Integration time for short broad band clicks in echolocating FM-bats (Eptesicus fuscus)

Vespertilionid FM-bats (four Eptesicus fuscus and one Vespertilio murinus) were trained in an electronic phantom target simulator to detect synthetic echoes consisting of either one or two clicks. The threshold sound pressure for single clicks was around 47 dB peSPL for all five bats corresponding to a threshold energy of -95 dB re 1 Pa² * s. By varying the interclick interval, T, for double clicks it was shown that the threshold intensity was around — 3 dB relative to the threshold for single clicks at T up to 2.4 ms, indicating perfect power summation of both clicks. A threshold shift of -13.5 dB for a 1 ms train of 20 clicks (0.05 ms interclick interval) confirmed that the bats integrated the power of the stimuli. At T longer than around 2.5 ms the threshold for double clicks was the same as for single clicks. Thus, the bats performed like perfect energy detectors with an integration time of approximately 2.4 ms. This integration time is an order of magnitude shorter than that reported for bats listening passively for pure tones. In our setup the bats emitted sonar signals with durations of 2–3 ms. Hence, the results may indicate that while echolocating the bats integration time is adapted to the duration of the sonar emissions.Abbreviations AGC automatic gain control - FM frequency modulated - peSPL peak equivalent sound pressure level - rms root mean square - SD standard deviation - SE standard error of mean - T interclick interval     

The detection of phantom targets in noise by serotine bats; negative evidence for the coherent receiver

Summary Using a target simulator three serotine bats,Eptesicus serotinus, were trained to judge whether a phantom target was present or absent. The echolocation sounds emitted by the bats during the detection were intercepted by a microphone, amplified and returned by a loudspeaker as an artificial echo, with a delay of 3.2 ms and a sound level determined by the overall gain and cry amplitude. The cry level of each pulse was measured and the echo level received by the bat was calculated. The target was presented in 50% of the trials and the gain adjusted using conventional up/down procedures. Under these conditions between 40 and 48 dB peSPL were required for 50% detection (Figs. 2, 3).In a subsequent experiment the phantom target was masked with white noise (N_o) with a spectrum level of –113 dB re. 1 Pa·Hz^–1/2. The thresholds were increased by 7–14 dB. Energy density (S) of a single pulse was measured and used to estimate S/N_o, which ranged from 36–49 dB at threshold. Theoretically the coherent receiver model predicts the ratio between hits and false alarms observed for the bats at a S/N_o of ca. 1–2 dB. Since the bats require 40–50 dB higher S/N_o (Fig. 3), this is taken as negative evidence for coherent reception (cross correlation).Furthermore, a strong sensitivity to clutter was found since there seemed to exist a fixed relationship between thresholds and clutter level.Abbreviations C clutter - Nbw noise in a specified bandwidth - No noise in i Hz bandwidth - peSPL peak equivalent sound pressure level - S signal energy - SD standard deviation - Y/N Yes/No psychometry - 2AFC two alternative forced choice psychometry     

Binaural influences on Doppler shift compensation of the horseshoe bat Rhinolophus rouxi

The flying horseshoe bat Rhinolophus rouxi compensates for Doppler shifts in echoes of their orientation pulses. By lowering the frequency of subsequent calls the echo's constant frequency is stabilized at the so-called reference frequency centered in a narrow and sensitive cochlear filter. This audio-vocal behaviour is known as Doppler shift compensation. To investigate whether the bats depend on binaural cues when compensating, three animals were tested for compensation on a swing before and after unilateral deafening. In each case compensation was severely impaired by unilateral deafening. Individual animals' compensation amplitude was reduced to 28–48% of the preoperational compensation of a +1.8 kHz shift. Doppler shift compensation performance did not recover to control levels during the observed period of 24 h after surgery. In contrast, unilateral middle ear removal which induces a unilateral auditory threshold increase of 9–14 dB does not impair compensation performance on the swing. To mimick Doppler shifts in a fixed setup, the frequencies of recorded echolocation calls were experimentally shifted between 0 and +2 kHz and played back via earphones to six animals. The bats completely compensated the experimental shifts only as long as the interaural intensity difference of the playback did not exceed 20 dB. No animal compensated with monaural playback. Accepted: 27 August 1999     

Physiology and tonotopic organization of auditory receptors in the cricketGryllus bimaculatus DeGeer

Summary Physiological recordings were obtained from identified receptors in the tympanal organ ofGryllus bimaculatus. By immersing the prothoracic leg in Ringer solution and removing the anterior tympanic membrane the auditory receptors were exposed without significantly altering the frequency response of the auditory organ (Fig. 1). Each receptor was tuned to a specific sound frequency. For sound frequencies below this characteristic frequency the roll-off in sensitivity decreased from 20–30 dB/octave to 10–15 dB/octave as the characteristic frequency of receptors increased from 3–11 kHz (Fig. 4A). For each individual receptor the slope, dynamic range and maximum spike response were similar for different sound frequencies (Fig. 9A). The receptors were tonotopically organized with the characteristic frequency of the receptors increasing from the proximal to the distal end of the array (Figs. 5, 6). Several receptors had characteristic frequencies of 5 kHz. These receptors were divided into two groups on the basis of their maximum spike response produced in response to pure tones of increasing intensity (Fig. 7). Independent of the tuning of the receptor no two-tone inhibition was observed in the periphery, thus confirming that such interactions are a property of central integration.     

Control of doppler shift compensation in the greater horseshoe bat,Rhinolophus ferrumequinum

Summary FlyingRhinolophus ferrumequinum lower the frequency of the constant frequency part (f _A ) of the emitted sounds in order to compensate for Doppler shifts caused by the flight speed. The echo frequency (f _E ) is kept constant within a frequency band of about 200 Hz, the center frequency of which is about 150 Hz above the average or resting frequency (f _R ) emitted by roosting bats shortly before take off. For the compensation they use a feedback control system in which the emission frequency is changed to hold the echo frequency at a criterion value. This feedback system was demonstrated by experiments with bats flying in an experimental wind tunnel and in a He-O₂-micture. In the wind tunnelRhinolophus lowers the emission frequency in order to compensate for Doppler shifts which are caused by the ground speed flown by the bat. In the He-O₂-mixtureRhinolophus compensates for Doppler shifts which correspond to the different sound speeds in the gas mixture.I would like to thank D. R. Griffin for his generous support and stimulating criticism. I express my appreciation to the New York Zoological Society for the use of its facilities and to R. Brown for technical assistance. The work was supported by grant number GB 7155 from the National Science Foundation to the New York Zoological Society. I also thank J. D. Pye for his suggestions.     

Directionality of phase locking in auditory nerve fibers of the leopard frog Rana pipiens pipiens

Summary A dorsal approach to the eighth nerve and free-field stimulation were used to investigate the effect of sound direction and intensity on phase locking in auditory nerve fibers of the leopard frog Rana pipiens pipiens.Tuning curves of 75 auditory neurons were analyzed (Fig. 2). Amphibian papillar neurons, but not basilar papillar neurons, exhibit significant phase locking to short tone bursts at the characteristic frequency (CF), the degree of phase locking (vector strength) decreasing with the neuron's CF (Figs. 3, 4 and 10E). Vector strength increases with sound pressure level to saturate about 20 dB above threshold, while the preferred firing phase is only slightly affected (Figs. 5 and 6).In contrast, sound direction hardly affects vector strength (Figs. 7, 8, 9A and 10A and C), but has a strong influence on the preferred firing phase (Figs. 7, 8, 9B and C, 10B and D): With respect to anterior tone presentation there are phase lags for ipsilateral and phase leads for posterior and contralateral presentation. Phase differences between both ears show a sinusoidal or cardioid/ovoidal directional characteristic; maximum differences are found with antero-lateral tone presentation (Fig. 11). The directionality of phase locking decreases with the neuron's CF (Fig. 10F) and only slightly changes with sound pressure level (Fig. 12). Thus, phase locking of amphibian papilla neurons can potentially provide intensity-independent information for sound localization.Abbreviations SPL sound pressure level - FTC frequency threshold curve - CF characteristic frequency - TF test frequency - VS vector strength - AP amphibian papilla - BP basilar papilla     

The acoustic role of tracheal chambers and nasal cavities in the production of sonar pulses by the horseshoe bat, Rhinolophus hildebrandti

Summary The acoustic role of the enlarged, bony, nasal cavities and rigid tracheal chambers in the horseshoe bat,Rhinolophus hildebrandti (Fig. 2) was investigated by determining the effect of their selective filling on the nasally emitted sonar pulse and on the sound traveling backwards down the trachea.Normal sonar signals of this bat contain a long constant frequency component with most energy in the second harmonic at about 48 kHz. The fundamental is typically suppressed 20 to 30 dB below the level of the second harmonic (Fig. 1).None of the experimental manipulations described affected the frequency of the sonar signal fundamental.Filling the dorsal and both lateral tracheal chambers had little effect on the emitted vocalization, but caused the level of the fundamental component in the trachea to increase 15 to 19 dB in most bats (Table 2). When only the dorsal chamber or only the two lateral chambers were filled, the effect was less striking and more variable (Tables 3 and 4), suggesting that the tracheal fundamental is normally suppressed by acoustic interaction between these three cavities.Filling the enlarged dorsal nasal cavities had no effect on the tracheal sound. The effect of this treatment on the nasally emitted sonar pulse was inconsistent. Sometimes the fundamental increased 10 to 12 dB, other times the intensity of all harmonics decreased; in still other cases the second, third or fourth harmonic increased, but the fundamental remained unchanged (Tables 5, 6, and 7).When bats were forced to vocalize through the mouth, by sealing the nostrils, there was a prominent increase in the level of the emitted fundamental (10 to 21 dB) and in the fourth harmonic (6 to 17 dB). In one instance there was also a significant increase in the level of the third harmonic (Tables 8 and 9). The supraglottal tract thus filters the fundamental from the nasally emitted sonar signal, although the role of the inflated nasal cavities in this process is unclear.We conclude that a high glottal impedance acoustically isolates the subglottal from the supraglottal vocal tract. The tracheal chambers do not affect the emitted sonar signal, but may attenuate the fundamental in the trachea and prevent it from being reflected from the lungs back towards the cochlea. It may be important to prevent the reflected fundamental from stimulating the cochlea, via tissue conduction, along multiple indirect pathways which would temporally smear cochlear stimulation.Tracheal and nasal chambers, by suppressing the internally reflected and externally radiated components, respectively, of the laryngeal fundamental, may enable horseshoe bats to rely on the tissue-conducted fundamental as a reference or marker of its own laryngeally generated sound which could be useful in processing sonar information.     

Echolocation and obstacle avoidance in the hipposiderid batAsellia tridens

Summary The echolocation sounds of the hipposiderid batAsellia tridens consist of a constant frequency (cf) component followed by a frequency modulated (fm) terminal downward sweep of 19–21 kHz. The cf-part constitutes about 7/10 of the entire signal. In individual roosting animals the frequencies of the cf-part of consecutive sounds (resting frequency) is kept very constant but varies from bat to bat. In 18Asellia tridens resting frequencies between 111–124 kHz have been measured.The sound duration in roosting and free flying bats is between 7–10 ms. In the approach and terminal phase of bats landing on a perch or flying through obstacles, the sound duration is reduced and the repetition rate increased the nearer the bat approaches the target. At the end of the terminal phase sound durations of a minimum of 3 ms have been measured. Flying bats lower their emission frequency in order to compensate for Doppler shifts caused by the flight movement. The echofrequency is therefore kept constant about 150–200 Hz above the resting frequency.In flights through obstacles consisting of vertically stretched wires with different diameters, the bats were able to avoid wires down to a diameter of 0.065 mm whereas at 0.05 mm the percentage of flights without collisions is far below the chance level. The results demonstrate that the echolocation behavior of the hipposiderid batAsellia tridens does not differ fundamentally from that of rhinolophid bats. As a result, a new suggestion for categorization of bats producing cf-fm orientation sounds is put forward.Abbreviations cf constant frequency component - fm frequency modulated component - P probability of collision-free flights through an obstacle of ertically tretched wires - I interval between wires - D minimal diameter of a bat with folded wings; , angle at which a bat approaches an obstacle - f _A frequency of the cf-component of the emitted sound - f _E frequency of the cf-component of the echo - f _M frequency of the cf-component of the sounds recorded with the microphone - c speed of sound Supported by the Deutsche Forschungsgemeinschaft grant no. Schn 138/6-9We thank W. Hollerbach for technical assistance.     

大蹄蝠多普勒正负补偿效应的声波特征与比较

利用单摆装置模拟大蹄蝠的飞行状态并实时记录其回声定位信号,以研究其多普勒频移补偿行为。与其静息状态下的超声波特征比较,发现大蹄蝠在接近目标的过程中有多普勒正补偿效应:叫声频率随相对速度改变而成正相关变化;当相对速度最大时,其叫声频率相对静息状态频率降低最多,而相对速度为零时,叫声频率回复到静息时频率。而当大蹄蝠远离目标时,有多普勒负补偿效应:叫声频率随相对速度改变成正相关变化,叫声频率在相对速度最大时,升高最多,但相同速度下升高之值较正补偿值低得多。另外,负补偿效应出现的频率较正补偿值低,这可能是由蝙蝠生理结构的限制以及自然状态下罕见的多普勒负补偿条件所决定。     

Ontogenesis of the echolocation system in the rufous horseshoe bat,Rhinolophus rouxi (Audition and vocalization in early postnatal development)

1. The development of vocalization and hearing was studied in Sri Lankan horseshoe bats (Rhinolophus rouxi) during the first postnatal month. The young bats were caught in a nursing colony of rhinolophids in which birth took place within a two week period. 2. The new-born bats emitted isolation calls through the mouth. At the beginning these calls consisted of pure tones with frequencies below 10 kHz (Fig. 1). During the first postnatal week the call frequency increased to about 15 kHz, and the fundamental was augmented by two to four harmonics. No evoked potentials to pure tone stimuli could be elicited in the inferior colliculus of this age group, i.e., auditory processing at the midbrain level was not demonstrable. 3. Evoked potentials were first recorded in the second week, broadly tuned to 15-45 kHz, with a maximum sensitivity between 15-25 kHz. In the course of the second week, however, higher frequencies up to 60 kHz became progressively incorporated into the audiogram (Fig. 3). The fundamental frequency of the multiharmonic isolation calls, emitted strictly through the mouth, increased to about 20 kHz. 4. In the bats' third postnatal week an increased hearing sensitivity (auditory filter) emerged, sharply tuned at frequencies between 57 and 60 kHz (Fig. 4e). The same individuals were also the first to emit long constant frequency echolocation calls through the nostrils (Fig. 4c). The energy of the calls was arranged in harmonic frequency bands with the second harmonic exactly tuned to the auditory filter. These young bats continued to emit isolation calls through the mouth, which were, however, not harmonically related to the echolocation calls (Fig. 4b, d). 5. During the fourth week, both the auditory filter and the matched echolocation pulses (the second harmonic) shifted towards higher frequencies (Fig. 5). During the fifth week the fundamental frequency of the calls was progressively attenuated, and both the second harmonic of the pulses and the auditory filter reached the frequency range typical for adult bats of 73-78 kHz (Fig. 6). 6. The development of audition and vocalization is discussed with regard to possible interactions of both subsystems, and their incorporation into the active orientation system of echolocation.     

Discrimination of insect wingbeat-frequencies by the batRhinolophus ferrumequinum

Summary Five Greater Horseshoe bats,Rhinolophus ferrumequinum, were trained in a two-alternative forced-choice procedure to discriminate between artificial echoes of insects fluttering at different wingbeat rates. The stimuli were electronically produced phantom targets simulating fluttering insects with various wingbeat frequencies (Figs. 3, 4). Difference thresholds for wingbeat rates of 50 Hz and 100 Hz were determined. For an S+ of 50 Hz the difference threshold values lay between 2.8 and 4.6 Hz for individual bats; with an S+ of 100 Hz they increased to between 9.8 and 12.0 Hz (Figs. 5, 6, Table 1).Three bats, previously trained to discriminate between a S+ of 50 Hz and a S– with a lower wingbeat rate, were tested with higher frequency stimuli. When they had to decide between their old S+ of 50 Hz and either a 60 or 70 Hz echo two bats continued to select the 50 Hz stimulus while the third bat now preferred the faster fluttering insects (Table 2).During the discrimination task the echolocation behavior of the bats was monitored. When the phantom targets were presented all bats increased their duty-cycle of sound emission from about 40% to sometimes near 70%. They did so by either emitting longer echolocation calls or by increasing the sound repetition rate (Figs. 7, 8).The results show that Greater Horseshoe bats can determine the wingbeat rate of flying insects with an accuracy between 6 and 12%. Possible cues for flutter rate determination by cf-fm bats from natural and artificial insect echoes are discussed.Abbreviations DC duty-cycle - PD pulse duration - PI pulse interval - cf constantfrequency - fm frequency modulation     

Laryngeal nerve activity during pulse emission in the CF-FM bat,Rhinolophus ferrumequinum

Summary The activity of the external (motor) branch of the superior laryngeal nerve (SLN), innervating the cricothyroid muscle, was recorded in the greater horseshoe bat,Rhinolophus ferrumequinum. The bats were induced to change the frequency of the constant frequency (CF) component of their echolocation signals by presenting artificial signals for which they Doppler shift compensated. The data show that the SLN discharge rate and the frequency of the emitted CF are correlated in a linear manner.Abbreviations SLN Superior laryngeal nerve - RLN Recurrent laryngeal nerve - DCS Doppler compensation system - CF Constant frequency - FM Frequency modulation Supported by grants of the Deutsche Forschungsgemeinschaft (DFG), Az.: Schu 390/1, /2 and SFB 45We are indebted to Dipl.-Ing. H. Zöller for providing the computer programs. We want to thank H. Hahn and A. Polotzek for technical help.     

A comparative study of the physiological properties of the inner ear in Doppler shift compensating bats (Rhinolophus rouxi andPteronotus parnellii)

Summary Cochlear microphonic (CM) and evoked neural (N-1) potentials were studied in two species of Doppler shift compensating bats with the aid of electrodes chronically implanted in the scala tympani. Potentials were recorded from animals fully recovered from the effects of anesthesia and surgery. InPteronotus p. parnellii andRhinolophus rouxi the CM amplitude showed a narrow band, high amplitude peak at a frequency about 200 Hz above the resting frequency of each species. InPteronotus the peak was 25–35 dB higher in amplitude than the general CM level below or above the frequency of the amplitude peak. InRhinolophus the amplitude peak was only a few dB above the general CM level but it was prominent because of a sharp null in a narrow band of frequencies just below the peak. The amplitude peak and the null were markedly affected by body temperature and anesthesia. InPteronotus high amplitude CM potentials were produced by resonance, and stimulated cochlear emissions were prominent inPteronotus but they were not observed inRhinolophus. InPteronotus the resonance was indicated by a CM afterpotential that occurred after brief tone pulses. The resonance was not affected by the addition of a terminal FM to the stimulus and when the ear was stimulated with broadband noise it resulted in a continual state of resonance. Rapid, 180 degree phase shifts in the CM were observed when the stimulus frequency swept through the frequency of the CM amplitude peak inPteronotus and the frequency of the CM null inRhinolophus. These data indicate marked differences in the physiological properties of the cochlea and in the mechanisms responsible for sharp tuning in these two species of bats.     

Ontogenesis of auditory fovea representation in the inferior colliculus of the Sri Lankan rufous horseshoe bat,Rhinolophus rouxi

Summary This report describes the ontogenesis of tonotopy in the inferior colliculus (IC) of the rufous horseshoe bat (Rhinolophus rouxi). Horseshoe bats are deaf at birth, but consistent tonotopy with a low-to-high frequency gradient from dorsolateral to ventromedial develops from the 2nd up to the 5th week. The representation of the auditory fovea is established in ventro-mediocaudal parts of the IC during the 3rd postnatal week (Fig. 3). Then, a narrow frequency band 5 kHz in width, comprising 16% of the bat's auditory range, captures 50–60 vol% of the IC (Fig. 3c). However, foveal tuning is 10–12 kHz (1/3 octave) lower than in adults; foveal tuning in females (65–68 kHz) is 2–3 kHz higher than in males (62–65 kHz). Thereafter, foveal tuning increases by 1–1.5 kHz per day up to the 5th postnatal week, when the adult hearing range is established (Figs. 4, 5). The increase of sensitivity and of tuning sharpness of single units also follows a low-to-high frequency gradient (Fig. 6).Throughout this development the foveal tuning matches the second harmonic of the echolocation pulses vocalised by these young bats. The results confirm the hypothesis of developmental shifts in the frequency-place code for the foveal high frequency representation in the IC.Abbreviations BF best frequency - CF constant frequency - FM frequency modulation - IC inferior colliculus - IHC inner hair cell; - OHC outer hair cell - RR Rhinolophus rouxi     

Responses and song pattern copying of Omega-type I-neurons in the cricket,Gryllus bimaculatus,at different prothoracic temperatures

Summary Omega-type I-neurons (ON/1) (Fig. 1A) were recorded intracellularly with the prothoracic ganglion kept at temperatures of either 8–9°, or 20–22° or 30–33 °C and the forelegs with the tympanal organs kept at ambient temperature (20–22 °C). The neurons were stimulated with synthetic calling songs (5 kHz carrier frequency) with syllable periods (SP in ms) varying between 20 and 100, presented at sound intensities between 40 and 80 dB SPL. The amplitude and duration of spikes as well as response latency decreased at higher temperatures (Figs. 1 B, 2, 6). At lower prothoracic temperatures (8–9 °C) the neuron's responses to songs with short SP (20 ms) failed to copy single syllables, or with moderate SP (40 ms) copied the syllable with low signal to noise ratio (Fig. 3). The auditory threshold of the ON/1 type neuron, when tested with the song model, was temperature-dependent. At 9° and 20 °C it was between 40 and 50 dB SPL and at 33 °C it was less than 40 dB SPL (Fig. 4). For each SP, the slope of the intensity-response function was positively correlated with temperature, however, at low prothoracic temperatures the slope was lower for songs with shorter SPs (Fig. 5). The poor copying of the syllabic structure of the songs with short SPs at low prothoracic temperatures finds a behavioral correlate because females when tested for phonotaxis on a walking compensator responded best to songs with longer SPs at a similar temperature.Abbreviations epsps excitatory postsynaptic potentials - ON/1 omega-type I-neuron - SP syllable period - SPL sound pressure level     

Effect of destruction of the inferior colliculus on function of the echolocation system in horseshoe bats

The effect of unilateral and bilateral destruction of the inferior colliculus on the sensitivity of the auditory system, on parameters of the sonor signals, and on Doppler shift compensation in echo signals was studied in experiments on horseshoe bats (Rhinolophus ferrum-equinum). The results show that complete bilateral destruction of the inferior colliculus in bats does not lead to total disturbance of function of the auditory system but it sharply reduces the sensitivity of that system, as shown by a decrease in the maximal obstacle detection range and inability to respond to an insect emitting a feeble sound. It can also be concluded that the inferior colliculus plays a direct part in maintenance of the emission frequency and that different parts of the inferior colliculus play different roles in this process. The Doppler shift compensation effect requires preservation of the integrity of not less than half of the central nucleus of at least one inferior colliculus.A. A. Ukhtomskii Physiological Institute, A. A. Zhdanov State University, Leningrad. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 375–381, July–August, 1980.     

Classification of insects by echolocating greater horseshoe bats

Summary Echolocating greater horseshoe bats (Rhinolophus ferrumequinum) detect insects by concentrating on the characteristic amplitude- and frequency modulation pattern fluttering insects impose on the returning echoes. This study shows that horseshoe bats can also further analyse insect echoes and thus recognize and categorize the kind of insect they are echolocating.Four greater horseshoe bats were trained in a twoalternative forced-choice procedure to choose the echo of one particular insect species turning its side towards the bat (Fig. 1). The bats were able to discriminate with over 90% correct choices between the reward-positive echo and the echoes of other insect species all fluttering with exactly the same wingbeat rate (Fig. 4).When the angular orientation of the reward-positive insect was changed (Fig. 2), the bats still preferred these unknown echoes over echoes from other insect species (Fig. 5) without any further training. Because the untrained bats did not show any prey preference, this indicates that the bats were able to perform an aspect-anglein-dependent classification of insects.Finally we tested what parameters in the echo were responsible for species recognition. It turned out that the bats especially used the small echo-modulations in between glints as a source of information (Fig. 7). Neither the amplitudenor the frequencymodulation of the echoes alone was sufficient for recognition of the insect species (Fig. 8). Bats performed a pattern recognition task based on complex computations of several acoustic parameters, an ability which might be termed cognitive.Abbreviations AM amplitude modulation - CF constant frequency - FM frequency modulation - S+ positive stimulus - S- negative stimulus     

Correlation between auditory thalamic area evoked responses and species-specific call characteristics

Evoked potentials were recorded from the posterior dorsal thalamus of green treefrogs (Hyla cinerea) in response to single tones and combinations of two and three tones. 1. The responses to two tones were largest when one of the component tones was 500 Hz and when the second component was between 2000 and 4000 Hz (Fig.3). 2. The response to 500 + 3000 Hz showed nonlinear facilitation; i.e., the amplitude of the response was greater than the sum of the responses to the component tones alone (Figs. 4, 5). This result provides evidence that cells functioning as 'AND' gates will be found in this center. 3. When a third tone around 1200 Hz was added to a stimulus of 500 + 3000 Hz a 65% decrease in the evoked response amplitude occurred (Fig. 6). 4. The largest evoked response amplitude to a two-tone stimulus (500 + 3000 Hz) occurred when the rise-time was less than 50 ms (Fig. 7). 5. The two-tone tuning was found to be temperature dependent. The optimal lower frequency tone shifted downward with decreasing temperatures (Fig. 8). 6. When the temperatures of the neurophysiological and the behavioral experiments are matched, the optimal stimuli for evoking a large response are closely correlated to the parameters of the acoustic stimuli preferred by gravid H. cinerea females in discrimination tests. This center therefore appears to be very important for the processing of complex species-specific sounds.