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Tuning properties of auditory cortex cells in the awake squirrel monkey.

Pure tone bursts elicited in primary auditory cortex (AI) cells of the awake squirrel monkey a wide range of response patterns which consisted of one or more excitatory or inhibitory temporal response components. In almost 60% of these cells, response patterns were frequency and/or intensity dependent. Response components such as early and late onset excitation, offset excitation and on-off excitation; as well as tonic excitation or inhibition often varied independently with changes in these stimulus parameters. Individual cells were therefore considered as multiple bandpass filters, and each discrete response component was analyzed separately for its tuning properties. A correlation between best frequencies of the various excitatory components (BEF), and between BEFs and best frequencies of inhibitory components (BIF), in cells which responded with more than one discrete response component, disclosed a significantly higher correlation between BEF/BIF pairs compared with BEF/BEF pairs, presumably reflecting certain "lateral inhibition like" processes. Applying Q10dB factor, and square root of Hf-square root of Lf bandwidth at 10 dB above threshold, as measures of the "sharpness" of response areas, revealed that approximately 65% of all response areas could be defined as "narrow" by either one of these 2 measures, with no distinction, in that regard, between excitatory and inhibitory components. The average response bandwidths of the narrowly and the broadly tuned components, at 10 dB above threshold, were 0.4 +/- 0.18 and 1.42 +/- 0.68 octaves respectively. A comparison with the medial geniculate body (MGB) of the squirrel monkey, applying the square root of Hf-square root of Lf measure of sharpness of tuning, showed a significantly higher proportion of narrow response areas in the AI. "Narrow" response areas in both these regions were equally narrow, whereas the "broad" response areas of MGB cells were significantly broader. These results suggest a sharpening of response areas throughout the geniculo-cortical transformation.

Animals↗

Improved auditory cortex imaging using clustered volume acquisitions.

The effects of the noise of echo-planar functional magnetic resonance imaging on auditory cortex responses were compared for two methods of acquiring functional MR data. Responses observed with a distributed volume acquisition sequence were compared to those obtained with a clustered volume acquisition sequence. In the former case, slices from the volume were acquired at equal intervals within the repetition time, whereas the latter acquired all slices in rapid succession at the end of the imaging period. The clustered volume acquisition provides a period of quiet during which a stimulus may be presented uninterrupted and uncontaminated by the noise of echo-planar imaging. Both sequences were implemented on a General Electric Signa imager retrofitted for echo-planar imaging by Advanced NMR Systems, Inc. The sequences were used to acquire 60 images per slice of a fixed volume of cerebral cortex while subjects were presented an instrumental music stimulus in an On vs. Off paradigm. Data were acquired for both sequences using TR values of 2, 3, 4, 6 and 8 sec. The clustered volume acquisition sequence was found to yield greater measures of dynamic range (percent signal change, mean statistical power per unit imaging time) across the tested range of TR values. Observations of more consistent spatial extent of responses, greater mean signal changes, and higher and more consistent values of mean t-statistic per unit imaging time demonstrate the efficacy of using a clustered volume acquisition for fMRI of auditory cortex.

Adult↗

Functional organization of auditory cortex in the mongolian gerbil (Meriones unguiculatus). I. Electrophysiological mapping of frequency representation and distinction of fields.

The frequency representation within the auditory cortex of the anaesthetized Mongolian gerbil (Meriones unguiculatus) was studied using standard microelectrode (essentially multiunit) mapping techniques. A large tonotopically organized primary auditory field (AI) was identified. High best frequencies (BFs) were represented rostrally and low BFs caudally along roughly dorsoventrally oriented isofrequency contours. Additional tonotopic representations were found adjacent to AI. Rostral to AI was a smaller field with a complete tonotopic gradient reversed with respect to that in AI (mirror image representation) and was termed the anterior auditory field (AAF). BFs in the range from 0.1 to 43 kHz, apparently covering the hearing range of the Mongolian gerbil, were found in AI and AAF. Units in these two core fields responded to narrow frequency ranges with short latencies. Ventral to the common high-frequency border to AAF and AI, a rapid transition to very low BFs suggested the presence of a ventral field (V). Caudal to AI two small tonotopically organized fields were identified, a dorsoposterior field (DP) and a ventroposterior field (VP). The VP showed a tonotopic organization mirror imaged to that of AI, i.e. low frequencies were represented rostrally near the caudal border of AI, and high frequencies caudally. The DP showed a concentric frequency organization with high BFs located in the centre. Units in DP and VP fired less strongly, with considerably longer latencies, and responded to a broader range of frequencies than units in AI and AAF. Dorsocaudal to AI a dorsal field (D) was identified, harbouring units that responded to very broad ranges of frequencies. A tonotopic organization of field D could not be discerned. In the border region of AI and D, low-frequency responses were similar to those found in parts of AI and AAF, but without a clear-cut tonotopic organization. This region was termed Ald. The two core fields AI and AAF appeared to be located within the koniocortex, while the remaining fields lay outside. Our data show that the organization of the gerbil auditory cortex is highly elaborate, with parcellation into fields as complex as in cat or primates.

Acoustic Stimulation↗

Critical period window for spectral tuning defined in the primary auditory cortex (A1) in the rat.

Experience-dependent plasticity during development results in the emergence of highly adapted representations of the external world in the adult brain. Previous studies have convincingly shown that the primary auditory cortex (A1) of the rat possesses a postnatal period of sensory input-driven plasticity but its precise timing (onset, duration, end) has not been defined. In the present study, we examined the effects of pure-tone exposure on the auditory cortex of developing rat pups at different postnatal ages with a high temporal resolution. We found that pure-tone exposure resulted in profound, persistent alterations in sound representations in A1 only if the exposure occurred during a brief period extending from postnatal day 11 (P11) to P13. We also found that postnatal sound exposure in this epoch led to striking alterations in the cortical representation of sound intensity.

Aging↗

Pathways from auditory cortex to the cochlear nucleus in guinea pigs.

The inferior colliculus (IC) and superior olivary complex (SOC) are important sources of descending pathways to the cochlear nucleus. The IC and SOC are also targets of direct projections from the auditory cortex but it is not known if cortical axons contact the cells that project to the cochlear nucleus. Multi-labeling techniques were used to address this question in guinea pigs. A fluorescent anterograde tracer was injected into temporal cortex to label corticofugal axons. Different fluorescent tracers were injected into one or both cochlear nuclei to label olivary and collicular cells. The brain was subsequently processed for fluorescence microscopy and the IC and SOC were examined for apparent contacts between cortical axons and retrogradely labeled cells. The results suggest that cortical axons contact cochlear nucleus-projecting cells in both IC and SOC. In both regions, contacts were more numerous on the side ipsilateral to the injected cortex. In the IC, the contacted cells projected ipsilaterally or contralaterally to the CN. In the SOC, the contacted cells projected ipsilaterally, contralaterally or bilaterally to the CN. We conclude that auditory cortex is in a position to modulate descending pathways from both the IC and SOC to the cochlear nucleus.

Animals↗

Age-related temporal processing speed deterioration in auditory cortex.

A common problem among the elderly is a difficulty in discriminating speech sounds. One factor that may contribute to this is a deterioration in the ability to process dynamic aspects of speech such as formant transitions. For the aging auditory system, this deterioration in temporal processing speed may be manifest as a deficit in encoding time-varying sounds that contain rapidly changing frequencies such as formant transitions. The primary goal of this study was to explore the neural basis of the effects of aging on temporal processing speed. To this end, single units were recorded from the auditory cortex of young and aged rats in response to frequency-modulated (FM) sweeps that changed from trial to trial in both direction and speed. Results showed that the majority of cells recorded from young rats responded most vigorously to fast and medium speeds. By contrast, the majority of units recorded from aged animals responded best to slow speeds. For preferred direction of FM sweep, similar results were observed for both age groups, namely, approximately half of the units exhibited a direction-selective response. The results of the present study demonstrate an age-related decrease in the rate of change of frequency that can be processed by the auditory cortex.

Aging↗

Gap detection threshold in the rat before and after auditory cortex ablation.

Gap detection threshold (GDT) was measured in adult female pigmented rats (strain Long-Evans) by an operant conditioning technique with food reinforcement, before and after bilateral ablation of the auditory cortex. GDT was dependent on the frequency spectrum and intensity of the continuously present noise in which the gaps were embedded. The mean values of GDT for gaps embedded in white noise or low-frequency noise (upper cutoff frequency 3 kHz) at 70 dB sound pressure level (SPL) were 1.57+/-0.07 ms and 2.9+/-0.34 ms, respectively. Decreasing noise intensity from 80 dB SPL to 20 dB SPL produced a significant increase in GDT. The increase in GDT was relatively small in the range of 80-50 dB SPL for white noise and in the range of 80-60 dB for low-frequency noise. The minimal intensity level of the noise that enabled GDT measurement was 20 dB SPL for white noise and 30 dB SPL for low-frequency noise. Mean GDT values at these intensities were 10.6+/-3.9 ms and 31.3+/-4.2 ms, respectively. Bilateral ablation of the primary auditory cortex (complete destruction of the Te1 and partial destruction of the Te2 and Te3 areas) resulted in an increase in GDT values. The fifth day after surgery, the rats were able to detect gaps in the noise. The values of GDT observed at this time were 4.2+/-1.1 ms for white noise and 7.4+/-3.1 ms for low-frequency noise at 70 dB SPL. During the first month after cortical ablation, recovery of GDT was observed. However, 1 month after cortical ablation GDT still remained slightly higher than in controls (1.8+/-0.18 for white noise, 3.22+/-0.15 for low-frequency noise, P<0.05). A decrease in GDT values during the subsequent months was not observed.

Acoustic Stimulation↗

Musicians with absolute pitch show distinct neural activities in the auditory cortex.

Meg responses from musicians who had absolute pitch and from non-musicians were measured while they received different auditory stimuli. The parameters of single equivalent current dipoles (ECDs) were calculated for the N1m responses occurring in the auditory cortex. The location of the ECD for the noise burst was significantly posterior to the ECDs for the tones in the two hemispheres of the musicians, but not for those of the non-musicians. Further, in the left hemisphere the ECDs for the musicians were significantly posterior to those for the non-musicians. These results suggest distinct neural activities in the auditory cortex of musicians, which may be the result of cortical plasticity produced by training and/or an inherent cortical structural specificity.

Acoustic Stimulation↗

Regulation of glutamate synapses by nicotinic acetylcholine receptors in auditory cortex.

Acetylcholine plays an important role in regulating the processing of sensory stimuli, and understanding its specific cellular actions is critical to understanding how sensory cortex develops and functions in different behavioral states. Here we review recent work on the cellular effects of nicotinic receptor activation in auditory cortex and describe how these actions could affect systems-level auditory function. In particular, we describe a novel function of nicotinic acetylcholine receptors to regulate glutamate synapses containing N-methyl-D-aspartate receptors during early postnatal development. The transient regulation of developing glutamate synapses also defines a window of vulnerability during which exposure to exogenous nicotine disrupts synapse development. Thus, it appears that nicotinic regulation of glutamate synapses is a critical feature of auditory cortex development.

Animals↗

Suppressed responses to self-triggered sounds in the human auditory cortex.

Humans are assumed to predict the sensory consequences of their own actions by means of forward models that enable discrimination between self-produced and external sensory signals. Here we tested whether responses in the human auditory cortex would differ to self-triggered versus externally triggered tones. The responses were recorded with a whole-scalp neuromagnetometer from 12 healthy subjects who either themselves triggered a tone by pressing a button once every 5 s or passively listened to externally triggered tones, presented in an identical sound sequence. Sources of the auditory N100m responses, peaking approximately 90 ms after sound onset in the supratemporal auditory cortex, were significantly weaker to self-triggered than to externally triggered sounds (suppressions 24 +/- 7% and 18 +/- 4% in the left and right hemispheres, respectively). These results support the existence of a forward model that predicts the auditory consequences of the subject's own motor acts on the environment--even with a tool--and thereby enables discrimination between self-produced and external sounds.

Acoustic Stimulation↗

Human auditory cortex responses to rising versus falling glides.

A 24-channel SQUID magnetometer was used to record signals from the right auditory cortex to tone glides with 16 different centre frequencies (ranging from 0.5 to 2 kHz), sweeping over one octave. The stimulus sequence, presented with an interstimulus onset interval of 0.6 s, consisted of infrequent 'deviant' rising (falling) glides and frequent 'standard' falling (rising) glides; the rising glides were identical to the falling glides but presented in the reverse direction. Deviant glides elicited significantly larger responses than standards at a latency of about 100 ms. This amplitude difference presumably arises from a mismatch response to the deviants. These results suggest that the auditory cortex extracts the direction of frequency transition even when the stimuli do not contain same frequencies.

Acoustic Stimulation↗

Perception and recognition discriminated in the mouse auditory cortex by c-Fos labeling.

The functions of the fields of the mammalian auditory cortex in sound perception and recognition are unknown. We used Fos (a protein of the inducible immediate-early gene c-fos) as a cellular marker of activated brain areas to show in the mouse (Mus domesticus) that sound is processed differentially in auditory cortical fields according to its actual significance in a behavioral context. Recognition, compared with perception of exactly the same sound, produced significantly less but well focused Fos-positive cells in a primary auditory cortical field and significantly more labeling in higher auditory and association fields. Thus, recognition means a state of distinctive spatial distribution of activity in auditory cortical fields with a predominance of activation in higher-order fields.

Acoustic Stimulation↗

Shared and private variability in the auditory cortex.

The high variability of cortical sensory responses is often assumed to impose a major constraint on efficient computation. In the auditory cortex, however, response variability can be very low. We have used in vivo whole cell patch-clamp methods to study the trial-to-trial variability of the subthreshold fluctuations in membrane potential underlying tone-evoked responses in the auditory cortex of anesthetized rats. Using methods adapted from classical quantal analysis, we partitioned this subthreshold variability into a private component (which includes synaptic, thermal, and other sources local to the recorded cell) and a shared component arising from network interactions. Here we report that this private component is remarkably small, usually about 1-3 mV, as quantified by the variance divided by the mean of the ensemble of tone-evoked response heights. The shared component can be much larger, and shows more heterogeneity across the population, ranging from about 0 to 10 mV. The remarkable fact that, at least 5 synapses from the auditory periphery, this variability remains so small raises the possibility that the intervening neural circuitry is organized so as to prevent private noise from accumulating as neural signals propagate to the cortex.

Acoustic Stimulation↗

Evidence for primary auditory cortex involvement in the echo suppression precedence effect: a 3CLT study.

An echo lagging shortly after a source and arising from another direction perceptually blends with the source, and the location of the fused 'source-echo' is dominated by the source location (the Precedence Effect). The neural substrates underlying the echo localization suppression are ambiguous. We recently suggested an auditory evoked potentials correlate of binaural echo lateralization suppression. A significant and specific reduction in binaural peak amplitude and area of the echo-evoked middle-latency component Pa was observed. The binaural echo-Pa suppression depended on echo lag and correlated with the psychophysical echo lateralization suppression. In this study, the echo-Pa generators were analyzed with 3CLT spatio-temporal analysis, in order to suggest the neural substrates involved in echo lateralization suppression. 3CLT enables reliable identification of components, based on rigid geometrical properties. The results suggest that the Pa1 subcomponent of Pa, associated with primary auditory cortex activity, fully accounts for the echo-Pa suppression. This physiological indication for primary auditory cortex involvement in the precedence effect is the first in humans.

Acoustic Stimulation↗

Activation of muscarinic receptors modulates NMDA receptor-mediated responses in auditory cortex.

The present study examines the ability of muscarinic receptor activation to modulate glutamatergic responses in the in vitro rat auditory cortex. Whole-cell patch-clamp recordings were obtained from layer II-III pyramidal neurons and responses elicited by either stimulation of deep gray matter or iontophoretic application of glutamate receptor agonists. Iontophoresis of the muscarinic agonist acetyl-beta-methylcholine (MCh) produced an atropine-sensitive reduction in the amplitude of glutamate-induced membrane depolarizations that was followed by a long-lasting (at least 20 min) response enhancement. Glutamate depolarizations were enhanced by MCh when elicited in the presence of alpha-amino-3-hydroxy-5-methyl-4-isoxazole propionic acid (AMPA)/kainate receptor antagonists 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) or 2,3-dihydroxy-6-nitro-7-sulfamoyl, benzo(F)quinoxaline (NBQX) but not the NMDA antagonists D-2-amino-5-phosphonovaleric acid (APV) or MK-801 hydrogen maleate. The magnitude of enhancement was voltage-dependent with the percentage increase greater at more depolarized membrane potentials. An involvement of NMDA receptors in these MCh-mediated effects was tested by using AMPA/kainate receptor antagonists to isolate the NMDA-mediated slow excitatory postsynaptic potential (EPSP) from other synaptic potentials. The slow EPSP and iontophoretic responses to NMDA were similarly modified by MCh, i.e., both being reduced during and enhanced (15-55 min) following MCh application. Cholinergic modulation of NMDA responses involves the engagement of G proteins, as enhancement was prevented by intracellular infusion with the nonhydrolyzable GDP analog guanosine-5'-O-(2-thiodiphosphate) trilithium salt (GDPbetaS). GDPbetaS was without effect on the early MCh-induced response suppression. Our results suggest that acetylcholine, acting at muscarinic receptors, produces a long-lasting enhancement of NMDA-mediated neurotransmission in auditory cortex, and that this modulatory effect is dependent upon a G protein-mediated event.

Acetylcholine↗

Induction of long-term receptive field plasticity in the auditory cortex of the waking guinea pig by stimulation of the nucleus basalis.

Learning induces neuronal receptive field (RF) plasticity in primary auditory cortex. This plasticity constitutes physiological memory as it is associative, highly specific, discriminative, develops rapidly, and is retained indefinitely. This study examined whether pairing a tone with activation of the nucleus basalis could induce RF plasticity in the waking guinea pig and, if so, whether it could be retained for 24 hr. Subjects received 40 trials of a single frequency paired with electrical stimulation of the nucleus basalis (NB) at tone offset. The physiological effectiveness of NB stimulation was assessed later while subjects were anesthetized with urethane by noting whether stimulation produced cortical desynchronization. Subjects in which NB stimulation was effective did develop RF plasticity and this was retained for 24 hr. Thus, activation of the NB during normal learning may be sufficient to induce enduring physiological memory in auditory cortex.

Acoustic Stimulation↗

Effect of stimulation on burst firing in cat primary auditory cortex.

1. Neural activity was recorded extracellularly with two independent microelectrodes aligned in parallel and advanced perpendicular to isofrequency sheets in cat primary auditory cortex. Multiunit activity was separated into single-unit spike trains using a maximum variance spike sorting algorithm. Only units that demonstrated a high quality of sorting and a minimum spontaneous firing rate of 0.2 spikes/s were considered for analysis. The primary aim of this study was to describe the effect of periodic click train and broadband noise stimulation on short-time-scale (< or = 50 ms) bursts in the spike trains of single auditory cortical units and to determine whether stimulation influenced the occurrence, spike count, and/or temporal structure of burst firing relative to a spontaneous baseline. 2. Extracellular recordings were made in 20 juvenile and adult cats from 69 single auditory cortical units during click train stimulation and silence, and from 30 single units during noise stimulation and in silence. In an additional 15 single units the effect of both click train and noise stimulation was investigated. The incidence, spike count, and temporal structure of short-time-scale burst firing in the first 100 ms following stimulus presentation was compared with burst firing in the period starting 500 ms after stimulus presentation and with spontaneous burst firing. In addition, the serial dependence of interspike intervals within a burst was tested during periods of stimulation. 3. Burst firing was present in the stimulation, poststimulation, and spontaneous conditions. Longer bursts (consisting of > or = 3 spikes) were more commonly observed in the poststimulation and spontaneous conditions than in the stimulation condition. This effect was most pronounced during click stimulation. A period of elevated firing activity was present in a subset of units 0.5-1.5 s after stimulus presentation, indicating prolonged effects of stimulation on single-unit firing behavior. 4. For both stimuli, the proportion of single-unit responses composed of bursts was significantly greater in poststimulation and spontaneous periods than during stimulation. Burst rate was higher in post-click-train stimulation and spontaneous periods than during periods of click stimulation. The isolated spike rate was significantly higher during periods of noise and click stimulation than in the poststimulation and spontaneous periods. 5. An examination of the autocorrelograms and higher-order interspike interval histograms of single-unit responses during click train stimulation indicated that 25% of single-unit spike trains contained an excess of brief first-order intervals and 14% of spike trains contained a shortage of long higher-order interspike intervals relative to a spontaneous baseline. During noise stimulation, 10% of single-unit responses contained an excess of short intervals relative to baseline. Interspike intervals of short-duration bursts were not serially dependent during periods of stimulation. 6. A comparison of the autocorrelograms and higher-order interval histograms of single-unit responses in the poststimulation and spontaneous conditions indicated that 20% of single-unit spike trains contained an excess of short first-, second-, and third-order intervals following stimulation. This subgroups of single units could not be distinguished on the basis of the age of the animal or the depth at which the recording was made. 7. The low incidence of burst firing during stimulation opposes the view that bursts serve as a mechanism to emphasize or amplify particular stimulus-related responses in the presence of ongoing spontaneous activity in the primary auditory cortex. Moreover, there is little evidence to support the notion that brief bursts represent neural codes, because intraburst intervals are not serially dependent. It is suggested that pyramidal burst firing may be an effective way to evoke postsynaptic firing in inhibitory interneurons and subsequ

Acoustic Stimulation↗