Free-field Stimulation Research Articles

The role of GABAergic inhibition on direction-dependent sharpening of frequency tuning in bat inferior collicular neurons

This study examined the role of GABAergic inhibition on direction-dependent sharpening of frequency tuning curves (FTCs) in bat inferior collicular (IC) neurons under free field stimulation conditions. The minimum threshold (MT) at the neurons best frequency (BF) and the sharpness ( Q 10, Q 20, Q 30) of FTCs of most IC neurons increased as the sound direction changed from contralateral azimuths to ipsilateral azimuths. The application of GABA A antagonist, bicuculline, lowered all MTs but the application did not abolish direction-dependent variation in MT. MTs determined during bicuculline application at 40 ipsilateral were still significantly higher than those determined at 40° contralateral (two-tailed paired t-test, P<0.0001). In contrast, although application of bicuculline essentially had no effect on the BFs of IC neurons, it differentially broadened neurons FTCs at different azimuths abolishing the direction-dependent sharpening of frequency tuning (i.e. Q n values, two-tailed paired t-test, P<0.01). These data indicate that GABAergic inhibition makes an important contribution to the direction-dependent frequency tuning of most IC neurons.

Temporally patterned pulse trains affect duration tuning characteristics of bat inferior collicular neurons

This study examines the effect of temporally patterned pulse trains on duration tuning characteristics of inferior collicular neurons of the big brown bat. Eptesicus fuscus, under free-field stimulation conditions. Using a 50% difference between maximal and minimal responses as a criterion, the duration tuning characteristics of inferior collicular neurons determined with pulse trains of different pulse durations are described as band-pass, long-pass, short-pass, and all-pass. Each band-pass neuron discharged maximally to a specific pulse duration that was at least 50% larger than the neuron's responses to a long- and a short-duration pulse. In contrast, each long- or short-pass neuron discharged maximally to a range of long- or short-duration pulses that were at least 50% larger than the minimal responses. The number of impulses of an all-pass neuron never differed by more than 50%. When pulse trains were delivered at different pulse repetition rates, the number of short-pass and band-pass neurons progressively increased with increasing pulse repetition rates. The slope of the duration tuning curves also became sharper when determined with pulse trains at high pulse repetition rates. Possible mechanisms underlying these findings are discussed.

Corticofugal regulation of excitatory and inhibitory frequency tuning curves of bat inferior collicular neurons

Corticofugal regulation of excitatory and inhibitory frequency tuning curves (FTCs) of neurons in the central nucleus of bat inferior colliculus (ICc) was studied by electrical stimulation of the primary auditory cortex (AC stimulation) under free field stimulation conditions using a two-tone inhibition paradigm. AC stimulation narrowed the excitatory FTCs and asymmetrically expanded the lateral inhibitory FTCs of corticofugally inhibited ICc neurons. The opposite results were observed for excitatory and inhibitory FTCs of corticofugally facilitated ICc neurons. These data support previous reports that corticofugal systems work together with widespread lateral inhibition to regulate subcortical frequency processing.

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 inter-stimulus 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 gamma-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.

GABAergic disinhibition affects responses of bat inferior collicular neurons to temporally patterned sound pulses.

Using the big brown bat, Eptesicus fuscus, as a model mammalian auditory system, we studied the effect of GABAergic disinhibition by bicuculline on the responses of inferior collicular (IC) neurons to temporally patterned trains of sound pulses delivered at different pulse repetition rates (PRRs) under free-field stimulation conditions. All 66 neurons isolated from eight bats either discharged one to two impulses (phasic on responders, n = 41, 62%), three to eight impulses (phasic bursters, n = 19, 29%), or many impulses throughout the entire duration of the stimulus (tonic responders, n = 6, 9%). Whereas 50 neurons responded vigorously to frequency-modulated (FM) pulses, 16 responded poorly or not at all to FM pulses. Bicuculline application increased the number of impulses of all 66 neurons in response to 4 ms pulses by 15-1,425%. The application also changed most phasic on responders into phasic bursters or tonic responders, resulting in 12 (18%) phasic on responders, 34 (52%) phasic bursters, and 20 (30%) tonic responders. Response latencies of these neurons were either shortened (n = 25, 38%) by 0.5-6.0 ms, lengthened (n = 9, 14%) by 0. 5-2.5 ms or not changed (n = 32, 48%) on bicuculline application. Each neuron had a highest response repetition rate beyond which the neuron failed to respond. Bicuculline application increased the highest response repetition rates of 62 (94%) neurons studied. The application also increased the highest 100% pulse-locking repetition rates of 21 (32%) neurons and facilitated 27 (41%) neurons in response to more pulses at the same PRR than predrug conditions. According to average rate-based modulation transfer functions (average rate MTFs), all 66 neurons had low-pass filtering characteristics both before and after bicuculline application. According to total discharge rate-based modulation transfer functions (total rate MTFs), filtering characteristics of these neurons can be described as band-pass (n = 52, 79%), low-pass (n = 12, 18%), or high-pass (n = 2, 3%) before bicuculline application. Bicuculline application changed the filtering characteristics of 14 (21%) neurons. According to synchronization coefficient-based modulation transfer functions, filtering characteristics of these neurons can be described as low-pass (n = 41, 62%), all-pass (n = 11, 17%), band-suppression (n = 7, 10.5%), and band-suppression-band-pass filters (n = 7, 10.5%). Bicuculline application changed filtering characteristics of 19 (29%) neurons.

Head-related transfer functions of the barn owl: measurement and neural responses

Sounds arriving at the eardrum are filtered by the external ear and associated structures in a frequency and direction specific manner. When convolved with the appropriate filters and presented to human listeners through headphones, broadband noises can be precisely localized to the corresponding position outside of the head (reviewed in Blauert, 1997). Such a `virtual auditory space' can be a potentially powerful tool for neurophysiological and behavioral work in other species as well. We are developing a virtual auditory space for the barn owl, Tyto alba, a highly successful auditory predator that has become a well-established model for hearing research. We recorded catalogues of head-related transfer functions (HRTFs) from the frontal hemisphere of 12 barn owls and compared virtual and free sound fields acoustically and by their evoked neuronal responses. The inner ca. 1 cm of the ear canal was found to contribute little to the directionality of the HRTFs. HRTFs were recorded by inserting probetube microphones to within about 1 or 2 mm of the eardrum. We recorded HRTFs at frequencies between 2 and 11 kHz, which includes the frequencies most useful to the owl for sound localization (3–9 kHz; Konishi, 1973). Spectra of virtual sounds were within ±1 dB of amplitude and ±10° of phase of the spectra of free field sounds measured near to the eardrum. The spatial pattern of responses obtained from neurons in the inferior colliculus were almost indistinguishable in response to virtual and to free field stimulation.

Spatial-tuning properties of auditory neurons in the optic tectum of the pigeon

We studied the auditory neurons in the optic tectum of the unanesthetized pigeon, using single-unit recordings and acoustic free-field stimulation. Most units showed spatial tuning, with best areas located in the contralateral hemifield. All units responded also to visual stimuli, the auditory best areas being in rough alignment with visual receptive fields.

The effect of pulse repetition rate, pulse intensity, and bicuculline on the minimum threshold and latency of bat inferior collicular neurons.

This study examines the effect of pulse repetition rate (PRR), pulse intensity, and bicuculline on the minimum threshold (MT) and latency of inferior collicular neurons of the big brown bat, Eptesicus fuscus, under free-field stimulation conditions. It tests the hypothesis that changes in MT and latency of collicular neurons are co-dependent on PRR. The number of impulses in inferior collicular neurons (n = 245) increased either monotonically (25%) or non-monotonically (75%) with pulse intensity. Latencies either decreased to a plateau (72%), fluctuated unpredictably within 3 ms (21%) or changed very little (7%) with increasing pulse intensity. Latencies and MTs of most collicular neurons increased by 1.5-24 ms (mean +/- SD = 4.8 +/- 3.3 ms) and 4-75 dB (mean +/- SD = 22.1 +/- 16.2 dB) with increasing PRR. In most neurons (94%), the latency increase was completely (42%) or partially (52%) eliminated when pulse intensity was compensated for the MT increase with PRR. Complete elimination of latency was achieved by bicuculline application. In a few neurons (6%), the latency increase with PRR was not affected by compensated pulse intensity or bicuculline application.

Auditory inattention in right-hemisphere-damaged patients with and without visual neglect

Fifteen right-hemisphere-damaged patients, eight with- and seven without visual neglect (N+ and N−, respectively), were assessed for the presence of auditory neglect using free-field bilateral simultaneous stimulation (BSS) and pseudorandom unilateral stimulation. Eight healthy subjects served as controls. Both N+ and N− groups extinguished left-sided sound stimuli in the BSS condition. N+ (but not N−) patients showed a right-side advantage in sound localization and were inferior, compared to normal individuals, in their ability to localize unilaterally administered sounds on the left side. Blindfolding significantly improved the localization performance. In a task demanding stimulus identification, both N+ and N− groups performed abnormally when auditory stimuli came from the left. Free-field stimulation is thus an adequate technique for the detection of auditory neglect. The results are consistent with the notion that left-sided neglect reflects a pathologically exaggerated attentional bias towards the right in normal individuals.

Cytoarchitecture and Sound Activated Responses in the Auditory Cortex of the Big Brown Bat, Eptesicus fuscus

Under free field and closed-system stimulation conditions, we studied the frequency threshold curves, auditory spatial sensitivity and binaurality of neurons in the primary auditory cortex (AC) of the big brown bat, Eptesicus fuscus. All 298 recorded AC neurons discharged phasically. They were recorded at depths less than 1,000 μm with response latencies of 7–25 ms, best frequencies (BFs) of 28–97 kHz and minimum thresholds (MTs) of 8–90 dB SPL. They received excitatory inputs from the contralateral ear and either excitatory (EE) or inhibitory (EI) inputs from the ipsilateral ear. These cortical neurons were tonotopically organized along the anteroposterior axis of the AC. High best frequency neurons were located anteriorly and low best frequency neurons posteriorly. They were most sensitive to sounds delivered from a restricted region of the contralateral frontal auditory space (0°-50° in azimuth and 2° up, 15° down in elevation). Frontal auditory space representation appears to be systematically arranged according to the tonotopic axis such that the lateral space is represented posteriorly and the middle space anteriorly. Cortical neurons sequentially isolated from an orthogonally penetrated electrode had similar frequency threshold curves, BFs, MTs, points of maximal auditory spatial sensitivity and binaurality. The EE and EI columns are organized concentrically such that the small number of centrally located EE columns were surrounded by an overwhelming number of El columns. Using Nissl and Golgi stains as well as c-fos immunocytochemistry, we studied the Cytoarchitecture, cell types and sound elicited Fos-like immunoreactivity in the primary AC of this bat species. The primary AC of this bat species can be described into molecular (137 μm), external granular (55 μm), external pyramidal (95 μm), internal granular (102 μm), internal pyramidal (191 μm) and multiform (120 μm) layers. The main type of cells distributed among these six layers are the small, medium and large pyramidal cells. Others include the stellate, horizontal, granular, fusiform, basket, and Martinotii cells. When stimulated with 30 kHz and 79 dB SPL sounds under natural conditions, bilaterally and symmetrically distributed Fos-like immunoreactive neurons were observed in about 20% of neurons in each AC. When stimulated under monaurally plugged conditions, 39–48% more of Fos-like immunoreactive neurons were observed in the ipsilateral AC. This finding supports the fact that the primary AC receives auditory inputs mainly from the contralateral ear.

Sound Pressure Transformation at the Pinna of Mus domesticus

Sound pressure transformation properties at the pinna of laboratory mice, Mus domesticus, were studied by measuring the sound pressure level of continuous tone at a series of frequencies at the tympanic membrane as a function of the position of a sound source under free-field stimulation conditions. Sound pressure transformation functions showed some prominent spectral notches throughout the frequency range of 10–80 kHz tested. When delivered from some angles within the ipsilateral frontal hemisphere, the sound pressure at the tympanic membrane of certain frequencies may be lower than that determined at the corresponding contralateral angles. For each sound frequency tested, there was an angle (the acoustic axis) within the ipsilateral frontal hemisphere from which the delivered sound reached a maximal pressure level at the tympanic membrane. However, sound delivered from the acoustic axis did not always generate a maximal pressure transformation. The isopressure contours determined within 2–5 dB of the maximal pressure were circumscribed, and their contained angular areas were found to decrease with increasing sound frequency. The 2 dB maximal pressure area may appear at more than one angular area for some test frequencies. Removal of the ipsilateral pinna or modification of pinna posture expanded isopressure contours irregularly and split the 2 dB maximal pressure area into several parts.

Binaural inhibition is important in shaping the free-field frequency selectivity of single neurons in the inferior colliculus.

1. We have shown previously that under free-field stimulation in the frontal field, frequency selectivity of the majority of inferior colliculus (IC) neurons became sharper when the loudspeaker was shifted to ipsilateral azimuths. These results indicated that binaural inhibition may be responsible for the direction-dependent sharpening of frequency selectivity. To test the above hypothesis directly, we investigated the frequency selectivity of IC neurons under several conditions: monaural stimulation using a semiclosed acoustical stimulation system, binaural stimulation dichotically also using a semiclosed system, free-field stimulation from different azimuths, and free-field stimulation when the ipsilateral ear was occluded monaurally (coated with a thick layer of petroleum jelly, which effectively attenuated acoustic input to this ear). 2. The binaural interaction pattern of 98 IC neurons of northern leopard frogs (Rana pipiens pipiens) were evaluated; of these neurons, there were 34 EE and 64 EO neurons. The majority of IC neurons (92 of 98) showed some degree of binaural inhibition (i.e., showing diminished response when the ipsilateral and contralateral ears were stimulated simultaneously) whether they were designated as EE or EO; these IC neurons thus were classified as EE-I or EO-I. Neurons were classified as exhibiting strong inhibition if the ILD function showed a pronounced response decrement, i.e., a decrease of > or = 50% of the response to monaural stimulation of the contralateral ear. Those neurons that showed smaller response decrements (decrease was > or = 25% but < 50%) were designated as showing weak inhibition. Most of these EE-I and EO-I neurons (n = 68) showed strong binaural inhibition. 3. In agreement with results from our earlier studies, frequency threshold curves (FTCs) of IC neurons were altered by sound azimuth. Independent of binaural interaction pattern, most IC neurons (59 of 98) showed a narrowing of the FTC as sound direction was changed from contralateral 90 deg (c90 degrees) to ipsilateral 90 deg (i90 degrees). IC neurons that exhibited the largest direction-dependent changes in frequency selectivity were typically those that displayed stronger binaural inhibition. Occlusion of the ipsilateral ear, which reduced the strength of binaural inhibition by this ear, abolished direction-dependent frequency selectivity. 4. FTCs of IC neurons that exhibited little to moderate direction-dependent effects on frequency selectivity were associated typically with neurons that displayed weak binaural inhibition. Associated with this weak binaural inhibition, central neural responses under monaural occlusion also displayed only small effects; the FTCs were only slightly broader than those derived in the intact condition, and as before, the experimental manipulation resulted in abolishment of direction-dependent frequency selectivity. 5. In contrast to most IC neurons, which showed direction-dependent narrowing of the FTC, about one-third (34 of 98) of IC neurons studied showed a broadening of the FTC when sound direction was shifted to ipsilateral azimuths. Interestingly, for 90% of these 34 neurons, monaural occlusion resulted in narrowing of the bandwidth at each azimuth instead of broadening of the FTC bandwidth. We have evidence to suggest that this direction-dependent broadening is actually a consequence of a truncation or loss of the tip of the FTC derived at c90 degrees, which results from strong binaural inhibition. 6. To compare the frequency threshold tuning in response to monaural stimulation of each ear with free-field FTCs, we measured FTCs for each of the 34 EE neurons to independent contralateral and ipsilateral stimulation. FTCs derived from ipsilateral monaural stimulation were significantly narrower than those resulting from contralateral monaural stimulation, independent of a neuron's direction-dependent changes in frequency selectivity.

Temporally patterned pulse trains affect directional sensitivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus.

The directional sensitivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus, was studied under free field stimulation conditions with 3 temporally patterned trains of sound pulses which differed in pulse repetition rate and duration. The directional sensitivity curves of 92 neurons studied can be described as hemifield, directionally-selective, or non-directional according to the variation in the number of impulses with pulse train direction. When these neurons were stimulated with all 3 pulse trains, the directional sensitivity curves of 50 neurons was unchanged but that of the other 42 neurons changed from one type into another. When these pulse trains were delivered at high pulse repetition rate and short pulse duration, they significantly sharpened the directional sensitivity of two thirds of the neurons examined by reducing the angular range and increasing the slope of their impulse directional sensitivity curves. These pulse trains also sharpened the slope of the threshold directional sensitivity curves of 25 neurons studied. However, when directional sensitivity of collicular neurons was determined with pulse trains that differed only in pulse repetition rate or in pulse duration, significant sharpening of directional sensitivity was rarely observed in all experimental conditions tested. Possible mechanisms underlying these findings are discussed.

Directionality of sound-pressure transformation at the pinna of Musdomesticus

Sound-pressure transformation properties at the pinna of laboratory mice, Musdomesticus, were studied by measuring the sound-pressure level of a continuous tone at a series of frequencies at the tympanic membrane as a function of the position of a sound source under free-field stimulation conditions. The spectral transformation, the interaural spectral difference, the isopressure contours and the interaural pressure difference contours were plotted. Sound-pressure transformation functions showed some prominent spectral notches throughout the frequency range of 10–80 kHz tested. When delivered from some angles within the ipsilateral frontal hemisphere, the sound pressure at the tympanic membrane of certain frequencies may be lower than that determined at the corresponding contralateral angles (i.e., the negative pressure). For each sound frequency tested, there was an angle (the acoustic axis) within the ipsilateral frontal hemisphere from which the delivered sound reached a maximal pressure level at the tympanic membrane. However, the acoustic axis often changed to a new angle after removal of the ipsilateral pinna. Removal of the ipsilateral pinna or modification of pinna posture expanded isopressure contours irregularly and split the 2-dB maximal pressure area into several parts. [Work supported by NIH and HFSP.]

Sound pressure transformation at the pinna of Mus domesticus.

Sound pressure transformation properties at the pinna of laboratory mice Mus domesticus were studied by measuring the sound pressure level of a continuous tone at a series of frequencies at the tympanic membrane as a function of the position of a sound source under free-field stimulation conditions. The spectral transformation, the interaural spectral difference, the isopressure contours and the interaural pressure difference contours were plotted. Sound pressure transformation functions showed some prominent spectral notches throughout the frequency range tested (10-80 kHz). However, the notch frequency did not appear to be systematically related to sound direction. The study of interaural pressure difference demonstrated that, when delivered from some angles within the ipsilateral frontal hemisphere, the sound pressure at the tympanic membrane of certain frequencies may be lower than that determined at the corresponding contralateral angles. For each sound frequency tested, there was an angle (the acoustic axis) within the ipsilateral frontal hemisphere from which the delivered sound reached a maximal pressure level at the tympanic membrane. However, the acoustic axis often changed to a new angle after removal of the ipsilateral pinna. In addition, sound delivered from the acoustic axis did not always generate a maximal pressure transformation. The isopressure contours determined within 2-5 dB of the maximal pressure were circumscribed, and their contained angular areas were found to decrease with increasing sound frequency. The 2 dB maximal pressure area may appear at more than one angular area for some test frequencies. Removal of the ipsilateral pinna or modification of pinna posture expanded isopressure contours irregularly and split the 2 dB maximal pressure area into several parts. The sound pressure difference determined between the angles of maximal and minimal sound pressure (the maximal directionality) increased with sound frequency regardless of pinna posture. Acoustic gain of the pinna at the acoustic axis reached 6-12 dB, depending upon sound frequency. However, the pinna gain was not always maximal at the acoustic axis for a given frequency.

Responses of pontine neurons of the big brown bat, Eptesicus fuscus, to temporally patterned sound pulses

Under free field stimulation conditions, this study examined the responses of pontine neurons of Eptesicus fuscus to temporally patterned sound pulses by means of repetitive single pulses and pulse trains. Among 93 pontine neurons isolated, 90 always discharged less than 5 impulses to sound pulses presented during this study and 3 discharged impulses throughout the whole duration of each presented pulse. Responses to sound pulses at different repetition rates were examined in 65 neurons. The number of impulses of individual neurons discharged to each pulse varied within a given repetition rate and among different repetition rates. Although these pontine neurons showed different degrees of habituation to high repetition rates, more than 25% could follow the highest repetition rate tested (100 pps). However, they did not always discharge maximal number of impulses to this repetition rate. The total number of impulses discharged by a neuron was also affected by pulse duration. Thus, each pontine neuron discharged maximally to a specific combination of pulse repetition rate and duration. Using a 50% difference between the maximal and minimal responses as a criterion, the response function with respect to repetition rate and duration can be described as band-pass, low-pass, high-pass, all-pass and irregular. These response properties reflect more those of inferior collicular neurons than auditory cortical neurons. This study also showed that response latencies of pontine neurons examined were lengthened by increasing pulse repetition rate and duration. In addition, whereas minimum thresholds of pontine neurons were elevated by increasing repetition rate, they were lowered by increasing pulse duration.

NMDA-receptor antagonists disrupt the formation of the auditory space map in the mammalian superior colliculus

In the ferret (Mustela putorius) the map of auditory space in the deeper layers of the superior colliculus (SC) matures over a period of several postnatal weeks, a process known to be guided by both visual and auditory experience. The auditory responses are initially very broadly tuned, and gradually become more selective for specific sound locations that coincide with the visual receptive fields recorded in the same region of the SC. To investigate the possible involvement of NMDA-type glutamate receptors in the postnatal development of this auditory representation, we have reared ferrets in which 400 microns thick sheets of the slow-release polymer Elvax, containing the NMDA receptor antagonists MK801 or APV, were placed on the dorsal surface of the SC. The Elvax was implanted on postnatal day (P) 25-27, just before the onset of hearing, and removed 5-6 weeks later, just prior to recording from the SC on around P61-70. In vitro measurements with Elvax containing 3H-MK801 revealed that the amount of drug released declined sharply over the first 10 d and then stabilized at a fairly constant rate for the following 5 weeks. These in vitro data were found to parallel the in vivo release of MK801 from implanted Elvax slices. Diffusion of MK801 from the implant was measured and significant levels were found within 800 microns of the SC surface, suggesting that the action of MK801 was restricted to the superficial and intermediate layers of the nucleus. Extracellular recordings were made from visual and auditory units in the SC in response to free-field stimulation. The visual responses of units recorded in the superficial layers appeared to be unaffected by either of the drug treatments, and formed a normal, adult-like map of visual azimuth along the rostrocaudal axis of the SC in all animals. Most of the auditory single-unit responses recorded at this age in normal, unoperated controls were spatially tuned and topographically organized, although the map of sound azimuth was less precise than that in adult ferrets. Data from age-matched control animals that had been reared with drug-free Elvax implants were not statistically different from the unoperated juvenile ferrets. However, in animals reared with APV or MK801 Elvax implants, there was an increase in the relative numbers of auditory units that were ambiguously tuned to two or more locations.(ABSTRACT TRUNCATED AT 400 WORDS)

Responses of inferior collicular neurons of the FM bat, Eptesicus fuscus, to pulse trains with varied pulse amplitudes

Under free field stimulation conditions, we studied the responses of inferior collicular neurons of the FM bat, Eptesicus fuscus, to pulse trains with varied pulse amplitudes. Each pulse train consisted of 7 pulses of 4 ms delivered at 24 ms interpulse-intervals (i.e. 42 pulses/s). For a control pulse train, all pulse amplitudes were equal to a neuron's best amplitude which, when delivered in single pulses, elicited maximal number of impulses from the neuron. The amplitudes of individual pulses of the remaining pulse trains were linearly increased or decreased at a slope of 0, 14, 28, 42, 56 and 69 dB/s. All 56 inferior collicular neurons discharged to pulse trains were of two main types. Type I ( N43, 77%) neurons discharged to each pulse within a train while type II ( N11, 20%) neurons discharged to the first pulse of a train stimulus only. Discharge patterns of the remaining ( N2, 3%) neurons changed between type I and type II when stimulated with different pulse trains. The number of impulses discharged by a neuron varied with different pulse trains. In addition, the number of impulses discharged to each pulse by type I neurons also varied among individual pulses within the train. Only 14 neurons (25%) discharged maximally to the control pulse train. Responses of the remaining neurons to other pulse trains were either 30%–120% larger than ( N17, 30%) or within 30% ( N25, 45%) of the control pulse train response. Furthermore, half of 56 neurons selectively discharged to a most preferred pulse train with a response magnitude which was at least 50% larger than the response to the least preferred pulse train. Possible mechanisms underlying the different discharge patterns are discussed in terms of a neuron's recovery cycle, minimum threshold and inhibitory period relative to the temporal characteristics (pulse repetition rate and amplitude) of the pulse trains.

Pulse repetition rate increases the minimum threshold and latency of auditory neurons

The effect of pulse repetition rate on auditory sensitivity of the big brown bat, Eptesicus fuscus, was studied by determining the minimum threshold, response latency and recovery cycle of inferior collicular neurons at different repetition rates under free field stimulation conditions. In general, collicular neurons shortened the response latency and increased the number of impulses monotonically or non-monotonically with stimulus intensity. They recovered at least 50% when the interpulse interval was 10–57 ms. In addition, they increased the minimum threshohold, lengthened the response latency, and reduced the number of impulses discharged to each pulse with increasing repetition rate. The increase in minimum threshold with repetition rate is partly because the neuron can not recover from previous stimulation when the interpulse interval is shortened. This increase reduces a neuron's response sensitivity and thus diminishes its number of impulses to each presented pulse. This increase also reduces the effectiveness of a given stimulus intensity which contributes to the lengthening of the neuron's response latency. Data obtained from single neuron recordings are used to highlight these observations. Implications of present findings regarding the bat's echolocation are also discussed.

Neurons in the inferior colliculus, auditory cortex and pontine nuclei of the FM bat, Eptesicus fuscus respond to pulse repetition rate differently

Single-neuron responses to pulse repetition rate in the inferior colliculus, auditory cortex and pontine nuclei of the FM bat, Eptesicus fuscus were studied under free-field stimulation conditions. The best frequency (BF) and minimum threshold (MT) of each neuron were first determined with a 4 ms pulse broadcast from a specific point (response center) of the bat's frontal auditory space at which the neuron had maximal spatial sensitivity. The neuron's intensity-rate function was then studied with a 4 ms BF pulse delivered at 10 dB increments above its MT in order to determine the best intensity to which the neuron discharged maximally. The neuron's discharge pattern and number of impulses to 32 trials of 300 ms train stimuli, which consisted of different number of 4 ms BF and best intensity pulses (1, 2, 3, 8, 12, 19, 24, 29 pulses/train) and delivered at an interpulse interval of 1000, 250, 170, 100, 40, 25, 15, 12 and 10 ms (i.e. at a pulse repetition rate of 1, 4, 6, 10, 25, 40, 67, 83, 100 pulses/s), were sequentially recorded. All neurons recorded from the inferior colliculus, auditory cortex and pontine nuclei discharged phasically (1–3 impulses) but they responded to the pulse repetition rate in different manners. More than 63% of 38 inferior collicular and 65 pontine neurons studied discharged impulses to each pulse within a train stimulus when the pulse repetition rate was up to 40 pulses/s. Although the number of neurons showing such a one-to-one following response decreased with increasing pulse repetition rate, at least 20% of neurons still maintained such a response pattern even at a pulse repetition rate of 100 pulses/s. In contrast, responses of 60% of 36 cortical neurons followed the pulse repetition rate only up to 10 pps beyond which all but one discharged impulses to the first pulse of a train stimulus. We suggest that the different discharge pattern to pulse repetition rate among individual neurons from these three brain centers is primarily due to their different recovery cycles.