电刺激蝙蝠中脑上丘对下丘听神经元电活动的影响

实验在24只鲁氏菊头蝠(Rhinolophus rouxi)上进行.使用玻璃微电极在中脑下丘中央核记录听神经元电反应.刺激点位于上丘核.共观察了294个对超声刺激产生反应的下丘听单位.当电刺激上丘时,有122个听单位的反应受到影响,占所观察总数的41.5%.其中96个单位表现为抑制性影响(占32.65%),26个单位表现为易代性效应(占8.84%).其余172个单位不受上丘刺激的影响(58.50%).实验中发现,上述抑制潜伏期一般在5毫秒以上,抑制时程较长.抑制程度与上丘刺激电流强度呈相关关系(r=0.99).实验中还发现,刺激上丘同样可抑制部分下丘神经元的自发放电活动,其抑制后效应相当长,有的可达120毫秒以上.     

Frequency and space representation in the primary auditory cortex of the frequency modulating batEptesicus fuscus

1. Frequency and space representation in the auditory cortex of the big brown bat, Eptesicus fuscus, were studied by recording responses of 223 neurons to acoustic stimuli presented in the bat's frontal auditory space. 2. The majority of the auditory cortical neurons were recorded at a depth of less than 500 microns with a response latency between 8 and 20 ms. They generally discharged phasically and had nonmonotonic intensity-rate functions. The minimum threshold, (MT) of these neurons was between 8 and 82 dB sound pressure level (SPL). Half of the cortical neurons showed spontaneous activity. All 55 threshold curves are V-shaped and can be described as broad, intermediate, or narrow. 3. Auditory cortical neurons are tonotopically organized along the anteroposterior axis of the auditory cortex. High-frequency-sensitive neurons are located anteriorly and low-frequency-sensitive neurons posteriorly. An overwhelming majority of neurons were sensitive to a frequency range between 30 and 75 kHz. 4. When a sound was delivered from the response center of a neuron on the bat's frontal auditory space, the neuron had its lowest MT. When the stimulus amplitude was increased above the MT, the neuron responded to sound delivered within a defined spatial area. The response center was not always at the geometric center of the spatial response area. The latter also expanded with stimulus amplitude. High-frequency-sensitive neurons tended to have smaller spatial response areas than low-frequency-sensitive neurons. 5. Response centers of all 223 neurons were located between 0 degrees and 50 degrees in azimuth, 2 degrees up and 25 degrees down in elevation of the contralateral frontal auditory space. Response centers of auditory cortical neurons tended to move toward the midline and slightly downward with increasing best frequency. 6. Auditory space representation appears to be systematically arranged according to the tonotopic axis of the auditory cortex. Thus, the lateral space is represented posteriorly and the middle space anteriorly. Space representation, however, is less systematic in the vertical direction. 7. Auditory cortical neurons are columnarly organized. Thus, the BFs, MTs, threshold curves, azimuthal location of response centers, and auditory spatial response areas of neurons sequentially isolated from an orthogonal electrode penetration are similar.     

电刺激蝙蝠小脑对中脑上丘神经元听反应的影响

实验在２３只成年中华鼠耳蝠（Ｍｙｏｌｉｓｃｈｉｎｅｎｓｉｓ）上进行。使用常规电生理学方法,观察了电刺激小脑对上丘神经元听反应的影响。在所观察的１７１个上丘神经元中,有１１６个（占６７．８４％）神经元听反应受到影响,其中７２个（占４２．１１％）表现为抑制效应,４４个（占２５．７３％）为易化效应。刺激小脑对上丘神经听反应的影响是双侧的。抑制或易化程度与电刺激强度、声刺激强度以及声、电刺激间隔有关。结果提示,小脑可以对上丘神经元听反应进行调制,这种调制作用可能是小脑调控回声定位过程中听觉－运动的中枢机制之一。     

Organization of somatic nerve projections in various cortical areas of the cat cerebellum

We conducted a layer analysis of evoked potentials arising in various cortical cerebellar areas (vermis and intermediate zones of the anterior lobe, and the ansiform lobe) of non-anesthetized cats upon stimulation of nerves in fore- and hindlimbs. This analysis yielded the conclusion on the arrival of stimuli at the cerebellar cortex over two types of moss fibers innervating two types of granule cells which we described earlier. Impulses transmitted over type I moss fibers stimulate Purkinje cells. The activation of type II moss fibers has no immediate effect on these cells. Type I moss fibers terminate in the vermis and the intermediate zone of the hemispheres and do not terminate in the lateral hemispheric region. While projections of type I moss fibers are somatotopically organized in the intermediate zone they are diffuse in the vermis. Type II moss fibers terminate in all the regions of the crebellar cortex under study, but their projections show no somatotopic organization. The question of the afferent pathways terminating as type I and II moss fibers is discussed.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 3, No. 2, pp. 166–174, March–April, 1971.     

幼小蝙蝠下丘神经元的听反应特性

实验在出生6—8天的8只幼龄鲁氏菊头蝠(Rhinolophus rouxi)上进行。使用玻璃微电极记录中脑下丘听神经元对超声信号的反应。共观察了162个听单位,它们对超声反应的最佳频率分布范围为25.8—60.9千赫,多数集中在43.0—47.0千赫。反应的潜伏期在6.0—38.0毫秒,平均为15.4±5.2毫秒。反应的最低阈值在25—84dB,平均为69.8±10.3dB.这些神经元对超声刺激的调谐曲线都较宽阔,故Q10-dB值都较小。当微电极由下丘表面垂直下插时,所记录到的神经元的最佳频率与记录深度之间不存在相关关系,即没有音调筑构现象。听神经元的这些特性与同种成年动物构成显著差异。     

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.     

Characteristics of unit responses of the cat cerebellar cortex to acoustic stimuli

Investigation of unit responses of the cerebellar cortex (lobules VI–VII of the vermis) to acoustic stimulation showed that the great majority of neurons responded by a discharge of one spike or a group of spikes with a latent period of 10–40 msec and with a low fluctuation value. Neurons identified as Purkinje cells responded to sound either by inhibition of spontaneous activity or by a "climbing fiber response" with a latent period of 40–60 msec and with a high fluctuation value. In 4 of 80 neurons a prolonged (lasting about 1 sec or more), variable response with a latent period of 225–580 msec was observed. The minimal thresholds of unit responses to acoustic stimuli were distributed within the range from –7 to 77 dB, with a mode from 20 to 50 dB. All the characteristics of the cerebellar unit responses studied were independent of the intensity, duration, and frequency of the sound, like neurons of short-latency type in the inferior colliculi. In certain properties — firing pattern, latent period, and threshold of response — the cerebellar neurons resemble neurons of higher levels of the auditory system: the medial geniculate body and auditory cortex.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 3–12, January–February, 1973.     

电刺激蝙蝠小脑对下丘神经元听反应的影响

实验在34只长翼蝠(Miniopterus schrebersi)上进行.使用玻璃微电极在中脑下丘中央核记录听神经元电反应.电刺激点分别位于小脑蚓部、半球和绒球小结叶共观察了515个对超声刺激产生反应的下丘听神经元.当电刺激小脑时,有171个(占33.2%)神经元听反应受到影响.其中126个(占24.5%)表现为抑制,45个(8.7%)表现为易化.抑制效应表现为神经元所反应放电频数降低和反应潜伏期延长.易化效应则相反.抑制与易化潜伏期一般都在6毫秒以上.抑制效应与电刺激强度、声刺激强度以及声刺激和电刺激的时间间隔有关.抑制和易化性影响都是双侧性的.     

Whole-head mapping of middle-latency auditory evoked magnetic fields

We recorded middle-latency auditory evoked magnetic fields from 9 healthy subjects with a 122-channel whole-head SQUID gradiometer. The stimuli were click triplets, 2.5 msec in total duration, delivered alternately to the two ears once every 333 msec. Contralateral clicks elicited P30m responses in 16 and P50m responses in 12 out of 18 hemispheres studied; ipsilateral clicks did so in 7 and 13 hemispheres, respectively. The field patterns were satisfactorily explained by current dipoles in 16 and 4 hemispheres for contra- and ipsilateral P30m, and in 4 and 10 hemispheres for contra- and ipsilateral P50m. The peak latencies of P30m and P50m were not affected by stimulation side. The results show that middle-latency auditory evoked responses receive a strong contribution from auditory cortical structures, and that differences of input latency to cortical auditory areas, evaluated from MLAEF latencies, do not explain the latency differences seen in late auditory evoked fields to contralateral vs. ipsilateral stimulation.     

Effects of Sound Frequency on Audiovisual Integration: An Event-Related Potential Study

A combination of signals across modalities can facilitate sensory perception. The audiovisual facilitative effect strongly depends on the features of the stimulus. Here, we investigated how sound frequency, which is one of basic features of an auditory signal, modulates audiovisual integration. In this study, the task of the participant was to respond to a visual target stimulus by pressing a key while ignoring auditory stimuli, comprising of tones of different frequencies (0.5, 1, 2.5 and 5 kHz). A significant facilitation of reaction times was obtained following audiovisual stimulation, irrespective of whether the task-irrelevant sounds were low or high frequency. Using event-related potential (ERP), audiovisual integration was found over the occipital area for 0.5 kHz auditory stimuli from 190–210 ms, for 1 kHz stimuli from 170–200 ms, for 2.5 kHz stimuli from 140–200 ms, 5 kHz stimuli from 100–200 ms. These findings suggest that a higher frequency sound signal paired with visual stimuli might be early processed or integrated despite the auditory stimuli being task-irrelevant information. Furthermore, audiovisual integration in late latency (300–340 ms) ERPs with fronto-central topography was found for auditory stimuli of lower frequencies (0.5, 1 and 2.5 kHz). Our results confirmed that audiovisual integration is affected by the frequency of an auditory stimulus. Taken together, the neurophysiological results provide unique insight into how the brain processes a multisensory visual signal and auditory stimuli of different frequencies.     

Corticofugal regulation of auditory sensitivity in the bat inferior colliculus

Under free-field stimulation conditions, corticofugal regulation of auditory sensitivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus, was studied by blocking activities of auditory cortical neurons with Lidocaine or by electrical stimulation in auditory cortical neuron recording sites. The corticocollicular pathway regulated the number of impulses, the auditory spatial response areas and the frequency-tuning curves of inferior colliculus neurons through facilitation or inhibition. Corticofugal regulation was most effective at low sound intensity and was dependent upon the time interval between acoustic and electrical stimuli. At optimal interstimulus intervals, inferior colliculus neurons had the smallest number of impulses and the longest response latency during corticofugal inhibition. The opposite effects were observed during corticofugal facilitation. Corticofugal inhibitory latency was longer than corticofugal facilitatory latency. Iontophoretic application of γ-aminobutyric acid and bicuculline to inferior colliculus recording sites produced effects similar to what were observed during corticofugal inhibition and facilitation. We suggest that corticofugal regulation of central auditory sensitivity can provide an animal with a mechanism to regulate acoustic signal processing in the ascending auditory pathway. Accepted: 15 July 1998     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

Paired associative stimulation of the auditory system: a proof-of-principle study

Background

Paired associative stimulation (PAS) consisting of repeated application of transcranial magnetic stimulation (TMS) pulses and contingent exteroceptive stimuli has been shown to induce neuroplastic effects in the motor and somatosensory system. The objective was to investigate whether the auditory system can be modulated by PAS.

Methods

Acoustic stimuli (4 kHz) were paired with TMS of the auditory cortex with intervals of either 45 ms (PAS(45 ms)) or 10 ms (PAS(10 ms)). Two-hundred paired stimuli were applied at 0.1 Hz and effects were compared with low frequency repetitive TMS (rTMS) at 0.1 Hz (200 stimuli) and 1 Hz (1000 stimuli) in eleven healthy students. Auditory cortex excitability was measured before and after the interventions by long latency auditory evoked potentials (AEPs) for the tone (4 kHz) used in the pairing, and a control tone (1 kHz) in a within subjects design.

Results

Amplitudes of the N1-P2 complex were reduced for the 4 kHz tone after both PAS(45 ms) and PAS(10 ms), but not after the 0.1 Hz and 1 Hz rTMS protocols with more pronounced effects for PAS(45 ms). Similar, but less pronounced effects were observed for the 1 kHz control tone.

Conclusion

These findings indicate that paired associative stimulation may induce tonotopically specific and also tone unspecific human auditory cortex plasticity.     

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.     

Neuronal responses of an isolated slab of cat auditory cortex to intracortical stimulation

Neuronal responses of an isolated slab of auditory cortex (area AI) to intracortical stimulation at the level IV were studied in curarized cats by extracellular recording 3 weeks after isolation. Dispersion of response latencies in the isolated slab was reduced (compared with that observed soon after isolation); the predominant responses were mono- and disynaptic, and the number of discharges consisting of bursts of spikes increased. However, despite simplification of the structural and functional organization of the chronically isolated slab of auditory cortex, the conditions for complex polysynaptic interaction between neurons of all layers were preserved in it, and in each layer the character of such interaction depended on the distance of the neuron from the focus of origin of the excitation. [In the chronically isolated slab of auditory cortex, just as in the acutely isolated slab, late reponses of over 40 msec were absent.]I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 14, No. 5, pp. 462–469, September–October, 1982.     

Bats and frogs and animals in between: evidence for a common central timing mechanism to extract periodicity pitch

Widely divergent vertebrates share a common central temporal mechanism for representing periodicities of acoustic waveform events. In the auditory nerve, periodicities corresponding to frequencies or rates from about 10 Hz to over 1,000 Hz are extracted from pure tones, from low-frequency complex sounds (e.g., 1st harmonic in bullfrog calls), from mid-frequency sounds with low-frequency modulations (e.g., amplitude modulation rates in cat vocalizations), and from time intervals between high-frequency transients (e.g., pulse-echo delay in bat sonar). Time locking of neuronal responses to periodicities from about 50 ms down to 4 ms or less (about 20–300 Hz) is preserved in the auditory midbrain, where responses are dispersed across many neurons with different onset latencies from 4–5 to 20–50 ms. Midbrain latency distributions are wide enough to encompass two or more repetitions of successive acoustic events, so that responses to multiple, successive periods are ongoing simultaneously in different midbrain neurons. These latencies have a previously unnoticed periodic temporal pattern that determines the specific times for the dispersed on-responses.     

Bilateral collicular interaction: modulation of auditory signal processing in amplitude domain

In the ascending auditory pathway, the inferior colliculus (IC) receives and integrates excitatory and inhibitory inputs from many lower auditory nuclei, intrinsic projections within the IC, contralateral IC through the commissure of the IC and from the auditory cortex. All these connections make the IC a major center for subcortical temporal and spectral integration of auditory information. In this study, we examine bilateral collicular interaction in modulating amplitude-domain signal processing using electrophysiological recording, acoustic and focal electrical stimulation. Focal electrical stimulation of one (ipsilateral) IC produces widespread inhibition (61.6%) and focused facilitation (9.1%) of responses of neurons in the other (contralateral) IC, while 29.3% of the neurons were not affected. Bilateral collicular interaction produces a decrease in the response magnitude and an increase in the response latency of inhibited IC neurons but produces opposite effects on the response of facilitated IC neurons. These two groups of neurons are not separately located and are tonotopically organized within the IC. The modulation effect is most effective at low sound level and is dependent upon the interval between the acoustic and electric stimuli. The focal electrical stimulation of the ipsilateral IC compresses or expands the rate-level functions of contralateral IC neurons. The focal electrical stimulation also produces a shift in the minimum threshold and dynamic range of contralateral IC neurons for as long as 150 minutes. The degree of bilateral collicular interaction is dependent upon the difference in the best frequency between the electrically stimulated IC neurons and modulated IC neurons. These data suggest that bilateral collicular interaction mainly changes the ratio between excitation and inhibition during signal processing so as to sharpen the amplitude sensitivity of IC neurons. Bilateral interaction may be also involved in acoustic-experience-dependent plasticity in the IC. Three possible neural pathways underlying the bilateral collicular interaction are discussed.     

Responses of cat auditory cortical neurons to tones of different frequencies and electrical stimulation of corresponding regions of the cochlea

Characteristic frequencies of neurons in the cat auditory cortex (area AI) whose receptive fields are located in different parts of the basilar membrane of the cochlea were determined in cats anesthetized with pentobarbital. The higher the characteristic frequency of a neuron in area AI, the nearer its receptive field lies to the base of the cochlea. Receptive fields of neurons with a characteristic frequency higher than 4 kHz lie on the first 10 mm of the basilar membrane. Receptive fields of neurons with a characteristic frequency below 4 kHz lie on the remaining 11–12 mm of the membrane. The effect of electrical stimulation of the center of the receptive field of a neuron corresponds to its response to a tone of characteristic frequency. The more the frequency of the acting tone differs from the characteristic frequency, or the further the point of stimulation from the center of the receptive field of the neuron, the less likely is the neuron to respond with an action potential. Neurons with a low characteristic frequency have wider receptive fields than neurons with a high characteristic frequency. Receptive fields of neurons with close characteristic frequencies on the basilar membrane overlap considerably. It was shown by the method of paired stimulation that excitation evoked in neurons in area AI by the action of a tone of a particular frequency is followed by long-lasting inhibition. This inhibition lasts longest and is most effective if a tone of the characteristic frequency is used.     

Synaptic effects induced in efferent neurons of the parietal associative cortex of the cat by stimulation of the cerebellar nuclei

Properties of EPSP, evoked in efferent neurons of the parietal associative cortex by stimulation of the cerebellar nuclei, were studied in acute experiments on anesthetized and immobilized cats; intracellular recording was used. The neurons were identified by their antidromic activation after stimulation of the motor cortex, pontinen.n. proprii, or red nucleus. The effects of stimulation of the cerebellar nuclei were of oligo- and polysynaptic nature. The latencies of cerebellofugal EPSP correlated with the latencies of antidromic activation, and correlations were significant both in the cases when the effects of stimulation of separate efferent projections (cortico-cortical, cortico-pontine, or cortico-rubral) and the effects of stimulation of separate cerebellar nuclei were analyzed. The functional role of the efferent systems of the parietal associative cortex and significance of functional parameters of the neurons constituting these systems are discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 3, pp. 190–198, May–June, 1995.     

Fast detection of unexpected sound intensity decrements as revealed by human evoked potentials

The detection of deviant sounds is a crucial function of the auditory system and is reflected by the automatically elicited mismatch negativity (MMN), an auditory evoked potential at 100 to 250 ms from stimulus onset. It has recently been shown that rarely occurring frequency and location deviants in an oddball paradigm trigger a more negative response than standard sounds at very early latencies in the middle latency response of the human auditory evoked potential. This fast and early ability of the auditory system is corroborated by the finding of neurons in the animal auditory cortex and subcortical structures, which restore their adapted responsiveness to standard sounds, when a rare change in a sound feature occurs. In this study, we investigated whether the detection of intensity deviants is also reflected at shorter latencies than those of the MMN. Auditory evoked potentials in response to click sounds were analyzed regarding the auditory brain stem response, the middle latency response (MLR) and the MMN. Rare stimuli with a lower intensity level than standard stimuli elicited (in addition to an MMN) a more negative potential in the MLR at the transition from the Na to the Pa component at circa 24 ms from stimulus onset. This finding, together with the studies about frequency and location changes, suggests that the early automatic detection of deviant sounds in an oddball paradigm is a general property of the auditory system.