Tonotopic representation and space map in the non-primary auditory cortex of the mustached bat

As auditory system has no sensory epithelium into which auditory space are projected, we studied the physiological map of the auditory space in the non-primary auditory cortex of the mustached bat by using the echo of their orientation sound. Ten bats were used as experimental subjects. Tungsten wire electrodes were inserted obliquely in the dorsomedial (DM) and ventroposterior (VP) areas of the non-primary auditory cortex. When single neuron was isolated, best frequency (BF), best azymuth (BAZ) and best elevation (BEL) were measured and were plotted on a schematic figure. To mimic its biosonar, one loudspeaker, delivering synthesized orientation sounds, was placed in front of the animal, and another loudspeaker delivering synthesized echo was mounted on a movable hoop. Tonotopic representation was observed but complicated in both areas, and those areas could be divided into several subdivisions consisting of the neuron groups characterized by three frequency bands. The neurons were thought to be related to the processing of biosonar informations from the facts that their BFs agreed with the scope of the FM sweep of each echo harmonics. The magnitude of the response showed rapid increase at their BAZ or BEL, so that the neurons seemed to tune to a certain direction in the auditory space. Especially in the DM area, neurons assumed a systematic arrangement of their BAZs on the cerebral surface and showed some tendency of a systematic arrangement of their BELs. The DM area was thought to have a kind of neural map of the auditory space.     

Directional sensitivity of the auditory midbrain in the mustached bat to free-field tones

To ascertain the directional characteristics of the auditory system in the mustached bat, Pteronotus parnellii, we measured the summated neural response at the lateral lemniscus (N4) in response to pure tones at 30, 60 and 90 kHz, frequencies that are typical of the harmonics of this species' biosonar signal. Stimuli were presented at various vertical and horizontal locations in the contralateral hemifield. Intensity-response functions were measured at different horizontal locations for the second harmonic, and showed no variation in shape with variations in azimuth. There was little difference in directionality measured from either threshold or amplitude of N4 potentials. Our results show that areas of maximum sensitivity (best areas) were significantly different for each of the harmonics (P less than 0.05). The centers of the best areas were: first harmonic (30 kHz), 39 degrees azimuth and -19 degrees elevation; second harmonic, 20 degrees azimuth and 0 degrees elevation; and third harmonic, 12 degrees azimuth and -11 degrees elevation. Thus, with increasing frequency best areas shifted toward the vertical midline. Directionality to first harmonic stimuli was broader than to either of the two higher harmonics.     

Development of functional organization of the pallid bat auditory cortex

The primary auditory cortex is characterized by a tonotopic map and a clustered organization of binaural properties. The factors involved in the development of overlain representation of these two properties are unclear. We addressed this issue in the auditory cortex of the pallid bat. The adult pallid bat cortex contains a systematic relationship between best frequency (BF) and binaural properties. Most neurons with BF<30 kHz are binaurally inhibited (EO/I), while most neurons with BF>30 kHz are monaural (EO). As in other species, binaural properties are clustered. The EO/I cluster contains a systematic map of interaural intensity difference (IID) sensitivity. We asked if these properties are present at the time the bat acquires its full audible range (postnatal day [P] 15). Tonotopy, relationship between BF and binaural properties, and the map of IID sensitivity are adult-like at P15. However, binaural facilitation is only observed in pups older than P25. Frequency selectivity shows a BF-dependent sharpening during development. Thus, overlain representation of binaural properties and tonotopy in the pallid bat cortex is remarkably adult-like at an age when the full audible range is first present, suggesting an experience-independent development of overlapping feature maps.     

Cochlear resonance in the mustached bat: Behavioral adaptations

Mustached bats. Pteronotus p. parnellii. use complex, multiharmonic biosonar signals with prominent approx. 60 kHz (CF) components. The sense of hearing is especially acute to sounds near 60 kHz and the cochlea shows a number of specializations in the 60 kHz region. Foremost is a remarkable degree of cochlear resonance. In this study it is shown that: 1) any sounds near the resonance frequency elicit a pronounced resonance that continues after the stimulus terminates; 2) Doppler-shifted echoes of the bat's own cries may cause resonance; 3) continuous resonance can be produced by stimulating the ear with broadband noise but such resonance does not interfere with the bat's ability to Doppler-shift compensate during simulated flight; 4) significant changes in the resonance frequency of the cochlea occur during and after flight; 5) the changes in resonance can be dependent or independent of body temperature changes; and 6) mustached bats continuously adjust the CF component of their pulses to keep the second harmonic echoes in a constant frequency band near the resonance frequency. Thus, mustached bats not only compensate for Doppler-shifts imposed by their movements relative to that of a target, but they cochlear resonance compensate to deal with small changes in the micromechanical properties of the cochlea.     

Mechanisms underlying azimuth selectivity in the auditory cortex of the pallid bat

A cochlear implant (CI) signal processing strategy named F0 modulation (F0mod) was compared with the advanced combination encoder (ACE) strategy in a group of four post-lingually deafened Mandarin Chinese speaking CI listeners. F0 provides an enhanced temporal pitch cue by amplitude modulating the multichannel electrical stimulation pattern at the fundamental frequency (F0) of the incoming speech signal. Word and sentence recognition tests were carried out in quiet and in noise. The responses for the word-recognition test were further segmented into phoneme and tone scores. Off-line implementations of ACE and F0mod were used, and electrical stimulation patterns were directly streamed to the CI subject's implant. To focus on the feasibility of enhanced temporal cues for tonal language perception, idealized F0 information that was extracted from speech tokens in quiet was used in the F0mod processing of speech-in-noise mixtures. The results indicated significantly better lexical tone perception with the F0mod strategy than with ACE for the male voice (p<0.05). No significant differences in sentence recognition were found between F0mod and ACE.     

Topographical distribution of delay-tuned responses in the mustached bat inferior colliculus

In the mustached bat, delay-tuned neurons respond best to specific delays between the first harmonic frequency modulated (FM) component (FM1; 24-29 kHz) of the emitted biosonar pulse and a higher harmonic FM component in returning echoes (e.g. FM3, 72-89 kHz). These delay-tuned, combinatorial responses predominate in the inferior colliculus (IC) of the mustached bat. This study examined the topographical distribution of delay-tuned neurons in the 72-89 kHz frequency representation of the IC. We recorded and histologically localized 163 single units. Ninety units were facilitated and 41 were inhibited by the combination of two frequencies in the 24-29 kHz and 72-89 kHz ranges. The facilitatory responses were selective for delays up to 20 ms between the two signals. To determine if delay-tuned neurons were topographically organized, we plotted the dorsomedio-ventrolateral and caudo-rostral positions of each unit versus its best delay. Best delay was not correlated with either location. Response latency to best frequency tones was topographically organized, but was not correlated with best delay. This indicates that the latency axis in the IC is unrelated to the delay tuning of these combinatorial neurons. Because delay-tuned neurons are not topographically organized in the IC but are in the auditory cortex, our findings suggest that the creation and organization of delay-tuned neurons occur at different stages in the ascending auditory system.     

Sound-evoked efferent effects on cochlear mechanics of the mustached bat

The influence of the crossed medial efferent system on cochlear mechanics of the mustached bat was tested by measuring delayed evoked otoacoustic emissions (DEOAEs), cochlear microphonics, distortion product otoacoustic emissions (DPOAEs) and stimulus frequency otoacoustic emissions. Contralaterally delivered sinusoids, broadband noise and bat echolocation calls were used for acoustic stimulation of the efferent system. With all four measures we found a level-dependent suppression under stimulation with both broadband noise and echolocation calls. In addition, the sharply tuned cochlear resonance of the mustached bat which is involved in processing echolocation signals at 61 kHz shifted upward in frequency by several 100 Hz. Presentation of sinusoids did not have any significant effect. DEOAEs and DPOAEs were in some cases enhanced during contralateral presentation of the bat calls at moderate intensities. The most important function of the efferent system in the mustached bat might be the control of the extraordinarily fine-tuned resonator of this species, which is close to instability as evident from the very pronounced evoked otoacoustic emissions which sometimes convert into spontaneous otoacoustic emissions of high level.     

The efferent cochlear projections of the superior olivary complex in the mustached bat

Following the placement of horseradish peroxidase in the scala tympani, labeled neurons were found in the ipsilateral interstitial nucleus (INT) and throughout the ipsilateral and contralateral dorsomedial periolivary nuclei (DMPO). The neurons in the INT were morphologically distinct from those in the DMPO. The INT neurons formed a thin shell over the lateral superior olivary nucleus (LSO) and their dendrites extended into the body and hilar region. The DMPO neurons had long, tapering dendrites that extended in every direction. Data indicate that the crossed fibers in the floor of the ventricle arise entirely from the DMPO while uncrossed olivocochlear fibers originate in the INT and DMPO. It was estimated that 75% of the efferent fibers arise from the INT and 25% from the DMPO. Approximately 70% of the efferent neurons in each DMPO project to the contralateral cochlea via the crossed olivocochlear bundle. The number of olivocochlear neurons associated with each ear was determined to be approximately 1585. This number is similar to that found in cats and guinea pigs, but the number of neurons per unit length of the basilar membrane is considerably higher in the mustached bat than in other species examined to date. The compact, restricted locations of the neurons in the INT and DMPO in the mustached bat are different from those described for most other mammals and the arrangement in the mustached bat offers advantages over other species for future anatomical and physiological studies.     

Spatial tuning of auditory neurons in the superior colliculus of the echolocating bat, Myotis lucifugus

The azimuthal selectivity of auditory neurons was examined in the superior colliculus of the little brown bat, Myotis lucifugus. Frequency-modulated (FM) sounds, synthesized to mimic biosonar signals the echolocating bat naturally hears, were delivered from a loudspeaker moving across the front of the unanesthetized animal. Neurons were classified on the basis of their spatial tuning into two general classes: (i) hemifield units (34%) were broadly tuned to the contralateral side irrespective of sound pressure level; (ii) azimuth-sensitive units (66%) were sharply tuned to different azimuths at sound pressures near their minimum thresholds (MTs). A distinguishing feature of these azimuth-sensitive neurons is that they responded maximally to a sound source located at a preferred azimuth (best azimuth) for levels as high as 30 dB above their MT. Mapping experiments provide evidence of a sequential representation of best azimuth along the rostrocaudal extent of the superior colliculus, with sounds originating from 0-10 degrees ipsilateral coded at the rostral end, and from 30-40 degrees contralateral coded at the caudal end. The highly directional echolocation system of Myotis probably accounts for the limited azimuthal representation of echo-source spanning mainly 40 degrees to either side of its line of flight.     

Properties of echo delay-tuning receptive fields in the inferior colliculus of the mustached bat

One role of the inferior colliculus (IC) in bats is to create neuronal delay-tuning, which is used for the estimation of target distance in the echolocating bat's auditory system. In this study, we describe response properties of IC delay-tuned neurons of the mustached bat (Pteronotus parnellii) and compare it with those of delay-tuned neurons of the auditory cortex (AC). We also address the question if frequency content of the stimulus (pure-tone (PT) or frequency-modulated (FM) pairs stimulation) affects combination-sensitive interaction in the same neuron. Sharpness and sensitivity of delay-tuned neurons in the IC are similar to those described in the AC. However, in contrast to cortical responses, in collicular neurons the delay at which the neurons show the maximum response does not change with changes in echo level. This tolerance to changes in the echo level seems to be a property of collicular delay-tuned neurons, which is modified along the ascending auditory pathway. In the IC we found neurons that showed a facilitated delay-tuned response when stimulated with FM components and did not show any delay-tuning with PT stimulation. This result suggests that not only is echo delay-tuning generated in the IC but also its FM-specificity observed in the cortex could be created to some extent in the IC and then topographically organized at higher levels.     

Excitatory and facilitatory frequency response areas in the inferior colliculus of the mustached bat

In the mustached bat's central nucleus of the inferior colliculus (ICC), many neurons display facilitatory or inhibitory responses when presented with two tones of distinctly different frequencies. Our previous studies have focused on spectral interactions between specific frequency bands contained in the bat's sonar vocalization. In this study, we describe excitatory and facilitatory frequency response areas across all frequencies in the mustached bat's audible range. We show that many neurons in the ICC have more extensive frequency interactions than previously documented. We recorded responses of 96 single units to single tones and combinations of two tones. Best frequencies of the units ranged from 59-15 kHz. Forty-one units had a single, excitatory frequency response area. The rest of the units had more complex frequency tuning that included multiple excitatory frequency response areas and facilitatory frequency response areas. Some of the facilitatory frequency interactions were between one sound with energy in a sonar frequency band and a second sound with energy in a non-sonar frequency band. We also found that neurons could be facilitated by more than one additional frequency band. Our findings of extensive frequency interactions in the ICC of the mustached bat suggest that some neurons may be well suited for the analysis of complex sounds, possibly including social communication sounds.     

Cochlear and CNS tonotopy: normal physiological shifts in the mustached bat.

The ear of the mustached bat (Pteronotus parnellii) shows marked cochlear resonance near 60 kHz and many sharply tuned neurons throughout the brain have best frequencies (BF) near the cochlear resonance frequency (CRF). Controlled changes in the normal physiological range of body temperature (approx 37-42 degrees C) were used to change the CRF and to study the tuning properties of neurons in the cochlear nucleus (CN) and inferior colliculus (IC). In all cases there were concomitant shifts in the CRF and the BFs. Results were the same for single and multi-units, and for CN and IC units. Although the BF reliably changed with shifts in the CRF, the majority of the units showed no change in minimum threshold or the sharpness (Q10 dB) of tuning. The temperature-induced effects on cochlear tuning were similar to those previously described in nonmammalian vertebrates. The physiological data reveal that, within a narrow frequency band, cochlear and CNS tonotopy are labile in the mustached bat. The lability of tuning is further substantiated by adaptations of biosonar emission behavior with shifts in CRF (Henson et al., 1990).     

Neuronal sensitivity to interaural time differences in the sound envelope in the auditory cortex of the pallid bat

Interaural time differences in the envelope of a sound (envelope ITDs) can potentially provide spatial information at high frequencies where interaural phase differences (IPDs) are not available. Interaural intensity differences (IIDs) also provide important spatial information at high frequencies. Both IIDs and envelope ITDs can influence spatial perception at high frequencies, but behavioral and physiological studies suggest that IIDs dominate perception. This study examines envelope ITD sensitivity in the auditory cortex of the pallid bat, a species that uses passive sound localization at the low end of its audible range to find prey. Its auditory system is entirely 'high-frequency' in that phase-locking does not occur at the low end of its audible range. If the bat uses ITDs, they must be derived from the envelope of the signal. A previous study of envelope ITD sensitivity in its inferior colliculus (IC) reported that neurons are sensitive to the small +/-70 micros range of available ITDs. This study extends these findings to the cortical level to assess the transformation of ITD sensitivity and the binaural response properties that underlie this sensitivity. Two measures of sensitivity were used. The dynamic ITD range measures the range of ITDs over which the maximum response of a neuron decreases by 80%. When presented with square-wave amplitude-modulated tones statically delayed in arrival time, the average dynamic ITD range in the IC is 304 micros, but dropped to 175 micros in auditory cortex. IC neurons average a 38% change in maximum response over the relevant ITD range, while cortical neurons average a 67% change. Also measured were time-intensity trading ratios, which index the extent to which a change in IID can cause a shift the dynamic ITD range. Average trading ratios are approximately the same in the IC and auditory cortex (17.9 micros/dB vs. 16.7 micros/dB, respectively). Binaural interactions changed from the IC to auditory cortex. In IC, ITD sensitivity is an inhibitory, subtractive process in which ITDs reduce the response evoked by contralateral monaural stimulation. In the auditory cortex, both binaural inhibition and facilitation occur. In the majority of cortical neurons, IID and ITD functions were remarkably similar in shape, having stepped, step-peaked or peaked functions. The binaural interactions (inhibition and/or facilitation) evoked by ITDs and IIDs were also typically similar. These results suggest that IIDs and envelope ITDs are having similar effects on output of the same binaural comparator system.     

Divergent response properties of layer-V neurons in rat primary auditory cortex

Layer-V pyramidal cells comprise a major output of primary auditory cortex (A1). At least two cell types displaying different morphology, projections and in vitro physiology have been previously identified in layer-V. The focus of the present study was to characterize extracellular receptive field properties of layer-V neurons to determine whether a similar breakdown of responses can be found in vivo. Recordings from 105 layer-V neurons revealed two predominant receptive field types. Thirty-two percent displayed strong excitatory V/U-shaped receptive field maps and spiking patterns with shorter stimulus-driven interspike intervals (ISIs), reminiscent of the bursting cells discussed in the in vitro literature. V/U-shaped maps remained relatively unchanged across the three sequential repetitions of the map run on each neuron. Neurons with V/U-shaped maps were also easily depolarized with extracellular current pulse stimulation. In contrast, 47% of the neurons displayed Complex receptive field maps characterized by weak and/or inconsistent excitatory regions and were difficult to depolarize with current pulses. These findings suggest that V/U-shaped receptive fields could correspond to previously described intrinsic bursting (IB) cells with corticotectal projections, and that neurons with Complex receptive fields might represent the regular spiking (RS) cells with their greater inhibitory input and corticocortical/corticostriatal projection pattern.     

Processing of pure-tone and FM stimuli in the auditory cortex of the FM bat, Myotis lucifugus.

FM bats perceive their surroundings during echolocation by analyzing frequency-modulated (FM) acoustic signals. Results from this study indicate a cortical organization in Myotis lucifugus which is largely made up of neurons sensitive to FM sounds (FM-sensitive neurons). Three types of neurons were distinguished by their responses to pure-tone and FM stimuli: (1) Type I FM-sensitive units (83%), Type II FM-sensitive units (13%) and pure-tone sensitive units (4%). Type I FM-sensitive units responded to pure tones, but exhibited greater response magnitudes to FM stimuli when the best FM swept through the BF. An orderly frequency representation was found when the frequencies of pure tones essential for response (EPTs) in Type I units were mapped along the cortical surface. The EPTs for Type I neurons were usually found within the last millisecond of a downward FM sweep. As outlined by two neuronal network models, both the responses of Type I and II units could likely result from the convergence of excitatory and inhibitory lower level neurons with slightly differing BFs. Type II units were selective for an FM sweep and showed negligible to no response to pure-tone stimuli. Pure-tone sensitive units exhibited weak or no responses to FM stimuli. These neurons were clustered in a small area located rostrodorsal to the tonotopic zone and had significantly lower best frequencies than adjacent EPT frequencies of Type I FM-sensitive neurons.     

The auditory cortex of the bat Molossus molossus: Disproportionate search call frequency representation

The extent of the auditory cortex in the bat Molossus molossus was electrophysiologically investigated. Best frequencies and minimum thresholds of neural tuning curves were analyzed to define the topography of the auditory cortex. The auditory cortex encompasses an average cortical surface area of 5 mm². Characteristic frequencies are tonotopically organized with low frequencies being represented caudally and high frequencies rostrally. However, a large interindividual variability in the tonotopic organization was found. In most animals, the caudal 50% was tonotopically organized. More anterior, a variable area was found. A distinct field with reversed topography was not consistently found. Within the demarcated auditory cortex, frequencies of 30–40 kHz, which correspond to the frequency range of search calls emitted during hunting, are overrepresented, occupying 49% of the auditory cortex surface. High minimum thresholds >50 dB SPL were found in a narrow dorsal narrow area. Neurons with multipeaked tuning curves (20%) preferentially were located in the dorsal part of the auditory cortex. In accordance with studies in other bat species, the auditory cortex of M. molossus is highly sensitive to the dominant frequencies of biosonar search calls.     

Specializations for sharp tuning in the mustached bat: the tectorial membrane and spiral limbus.

The sense of hearing in the mustached bat, Pteronotus parnellii, is specialized for fine frequency analysis in three narrow bands that correspond to approx 30, 60 and 90 kHz constant frequency harmonics in the biosonar signals used for Doppler-shift compensation and acoustic imaging of the environment. Previous studies have identified anatomical specializations in and around the area of the cochlea that processes the dominant second harmonic component, but similar features have not been found in areas related to sharp tuning and high sensitivity for the first or third harmonics. In this report we call attention to the large size of the tectorial membrane and spiral limbus in all three areas that appear to process the harmonically related constant frequency components. These structures are especially pronounced in the regions of the cochlea that respond to the approx 61 kHz, second harmonic and 91.5 kHz, third harmonic bands; they correspond specifically to areas where the density of afferent nerve fibers is high and where very sharply tuned neurons occur. These data for cochleae with multiple specializations lend strong support to the idea that the mass of the tectorial membrane can be an important factor in establishing the response properties of the cochlea.     

Sodium salicylate reduces inhibitory postsynaptic currents in neurons of rat auditory cortex

Sodium salicylate (SS) is a medicine for anti-inflammation and for chronic pain relief with a side effect of tinnitus. To understand the cellular mechanisms of tinnitus induced by SS in the central auditory system, we examined effects of SS on evoked and miniature inhibitory postsynaptic currents (eIPSCs and mIPSCs) recorded from layer II/III pyramidal neurons of rat auditory cortex in a brain slice preparation with whole-cell patch-clamp techniques. Both eIPSCs and mIPSCs recorded from the auditory cortex could be completely blocked by bicuculline, a selective GABA(A) receptor antagonist. SS did not change the input resistance of neurons but was found to reversibly depress eIPSCs in a concentration-dependent manner. SS reduced eIPSCs to 82.3% of the control level at 0.5 mM (n=7) and to 60.9% at 1.4 mM (n=12). In addition, SS at 1.4 mM significantly reduced the amplitude of mIPSCs from 24.12+/-1.44 pA to 19.92+/-1.31 pA and reduced the frequency of mIPSCs from 1.34+/-0.23 Hz to 0.89+/-0.13 Hz (n=6). Our results demonstrate that SS attenuates inhibitory postsynaptic currents in the auditory cortex, suggesting that the alteration of inhibitory neural circuits may be one of the cellular mechanisms for tinnitus induced by SS in the central auditory region.     

Interaction between excitation and inhibition affects frequency tuning curve,response size and latency of neurons in the auditory cortex of the big brown bat,Eptesicus fuscus

Neurons in the auditory cortex (AC) receive convergent excitatory and inhibitory inputs from the lower auditory nuclei. Interaction between these two opposing inputs shapes different response properties of AC neurons. In this study, we examined how this interaction might affect the frequency tuning curves (FTCs), number of impulses and latency of AC neurons in the big brown bat, Eptesicus fuscus, using a probe (excitatory tone) and a masker (inhibitory tone) under different stimulation conditions. Excitatory FTCs of AC neurons were either V-shaped, closed (i.e. upper threshold) or double-peaked. Inhibitory FTCs were obtained either at both flanks or only at the low or high flank of excitatory FTCs. Application of bicuculline, an antagonist for gamma-aminobutyric acid A receptors, produced expansion of excitatory FTCs into predrug inhibitory FTCs. Inhibition of probe-elicited responses occurred when a masker was presented at certain intertone intervals. Maximal inhibition typically took place when a masker was presented within 4 ms prior to the probe. During maximal inhibition, a neuron had the minimal number of impulses and the longest response latency. Inhibition became stronger with increasing masker intensity but became weaker with increasing intertone interval. Biological significance of these data is discussed.     

蹄蝠听性脑干诱发电位特点分析

目的对野生蹄蝠进行短纯音（ToneBurst）和短声（Click）诱导听性脑干反应（ABR）分析,对其ABR特点与已有的研究进行比较,分析其中差异,加深我们对回声定位蝙蝠听觉功能的认识。方法捕捉并饲养野生蹄蝠,运用美国TDT公司（Tucker-Davis Technologies）TDT System Ⅲ诱发电位仪对蝙蝠进行短声和短纯音的听阈测定,分析其波形变化规律,确定其短纯音听阈阈值,分析听力图的特点。结果（1）蹄蝠听阈呈“W”形分布,经统计学分析各听阈区段有统计学意义（P〈0．05）。存在2个听敏区：28kHz处为最敏感的第一听敏区,阈值最低可达15dBSPL,平均29.1dBSPL;40kHz附近出现第二个听敏区,阈值最低25dBSPL,平均35．9dBSPL;在12kHz～64kHz听阈不超过55．9dBSPL。（2）短声引出的蹄蝠ABR可见4个相对稳定的波峰：波I的潜伏期为1．26ms：波Ⅱ较高且尖,潜伏期为1．94ms;波Ⅲ相对稳定,潜伏期为2．47ms;波Ⅴ潜伏期为3．40ms。随着短声强度的上升,在20～80dBSPL声强范围,各个波振幅逐渐增大、潜伏期缓慢缩短。（3）短纯音引出的蹄蝠ABR各波随刺激频率的提高而潜伏期缓慢地缩短,在64kHz以上潜伏期又逐渐延长;在28kHz、80dBSPL条件下,各波潜伏期分别为I波1．74ms,Ⅱ波2．72ms,Ⅲ波3．60ms,Ⅳ波4．58ms,Ⅴ波5．76ms。频率28kHz、声强在45～100dBSPL范围内上升时,主波振幅逐渐增大;但在部分蝙蝠出现主波振幅在达到最大后,随着声强的上升．振幅逐渐减小。（4）在高声强诱发下,短纯音和短声诱发的ABR均可见Ⅰ波分化为2个波峰,Ⅰa和Ⅰb波。（5）越接近敏感频率,ABR波形就越典型,振幅越大。结论（1）蹄蝠听闯呈“W”形分布,类似我们已知的其他回声定位蝙蝠,存在2个听敏区：28kHz处达到最敏感的第一听敏区,40kHz附近出现第二个昕敏区,在12kHz。64kHz听阈不超过55．9dBSPL。（2）从蹄蝠主波与声强、频率的关系可以看出,蝙蝠主波振幅与声强存在一定的正相关。随着接近其敏感听觉频率,波形逐渐变得更为典型,波幅也愈来愈高,反应阈值愈来愈低。但部分蹄蝠在声强高于80dBSPL时,出现振幅随声强增加而下降的趋势,提示可能与中枢某种调节性抑制有关。（3）在高声强诱发下,蹄蝠ABR可以看到Ⅰ波分化为2个波峰,Ⅰa和Ⅰb波,这可能与蝙蝠耳蜗的某些特化结构的“听黄斑”相关。