Fields In Auditory Cortex Research Articles

Absence of eye position effects in the early auditory cortex of monkeys.

This study aims to investigate whether the position of the eyes affects the neuronal activity in auditory cortex in a condition in which not the active control of eye position but the execution of hand movements was required relative to stimuli. Two monkeys were trained to perform audio-visual tasks in which they had to use their hand to respond to both the visual and the auditory stimuli to earn a reward. We recorded the spiking activity and the local field potentials from the core fields of auditory cortex, along with the eye position of the monkeys while they performed the tasks. We found that both the spiking activity and the local field potentials did not significantly vary with the eye position. This was the case both during the presentation of sounds and during other periods of the tasks. Our results indicate that eye position did not affect the neuronal activity in auditory cortex during the audio-visual tasks. Our results, together with the previous finding that eye position affects the neuronal activity in auditory cortex during eye fixation tasks, suggest that the presence of eye position effects in auditory cortex depends on the specific behavior a subject has to exhibit to obtain a reward.

Optimising a computational model of human auditory cortex with an evolutionary algorithm

We demonstrate how the structure of auditory cortex can be investigated by combining computational modelling with advanced optimisation methods. We optimise a well-established auditory cortex model by means of an evolutionary algorithm. The model describes auditory cortex in terms of multiple core, belt, and parabelt fields. The optimisation process finds the optimum connections between individual fields of auditory cortex so that the model is able to reproduce experimental magnetoencephalographic (MEG) data. In the current study, this data comprised the auditory event-related fields (ERFs) recorded from a human subject in an MEG experiment where the stimulus-onset interval between consecutive tones was varied. The quality of the match between synthesised and experimental waveforms was 98%. The results suggest that neural activity caused by feedback connections plays a particularly important role in shaping ERF morphology. Further, ERFs reflect activity of the entire auditory cortex, and response adaptation due to stimulus repetition emerges from a complete reorganisation of AC dynamics rather than a reduction of activity in discrete sources. Our findings constitute the first stage in establishing a new non-invasive method for uncovering the organisation of the human auditory cortex.

High resolution recording of local field currents simultaneously with sound-evoked calcium signals by a photometric patch electrode in the auditory cortex field L of the chick

BackgroundFunctioning of the brain is based on both electrical and metabolic activity of neural ensembles. Accordingly, it would be useful to measure intracellular metabolic signaling simultaneously with electrical activity in the brain in vivo. New methodWe innovated a PhotoMetric-patch-Electrode (PME) recording system that has a high temporal resolution incorporating a photomultiplier tube as a light detector. The PME is fabricated from a quartz glass capillary to transmit light as a light guide, and it can detect electrical signals as a patch electrode simultaneously with a fluorescence signal. ResultsWe measured the sound-evoked Local Field Current (LFC) and fluorescence Ca2+ signal from neurons labeled with Ca2+-sensitive dye Oregon Green BAPTA1 in field L, the avian auditory cortex. Sound stimulation evoked multi-unit spike bursts and Ca2+ signals, and enhanced the fluctuation of LFC. After a brief sound stimulation, the cross-correlation between LFC and Ca2+ signal was prolonged. D-AP5 (antagonist for NMDA receptors) suppressed the sound-evoked Ca2+ signal when applied locally by pressure from the tip of PME. Comparison with existing methodsIn contrast to existing multiphoton imaging or optical fiber recording methods, the PME is a patch electrode pulled simply from a quartz glass capillary and can measure fluorescence signals at the tip simultaneously with electrical signal at any depth of the brain structure. ConclusionThe PME is devised to record electrical and optical signals simultaneously with high temporal resolution. Moreover, it can inject chemical agents dissolved in the tip-filling medium locally by pressure, allowing manipulation of neural activity pharmacologically.

Predictive encoding of pure tones and FM-sweeps in the human auditory cortex.

Expectations substantially influence perception, but the neural mechanisms underlying this influence are not fully understood. A prominent view is that sensory neurons encode prediction error with respect to expectations on upcoming sensory input. Although the encoding of prediction error has been previously demonstrated in the human auditory cortex (AC), previous studies often induced expectations using stimulus repetition, potentially confounding prediction error with neural habituation. These studies also measured AC as a single population, failing to consider possible predictive specializations of different AC fields. Moreover, the few studies that considered prediction error to stimuli other than pure tones yielded conflicting results. Here, we used functional magnetic resonance imaging (fMRI) to systematically investigate prediction error to subjective expectations in auditory cortical fields Te1.0, Te1.1, Te1.2, and Te3, and two types of stimuli: pure tones and frequency modulated (FM) sweeps. Our results show that prediction error is elicited with respect to the participants' expectations independently of stimulus repetition and similarly expressed across auditory fields. Moreover, despite the radically different strategies underlying the decoding of pure tones and FM-sweeps, both stimulus modalities were encoded as prediction error in most fields of AC. Altogether, our results provide unequivocal evidence that predictive coding is the general encoding mechanism in AC.

Deviance detection in physiologically identified cell types in the rat auditory cortex

Auditory deviance detection is a function of the auditory system that allows reduction of the processing demand for repetitive stimuli while stressing unpredictable ones, which are potentially more informative. Deviance detection has been extensively studied in humans using the oddball paradigm, which evokes an event-related potential known as mismatch negativity (MMN). The same stimulation paradigms are used in animal studies that aim to elucidate the neuronal mechanisms underlying deviance detection. In order to understand the circuitry responsible for deviance detection in the auditory cortex (AC), it is necessary to determine the properties of excitatory and inhibitory neurons separately. Measuring the spike widths of neurons recorded extracellularly from the anaesthetized rat AC, we classified them as fast spiking or regular spiking units. These two neuron types are generally considered as putative inhibitory or excitatory, respectively. In response to an oddball paradigm, we found that both types of units showed similar amounts of deviance detection overall. When considering each AC field separately, we found that only in A1 fast spiking neurons showed higher deviance detection levels than regular spiking neurons, while in the rest of the fields there was no such distinction. Interpreting these responses in the context of the predictive coding framework, we found that the responses of both types of units reflect mainly prediction error signaling (i.e., genuine deviance detection) rather than repetition suppression.

Streaming of Repeated Noise in Primary and Secondary Fields of Auditory Cortex.

Statistical regularities in natural sounds facilitate the perceptual segregation of auditory sources, or streams. Repetition is one cue that drives stream segregation in humans, but the neural basis of this perceptual phenomenon remains unknown. We demonstrated a similar perceptual ability in animals by training ferrets of both sexes to detect a stream of repeating noise samples (foreground) embedded in a stream of random samples (background). During passive listening, we recorded neural activity in primary auditory cortex (A1) and secondary auditory cortex (posterior ectosylvian gyrus, PEG). We used two context-dependent encoding models to test for evidence of streaming of the repeating stimulus. The first was based on average evoked activity per noise sample and the second on the spectro-temporal receptive field. Both approaches tested whether differences in neural responses to repeating versus random stimuli were better modeled by scaling the response to both streams equally (global gain) or by separately scaling the response to the foreground versus background stream (stream-specific gain). Consistent with previous observations of adaptation, we found an overall reduction in global gain when the stimulus began to repeat. However, when we measured stream-specific changes in gain, responses to the foreground were enhanced relative to the background. This enhancement was stronger in PEG than A1. In A1, enhancement was strongest in units with low sparseness (i.e., broad sensory tuning) and with tuning selective for the repeated sample. Enhancement of responses to the foreground relative to the background provides evidence for stream segregation that emerges in A1 and is refined in PEG.SIGNIFICANCE STATEMENT To interact with the world successfully, the brain must parse behaviorally important information from a complex sensory environment. Complex mixtures of sounds often arrive at the ears simultaneously or in close succession, yet they are effortlessly segregated into distinct perceptual sources. This process breaks down in hearing-impaired individuals and speech recognition devices. By identifying the underlying neural mechanisms that facilitate perceptual segregation, we can develop strategies for ameliorating hearing loss and improving speech recognition technology in the presence of background noise. Here, we present evidence to support a hierarchical process, present in primary auditory cortex and refined in secondary auditory cortex, in which sound repetition facilitates segregation.

Stimulus dependent transformations between synaptic and spiking receptive fields in auditory cortex

Auditory cortex neurons nonlinearly integrate synaptic inputs from the thalamus and cortex, and generate spiking outputs for simple and complex sounds. Directly comparing synaptic and spiking activity can determine whether this input-output transformation is stimulus-dependent. We employ in vivo whole-cell recordings in the mouse primary auditory cortex, using pure tones and broadband dynamic moving ripple stimuli, to examine properties of functional integration in tonal (TRFs) and spectrotemporal (STRFs) receptive fields. Spectral tuning in STRFs derived from synaptic, subthreshold and spiking responses proves to be substantially more selective than for TRFs. We describe diverse spectral and temporal modulation preferences and distinct nonlinearities, and their modifications between the input and output stages of neural processing. These results characterize specific processing differences at the level of synaptic convergence, integration and spike generation resulting in stimulus-dependent transformation patterns in the primary auditory cortex.

ON-OFF receptive fields in auditory cortex diverge during development and contribute to directional sweep selectivity

Neurons in the auditory cortex exhibit distinct frequency tuning to the onset and offset of sounds, but the cause and significance of ON and OFF receptive field (RF) organisation are not understood. Here we demonstrate that distinct ON and OFF frequency tuning is largely absent in immature mouse auditory cortex and is thus a consequence of cortical development. Simulations using a novel implementation of a standard Hebbian plasticity model show that the natural alternation of sound onset and offset is sufficient for the formation of non-overlapping adjacent ON and OFF RFs in cortical neurons. Our model predicts that ON/OFF RF arrangement contributes towards direction selectivity to frequency-modulated tone sweeps, which we confirm by neuronal recordings. These data reveal that a simple and universally accepted learning rule can explain the organisation of ON and OFF RFs and direction selectivity in the developing auditory cortex.

Neural Representations of the Full Spatial Field in Auditory Cortex of Awake Marmoset (Callithrix jacchus).

Unlike visual signals, sound can reach the ears from any direction, and the ability to localize sounds from all directions is essential for survival in a natural environment. Previous studies have largely focused on the space in front of a subject that is also covered by vision and were often limited to measuring spatial tuning along the horizontal (azimuth) plane. As a result, we know relatively little about how the auditory cortex responds to sounds coming from spatial locations outside the frontal space where visual information is unavailable. By mapping single-neuron responses to the full spatial field in awake marmoset (Callithrix jacchus), an arboreal animal for which spatial processing is vital in its natural habitat, we show that spatial receptive fields in several auditory areas cover all spatial locations. Several complementary measures of spatial tuning showed that neurons were tuned to both frontal space and rear space (outside the coverage of vision), as well as the space above and below the horizontal plane. Together, these findings provide valuable new insights into the representation of all spatial locations by primate auditory cortex.

Auditory-visual integration in fields of the auditory cortex

While multimodal interactions have been known to exist in the early sensory cortices, the response properties and spatiotemporal organization of these interactions are poorly understood. To elucidate the characteristics of multimodal sensory interactions in the cerebral cortex, neuronal responses to visual stimuli with or without auditory stimuli were investigated in core and belt fields of guinea pig auditory cortex using real-time optical imaging with a voltage-sensitive dye. On average, visual responses consisted of short excitation followed by long inhibition. Although visual responses were observed in core and belt fields, there were regional and temporal differences in responses. The most salient visual responses were observed in the caudal belt fields, especially posterior (P) and dorsocaudal belt (DCB) fields. Visual responses emerged first in fields P and DCB and then spread rostroventrally to core and ventrocaudal belt (VCB) fields. Absolute values of positive and negative peak amplitudes of visual responses were both larger in fields P and DCB than in core and VCB fields. When combined visual and auditory stimuli were applied, fields P and DCB were more inhibited than core and VCB fields beginning approximately 110 ms after stimuli. Correspondingly, differences between responses to auditory stimuli alone and combined audiovisual stimuli became larger in fields P and DCB than in core and VCB fields after approximately 110 ms after stimuli. These data indicate that visual influences are most salient in fields P and DCB, which manifest mainly as inhibition, and that they enhance differences in auditory responses among fields.

Scopolamine attenuates auditory cortex response

Conclusion: Scopolamine, a tropane alkaloid drug that mainly acts as an antagonist of muscarinic acetylcholine receptors, was found to reduce the local field potentials (LFP) of auditory cortex (AC) evoked by tone and gap-offsets whose effects may compensate the cortical hyperexcitability related to tinnitus. Objective: To study the effects of scopolamine on the AC and the inferior colliculus (IC) of awake rats in order to understand scopolamine’s effect on tinnitus and gap detection. Method: Silent gaps (duration varied from 2–100 ms) embedded in otherwise continuous noise were used to elicit AC and IC response. Gap evoked AC and IC field potentials were recorded from awake rats before and after treatment of scopolamine (3 mg/kg, i.m.). Results: Acute injection of scopolamine (3 mg/kg, i.m.) induced a significant reduction of the AC response, but not the IC response, to the offset of the gaps embedded in white noise. The results suggest that scopolamine may reduce AC neural synchrony.

Neuronal activity in primate auditory cortex during the performance of audiovisual tasks.

This study aimed at a deeper understanding of which cognitive and motivational aspects of tasks affect auditory cortical activity. To this end we trained two macaque monkeys to perform two different tasks on the same audiovisual stimulus and to do this with two different sizes of water rewards. The monkeys had to touch a bar after a tone had been turned on together with an LED, and to hold the bar until either the tone (auditory task) or the LED (visual task) was turned off. In 399 multiunits recorded from core fields of auditory cortex we confirmed that during task engagement neurons responded to auditory and non-auditory stimuli that were task-relevant, such as light and water. We also confirmed that firing rates slowly increased or decreased for several seconds during various phases of the tasks. Responses to non-auditory stimuli and slow firing changes were observed during both the auditory and the visual task, with some differences between them. There was also a weak task-dependent modulation of the responses to auditory stimuli. In contrast to these cognitive aspects, motivational aspects of the tasks were not reflected in the firing, except during delivery of the water reward. In conclusion, the present study supports our previous proposal that there are two response types in the auditory cortex that represent the timing and the type of auditory and non-auditory elements of a auditory tasks as well the association between elements.

BRAIN STEM AUDITORY AGNOSIA: AUDITORY CORTEX ACTIVATED TDCS

fMRI was used to study frationate auditory field activations in a young woman with auditory agnosia due to traumatic avulsion of the left inferior colliculus. She was unable to recognize...

Functional lateralization in auditory cortex under informational masking and in silence

The N(1) m is an evoked magnetic field in auditory cortex that is automatically elicited by tones in silence but not in the context of multiple other tones: when listeners are unaware of a tone stream because of informational masking, no N(1) m-like activity is observed. In contrast, N(1) m-like activity is evoked when listeners are aware of the regular tone stream in the same context but in another trial. Here we compared this awareness-related negativity (ARN) with the automatic N(1) m. First, we evaluated whether stimulus lateralization by ear or interaural time differences modulates hemispheric lateralization of the response, as a putative marker of sensory processing. Second, we evaluated the stimulus-independent hemispheric balance thought to indicate higher level cortical processing. The results dissociate three, partly overlapping, time intervals: the P(1) m (45-85 ms) was evoked by missed and detected target tones alike. Subsequent negative activity was only observed when listeners indicated awareness of the target stream inside the multi-tone masker. In the N(1) m time interval (75-175 ms), hemispheric balance of the ARN and N(1) m was modulated by stimulus lateralization. In the subsequent time interval (175-275 ms), auditory-cortex activity was generally right-lateralized in silence and balanced under informational masking, but was not modulated by stimulus lateralization. These results suggest that the same auditory-cortex activity that varies with perceptual awareness also shows sensory response features. This is in accordance with models for visual perception, suggesting that sensory competition determines whether midlevel visual responses occur automatically or vary with perceptual state.

Spatiotemporal dynamics of neural activity related to auditory induction in the core and belt fields of guinea-pig auditory cortex

Auditory induction is a continuity illusion in which missing sounds are perceived under appropriate conditions, for example, when noise is inserted during silent gaps in the sound. To elucidate the neural mechanisms underlying auditory induction, neural responses to tones interrupted by a silent gap or noise were examined in the core and belt fields of the auditory cortex using real-time optical imaging with a voltage-sensitive dye. Tone stimuli interrupted by a silent gap elicited responses to the second tone following the gap as well as early phasic responses to the first tone. Tone stimuli interrupted by broad-band noise (BN), considered to cause auditory induction, considerably reduced or eliminated responses to the tone following the noise. This reduction was stronger in the dorsocaudal field (field DC) and belt fields compared with the anterior field (the primary auditory cortex of guinea pig). Tone stimuli interrupted by notched (band-stopped) noise centered at the tone frequency, considered to decrease the strength of auditory induction, partially restored the second responses from the suppression caused by BN. These results suggest that substantial changes between responses to silent gap-inserted tones and those to BN-inserted tones emerged in field DC and belt fields. Moreover, the findings indicate that field DC is the first area in which these changes emerge, suggesting that it may be an important region for auditory induction of simple sounds.

Comparison of auditory deficits associated with neglect and auditory cortex lesions

In contrast to lesions of the visual and somatosensory cortex, lesions of the auditory cortex are not associated with self-evident contralesional deficits. Only when two or more stimuli are presented simultaneously to the left and right, contralesional extinction has been observed after unilateral lesions of the auditory cortex. Because auditory extinction is also considered a sign of neglect, clinical separation of auditory neglect from deficits caused by lesions of the auditory cortex is challenging. Here, we directly compared a number of tests previously used for either auditory-cortex lesions or neglect in 29 controls and 27 patients suffering from unilateral auditory-cortex lesions, neglect, or both. The results showed that a dichotic-speech test revealed similar amounts of extinction for both auditory cortex lesions and neglect. Similar results were obtained for words lateralized by inter-aural time differences. Consistent extinction after auditory cortex lesions was also observed in a dichotic detection task. Neglect patients showed more general problems with target detection but no consistent extinction in the dichotic detection task. In contrast, auditory lateralization perception was biased toward the right in neglect but showed considerably less disruption by auditory cortex lesions. Lateralization of auditory-evoked magnetic fields in auditory cortex was highly correlated with extinction in the dichotic target-detection task. Moreover, activity in the right primary auditory cortex was somewhat reduced in neglect patients. The results confirm that auditory extinction is observed with lesions of the auditory cortex and auditory neglect. A distinction can nevertheless be made with dichotic target-detection tasks, auditory-lateralization perception, and magnetoencephalography.

Sound-evoked network calcium transients in mouse auditory cortexin vivo

Population calcium signals generated by the action potential activity of local clusters of neurons have been recorded in the auditory cortex of mice using an optical fibre-based approach. These network calcium transients (NCaTs) occurred spontaneously as well as in response to sound stimulation. Two-photon calcium imaging experiments suggest that neurons and neuropil contribute about equally to the NCaT. Sound-evoked calcium signals had two components: an early, fast increase in calcium concentration, which corresponds to the short-latency spiking responses observed in electrophysiological experiments, and a late, slow calcium transient which lasted for at least 1 s. The slow calcium transients evoked by sound were essentially identical to spontaneous NCaTs. Their sizes were dependent on the spontaneous activity level at sound onset, suggesting that spontaneous and sensory-evoked NCaTs excluded each other. When using pure tones as stimulus, the early evoked calcium transients were more narrowly tuned than the slow NCaTs. The slow NCaTs were correlated with global ‘up states' recorded with epidural potentials, and sound presented during an epidural ‘down state' triggered a calcium transient that was associated with an epidural ‘up state'. Essentially indistinguishable calcium transients were evoked by optogenetic activation of local clusters of layer 5 pyramidal neurons in the auditory cortex, indicating that these neurons play an important role in the generation of the calcium signal. Taken together, our results identify sound-evoked slow NCaTs as an integral component of neuronal signalling in the mouse auditory cortex, reflecting the prolonged neuronal activity of local clusters of neurons that can be activated even by brief stimuli.

GABAergic neural activity involved in salicylate-induced auditory cortex gain enhancement

Although high doses of sodium salicylate impair cochlear function, it paradoxically enhances sound-evoked activity in the auditory cortex (AC) and augments acoustic startle reflex responses, neural and behavioral metrics associated with hyperexcitability and hyperacusis. To explore the neural mechanisms underlying salicylate (SS)-induced hyperexcitability and "increased central gain," we examined the effects of GABA receptor agonists and antagonists on SS-induced hyperexcitability in the AC and startle reflex responses. Consistent with our previous findings, local or systemic application of SS significantly increased the amplitude of sound-evoked AC neural activity, but generally reduced spontaneous activity in the AC. Systemic injection of SS also significantly increased the acoustic startle reflex. S-baclofen or R-baclofen, GABA-B agonists, which suppressed sound-evoked AC neural firing rate and local field potentials, also suppressed the SS-induced enhancement of the AC field potential and the acoustic startle reflex. Local application of vigabatrin, which enhances GABA concentration in the brain, suppressed the SS-induced enhancement of AC firing rate. Systemic injection of vigabatrin also reduced the SS-induced enhancement of acoustic startle reflex. Collectively, these results suggest that the sound-evoked behavioral and neural hyperactivity induced by SS may arise from a SS-induced suppression of GABAergic inhibition in the AC.

Fast Propagating Waves within the Rodent Auditory Cortex

Central processing of acoustic signals is assumed to take place in a stereotypical spatial and temporal pattern involving different fields of auditory cortex. So far, cortical propagating waves representing such patterns have mainly been demonstrated by optical imaging, repeatedly in the visual and somatosensory cortex. In this study, the surface of rat auditory cortex was mapped by recording local field potentials (LFPs) in response to a broadband acoustic stimulus. From the peak amplitudes of LFPs, cortical activation maps were constructed over 4 cortical auditory fields. Whereas response onset had same latencies across primary auditory field (A1), anterior auditory field (AAF), and ventral auditory field and longer latencies in posterior auditory field, activation maps revealed a reproducible wavelike pattern of activity propagating for ∼45 ms poststimulus through all cortical fields. The movement observed started with 2 waves within the primary auditory fields A1 and AAF moving from ventral to dorsal followed by a motion from rostral to caudal, passing continuously through higher-order fields. The pattern of propagating waves was well reproducible and showed only minor changes if different anesthetics were used. The results question the classical "hierarchical" model of cortical areas and demonstrate that the different fields process incoming information as a functional unit.

Intention and attention: Top-down influences on the representation of task-relevant sounds.

To explore the role of top-down projections from frontal cortex (FC) to auditory cortex (AC), which may play a role in adaptive reshaping of A1 receptive fields to attended acoustic stimuli during behavior, simultaneous recordings were made from single neurons in FC and AC in behaving ferrets, trained on multiple auditory detection and discrimination tasks in positive and negative reinforcement paradigms. Performance required selective attention to different salient spectral frequency and/or temporal cues. Previous observations revealed task-specific transformations in receptive fields in AC [Fritz et al. (2003); (2005); (2007); (2009); Atiani et al. (2009)]. In contrast, FC neurons showed recognition responses, which categorically distinguished between acoustic foreground and background stimuli. FC responses to targets were often independent of the acoustic properties of the target and thus encoded an abstract representation of the class of target stimuli [Fritz et al. (2009)]. Stimulation of FC, paired with tones, lead to receptive field transformations similar to those observed in behavior. These results emphasize the importance of interactions between multiple areas during selective attention, and the tight coupling of target recognition and auditory attention that enhances auditory cortical filters for attended acoustic stimuli, thus creating a functional representation of task-salient sounds during behavior. [This work was supported by grants from NIDCD, NIH.]