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The auditory cortex of the mouse: connections of the ultrasonic field.

The cortical and subcortical connections of the ultrasonic field (UF) of the auditory cortex of the house mouse (Mus musculus) were studied by using retrograde and anterograde transport of horseradish peroxidase (HRP). Small amounts of HRP were locally injected into the electrophysiologically defined UF. Superficial (layer I-IV) and deep (layer IV-VI) injections were prepared. Superficial injections led to labelling of both cells (retrograde) and terminals (anterograde) in areas of the ipsilateral primary and secondary auditory cortex and in its dorsoposterior field, in an ipsilateral dorsal association area (patches of label), probably in ipsilateral secondary somatosensory cortex, in the contralateral homotopic UF, and in the ipsilateral medial geniculate body (MGBv, MGBd, and MGBm) and caudal posterior nucleus complex. Deep injections showed the same connectivities as superficial ones and, in addition, terminals in the very caudal caudatoputamen, in the nucleus limitans and the nucleus reticularis of the thalamus, in the rostral pole, the dorsomedial, and lateral nucleus of the inferior colliculus, in the stratum griseum intermediale of the superior colliculus, and in a pontine nucleus ventromedial of the lateral lemniscus. All these projections occurred only ipsilaterally. The majority of connections, except those with the nucleus limitans, superior colliculus and pontine nucleus, suggest that UF is part of the primary anditory cortex (AI) and/or of the anterior anditory field (AAF) of the auditory cortex. Since UF has no regular tonotopy, this has important implications for the functional role that AI/AAF can have in communication-sound analysis.

Animals↗

Functional MRI of human auditory cortex using block and event-related designs.

fMRI of human auditory cortex response to sinusoidal tones of 200, 1000, and 3000 Hz was evaluated using block design and conventional and "silent" event-related designs. Conventional event-related fMRI revealed the timecourse of the BOLD response (approximately 5 sec to peak, approximately 4 sec full-width-half-max, and approximately 14 sec recovery to baseline). Both event-related, but not block, designs provided evidence for tonotopic organization in auditory cortex. Sources of low-frequency activation were more lateral and anterior than the sources of high-frequency activation (P < or = 0.05). In the block designs, repeated rapid stimulus presentation and the co-incidence of scanner noise preclude definition of tonotopic organization revealed in event-related approaches. Magn Reson Med 45:254-260, 2001.

Acoustic Stimulation↗

Location of rabbit auditory cortex and description of single unit activity.

The following data were obtained from microelectrode exploration of rabbit auditory cortex (AC). (1) Location of AC--Major portions of the rabbit's right hemisphere were stereotaxically mapped for auditory responsive and non-responsive regions. Auditory responsive regions were found to lie behind the rhinal sulcus, extending from its midpoint to its inferior border. (2) Frequency selectivity--Characteristic frequencies (CFs) ranged from 0.4 to 33 kHz. Sharp, multipeaked and broad tuning curves were described. (3) Response patterns--A variety of post-stimulus-time (PST) histogram shapes were obtained in response to tonal stimuli. The response of a given neuron often varied with changes in stimulus frequency or intensity. (4) Latency--Neural response latencies ranged from 10 to 100 msec following stimulus onset. (5) Spontaneous rate--In the absence of acoustic stimulation, spike discharges ranged from less than 1 spike/sec to 48 spikes/sec. Spontaneous rates of less than 5 spikes/sec were measured in 42% of the neurons. Rates in excess of 15 spikes/sec occurred in fewer than 18% of the cells examined. (6) Response variability--Stimulus-evoked activity was seen to vary over time. Changes consisted of an overall increase or attenuation of discharge rate, as well as modifications of selected portions of the response pattern (PST histogram shape). The characteristics of single unit activity in rabbit auditory cortex are consistent with findings reported in other species.

Animals↗

The personalized auditory cortex of the mustached bat: adaptation for echolocation.

1. In the mustached bat, Pteronotus parnellii, the "resting" frequency of the constant-frequency component of the second harmonic (CF2) of the orientation sound (biosonar signal) is different among individuals within a range from 59.69 to 63.33 kHz. The standard deviation of CF2 resting frequency is 0.091 kHz on the average for individual bats. The male's CF2 resting frequency (61.250 +/- 0.534 kHz, n = 58) is 1.040 kHz lower than the female's (62.290 +/- 0.539 kHz, n = 58) on the average. Females' resting frequencies measured in December are not different from those measured in April when almost all of them are pregnant. Therefore, the orientation sound is sexually dimorphic. 2. In the DSCF (Doppler-shifted CF processing) area of the auditory cortex, tonotopic representation differs among individual bats. The higher the CF2 resting frequency of the bat's own sound, the higher the frequencies represented in the DSCF area of that bat. There is a unique match between the tonotopic representation and the CF2 resting frequency. This match indicates that the auditory cortex is "personalized" for echolocation and that the CF2 resting frequency is like a signature of the orientation sound. 3. If a bat's resting frequency is normalized to 61.00 kHz, the DSCF area overrepresents 60.6-62.3 kHz. The central region of this overrepresented band is 61.1-61.2 kHz. This focal band matches the "reference" frequency to which the CF2 frequency of a Doppler-shifted echo is stabilized by Doppler-shift compensation. 4. Since DSCF neurons are extraordinarily sharply tuned in frequency, the personalization of the auditory cortex or system is not only suited for the detection of wing beats of insects, but also for the reduction of the masking effect on echolocation of consepecific's biosonar signals. 5. Because the orientation sound is sexually dimorphic and the auditory cortex is personalized, the tonotopic representation of the auditory cortex is also sexually dimorphic.

Acoustic Stimulation↗

Comparison of evoked potentials and high-frequency (gamma-band) oscillating potentials in rat auditory cortex.

1. Transient and steady-state (40 Hz) evoked potentials, as well as spontaneous and click-evoked gamma-band oscillations, were recorded from 15 lightly anesthetized rats using an 8 x 8 electrode epipial array covering auditory cortex and adjacent areas to determine and compare the spatiotemporal distributions of these four phenomena. 2. The transient evoked response replicated earlier findings in our laboratory, consisting of an initial biphasic sharp wave in area 41, a similar but delayed biphasic sharp wave in area 36, and more widely distributed slow-wave components. Spatiotemporal analysis supported a model of parallel and asynchronous activation of distinct groups of thalamocortical projections underlying the neurogenesis of these temporal components of the middle-latency auditory evoked potential (MAEP) complex. 3. The 40-Hz response to click trains was superimposed on a steady potential shift (SP), both of which were localized within primary auditory cortex. Epipial distributions of the SP were similar to those of the shortest-latency negative peak in area 41 recorded in the same animals, suggesting similar neural generators. The 40-Hz response was more focal and dissimilar from the SP and any other temporal components of the MAEP complex, suggesting that a unique subpopulation of cells underlies its neurogenesis. 4. Spontaneous gamma-band activity, as assessed by power spectrum analysis, was localized to primary and secondary auditory cortex but had a variable distribution between rats that did not conform to the cytoarchitectonic boundaries within subdivisions of this region. Digital movies computed for individual bursts of gamma-activity indicated a high degree of spatiotemporal variability within and between bursts. 5. Single-trial spectral analysis of click responses indicated an inhibition of gamma-band oscillations during most of the MAEP complex, with subsequent enhanced gamma-activity during the 300- to 350-ms slow-wave component that outlasted the MAEP by approximately 500 ms. The epipial distributions of prestimulus and enhanced poststimulus gamma-oscillations were the same. In contrast to the 40-Hz response to click trains, phase-locking of gamma-oscillations by the single click stimulus was not observed. 6. These results suggest that both the MAEP complex and the steady-state 40-Hz response with its associated SP are highly stereotyped in lightly anesthetized rodent cortex. Their spatiotemporal distributions are probably determined in large part by asynchronous activation of parallel thalamocortical projection systems. Our data suggest no direct link between either the MAEP or the steady-state 40-Hz response to spontaneous or evoked gamma-band oscillations in auditory cortex.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Functional specialization in auditory cortex: responses to frequency-modulated stimuli in the cat's posterior auditory field.

The mammalian auditory cortex contains multiple fields but their functional role is poorly understood. Here we examine the responses of single neurons in the posterior auditory field (P) of barbiturate- and ketamine-anesthetized cats to frequency-modulated (FM) sweeps. FM sweeps traversed the excitatory response area of the neuron under study, and FM direction and the linear rate of change of frequency (RCF) were varied systematically. In some neurons, sweeps of different sound pressure levels (SPLs) also were tested. The response magnitude (number of spikes corrected for spontaneous activity) of nearly all field P neurons varied with RCF. RCF response functions displayed a variety of shapes, but most functions were of low-pass characteristic or peaked at rather low RCFs (<100 kHz/s). Neurons with strong responses to high RCFs (high-pass or nonselective RCF response function characteristics) all displayed spike count-SPL functions to tone burst onsets that were monotonic or weakly nonmonotonic. RCF response functions and best RCFs often changed with SPL. For most neurons, FM directional sensitivity, quantified by a directional sensitivity (DS) index, also varied with RCF and SPL, but the mean and width of the distribution of DS indices across all neurons was independent of RCF. Analysis of response timing revealed that the phasic response of a neuron is triggered when the instantaneous frequency of the sweep reaches a particular value, the effective Fi. For a given neuron, values of effective Fi were independent of RCF, but depended on FM direction and SPL and were associated closely with the boundaries of the neuron's frequency versus amplitude response area. The standard deviation (SD) of the latency of the first spike of the response decreased with RCF. When SD was expressed relative to the rate of change of stimulus frequency, the resulting index of frequency jitter increased with RCF and did so rather uniformly in all neurons and largely independent of SPL. These properties suggest that many FM parameters are represented by, and may be encoded in, orderly temporal patterns across different neurons in addition to the strength of responses. When compared with neurons in primary and anterior auditory fields, field P neurons respond better to relatively slow FMs. Together with previous studies of responses to modulations of amplitude, such as tone onsets, our findings suggest more generally that field P may be best suited for processing signals that vary relatively slowly over time.

Acoustic Stimulation↗

Attention modulates activity in the primary and the secondary auditory cortex: a functional magnetic resonance imaging study in human subjects.

Using functional magnetic resonance imaging, ten healthy subjects were scanned whilst listening to consonant-vowel syllables under three different conditions: (i) a 'no-attention' condition required subjects to ignore the stimuli; (ii) an 'attend' condition requiring attentive listening to stimuli; (iii) a 'detect' condition requiring detection of a specific target syllable. Hemodynamic responses were measured in the primary and secondary auditory cortex. These three conditions were associated with significantly different activations in the primary and secondary auditory cortex. The strongest activations were found for the 'detect' condition, followed by the 'attend' condition. The weakest activation was evident during the 'no-attention' condition. There were also stronger activations in the left hemisphere and within the primary auditory cortex. These results suggest that the primary and secondary auditory cortex play a main role in the selective attention.

Acoustic Stimulation↗

Spectral envelope coding in cat primary auditory cortex: linear and non-linear effects of stimulus characteristics.

Electrophysiological studies in mammal primary auditory cortex have demonstrated neuronal tuning and cortical spatial organization based upon spectral and temporal qualities of the stimulus including: its frequency, intensity, amplitude modulation and frequency modulation. Although communication and other behaviourally relevant sounds are usually complex, most response characterizations have used tonal stimuli. To better understand the mechanisms necessary to process complex sounds, we investigated neuronal responses to a specific class of broadband stimuli, auditory gratings or ripple stimuli, and compared the responses with single tone responses. Ripple stimuli consisted of 150-200 frequency components with the intensity of each component adjusted such that the envelope of the frequency spectrum is sinusoidal. It has been demonstrated that neurons are tuned to specific characteristics of those ripple stimulus including the intensity, the spacing of the peaks, and the location of the peaks and valleys (C. E. Schreiner and B. M. Calhoun, Auditory Neurosci., 1994; 1: 39-61). Although previous results showed that neuronal response strength varied with the intensity and the fundamental frequency of the stimulus, it is shown here that the relative response to different ripple spacings remains essentially constant with changes in the intensity and the fundamental frequency. These findings support a close relationship between pure-tone receptive fields and ripple transfer functions. However, variations of other stimulus characteristics, such as spectral modulation depth, result in non-linear alterations in the ripple transformation. The processing between the basilar membrane and the primary auditory cortex of broadband stimuli appears generally to be non-linear, although specific stimulus qualities, including the phase of the spectral envelope, are processed in a nearly linear manner.

Acoustic Stimulation↗

Cellular mechanisms of thalamically evoked gamma oscillations in auditory cortex.

The purpose of this study was to clarify the neurogenesis of thalamically evoked gamma frequency (approximately 40 Hz) oscillations in auditory cortex by comparing simultaneously recorded extracellular and intracellular responses elicited with electrical stimulation of the posterior intralaminar nucleus of the thalamus (PIL). The focus of evoked gamma activity was located between primary and secondary auditory cortex using a 64-channel epipial electrode array, and all subsequent intracellular recordings and single-electrode field potential recordings were made at this location. These data indicate that PIL stimulation evokes gamma oscillations in auditory cortex by tonically depolarizing pyramidal cells in the supra- and infragranular layers. No cells revealed endogenous membrane properties capable of producing activity in the gamma frequency band when depolarized individually with injected current, but all displayed both sub- and supra-threshold responses time-locked to extracellular fast oscillations when the population was depolarized by PIL stimulation. We propose that cortical gamma oscillations may be produced and propagated intracortically by network interactions among large groups of neurons when mutually excited by modulatory input from the intralaminar thalamus and that these oscillations do not require specialized pacemaker cells for their neurogenesis.

Acoustic Stimulation↗

Learning-induced physiological memory in adult primary auditory cortex: receptive fields plasticity, model, and mechanisms.

It is well established that the functional organization of adult sensory cortices, including the auditory cortex, can be modified by deafferentation, sensory deprivation, or selective sensory stimulation. This paper reviews evidence establishing that the adult primary auditory cortex develops physiological plasticity during learning. Determination of frequency receptive fields before and at various times following aversive classical conditioning and instrumental avoidance learning in the guinea pig reveals increased neuronal responses to the pure tone frequency used as a conditioned stimulus (CS). In contrast, responses to the pretraining best frequency and other non-CS frequencies are decreased. These opposite changes are often sufficient to shift cellular tuning toward or even to the frequency of the CS. Learning-induced receptive field (RF) plasticity (i) is associative (requires pairing tone and shock), (ii) highly specific to the CS frequency (e.g., limited to this frequency +/- a small fraction of an octave), (iii) discriminative (specific increased response to a reinforced CS+ frequency but decreased response to a nonreinforced CS- frequency), (iv) develops extremely rapidly (within 5 trials, the fewest trials tested), and (v) is retained indefinitely (tested to 8 weeks). Moreover, RF plasticity is robust and not due to arousal, but can be expressed in the deeply anesthetized subject. Because learning- induced RF plasticity has the major characteristics of associative memory, it is therefore referred to as "physiological memory". We developed a model of RF plasticity based on convergence in the auditory cortex of nucleus basalis cholinergic effects acting at muscarinic receptors, with lemniscal and nonlemniscal frequency information from the ventral and magnocellular divisions of the medial geniculate nucleus, respectively. In the model, the specificity of RF plasticity is dependent on Hebbian rules of covariance. This aspect was confirmed in vivo using microstimulation techniques. Further, the model predicts that pairing a tone with activation of the nucleus basalis is sufficient to induce RF plasticity similar to that obtained in behavioral learning. This prediction has been confirmed. Additional tests of the model are described. RF plasticity is thought to translate the acquired significance of sound into an increased frequency representation of behaviorally important stimuli.

Adult↗

Parallels between timing of onset responses of single neurons in cat and of evoked magnetic fields in human auditory cortex.

Sound onsets constitute particularly salient transients and evoke strong responses from neurons of the auditory system, but in the past, such onset responses have often been analyzed with respect to steady-state features of sounds, like the sound pressure level. Recent electrophysiological studies of single neurons from the auditory cortex of anesthetized cats have revealed that the timing and strength of onset responses are shaped by dynamic stimulus properties at their very onsets. Here we demonstrate with magnetoencephalography that stimulus-response relationships very similar to those of the single neurons are observed in two onset components, N100m and P50m, of auditory evoked magnetic fields (AEFs) from the auditory cortex of awake humans. In response to tones shaped with cosine-squared rise functions, N100m and P50m peak latencies vary systematically with tone level and rise time but form a rather invariant function of the acceleration of the envelope at tone onset. Hence N100m and P50m peak latencies, as well as peak amplitudes, are determined by dynamic properties of the stimuli within the first few milliseconds, though not necessarily by acceleration. The changes of N100m and P50m peak latencies with rise time and level are incompatible with a fixed-amplitude threshold model. The direct comparison of the neuromagnetic and single-neuron data shows that, on average, the variance of the neuromagnetic data is larger by one to two orders of magnitude but that favorable measurements can yield variances as low as those derived from neurons with mediocre precision of response timing. The striking parallels between the response timing of single cortical neurons and of AEFs provides a stronger link between single neuron and population activity.

Acceleration↗

Basal forebrain stimulation modifies auditory cortex responsiveness by an action at muscarinic receptors.

We have hypothesized that auditory cortex plasticity involves modification of thalamocortical transmission by basal forebrain (BF) cholinergic neurons, and that this action may involve muscarinic receptors. In a first test of this hypothesis, we report that BF stimulation can suppress or facilitate, depending on the intensity of stimulation, auditory cortical responses elicited by thalamic stimulation. BF-mediated facilitation is antagonized by atropine, implicating muscarinic receptors. These data suggest that BF cholinergic neurons functionally modify auditory cortex by regulating thalamocortical transmission.

Animals↗

Effects of salicylate on neural activity in cat primary auditory cortex.

The effect of systemically applied salicylate on single-unit firing activity in primary auditory cortex was investigated in six cats. A dose of 200 mg/kg sodium salicylate was administered intraperitoneally, and recordings from the same units were performed prior to application and continuously up to, on average. 6 h after administration. Local field potentials were used to track the threshold shifts and general input-output (I/O) behavior following salicylate administration. All animals showed 20-30 dB of threshold shift about 2 h after administration and showed no recovery during the following 4 h. I/O curves were invariably of the recruitment type. Significant changes were found in spontaneous firing rates for two groups of unit separately. Low-spontaneous rate units (initial firing rate < 1 spike/s) showed an increase in spontaneous rate and high-spontaneous rate units (initial firing rate > 1 spike/s) showed a decrease in spontaneous firing rate. There were no significant changes in modal and mean values for interspike-interval (ISI) histograms. The duration-to-rebound peak in the autocorrelation function for spontaneous firings was prolonged significantly after salicylate administration. Peak cross-correlation coefficients for the firing patterns of simultaneously recorded cells showed no significant change but the correlogram's central peak was significantly narrower after salicylate application. The percentage of firings occurring in bursts showed no significant change after administration of salicylate. The best modulation frequency in response to stimulation with periodic click trains decreased after administration. Both the changes in the spontaneous autocorrelogram and in the temporal modulation transfer function suggest a prolongation in the duration of the Ca(2+)-activated K+ conductance of the cortical pyramidal cells following salicylate. This suggests that salicylates affect both the auditory periphery and the auditory cortex.

Acoustic Stimulation↗

Effects of unilateral ablation of auditory cortex on monaural cat's ability to localize sound.

1. Cats with one cochlea destroyed were trained to localize sound. After behavioral measures of the animal's accuracy of localization were made, cortical auditory areas were ablated unilaterally. 2. The results showed: a) like binaural localization, monaural localization of sound in space, as measured by the ability of an animal to move toward a sound source, depends on integrity of auditory cortex; b) it is only ablation of cortex contralateral to the functional ear that seriously affects localizing behavior; ablation of cortex ipsilateral to the intact cochlea has little or no effect on localizing behavior. 3. To explain the results, we suggest that auditory cortex is essential for an organized perception of space including the relation of the animal's position to other objects in space. We also suggest that auditory cortex contralateral to a given ear is necessary in order for the animal to recognize that a stimulus is presented to that ear of, when both ears are intact, to recognize that the stimulus to the given ear differs in some way (intensity, time of arrival, sequential arrangement of sounds) from the stimulus to the opposite ear.

Animals↗

Context modulates processing of speech sounds in the right auditory cortex of human subjects.

Using whole-head magnetoencephalography, we studied how context affects processing of speech sounds by recording the magnetic counterpart (MMNm) of the mismatch negativity (MMN) event-related potential, which is elicited in the auditory cortex by a contrast between infrequent and frequent auditory stimuli. In Experiment 1, in the right, but not in the left, hemisphere auditory cortex, a contrast of consonant-vowel syllables elicited a stronger MMNm when presented within a word context than when occurring alone. Further, in Experiment 2, these same syllable contrasts were processed in the right auditory cortex more strongly when embedded in a non-speech sound context than when presented alone. This facilitation thus was insensitive to the phonetic and semantic features of the context but might nevertheless reflect the participation of the right hemisphere in speech perception.

Acoustic Stimulation↗

Native language, gender, and functional organization of the auditory cortex.

Whole-head magnetoencephalography was employed in 40 normal subjects to investigate whether the basic functional organization of the auditory cortex varies with linguistic environment. Robust activations of the bilateral supratemporal auditory cortices to 1-kHz pure tones, maximum at about 100 ms after stimulus onset, were studied in Finnish and German female and male subject groups with monolingual background. Activations elicited by the tones were mutually indistinguishable in German and Finnish women. In contrast, German men showed significantly stronger auditory responses to pure tones in the left, language-dominant hemisphere than Finnish men. We discuss the possibility that the prominent left-hemisphere activation in German males reflects higher frequency resolution required for distinguishing between German than Finnish vowels and that the clear effect of native language in male but not in female auditory cortex derives from more pronounced functional lateralization in men. The present data suggest that the influence of native language can extend to auditory cortical processing of pure-tone stimuli with no linguistic content and that this effect is conspicuous in the male brain.

Acoustic Stimulation↗

Cytoarchitectonic organization of the human auditory cortex.

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.

Animals↗

[Effect of white noise on the rat auditory cortex ultrastructure].

Changes in ultrastructure of neurones and glial cells of rat auditory cortex were studied after exposure to white noise for 21 days. Appearance of light neurones with peripheral chromatolysis and the increased number of lysosomes, pigment granules and vacuoles were recorded both in the neurones and glial cells. It is concluded that exposure to white noise in chronic experiments is the pathogenic factor which causes destruction in the neurons and glial cells of the auditory cortex.

Auditory Cortex↗