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Responses to frequency modulated (FM) sweeps were recorded in rat primary auditory cortex. Forty-four percent of the cells were direction-selective. For speed selectivity, the majority of the cells preferred faster sweeps. The results suggest that rat auditory cortex may be used for processing communication signals of their predators or for detecting spectral changes in acoustic signals.
Motor alimentary conditioned reflexes to acoustic stimuli were elaborated in 27 laboratory rats. After bilateral ablation of the auditory cortex, differentiation of tonal stimuli from amplitude-modulated (AM) signals with a 5 c/s modulation frequency was completely abolished, without its subsequent restoration in six months after the operation. Differentiation of tonal from AM-stimuli with a 50 and 500 c/s modulation frequency, however, not only persists, but is elaborated in animals with a preliminarily ablated auditory cortex. Results of the investigation permit to draw the conclusion that the auditory cortex is the centre where the coding of stimuli with a low frequency of amplitude modulation is completed. Correspondingly, the structural organization of the sensory part of the arc of conditioned responses to different classes of AM-stimuli is discussed.
The sequential development and growth of the subcortico-cortical fibres have been studied by means of acetylcholinesterase (AChE; acetylcholine acetylhydrolase; EC 3.1.1.7) histochemistry on the serial sections obtained from brains of human fetuses ranging from 10 to 28 weeks of gestation. It was found that thalamic and basal telencephalic fibres approaching the fetal auditory cortex have very strong AChE reactivity during a prolonged period of growth. In the youngest fetuses (10.5 weeks) a prominent fibre system was seen emerging from the ventroposterior thalamic territory while the auditory neopallium was free of AChE staining. In older fetuses (16-18 weeks) AChE-positive fibres originating in the ventroposterior thalamus and basal telencephalon penetrate the 'subplate layer' of the auditory cortex. On the basis of AChE reactivity of the 'subplate layer' the auditory cortex can be delineated from the surrounding neocortical areas. In the oldest fetuses (22-28 weeks) AChE-positive fibres of thalamic origin penetrate the developing cortical plate of the auditory cortex.
Understanding the functional organization of the human primary auditory cortex (PAC) is an essential step in elucidating the neural mechanisms underlying the perception of sound, including speech and music. Based on invasive research in animals, it is believed that neurons in human PAC that respond selectively with respect to the spectral content of a sound form one or more maps in which neighboring patches on the cortical surface respond to similar frequencies (tonotopic maps). The number and the cortical layout of such tonotopic maps in the human brain, however, remain unknown. Here we use silent, event-related functional magnetic resonance imaging at 7 Tesla and a cortex-based analysis of functional data to delineate with high spatial resolution the detailed topography of two tonotopic maps in two adjacent subdivisions of PAC. These maps share a low-frequency border, are mirror symmetric, and clearly resemble those of presumably homologous fields in the macaque monkey.
The orientation sound emitted by the Panamanian mustached bat, Pteronotus parnellii rubiginosus, consists of four harmonics. The third harmonic is 6-12 dB weaker than the predominant second harmonic and consists of a long constant-frequency component (CF3) at about 92 kHz and a short frequency-modulated component (FM3) sweeping from about 92 to 74 kHz. Our primary aim is to examine how CF3 and FM3 are represented in a region of the primary auditory cortex anterior to the Doppler-shifted constant-frequency (DSCF) area. Extracellular recordings of neuronal responses from the unanesthetized animal were obtained during free-field stimulation of the ears with pure tones. FM sounds, and signals simulating their orientation sounds and echoes. Response properties of neurons and tonotopic and amplitopic representations were examined in the primary and the anteroventral nonprimary auditory cortex. In the anterior primary auditory cortex, neurons responded strongly to single pure tones but showed no facilitative responses to paired stimuli. Neurons with best frequencies from 110 to 90 kHz were tonotopically organized rostrocaudally, with higher frequencies located more rostrally. Neurons tuned to 92-94 kHz were overpresented, whereas neurons tuned to sound between 64 and 91 kHz were rarely found. Consequently a striking discontinuity in frequency representation from 91 to 64 kHz was found across the anterior DSCF border. Most neurons exhibited monotonic impulse-count functions and responded maximally to sound pressure level (SPL). There were also neurons that responded best to weak sounds but unlike the DSCF area, amplitopic representation was not found. Thus, the DSCF area is quite unique not only in its extensive representation of frequencies in the second harmonic CF component but also in its amplitopic representation. The anteroventral nonprimary auditory cortex consisted of neurons broadly tuned to pure tones between 88 and 99 kHz. Neither tonotopic nor amplitopic representation was observed. Caudal to this area and near the anteroventral border of the DSCF area, a small cluster of FM-FM neurons sensitive to particular echo delays was identified. The responses of these neurons fluctuated significantly during repetitive stimulation.
Ten Japanese macaques were trained to discriminate between two types of Japanese macaque coo vocalizations before and after auditory cortex ablation. Five of the animals were tested following left unilateral ablation, whereas the other five were tested following right unilateral ablation. After postoperative testing, symmetrical lesions were made in the remaining hemisphere in two animals from each group and the effect of bilateral lesions was assessed. The animals were tested using a shock avoidance procedure. Unilateral ablation of left auditory cortex consistently resulted in an initial impairment in the ability to discriminate between the vocalizations with the animals regaining normal performance in 5-15 sessions. In contrast, right unilateral ablation had no detectable effect on the discrimination. Bilateral auditory cortex ablation rendered the animals permanently unable to discriminate between the coos. Although the monkeys could learn to discriminate the coos from noise and from 2- and 4-kHz tones, they had great difficulty in discriminating between the coos and tones in the same frequency range as the coos (i.e., 500 Hz and 1 kHz). The initial impairment following left unilateral lesions indicates that the ability to perceive species-specific vocalizations is lateralized to the left hemisphere. The observation that bilateral lesions abolish the discrimination indicates that the recovery in the left lesion cases was the result of the right hemisphere mediating the discrimination.
Muscarinic acetylcholine receptors are extensively involved in cortical cognition and learning-induced or experience-dependent cortical plasticity. The most abundant muscarinic receptor subtype in the cerebral cortex is the M1 receptor, but little is known about its contribution to experience-dependent plasticity of the adult auditory cortex. We have examined the role of the M1 receptor in experience-dependent plasticity of the auditory cortex in mice lacking the M1 (chrm1) gene. We show here that electrical stimulation of the basal forebrain, a major source of cortical cholinergic inputs, facilitated the auditory responses of cortical neurons in both wild types and M1 mutants. The basal forebrain stimulation alone caused change in the best frequencies of cortical neurons that were significantly greater in M1 mutants. When animals received the paired stimuli of electrical stimulation of the basal forebrain and tone, the frequency tuning of cortical neurons systematically shifted toward the frequency of the paired tone in both wild types and M1 mutants. However, the shift range in M1 mutants was much smaller than that in wild-type mice. Our data suggest that the M1 receptor is important for the experience-dependent plasticity of the auditory cortex.
BACKGROUND AND PURPOSE: Responses of the auditory cortex to sound during sleep have been explored with somewhat discrepant results. The purpose of this study was to investigate the effect of stage 1 sleep on signal intensity changes in the auditory cortex in response to pure tone stimulus measured by functional MR imaging. METHODS: Six sleep-deprived subjects were exposed to a series of echo-planar images for 30-40 minutes. No medication was used to help the subjects go to sleep. A long repetition time of 12 seconds and a 1.9-second clustered multisection acquisition were used to minimize the effect of imager acoustic noise from the preceding acquisition and to make it possible to obtain electroencephalographs between image acquisitions. A pure tone stimulus (beep, 1,000-Hz sine waves, 30-millisecond duration, five beeps per second) was alternated with the baseline every 36 seconds. RESULTS: All subjects fell asleep. The effect of habituation evaluated by comparing the percentage of signal intensity change between the first and second half was not significant. The percentage of signal intensity changes in the right and left transverse temporal gyri were 0.49% and 0.43% during wakefulness and 0.05% and 0.07% during stage 1 sleep. The differences between wakefulness and stage 1 sleep were significant. CONCLUSION: Transition to stage 1 sleep coincides with a decrease in functional MR imaging-determined signal intensity changes in the auditory cortex in response to pure tone stimulus. The limited response of the brain at this stage may protect the brain from sound and facilitate deepening of the sleep stage.
The prospective auditory cortex was analysed in human fetuses at 11--13,5 post-ovulatory weeks with Nissl, Golgi and E.M. techniques. At 11--12 weeks, marginal, cortical plate, intermediate, subventricular and ventricular layers were recognized. Post-migratory neurons with developing dendrites were seen in marginal layer, cortical plate and superficial part of the intermediate zone only. At 12--13,5 weeks the superficial part of the intermediate zone is transformed into the true cortical layer--"subplate layer"--characterized by maturing neurons with growing dendrites, fine axonal arborization and low cell density. The neuronal circuitry elements are thus present very early in the prospective auditory cortex and distributed throughout the deep cortical plate of the "subplate layer" corresponding to the synaptic territory of other areas of the human fetal cortex.
The Zwicker tone (ZT) is defined as an auditory negative afterimage, perceived after the presentation of an appropriate inducer. Typically, a notched noise (NN) with a notch width of 1/2 octave induces a ZT with a pitch falling in the frequency range of the notch. The aim of the present study was to find potential neural correlates of the ZT in the primary auditory cortex of ketamine-anesthetized cats. Responses of multiunits were recorded simultaneously with two 8-electrode arrays during 1 s and over 2 s after the presentation of a white noise (WN) and three NNs differing by the width of the notch, namely, 1/3 octave (NN1), 1/2 octave (NN2), and 2/3 octave (NN3). Both firing rate (FR) and peak cross-correlation coefficient (p) were evaluated for time windows of 500 ms. The cortical units were grouped according to whether their characteristic frequency (CF) was inside ("In" neurons) or outside ("Out" neurons) a 1-octave-wide frequency band centered on the notch center frequency. The ratios between the FRs and the rhos for each NN and the WN condition and for each group of neurons were then statistically evaluated. The ratios of FRs were significantly increased during and after the presentation of the NN for the "In" neurons. In contrast, the changes for the t" neurons were small and most often insignificant. The ratios of the p values differed significantly from 1 in the "In-In" and "In-Out" groups during stimulation as well as after it. We also found that the ps of "Out" neurons were dependent on the type of NN. Potentially, a combination of increased p and increased FR might be a neurophysiological correlate of the ZT.
We recorded evoked magnetic fields from the human auditory cortex to noise/square wave sequences. Two prominent deflections were observed: one 100 msec after the noise onset (N100m) and another 100 msec after the noise/square wave transition (N100m'). The amplitude of N100m' increased with decrease in square wave frequency from 2 kHz to 0.125 kHz and with increase in square wave duration from 4 msec to 200 msec. The latency of N100m' was on the average 23 msec longer for noise durations of 60 msec than 310 msec, whereas the amplitude of N100m' did not change. Increase in interstimulus interval from 1.1 to 8.8 sec enhanced the amplitude of N100m significantly more than that of N100m'. The small interaction between N100m and N100m' and their different recovery cycles suggest that different activation patterns underlie these two 100 msec responses at the auditory cortex.
1. Glutamatergic excitation and gamma-aminobutyric acid (GABA)-ergic inhibition in layers II and III of the auditory cortex of anaesthetized guinea-pigs were recorded optically using a voltage-sensitive dye RH795 and a 12 x 12 photodiode array. 2. After contralateral ear stimulation with pure tones, transient excitatory responses followed by inhibitory responses were observed in fields A (primary) and DC of the auditory cortex. The area of the excitatory responses was sandwiched or surrounded by the areas of the inhibitory responses. 3. Optically recorded excitatory responses to pure tones had two components: a component sensitive to 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX), a non-N-methyl-D-aspartate (non-NMDA) receptor antagonist, and a component sensitive to 2-amino-5-phosphono-valerate (APV), an NMDA receptor antagonist. Application of CNQX (5 microM) to the auditory cortex suppressed an early, but not a late, phase of the excitation; application of APV (100 microM) had the opposite effect. Concomitant application of CNQX and bicuculline methiodide (BMI, 4 microM), a GABAA receptor antagonist, increased the amplitude of the late phase 4-fold. This enhanced response was suppressed by APV. 4. These results indicate that (i) auditory cortical excitatory responses are mediated by both non-NMDA and NMDA receptors, (ii) inhibition is mediated by GABAA receptors, (ii) the excitatory bands are sandwiched or surrounded by GABAA receptor-mediated inhibitory areas and (iv) GABAA receptors effectively inhibit the NMDA, but not the non-NMDA, receptor-mediated excitation.
Mongolian gerbils (Meriones unguiculatus) were trained in a shuttle box to discriminate the direction in frequency-modulated tones (FM). Whereas control animals easily acquired FM discrimination, animals with auditory cortex lesion on the right side showed considerable difficulties in learning this task. The discrimination performance of gerbils with left auditory cortex lesion, however, was not different from controls. This study, suggesting that the right auditory cortex plays a dominant role in FM discrimination learning in gerbils, describes a useful animal model for investigation of the basic mechanisms underlying hemispheric asymmetries in auditory perception.
Single-unit responses to tone pip stimuli were isolated from numerous microelectrode penetrations of auditory cortex (under ketamine anesthesia) in the developing chinchilla (laniger), a precocious mammal. Results are reported at postnatal day 3 (P3), P15, and P30, and from adult animals. Hearing sensitivity and spike firing rates were mature in the youngest group. The topographic representation of sound frequency (tonotopic map) in primary and secondary auditory cortex was also well ordered and sharply tuned by P3. The spectral-temporal complexity of cortical receptive fields, on the other hand, increased progressively (past P30) to adulthood. The (purported) refinement of initially diffuse tonotopic projections to cortex thus seems to occur in utero in the chinchilla, where external (and maternal) sounds are considerably attenuated and might not contribute to the mechanism(s) involved. This compares well with recent studies of vision, suggesting that the refinement of the retinotopic map does not require external light, but rather waves of (correlated) spontaneous activity on the retina. In contrast, it is most probable that selectivity for more complex sound features, such as frequency stacks and glides, develops under the influence of the postnatal acoustic environment and that inadequate sound stimulation in early development (e.g., due to chronic middle ear disease) impairs the formation of the requisite intracortical (and/or subcortical) circuitry.
Induction of c-Fos in the auditory cortex of gerbils was investigated immunocytochemically 1 h after single, triple or 1 h continuous stimulation with a series of narrow band frequency-modulated tone bursts. With single stimulation c-Fos immunoreactive neurons were chiefly found in the primary auditory field (AI), where they formed a narrow frequency-specific column across layers II-VI. Side-band-like patterns adjacent to this column appeared characteristically with triple stimulation. Immunoreactive cell density in the anterior auditory field and the caudal fields was sparse and location not frequency specific with single or triple stimulation. Spatial comparisons of c-Fos immunoreactive neuron density with 2-deoxy-2-fluoro-D-glucose (FDG) autoradiography in the same animals after 1 h of stimulation revealed spreading of c-Fos expression in neurons across the tonotopic maps of the AI and in the rostral and caudal fields of the auditory cortex. The pattern of the highest density of c-Fos labelled cells in the AI still matched the peak labelling of FDG autoradiographs. The results show that the postsynaptic marker c-Fos reflects the frequency representation in the AI with single or triple stimulation yet with a higher spatial resolution than the deoxyglucose technique. Longer stimulation causes nontonotopic intracortical spreading of the c-Fos-inducing message, a phenomenon potentially reflecting the effects of cooperativity in the maps.
"Physiological memory" is enduring neuronal change sufficiently specific to represent learned information. It transcends both sensory traces that are detailed but transient and long-term physiological plasticities that are insufficiently specific to actually represent cardinal details of an experience. The specificity of most physiological plasticities has not been comprehensively studied. We adopted receptive field analysis from sensory physiology to seek physiological memory in the primary auditory cortex of adult guinea pigs. Receptive fields for acoustic frequency were determined before and at various retention intervals after a learning experience, typified by single-tone delay classical conditioning, e.g., 30 trials of tone-shock pairing. Subjects rapidly (5-10 trials) acquire behavioral fear conditioned responses, indexing acquisition of an association between the conditioned and the unconditioned stimuli. Such stimulus-stimulus association produces receptive field plasticity in which responses to the conditioned stimulus frequency are increased in contrast to responses to other frequencies which are decreased, resulting in a shift of tuning toward or to the frequency of the conditioned stimulus. This receptive field plasticity is associative, highly specific, acquired within a few trials, and retained indefinitely (tested to 8 weeks). It thus meets criteria for "physiological memory." The acquired importance of the conditioned stimulus is thought to be represented by the increase in tuning to this stimulus during learning, both within cells and across the primary auditory cortex. Further, receptive field plasticity develops in several tasks, one-tone and two-tone discriminative classical and instrumental conditioning (habituation produces a frequency-specific decrease in the receptive field), suggesting it as a general process for representing the acquired meaning of a signal stimulus. We have proposed a two-stage model involving convergence of the conditioned and unconditioned stimuli in the magnocellular medial geniculate of the thalamus followed by activation of the nucleus basalis, which in turn releases acetylcholine that engages muscarinic receptors in the auditory cortex. This model is supported by several recent findings. For example, tone paired with NB stimulation induces associative, specific receptive field plasticity of at least a 24-h duration. We propose that physiological memory in auditory cortex is not "procedural" memory, i.e., is not tied to any behavioral conditioned response, but can be used flexibly.
The peculiarities of the AChE distribution were investigated in the intact cat auditory cortex and during early period of its neuronal isolation. It is shown that in the isolated cortex slab the staining of the AChE containing fibre disappeared from the neuropile, while in the intact cortex it was well pronounced. AChE accumulation was observed in the proximal parts of the transsected thalamo-cortical fibres. It is supposed that the AChE-containing fibres in the auditory cortex belong to nonspecific thalamic inputs.
Spiking activity was recorded from cat auditory cortex using multi-electrode arrays. Cross-correlograms were calculated for spikes recorded on separate microelectrodes. The pair-wise cross-correlation matrix was constructed for the peak values of the correlograms. Hierarchical clustering was performed on the cross-correlation matrix for six stimulus conditions. These were silence, three multi-tone stimulus ensembles with different spectral densities, low-pass amplitude-modulated noise, and Poisson-distributed click trains that each lasted 15 min. The resulting neuron clusters reflect patches in cortex of up to several mm(2) in size that expand and contract in response to different stimuli. Cluster positions and size were very similar for spontaneous activity and multi-tone stimulus-evoked activity but differed between those conditions and the noise and click stimuli. Cluster size was significantly larger in posterior auditory field (PAF) compared with primary auditory cortex (AI), whereas the fraction of common spikes (within a 10-ms window) across all electrode activity participating in a cluster was significantly higher in AI compared with PAF. Clusters crossed area boundaries in <5% of the cases were simultaneous recording were made in AI and PAF. Clusters are therefore similar to but not synonymous with the traditional view of neural assemblies. Common-spike spectrotemporal receptive fields (STRFs) were obtained for common-spike activity and all-spike activity within a cluster. Common-spike STRFs had higher signal-to-noise ratio than all-spike STRFs and showed generally spectral and temporal sharpening. The coincident and noncoincident output of the clusters could potentially act in parallel and may serve different modes of stimulus coding.