Time course of rate responses to two-tone stimuli in auditory nerve fibres in the guinea pig.

Peri-stimulus time histograms (PSTHs) were constructed from responses of auditory nerve fibres in anaesthetized guinea pigs. Acoustic stimuli consisted of pure tones, presented either as tone bursts, or in two-tone combinations in which a gated test tone was superimposed on a continuous excitatory tone at characteristic frequency (CF). The majority of the sample of fibres displayed two-tone rate suppression (2TRS). The suppression was either a monotonic or a non-monotonic function of the level of the superimposed test tone. Monotonic suppression of CF-driven rate occurred only for test tones at frequencies higher than CF, presented at levels up to the maximum available (approx. 100 dB SPL). For test tones below CF, 2TRS initially increased, then reverted towards excitation for higher levels of the test tone. Three levels were identified in non-monotonic, two-tone rate functions; (1) the threshold for rate suppression, (2) the maximally suppressing level and (3) the level (referred to as the balance point) at which average firing rate was restored to the background, CF-driven rate. PSTHs for two-tone responses obtained for test tone levels between the maximally-suppressing level and the balance point typically showed brief decrements (notches) in spike rate, at the onset and following the offset of the test tone. The latency, depth and duration of notches, however, depended on the level of the test tone, in a different manner for onset and offset. In some cases, without overt rate excitation above the probe-driven rate, the offset notch became more pronounced and of extended duration with increased level of the test tone, suggestive of adaptation to the test tone. Two-tone responses, in which rate exceeded the background, CF-driven rate, in general were preceded by a reduced onset notch and were followed by a longer-lasting depression of the background spike rate, typical of post-excitatory depression. Relative to responses obtained to the test tones presented alone, excitatory two-tone responses were of lower rate and were delayed by the onset notch. Onset notches sometimes preceded rate excitation in responses to single tones. Some features of the time course of rate suppression and excitation displayed in PSTHs for responses to one and two-tone stimuli seem inconsistent with current models of 2TRS.     

Two-tone rate suppression in auditory-nerve fibers: dependence on suppressor frequency and level

The growth of two-tone rate suppression with suppressor level was studied for auditory-nerve fibers in anesthetized cats. The level of a tone at the characteristic frequency (CF) was adjusted by an adaptive procedure (PEST) so that, when presented with a suppressor tone, the CF tone would produce a criterion discharge rate. Suppression (in dB) was defined as the CF-tone level that met criterion in the presence of a suppressor minus the level that met criterion in quiet. The growth of suppression with suppressor level was well characterized by a straight line whose slope (in dB-excitor/dB-suppressor) varied with suppressor frequency by as much as a factor of 10 in the same fiber. These slope differences were systematically related to the position of the suppressor frequency relative to the fiber CF: for below-CF suppressors, slopes ranged from 1 to 3 dB/dB, while, for above-CF suppressors, they were between 0.15 and 0.7 dB/dB. Slopes decreased rapidly with increasing suppressor frequency near the CF, but, for frequencies well below the CF, the slope reached a maximum that increased gradually with CF. These results resemble psychophysical data on the growth of masking and psychophysical suppression, and pose difficulties for existing models of two-tone suppression.     

Neural correlates of cubic difference tones in the medial geniculate body of the cat

Single unit responses to the cubic difference tone CDT (2f₁ ? f₂ = CF) and the difference tone DT (f₂ ? f₁ = CF) were studied in the medial geniculate body (MGB) of the cat. Out of 66 units tested with CDT stimuli and having characteristic frequencies (CF) below 10 kHz, 77% gave a response to the two-tone combination stimulus. The component tones when presented alone evoked no responses, or in some cases a response pattern that was different from the one observed for the combination tone. The CDT response pattern was always similar to that seen for a pure tone at the CF. The threshold of response for the CDT was 10–70 dB higher than for a pure tone stimulus at the CF.The few units which were phase-locked could be synchronised with the CF, CDT, or DT, depending on the particular stimulus conditions. The index of synchrony was in many cases found to be higher for CDT responses than for a pure tone at CF.     

Lateral inhibition in the anteroventral cochlear nucleus of the cat: a microiontophoretic study

The spontaneous firing rates of non-prepotential (NPP) units of the anteroventral cochlear nucleus are quite low so it has not been possible to determine whether side band tones are inhibitory when presented alone. Microiontophoretically-applied excitatory amino acids can be used to excite non-spontaneous cells directly. Using this technique it can be shown that side band tone bursts 1/2 to 3/4 octave above the characteristic frequency (CF) of a NPP unit inhibit the amino acid-induced firing. Side band tones which inhibited the amino acid-induced firing were beyond the tuning curve. Side band tones within the tuning curve produced excitation. Both, however, usually reduced the activity evoked by a CF tone burst (i.e., two-tone interaction). The data suggests that lateral inhibition and two-tone interactions are separate phenomena in the auditory system and that lateral inhibition may play a critical role in determining the shape of the tuning curve of NPP units.     

Growth of pulsation threshold of a suppressed tone as a function of its level

The pulsation threshold (PT) was measured at the frequency of a probe tone in a two-tone stimulus. A suppressor tone was higher in frequency than the probe tone and was fixed in level. As the level of the probe tone was increased, three regions of performance were observed: (1) for probe tone levels below simultaneous masked threshold (SMT), PT was the same as that measured for the suppressor alone, (2) for levels above SMT, PT increased linearly with level, indicating a constant amount of suppression in dB, and (3) for higher levels a recruitment-like phenomenon was observed, in which the PT increased faster than the probe level. The maximum amount of suppression observed was equal to the difference between the PT and SMT for the suppressor alone. One interpretation is that the suppressor reduces excitation on the slopes of its own excitation pattern by the same amount that it reduces the additional excitation from a probe tone. These results are consistent with physiological data, where the amount of suppression is determined by the suppressor and is independent of the level of the probe tone.     

Induced suppression in spike responses to tone-on-noise stimuli in the auditory nerve of the pigeon

Spike potentials were recorded from single, afferent fibres in the pigeon auditory nerve. Pure-tone stimuli were presented in quiet and in combination with wide band noise. Presented alone, tones produced tuned response areas; noise generally drove spike rate to well above the spontaneous rate measured in quiet. When presented in combination with noise, tones up to 75 dB SPL at frequencies far from the fibre's response area had no effect on the noise-driven spike rate. As the tone frequency was shifted towards the response area, from above or below CF, suppression of the noise-driven spike rate became stronger until the tone reached the edge of the response area. Suppression of the noise-driven rate was directly proportional to the level of the tone. Within the area of response to the tone, tone-driven spike rates generally were unchanged or variably decreased (occasionally slightly increased) by tone-on-noise stimulation, depending on the relation of the tone frequency to CF and the level of the tone relative to that of the noise. Tuning properties were unaffected. It is suggested that in the pigeon, the suppression of driven spike rate during presentation of combination stimuli, which is common to all fibres, depends on the same mechanism as the suppression of spontaneous firing by tones that is observed in a proportion of fibres (Temchin, A.N. (1988), J. Comp. Physiol. A 163, 99-115; Hill et al., (1989) Hear. Res. 39, 37-48).     

Interaural delay sensitivity to tones and broad band signals in the guinea-pig inferior colliculus

We have measured the sensitivity of 243 low-frequency cells in the central nucleus of the guinea pig to the interaural time delay of best frequency (BF) tones, wideband noise and synthetic vowels. The highest rate of firing for the majority of cells occurred when the stimulus to the contralateral ear arrived 100–400 μs before that to the ipsilateral ear. The best delays for tones and noise measured in the same cell were highly correlated. In contrast to the tone delay functions, the majority of the delay functions obtained in response to wideband signals did not cycle, but were characterized by a single dominant peak or trough. The response frequency calculated from the delay functions to the vowel often did not correspond to the unit's BF, suggesting that the unit was responding to a component close to the first formant frequency (730 Hz) of the vowel. Phase-locked responses, on the other hand, only occurred to the fundamental frequency of the vowel (100 Hz) and not to higher frequency components. The responses to delayed tone and noise signals in the guinea pig are very like those obtained in the cat and other mammals. The similarity of the range of best delays for the guinea-pig with those reported for the cat, despite the difference in head size in these two species, suggests that the sensitivity to interaural delays reflects the properties of the binaural pathways rather than an adaptation to the delays normally experienced by the animal.     

Auditory nerve rate-level functions for two-tone stimuli: possible relation to basilar membrane nonlinearity

Rate versus level functions are presented for auditory-nerve fiber responses to best frequency (BF) tones presented alone and in the presence of a constant level suppressor tone. The paper focuses on units with sloping saturation rate functions for BF tones presented alone. Two-tone rate functions are generally parallel to BF functions at firing rates below those in the sloping saturation region of the BF function. At rates in the sloping saturation range the two-tone function grows with a greater slope than that of the BF function until the two functions meet. This result is discussed in terms of current knowledge of two-tone suppression on the basilar membrane.     

Effects of excitatory and non-excitatory suppressor tones on two-tone rate suppression in auditory nerve fibers

Recordings were obtained from individual auditory nerve fibers in anesthetized chinchillas. Rate versus level functions were obtained for best frequency (BF) tones alone and for simultaneously-gated tone pairs comprising a BF tone and a second tone at a fixed intensity that produced evidence of two-tone rate suppression. Care was taken in selecting a range of suppressor tone levels that included excitatory (i.e., the suppressor tone evoked a rate change by itself) and non-excitatory (i.e., no suppressor tone-evoked rate increase) suppressor tone levels. Addition of a suppressor tone produced a shift of the dynamic range portion of the BF rate versus level function to higher test intensities. A parallel shift of the dynamic range portion of the rate versus level function was associated with a non-excitatory suppressor tone. The shift produced by an excitatory suppressor tone was characterized by a decrease in slope. Results indicated that the magnitude of shift increased monotonically as suppressor tone intensity was raised and that there was a gradual transition from a non-excitatory response to an excitatory response. The rate of shift (i.e., dB of shift per dB change in suppressor tone intensity) did not differ for non-excitatory versus excitatory responses, but was substantially greater for below-BF suppressor tones (1.38 dB/dB) than for above-BF suppressor tones (0.54 dB/dB). The rate of shift did not, however, appear to be related systematically to suppressor tone frequency separation from BF. Above- and below-BF suppression was noted for fibers over the range of best frequencies tested (110 Hz to 16.4 kHz).     

On the prediction of sweep rate and directional selectivity for FM sounds from two-tone interactions in the inferior colliculus

Two-tone stimuli have traditionally been used to reveal regions of inhibition in auditory spectral receptive fields, particularly for neurons with low spontaneous rates. These techniques reveal how different frequencies excite or suppress the response to an excitatory frequency of a cell, but have often been assessed at a fixed masker–probe time interval. We used a variation of this methodology to determine whether two-tone spectrotemporal interactions can account for rate-dependent directional selectivity for frequency modulations (FM) in the mustached bat inferior colliculus (IC). First, we quantified the response to upward and downward sweeping, linear, fixed-bandwidth FM tones centered at a unit’s characteristic frequency (CF) at 6 sweep durations ranging from 2 to 64 ms. Then, to examine how responses to instantaneous frequencies contained within the sweeps might interact in time, we varied the frequency and relative onset of a brief (4 ms) “conditioner” tone paired with a fixed 4-ms CF probe tone. We constructed “conditioned response areas” (CRA) depicting regions of suppression and facilitation of the probe tone caused by the conditioning tone. We classified the CRAs as predominantly excitatory (40.9%), inhibitory (22.7%), or mixed (36.4%). To generate FM response predictions, the CRAs were multiplied with spectrograms of the same sweeps used to assess response to FM. The predictions of FM rate and directionality were accurate by our criteria in approximately 20% of units. Conversely, the CRAs from the remaining units failed to predict FM responses as accurately, suggesting that most IC units respond differently to FM sweeps than they do to tone-pairs matched to the instantaneous frequencies contained in those sweeps. The implications of these results for models of FM directionality are discussed.     

Distortion in the cochlea: Acoustic f2−f1 at low stimulus levels

The stimulus level and frequency dependence of the quadratic difference tone (QDT) measured as an otoacoustic emission in the ear canal has been investigated in the guinea pig and compared with simultaneously measured cubic difference tone (CDT) and with the round window electrical response. Acoustic QDT level tended to be highly labile. Growth of the ear canal response with covaried stimuli was very gradual (slope < 0.5). Acoustic and CM responses showed similar behaviour when f₂ alone was incremented. The QDT was strongly dependent on stimulus frequency separation for high frequency stimuli. It is suggested that, at low stimulus levels and high frequencies, the acoustic QDT may originate in the ‘tonic’ motile responses of outer hair cells as they follow the envelope of the two-tone stimulus.     

Origin of suppression of otoacoustic emissions evoked by two-tone bursts

Otoacoustic emission (OAE) data recorded for tone bursts presented separately and as a two-tone burst complex, that had been reported previously [Yoshikawa, H., Smurzynski, J., Probst R., 2000. Suppression of tone burst evoked otoacoustic emissions in relation to frequency separation. Hear. Res. 148, 95-106], were re-processed using the method of adaptive approximations by matching pursuit (MP). Two types of stimuli were applied to record tone burst OAEs (TBOAEs): (a) cosine-windowed tone bursts of 5-ms duration with center frequencies of 1, 1.5, 2 and 3kHz, (b) complex stimuli consisting of a digital addition of the 1-kHz tone burst together with either the 1.5-, 2- or 3-kHz tone burst. The MP method allowed decomposition of signals into waveforms of defined frequency, latency, time span, and amplitude. This approach provided a high time-frequency (t-f) resolution and identified patterns of resonance modes that were characteristic for TBOAEs recorded in each individual ear. Individual responses to single-tone bursts were processed off-line to form 'sum of singles' responses. The results confirmed linear superposition behavior for a frequency separation of two-tone bursts of 2kHz (the 1-kHz and 3-kHz condition). For the 1, 1.5-kHz condition, the MP results revealed the existence of closely positioned resonance modes associated with responses recorded individually with the stimuli differing in frequency by 500Hz. Then, the differences between t-f distributions calculated for dual (two-tone bursts) and sum-of-singles conditions exhibited mutual suppression of resonance modes common to both stimuli. The degree of attenuation depended on the individual pattern of characteristic resonance modes, i.e., suppression occurred when two resonant modes excited by both stimuli overlapped. It was postulated that the suppression observed in case of dual stimuli with closely-spaced components is due to mutual attenuation of the overlapping resonance modes.     

Sensitivity of neurons in cat primary auditory cortex to tones and frequency-modulated stimuli. II: Organization of response properties along the 'isofrequency' dimension.

The spatial distribution of neuronal responses to tones and frequency-modulated (FM) stimuli was mapped along the 'isofrequency' dimension of the primary auditory cortex (AI) of barbiturate-anesthetized cats. In each cat, electrode penetrations roughly orthogonal to the cortical surface were closely spaced (average separation approximately 130 microns) along the dorsoventral extent of a single 'isofrequency' strip in high frequency parts of AI (> 15 kHz). Characteristic frequency (CF), minimum threshold, sharpness of frequency tuning (Q10 and Q20), the dynamic range of the spike count-intensity function at CF, sensitivity to the rate of change of frequency (RCF) and to the direction of frequency-modulation (DS) were determined for contralaterally-presented tone and FM stimuli. Sharpness of tuning attained maximum values at central loci along the dorsoventral 'isofrequency' axis and values declined towards more dorsal and more ventral locations. Minimum threshold and dynamic range varied between high and low values in a similar and correlated periodic fashion. Their combined organization yielded an orderly spatial representation of response strength, relative to maximum, as a function of stimulus amplitude. The distributions of the most common forms of FM rate sensitivity (RCF response categories) and best RCF along 'isofrequency' strips were significantly non-random although there was a considerable degree of variability between cats. FM directional preference and sensitivity appeared to be randomly distributed. Sharpness of tuning may be related to the analysis of the spectral content of an acoustic stimulus, both minimum threshold and dynamic range are related to the encoding of stimulus intensity, and measures of FM rate and directional sensitivity assess the coding of temporal changes of stimulus spectra. The independent, or for minimum threshold and dynamic range dependent, topographic organizations of these neuronal parameters therefore suggest parallel and independent processing of these aspects of acoustic signals in AI.     

Suppression of tone burst evoked otoacoustic emissions in relation to frequency separation

Tone burst evoked otoacoustic emissions (TBEOAEs) were measured for two tone bursts presented separately and as a two-tone burst complex to examine the linearity of TBEOAE generators for different frequency separations of the stimuli. The stimuli were: (a) tone bursts of 5-ms duration and center frequencies of 1, 1.5, 2 and 3 kHz; (b) complex stimuli with the 1-kHz tone burst combined digitally with each of the other specified tone bursts. Signals were delivered at 70 dB SPL using a non-linear processing method and at 60 dB SPL using a linear method to 21 ears of normally hearing adults. Spectra of TBEOAEs obtained with single-tone bursts were superimposed (composite) and compared to those of the two-tone burst complex. A close correspondence between the composite and complex spectra was present in all ears. However, the components on the higher-frequency slope of the 1-kHz spectral peak were reduced in the complex spectra obtained with a frequency separation of 0.5 kHz when compared to the corresponding composite spectra. The reduction was greater at a stimulus level of 70 dB SPL than with 60 dB SPL. The effect was smaller for a frequency separation of 1 kHz, and almost absent for the tone burst separation of 2 kHz. Thus, suppression leads to weak non-linear frequency superposition for higher-level, closely spaced stimuli.     

Delay analysis in the auditory brainstem of the rat: comparison with click latency

Many cells in the auditory brainstem 'phase lock' to tone stimuli. From the changing phase relationship between the stimulus and the neural response in phase-locking cells, the delay between them can be estimated. This delay, however, is consistently greater than the latency measured in response to click stimuli, an important discrepancy. In this paper the different measures of delay, namely phase delay, group delay and signal-front delay are re-examined. An improved method for computing the average group delay is presented, which accounts for the cyclical nature of the phase data. Data were collected from units in successive processing sites of auditory pathway: the auditory nerve, the cochlear nucleus, the trapezoid body and the medial nucleus of the trapezoid body. Low-characteristic frequency (CF) units gave multimodal post-stimulus-time histograms in response to clicks, and showed stepwise decreases in latency with increasing intensity, with the appearance of earlier peaks in the response, rather than shifts in the timing of the peaks. The separation of peaks corresponded to the inverse of the unit's CF. High-CF units also showed a decline in click latency with intensity, but to a lesser degree than low CF units. We present an analysis which explains the difference between click latency and delay, and which in contrast to previous accounts is experimentally testable. We demonstrate that this new framework accounts for the discrepancy between the two measures of delay, and in addition accounts for the observed stepwise shifts in click latency for low-CF units.     

Patterns of inhibition in auditory cortical cells in awake squirrel monkeys

Two-tone interactions are recorded in the responses of single units in the superior temporal gyrus to contralateral acoustic stimulation of the awake squirrel monkey. Four response types are distinguished based primarily on the nature of the inhibitory responses elicited by two-tone stimuli, and secondarily on such criteria as the patterns of response to single tones and noise stimuli, thresholds, and spontaneous activity levels. Type A units display strong lateral inhibitory influences which may extend up to 2 octaves on either side, or both sides, of the BF. They are sharply tuned at all intensities and exhibit sustained response to single tone stimuli at the BF. The units have nonmonotonic rate-level functions, and show little or no response to broad band noise. Type A units have low spontaneous rates (less than 3 spikes/s) and relatively high thresholds (greater than or equal to 30 dB SPL). Type B units are characterized by relatively high spontaneous rates of activity (greater than 20 spikes/s) and inhibitory responses to single tone stimuli. Broad band noise may evoke strong excitatory response. Type C units summate the responses to the two-tone stimulus, and show little or no inhibitory influences. They have V-shaped tuning curves, monotonic rate-level functions, low thresholds (greater than or equal to 30 dB SPL), moderate spontaneous rates (ca. 10 spikes/s), and a strong and sustained response to noise and single tone stimuli. Type D units show 'temporal inhibition' to two-tone stimuli, in that an excitatory response to the first tone suppresses (adapts or inhibits) the response to the second tone. These units generally have moderate to broad frequency tuning and phasic responses to single tone stimuli. Histological examination of electrode tracks suggests that Type A units are restricted to A1 (and possibly the rostral field) while other types are distributed over all auditory fields.     

Excitation and suppression of primary auditory fibres in the pigeon

Spike potentials were recorded from single fibres in the auditory nerve of the pigeon. In fibres with recognizable responses to sound, spontaneous activity and properties of responses to tonal stimuli were studied in quiet background conditions. Mean spontaneous rate in the sample of fibres was 35 spikes/s. Tuning of spike response to tones was manifest as a single peak in rate at each sound pressure level (SPL) in the frequency-intensity plane. The majority of fibres showed only excitation of spike rate above spontaneous rate. Post stimulus time histograms (PSTs) in such cases were typical of excitatory responses, previously described in birds and mammals showing pronounced adaptation and post-stimulus suppression of spike rate. In most cases of excitation-only responses, however, slopes of rate functions depended on stimulus frequency. Close to characteristic frequency (CF), slopes tended to decrease with increasing SPL, whereas away from CF, slopes tended to increase with SPL. In a minority of excitation-only responses, slopes of rate functions were parallel. In some fibres, tones adjacent to the response area caused overt suppression of spontaneous firing. For these fibres, the slopes of rate functions were more-strongly frequency-dependent, being negative at low SPL when rate suppression occurred. Suppression of spontaneous activity at low SPL was non-monotonic and quite different from suppression of spike rate at stimulus intensities above rat saturation. In PSTs of suppressed spontaneous activity, rebound occurred at the termination of the tone. The results clarify previous observations of suppression of primary auditory responses in birds. We conclude that responses in the majority of auditory fibres in the pigeon are the product of opposing excitatory and suppressive influences in the cochlea, generated by single tones in quite.     

Masking and suppression in auditory nerve fibers of the goldfish, Carassius auratus.

The responses of single fibers of the auditory nerve of the goldfish (Carassius auratus) were recorded in response to two tones of different duration (20 ms 'signals' and 200 ms 'maskers') presented simultaneously or non-simultaneously. A single tone may produce excitation, adaptation, and suppression in auditory nerve fibers. For fibers with characteristic frequencies (CF) in the 200 to 400 Hz range, frequencies well above CF tend to produce suppression. If the net response to the masker tone is excitation, an added excitatory signal tone tends to increment the response in a way predictable from the rate-level function for the masker. A masker can attenuate the response to a signal as a result of a compressive and saturating response to the masker, and as a result of a low signal-to-masker ratio. If the net response to a masker tone is suppression, it effectively subtracts from signal excitation, causing 'suppressive masking.' In non-spontaneous fibers, suppression, additive excitatory effects, and adaptation can be revealed by responses to the signal in the absence of spike responses to the masker. In general, the ability of one tone (the masker) to reduce the response to a second tone (the signal) is greater in non-spontaneous fibers than in spontaneous fibers. These results also show that estimates of the frequency selectivity of many goldfish auditory nerve fibers will depend on whether the response of the fiber is defined by excitation, suppression, or both. The response of many fibers with CF in the 200-400 Hz region, as defined by excitation, can be masked or suppressed by a broad range of frequencies covering the effective hearing range of the goldfish.     

Responses of inferior colliculus neurons to SAM tones located in inhibitory response areas

In order to examine the effect of inhibition on processing auditory temporal information, responses of single neurons in the inferior colliculus of the chinchilla to sinusoidally amplitude-modulated (SAM) tones alone and the presence of a steady-state tone were obtained. The carrier frequency of the SAM tone was either the characteristic frequency (CF) or a frequency in the inhibitory response area of a studied neuron. When the carrier frequency was set to the neuron’s CF, neurons responded in synchrony to the SAM-tone envelope, as expected. When the carrier frequency was set to a frequency at which pure tones produced inhibition, SAM tones elicited little or no response, also as expected. However, when the same SAM tone was paired with a pure tone whose frequency was set to the neuron’s CF, responses synchronized to the SAM tone envelope were obtained. These modulated responses were typically one-half cycle out-of-phase with the response to the SAM tone at CF, suggesting that they arose from cyclic inhibition and release from inhibition by the SAM tone. The results demonstrate that the representation of temporal information by inferior colliculus neurons is influenced by temporally-patterned inhibition arising from locations remote from CF.     

Adjustable frequency selectivity of auditory forebrain neurons recorded in a freely moving songbird via radiotelemetry

One of the hearing system's basic properties that determines the detection of signals is its frequency selectivity. In the natural environment, a songbird may achieve an improved detection ability if the neuronal filters of its auditory system could be sharpened to adapt to the spectrum of the background noise. To address this issue, we studied 35 multi-unit clusters in the input layer of the primary auditory forebrain of nine European starlings (Sturnus vulgaris). Microelectrodes were chronically implanted in this songbird's cortex analogue and the neuronal activity was transmitted from unrestrained birds via a miniature FM transmitter. Frequency tuning curves (FTCs) and inhibitory sidebands were determined by presenting a matrix of frequency-level combinations of pure tones. From each FTC, the characteristic frequency (CF) and several parameters describing the neurons' filter characteristics were derived and compared to the same recording site's filter function while simultaneously stimulating with a continuous CF tone 20 dB above the response threshold. Our results show a significant improvement of frequency selectivity during two-tone stimulation, indicating that spectral filtering in the starling's auditory forebrain depends on the acoustic background in which a signal is presented. Moreover, frequency selectivity was found to be a function of the time over which the stimulus persisted, since FTCs were much sharper and inhibitory sidebands were largely expanded several milliseconds after response onset. Neuronal filter bandwidths during two-tone stimulation in the auditory forebrain are in good agreement with psychoacoustically measured critical bandwidths in the same species. Radiotelemetry proved to be a powerful tool in studying neuronal activity in freely behaving birds.