cortical streams, that are assumed to be preferably concerned with analysis of spatial and non-spatial sound features, respectively. The finding of activations also in the ventral stream could, on the one hand, reflect the well-known functional duality of auditory spectral analysis, that is, the concurrent extraction of information based on location (due to the spectrotemporal distortions caused by head and pinnae) and spectral characteristics of a sound source. On the other hand, this result may suggest the existence of shared neural networks, performing analyses of auditory 'higher-order' cues for both localization and identification of sound sources.
... In humans, communication through a learned spoken language provides an opportunity to design auditory stimulation paradigms ranging from simple sound stimulation to word or sentence recognition, interpretation, and memorization. The elicited neural responses are a combination of complex hierarchical sensory processes possibly influenced by previous experience (Bernal and Altman, 2001; Seifritz et al., 2001). Introduction of an animal model that evolved a similar type of communication would provide new opportunities for the validation of BOLD responses observed during auditory stimulation in humans. ...
Article
Auditory fMRI in humans has recently received increasing attention from cognitive neuroscientists as a tool to understand mental processing of learned acoustic sequences and analyzing speech recognition and development of musical skills. The present study introduces this tool in a well-documented animal model for vocal learning, the songbird, and provides fundamental insight in the main technical issues associated with auditory fMRI in these songbirds. Stimulation protocols with various listening tasks lead to appropriate activation of successive relays in the songbirds' auditory pathway. The elicited BOLD response is also region and stimulus specific, and its temporal aspects provide accurate measures of the changes in brain physiology induced by the acoustic stimuli. Extensive repetition of an identical stimulus does not lead to habituation of the response in the primary or secondary telencephalic auditory regions of anesthetized subjects. The BOLD signal intensity changes during a stimulation and subsequent rest period have a very specific time course which shows a remarkable resemblance to auditory evoked BOLD responses commonly observed in human subjects. This observation indicates that auditory fMRI in the songbird may establish a link between auditory related neuro-imaging studies done in humans and the large body of neuro-ethological research on song learning and neuro-plasticity performed in songbirds.
... Dorsal view of the brain stem with superimposed schematic organization of subcortical relay stations of the ascending auditory system. Reprinted with permission fromSeifritz E et al. (2001) Auditory system: functional magnetic resonance imaging. Neuroimaging Clinics of North America 11( ...
... Given the favourable tracer logistics in terms of brain uptake (for example, about 20-30 min for 18 F-FDG), most of the patient studies do not have to be discarded after injection, unless the post-injection PSG-monitoring indicates this within the tracer-specific timeframe. Similar constraints with fMRI may be found, e.g., in auditory stimulation studies (Seifritz et al., 2001), and in studies on deep relaxation and trance induction (e.g., Halsband, 2004). ...
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Article
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Neurons in the superior temporal gyrus of anesthetized rhesus monkeys were exposed to complex acoustic stimuli. Bandpassed noise bursts with defined center frequencies evoked responses that were greatly enhanced over those evoked by pure tones. This finding led to the discovery of at least one new cochleotopic area in the lateral belt of the nonprimary auditory cortex. The best center frequencies of neurons varied along a rostrocaudal axis, and the best bandwidths of the noise bursts varied along a mediolateral axis. When digitized monkey calls were used as stimuli, many neurons showed a preference for some calls over others. Manipulation of the calls' frequency structure and playback of separate components revealed different types of spectral integration. The lateral areas of the monkey auditory cortex appear to be part of a hierarchical sequence in which neurons prefer increasingly complex stimuli and may form an important stage in the preprocessing of communication sounds.
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This study investigated the cortical response to hearing threat-related and neutral words using functional magnetic resonance imaging (fMRI) in 16 coronal planes. Right-handed volunteers listened to (i) neutral words alternating with no words as the control condition, and (ii) neutral words alternating with threat-related words as the experimental condition. Threat-related words compared to neutral words activated left posterior cingulate gyrus in eight of 10 subjects with activation most prominent in the retrosplenial region. Patterns of activation produced by neutral words compared to no words included bilateral temporal and frontal regions but not posterior cingulate. The retrosplenial cingulate region has recently been implicated in episodic memory processes. We discuss the possible role of the posterior cingulate cortex in processes involving emotion and memory and in anxiety disorders.
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An event-related protocol was designed to permit auditory fMRI studies minimally affected by the echo-planar noise artifact; a long time interval (TR = 10 s) between each cerebral volume acquisition was combined with stroboscopic data acquisition, and event-related curves were reconstructed with a 1-s resolution. The cerebral hemodynamic-response time course to a target auditory stimulus was measured in five individual subjects using this method. Clear bell-shaped event-related responses were observed bilaterally in all individuals in primary auditory cortex (A1) as well as in laterally extending secondary cortical fields. Group-average event-related curves attained their maxima (0.5–0.7%) 3 s after stimulus onset in A1 (4 s for more anterior and lateral regions of auditory cortex), and signal had returned to near-baseline level 6 s after stimulus onset. The stroboscopic event-related method appeared effective in minimizing effects of the interaction between scanning noise and experimental auditory stimulation; it adds useful temporal information to the spatial resolution afforded by fMRI in studies of human auditory function, while allowing presentation of auditory stimuli on a silent background.
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Apart from being a common feature of mental illness, auditory hallucinations provide an intriguing model for the study of internally generated sensory perceptions that are attributed to external sources. Until now, the knowledge about the cortical network that supports such hallucinations has been restricted by methodological limitations. Here, we describe an experiment with paranoid schizophrenic patients whose on- and offset of auditory hallucinations could be monitored within one functional magnetic resonance imaging (fMRI) session. We demonstrate an increase of the blood oxygen level–dependent (BOLD) signal in Heschl’s gyrus during the patients’ hallucinations. Our results provide direct evidence of the involvement of primary auditory areas in auditory verbal hallucinations and establish novel constraints for psychopathological models.
Article
Gross inspection of 30 human brains (60 hemispheres) from patients without hearing impairment revealed the following configurations of gyri: one on left and two on right (14 of 30); two on left and two on right (11 of 30); one on left and one on right (3 of 30); two on left and one on right (1 of 30); and two on left and three on right (1 of 30). The last group has since been extended to include all combinations other than the first four and the proportions of these combinations appear to persist in larger tallies. Thirteen of the 30 brains displayed double gyri on the left while 25 had double gyri on the right. Seventeen of the 30 brains possessed a single gyrus on the left and four specimens had a single gyrus on the right. In general, the left side has a tendency to fewer gyral markings. The average cortical surface area on the left is 1198 mm2 compared to 1380 on the right.
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In a series of experiments on New World and Old World monkeys, architectonic features of auditory cortex were related to tone frequency maps and patterns of connections to generate and evaluate theories of cortical organization. The results suggest that cortical processing of auditory information involves a number of functionally distinct fields that can be broadly grouped into four or more levels of processing. At the first level, there are three primary-like areas, each with a discrete pattern of tonotopic organization, koniocortical histological features, and direct inputs from the ventral division of the medial geniculate complex. These three core areas are interconnected and project to a narrow surrounding belt of perhaps seven areas which receive thalamic input from the major divisions of the medial geniculate complex, the suprageniculate/limitans complex, and the medial pulvinar. The belt areas connect with a lateral parabelt region of two or more fields that are almost devoid of direct connections with the core and the ventral division of the medial geniculate complex. The parabelt fields connect with more distant cortex in the superior temporal gyrus, superior temporal sulcus, and prefrontal cortex. The results indicate that auditory processing involves 15 or more cortical areas, each of which is interconnected with a number of other fields, especially adjoining fields of the same level.
Article
We present an approach to characterizing the differences among event-related hemodynamic responses in functional magnetic resonance imaging that are evoked by different sorts of stimuli. This approach is predicated on a linear convolution model and standard inferential statistics as employed by statistical parametric mapping. In particular we model evoked responses, and their differences, in terms of basis functions of the peri-stimulus time. This facilitates a characterization of the temporal response profiles that has a high effective temporal resolution relative to the repetition time. To demonstrate the technique we examined differential responses to visually presented words that had been seen prior to scanning or that were novel. The form of these differences involved both the magnitude and the latency of the response components. In this paper we focus on bilateral ventrolateral prefrontal responses that show deactivations for previously seen words and activations for novel words.
Article
A method is introduced by which brain activation caused by the acoustic noise associated with echo planar imaging (EPI) is mapped. Two types of time series were compared. The first time series, considered the "task," involved applying only EPI gradients for 20 s without the application of RF pulses, then, without pause, starting image collection. The second, considered the "control," involved typical sequential image acquisition without the prior gradient pulses. Subtraction of the first 5 s of the two time series revealed signal enhancement mainly in the primary auditory cortex. The technique was validated using a motor cortex task that mimicked the hypothesized scanner noise induced activation.
Article
This study analyzes the distribution of the intrinsic and commissural fiber plexuses originating in the central nucleus of the inferior colliculus in the rat. The anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) was injected iontophoretically at different places along the tonotopic axis of the central nucleus and visualized immunohistochemically. In coronal sections the terminal fields of axons originating at each injection site are seen to create four well-defined bands across the rostrocaudal extent of the inferior colliculus, two in the ipsilateral and two in the contralateral side. The “ipsilateral main band” extends dorsomedially and ventrolaterally from the injection site, in register with the known isofrequency contours of the central nucleus, spanning this nucleus and extending into the dorsal cortex of the inferior colliculus. The “ipsilateral external band” is located in the external cortex, where it is oriented dorsoventrally, slightly oblique to the pial surface. In caudal sections, the ventral portion of these two bands appear to join. The two bands in the contralateral inferior colliculus occupy a symmetric position to those of the ipsilateral side, forming a mirror-like image. The position of the four bands changes as the position of the injection site is varied along the frequency gradient axis of the central nucleus. After ventromedial (high frequency area) injections, the main band is ventral and medial, and the external band ventral and lateral. After more dorsolateral (lower frequency) injections, the main band is more dorsal and lateral, whereas the external band is more dorsal but more medial. Thus, the change in the position of the external band is separate and opposite to that of the main band. We suggest that the main bands represent isofrequency contours. Since the projection from the central nucleus to the external cortex of the inferior colliculus also appears to be tonotopic, we also propose a tonotopic organization for the external cortex. The main bands overlap the terminal field of the lemniscal fibers in the central nucleus; thus, it is concluded that the intracollicular fibers contribute to the formation of the known fibrodendritic laminae of the central nucleus. A possible role in preservation of frequency information and integration of other different acoustic parameters is proposed for the main bands. The external bands could participate in polysensory integration, and the commissural connections could be involved in hitherto unknown stages of binaural processing of sound. Based on our results, several modifications are proposed for delineating the subdivisions of the inferior colliculus.
Article
A 14-channel SQUID (superconducting quantum interference device) system has been used to record the magnetic signal from the human brain in response to an auditory stimuli (750, 1,000, 1,250 and 1,500 Hz, 70, 76 and 82 dB SPL, 500 ms duration). Three individuals with normal hearing were studied. The locations of magnetic response at the latency of 70 ms (P70), 100 ms (N100) and 160 ms (P160) from the onset of the auditory stimulus were identified. The location for N100 response corresponded to the primary auditory cortex (area 41), where a clear tonotopic organization was demonstrated. The amplitopic organization was less evident. These results suggest a flow of auditory signals in the temporal lobe and tonotopic organization in the auditory cortex.
Article
Paramagnetic deoxyhemoglobin in venous blood is a naturally occurring contrast agent for magnetic resonance imaging (MRI). By accentuating the effects of this agent through the use of gradient-echo techniques in high fields, we demonstrate in vivo images of brain microvasculature with image contrast reflecting the blood oxygen level. This blood oxygenation level-dependent (BOLD) contrast follows blood oxygen changes induced by anesthetics, by insulin-induced hypoglycemia, and by inhaled gas mixtures that alter metabolic demand or blood flow. The results suggest that BOLD contrast can be used to provide in vivo real-time maps of blood oxygenation in the brain under normal physiological conditions. BOLD contrast adds an additional feature to magnetic resonance imaging and complements other techniques that are attempting to provide positron emission tomography-like measurements related to regional neural activity.
The tonotopic organization of the human auditory cortex has been investigated by systematic measurements of magnetic fields evoked by tone-bursts with carrier frequencies of 250, 500, 1000, 2000 and 4000 Hz. The measured field distribution changes with both time elapsed since stimulus onset and frequency of the stimulus. Nevertheless, the field distribution has always the same overall features and can be approximated by that of an equivalent current dipole located in a semi-infinite volume. This model can be described in terms of 5 parameter values: 3 orthogonal coordinates specifying the dipole location, and amplitude and angle of the dipole moment. The amplitude of the dipole moment is maximal at about 100 msec ('component 100m') and 160 msec ('component 160m') after stimulus onset. The depth estimated for the generator site of the 100m component shows a logarithmic dependence on test frequency whereas no similar behaviour could be observed for the 160m component. Anatomical studies performed in cadaver heads suggest that the equivalent current dipoles of both the 100m and the 160m component are located in the transverse temporal gyri.
Article
Coupling between cerebral blood flow (CBF) and cerebral metabolic rate of oxygen (CMRO2) was studied using multiple sequential administrations of 15O-labeled radiotracers (half-life, 123 sec) and positron emission tomography. In the resting state an excellent correlation (mean r, 0.87) between CBF and CMRO2 was found when paired measurements of CBF and CMRO2 from multiple (30-48) brain regions were tested in each of 33 normal subjects. Regional uncoupling of CBF and CMRO2 was found, however, during neuronal activation induced by somatosensory stimulation. Stimulus-induced focal augmentation of cerebral blood flow (29% mean) far exceeded the concomitant local increase in tissue metabolic rate (mean, 5%), when resting-state and stimulated-state measurements were obtained in each of 9 subjects. Stimulus duration had no significant effect on response magnitude or on the degree of CBF-CMRO2 uncoupling observed. Dynamic, physiological regulation of CBF by a mechanism (neuronal or biochemical) dependent on neuronal firing per se, but independent of the cerebral metabolic rate of oxygen, is hypothesized.
Article
Positron emission tomography (PET) was used to map alterations in local neuronal activity induced in human primary auditory cortex by pure-tone stimulation. Patterns of blood flow were observed in specific regions on the superior temporal plane showing systematic changes in activity depending on the frequency of a stimulating pure tone. The orientation of these regions agrees well with data for non-human primates.
Article
The distribution of best frequencies of neurons and neuron clusters was mapped on the superior temporal plane of the macaque monkey. Primary auditory cortex (A1) comprises a complete and orderly cochlear representation. It is coextensive with a cytoarchitectonic field often referred to as koniocortex. Cochlear apex (low frequency) is represented rostrolaterally within the field and cochlear base (high frequency) caudomedially. A small region of the cochlear partition is found represented in a band of auditory cortex; that is neurons with very similar best frequencies are arrayed both vertically and horizontally. Surrounding A1 4 other fields are found that on both anatomical and physiological grounds stand apart from the primary field. In two of them the data suggest the way in which they are topographically organized. In addition there appear to be auditory areas extending further rostrally on the plane and onto the lateral surface of the superior temporal gyrus.
Frequency glides from a continuous tone have been shown to produce activity from the human cortex that can be recorded as time-varying magnetic fields outside the scalp in the same way as simpler auditory stimuli such as clicks and tone bursts. Data analysis has been based on a model assuming an equivalent current dipole localized close to the skull surface. Recorded data have shown good agreement with such a model. Interhemispheric differences have been shown in the location of this dipole, as well as with regard to dipole moment and latencies of responses to contralateral stimulation. The location of the equivalent dipole for frequency glide stimulation is close to that previously reported for tone pulse stimulation. However, the results indicate that differences in location of the order of 10 mm may exist. Comparing previously reported electric responses to frequency glides indicates essentially qualitative agreement although some significant differences have also been found. This is interpreted as evidence that at least the major contributions to the two types of response are produced by the same generator in the temporal lobe of the human cortex.
Article
A new cytoarchitectonic study of the human auditory cortex was undertaken in the light of recent knowledge concerning the architecture, fiber connectivity, and physiology of this region in the monkey. The survey of three normative human brains (six hemispheres) processed in whole-brain serial sections disclosed a cytoarchitectonic organization of the cortical auditory region similar to that in the macaque. Unlike the monkey, auditory-related cortex was found in parietal operculum and inferior parietal lobule. Similarities in cortical architectonics between human and monkey brains may provide a rationale for the application of knowledge concerning animal physiology and connectivity to man.
Article
The late, acoustically evoked, averaged magnetic field from the right hemisphere of the human brain is composed of two signals. One is dominant, appears generated by an equivalent current dipole within or near the primary auditory cortex and shows a frequency dependent location and/or orientation (tonotopical organization). The other, denoted the 'residual' signal, resembles the electric T-complex and is possibly generated more diffusely in the auditory and adjacent cortical areas.
Article
Afferents from the hindbrain auditory system to the nuclei of the lateral lemniscus were analyzed by the use of orthograde and retrograde axon-tracing techniques. Three divisions of the nuclei of the lateral lemniscus, a dorsal, an intermediate, and a ventral division are discussed. The dorsal nucleus of the lateral lemniscus is a recipient of afferents from cells located mainly in the superior olivary complex and the contralateral dorsal nucleus of the lateral lemniscus. It receives direct afferents from only a few cells in the cochlear nuclei. In sharp contrast, the ventral nucleus of the lateral lemniscus is the recipient of afferents from many cells in the contralateral ventral cochlear nucleus and from only a few cells in the superior olivary complex. Further, it receives no afferents from cells in the contralateral nuclei of the lateral lemniscus. The intermediate nucleus of the lateral lemniscus receives afferents from some cells in the cochlear nucleus and the superior olivary complex. It is unique among the three nuclei of the lateral lemniscus in that it receives a substantial projection from the medial nucleus of the trapezoid body.
This paper presents data concerning auditory evoked responses in the middle latency range (wave Pam/Pa) and slow latency range (wave N1m/N1) recorded from 12 subjects. It is the first group study to report multi-channel data of both MEG and EEG recordings from the human auditory cortex. The experimental procedure involved potential and current density topographical brain mapping as well as magnetic and electric source analysis. Responses were compared for the following 3 stimulus frequencies: 500, 1000 and 4000 Hz. It was found that two areas of the auditory cortex showed mirrored tonotopic organization; one area, the source of N1m/N1 wave, exhibited higher frequencies at progressively deeper locations, while the second area, the source of the Pam/Pa wave, exhibited higher frequencies at progressively more superficial locations. The Pa tonotopic map was located in the primary auditory cortex anterior to the N1m/N1 mirror map. It is likely that N1m/N1 results from activation of secondary auditory areas. The location of the Pa map in A1, and its N1 mirror image in secondary auditory areas is in agreement with observations from animal studies.
The tonotopic organization of the human auditory cortex has been investigated by means of scalp potential mapping and dipole modelling of the evoked response occurring around 100 msec after the stimulus onset. The major characteristics of the topographical changes observed with increasing stimulus frequency were statistically demonstrated. Using a 3-concentric sphere head model, the scalp potential distributions can be explained in first approximation by two equivalent current dipoles, located in the supratemporal plane and mimicking the activity of both auditory cortices. To take into account the temporal aspects of the brain activities, 3 time-varying dipole strategies were tested. Frequency dependence of the dipole orientation has been evidenced in both hemispheres with the 3 models, whereas no significant change in dipole position was found. The tilt in dipole orientation could be related to the folding geometry of Heschl's gyrus, which varies with depth. In agreement with previous MEG findings, this brings new evidence for a tonotopic organization of the auditory cortical area involved in the N100 wave generation. Moreover, distinct frequency dependences of the equivalent current dipoles were observed in the early and the late parts of the N100. This study demonstrates that simple dipolar models, applied on electrical data, make it possible to reveal functionally distinct cortical areas.
Article
Microelectrode recordings were used to investigate the tonotopic organization of auditory cortex of macaque monkeys and guide the placement of injections of wheat germ agglutinin-horse radish peroxidase (WGA-HRP) and fluorescent dyes. Anatomical and physiological results were later related to histological distinctions in the same brains after sections were processed for cytoarchitecture, myeloarchitecture, acetylcholinesterase (AchE), or cytochrome oxidase (CO). The experiments produced several major findings. (1) Neurons throughout a broad expanse of cortex were highly responsive to pure tones, and best frequencies could be determined for neurons in arrays of recording sites. (2) The microelectrode recordings revealed two systematic representations of tone frequencies, the primary area (AI) and a primary-like rostral field (R) as previously described. The representation of high to low frequency tones in A1 was largely caudorostral along the plane of the sulcus. A reversal of the order of representation of frequencies occurred in R. (3) AI and R together were coextensive with a koniocellular, densely myelinated zone that expressed high levels of AchE and CO. These architectonic features were somewhat less pronounced in R than AI, but a clear border between the two areas was not apparent. (4) Cortex bordering AI and R was less responsive to tones, but when best frequencies for neurons could be determined, they matched those for adjoining parts of AI and R. (5) Architectonically distinct regions were apparent within some of the cortex bordering AI and R. (6) The major ipsilateral cortical connections of AI were with R and cortex immediately lateral and medial to AI. (7) Callosal connections of AI were predominantly with matched locations in the opposite AI, but they also included adjoining fields. (8) Neurons in the ventral (MGV), medial (MGM), and dorsal (MGD) nuclei of the medial geniculate complex projected to AI and cortex lateral to AI. (9) Injections in cortex responsive to high frequency tones labeled more dorsal parts of MGV than injections in cortex responsive to low frequency tones.
Article
Neuromagnetic responses were recorded over the whole head with a 122-channel gradiometer. A pair of 150-ms 1-kHz tones separated by an interval of 150 ms was presented to one ear every 2 s. The other ear received either no input, an identical pair simultaneous to the first, an identical pair alternating with the first or a continuous 600-ms tone. The 'monaural shift' condition in which stimuli alternated between ears produced a clear perception of changing lateralisation, but the evoked response could be explained as merely the sum of simple monaural onset and offset responses; thus we found no evidence for a separate response to interaural intensity difference in this condition. The 'binaural shift' condition, in which intensity changed in one ear while the other received a continuous tone, evoked a transient response (N130m) at a latency of about 130 ms. N130m was larger over the hemisphere contralateral to the direction of shift, and larger than the corresponding monaural response, whether to an onset or an offset. We concluded that N130m also was not a separate directional response, but was analogous to a simple monaural response, the prolonged latency being due to masking and the enhanced amplitude to facilitation by the sustained response to the continuous tone.
Article
Functional magnetic resonance imaging (FMRI) detects focal MRI signal changes in brain tissue that are believed to result from changes in neuronal activity. We describe the dependence of this response in auditory cortex on the rate of presentation of simple speech stimuli. Speech syllables were presented to five normal subjects at rates ranging from 0.17 to 2.5 Hz, while the subjects performed a phoneme discrimination task. Regions studied with FMRI during this task included the lateral aspect of both temporal lobes. All subjects showed bilateral superior temporal lobe MRI signal increases that were coincident with stimulus presentation and performance of the task. The magnitude of this response increased in a monotonic, non-linear manner with increasing stimulus rate. This rate-response relationship was nearly identical in right and left hemispheres. The relationship may reflect metabolic activity integrated over time and subject to non-linear characteristics of neuronal recovery or blood flow regulation. The dependence of response magnitude on stimulation rate supports the hypothesis that the FMRI phenomenon indirectly reflects neuronal metabolic activity. The measures provided here should assist in the design of optimal activation strategies for the human auditory cortex.
Article
Corticothalamic connections of auditory areas of the superior temporal regions (STR) were investigated in the rhesus monkey. These connections are organized according to the recently described architectonic parcellation of the STR. The core line regions of the supratemporal plane (STP) project to the medial geniculate nucleus (MGN). All regions except the primary auditory area also have projections to additional thalamic nuclei. The rostral STP has strong connections with the caudal part of the ventral subdivision (MGv) of MGN as well as with medial pulvinar (PM). In contrast the primary auditory area projects mainly to rostral MGv. The caudalmost STP projects mainly to the dorsal subdivision (MGd) and to the magnocellular subdivision (MGmc) as well as to the PM and the lateral (PL) and oral (PO) pulvinar, nucleus limitans (Li), and mediodorsal (MD) nucleus. The belt line regions of the superior temporal gyrus (STG) project mainly to the pulvinar but also have projections to MGd and MGmc. Specifically, rostral STG projects to the caudal part of PM, to MGmc, and to the suprageniculate (SG) nucleus, whereas caudal STG projects to the rostral part of PM and to PL, PO, MGd, MGmc, SG-Li and MD nuclei. The root line areas in the circular sulcus of the Sylvian fissure project mainly to MGmc but also to MGd, PM, and SG-Li nuclei. These connections originate mainly from neurons in cortical layer VI, with some from layer Vb. It is suggested that these connections may be involved in different aspects of auditory information processing.
Article
Stimulus-related signal changes in functional MRI of human brain activation not only reflect associated adjustments of cerebral blood flow and oxygen consumption, but strongly depend on the MRI technique chosen and the actual experimental setting. A list of relevant parameters includes static field homogeneity of the magnet, MR pulse sequence and signal type, TE, TR, flip angle, gradient strengths, gradient waveforms, receiver bandwidth and voxel size. In principle, a local signal increase during functional activation may reflect a regional change in cerebral blood flow or deoxyhemoglobin concentration or both. This ambiguity was demonstrated using long TE FLASH MRI at high spatial resolution. Subsequently, experimental strategies were evaluated that either discriminate MRI effects in large vessels from those in the cortical microvasculature or separate changes in blood flow velocity from those in blood oxygenation. Examples comprise studies of the human visual and motor cortex.
Article
Magnetic resonance imaging methods recently demonstrated regional cerebral signal changes in response to limb movement and visual stimulation, attributed to blood flow enhancement. We studied 5 normal subjects scanned while listening to auditory stimuli including nonspeech noise, meaningless speech sounds, single words, and narrative text. Imaged regions included the lateral aspects of both hemispheres. Signal changes in the superior temporal gyrus and superior temporal sulcus were observed bilaterally in all subjects. Speech stimuli were associated with significantly more widespread signal changes than was the noise stimulus, while no consistent differences were observed between responses to different speech stimuli. Considerable intersubject variability in the topography of signal changes was observed. These observations confirm the utility of magnetic resonance imaging in the study of human brain structure-function relationships and emphasize the role of the superior temporal gyrus in perception of acoustic-phonetic features of speech, rather than processing of semantic features.
Article
Conventional gradient-echo magnetic resonance imaging (MRI) at 4 Tesla was used successfully to study the activity of Broca's area during internal speech word generation in healthy right-handed volunteers. Activity was demonstrated in the internal gray matter surrounding the ascending ramus of the lateral sulcus, deep to the cortical surface representation of Broca's area, in all the subjects. These studies demonstrate the capability of functional MRI to non-invasively map language related cognitive functions. Such functional mapping has value for both the study of basic neuroscience and neurosurgical planning.
Article
Two tone stimuli, one frequent (standard) and the other infrequent (a slightly higher, deviant tone), were presented in random order and at short intervals to subjects reading texts they had selected. In different blocks, standards were either 250, 1,000, or 4,000 Hz, with the deviants always being 10% higher in frequency than the standards of the same blocks. Magnetic responses elicited by the standard and deviant tones included N1m, the magnetoencephalographic equivalent of the electrical N1 (its supratemporal component). In addition, deviant stimuli elicited MMNm, the magnetic equivalent of the electrical mismatch negativity, MMN. The equivalent dipole sources of the two responses were located in supratemporal auditory cortex, with the MMNm source being anterior to that of N1m. The dipole orientations of both sources in teh sagittal plane depended on stimulus frequency, suggesting that the responses are generated by tonotopically organized neuronal populations. The tonotopy reflected by the frequency dependence of the MMNm source might be that of the neural trace system underlying frequency representation of auditory stimuli in sensory memory.
Article
Percepts unaccompanied by a veridical stimulus, such as hallucinations, provide an opportunity for mapping the neural correlates of conscious perception. Functional magnetic resonance imaging (fMRI) can reveal localized changes in blood oxygenation in response to actual as well as imagined sensory stimulation. The safe repeatability of fMRI enabled us to study a patient with schizophrenia while he was experiencing auditory hallucinations and when hallucination-free (with supporting data from a second case). Cortical activation was measured in response to periodic exogenous auditory and visual stimulations using time series regression analysis. Functional brain images were obtained in each hallucination condition both while the patient was on and off antipsychotic drugs. The response of the temporal cortex to exogenous auditory stimulation (speech) was markedly reduced when the patient was experiencing hallucinating voices addressing him, regardless of medication. Visual cortical activation (to flashing lights) remained normal over four scans. From the results of this study and previous work on visual hallucinations we conclude that hallucinations coincide with maximal activation of the sensory and association cortex, specific to the modality of the experience.
Article
We investigated the functional organization of human auditory cortex using a new chronic microelectrode technique. Tonotopic mapping data was obtained at the single unit level for the first time in humans. All sound-driven units were noted to have frequency-dependent response patterns. The majority of units (73%) demonstrated sharply tuned excitatory best-frequency responses. Twenty seven percent of units showed wide receptive fields, representing excitatory responses to almost the entire range of frequencies presented. A tonotopic pattern was observed with best frequencies systematically increasing as more medial-caudal recording sites were sampled.
Article
Gradient acoustic noise has been measured and characterized for an epoxy-potted, shielded gradient assembly in a 1.5 T MRI system. Noise levels vary by 10 dB or more as a function of longitudinal position in the scanner and reflect the pattern of forces applied to the gradient assembly. The noise level increases slightly (1-3 dB) with a patient in the scanner. The spectrum of the noise is similar (but not identical) to the spectrum of the input signal. A gradient-pulse-to-acoustic-noise transfer function was obtained by using a white noise voltage input to the gradient system. The transfer function enabled us to accurately predict acoustic noise output for a pulse sequence consisting of a series of trapezoidal pulses on a single axis and for a clinical fast spin echo sequence with gradients present on all three axes.
Article
The amygdalar complex is a medial temporal lobe structure in the brain which is widely considered to be involved in the neural substrates of emotion. Selective bilateral damage to the human amygdala is rare, offering a unique insight into its functions. There is impairment of social perception after amygdala damage, with defective recognition of facial expressions of emotion. Among the basic emotions, the processing of fear and anger has been shown to be disrupted by amygdala damage. Although it remains puzzling why this not found in all cases, the importance of the amygdala in negative emotion, and especially fear, has been confirmed by conditioning, memory and positron emission tomography (PET) experiments. Central to our understanding of these findings is the question of whether the amygdala is involved specifically in the perception of visual signals of emotion emanating from the face, or more widely in the perception of emotion in all sensory modalities. We report here a further investigation of one of these rare cases, a woman (D.R.) who has impaired perception of the intonation patterns that are essential to the perception of vocal affect, despite normal hearing. As is the case for recognition of facial expressions, it is recognition of fear and anger that is most severely affected in the auditory domain. This shows that the amygdala's role in the recognition of certain emotions is not confined to vision, which is consistent with its being involved in the appraisal of danger and the emotion of fear.
Article
Using a model of the functional MRI (fMRI) impulse response based on published data, we have demonstrated that the form of the fMRI response to stimuli of freely varied timing can be modeled well by convolution of the impulse response with the behavioral stimulus. The amplitudes of the responses as a function of parametrically varied behavioral conditions are fitted well using a piecewise linear approximation. Use of the combined model, in conjunction with correlation analysis, results in an increase in sensitivity for the MRI study. This approach, based on the well-established methods of linear systems analysis, also allows a quantitative comparison of the response amplitudes across subjects to a broad range of behavioral conditions. Fit parameters, derived from the amplitude data, are relatively insensitive to a variety of MRI-related artifacts and yield results that are compared readily across subjects.