Brain projections and information processing of biologically significant sounds by two large ventral-cord neurons ofGryllus bimaculatus DeGeer (Orthoptera,Gryllidae)

Summary Two ventral-cord neurons in the auditory system ofGryllus bimaculatus were studied electrophysiologically by stimulation with pulses of sound at a single frequency (sine-wave pulses), stridulatory songs, and artificial sounds constructed to imitate the conspecific songs. The sine-wave pulses were varied in frequency, sound intensity, duration, and repetition rate. The stridulatory songs were the conspecific calling, aggressive, and courtship songs and the calling songs of 8 sympatric gryllids (played back at different sound intensities). The artificial songs were varied in carrier frequency, pulse rate, chirp rate, and sound intensity.The LF₁ neuron precisely duplicates the temporal structure of the conspecific calling (and aggressive) song over the whole intensity range (Figs. 7, 8, 10). It is sharply tuned to the carrier frequency of the song (5 kHz) and shows little or no response above 10 kHz and below 3 kHz (Figs. 1, 2). By variation of the calling song's temporal structure it can be demonstrated that the LF₁ neuron is particularly suited to respond to the pulse duration and the pulse and chirp repetition rates of this song pattern (Figs. 6, 9).On the other hand, the HF₁ neuron is a broad-band neuron with a maximal sensitivity at 16 kHz (Figs. 1, 4); it is tuned to the conspecific courtship song with respect to carrier frequency, the short pulse duration, and the very low pulse repetition rate (Figs. 6, 7, 8).The results demonstrate that the two ventral-cord neurons represent highly evolved channels of the auditory pathway in gryllids, each of which transmits important features of the corresponding conspecific songs to several areas of the brain (Fig. 11). But they are not ideal filters for these conspecific songs, since they also respond to many other sound signals (Fig. 10).Supported by the Deutsche Forschungsgemeinschaft as part of the program Sonderforschungsbereich 114 (Bionach), BochumUnder the auspices of the scientist exchange program of the Deutsche Forschungsgemeinschaft and the Academy of Sciences, USSRWe thank Prof. Dr. Schwartzkopff for his help and support; it was due to his initiative and organization that this work could be done in collaboration between the Sechenov Institute, Leningrad, and the Lehrstuhl für Allgemeine Zoologie, Ruhr University, Bochum. We are grateful to Mrs. I. Klotz and Mrs. B. Brücher for technical assistance.     

Auditory nerve and interneurone responses to natural sounds in several species of cicadas

ABSTRACT. The calling and courtship songs of 17-year cicadas and of Say's cicadas differ both in the sound frequency spectrum and in temporal pattern. Multiunit recordings with hook electrodes from the whole auditory nerve show that the hearing organs are especially sensitive to transient stimuli occurring in natural sounds. Artificially produced clicks elicit bursts of spikes synchronized among various primary sensory fibres. These fibres respond to natural calling and courtship songs with a specificity dependent on carrier frequency, rhythm and transient content of the sound, following sound pulses (i.e. tymbal actions) up to repetition rates of 200 Hz. An ascending, plurisegmental interneurone was characterized by intracellular recording and simultaneously stained with cobalt. Its main arborization spatially overlaps the anterior part of the sensory auditory neuropile, and the axon was traced as far as the prothoracic ganglion. Direct input from primary auditory fibres was suggested by latency measurements. Intracellular recordings from such neurons in different species show distinct auditory input, with phasic-tonic spike responses to tones. In general, the interneurone response is more species-specific to calling than to courtship songs, and the preferential response to the conspecific calling song is based primarily upon sound frequency content.     

Topography of estradiol‐modulated genomic responses in the songbird auditory forebrain

Sex steroids facilitate dramatic changes in behavioral responses to sociosexual signals and are increasingly implicated in the sensory processing of those signals. Our previous work demonstrated that in female white‐throated sparrows, which are seasonal breeders, genomic responses in the auditory forebrain are selective for conspecific song over frequency‐matched tones only when plasma estradiol (E2) reaches breeding levels. Here, we sought to map this E2‐dependent selectivity in the best‐studied area of the auditory forebrain, the caudomedial nidopallium (NCM). Nonbreeding females with low endogenous levels of E2 were treated with E2 or a placebo and exposed to conspecific song, tones, or no sound playback. Immunoreactive protein product of the immediate early gene zenk (egr‐1) was then quantified within seven distinct subregions, or domains, of NCM. We report three main findings: (1) regardless of hormone treatment, the zenk response is significantly higher in dorsal than in ventral NCM, and higher in medial than in lateral NCM; (2) E2‐dependent selectivity of the response is limited to the rostral and medial domains of NCM; in the more caudal domains, song induces more zenk expression than tones regardless of hormone treatment; (3) even when no sound stimuli were presented, E2 treatment significantly increased zenk expression in the rostral, but not the caudal, domains of NCM. Together, the latter two findings suggest that E2‐dependent plasticity in NCM is concentrated in rostral NCM, which is hodologically and neurochemically distinct from caudal NCM. Activity in rostral NCM may therefore be seasonally regulated in this species. © 2009 Wiley Periodicals, Inc. Develop Neurobiol 2010     

Species Recognition During Substrate-Borne Communication in <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.) (Pentatomidae: Heteroptera)

The information code in the temporal and spectral characteristics of the substrate-borne communication signals produced by insects has been primarily studied in insects in the suborder Auchenorrhyncha. In the present study we investigated which of the female calling song (FCS) parameters in Nezara viridula (L.) (Heteroptera, Pentatomidae) are essential for recognition by conspecific males. In playback experiments we measured male vibrational responsiveness to FCS signals varying in the durations of pulse trains and inter-pulse train intervals, repetition times, duty cycles, and dominant frequencies, and determined the preference range for each specific parameter. Males were able to distinguish songs of different temporal and frequency parameters and responded best to values characteristic of the song of conspecific females. Signal recognition is achieved on the basis of two temporal filters tuned to the durations of the pulse train and inter-pulse train interval. Males responded best to the dominant frequency characteristic of conspecific songs, which are tuned to the resonant properties of the herbaceous plants used for intraspecific signal transmission during communication.     

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.     

Development of Tonal Quality in Birdsong: Further Evidence from Song Sparrows

Song sparrow (Melospiza melodia) songs are composed largely of pure-tone notes. Song sparrows raised in acoustic isolation (i.e. never hearing conspecific songs) tend to produce half of their notes with harmonic overtones, an atypical tonal structure, suggesting that exposure to pure tones is necessary for the development of normal tonal quality. The experiment presented here directly investigated the influence of early exposure to songs with different tonal qualities on subsequent production. Seven young song sparrow males were exposed to 16 normal, pure-tone song sparrow songs and 6 males were exposed to the same 16 song types with added harmonic overtones. Birds in both groups learned equally well, confirming an earlier finding that tonal quality does not influence selection of models during the sensitive period for song acquisition. Birds exposed exclusively to harmonic song models, however, produced over 85% of their learned notes in a pure-tone fashion, even though they had never heard pure-tone sounds. Thus, pure tones do not need to be experienced directly by song sparrows, but exposure to some features of species-typical song models appears to facilitate the reproduction of models in a pure-tone fashion.     

Broad Selectivity for Courtship Song in the Cricket Gryllus bimaculatus

Various characteristics of a long‐distance acoustic signal have been shown to vary to different degrees. It has been suggested that female preferences based on stable song parameters are stabilising or weakly directional, and preferences based on variable parameters are strongly directional. We tested this hypothesis based on a short‐distance signal (courtship song) produced by the field cricket, Gryllus bimaculatus. We studied the degree of variability of different courtship song parameters and the behavioural importance of several parameters using synthesised song models in playback experiments. We found that most of the courtship song elements of G. bimaculatus were quite variable (coefficient of variation, CV, in the range of 20–53%). The most variable parameter of the courtship song was the relative amplitude of two elements: high‐amplitude ticks and low‐amplitude pulses. Because songs containing only ticks (of rare occurrence) appeared to be more effective than songs with both ticks and pulses (of frequent occurrence), we consider female preferences to be directional. Alteration of less variable traits, such as the carrier frequency and duration of ticks (CV = 20–25%), had a different effect on female responsiveness. The synthesised songs with different carrier frequencies of ticks were as attractive to females as the positive control (courtship of muted males accompanied by playback of the recorded song). Altering the duration of ticks had a crucial effect on the female response rate, decreasing female responsiveness to the level observed in the negative control (courtship of muted males). Thus, we did not find a strong relationship between the variability of individual song parameters and their potential importance in song recognition and the evaluation of male quality. The partial inconsistency of our results with the data of other authors may be due to different patterns of past and current selection on long‐distance and short‐distance acoustic signals.     

Commensal auditory communication in two species of Neoconocephalus (Orthoptera)

Auditory communication in Neoconcephalus ensiger and Neoconocephalus robustus, two stridulating tettigoniids that inhabit the same geographical area, was examined to determine how these two species interact. Computer and electrophysiological techniques were used to analyze sound production and reception. Although similar in appearance, the males of these species produce easily distinguishable acoustic communication signals (call songs) that serve to attract conspecific females. Both wave form and spectral analyses were done on selected segments of the species-specific call songs by using discrete Fourier transform techniques. Sound production and reception capacities were measured by octave-band analyzers and extracellular electrode techniques. The results show that each of these species has a distinct, primary carrier frequency band. Secondly, the log magnitude spectra revealed a significant secondary component in the call song of N. robustus which corresponds to the primary energy band in the call song of N. ensiger. This overlap in acoustic signals and other findings suggest that males of N. ensiger might use the louder stridulation of the more metabolically active N. robustus to attract conspecific females over great distances. N. ensiger males have a sound production capacity like N. robustus, but conserve energy for soft, near-field signaling. The audiogram representing sensitivity of the tympano-receptor in the foreleg showed sound reception to be matched precisely with sound production curves.     

Neuronal basis of phonotactic behaviour in Tettigonia viridissima : processing of behaviourally relevant signals by auditory afferents and thoracic interneurons

Information transmission in the auditory pathway of Tettigonia viridissima was investigated using song models and artificial stimuli. Receptor cells respond tonically to song models and copy the syllable pattern within a wide intensity range. The omega-neuron responds tonically to soma-ipsilateral stimuli. Contralateral stimuli elicit IPSPs both within dendritic (ipsilateral) and axonal (contralateral) branches, thereby emphasizing directionality. Both AN1 and AN2 respond with tonic, non-adapting responses, precisely copying the syllable pattern of the song. While AN1 is excited by sonic frequencies and inhibited by ultrasonic frequencies, AN2 responds predominantly to ultrasound. The TN1 only responds to the ultrasonic components of the song, with phasic responses, which adapt quickly. In the adapted state, it responds selectively to the time pattern of the conspecific song, but not to the song patterns of two syntopic Tettigonia species. TN2, which has not been described up until now, is tuned to ultrasonic frequencies. Its responses to song models vanish after a few syllables, because of quick adaptation. The morphology is unusual with the axon running contralateral to the input site. The behavioural relevance of auditory interneurons is discussed and compared with the auditory system of crickets. Accepted: 3 November 1996     

SOUND PRODUCTION AND SOUND EMISSION IN SEVEN SPECIES OF EUROPEAN TETTIGONIIDS. PART I. THE DIFFERENT PARAMETERS OF THE SONG; THEIR RELATION TO THE MORPHOLOGY OF THE BUSHCRICKET

Abstract

Comparative studies of sound production and sound emission in seven species of European tettigoniids have been carried out. The species chosen were two Tettigoniines (Tettigonia cantans, Tettigonia viridissima), two Ephippigerines (Ephippiger discoidalis, Ephippiger ephippiger), and three Decticines (Decticus albifrons, Decticus verrucivorus, Psorodonotus illyricus). The factors which determined the choice of species were the different morphology (for example body shape and weight, and wing size) of the three subfamilies. The parameters of the different songs (e.g. dominant frequency, intensity) are normally not correlated to any of the investigated morphological characteristics of the animals. In the brachypterous species intraspecific correlations exist between wing size and the dominant low frequency band of the call. This frequency band is also observable at related higher frequencies in the ultrasonic range (20–60 kHz), the observed band width increasing with frequency. Sound emission in all species is to some extent directional. This directionality is related to body size and wing structure. The song structure of the different species does not appear to be related to any observable characteristic of the habitat of die animals. A possible exception may be the song of Psorodonotus illyricus with a particularly low dominant frequency band. The phylogenetic development of the songs seems to be determined by relationships between the different species rather than to any factors contributed by the habitat.     

The function of auditory neurons in cricket phonotaxis

Summary The activity of auditory receptor cells and prothoracic auditory neurons of the cricket,Gryllus bimaculatus, was recorded intracellularly while the animal walked on a sphere or while passive movement was imposed on a foreleg.During walking the responses to simulated calling song is altered since (i) the auditory sensory cells and interneurons discharged impulses in the absence of sound stimuli (Figs. 1, 3) and (ii) the number of action potentials in response to sound is reduced in interneurons (Figs. 2, 3).These two effects occurred in different phases of the leg movement during walking and therefore masked, suppressed or did not affect the responses to auditory stimuli (Figs. 3, 4). Hence there is a time window within which the calling song can be detected during walking (Fig. 5).The extra excitation of receptors and interneurons is probably produced by vibration of the tympanum because (i) the excitation occurred at the same time as the leg placement (Fig. 4), (ii) during walking on only middle and hindlegs, no extra action potentials were observed (Fig. 6), (iii) in certain phases of passive movements receptor cells and interneurons were excited as long as the ipsilateral ear was not blocked (Figs. 8, 9).Suppression of auditory responses seems to be peripheral as well as central in origin because (i) it occurred at particular phases during active and passive leg movements in receptor cells and interneurons (Figs. 1, 4, 9), (ii) it disappeared if the ear was blocked during passive leg movements (Fig. 9) and (iii) it persisted if the animal walked only on the middle and hind legs (Fig. 6).     

Asymmetric song responses in coexisting Japanese cicadas

Diel partitioning of singing, which has been observed from the tropics to the temperate regions, is among the various mechanisms of sound partitioning found in multispecies cicada assemblages and is regarded as an important mechanism of coexistence. Using playback experiments, we studied interspecific interference between two Japanese cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata, which exhibit similar peak frequencies in their calling songs and different diel patterns of song activity. We found that the males of the two species had different song response patterns: C. facialis responded significantly more to conspecific calling songs than G. nigrofuscata. These results suggest that male C. facialis have evolved more intense conspecific song recognition than G. nigrofuscata. This may cause male G. nigrofuscata to avoid destructive acoustic interference during their active singing time.     

Temporal patterns of intra- and interspecific acoustic signals differ in two closely related species of Acanthoplus (Orthoptera: Tettigoniidae: Hetrodinae)

Males of the closely related African tettigoniids Acanthoplus discoidales and Acanthoplus longipes produce a long-lasting calling song and a short disturbance sound. The temporal patterns of the sounds were analysed in respect to species differences and song type differences. The calling songs of both species consist of impulses which are separated into verses of two syllables, with fewer impulses in the first syllable. A. longipes produces more impulses in each syllable than A. discoidales and has longer verse durations, verse intervals and syllable intervals. Also, the disturbance sounds, produced after mechanical stimulation, contain distinct verses of impulses. The disturbance sound of A. longipes has a higher number of impulses per verse than that of A. discoidales. The frequency spectra of the songs in both species have similar peak frequencies (around 12.5 kHz) and both species have their greatest hearing sensitivity in the range between 5 and 10 kHz. Females of A. longipes perform phonotaxis only to songs with a species-specific temporal pattern. By contrast, females of A. discoidales react positively to calling songs of both species.     

Sex-specific spectral tuning for the partner's song in the duetting bushcricket Ancistrura nigrovittata (Orthoptera: Phaneropteridae)

The song of the male bushcricket Ancistrura nigrovittata consists of a sequence of verses. Each verse comprises a syllable group, plus, after about 400 ms a single syllable serving as a trigger for the female response song. The carrier frequency of the male song spectrum peaks at around 15 kHz, while the female song peaks at around 27 kHz. The thresholds of female responses to models of male songs are lowest for song frequencies between 12 and 16 kHz and therefore correspond to the male song spectrum. The threshold curve of the female response to the trigger syllable at different frequencies has the same shape as the tuning for the syllable group. Phonotactic thresholds of male Ancistrura nigrovittata to synthetic female responses at different frequencies are lowest between 24 and 28 kHz and thereby correspond to the female song spectrum and clearly differ from female response thresholds. Activity of the tympanic fibre bundle of both sexes is most sensitive between 15 and 35 kHz and therefore not specifically tuned to the partner's song. Individual behavioural thresholds have their minimum within 10 dB of the values of tympanic thresholds.     

Variation in male wing song characters inDrosophila plantibia (Hawaiian picture-wingedDrosophila group)

The males of most species of the HawaiianDrosophila planitibia group produce songs when vibrating their wings during courtship. In four of the most recently evolved species,D. differens, D. planitibia, D. heteroneura, andD. silvestris, these songs are simple in structure and possess a higher and more variable carrier frequency than the songs of the more ancestral species of the group. In the present paper, we studied the variation in wing song production and song characters in aD. planitibia population. Some males vibrated their wings at a very low amplitude or slow speed, producing no detectable sound. Other males produced sound bursts varying in carrier frequency and burst length. The carrier frequency of the song depended mainly on the wing posture of the male during wing vibrations and was consistent for individual males. Variation between males of different isofemale progenies was not significant in any measured song trait. Songs of the males recorded in the present study differed, however, from songs of the males of a laboratory strain recorded earlier.     

Calling song and selective phonotaxis in the field crickets,Gryllus firmus andG. pennsylvanicus (Orthoptera: Gryllidae)

The field cricket species, Gryllus firmusand G. pennsylvanicus,occur in a mosaic hybrid zone that roughly parallels the eastern slope of the Appalachian mountains in the northeastern United States. It is important to know what role, if any, the calling song plays in mate choice in sympatric and allopatric populations. In this report, we present results on the variability of calling song properties along transects across this hybrid zone. We also present the results of experiments on phonotactic selectivity of females from an allopatric population of G. firmus.The male calling song of allopatric G. firmuswas significantly slower in temporal rhythm (i. e., chirp and pulse repetition rates) and lower in pitch (i.e., dominant frequency) than that of allopatric G. pennsylvanicus.Calling song properties of males recorded in the hybrid zone varied considerably in temporal and spectral properties. In two-stimulus (choice) phonotaxis experiments, allopatric females of G. firmuspreferred synthetic calling songs with conspecific pulse repetition rates over songs that had lower and higher pulse rates. This preference persisted even when the sound pressure levels of alternative stimuli were unequal. Therefore, allopatric females of G. firmuscan discriminate between conspecific and heterospecific calling songs. Whether or not this same selectivity is present in sympatric populations remains unclear. Investigations of phonotactic selectivity in other allopatric and sympatric populations of both species are currently under way.     

Response properties of primary auditory fibers in the cricketTeleogryllus oceanieus (Le Guillou)

Summary Single unit recordings of primary auditory fibers ofTeleogryllus oceanicus show responses to frequencies over the range 0.5 kHz to 42 kHz. The characteristic frequencies (ChFs) of units were distributed over much of the bandwidth investigated although few units were recorded with ChFs below 4 kHz or in the region 7 kHz to 10 kHz. Some units showed more than one peak of sensitivity and others were broad-banded with no tuning to a particular frequency. Units whose ChFs approximated to the carrier frequency (CF) of the proclamation song were the most highly tuned. The majority of units had a tonic response pattern and were not spontaneously active. The implications of these findings are discussed.Abbreviations ChF characteristic frequency - CF carrier frequency We thank Mr. P. Foster for techninical help.     

Encoding properties of auditory neurons in the brain of a soniferous damselfish: response to simple tones and complex conspecific signals

The fish auditory system encodes important acoustic stimuli used in social communication, but few studies have examined response properties of central auditory neurons to natural signals. We determined the features and responses of single hindbrain and midbrain auditory neurons to tone bursts and playbacks of conspecific sounds in the soniferous damselfish, Abudefduf abdominalis. Most auditory neurons were either silent or had slow irregular resting discharge rates <20 spikes s⁻¹. Average best frequency for neurons to tone stimuli was ~130 Hz but ranged from 80 to 400 Hz with strong phase-locking. This low-frequency sensitivity matches the frequency band of natural sounds. Auditory neurons were also modulated by playbacks of conspecific sounds with thresholds similar to 100 Hz tones, but these thresholds were lower than that of tones at other test frequencies. Thresholds of neurons to natural sounds were lower in the midbrain than the hindbrain. This is the first study to compare response properties of auditory neurons to both simple tones and complex stimuli in the brain of a recently derived soniferous perciform that lacks accessory auditory structures. These data demonstrate that the auditory fish brain is most sensitive to the frequency and temporal components of natural pulsed sounds that provide important signals for conspecific communication.     

Response properties of the prothoracic AN2 auditory interneurone to model calling songs in the cricket Gryllus bimaculatus

Sound processing properties for calling song (CS) models, as described for the prothoracic L3 auditory neurone in Acheta domesticus, are investigated for the homologous auditory neurone 2 (AN2) in female Gryllus bimaculatus De Geer. AN2 of G. bimaculatus responds selectively to the syllable period (SP) of models of a male CS. The selectiveness of this response parallels the selectivity of phonotaxis females perform in response to the same SPs. Both, the responses of AN2 and female behaviour show clear interindividual variability. The SP‐selective responses of AN2 result from an SP‐dependent reduction in the spiking to subsequent syllables of the model CSs, measured as the percentage decrement. This SP‐dependent response does not primarily result from inbuilt properties of the AN2 membrane. Rather, it is dependent on inhibitory input to the AN2. However, clear inhibitory postsynaptic potentials in dendritic recordings of the AN2 are not encountered. This immediate response of AN2 to CSs is followed by an increased rate of tonic firing between stimulus CSs, which is termed the prolonged response, and is dependent on the carrier frequencies that make up the male CSs. With stimulation on the contralateral side of the soma of AN2s, more than 50% of AN2s exhibit a prolonged response. However, with stimulation from the ipsilateral side of the soma, most AN2s exhibit a prolonged response. The prolonged response of AN2 at 5 kHz may be even more sensitive than the immediate response. Thus, the AN2 neurone could provide a basis for phonotaxis that is selective for both the SPs and the carrier frequencies of potentially attractive calling songs.     

Spectral and temporal gating mechanisms enhance the clutter rejection in the echolocating bat,Rhinolophus rouxi

Summary Doppler shift compensation behaviour in horseshoe bats, Rhinolophus rouxi, was used to test the interference of pure tones and narrow band noise with compensation performance. The distortions in Doppler shift compensation to sinusoidally frequency shifted echoes (modulation frequency: 0.1 Hz, maximum frequency shift: 3 kHz) consisted of a reduced compensation amplitude and/or a shift of the emitted frequency to lower frequencies (Fig. 1).Pure tones at frequencies between 200 and 900 Hz above the bat's resting frequency (RF) disturbed the Doppler shift compensation, with a maximum of intererence between 400 and 550 Hz (Fig. 2). Minimum duration of pure tones for interference was 20 ms and durations above 40 ms were most effective (Fig. 3). Interfering pure tones arriving later than about 10 ms after the onset of the echolocation call showed markedly reduced interference (Fig. 4). Doppler shift compensation was affected by pure tones at the optimum interfering frequency with sound pressure levels down to –48 dB rel the intensity level of the emitted call (Figs. 5, 6).Narrow bandwidth noise (bandwidth from ± 100 Hz to ± 800 Hz) disturbed Doppler shift compensation at carrier frequencies between –250 Hz below and 800 Hz above RF with a maximum of interference between 250 and 500 Hz above resting frequency (Fig. 7). The duration and delay of the noise had similar influences on interference with Doppler shift compensation as did pure tones (Figs. 8, 9). Intensity dependence for noise interference was more variable than for pure tones (-32 dB to -45 dB rel emitted sound pressure level, Fig. 10).The temporal and spectral gating in Doppler shift compensation behaviour is discussed as an effective mechanism for clutter rejection by improving the processing of frequency and amplitude transients in the echoes of horseshoe bats.Abbreviations CF constant frequency - FM frequency modulation - RF resting frequency - SPL sound pressure level