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[Comparative study of interneuronal relations in the auditory cortex of awake and anesthetized cats].

The character of interneuronal relations in the auditory cortex of alert and anaesthetized cats (nembutal) with chronicly inplanted electrodes was studied with the method of statistic analysis of cross-intervals of the two impulse series. The analysis of the histograms, obtained by means of processing a neuronal activity, showed that nembutal did not eliminate the dependent relations between neurones and that in the majority of cases the types of these relations are either retained or supplemented with new components. Experiments with a reduced dose of nembutal permitted to trace in time the changes in the amount of the inhibitory and excitatory interrelations in the anaesthetized state, and to compare these changes to the changes in the frequency of spike activity. It was found that nembutal predominantly suppresses the activity of the neurones, generating small spikes. The number of inhibitory connections is reduced simultaneously. Such synchroneity permits to assume the participation of the neurones generating small spikes in the establishment of inhibitory interrelations in the cat auditory cortex.

Anesthesia, General

Ferrier and the study of auditory cortex.

David Ferrier was a British physician who studied the localization of function in the cerebral hemispheres during the latter half of the 19th century. Using stimulation and ablation techniques, Ferrier demonstrated that auditory cortex was located in the superior temporal gyrus of the monkey and that ablation of auditory cortex resulted in deafness. Although he was substantially correct, Ferrier's location of auditory cortex was not accepted by his contemporaries, and his observations of cortical deafness were, until recently, discounted by modern researchers. Just why his findings were rejected is of interest to the study of cortical function.

Auditory Cortex

Role of context in the expression of learning-induced plasticity of single neurons in auditory cortex.

Classical conditioning produces frequency-specific plasticity of receptive fields (RFs) of single neurons in cat auditory cortex (Diamond & Weinberger, 1986). In this article we show that although plasticity may be observed during both training trials and determination of RFs, it is usually expressed in a qualitatively different form (e.g., decreased response during conditioning vs. increased response to this same conditioned stimulus in the postconditioning RF). This differential expression of learning-induced plasticity provides evidence for a role of context in neurophysiological mechanisms of learning in auditory cortex. A model of cortical neurons functioning within a mosaic of influences is presented. The Functional Mosaic model views the induction and expression of plasticity as separate processes.

Animals

Extrathalamic ascending projections to physiologically identified fields of the cat auditory cortex.

The neurons of origin of ascending extrathalamic projections to the auditory cortex were labeled retrogradely with WGA-HRP injected in physiologically identified auditory cortical fields of the cat (anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) fields). After injection in the tonotopically organized auditory cortical fields (AAF, AI and PAF), labeled neurons were distributed in 7 extrathalamic subcortical regions included in one or the other of 2 distinct systems of ascending projections to the neocortex. In the 'diffuse' system of projection, labeled neurons were observed bilaterally in the locus coeruleus, the nuclei of the raphe, the lateral hypothalamus, ipsilaterally in the ventromedial mesencephalic tegmentum and the basal forebrain; in the 'accessory sensory' system of projection, labeled neurons were found ipsilaterally in the nucleus of the brachium of the inferior colliculus and bilaterally in the claustrum. After injection in AII, labeled neurons were seen only in the 'diffuse' system of projection. For AAF and AI, the major contribution to the total extrathalamic ascending input originated from the lateral hypothalamus, whereas for AII it was the locus coeruleus. In contrast, PAF received extrathalamic ascending inputs mainly from the claustrum. Anterogradely labeled corticofugal terminal fields were found only in the nucleus of the brachium of the inferior colliculus and, after injection in PAF, in the claustrum.

Acetylcholine

Foetal and neonatal development of evoked responses in guinea-pig auditory cortex.

Development of the response of the auditory cortex to unilateral acoustic stimulation by a chick was studied in guinea-pig foetuses from the 50th day to the end of gestation and in newborn animals. The first cortical response appeared on the 52nd to 53rd day of gestation. The maximum responses were concentrated in the temporal cortex, between the somatosensory (parietal) and optic (occipital) area. The progressive development of the latent period of the cortical response and of its various components distinctly slowed down on the last days of gestation. At the same time, the amplitude of the cortical response was temporarily augmented. The cortical response developed from a simple negative wave in the youngest embryos into an intricate complex with an initial positive component in newborn guinea-pigs. The basic components of this complex were already discernible on the 64th to 65th day of gestation. The ability to react to repeated peripheral stimulation of 0.1-2 c/s frequency increased with foetal age, with temporary deterioration on the last days of gestation. Resistance of the cortical auditory response to cerebral anoxia rose up to term, with a temporary drop from the 64th day of gestation. After the initiation of independent respiration, cerebral hypoxia and bilateral vagotomy chiefly influenced the stability of the more recent components of the cortical auditory response in mature foetuses.

Acoustic Stimulation

Modification of neuromagnetic responses of the human auditory cortex by masking sounds.

We have studied the effects of masking sounds on auditory evoked magnetic fields (AEFs) of healthy humans. The AEFs were elicited by 25-ms tones presented randomly to the left or to the right ear, and the responses were recorded over the right auditory cortex. Without masking, the 100-ms deflection (N100m) was of somewhat higher amplitude and of shorter latency for contra- than ipsilateral stimuli. Continuous speech, music, or intermittent noise, delivered to the left ear, dampened N100m to stimulation of both ears without correlated changes in sensation. Intermittent noise had a weaker effect on N100m than speech or music. Continuous noise fed to the left ear dampened both the sensation of and the responses to the left-ear stimuli, with no significant effect on the responses to the right-ear stimuli. The results suggest that the masking effects of continuous noise, seen at the auditory cortex, derive mainly from the periphery whereas the effects of sounds with intensity and frequency modulations take place at more central auditory pathways.

Acoustic Stimulation

Age-related deterioration of pyramidal cell basal dendrites in rat auditory cortex.

The basal dendritic trees of layer V pyramidal cells in the rat auditory cortex were examined quantitatively in a group of 3-month-old and a group of 34- and 36-month-old rats. Two forms of analysis were used on the Golgi preparations: (1) the number of intersections between the basal dendrites and a series of concentric circles whose common center lies over the perikaryon center, and (2) the number of dendritic branches, by order, per neuron. The data indicate that in the old animals the density of the dendritic tree has decreased significantly within a radius of about 150mu of the perikaryon, yet the extent of the dendritic domain has not changed appreciably. Analysis of the dendritic branching suggests that there has been a deterioration not only in the peripheral branches of the dendritic tree, but also that entire dendrites have been lost. This loss of primary branches was confirmed through the reconstruction of layer V neuronal perikarya and their proximal dendrites from 1-mu plastic serial sections of auditory cortex. Concomitant with the loss of dendrites which accompanies advancing age is a tendency for the perikaryon to be smaller, but not distorted, in the old animals.

Aging

A study on the tonotopic organization in the auditory cortex of the cat; an application of the glycine labelling method.

3H-glycine was locally applied to the auditory cortex of chloralose anaesthetized cats. Upon tonal stimulation the 3H-glycine was taken up and incorporated into the proteins of nerve cells. The selectively activated neurons were visualized by serial light microscopic autoradiography. Systematic application of this experimental setup revealed tonotopic organization in the primary auditory cortex. The distribution of nerve cells responding to spectrally pure, continuous tones of 0.34, 3.3, 8.0, 16 and 30 kHz was mapped. At these frequencies, distinct but overlapping representations were found, whose area increased in parallel with the elevation of frequencies. Tone pips and ramp stimuli resulted in generalized labelling, independently of pitch.

Animals

[The connections of the parietal cortex with the lateral suprasylvian gyrus (the Clare-Bishop field) and the auditory cortex in the cat].

It has been shown that a parietal projection to the Clare--Bishop area is moderate and organized in a topographic manner. Associative fibres of area 5 terminate in the anterior part of the Clare--Bishop area, which corresponds to the intermediate and anterior part of the posterior suprasylvian sulcus belt. Area 7 projects to the posterior part of the intermediate and posterior suprasylvian sulcus belt. Area 5 and 7 send a few fibres to the auditory cortex. Associative fibres of area 5 terminate in the middle ectosylvian and sylvian gyri: areas 22, 50. Area 7 is connected only with the superior extremity of the middle ectosylvian gyrus or of areas 22, 50.

Afferent Pathways

Auditory cortex of the long-eared hedgehog (Hemiechinus auritus): II. Tuning properties.

The forebrain of hedgehogs is considered by many investigators as one of the simplest and most primitive among extant placental mammals. In a recent study we have shown that the auditory cortex of the long-eared hedgehog (Hemiechinus auritus) comprises two distinct auditory fields, which are tonotopically organized. In this study, we describe tuning properties of single cells in these two fields. Application of the Q10dB and Square Root measures for determining sharpness of tuning revealed that, although most of the cells in the more anterior field, which is considered primary, are sharply tuned, on the average they are more broadly tuned than cells in the primary auditory cortex of other mammals. In the posterior field, the distribution of narrowly and broadly tuned cells is equal. Narrowly tuned cells in both fields are equally narrow, as are the broadly tuned cells. Latencies of single cells in both fields are frequency and intensity dependent and are somewhat longer than these found in other mammals. The distribution of BFs vs. threshold intensity matches fairly well the behavioral audiogram previously described. Our findings suggest that, in spite of the view that the isocortex of hedgehogs represents a 'primitive' condition, some basic tuning properties of their auditory cortex cells are comparable to those of other mammals.

Animals

Effect of bilateral auditory cortex lesions on sound localization in Japanese macaques.

1. The ability of four Japanese macaques (Macaca fuscata) to localize sound was determined after bilateral ablation of auditory cortex. The animals were given two tests: a "midline" test in which they had to discriminate noise bursts presented from a loudspeaker located to the left from identical noise bursts presented from a loudspeaker located to the right of midline, and a "hemifield" test in which both loudspeakers were located in their right hemifield. 2. Both of the tests were administered by the use of two different behavioral tasks: a conditioned-avoidance task in which the animals were trained to make or break contact with a water spout to indicate the location of a sound source, and a two-choice task that required the animals to walk to the source of the sound. 3. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were able to perform the midline discrimination although their localization acuity was reduced. However, the animals had great difficulty in learning to walk to the source of a sound in spite of the fact that they had received previous sound-localization training in the conditioned-avoidance task. This difficulty suggested that the monkeys no longer associated the sound with a location in space. 4. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were unable to discriminate the locus of a sound source when both loudspeakers were located in the same hemifield. 5. Bilateral ablation of auditory cortex results in both sensory and perceptual deficits. The presence of sensory deficits is indicated by the decreased acuity in the left-right discrimination and the inability to discriminate between two loudspeakers located in the same hemifield. The deficit in the perception of the locus of sound is indicated by the difficulty in learning to approach the source of a sound, an ability which normal monkeys exhibit without training. 6. There appear to be species' differences in the effect of auditory cortex lesions on sound localization. Although cortical lesions result in a sound-localization deficit in several species of primates and carnivores, they have little or no effect on rats.

Animals

[Role of the auditory cortex in animal recognition of synthesized vowels].

Discrimination of synthesized vowels [a] and [i] was studied in intact dogs and animals with an ablated auditory cortex. Electro-defensive and conditioned reflex methods were used. It has been found that as a result of learning an auditory image of a stationary vowel can be formed in intact dogs, which is invariant relatively to any change in the basic frequency of the voice, the intensity and duration of presentation (300 to 75 msec), and that two formants are sufficent for discrimination. The auditory image of the vowel is preserved after a bilateral ablation of the auditory cortical projection zone. Discrimination of vowels is disturbed in animals with a removed auditory cortex when information redundancy is reduced (diminished number of formants and reduced signal duration).

Animals

[Reactions of neurons of the auditory cortex of unanesthetized cats to tones of a characteristic frequency].

Extra- and intracellular responses of primary auditory cortex (AI) neurons were studied in acute experiments on non-anaesthetized cats. It was found that auditory cortex neurons having similar best frequencies revealed various forms of responses to corresponding frequency tones. Neurons responded to the tone by on reactions constituted about 40% of nerve cells studied. 27% of neurons revealed responses of on-off and off types. 27% of cortical neurons responded by steady excitation or by inhibition of background activity. About 6% of neurons did not respond to the tone. During intracellular recordings about 85% of neurons studied responded to switching on and/or off of the tone by a spike-IPSP sequence. 96% of cortical neurons generated IPSP as a constant component of the response to tone. Tonic responses of auditory cortical neurons were the result of powerful and lasting depolarization of the postsynaptic membranes. The conclusion is made that interaction of excitatory and inhibitory processes is the most significant in any kind of responses of the auditory cortical neurons to tones.

Animals

Responses of the human auditory cortex to vowel onset after fricative consonants.

Neuromagnetic responses to different auditory stimuli (noise bursts and short speech stimuli) were mapped over both hemispheres of seven healthy subjects. The results indicate that a particular acoustic feature of speech, vowel onset after voiceless fricative consonants, evokes a prominent response in the human supratemporal auditory cortex. Although the observed response seems to be specific to acoustic rather than phonetic characteristics of the stimuli, it might reflect feature detection essential for further speech processing.

Auditory Cortex

Acquired word deafness, and the temporal grain of sound representation in the primary auditory cortex.

This paper explores the nature of the processing disorder which underlies the speech discrimination deficit in the syndrome of acquired word deafness following from pathology to the primary auditory cortex. A critical examination of the evidence on this disorder revealed the following. First, the most profound forms of the condition are expressed not only in an isolation of the cerebral linguistic processor from auditory input, but in a failure of even the perceptual elaboration of the relevant sounds. Second, in agreement with earlier studies, we conclude that the perceptual dimension disturbed in word deafness is a temporal one. We argue, however, that it is not a generalized disorder of auditory temporal processing, but one which is largely restricted to the processing of sounds with temporal content in the milliseconds to tens-of-milliseconds time frame. The perceptual elaboration of sounds with temporal content outside that range, in either direction, may survive the disorder. Third, we present neurophysiological evidence that the primary auditory cortex has a special role in the representation of auditory events in that time frame, but not in the representation of auditory events with temporal grains outside that range.

Animals

Neuronal connections in the primary auditory cortex: an electrophysiological study in the cat.

Neuronal connections in the primary auditory cortex (AI) of the cat were studied electrophysiologically by using intracellular recording techniques. Fast-conducting fibers from the medial geniculate nucleus (MG) projected monosynaptically onto AI neurons in layers III-VI (mainly in layer IV), whereas slow-conducting MG-fibers projected monosynaptically onto AI neurons in layer I. AI neurons which received monosynaptic inputs from the auditory association cortices (AII and Ep) and/or from the contralateral AI were distributed in all layers of the AI; the commissural fibers from the contralateral AI were divided into fast- and slow-conducting ones. AI neurons were categorized into seven types: type I neurons which received monosynaptic inputs from slow-conducting MG-fibers were located in layer I. Type II neurons which received polysynaptic inputs from the MG were located in layers II-VI. Type III neurons which sent their axons to the AII or Ep were mainly located in layer III. Type IV neurons which sent their axons to the contralateral AI were located mainly in layer III. Type V neurons which received monosynaptic inputs from fast-conducting MG-fibers were located mainly in layer IV. Type VI neurons which projected onto the inferior colliculus were located in the upper part of the layer V. Type VII neurons which projected onto the MG were located in layers V and VI.

Animals

Habituation produces frequency-specific plasticity of receptive fields in the auditory cortex.

Associative learning produces conditioned stimulus (CS)-specific plasticity of frequency receptive fields (RFs) in the auditory cortex; responses to the CS frequency are increased, whereas responses to other frequencies are decreased. This study determined the effects of habituation on the RF of neurons in the auditory cortex of the guinea pig (Cavia porcellus). One frequency was presented repeatedly (REP) followed by redetermination of the RF. After REP, 26/36 (72%) RFs exhibited a substantial reduction (70-75%) of response to the repeated frequency, and this was highly specific (bandwidth less than 0.125 octave). This RF plasticity involves an initial decrease in response during REP but does not require attenuated responses at the end of REP. Incubation (i.e., development over time after cessation of REP) and long-term frequency-specific effects are evident. Thus, habituation induces a specific change in the processing of frequency information rather than a general reduction in responsivity.

Acoustic Stimulation

Neural representation of sound amplitude in the auditory cortex: effects of noise masking.

Single auditory cortical neurons express their sensitivity to the amplitude of a preferred-frequency tone pulse as either a monotonic, saturating intensity profile or as a non-monotonic, bell-shaped intensity function. In the presence of continuous, wideband noise masking, the tone intensity profile is displaced toward higher tone levels. The magnitude of the tone threshold adjustments brought about by increments in noise level very closely match the elevations in noise amplitude. The mechanisms underlying the threshold adjustments likely include neural adaptation. This is because the tone threshold shifts seen in the spike count data are paralleled by spike latency data, and because recovery of tonal sensitivity following noise offset proceeds in a negatively-accelerating fashion. In some instances, the slope of the masked tone intensity profile is greater than that for unmasked tones. For masked tone levels evoking submaximal responses, this has the consequence that cortical responses to masked tones are somewhat more salient than those for unmasked tones of comparable suprathreshold level. These observations bolster our understanding of the psychophysics of noise-masking in normal listeners, and they provide a partial explanation of the difficulty shown by patients with temporal lobe lesions in discriminating signals in noise.

Animals