Responses of neurons in the rat auditory and associative cortices to tonal stimuli

Activity of single neurons and mass evoked potentials (EP) were recorded from the auditory (area 41) and associative (area 39) cortices in acute experiments on rats anesthetized with urethane, nembutal, or chloralose; pure tones were used as acoustic stimuli. The EP appearing in response to a wide range of sound tones on the surface of the auditory and associative cortices were dissimilar in their latency and shape. For neurons exhibiting stable responses, the frequency-threshold curves (FTC) were plotted.Weak and variable responses of neurons were observed under slight urethane anesthesia. Nembutal anesthesia increased the responsiveness of neurons and the probability of appearing of late components in the responses. Chloralose anesthesia was characterized by extension of frequency range perceived by a neuron, while its sharpness of tuning remained unchanged. Under all types of anesthesia employed, the responses recorded from the associative cortex neurons had longer latencies than those recorded from the auditory cortex neurons. Neurons exhibiting the frequency selectivity were much less numerous in the associative cortex than in the auditory cortex. The former neurons were often characterized by intermittent FTC and they responded to a more extended frequency range. No clear tonotopic organization was found in the associative cortex.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 343–349, September–October, 1993.     

Response of parietal association cortex neurons to acoustic stimulation before and after auditory cortex blockage in the cat

The effect of auditory cortex blockade on response patterns of parietal association cortex neurons responding to different frequency tones was investigated in the cat. Blockade was produced by two methods: bilateral isolation and application of a 6% Nembutal solution to the auditory cortex surface. Frequency threshold curves were plotted for all test neurons. The majority of test neurons (84%) displayed one or two characteristic frequencies before blockade, as against only 63% of all neurons responding following blockade. Changes also affect the range of frequencies at which the cells could respond. Virtually all test neurons responded to application of a broad spectrum of frequencies under normal conditions. After blockade of the auditory cortex 69% of neurons no longer responded to tones above 8–10 kHz. This would suggest that mainly information on high frequency tones is transmitted via the auditory cortex. The question of where acoustic information for parietal association cortex neurons mostly originates is also discussed; association thalamic nuclei are thought to be the main source.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 354–360, May–June, 1986.     

Interaction between impulses evoked by stimuli of different modalities in neurons of the parietal associative cortex

In acute experiments on cats anesthetized with chloralose and nembutal interaction between visual, auditory, and electrodermal stimuli in neurons of the parietal association cortex was studied. Two types of interaction were found; the first characterized by inhibition or by inhibition followed by facilitation of the response to the test stimulus, the second by facilitation or by facilitation followed by inhibition of spontaneous impulses. Interaction between stimuli of different modalities was shown to depend on the properties of the neuron. In polysensory neurons ability to interact was much higher than in bimodal or monomodal neurons.M. Gorkii Donetsk Medical Institute. Kemerovo Medical Institute. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 223–229, May–June, 1976.     

Retrosplenial and hippocampal projections of structures of the thalamoparietal associative system in rats

Afferent connections of the nucleus lateralis posterior (NLP) of the thalamus and area 7 of the parietal cortex with the retrosplenial region of the limbic cortex and hippocampus were studied in rats with retrograde axon transport of horseradish peroxidase. It was shown that the NLP receives ipsilateral projections from area 29d neurons, while area 7 receives ipsilateral axons from area 29d and 29c neurons. It was found that associations of the retrosplenial region with associative cortex are far more pronounced than with associative thalamus. Moreover, the afferent connections of area 7 with area 29d are more numerous than with area 29c. We disclosed no projections of areas 29a and 29b to thalamoparietal system structures. In addition to neocortical input from the limbic cortex, area 7 receives afferent fibers from the archicortex; neurons situated in hippocampus area CA1 are the source of these projections.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Russian Academy Academy of Sciences, Leningrad. Translated from Neirofiziologiya, Vol. 23, No. 6, pp. 647–655, November–December, 1991.     

Comparison of spontaneous cortical unit activity in several brain areas of unanesthetized cats

Spontaneous unit activity in association area 5 and some projection areas of the cortex (first somatosensory, first and second auditory areas) were studied in cats immobilized with D-tubocurarine in which the index of specific spontaneous activity, the mean frequency, types of spontaneous activity, and statistical parameters — distribution of interspike intervals and autocorrelation function — were determined. The results showed that spontaneous unit activity in the association area differs from that in the projection areas in both intensity and character. A special feature of the spontaneous activity of the auditory areas was a well-marked volley distribution of activity. In the somatosensory area the level of spontaneous activity as reflected in all indices was the lowest. In the association cortex the largest number of neurons with spontaneous activity lay at a depth of 500–1000 µ corresponding to cortical layers III–IV. In the first auditory area neurons with spontaneous activity were concentrated at a depth of 1400 µ (layer V) and in the somatosensory area at a depth of 1000–1400 µ (alyers IV–V). The possible functional significance of these differences is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 13–21, January–February, 1976.     

Tonotopic organization of the dorsocaudal zone of cortical area aii in the cat

The tonotopic organization of the dorsocaudal (DC) auditory cortex area AII was investigated during acute experiments on cats anesthetized with Nembutal. A capacity for selective response to presentation of auditory stimuli at a certain frequency was found in 93% of the neurons investigated. It was further observed that 75% of these cells were characterized by their fine tuning to one characteristic frequency (CF), the remaining 26% had several CF, and 7% reacted with a spike response to acoustic stimulation at all test frequencies and had no clearcut CF. A relationship was found between the location of a unit within the DC zone and its CF level. Neurons with the lowest CF were located in the upper position of the sylvian gyrus near the posterior ectosylvian sulcus. The CF of neurons rose progressively in step with increasing distance between the site of microelectrode recording and the low frequency focus of the DC zone travelling along the sylvian gyrus in a ventrorostral direction. Distance between low and high frequency foci of the DC zone measured 2.5–3.5 mm. Location of this zone in relation to the auditory cortex sulci varied considerably from one animal to another. Neurons with similar CF levels and arranged on this basis in vertical cortical columns could vary substantially in the dimensions of their receptive fields, sharpness of tunining to their own CF, and firing response pattern.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 2, pp. 220–227, March–April, 1988.     

Auditory Cortical Areas Activated by Slow Frequency-Modulated Sounds in Mice

Species-specific vocalizations in mice have frequency-modulated (FM) components slower than the lower limit of FM direction selectivity in the core region of the mouse auditory cortex. To identify cortical areas selective to slow frequency modulation, we investigated tonal responses in the mouse auditory cortex using transcranial flavoprotein fluorescence imaging. For differentiating responses to frequency modulation from those to stimuli at constant frequencies, we focused on transient fluorescence changes after direction reversal of temporally repeated and superimposed FM sweeps. We found that the ultrasonic field (UF) in the belt cortical region selectively responded to the direction reversal. The dorsoposterior field (DP) also responded weakly to the reversal. Regarding the responses in UF, no apparent tonotopic map was found, and the right UF responses were significantly larger in amplitude than the left UF responses. The half-max latency in responses to FM sweeps was shorter in UF compared with that in the primary auditory cortex (A1) or anterior auditory field (AAF). Tracer injection experiments in the functionally identified UF and DP confirmed that these two areas receive afferent inputs from the dorsal part of the medial geniculate nucleus (MG). Calcium imaging of UF neurons stained with fura-2 were performed using a two-photon microscope, and the presence of UF neurons that were selective to both direction and direction reversal of slow frequency modulation was demonstrated. These results strongly suggest a role for UF, and possibly DP, as cortical areas specialized for processing slow frequency modulation in mice.     

Microelectrode analysis of neuronal organization of the parietal association cortex

Responses of 304 neurons in the anterior part of the middle suprasylvian gyrus in cats to acoustic, photic, and somatosensory stimulation and also to simultaneous presentation of 2 or 3 stimuli of these modalities. Three groups of neurons were distinguished: those responding by an increased firing rate (187) or by inhibition (22) and those not responding (95). The first group comprised mono- (64) and polysensory (105) and well as neurons responding only to a combination (18). On the basis of the convergent properties of the polysensory neurons in this region it is postulated that the parietal cortex performs principally integrative processes based on iteraction of visual-cutaneous and cutaneo-auditory afferent information.State Medical Institute, Kemerovo. Translated from Neirofiziologiya, Vol. 4 No. 1, pp. 54–60, January–February, 1972.     

Electric reactions of the auditory cortex areas in bats to ultrasonic stimuli with various fill frequencies

The overall electric reactions and action potentials of single neurons in the auditory cortex were investigated for Vespertilionidae (Myotis oxygnathus) and Rhinolophidae (Rhinolophus ferrum equinum) narcotized with Hexenal. In the Vespertilionidae the greatest sensitivity to ultrasound is manifest at frequencies from 10 to 50 kHz, and in the Rhinolophidae for the ranges from 10 to 40 and from 82 to 84 kHz. The shapes of the response areas of single neurons in both types of bats are similar except for neurons discovered in Rhinolophidae that have three response areas with characteristic frequencies in the ranges 27–28, 40–42, and 80–84kHz. Narrow response areas with characteristic frequencies in the range from 70 to 90kHz appear on a considerable proportion of the neurons in the Rhinolophidae, but not the Vespertilionidae. Low thresholds are recorded to the stimulus cutoff in the range from 76 to 86 kHz.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 3, No. 5, pp. 526–532, September–October, 1971.     

Morphological characteristics of the neuronal population in the feline auditory cortex projecting into the medial geniculate body

The location and morphological profile of auditory cortex neurons projecting to the medial geniculate body were investigated in adult cats using horseradish peroxidase retrograde axonal transport techniques. Sources of descending projections to the medial geniculate body from auditory cortex areas I and II were found to be neurons belonging to deep-lying layers (layer VI and layer V to a lesser extent). By far the majority of corticogeniculate neurons in the auditory cortex were pyramidal cells. In layer VI of the primary auditory area (A1), the number of corticogeniculate neurons reaches 60% of all cells belonging to that layer. The average area (M±m) of the profile of perikarya of corticogeniculate neurons in layer VI, area Al equaled 139.3±2.5 µm² and 219.5±7.0 µm² in layer V neurons; average size of long diameter: 15.0±0.19 and 18.3±0.4 µm respectively. The lower regions of layers III and IV in area Al were found to be the termination point of the greater mass of anterogradely-labeled geniculocortical fibers (terminals of relay neuron axons belonging to the medial geniculate body).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 4, July–August, pp. 513–521, 1989.     

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.     

The role of inhibition in shaping the type of single neuron responses in the auditory system of the frog

The responses of single neurons of the auditory center in the frog mesencephalon to tonal stimuli of varying frequencies have been studied. It has been found that some neurons which respond to the signal of the characteristic frequency (CF) by a long-lasting discharge respond to tones of higher frequencies only at the start of stimulation. It is shown, that the tones giving rise to a phasic response inhibit impulsation brought about by the action of the CF tone.Acoustics Institute, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 236–241, May–June, 1970.     

Tonotopic organization of the ventrorostral zone of the auditory cortex region AII in cats

The tonotopic organization of the ventrorostral (VR) zone of cortical auditory area AII was investigated in acute experiments on cats anesthetized with nembutal and unanesthetized immobilized animals. Response with the lowest threshold arose in 92% of test neurons to presentation of one or several sound frequencies. The majority (54%) were "tuned" to one characteristic frequency (CF), 38% to several frequencies, and 8% had no clear-cut CF. A connection was found between location of a unit within the VR zone and its CF. Neurons with the highest CF were located in the ventrocaudal AII. An increase was noted in numbers of neurons with the lowest CF with increasing distance (rost-rally) from the VR location zone of neurons tuned to a high frequency. Going by response to acoustic stimuli of frequencies ranging between 1 and 24 kHz, length of the VR projection zone of the AII was found to measure 1.8–2.0 mm. Location of the test zone in relation to auditory cortex sulci varied substantially from one animal to the next.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 178–184, March–April, 1990.     

Cholinergic structures in the cat auditory cortex (area AI)

Plexuses of cholinergic varicose fibers, differing in density in different layers of the neuropil, were found in area AI of the cat's auditory cortex by the histochemical reaction for acetylcholinesterase: Their density was maximal or average in layer I or deeper layers and minimal in layers II and III. Among cells in area AI those which are cholinergic are a few stellate neurons located in layers II–VI. Axons of some neurons terminate on neighboring cells, those of others (some neurons in layer VI) run into the subcortical layer of arcuate association fibers. Cholinergic terminals are located on the bodies and proximal areas of dendrites of neurons most of which do not contain acetylcholinesterase. Choliniceptive neurons of different sizes and shapes are found in all layers of this region of the auditory cortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 75–81, January–February, 1984.     

Enhancement and Distortion in the Temporal Representation of Sounds in the Ventral Cochlear Nucleus of Chinchillas and Cats

A subset of neurons in the cochlear nucleus (CN) of the auditory brainstem has the ability to enhance the auditory nerve''s temporal representation of stimulating sounds. These neurons reside in the ventral region of the CN (VCN) and are usually known as highly synchronized, or high-sync, neurons. Most published reports about the existence and properties of high-sync neurons are based on recordings performed on a VCN output tract—not the VCN itself—of cats. In other species, comprehensive studies detailing the properties of high-sync neurons, or even acknowledging their existence, are missing.Examination of the responses of a population of VCN neurons in chinchillas revealed that a subset of those neurons have temporal properties similar to high-sync neurons in the cat. Phase locking and entrainment—the ability of a neuron to fire action potentials at a certain stimulus phase and at almost every stimulus period, respectively—have similar maximum values in cats and chinchillas. Ranges of characteristic frequencies for high-sync neurons in chinchillas and cats extend up to 600 and 1000 Hz, respectively. Enhancement of temporal processing relative to auditory nerve fibers (ANFs), which has been shown previously in cats using tonal and white-noise stimuli, is also demonstrated here in the responses of VCN neurons to synthetic and spoken vowel sounds.Along with the large amount of phase locking displayed by some VCN neurons there occurs a deterioration in the spectral representation of the stimuli (tones or vowels). High-sync neurons exhibit a greater distortion in their responses to tones or vowels than do other types of VCN neurons and auditory nerve fibers.Standard deviations of first-spike latency measured in responses of high-sync neurons are lower than similar values measured in ANFs'' responses. This might indicate a role of high-sync neurons in other tasks beyond sound localization.     

Associative connections of the cat parietal cortex

Interneuronal connections of area 7 of the cat parietal cortex with projection areas of the visual, auditory, and somatosensory cortex were analyzed by orthograde degeneration and retrograde transport of horseradish peroxidase methods. By combined investigation the cortico-cortical sources of afferentation of parietal area 7 could be precisely identified and concentration sites of neurons sending their axons into this area identified, and the morphological characteristics of these neurons could also be determined.A. A. Ukhtomskii Physiological Institute, A. A. Zhdanov Leningrad State University. Donetsk Medical Institute. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 13–17, January–February, 1980.     

Responses of parietal associative neurons to stimulation of primary sensory areas

Responses of 251 neurons in the anterior part of the middle suprasylvian gyrus to stimulation of primary sensory (auditory, visual, somatosensory) areas and also to acoustic, visual, and somatosensory stimuli were studied in acute experiments on cats anesthetized with chloralose (40 mg/kg) and pentobarbital (20 mg/kg). Three groups of neurons were distinguished by their responses to stimulation of the primary sensory areas: those responding by an increased firing rate (117) or by inhibition (35) and those not responding (99). Responses of 193 neurons to stimulation of the peripheral afferent systems were analyzed. Neurons of the parietal associative cortex responded more frequently to cortical stimulation than to peripheral. By the duration of the latent period of their response to cortical stimulation the neurons were divided into three groups: those with short (less than 20 msec), medium (20–30 msec), and long latent periods (over 30 msec). The first group was the largest.Kemerovo State Medical Institute. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 524–530, September–October, 1972.     

Responses of secondary sensomotor cortical neurons to electrodermal and acoustic stimulation in waking cats

Unit responses in the second somatosensory cortical projection area (SII) to clicks and electric shocks applied to the contralateral limb were investigated in chronic experiments on cats. In response to specific stimulation for the cortical region studied the discharge frequency of 75% of neurons increased, spontaneous activity of 18% was reduced in frequency or the discharges ceased altogether, and 25% of cells did not respond. In response to "nonspecific" stimulation (clicks) 30% of neurons were activated; the discharge of 25% of cells was inhibited and 45% did not respond. The results of investigation of intersensory convergence of stimuli from different sensory systems showed that a high proportion (55%) of SII neurons give bimodal responses. Another 18% of neurons give a specific response to both adequate and inadequate stimulation. It is suggested that the presence of polysensory convergence of SII neurons and of short pathways for the conduction of sensory information, and also the ability of neurons to acquire polysensory properties during stimulus presentation are evidence of the important role of this cortical region in conditioning.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 5, pp. 453–459, September–October, 1977.     

Neuronal and synaptic mechanisms of cortical inhibition

The latent periods, amplitude, and duration of IPSPs arising in neurons in different parts of the cat cortex in response to afferent stimuli, stimulation of thalamocortical fibers, and intracortical microstimulation are described. The duration of IPSPs evoked in cortical neurons in response to single afferent stimuli varied from 20 to 250 msec (most common frequency 30–60 msec). During intracortical microstimulation of the auditory cortex, IPSPs with a duration of 5–10 msec also appeared. Barbiturates and chloralose increased the duration of the IPSPs to 300–500 msec. The latent period of 73% of IPSPs arising in auditory cortical neurons in response to stimulation of thalamocortical fibers was 1.2 msec longer than the latent period of monosynaptic EPSPs evoked in the same way. It is concluded from these data that inhibition arising in most neurons of cortical projection areas as a result of the arrival of corresponding afferent impulsation is direct afferent inhibition involving the participation of cortical inhibitory interneurons. A mechanism of recurrent inhibition takes part in the development of inhibition in a certain proportion of neurons. IPSPs arise monosynaptically in 2% of cells. A study of responses of cortical neurons to intracortical microstimulation showed that synaptic delay of IPSPs in these cells is 0.3–0.4 msec. The length of axons of inhibitory neurons in layer IV of the auditory cortex reaches 1.5 mm. The velocity of spread of excitation along these axons is 1.6–2.8 msec (mean 2.2 msec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 394–403, May–June, 1984.     

Interaction of influences from the auditory and somatosensory afferent systems on neurons of the primary auditory cortex

In experiments on anesthetized cats, 80 neurons of the primary auditory cortex (A1) were studied. Within the examined neuronal population, 66 cells (or 82.5%) were monosensory units, i.e., they responded only to acoustic stimulations (sound clicks and tones); 8 (10.1%) neurons responded to acoustic stimulation and electrocutaneous stimulation (ECS); the rest of the units (7.4%) were either trisensory (responded also to visual stimulation) or responded only to non-acoustic stimulations. In the A1 area, neurons responding to ECS with rather short latencies (15.6–17.0 msec) were found. ECS usually suppressed the impulse neuronal responses evoked by sound clicks. It is concluded that somatosensory afferent signals cause predominantly an inhibitory effect on transmission of an acoustic afferent volley to the auditory cortex at a subcortical level; however, rare cases of excitatory convergence of acoustic and somatosensory inputs toA1 neurons were observed.