Secondary Auditory Areas Research Articles

Heschl's gyrus is more sensitive to changes in sound level than secondary auditory areas

Heschl's gyrus is more sensitive to changes in sound level than secondary auditory areas

Separation of mismatch negativity and the N1 wave in the auditory cortex of the cat: a topographic study

Objective: The amplitude distribution of the frequency mismatch negativity (MMN) and that of P1 and N1 components were investigated in cats to reveal their sources in the auditory areas of the neocortex. Methods: Pure tone stimuli were given in a passive oddball paradigm with different degrees of deviance between the standard and deviant tones. Amplitude maps of event-related potential (ERP) components were generated from the responses, recorded in awake, freely moving animals by a chronically implanted epidural electrode matrix, covering both the primary and secondary auditory fields. Results: The P1 and N1 components appeared with highest amplitude on the middle ectosylvian gyrus, while the amplitude maximum of the MMN was ventral and rostral to them on the AII area. Both the latency and the peak amplitude of the MMN depended on the degree of deviance. Conclusions: The MMN is generated in the rostroventral part of the secondary auditory area, well separated from the sources of the P1 and N1 components.

Perception of audiovisual speech stimuli

In face-to-face communication, speech is seen as well as heard. The important role of visual speech becomes evident under noisy conditions. The efficacy of visual speech is also nicely demonstrated by the so-called McGurk effect: when the auditory syllable /aba/ is dubbed on to the video-taped visual articulation of /aga/, most subjects hear a syllable /ada/. Recent neuromagnetic recordings have revealed a specific integration signal generated at or close to the primary auditory cortex. The onset of this signal is 150–250 ms. In a similar vein, fMRl studies have indicated that the primary and/or secondary auditory cortical areas are active during silent lipreading. It was recently demonstrated in our laboratory that the integration is similar when nonwords and real words are used as stimuli, suggesting that the integration is not much influenced by lexical representations and probably occurs at the phonetical-perceptual information processing stage. New data from our laboratory preliminarily suggests that the perception of audiovisual stimuli is influenced by attention. This indicates that the integration is not, at least totally, automatic.

Functional Differentiation in the Human Auditory and Language Areas Revealed by a Dichotic Listening Task

The human auditory cortex plays a special role in speech recognition. It is therefore necessary to clarify the functional roles of individual auditory areas. We applied functional magnetic resonance imaging (fMRI) to examine cortical responses to speech sounds, which were presented under the dichotic and diotic (binaural) listening conditions. We found two different response patterns in multiple auditory areas and language-related areas. In the auditory cortex, the medial portion of the secondary auditory area (A2), as well as a part of the planum temporale (PT) and the superior temporal gyrus and sulcus (ST), showed greater responses under the dichotic condition than under the diotic condition. This dichotic selectivity may reflect acoustic differences and attention-related factors such as spatial attention and selective attention to targets. In contrast, other parts of the auditory cortex showed comparable responses to the dichotic and diotic conditions. We found similar functional differentiation in the inferior frontal (IF) cortex. These results suggest that multiple auditory and language areas may play a pivotal role in integrating the functional differentiation for speech recognition.

Functional brain imaging of tinnitus-like perception induced by aversive auditory stimuli.

Tinnitus is an aversive auditory percept of unknown origin. We tested the speculation that tinnitus may share neuronal processing mechanisms with aversive auditory percepts of known origin. This study revealed the functional neuroanatomy of the perception of aversive auditory stimuli. The stimuli were presented to 12 healthy volunteers so as to mimic the psychoacoustical features of tinnitus and its affective response in tinnitus sufferers. The regional cerebral blood flow distribution was measured by PET during four auditory processing conditions and one control condition. The aversive auditory stimuli activated primary and secondary auditory areas bilaterally, dorsolateral prefrontal attention areas, and structures in the limbic system which subserve emotional processing. Based on these results and findings from other functional neuroimages of tinnitus, we hypothesize that the perception of tinnitus may involve the functional linkage of these brain areas: secondary auditory cortex, dorsolateral prefrontal cortex, and limbic system.

The organization of the ipsilateral corticocortical projections of the middle sylvian gyrus (AII) in the cat cerebral cortex: an anterograde and retrograde tracing study

The primary auditory area and surrounding cortical areas including the secondary auditory area subserve auditory information processing. In this study, corticocortical ipsilateral connections of the secondary auditory area were investigated in the cat by injections of axonally transported tracers into this area. Retrogradely labeled cells and anterogradely labeled terminals were observed within the secondary auditory area itself, and the primary, posterior, and ventroposterior auditory areas, in the insular cortex and in the temporal auditory area. These labeled terminals appeared in patches or bands. In the ventroposterior auditory area, a transverse band of anterograde and retrograde labeling, which extended from the posterior ectosylvian sulcus to the surface of the posterior ectosylvian gyrus was observed. In addition, scattered labeling was detected in the posterior auditory area. Although different parts of the secondary auditory area were found to send axons to different zones within the temporal cortex, no precise topographical arrangement was detected. When two different tracers were injected into the secondary auditory area and the anterior auditory area, respectively, some terminal aggregates anterogradely labeled from these two areas converged within the posterior auditory area. The organization of these connections is discussed in relationship to the functional role of the auditory cortical areas. © 1998 Elsevier Science Inc.

Corticocortical projections from the secondary auditory area to cortices around the anterior ectosylvian sulcus in the cat

Corticocortical projections from the secondary auditory area to cortices around the anterior ectosylvian sulcus in the cat

Complementary expression of parvalbumin and calbindin D-28k delineates subdivisions of the rabbit medial geniculate body.

The complementary pattern of immunohistochemical staining for the calcium-binding proteins parvalbumin (PV) and calbindin D-28k (CB) was used to delineate four major subdivisions of the rabbit medial geniculate body (MGB). PV immunoreactivity predominates in the ventral and medial divisions, whereas CB-immunoreactive cells characterize the dorsal and internal divisions. The ventral nucleus is strongly PV+ due to dense neuropil labeling and moderately labeled somata. The medial nucleus contains both medium-sized and large PV+ somata, as well as thick PV+ axons and terminals. The wedge-shaped internal nucleus composed of densely labeled CB+ cells, separates the dorsal and ventral nuclei rostrally, and expands caudally to encapsulate the posterior MGV. Large multipolar CB+ neurons with radiate dendrites characterize the dorsal nucleus. The differential expression of PV and CB also distinguishes the deep dorsal and superficial dorsal subnuclei in the dorsal division and a ventrolateral component in the ventral division. A comparison with studies of MGB connectivity in a variety of species suggests that PV immunoreactivity is highest in subdivisions that receive a substantial input from the central nucleus of the inferior colliculus and that project to primary auditory cortex. In contrast, CB immunoreactivity characterizes nuclei that receive input primarily from other sources, such as the paracentral nuclei of the inferior colliculus, the lateral tegmentum, and the spinal cord, and that project to secondary auditory areas. The ability of calcium-binding protein immunohistochemistry to delineate neuronal compartments across indistinct cytoarchitectonic borders makes it a powerful tool for guiding future connectional and physiological studies of the MGB.

Sensory cortical processing and the biological basis of personality

Action-oriented personality traits such as sensation seeking, extraversion, and impulsivity have been related to a pronounced amplitude increase of auditory evoked scalp potentials with increasing stimulus intensity. Dipole source analysis represents a crucial methodological progress in this context, because overlapping subcomponents of the scalp potentials can be separated and can be related to their generating cortical structures. In a study on 40 healthy subjects, it was found that sensation seeking is clearly related to the auditory evoked response pattern (N1/P2-component, stimulus intensities: 60, 70, 80, 90, 100 dB SPL) of the superior temporal plane including primary auditory cortex, but not to that of secondary auditory areas in the lateral temporal cortex. These results support the concept that the serotonergic brain system, which is supposed to modulate sensory processing in primary auditory cortices, is an important factor underlying individual differences in sensation seeking.

The tridimensional personality questionnaire and the intensity dependence of auditory evoked dipole source activity

The relationship between the tridimensional personality questionnaire's (TPQ) dimensions "novelty seeking," "harm avoidance," and "reward dependence" and the intensity dependence of the auditory evoked N1/P2-component was investigated in healthy subjects. Using dipole source analysis, evoked activity of the primary auditory cortex (tangential dipole) could be analyzed at least in part separately from that of secondary auditory areas (radial dipole). It was found that the intensity dependence of the tangential dipole was positively correlated to the TPQ dimension "novelty seeking," but not to "harm avoidance" and "reward dependence." This is in line with findings concerning similar personality traits like "sensation seeking," "impulsivity," or "extraversion." It is therefore concluded that a strong intensity dependence may characterize subjects with an action-oriented and extroverted personality style. The results are discussed within the concept that a low central serotonergic neurotransmission is underlying both an impulsive personality type and a strong intensity dependence of the tangential dipole.

Specific tonotopic organizations of different areas of the human auditory cortex revealed by simultaneous magnetic and electric recordings

This paper presents data concerning auditory evoked responses in the middle latency range (wave Pam/ Pa) and slow latency range (wave Nlm/Nl) 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 Al, and its N1 mirror image in secondary auditory areas is in agreement with observations from animal studies.

Responses of the neurons of the cortical secondary auditory area to acoustic and non-acoustic stimuli in cats

The responses of the cortical secondary auditory area (AII) to the non-acoustic stimuli (electrical stimulation of the skin in the vibrissae area and light flash) and their combination with acoustic stimulation (sound click or tone) were studied in experiments on cats anesthetized by kalipsol using extra- and intracellular recording. Of the total number of neurons, 69% of the units generating spike responses to the acoustic stimulation responded to the non-acoustic stimulation too. The responses to the modal-nonspecific stimulation, as a rule, were weak and variable; they were mostly represented by a tonic change in the neuronal discharge frequency. The nonspecific stimulation evoked primary excitatory and inhibitory postsynaptic potentials in 77% and 20% of the examined neurons, respectively. We found that synaptic effects of the nonspecific and specific stimulations interact with each other, ensuring considerable modulation of the latter (mostly a significant facilitation resulting from the EPSP summation and suppression of an inhibitory component of the response to acoustic stimulation). Possible participation of the midbrain reticular formation in the transmission of the modal-nonspecific influences to the cortical neurons is considered; stimulation of this structure evoked responses similar to those evoked by the modal-nonspecific sensory stimuli.

Intensity dependence of auditory evoked dipole source activity

The spatio-temporal approach of dipole source analysis represents a crucial methodological progress in research on individual differences in the dependence of auditory evoked potential amplitude on stimulus intensity (augmenting /reducing) because overlapping subcomponents of the N1/P2-component can be separated and can be related to their generating cortical structures. Basic aspects of the intensity dependence of auditory evoked dipole source activity were analysed in 40 healthy subjects. The evoked responses to binaural 1000-Hz tones at five levels of intensity (60, 70, 80, 90, 100 dB sound pressure level) were recorded at 33 sites across the scalp. The dipole source analysis of the grand average data confirms the reports in the literature that the N1/P2 potentials at the scalp can be explained by two dipoles per hemisphere: a tangential dipole, representing activity of the superior temporal cortex (including primary auditory cortex), and a radial dipole, representing activity of the lateral temporal cortex (secondary auditory areas). The intensity dependence of the tangential dipole activity was significantly more pronounced than that of the radial dipoles, supporting the assumption that radial and tangential dipoles represent different physiological processes. A high reliability of the intensity dependence of the tangential dipole (Pearson correlation: r = 0.88) was found when retesting the subjects after three weeks. Age was negatively correlated with the intensity dependence of the tangential dipole. Dipole source analysis proved to be a reliable method which allows, at least in part, to study separately the intensity dependence of the evoked responses from primary and secondary auditory cortices. This is of importance with regard to the hypothesis that the central serotonergic system modulates the intensity dependence of the evoked N1/P2-response of primary auditory cortex.

Learning-induced changes of auditory receptive fields

Classical conditioning specifically modifies receptive fields in primary and secondary auditory cortical areas to favor the frequency of a tone signal over other frequencies, including tuning shifts toward, or to, this frequency. This plasticity of receptive fields is associative and highly specific, can develop very rapidly, can be expressed under anesthesia and can be maintained indefinitely. Muscarinic receptors in the cortex may be involved. Non-lemniscal thalamic nuclei also develop receptive field plasticity that may contribute to cortical plasticity.

Intensity dependence of auditory evoked potentials as an indicator of central serotonergic neurotransmission: A new hypothesis

Because of the increasing importance of the central serotonergic neurotransmission for pathogenetic concepts and its role as a target of pharmacotherapeutic interventions in psychiatry, reliable indicators of this system are needed. It is proposed that the stimulus intensity dependence of auditory evoked N1/P2-component, which is probably modulated by cortical serotonergic innervation, may be a useful and noninvasive indicator of behaviorally relevant aspects of serotonergic activity. Converging evidence from our own studies as well as from the literature suggests that a pronounced intensity dependence of auditory evoked N1/P2-component reflects low central serotonergic neurotransmission. Recent findings concerning general functional aspects of the brain serotonin system reveal that this system is well qualified for adjusting individuals levels of sensory processing (“set the tone”), especially in the primary auditory cortex in which the N1/P2-component is mainly generated. Dipole source analysis represents an important methodological advance in this context because it allows the separation of N1/P2-subcomponents generated in the primary auditory cortex from those generated in secondary auditory areas.

Auditory corticocortical interconnections in the cat: evidence for parallel and hierarchical arrangement of the auditory cortical areas

The origin and laminar arrangement of the homolateral and callosal projections to the anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) auditory cortical areas were studied in the cat by means of electrophysiological recording and WGA-HRP tracing techniques. The transcallosal projections to AAF, AI, PAF and AII were principally homotypic since the major source of input was their corresponding area in the contralateral cortex. Heterotypic transcallosal projections to AAF and AI were seen, originating from the contralateral AI and AAF, respectively. PAF received heterotypic commissural projections from the opposite ventroposterior auditory cortical field (VPAF). Heterotypic callosal inputs to AII were rare, originating from AAF and AI. The neurons of origin of the transcallosal connections were located mainly in layers II and III (70-92%), and less frequently in deep layers (V and VI, 8-30%). Single unit recordings provided evidence that both homotypic and heterotypic transcallosal projections connect corresponding frequency regions of the two hemispheres. The regional distribution of the anterogradely labeled terminals indicated that the homotypic and heterotypic auditory transcallosal projections are reciprocal. The present data suggest that the transcallosal auditory interconnections are segregated in 3 major parallel components (AAF-AI, PAF-VPAF and AII), maintaining a segregation between parallel functional channels already established for the thalamocortical auditory interconnections. For the intrahemispheric connections, the analysis of the retrograde tracing data revealed that AAF and AI receive projections from the homolateral cortical areas PAF, VPAF and AII, whose neurons of origin were located mainly in their deep (V and VI) cortical layers. The reciprocal interconnections between the homolateral AAF and AI did not show a preferential laminar arrangement since the neurons of origin were distributed almost evenly in both superficial (II and III) and deep (V and VI) cortical layers. On the contrary, PAF received inputs from the homolateral cortical fields AAF, AI, AII and VPAF, originating predominantly from their superficial (II and III) layers. The homolateral projections reaching AII originated mainly from the superficial layers of AAF and AI, but from the deep layers of VPAF and PAF. The laminar distribution of anterogradely labeled terminal fields, when they were dense enough for a confident identification, was systematically related to the laminar arrangement of neurons of origin of the reciprocal projection: a projection originating from deep layers was associated with a reciprocal projection terminating mainly in layer IV, whereas a projection originating from superficial layers was associated with a reciprocal projection terminating predominantly outside layer IV.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent and Efferent Connections of Temporal Association Cortex in the Rat: A Horseradish Peroxidase Study.

We studied the afferent and efferent connections of the caudal temporal cortex in rat using the tracer wheat germ agglutinin - horseradish peroxidase (WGA - HRP). This area is reciprocally connected with primary and secondary visual and auditory areas of cortex. The connections with primary visual cortex are restricted to the ventral and caudal parts of the caudal temporal area. Caudal temporal cortex has reciprocal connections with the perirhinal cortex and projects to the caudate - putamen and lateral and basolateral nuclei of the amygdala. It also has reciprocal connections with the nucleus lateralis posterior, the dorsal and medial divisions of the medial geniculate nucleus and the caudal part of the posterior nucleus of the thalamus. It projects to the deep layers of the superior colliculus, the pericentral nucleus of the inferior colliculus and to the ventral nucleus of the basilar pons. Our results suggest that the rat caudal temporal cortex forms part of a pathway that connects visual and auditory cortex with the limbic system, by the way of the amygdala and perirhinal cortex.

Ventral temporal cortex in the rat: Connections of secondary auditory areas Te2 and Te3

The present study in the rat deals with the hodological organization of two cytoarchitectonically distinct areas lying caudoventrally (Te2) or ventrally (Te3) to the primary auditory area (Te1). The afferent and efferent systems of connections were identified by using the properties of retrograde and anterograde transport of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP). Large tracer deposits in the ventral temporal cortex involving Te2, Te3, and the dorsal bank of the perirhinal cortex induced a dense retrograde and anterograde pattern of labeling in the following nuclei of the medial geniculate (MG) complex: caudodorsal (MGCD), dorsal (MGD), medial (MGM), suprageniculate (SG), and peripeduncular area (PPA). The ventral nucleus (MGV) was only slightly labeled in its caudal division. Several extrageniculate structures were also labeled. Retrograde cell labeling occurred in centers giving rise to ascending systems of diffuse projections: locus coeruleus (LC), dorsal raphe nucleus (DR), and basal magnocellular nucleus (B). Slight anterograde labeling was present in the dorsal and external cortices of the inferior colliculus (IC), central gray, deep layers of the superior colliculus (SC), reticular thalamic nucleus (RT), and caudate putamen (CPU). Callosal connections were also noted with the contralateral homotopic cortex. In the cases in which there was a notable extension of the zone of diffusion of the tracer into the dorsal bank of the perirhinal cortex, a characteristic pattern of labeling in the subparafascicular, reuniens and paraventricular thalamic nuclei, mammillary complex, lateral and dorsal hypothalamic nuclei, amygdaloid complex, laterodorsal tegmental nucleus, subiculum, and retrosplenial cortex was displayed. Tracer deposits restricted to Te2 induced a dense labeling of the caudal, ventrolateral MGD, lateral PPA and, to a lesser extent, MGCD. The MGM and SG were only slightly labeled. Extrageniculate afferents essentially consist of sparse projections from LC, DR, and B, whereas efferent fibers are directed to the dorsal cortex of the IC, central gray, deep SC layers, and CPU. Callosal connections were also identified. Following tracer deposits restricted to Te3, dense labeling occurred in the MGD, mostly in its medial division, in the caudal MGM, and in the PPA. The MGCD, SG, and MGV were only sparsely labeled. Extrageniculate afferents arise from LC, DR, and B, and efferents are directed to the RT and dorsal cortex of the IC. Contralateral connections with the homotopic cortical area were also noted. Te2 and Te3 share some degree of similitude in their pattern of connections with the MG complex.(ABSTRACT TRUNCATED AT 400 WORDS)

Cortical responses to speech sounds and their formants in normal infants: maturational sequence and spatiotemporal analysis

Cortical auditory evoked potentials (AEPs) to the synthesized syllables [da[ and [ta[ and to the isolated 3 formants of [da[ were obtained from 32 normal infants studied at monthly intervals from birth through 3 months and at 6 months of age. A bilateral array of 16 electrodes referenced to the mid-occiput permitted a topographic analysis of the cortical AEPs at selected latencies. A differential maturational sequence was seen: a predominantly negative cortical AEP wave form became positive, first over the frontocentral region (around term), and then over the temporal region (at 1-2 months). The timing of these electrophysiological changes coincides with a differential anatomical maturational sequence in the auditory cortex, as myelination and synaptogenesis are more advanced in primary than secondary auditory areas at term. All infants in this study followed this developmental sequence. However, there was no systematic effect of the center frequency of the formant stimuli on the maturational level of their respective cortical AEPs, suggesting a relative maturational equivalence in those regions of auditory cortex responding to stimuli across the frequency range present in human speech. In term infants, an initial midline positivity and bitemporal negativities were asynchronous in their peak and offset latencies. suggesting independent generators for each of these components. In infants from 3 to 6 months of age, cortical AEP wave forms consisted of 2 initial positive peaks followed by a negative peak. While the wave forms were similar over midline and lateral scalp, spatiotemporal analysis revealed differences in the latency of onset, duration and in the spatial extent of these components, again suggesting that 2 bilateral, temporally overlapping generators contribute to the cortical AEP. No systematic topographic difference was observed in the cortical AEPs elicited by each of the 3 formants, which differed in center frequencies.

Effect of restricted cortical lesions on absolute thresholds and aphasia-like deficits in Japanese macaques.

The effect of small bilateral cortical lesions on pure-tone audiograms and on the ability to discriminate between two types of Japanese macaque coo vocalizations was determined in four Japanese macaques (Macaca fuscata). A lesion that included the middle portion of the superior temporal gyrus of both hemispheres, that is, the primary and secondary auditory areas, resulted in a partial hearing loss as well as an inability to discriminate the vocalizations. Lesions that included the ventral portions of the superior temporal gyrus of both hemispheres but spared auditory cortex on one side also resulted in a partial hearing loss but had either a small effect or no effect on the ability to discriminate the vocalizations. Bilateral ablation of the dorsal superior temporal gyrus and adjacent parietal and occipital areas did not appear to result in a hearing loss and had no effect on the ability to discriminate the vocalizations. These results suggest that a hearing loss may be produced by lesions that involve small portions of the ventral two-thirds of the superior temporal gyrus bilaterally although the resulting loss is not as great as that observed with larger lesions. However, the aphasia-like deficit appears to result from a lesion of primary and/or secondary auditory cortex.