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PubMed1975

Representation of cochlea within primary auditory cortex in the cat.

The representation of sound frequency (and of the cochlear partition) within primary auditory cortex has been investigated with use of microelectrode-mapping techniques in a series of 25 anesthetized cats. Among the results were the following: 1) Within vertical penetrations into AI, best frequency and remarkably constant for successively studied neurons across the active middle and deep cortical layers. 2) There is an orderly representation of frequency (and of represented cochlear place) within AI. Frequency is rerepresented across the mediolateral dimension of the field. On an axis perpendicular to this plane of rerepresentation, best-frequency (represented cochlear place) changes as a simple function of cortical location. 3) Any given frequency band (or sector of the cochlear partition) is represented across a belt of cortex of nearly constant width that runs on a nearly straight axis across AI. 4) There is a disproportionately large cortical surface representation of the highest-frequency octaves (basal cochlea) within AI. 5) The primary and secondary field locations were somewhat variable, when referenced to cortical surface landmarks. 6) Data from long penetrations passing down the rostral bank of the posterior ectosylvian sulcus were consistent with the existence of a vertical unit of organization in AI, akin to cortical columns described in primary visual and somatosensory cortex. 7) Responses to tonal stimuli were encountered in fields dorsocaudal, caudal, ventral, and rostral to AI. There is an orderly representation of the cochlea within the field rostal to AI, with a reversal in best frequencies across its border with AI. 8) Physiological definitions of AI boundaries are consistent with their cytoarchitectonic definition. Some of the implications of these findings are discussed.

Acoustic Stimulation↗
PubMed1976

Evoked unit activity in auditory cortex of monkeys performing a selective attention task.

Single-unit responses were recorded from the auditory cortex of rhesus monkeys performing a selective atteintion task which used combined light and sound stimuli. The animals were first trained to push a lever to the left for a noise burst and to the right for a tone burst, and then trained to push left for a left light and right for a right light. Subsequently, one of the four possible light and sound stimulus combinations (Noise + Left Light, Noise + Right Light, Tone + Left Light, Tone + Right Light) was randomly presented on each trial. In blocks of 100 trials only one part of the combined stimulus (either the light or the sound) determined the direction of lever push that would be reinforced. Responses of single units to identical sound stimuli were compared for blocks in which sound was the relevant cue and blocks in which light was the relevant cue. Typically, differences were in response strength without alteration of response pattern. Even the earliest response components (15-20 msec latency) could show changes. Two-thirds of the response comparisons showed differences in strength depending upon whether sound or light was the relevant cue, with about as many responses stronger for light relevant as for sound relevant. Independent of which modality was relevant, unit responses for trials in which both parts of the combined light and sound stimulus indicated the same direction of lever push were generally stronger than unit responses for trials in which the two parts of the combined stimulus signalled opposite directions of lever pushes.

Animals↗
PubMed1976

Relation of unit spike discharges and evoked potentials in the auditory cortex of cats.

Extracellular microelectrode recordings were made from the auditory cortex of anaesthetized cats during acoustic click stimulation. The microelectrode of low resistance allowed to record evoked field potentials and unit discharges simultaneously. In distant extracellular leads the relation of unit discharges and field potentials was equivocal. Near extracellular leads revealed that the antidromic invasion of the somadendritic membrane by excitation is a frequency dependent process (just as evoked field potentials) while spike potentials can reliably be elicited from the initial segment at high frequencies. It is assumed that the excitation spreading from the initial segment to the soma-dendritic membrane represents an important component of the evoked potentials, and their frequency dependence may be traced back to inhibitions activated by afferent impulses.

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↗
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↗
PubMed1975

A Golgi and electron microscopical study of nerve cells in layer I of the cat auditory cortex.

The nerve cells of the plexiform layer of the cat auditory cortex were studied in Golgi rapid preparations, and by electron microscopy. In the Golgi material 178 cells were found. Thirty were horizontal cells (Cajal's cells) with spiny dendrites and a long axon with horizontal trajectory. Seventy-two small cells had smooth dendrites and a short axon branching profusely around the perikaryon, within the plexiform layer. Twenty-eight were slightly larger cells, morphologically similar to the small ones, but their axons and dendrites were longer, extending to layer II. Twenty-five cells had large bottle-shaped perikarya, spiny dendrites reaching layer II, and a vertically oriented axon branching in layer II, and sometimes reaching layer III. In addition, 23 small modified pyramidal cells, similar to those of layer II, were found within the boundaries of layer I. In the most superficial 50 mum of the cortex horizontal cells were found, but the other categories were very rare. The 3 categories of short axon cells were preferentially localized between 50 and 150 mum from the pial surface. By electron microscopy two classes of cells were found. The first had abundant mitochondria, well defined Nissl bodies and few axosomatic synapses. The second had few mitochondria, undefined Nissl bodies and 6-8 axosomatic synapses per profile. Desmosome-like membrane specializations were found between perikarya of the second class and pyramidal dendrites or their spines. It is suggested that the first class of cells found by electron microscopy may correspond to the horizontal cells while the second class may correspond to the 3 categories of short axon cells.

Age Factors↗
PubMed1975

Effects of putative neurotransmitters on neuronal activity in monkey auditory cortex.

The effects of the putative neurotransmitters norepinephrine (NE), gamma-aminobutyric acid (BAGA), and acetylcholine (ACh) were tested on auditory cortex neurons which were activated acoustically by species-specific vocalizations in awake squirrel monkeys. Five-barrel glass electrodes were used to record the activity single neurons in the superior temporal gyrus and to apply NE, GABA, or ACh microiontophoretically. Poststimulus time histograms and raster displays of neuronal responses to the vocalizations were computed before, during, and after iontophoresis. Dose-dependent inhibition of spontaneous and vocalization-evoked discharge rates was seen with NE and GABA. Generally, excitation was observed with ACh. A given dose of NE or GABA reduced spontaneous activity by a greater proportion than it reduced activity evoked by the vocalizations. During excitatory responses, segments with lower discharge rates were reduced proportionately more than segments with higher discharge rates. Usually, response 'pattern' was not altered by iontophoresis of any of the substances. However, in some cases the differential inhibition of slow activity produced by NE or GABA did result in a 'patern' change. The demonstration that small amounts of locally applied NE and GABA substantially alter the specific neuronal activation produced by vocalizations provides additional evidence that these agents may function as neurotransmitters in this neocortical area and offers clues about their functional significance.

Acetylcholine↗
PubMed1977

Organization of auditory cortex in the owl monkey (Aotus trivirgatus).

The region of cerebral cortex in the owl monkey that is responsive to acoustic stimulation is located on the dorsal and lateral surfaces of the rostral half of the superior temporal gyrus. Systematic microelectrode mapping of this area has revealed multiple frequency representations. The boundaries of these fields determined electrophysiologically correlate with the architectural boundaries apparent in Nissl stained material. On the basis of combined cytoarchitectonic and electrophysiological maps we have divided auditory cortex into five fields. Two of them, the primary field (AI) and the field rostral to it (R) are somewhat similar architectonically and constitute the central core of auditory cortex. Each of these two fields has a complete and orderly representation of the audible frequency spectrum within it. Surrounding these fields is a belt of cortex in which units are generally less responsive to acoustic stimulation and the frequency organization is more complex than in AI or R. Electrophysiological and cytoarchitectonic evidence suggest that this belt is composed of at least three and possibly four separate auditory fields.

Acoustic Stimulation↗
PubMed1978

Projections of auditory cortex upon the thalamus and midbrain in the owl monkey.

Two tonotopically organized cortical fields, the primary (AI) and the rostral (R) fields, comprise the core of auditory cortex in the owl monkey. Injections of tritiated proline were made into each of these fields to determine their efferent projections using autoradiographic methods. Both AI and R project to the principal and magnocellular divisions of the medial geniculate body. In addition, R projects to the posterior part of the dorsal division of the medial geniculate. AI sends axons to the dorsomedial region and laminated portion of the central nucleus of the inferior colliculus. Labeling in the central nucleus following AI injections appears as a band of silver grains oriented parallel to isofrequency contours. Axons from R terminate in the dorsomedial region of the central nucleus of the inferior colliculus and in the pericentral and external nuclei of the inferior colliculus. In addition, the rostral field projects to a small area of the medial pulvinar just anterior to the brachium of the superior colliculus.

Animals↗
PubMed1976

Evoked potential decrements in auditory cortex. I. Discrete-trial and continual stimulation.

In experiment 1 cats were exposed to sets of clicks (trials) with 1 min inter-trial-intervals to determine if the effects of repetitive stimulation on potentials evoked in the auditory cortex would be cumulative despite discrete-trial stimulation. Evoked potentials were averaged to give one average evoked potential (AEP) for each trial for each electrode; there were four cortical electrodes per subject. To test for dishabituation pawshocks were given between trials 60 and 61. Subjects were paralyzed to insure stimulus constancy. The latency and peak-to-peak amplitude of each component of each AEP was measured; significant amplitude decremented; and decrements were more frequent in components with latencies greater than 15 msec. A few amplitude increments and latency changes were also observed...

Animals↗
PubMed1978

Aural representation in the Doppler-shifted-CF processing area of the auditory cortex of the mustache bat.

In the mustache bat (Pteronotus pamellii rubiginosus) the frequency and amplitude of an acoustic signal are represented in the coordinates parallel to the surface of the Doppler-shifted-CF (constant frequency) processing area ofthe primary auditory cortex. In this area all cortical neurons studied were excited by contralateral stimuli, and almost all of them were either excited or inhibited by ipsilateral stimuli. These are called E-E (ipsilateral and contralateral excitatory) and I-E (ipsilateral inhibitory and contralateral excitatory) neurons, respectively. The I-E neurons are directionally sensitive, while the E-E neurons are not. The E-E neurons are equally sensitive to echoes between 30 degrees contralateral and 30 degrees ipsilateral. Of the electrode penetrations orthogonal to the Doppler-shifted-CF processing area, 57 percent were characterized by either E-E or I-E neurons. Thus, there are at least two types of binaural columns: E-E columns, mainly located in a ventral part of the Doppler-shifted-CF processing area, where neurons are tuned to weak echoes; and IE columns, mainly distributed in a dorsal part, where neurons are tuned to moderate to intense echoes. Therefore, neurons tuned to weaker echoes integrate or even multiply faint signals from both ears for effective detection of a distant small target, while neurons tuned to moderate to intense echoes are suited for processing directional information and are stimulated when a bat approaches a target at short range. The Doppler-shifted-CF processing area may be considered to consist of two functional subdivisions.

Animals↗
PubMed1979

Harmonic-sensitive neurons in the auditory cortex of the mustache bat.

Human speech and animal sounds contain phonemes with prominent and meaningful harmonics. The biosonar signals of the mustache bat also contain up to four harmonics, and each consists of a long constant-frequency component followed by a short frequency-modulated component. Neurons have been found in a large cluster within auditory cortex of this bat whose responses are facilitated by combinations of two or more harmonically related tones. Moreover, the best frequencies for excitation of these neurons are closely associated with the constant-frequency components of the biosonar signals. The properties of these neurons make them well suited for identifying the signals produced by other echolocating mustache bats. They also show how meaningful components of sound are assembled by neural circuits in the central nervous system and suggest a method by which sounds with important harmonics (or formants) may be detected and recognized by the brain in other species, including humans.

Action Potentials↗
PubMed1979

Neural axis representing target range in the auditory cortex of the mustache bat.

In echolocating bats, the primary cue for determining distance to a target is the interval between an emitted orientation sound and its echo. Whereas frequency is represented by place in the bat cochlea, no anatomical location represents of primary range. Target range is coded by the time interval between grouped discharges of primary auditory neurons in response to both the emitted sound and its echo. In the frequency-modulated-signal processing area of the auditory cortex of the mustache bat (Pteronotus parnellii rubiginosus), neurons respond poorly or not at all to synthesized orientation sounds or echoes alone but respond vigorously to echoes following the emitted sound with a specific delay from targets at a specific range. These range-tuned neurons are systemically arranged along the rostrocaudal axis of the frequency-modulated-signal processing area according to the delays to which they best respond, and thus represent target range in terms of cortical organization. The frequency-modulated-signal processing area therefore shows odotopic representation.

Animals↗
PubMed1979

Ultrastructure of neurons in the auditory cortex of ageing rats: a morphometric study.

The cell bodies of pyramidal cells in layers II and V of rat auditory cortex were quantitatively examined in groups of rats 3, 6, 15, 23, 27 and 34-36 months of age. The mean diameters of cell bodies of both layer II and layer V neurons, as measured in 1 micron plastic sections, increased between 3 and 15 months of age, then decreased to a diameter that was less in the 36-month-old than in the 3-month-old rats. Morphometry of the nuclei of the cells was done by measuring nuclear area and nuclear envelope length directly on electron micrographs. In the layer II cells, neither parameter changed with advancing age. In the layer V cells, the mean nuclear area decreased significantly in the old animals and the mean envelope length increased. Point-counting techniques were applied to electron micrographs of cell bodies to determine the relative volume of selected organelles, inclusions and ground substance in the perikaryal cytoplasm. In this part of the study the chronological pattern of change in layer II and layer V pyramidal cells was similar. The relative volume of dense bodies increased linearly with advancing age, with a slightly more accelerated rate in layer II cells. The relative volume of ground substance remained essentially constant through 27 months, and then at 34-36 months decreased to 83% and 89% of the three-month level in layer V and layer II, respectively. The relative volume of the rough endoplasmic reticulum (RER) did not change significantly until after 15 months, at which time it began to occupy increasingly a larger fraction of the perikaryal cytoplasm. Finally, the relative volumes of mitochondria, multivesicular bodies and Golgi apparatus did not show clear trends of change during the 33-month period.

Aging↗
PubMed1979

Tangential orientation and spatial order in dendrites of cat auditory cortex: a computer microscope study of Golgi-impregnated material.

In the tangential plane (parallel to the pial surface) dendrites in the primary auditory cortex (A1) of cat were found to exhibit preferentially oriented growth. This was shown by means of a computer microscope study of Golgi-Cox stained neurons as seen in 100 micrometers and 300 micrometers thick tangential sections. Two techniques were used to represent the 3-dimensional structure of dendrites: the "dendritic stick" and the "dendritic trumpet". The former dismembers a dendrite into its individual segments; the latter considers a dendrite as an entity and represents it by its centroid, its moments and the spatial dispersionof its branches. Both statistical and Fourier analyses of the data show that within the tangential plane there is a significant and consistent orientation of the dendritic sticks in a dorso-ventral direction which seems correlated with the cortical isofrequency contours observed in electrophysiological maps of the A1 region. The dendritic trumpet analyses also show a distinctly non-random vertical distribution of pyramidal cell basal dendrites but not of stellate cell dendrites.

Animals↗
PubMed1979

Detection versus discrimination of brief tones by cats with auditory cortex lesions.

In recent years, a number of investigators have provided evidence that the auditory cortex has a critical role in both the detection and discrimination of brief sounds. Dogs and humans with lesions of the neocortical auditory centers have been reported to exhibit significantly elevated detection thresholds for signals shorter than 16 ms in duration. In tests of frequency discrimination, the same subjects also exhibited severe deficits whenever tonal signals were less than 20--40 mn in lengths. In the present report, we present evidence brief tones. Operated cats, while exhibiting normal difference limens for 1-kHz tones of 100-ms duration, have significantly elevated limens for discriminating tones of 8- and 2-ms duration. With further testing, the same operated cats can be shown to have normal absolute thresholds for detecting brief tones.

Animals↗
PubMedDate not supplied

[Visual evoked responses on the primary auditory cortex in the cat after an early suppression of cochlear receptors (author's transl)].

In the adult cat, after an early degeneration (hereditary deaf white cats) or surgical removal of auditory receptors, it is possible to record visual evoked responses on the primary auditory cortex. The appearance of such potentials depends on two conditions: a complete (bilateral) and early (first postnatal month) deafferentation. This can be explained by an early multi-afferentation of the primary cortical areas. The early suppression of the primary modality may keep the other modalities still working in the adult.

Afferent Pathways↗
PubMedDate not supplied

Comparison of electrically and acoustically evoked responses in the auditory cortex of the guinea pig: implications for a cochlear prosthesis.

Guinea pigs have been used to develop an animal model for evaluation of electric stimulation of the cochlea. Techniques were developed for recording from single neural units in the auditory cortex. Acoustic responses exhibit similar patterns of neural discharge ("on", "off", sustained excitation and inhibition, etc) to those reported for auditory neurons in other species. Similar response patterns are obtained when electric stimulation is used. However, cortical units display characteristic frequency tuning with acoustic stimulation, whereas it becomes increasingly difficult to evoke electric responses as stimulus frequency increases. Recording of gross evoked responses from the same cortical area indicates that the intensity function for electric stimulation is much steeper than that seen for acoustic stimulation.

Acoustic Stimulation↗