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[Advances in the study of primary auditory cortex. Demostration of its activation by functional magnetic resonance imaging].

PURPOSE: The purpose of this preliminary study has been to demonstrate and investigate the activation patterns of the primary auditory cortex (Heschl's gyrus = HG) using functional magnetic resonance imaging (fMRI). MATERIAL AND METHODS: A 2500 Hz tone stimulus was delivered monoaurally to the right and left ear of 15 normal-hearing right-handed volunteers in 20-second on-off cycles. FMRI data were obtained using a 1.5-Tesla scanner and processed with SPM2. RESULTS: Activated pixels were identified in the transverse temporal gyrus (Heschl's gyrus) of both hemispheres in response to pure tone stimuli using cross-correlation analysis (P < 0.001). Bilateral hemispheric activation was observed in all subjects and there was a trend towards contralateral HG activation to the stimulated ear. CONCLUSION: These results demonstrate directly that fMRI is a new and useful imaging technique to study the complex auditory cortex and it will have potential clinical applications in the next future.

Adult↗
PubMed1994

Topographic representation of tone intensity along the isofrequency axis of cat primary auditory cortex.

The sound pressure level (SPL), henceforth termed intensity, of acoustic signals is encoded in the central auditory system by neurons with different forms of intensity sensitivity. However, knowledge about the topographic organization of neurons with these different properties and hence about the spatial representation of intensity, especially at higher levels of the auditory pathway, is limited. Here we show that in the tonotopically organized primary auditory cortex (AI) of the cat there are orderly topographic organizations, along the isofrequency axis, of several neuronal properties related to the coding of the intensity of tones, viz. minimum threshold, dynamic range, best SPL, and non-monotonicity of spike count--intensity functions to tones of characteristic frequency (CF). Minimum threshold, dynamic range, and best SPL are correlated and alter periodically along isofrequency strips. The steepness of the high-intensity descending slope of spike count--intensity functions also varies systematically, with steepest slopes occurring in the regions along an isofrequency strip where low thresholds, narrow dynamic ranges and low best SPLs are found. As a consequence, CF-tones of various intensities are represented by orderly and, for most intensities, periodic, spatial patterns of distributed neuronal activity along an isofrequency strip. For low--to--moderate intensities, the mean relative activity along the entire isofrequency strip increases rapidly with intensity, with the spatial pattern of activity remaining quite constant along the strip. At higher intensities, however, the mean relative activity along the strip remains fairly constant with changes in intensity, but the spatial patterns change markedly. As a consequence of these effects, low- and high-intensity tones are represented by complementary distributions of activity alternating along an isofrequency strip. We conclude that in AI tone intensity is represented by two complementary modes, viz. discharge rate and place. Furthermore, the magnitude of the overall changes in the representation of tone intensity in AI appears to be closely related to psychophysical measures of loudness and of intensity discrimination.

Acoustic Stimulation↗
PubMed2005

Neuromagnetic correlates of streaming in human auditory cortex.

The brain is constantly faced with the challenge of organizing acoustic input from multiple sound sources into meaningful auditory objects or perceptual streams. The present study examines the neural bases of auditory stream formation using neuromagnetic and behavioral measures. The stimuli were sequences of alternating pure tones, which can be perceived as either one or two streams. In the first experiment, physical stimulus parameters were varied between values that promoted the perceptual grouping of the tone sequence into one coherent stream and values that promoted its segregation into two streams. In the second experiment, an ambiguous tone sequence produced a bistable percept that switched spontaneously between one- and two-stream percepts. The first experiment demonstrated a strong correlation between listeners' perception and long-latency (>60 ms) activity that likely arises in nonprimary auditory cortex. The second demonstrated a covariation between this activity and listeners' perception in the absence of physical stimulus changes. Overall, the results indicate a tight coupling between auditory cortical activity and streaming perception, suggesting that an explicit representation of auditory streams may be maintained within nonprimary auditory areas.

Acoustic Stimulation↗
PubMed1990

Steady-state responses from the cat auditory cortex.

Responses to 350 ms trains of clicks with 10-100 Hz repetition rate were recorded from the auditory cortices of six cats. Click trains of 30-50 and 90-100 Hz elicited a clear steady-state response (SSR) in awake state. SSRs were small or absent below 30 Hz and in 60-70 Hz stimulus range. In slow wave sleep the optimal rate to elicit SSR shifted towards lower frequencies. 90 Hz SSR was largest in paradoxical sleep. SSRs were strongly suppressed by barbiturate anesthesia. The amplitude of the SSR from the medial geniculate body (MGB) in two cats gradually decreased from 20 to 100 Hz and was more resilient to barbiturate anesthesia than the cortical SSRs. Only low amplitude or no SSRs could be recorded from vertex, visual and association cortices and from the hippocampus in control recordings. The results suggest different generation mechanisms for SSRs recorded from cat auditory cortex and MGB. Human auditory SSRs resemble cat auditory cortical SSRs more than those recorded from cat MGB. The results imply that auditory SSRs in humans are generated in the cortex.

Acoustic Stimulation↗
PubMed1993

Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. I. Evidence for monaural directional cues.

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Neurons, sensitive to sound direction in the horizontal plane (azimuth), were identified by their responses to noise bursts, presented in the free field, that varied in azimuth and sound pressure level (SPL). SPLs typically varied between 0 and 80 dB and were presented at each azimuth that was tested. Each azimuth-sensitive neuron responded well to some SPLs at certain azimuths and did not respond well to any SPL at other azimuths. This report describes AI neurons that were sensitive to the azimuth of monaurally presented noise bursts. 2. Unilateral ear plugging was used to test each azimuth-sensitive neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the concha and ear canal, attenuated sound reaching the tympanic membrane by 25-70 dB. Binaural interactions were inferred by comparing responses obtained under binaural (no plug) and monaural (ear plug) conditions. 3. Of the total sample of 131 azimuth-sensitive cells whose responses to ear plugging were studied, 27 were sensitive to the azimuth of monaurally presented noise bursts. We refer to these as monaural directional (MD) cells, and this report describes their properties. The remainder of the sample consisted of cells that either required binaural stimulation for azimuth sensitivity (63/131), because they were insensitive to azimuth under unilateral ear plug conditions or responded too unreliably to permit detailed conclusions regarding the effect of ear plugging (41/131). 4. Most (25/27) MD cells received either monaural input (MD-E0) or binaural excitatory/inhibitory input (MD-EI), as inferred from ear plugging. Two MD cells showed other characteristics. The contralateral ear was excitatory for 25/27 MD cells. 5. MD-E0 cells (22%, 6/27) were monaural. They were unaffected by unilateral ear plugging, showing that they received excitatory input from one ear, and that stimulation of the other ear was without apparent effect. On the other hand, some monaural cells in AI were insensitive to the azimuth of noise bursts, showing that sensitivity to monaural directional cues is not a property of all monaural cells in AI. 6. MD-EI cells (70%, 19/27) exhibited an increase in responsiveness on the side of the plugged ear, showing that they received excitatory drive from one ear and inhibitory drive from the other. MD-EI cells remained azimuth sensitive with the inhibitory ear plugged, showing that they were sensitive to monaural directional cues at the excitatory ear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed2006

Non-Gaussian membrane potential dynamics imply sparse, synchronous activity in auditory cortex.

Many models of cortical dynamics have focused on the high-firing regime, in which neurons are driven near their maximal rate. Here we consider the responses of neurons in auditory cortex under typical low-firing rate conditions, when stimuli have not been optimized to drive neurons maximally. We used whole-cell patch-clamp recording in vivo to measure subthreshold membrane potential fluctuations in rat primary auditory cortex in both the anesthetized and awake preparations. By analyzing the subthreshold membrane potential dynamics on single trials, we made inferences about the underlying population activity. We found that, during both spontaneous and evoked responses, membrane potential was highly non-Gaussian, with dynamics consisting of occasional large excursions (sometimes tens of millivolts), much larger than the small fluctuations predicted by most random walk models that predict a Gaussian distribution of membrane potential. Thus, presynaptic inputs under these conditions are organized into quiescent periods punctuated by brief highly synchronous volleys, or "bumps." These bumps were typically so brief that they could not be well characterized as "up states" or "down states." We estimate that hundreds, perhaps thousands, of presynaptic neurons participate in the largest volleys. These dynamics suggest a computational scheme in which spike timing is controlled by concerted firing among input neurons rather than by small fluctuations in a sea of background activity.

Acoustic Stimulation↗
PubMed1998

Tonotopic organization of the human auditory cortex as detected by BOLD-FMRI.

Functional magnetic resonance imaging is a noninvasive and nonradioactive method for the detection of focal brain activity. In the present study the auditory cortex was investigated in nine normal subjects who were binaurally stimulated using pulsed sine tones of 500 Hz and 4000 Hz. The BOLD (blood oxygenation level dependent) signal change coincided with the stimulation paradigm and was detected in the plane of the superior temporal gyrus. The comparison of the spatial distribution of activated areas revealed a different behavior for the two frequencies. The present findings underline the existence of a frequency specific organization in the medio-lateral, fronto-occipital and cranio-caudal extension in both hemispheres of the auditory cortex in human. The activated areas for the high tone were found more frontally and medially orientated than the low tone stimulated areas. Furthermore, a slight cranio-caudal shift was observed for the higher frequency, more pronounced in the right than in the left temporal lobe. Finally, for most of the subjects investigated the BOLD activation area of the 500 Hz sine tone was larger than that of the 4000 Hz stimulation. Both frequencies showed a lateralization of signal response to the left temporal lobe.

Acoustic Stimulation↗
PubMed1975

Detection of natural complex sounds by cells in the primary auditory cortex of the cat.

The neural mechanisms involved in the detection of natural complex sounds were studied by recording single-neuron responses from 132 cells in the primary auditory cortex of the cat. The cats were paralyzed and under neuroleptanalgesia (NLA). The cells were first stimulated with pure tones; the responses were then compared with those evoked by many different types of complex sounds, most of which were animal vocalizations. Per-stimulus-time (PST) histograms constructed from the responses to repetitive stimuli were compared with the corresponding sound spectrograms formed from the sounds used as stimuli. Of 100 cells 68 per cent gave predictable responses to complex sounds on the basis of their responses to different pure tone frequencies. In 32 per cent of the cells the responses were unpredictable. Half of these cells did not react to pure tones at all but responded to one or more animal vocalizations or generator sounds with different patterns. Some cells reacted to pure tones in quite a different way than to certain complex sounds, e.g. with inhibition instead of excitation. These results indicate that cells in the primary auditory cortex of the cat reacting in an unpredictable way to sounds with a complex structure have a more or less specialized function, in detecting and analyzing natural and other complex sound patterns. Cells reacting phasically to pure tones seem to be involved in the detection of transient sound elements.

Acoustic Stimulation↗
PubMed2002

Long-term consolidation and retention of learning-induced tuning plasticity in the auditory cortex of the guinea pig.

The major goal of this study was to determine whether classical conditioning produces long-term neural consolidation of frequency tuning plasticity in the auditory cortex. Local field potentials (LFPs) were obtained from chronically implanted adult male Hartley guinea pigs that were divided into conditioning (n = 4) and sensitization control (n = 3) groups. Tuning functions were determined in awake subjects for average LFPs (approximately 0.4 to 36.0 kHz, -20 to 80 dB) immediately before training as well as 1 h and 1, 3, 7, and 10 days after training; sensitization subjects did not have a 10-day retention test. Conditioning consisted of a single session of 30 to 45 trials of a 6-s tone (CS, 70 dB) that was not the best frequency (BF, peak of a tuning curve), followed by a brief leg shock (US) at CS offset. Sensitization control animals received the same density of CS and US presentations unpaired. Heart rate recordings showed that the conditioning group developed conditioned bradycardia, whereas the sensitization control group did not. Local field potentials in the conditioning group, but not in the sensitization group, developed tuning plasticity. The ratio of responses to the CS frequency versus the BF were increased 1 h after training, and this increase was retained for the 10-day period of the study. Both tuning plasticity and retention were observed across stimulus levels (10-80 dB). Most noteworthy, tuning plasticity exhibited consolidation (i.e., developed greater CS-specific effects across retention periods), attaining asymptote at 3 days. The findings indicate that LFPs in the auditory cortex have three cardinal features of behavioral memory: associative tuning plasticity, long-term retention, and long-term consolidation. Potential cellular and subcellular mechanisms of LFP tuning plasticity and long-term consolidation are discussed.

Animals↗
PubMed2000

Induction of visual orientation modules in auditory cortex.

Modules of neurons sharing a common property are a basic organizational feature of mammalian sensory cortex. Primary visual cortex (V1) is characterized by orientation modules--groups of cells that share a preferred stimulus orientation--which are organized into a highly ordered orientation map. Here we show that in ferrets in which retinal projections are routed into the auditory pathway, visually responsive neurons in 'rewired' primary auditory cortex are also organized into orientation modules. The orientation tuning of neurons within these modules is comparable to the tuning of cells in V1 but the orientation map is less orderly. Horizontal connections in rewired cortex are more patchy and periodic than connections in normal auditory cortex, but less so than connections in V1. These data show that afferent activity has a profound influence on diverse components of cortical circuitry, including thalamocortical and local intracortical connections, which are involved in the generation of orientation tuning, and long-range horizontal connections, which are important in creating an orientation map.

Animals↗
PubMed1999

Organization of inhibitory frequency receptive fields in cat primary auditory cortex.

Based on properties of excitatory frequency (spectral) receptive fields (esRFs), previous studies have indicated that cat primary auditory cortex (A1) is composed of functionally distinct dorsal and ventral subdivisions. Dorsal A1 (A1d) has been suggested to be involved in analyzing complex spectral patterns, whereas ventral A1 (A1v) appears better suited for analyzing narrowband sounds. However, these studies were based on single-tone stimuli and did not consider how neuronal responses to tones are modulated when the tones are part of a more complex acoustic environment. In the visual and peripheral auditory systems, stimulus components outside of the esRF can exert strong modulatory effects on responses. We investigated the organization of inhibitory frequency regions outside of the pure-tone esRF in single neurons in cat A1. We found a high incidence of inhibitory response areas (in 95% of sampled neurons) and a wide variety in the structure of inhibitory bands ranging from a single band to more than four distinct inhibitory regions. Unlike the auditory nerve where most fibers possess two surrounding "lateral" suppression bands, only 38% of A1 cells had this simple structure. The word lateral is defined in this sense to be inhibition or suppression that extends beyond the low- and high-frequency borders of the esRF. Regional differences in the distribution of inhibitory RF structure across A1 were evident. In A1d, only 16% of the cells had simple two-banded lateral RF organization, whereas 50% of A1v cells had this organization. This nonhomogeneous topographic distribution of inhibitory properties is consistent with the hypothesis that A1 is composed of at least two functionally distinct subdivisions that may be part of different auditory cortical processing streams.

Acoustic Stimulation↗
PubMed2000

Complex tone processing in primary auditory cortex of the awake monkey. II. Pitch versus critical band representation.

Noninvasive neurophysiological studies in humans support the existence of an orthogonal spatial representation of pure tone frequency and complex tone pitch in auditory cortex [Langner et al., J. Comp. Physiol. A 181, 665-676 (1997)]. However, since this topographic organization is based on neuromagnetic responses evoked by wideband harmonic complexes (HCs) of variable fundamental frequency (f0), and thus interharmonic frequency separation (deltaF), critical band filtering effects due to differential resolvability of harmonics may have contributed to shaping these responses. To test this hypothesis, the present study examined responses evoked by three-component HCs of variable f0 in primary auditory cortex (A1) of the awake monkey. The center frequency of the HCs was fixed at the best frequency (BF) of the cortical site. Auditory evoked potential (AEP), multiunit activity, and current source density techniques were used to evaluate A1 responses as a function of f0 (=deltaF). Generally, amplitudes of nearly all response components increased with f0, such that maximal responses were evoked by HCs comprised of low-order resolved harmonics. Statistically significant increases in response amplitude typically occurred at deltaFs between 10% and 20% of center frequency, suggestive of critical bandlike behavior. Complex tone response amplitudes also reflected nonlinear summation in that they could not be predicted by the pure tone frequency sensitivity curves of the cortical sites. A mechanism accounting for the observed results is proposed which involves mutual lateral inhibitory interactions between responses evoked by stimulus components lying within the same critical band. As intracortical AEP components likely to be propagated to the scalp were also strongly modulated by deltaF, these findings indicate that noninvasive recordings of responses to complex sounds may require a consideration of critical band effects in their interpretation.

Acoustic Stimulation↗
PubMed1979

Lability in the responses of cells in the auditory cortex of squirrel monkeys to species-specific vocalizations.

The activity of 28 cells located mainly in the secondary auditory cortex (A II) of awake squirrel-monkeys, was extracellularly recorded for periods of up to 6 h. Seven different species-specific vocalizations, which were repeatedly presented to the monkey, were used as auditory stimuli. Twenty-six cells responded, at least once, to one or more vocalizations; 22 cells revealed some change in their response (pattern or strength) to at least one vocalization ("change in response"). Twenty-one cells exhibited a change in the number and/or type of vocalization to which they responded during the recording period ("change in selectivity"). At some time during the recording period all the responding cells exhibited a "change in response" and/or a "change in selectivity" ("change in responsiveness"). A "change in response" of a cell to a vocalization did not necessarily exclude a "change in selectivity", associated with the same vocalization, later in time and vice-versa. A "change in responsiveness" to one vocalization was not necessarily correlated with "changes in responsiveness" to other vocalizations.

Acoustic Stimulation↗
PubMed2006

Spectral resolution of monkey primary auditory cortex (A1) revealed with two-noise masking.

An important function of the auditory nervous system is to analyze the frequency content of environmental sounds. The neural structures involved in determining psychophysical frequency resolution remain unclear. Using a two-noise masking paradigm, the present study investigates the spectral resolution of neural populations in primary auditory cortex (A1) of awake macaques and the degree to which it matches psychophysical frequency resolution. Neural ensemble responses (auditory evoked potentials, multiunit activity, and current source density) evoked by a pulsed 60-dB SPL pure-tone signal fixed at the best frequency (BF) of the recorded neural populations were examined as a function of the frequency separation (DeltaF) between the tone and two symmetrically flanking continuous 80-dB SPL, 50-Hz-wide bands of noise. DeltaFs ranged from 0 to 50% of the BF, encompassing the range typically examined in psychoacoustic experiments. Responses to the signal were minimal for DeltaF = 0% and progressively increased with DeltaF, reaching a maximum at DeltaF = 50%. Rounded exponential functions, used to model auditory filter shapes in psychoacoustic studies of frequency resolution, provided excellent fits to neural masking functions. Goodness-of-fit was greatest for response components in lamina 4 and lower lamina 3 and least for components recorded in more superficial cortical laminae. Physiological equivalent rectangular bandwidths (ERBs) increased with BF, measuring nearly 15% of the BF. These findings parallel results of psychoacoustic studies in both monkeys and humans, and thus indicate that a representation of perceptual frequency resolution is available at the level of A1.

Acoustic Stimulation↗
PubMed1990

Visual projections induced into the auditory pathway of ferrets. I. Novel inputs to primary auditory cortex (AI) from the LP/pulvinar complex and the topography of the MGN-AI projection.

The organization of cortical circuitry responsible for processing sensory information is a subject of intense examination. However, it is not known whether cortical cells in different sensory cortices process information in a way that is specific to the modality of their input, or whether there are commonalities in processing circuitry across different cortices. In our laboratory, this question has been investigated at the level of the geniculocortical pathway by routing information of one sensory modality into the processing circuitry of another modality. Appropriate early lesions cause growth of retinal axons into the auditory thalamus (MGN) (Sur et al., Science 242:1437, '88). Previously, we have established that the MGN carries the resulting visual information on to primary auditory cortex (AI), which thus contains visually responsive neurons and a topographic representation of the retina (Roe et al., Soc. Neurosci. Abstr. 14:460, '88; Sur et al., Science 242:1437, '88). In this paper, we describe anomalous projections from the dorsal part of the thalamus, specifically the lateral posterior/pulvinar complex, into AI. This result demonstrates that thalamic neurons belonging to one modality can be induced to project to cortex that is normally of a different modality. In addition, we have studied in detail the nature of the MGN to AI projection in these animals as compared to the normal projection. The MGN to AI projection appears to be unaltered by the lesions; the location and topography of labelled cells are similar to that in normal animals. Because the MGN to AI projection is still highly divergent along the "isofrequency" dimension when compared to the tonotopic dimension, our data suggest that visual topography in the cortical map is created within the auditory cortex, perhaps by activity-dependent sharpening of the retinal representation during development.

Animals↗
PubMed2001

Persistent and specific influences of early acoustic environments on primary auditory cortex.

This study demonstrates that the adult form of 'tonotopic maps' of sound frequency in the rat primary auditory cortex (A1) arises from parallel developmental processes involving two cortical zones: the progressive differentiation and refinement of selectively tone-responsive receptive fields within an initially broadly-tuned posterior zone, and the progressive loss of tone-evoked, short-latency response over an initially large, very broadly tuned anterior zone. The formation of tonotopic maps in A1 was specifically influenced by a rat pup's early acoustic environments. Exposure to pulsed tones resulted in accelerated emergence and an expansion of A1 representations of those specific tone frequencies, as well as a deteriorated tonotopicity and broader-than-normal receptive fields. Thus, auditory experiences during early postnatal development are important in shaping the functional development of auditory cortical representations of specific acoustic environments.

Acoustic Stimulation↗
PubMed1981

[Evoked auditory cortex potentials of wakeful dogs in response to paired clicks].

EP to a test click appears in the auditory cortex of alert intact dogs in 30--40 ms after the conditioning click (period of the so called absolute refractory state). The curve of the recovery of the second EP has a complex periodic character with two stages of complete recovery of the early EP complex amplitude: rapid initial after 80--180 ms, and final, 280--430 ms after presentation of the conditioning click.

Animals↗
PubMed2003

[Effect of forward masking on evoked potentials of auditory cortex in guinea pigs].

Thresholds of the event-related potentials (ERPs) appearance were measured for one stationary and four moving auditory images presented in silence or under forward masking conditions. The difference between thresholds in silence and after noise masker was considered as masking level. Under the forward masking, the amplitude of the ERP to the first click in the test series decreased in guinea pig auditory cortex. Masking level decreased with the time lag between signal and masker and didn't depend on the fused auditory image localization that corresponded to the first click in different test signals. This fact can support the hypothesis that for the long test signals the initial part can be masked more than the final one. The ERPs amplitude to next clicks in test series depended on interaction of two factors: forward masking in the "masker-signal" system and interaction of separate ERPs in the series evoked by the test signal.

Acoustic Stimulation↗