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Interhemispheric asymmetry of homotopic transcallosal responses of the auditory cortex in the cat.

In acute experiments on 29 d-tubocurarine immobilized cats, with topographical pick-up of potentials from 150-170 symmetrical points of the auditory cortex, the interhemispheric asymmetry of homotopic transcallosal responses were studied. It was established that in the majority of the animals the potentials in the right hemisphere had a greater amplitude and shorter latent periods as compared with the responses of the left hemisphere. A dependence of the properties of the interhemispheric asymmetry on the sex of the animal was noted. In females the asymmetry was characterized by a right hemispheric tendency and by a greater magnitude as compared with males. The maximal magnitude of the asymmetry was detected in the group of pregnant females. The hypothesis is proposed to the effect that the established polarity of transcallosal influenced in the auditory cortex is associated with the dominance of the right hemisphere in the organization of spatial audition.

Acoustic Stimulation↗

Visual projections induced into the auditory pathway of ferrets: II. Corticocortical connections of primary auditory cortex.

Although the development of corticocortical projections has been well studied, less is known about the role of sensory inputs in the specification of these connections. As part of an ongoing series of studies in our laboratory, we have examined the role of thalamic input modality in the development of corticocortical connections. These studies involve making unilateral lesions and inducing retinal inputs into the auditory thalamus (MGN) during early development in ferrets, thereby conferring visual responsiveness on primary auditory cortex (AI). In this way we can examine the role of input identity in cortical specification in general, and connectivity patterns specifically. A previous paper (Pallas et al. [1990] J. Comp. Neurol. 298:50-68) described the pattern of thalamocortical and corticothalamic connections of auditory cortex in normal and lesioned animals. This study compares the pattern of auditory corticocortical connections in normal and lesioned animals. We injected neuroanatomical tracers into AI and mapped out the distribution of retrogradely labelled cells in the cortex. We report that the cortical inputs to ferret AI resembled those in cats, and that the pattern of ipsi- and contralateral corticocortical connections of ferret AI with visual input was similar to the normal pattern. Auditory cortex with visual input did not make ectopic connections with visual cortex, but maintained its connections with other auditory cortical areas. These results suggest that the overall corticocortical connections of an area are not influenced by the modality or activity pattern of its inputs. In particular, altering the input activity to a cortical area does not seem to promote the formation of entirely new connections, although small changes in the strength of existing connections are possible (Sur et al. [1990] Trends Neurosci. 13:227-233).

Animals↗

Role of cat primary auditory cortex for sound-localization behavior.

Small lesions designed to completely destroy the cortical zone of representation of a restricted band of frequency were introduced within the primary auditory cortex (AI) in adult cats. Physiological mapping was used to guide placement of lesions. Sound-localization performance was evaluated prior to and after induction of these lesions in a seven-choice free-sound-field apparatus. All tested cats had profound contralateral hemifield deficits for the localization of brief tones at frequencies roughly corresponding to those whose representations were destroyed by the lesion. Sound-localization performance was normal at all other test frequencies. In a single adult cat, a massive lesion destroyed nearly all auditory cortex unilaterally, with only the representation of a narrow band of frequency within AI spared by the lesion. This cat had normal abilities for azimuthal sound localization across that frequency band but a profound contralateral deficit for the azimuthal localization of brief sounds at all other frequencies. Recorded sound-localization deficits were permanent. Localization of long-duration tones was not affected by a unilateral AI lesion. These studies indicate that, at least in cats, AI is necessary for normal binaural sound-localization behavior; among auditory cortical fields, AI is sufficient for normal binaural sound-localization behavior; sound-location representation is organized by frequency channel in the auditory forebrain; and AI in each hemisphere contributes to only contralateral free-sound-field location representation.

Animals↗

Binary spiking in auditory cortex.

Neurons are often assumed to operate in a highly unreliable manner: a neuron can signal the same stimulus with a variable number of action potentials. However, much of the experimental evidence supporting this view was obtained in the visual cortex. We have, therefore, assessed trial-to-trial variability in the auditory cortex of the rat. To ensure single-unit isolation, we used cell-attached recording. Tone-evoked responses were usually transient, often consisting of, on average, only a single spike per stimulus. Surprisingly, the majority of responses were not just transient, but were also binary, consisting of 0 or 1 action potentials, but not more, in response to each stimulus; several dramatic examples consisted of exactly one spike on 100% of trials, with no trial-to-trial variability in spike count. The variability of such binary responses differs from comparably transient responses recorded in visual cortical areas such as area MT, and represent the lowest trial-to-trial variability mathematically possible for responses of a given firing rate. Our study thus establishes for the first time that transient responses in auditory cortex can be described as a binary process, rather than as a highly variable Poisson process. These results demonstrate that cortical architecture can support a more precise control of spike number than was previously recognized, and they suggest a re-evaluation of models of cortical processing that assume noisiness to be an inevitable feature of cortical codes.

Action Potentials↗

Selective listening modifies activity of the human auditory cortex.

We have studied the effect of selective listening on the neuromagnetic evoked activity of the human auditory cortex. In the word categorization experiment the stimuli were 5-letter words, each beginning with (k). Half of them were targets, i.e., names of animals or plants, and half other meaningful Finnish words. In the duration discrimination experiment equiprobable tones of 425 ms (targets) or 600 ms duration were presented. In both experiments the interstimulus interval (ISI) was 2.3 s and the stimuli of the two classes were presented randomly. Subjects either ignored the stimuli (reading condition) or counted the number of targets (listening condition). The magnetic field over the head was measured with a 7-channel 1st-order SQUID-gradiometer. The stimuli evoked a transient response followed by a sustained field. The transient response did not differ between the two conditions but the sustained field was significantly larger in the listening than reading condition; the increase began 120-200 ms after stimulus onset and continued for several hundred milliseconds. The equivalent source locations of both transient and sustained responses agreed with activation of the supratemporal auditory cortex. In the dichotic listening experiment 25-ms square-wave stimuli were presented randomly and equiprobably either to the left or to the right ear at an ISI of 0.8-1 s, either alone or in presence of a speech masker. Counting the stimuli of either ear resulted in differences between responses to relevant and irrelevant sounds. The difference began 140-150 ms after stimulus onset and peaked at 200-240 ms. During monaural speech masking, N100m was larger for attended than ignored stimuli.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

fMRI of the auditory cortex in patients with unilateral carotid artery steno-occlusive disease.

PURPOSE: To examine whether an internal carotid artery (ICA) steno-occlusive disease leads to a reduced blood oxygenation level dependent (BOLD)-signal change due to a decreased vasodilatory reserve capacity. MATERIALS AND METHODS: Patients suffering from unilateral ICA stenosis or occlusion were examined using functional magnetic resonance imaging (fMRI) of the auditory cortex with a bilateral stimulation applying a pulsed 1000 Hz sine-tone. RESULTS: Compared to control subjects, who showed symmetric bilateral BOLD-responses within the auditory cortex, the ICA patients revealed either a normal bilateral cortical activation pattern or a reduced cortical activation on the steno-occluded side. This latter decrease of BOLD-signal change might indicate a depressed vasomotor reserve capacity. The observed coincidence between this asymmetric reduction in BOLD-signal and ischemic borderzone lesions on the affected side in this subgroup of patients strongly supports this assumption. CONCLUSION: This study shows that fMRI of the auditory cortex appears to have clinical merit in the workup of cerebrovascular conditions.

Acoustic Stimulation↗

Centripetal and centrifugal reorganizations of frequency map of auditory cortex in gerbils.

As repetitive acoustic stimulation and auditory conditioning do, electric stimulation of the primary auditory cortex (AI) evokes reorganization of the frequency map of AI, as well as of the subcortical auditory nuclei. The reorganization is caused by shifts in best frequencies (BFs) of neurons either toward (centripetal) or away from (centrifugal) the BF of stimulated cortical neurons. In AI of the Mongolian gerbil, we found that focal electrical stimulation evoked a centripetal BF shift in an elliptical area centered at the stimulated neurons and a centrifugal BF shift in a zone surrounding it. The 1.9-mm long major and 1.1-mm long minor axes of the elliptical area were parallel and orthogonal to the frequency axis, respectively. The width of the surrounding zone was 0.2-0.3 mm. Such "center-surround" reorganization has not yet been found in any sensory cortex except AI of the gerbil. The ellipse is similar to the arborization pattern of pyramidal neurons, the major source of excitatory horizontal connections in AI, whereas the surrounding zone is compatible to the arborization range of small basket cells (inhibitory neurons) in AI.

Animals↗

Neurotoxic effects of mercury on auditory cortex networks growing on microelectrode arrays: a preliminary analysis.

Mercury is known to cause sensorineural hearing loss and impaired speech perception. However, there is still a lack of a quantitative description of mercury toxicity on central auditory structures. This is a preliminary study using the novel technique of microelectrode array (MEA) recordings to evaluate acute and chronic neurotoxic effects of mercury on auditory cortex networks (ACNs) in vitro. Morphological and electrophysiological effects of mercuric chloride (HgCl(2)) were studied. Neurons dissociated from auditory cortices of 14-day-old mouse embryos were grown on photoetched MEAs containing 64 transparent indium-tin oxide (ITO) electrodes. For acute electrophysiological experiments, the spontaneous spiking and bursting activity from ACNs were compared before and after application of HgCl(2). For chronic electrophysiological experiments, auditory cortex cultures were treated with various concentrations of HgCl(2) from the day of seeding, and were tested 4 weeks later for the presence of spontaneous activity. Morphological analysis was conducted on 8-day-old ACNs treated with HgCl(2) for 3 days. Results of acute experiments indicated that <75 mM of HgCl(2) had an excitatory effect of variable magnitude on the spontaneous activity of ACNs; however, concentrations above 100 microM completely and irreversibly inhibited spike and burst activity. Chronic exposure of ACNs to 10 microM HgCl(2) completely blocked the spontaneous activity. Morphological analysis indicated that 10 microM HgCl(2) caused neuronal cell death in 3 days. It is concluded that HgCl(2) has a more toxic effect on auditory networks when exposed chronically, and the levels of mercury showing toxic effects on ACNs are within the dose range shown to cause neurologic symptoms in humans.

Action Potentials↗

Single unit activity vs. amplitude of the epidural evoked potential in primary auditory cortex of awake cats.

The study investigated, in primary auditory cortex (AI) of awake cats, the relationship over a range of stimuli between the amplitude and latency of the initial positive deflection (P1) of the primary evoked potential and the intensity of concurrent underlying evoked single unit activity. Epidural evoked potentials and extracellular responses of 155 single units to monaural 100 musec clicks ranging from 45 to 110 dB were recorded. At low stimulus levels, considerable unit response could occur with a very small P1. At middle stimulus levels, unit response was directly proportional to P1 amplitude. At higher stimulus levels, P1 amplitude continued to increase while unit response began to saturate.

Animals↗

Descending projections from the auditory cortex to the inferior colliculus in the gerbil, Meriones unguiculatus.

Corticofugal projections to the auditory midbrain, the inferior colliculus (IC), influence the way in which specific sets of IC neurons process acoustic signals. We used retrograde tracer (Fluorogold, Fluororuby, microbeads) injections in the IC to study the morphology and location of cortico-collicular projecting neurons and anterograde tracer (dextran biotin) injections in auditory cortical fields to describe the distribution of terminals in the IC. Nissl staining, cytochrome oxidase activity, and neurofilament SMI32 immunostaining were used to delimit the different auditory areas. We defined a primary or "core" auditory cortex and a secondary "caudal" auditory area containing layer V pyramidal neurons that project to the IC. These projections target the central nucleus of the IC (CNIC) ipsilaterally and the IC cortices bilaterally, with the ipsilateral component predominant. Other secondary auditory areas, dorsal and ventral to the core, do not directly participate in this projection. The ventral secondary cortex targets midbrain periaqueductal gray. The projection from the core cortex originates from two classes of layer V pyramidal cells. Cells presenting a tufted apical dendrite in layer I have dense terminal fields in the IC cortices. Pyramids lacking layer I dendritic tufts target the CNIC in a less dense but tonotopic manner. The caudal cortex projection originates from smaller layer V pyramids and targets the IC cortices with dense terminal fields. Descending auditory inputs from the core and caudal areas converge in the dorsal and external cortices of the IC. Descending connections to the gerbil IC form a segregated system in which multiple descending channels originating from different neuronal subpopulations may modulate specific aspects of ascending auditory information.

Animals↗

Time course of tonal frequency-response-area of primary auditory cortex neurons in alert cats.

Cells in the A1 auditory cortex of alert animals show various response time-courses during pure-tone stimuli: tonic, phasic-tonic, and phasic. Previously the time course of the spike firing rates was examined at a characteristic frequency (CF) or in a range of frequencies including CF. We investigated time-course of the frequency-response-area (FRA) during pure-tone stimuli in A1 cells of alert cats. The short rise/fall time (0.1-2 ms) and long stimulus duration (0.5 s) was used for investigation of the time course. FRA changed with time drastically in the phasic cells, mildly in the phasic-tonic cells, but not in the tonic cells. The best-response frequency (BF) within FRA was constant throughout the stimulus duration in the tonic and phasic-tonic cells, but was difficult to define in the phasic cells. The phasic firing properties of the phasic cells were preserved even during the bandnoise stimuli at various bandwidth and spectral locations. The variability of FRA time-course between cell types may play a role for analyzing auditory spectral cues that vary with a wide range of time constant.

Acoustic Stimulation↗

Heterosynaptic long-term facilitation of sensory-evoked responses in the auditory cortex by stimulation of the magnocellular medial geniculate body in guinea pigs.

The magnocellular nucleus of the medial geniculate body (MGm) develops physiological plasticity during classical conditioning and may be involved in learning-induced receptive field plasticity in the auditory cortex. To determine the ability of the MGm to produce long-term modification of evoked activity in the auditory cortex, the experimenters paired electrical stimulation of the MGm with preceding clicks in adult guinea pigs under barbiturate anesthesia. The amplitudes of average click-evoked potentials were significantly facilitated in all subjects. Facilitation endured for 2 hr, the maximum duration of recording. Sham-stimulated control guinea pigs did not develop facilitation. Thus, a nonlemniscal thalamic sensory nucleus can produce enduring facilitation of sensory-evoked activity in primary sensory cortex, suggesting that long-term physiological plasticity in the sensory cortex during learning may involve nonlemniscal thalamic mechanisms.

Animals↗

Morphometry of spine-free nonpyramidal neurons in rabbit auditory cortex.

A study of the morphometry and laminar distribution of spine-free nonpyramidal neurons in electrophysiologically verified primary auditory cortex was carried out in adult rabbits. By using image-combining computer microscopy, the locations of all impregnated neurons in 300-micrometers Golgi-Cox Nissl sections through the auditory cortex were determined. Spine-free non-pyramidal neurons constitute nearly 72% of the nonpyramidal neurons present. They are distributed in a band extending from 450 to 750 micrometers beneath the pial surface corresponding to laminae III and IV. A combination of dendritic stick, Fourier, and statistical analyses revealed a highly significant spatial orientation of their dendrite systems along a vertical axis parallel to the apical dendrites of pyramidal neurons. A significant tangential orientation of dendrites along a dorsal-ventral axis was also found. A radial analysis of the dendrite systems revealed that the pronounced vertical orientation of spine-free nonpyramidal neurons is due to (1) directed dendritic growth along the vertical axis, (2) decreased branching and rapid termination of tangentially oriented dendrites, and (3) increased branching of vertically growing dendrites. The radial analysis also revealed that the longest branches are those directed toward the white matter.

Animals↗

Postnatal shift of tonotopic organization in the chick auditory cortex analogue.

The existence of an ontogenetic shift of tonotopic organization throughout the auditory pathway concomitant with cochlea maturation is a matter of controversy. Using the 2-deoxyglucose method we demonstrate here for the first time the shift phenomenon in an auditory forebrain structure, field L, the auditory cortex analogue of the chick. During the first postnatal month isofrequency contours move to positions where, in younger chicks, lower frequencies (up to half an octave) are represented. This developmentally changing place code of sound frequencies at the forebrain level is similar to the one previously reported for brain stem auditory nuclei. It raises the question of constancy of frequency-related pitch perception during development and may be a complication of early auditory learning and memory.

Acoustic Stimulation↗

Associative retuning in the thalamic source of input to the amygdala and auditory cortex: receptive field plasticity in the medial division of the medial geniculate body.

The medial division of the medial geniculate body (MGm) projects to the lateral amygdala and the upper layer of auditory cortex and develops physiological plasticity rapidly during classical conditioning. The effects of learning on frequency receptive fields (RFs) in the MGm of the guinea pig have been determined. Classical conditioning (tone-footshock), as indexed by rapid development of conditioned bradycardia, produced conditioned stimulus (CS)-frequency specific RF plasticity: increased response at the CS frequency with decreased responses at other frequencies, both immediately and after a 1-hr retention period. Sensitization training produced only general changes in RFs. These findings are considered with reference to both the elicitation of amygdala-mediated, fear-conditioned responses and the mechanism of retrieval of information stored in the auditory cortex during acquisition.

Amygdala↗

Effect of bilateral auditory cortex lesions on absolute thresholds in Japanese macaques.

1. The behavioral audiograms of four Japanese macaques (Macaca fuscata) were assessed before and after receiving two-stage bilateral lesions of auditory cortex. Thresholds were assessed for each ear with the use of insertion earphones. 2. The bilateral lesions resulted in a large initial hearing loss followed by partial recovery that left the animals with a permanent hearing loss in both ears. 3. The initial hearing loss consisted of a total insensitivity to sound in the ear contralateral to the second lesion with limited hearing in the other ear. However, the animal with the most complete lesion was initially unable to hear sound in either ear. Broadband noise was often more effective in eliciting a behavioral response than tones. 4. Partial recovery occurred in all animals and was observed as early as the first week after surgery. Most of this recovery occurred during the first 3-7 wk after surgery. This rapid phase of recovery was sometimes followed by a more gradual phase although thresholds were still elevated after 94 wk. 5. The permanent hearing loss, which averaged from 30 to 44 dB, was not constant across frequency. Threshold shifts were smallest at 63 Hz and progressively increased with frequency to a maximum loss from 8 to 25 kHz with slightly less loss at 32 kHz. 6. Analysis of the psychophysical functions and threshold stability gave no indication of any nonsensory deficits in attention or vigilance. 7. These results, taken with those of previous experiments, indicate that each hemisphere is primarily involved in the detection of sound in the contralateral ear and secondarily involved in detection in the ipsilateral ear. This arrangement differs from that seen in sound localization where each hemisphere is involved with the contralateral hemifield as opposed to the contralateral ear. Thus it appears that the functional organization of auditory cortex for sound localization is different from that for the detection and identification of sound itself.

Animals↗

Seeing speech: visual information from lip movements modifies activity in the human auditory cortex.

Neuromagnetic responses were recorded over the left hemisphere to find out in which cortical area the heard and seen speech are integrated. Auditory stimuli were Finnish/pa/syllables presented together with a videotaped face articulating either the concordant syllable/pa/(84% of stimuli, V = A) or the discordant syllable/ka/(16%, V not equal to A). In some subjects the probabilities were reversed. The subjects heard V not equal to A stimuli as/ta/ or ka. The magnetic responses to infrequent perceptions elicited a specific waveform which could be explained by activity in the supratemporal auditory cortex. The results show that visual information from articulatory movements has an entry into the auditory cortex.

Adult↗

Azimuth coding in primary auditory cortex of the cat. I. Spike synchrony versus spike count representations.

The neural representation of sound azimuth in auditory cortex most often is considered to be average firing rate, and azimuth tuning curves based thereupon appear to be rather broad. Coincident firings of simultaneously recorded neurons could provide an improved representation of sound azimuth compared with that contained in the firing rate in either of the units. In the present study, a comparison was made between local field potentials and several measures based on unit firing rate and coincident firing with respect to their azimuth-tuning curve bandwidth. Noise bursts, covering a 60-dB intensity range, were presented from nine speakers arranged in a semicircular array with a radius of 55 cm in the animal's frontal half field. At threshold intensities, all local field potential (LFP) recordings showed preferences for contralateral azimuths. Multiunit recordings showed in 74% a threshold for contralateral azimuths, in 16% for frontal azimuths, and in only 5% showed an ipsilateral threshold. The remaining 5% were not spatially tuned. Representations for directionally sensitive units based on coincident firings provided significantly sharper tuning (50-60 degrees bandwidth at 25 dB above the lowest threshold) than those based on firing rate (bandwidths of 80-90 degrees). The ability to predict sound azimuth from the directional information contained in the neural population activity was simulated by combining the responses of the 102 single units. Peak firing rates and coincident firings with LFPs at the preferred azimuth for each unit were used to construct a population vector. At stimulus levels of >/=40 dB SPL, the prediction function was sigmoidal with the predicted frontal azimuth coinciding with the frontal speaker position. Sound azimuths >45 degrees from the midline all resulted in predicted values of -90 or 90 degrees, respectively. No differences were observed in the performance of the prediction based on firing rate or coincident firings for these intensities. This suggests that although coincident firings produce narrower azimuth tuning curves, the information contained in the overall neural population does not increase compared with that contained in a firing rate representation. The relatively poor performance of the population vector further suggests that primary auditory cortex does not code sound azimuth by a globally distributed measure of peak firing rate or coincident firing.

Acoustic Stimulation↗