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N Suga

Publications and source records attributed to N Suga.

At least 91 records · Page 5Linked to original sources

Disproportionate tonotopic representation for processing CF-FM sonar signals in the mustache bat auditory cortex.

The extent of cortical representation of the peripheral sensory field depends on its importance for species behavior. The orientation sound of the mustache bat (Pteronotus parnellii rubiginosus) invariably consists of long constant-frequency and short frequency-modulated components and is indispensable for its survival. A disproportionately large part of the auditory cortex of this bat is occupied by neurons processing the predominant components in the orientation signal and Doppler-shifted echoes. This disproportionate cortical representation related to features of biologically significant signals is comparable to that in the somatosensory and visual systems in many mammals, but it has not previously been observed in the auditory system.

Action Potentials↗

Coordinated activities of middle-ear and laryngeal muscles in echolocating bats.

The middle-ear muscles and laryngeal muscles of the little brown bat (Myotis lucifugus) are highly developed. When the bat emits orientation sounds, action potentials of middle-ear muscles appear approximately 3 milliseconds after those of the laryngeal muscles; this activity of middle-ear muscles attenuates the vocal self-stimulation and improves the performance of the echolocation system. When an acoustic stimulus is delivered, both types of muscles contract; action potentials of the laryngeal muscles appear approximately 3 milliseconds after those of the middle-ear muscles. These two groups of muscles are apparently activated in a coordinated manner not only by the nerve impulses from the vocalization center, but also by those from the auditory system.

Action Potentials↗

Peripheral control of acoustic signals in the auditory system of echolocating bats.

Many species of echolocating bats emit intense orientation sounds. If such intense sounds directly stimulated their ears, detection of faint echoes would be impaired. Therefore, possible mechanisms for the attenuation of self-stimulation were studied with Myotis lucifugus. The acoustic middle-ear-muscle reflex could perfectly and transiently regulate the amplitude of an incoming signal only at its beginning. However, its shortest latency in terms of electromyograms and of the attenuation of the cochlear microphonic was 3-4 and 4-8 msec, respectively, so that these muscles failed to attenuate orientation signals by the reflex. The muscles, however, received a message from the vocalization system when the bat vocalized, and contracted synchronously with vocalization. The duration of the contraction-relaxation was so short that the self-stimulation was attenuated, but the echoes were not. The tetanus-fusion frequency of tha stapedium muscle ranged between 260 and 320/sec. Unlike the efferent fibres in the lateral-line and vestibular systems, the olivo-cochlear bundle showed no sign of attenuation of self-stimulation.

Acoustic Stimulation↗

Peripheral specialization for fine analysis of doppler-shifted echoes in the auditory system of the "CF-FM" bat Pteronotus parnellii.

Pteronotus parnellii uses the second harmonic (61-62 kHz) of the CF component in its orientation sounds for Doppler-shift compensation. The bat's inner ear is mechanically specialized for fine analysis of sounds at about 61-62 kHz. Because of this specialization, cochlear microphonics (CM) evoked by 61-62 kHz tone bursts exhibit prominent transients, slow increase and decrease in amplitude at the onset and cessation of these stimuli. CM-responses to 60-61 kHz tone bursts show a prominent input-output non-linearity and transients. Accordingly, a summated response of primary auditory neurones (N1) appears not only at the onset of the stimuli, but also at the cessation. N1-off is sharply tuned at 60-61 kHz, while N1-on is tuned at 63-64 kHz, which is 2 kHz higher than the best frequency of the auditory system because of the envelope-distortion originating from sharp mechanical tuning. Single peripheral neurones sensitive to 61-62 kHz sounds have an unusually sharp tuning curve and show phase-locked responses to beats of up to 3 kHz. Information about the frequencies of Doppler-shifted echoes is thus coded by a set of sharply tuned neurones and also discharges phase-locked to beats. Neurones with a best frequency between 55 and 64 kHz show not only tonic on-responses but also off-responses which are apparently related to the mechanical off-transient occuring in the inner ear and not to a rebound from neural inhibition.

Action Potentials↗

Site of neural attenuation of responses to self-vocalized sounds in echolocating bats.

Bats of the genus Myotis emit intense orientation sounds for echolocation. If such sounds directly stimulated their ears, the detection of echoes from short distances would be impaired. In addition to the muscular mechanism in the middle ear, the bat has a neural mechanism in the brain for attenuation of responses to self-vocalized orientation and nonorientation sounds. This neural attenuating mechanism operates in the nucleus of the lateral lemniscus, reducing its activity by about 15 decibels, and it is synchronized with vocalization.

Acoustic Stimulation↗

Neural attenuation of responses to emitted sounds in echolocating rats.

Bats of the family Vespertilionidae enmit strong ultrasonic pulses for echolocation. If such sounds directly stimulate their ears, the detection of echoes from short distances would be impaired. The responses of lateral lemniscal neurons to emitted sounds were found to be much smaller than those to playback sounds, even when the response of the auditory nerve was the same to both types of sounds. Thus, responses to self-vocalized sounds were attenuated between the cochlear nerve and the inferior colliculus. The mean attenuation was 25 decibels. This neural attenuating mechanism is probably a part of the mechanisms for effective echo detection.

Animals↗

Properties of 'two-tone inhibition' in primary auditory neurones.

1. Properties of two-tone inhibition in primary auditory neurones of cats were studied with phase-locked sound stimuli. One sound was a continuous tone at the best frequency of a given neurone, and the other, a tone burst which was changed in amplitude, frequency, and phase relative to the continuous tone.2. The tone burst which caused two-tone inhibition had either an excitatory or no effect when it was delivered alone. Inhibitory areas commonly appeared on both sides of the excitatory area when the best frequency was higher than a few kc/s.3. Two-tone inhibition began and ceased within a few milliseconds of the onset and termination of the excitation caused by a tone burst. The degree of inhibition was greatest at the beginning of the tone burst and reached a plateau within 500 msec. The discharge rate during inhibition could be lower than the rate for either tone alone or for spontaneous activity. At the termination of inhibition, prominent rebound in the discharge rate was found.4. With an increase in amplitude of a tone burst, for either a fixed or equally increased continuous tone, the discharge rate during inhibition decreased to a minimum and then began to increase. That is, the degree of inhibition was non-monotonically related to the sound level.5. Compound period histograms of discharges during inhibition showed that single neurones usually carried information about the combined wave form of the two tones. The information about each tone was, however, modified by the inhibitory phenomenon in both amplitude and phase from that indicated by the compound period histograms for the individual tones.6. Possible mechanisms and functional significance of two-tone inhibition are discussed.

Animals↗

Responses of inferior collicular neurones of bats to tone bursts with different rise times.

1. A study was made of the responses of single neurones in the inferior colliculus of bats to tone bursts with different rise times (or with different rates of amplitude increase) in order to determine whether the neurones were specialized for analysis of amplitude-modulated sound, especially the rising phase in amplitude.2. The response patterns of neurones which showed phasic on-responses usually did not change with rise time, although the response patterns of some neurones changed from phasic on-responses to inhibitory responses.3. The thresholds of responses to tone bursts increased when the rise time was lengthened. The amount of increase greatly differed from neurone to neurone. For the excitation of neurones which showed a large increase in threshold, the stimulus amplitude should quickly increase in amplitude. For tone bursts with a short rise time, some neurones showed an upper-threshold above which the sounds failed to excite them. The upper-threshold usually disappeared when the rise time was lengthened. For the excitation of neurones which showed an upper-threshold, the rate of amplitude increase and its extent were very important parameters.4. Lengthening the rise time of a tone burst from 0.5 to 98 msec caused various types of change in the excitatory area. In some cases, there was a diminution of the area and, in others, there was an expansion of the area. Neurones showing these changes may be considered to be specialized for responding to tone bursts with either a rapid or slow increase in amplitude.5. Changes with rise time occurred not only in the excitatory area, but also in the inhibitory area. The change in the inhibitory area of a given neurone, however, was not necessarily the same as that in the excitatory area.6. The latency of response usually changed as a function of stimulus amplitude and rise time. In 26% of the neurones studied, however, the latency was relatively constant regardless of stimulus amplitude and rise time. Some of the latency-constant neurones appeared to have properties suited for echo-ranging.7. The changes in excitatory area with rise time appear to involve neural inhibition and/or accommodation. Possible neural mechanisms for the changes in threshold, upper-threshold and response pattern and the properties of latency-constant neurones are discussed.

Animals↗

Echo-ranging neurons in the inferior colliculus of bats.

Bats measure the distance to an object in terms of the time lag between their outgoing orientation sounds and the returning echo. For measurement of the time lag, the latency of response of a neuron to a stimulus must be nearly constant regardless of the stimulus amplitude and envelope. Otherwise, a large error would be introduced into the measurement. Bats have neurons that are specialized for echo ranging.

Animals↗

Classification of inferior collicular neurones of bats in terms of responses to pure tones, FM sounds and noise bursts.

1. Single unit activity in the inferior colliculus of bats was studied with pure tones, FM (frequency-modulated) sounds and noise bursts which are the most basic three components of the complex sounds produced by many different animals including man. Neurones were divided into three groups, (i) ;generalized' units responding to all three elements, (ii) ;deaf' units responding to two out of the three and (iii) ;specialized' units responding to only one of the three. Each of these was divided into three subgroups.2. Three subgroups of generalized units were called ;symmetrical', ;asymmetrical' and ;upper-threshold' units. The symmetrical unit had a wide excitatory area and responded to any sounds which had components falling in this area. The asymmetrical unit had a narrow excitatory area abutting on a large inhibitory one and showed different responses to FM sounds depending on the directions of frequency sweep. The upper-threshold unit had not only an excitatory area, but also an inhibitory one. The neurone failed to respond to strong sounds, i.e. these showed upper thresholds.3. Three subgroups of deaf units were called ;pure tone-deaf', ;FM-deaf' and ;noise-deaf' units. The pure tone-deaf unit did not respond to any pure tones but did respond to FM sounds and noise bursts. The noise-deaf unit did not respond to noise bursts but did respond to pure and FM tone pulses. An FM-deaf unit has not yet been confirmed.4. Three subgroups of specialized units were called ;pure tone-specialized', ;FM-specialized' and ;noise-specialized' units which responded exclusively to either pure tones, FM sounds or noise bursts, respectively. In the FM- and noise-specialized units, pure tone pulses caused only inhibitory processes.5. About 95% of neurones studied showed phasic on-responses to sound stimuli and almost no spontaneous discharges. Only a small percent of neurones showed spontaneous discharges higher than a few impulses per second. Response patterns of some of these neurones changed with frequency, intensity, duration and repetition rate of sound stimuli. These neurones responded to any of pure tones, FM sounds and noise bursts.6. All neurones except the symmetrical ones had inhibitory areas, in which sounds inhibited responses to excitatory ones when these are delivered simultaneously. Therefore the structure in complex sounds is very important in the excitation of these neurones.

Animals↗

Echo-location and evoked potentials of bats after ablation of inferior colliculus.

1. Echo-location and evoked potentials of blinded Yuma bats (Myotis yumanensis) were studied before and after ablation of the inferior colliculus (I.C.). A task of obstacle-avoidance was given to the bats: hits and misses of strands in the flight path were counted. Orientation sounds emitted by the bats during flight were recorded.2. Bilateral ablation of the dorso-medial region of I.C. including the internuclear cortex and commissure had no effect on obstacle-avoidance performance. The bats avoided even strands of 0.2 mm diameter with orientation sounds.3. Bilateral ablation of the dorsal half of I.C. including the external nucleus (lateral cortex) also had no effect on echo-location.4. Bilateral ablation of the ventral half of I.C. caused severe deficiency in ability to avoid obstacles. The main nucleus appeared to be very important for echo-location. When bilateral ablation including the main nucleus was moderate, the bats failed to avoid strands of less than 0.5 mm diameter in spite of detecting them, but avoided large obstacles such as 3.7 mm strands. With severe bilateral ablation including the main nucleus, the bats did not avoid even the 3.7 mm strands in spite of frequent emission of orientation sounds, but often avoided crashing into the wall.5. Severe unilateral ablation of I.C. including the main nucleus and a part of the lateral lemniscus had no effect on ability to avoid obstacles. Since sound localization by such bats are not explained by Van Bergeijk's model based on Békésy's, a modification of Van Bergeijk's model has to be considered.6. Of the positive evoked potentials recorded with an active electrode placed at the dorsal surface of I.C., the slow component with a 7-9 msec peak latency reflected activity of inferior collicular neurones, while the fast component (N(4)) with a 3 msec peak latency represented activity of ascending lateral lemniscal fibres.

Animals↗

Echo-location of bats after ablation of auditory cortex.

1. Echo-location of blinded Yuma bats (Myotis yumanensis) was studied after ablation of the auditory cortex (A.C.). A task of obstacle-avoidance was given to the bats. Hits and misses of strands were counted, and orientation sounds emitted by the bats during flight were recorded.2. After bilateral ablation of A.C., two bats out of six failed to avoid even large obstacles such as 3.7 mm strands and wall. These bats emitted orientation sounds at a rate of 10-15/sec during flight, but did not change that rate before crossing the obstacles and crashed into them. In these bats, other cortical areas in addition to A.C. were probably ablated.3. In three bats out of six, obstacle-avoidance performance was quite normal. These bats avoided even 0.2 mm strands with orientation sounds, the repetition rate of which was systematically increased before crossing the obstacles. In two of them, the dorsal half of the inferior colliculus (I.C.) was bilaterally ablated in addition to A.C. But ability to avoid the obstacles was not impaired at all. Their cerebral cortices did not show the normal positive-negative diphasic potential change in response to tonal stimuli, although the normal diphasic potential change was retained in A.C. of bats which could not echo-locate as a result of bilateral ablation of the main nucleus of I.C. A.C. appeared to be not essential for echo-location.4. Unilateral ablation of A.C. and the internal capsule had no effect on echo-location, but bilateral ablation of them usually resulted in death from operational trauma.5. It was suggested that A.C. was less important for sound localization in bats than in cats.

Journal Article↗

Analysis of frequency-modulated and complex sounds by single auditory neurones of bats.

1. Single unit activity in the inferior colliculus of bats was studied in relation to the analysis of frequency-modulated (FM) and complex sounds. Complex sounds were composed of tone pulse I (pure or FM tone) delivered simultaneously with tone pulse II (pure or sometimes FM tone). It was assumed that in relevant complex sounds produced by animals, an important component (e.g. a formant in human speech) occurred at the best frequency (BF) of a given neurone. Tone pulse I represented such a component (called BF component). Tone pulse II was assumed to correspond to higher or lower components according to its relation to BF. Depending on characteristics of responses to tonal stimuli, collicular neurones were classified into five types: symmetrical, asymmetrical, FM-insensitive, FM-sensitive (or FM-specialized) and upper-threshold units.2. The symmetrical unit had a wide excitatory area and no inhibitory areas and it responded with equal thresholds to FM tone pulses sweeping in either directions. This type of neurone responded to all frequency modulations (e.g. transition in human speech) of the BF component, and the response was scarcely inhibited by other components.3. In the asymmetrical unit, the extent of frequency modulation of the BF component which could excite the neurone was limited by inhibitory areas on one or both sides of an excitatory area. Inhibitory areas on the lower frequency side tended to be larger than those on the high frequency side. The limitation was more severe for frequency sweeps toward the best frequency than for sweeps starting from it. The response to the BF component was inhibited by lower and/or higher components unless these were outside the inhibitory areas. In most of the asymmetrical units, lower components were more important than higher ones in determining whether the response to the BF component could occur.4. In the FM-insensitive unit with a narrow excitatory area, inhibitory areas on both sides of the excitatory area restricted the extent of frequency modulation of the BF components which could activate the neurone. Responses to frequency sweeps toward the best frequency were strongly limited by the inhibitory areas. When the lower and/or higher components were within the inhibitory areas, the response to the BF component was inhibited. This type of neurone responded to more restricted combinations of components than did the asymmetrical units.5. The FM-sensitive unit which had no excitatory area but a large inhibitory area responded only to FM components in a certain range. Evidence was obtained that not only the range and direction but rate and functional form of frequency sweep were important in determining the excitation of the neurone. Noise bursts with various band widths did not activate the neurones. Responses of the neurones were commonly inhibited by tones within an inhibitory area so wide as to involve even frequencies in the FM component which excited the neurone. Thus, the response of the FM-sensitive unit depended not only upon the characteristics of the FM component, but also on the frequencies of other components.6. The asymmetrical, FM-insensitive and FM-sensitive units required for their activation a certain structure in the complex sound. Some of the upper-threshold units did not respond to a sufficiently strong BF component and/or its frequency modulation. Furthermore, the response of the neurone to a weak BF component was inhibited by strong lower and/or higher components in a certain range. Some upper-threshold units also had asymmetrical or FM-sensitive characteristics. Those neurones appeared to be specialized for the analysis of sound structure not only in frequency, but in intensity.7. Although various types of behaviour of single neurones were found in the inferior colliculus, a strong tendency in the neural analysis of complex sound was the restriction of conditions under which single neurones were activated. Neurones at higher levels responded to more restricted sequences or sets of sound stimuli than did those at lower levels.

Action Potentials↗