Frequency specific susceptibility to acoustic trauma in the budgerigar (Melopsittacus undulatus)

The effects of intense noise exposure on hearing in the budgerigar were examined by behavioral audiometry. After binaural exposure to an intense broadband noise, auditory threshold shifts (TS) of the birds were continuously measured at frequencies between 0.125 and 8 kHz using an avoidance conditioning technique. Temporary threshold shifts (TTS) were observed at frequencies higher than 1.5 kHz and considerable permanent threshold shifts (PTS) were observed at frequencies below 1 kHz. This pattern of threshold shifts is contrary to that observed in mammals.     

Lateralization of acoustic signals by dichotically listening budgerigars (Melopsittacus undulatus)

Sound localization allows humans and animals to determine the direction of objects to seek or avoid and indicates the appropriate position to direct visual attention. Interaural time differences (ITDs) and interaural level differences (ILDs) are two primary cues that humans use to localize or lateralize sound sources. There is limited information about behavioral cue sensitivity in animals, especially animals with poor sound localization acuity and small heads, like budgerigars. ITD and ILD thresholds were measured behaviorally in dichotically listening budgerigars equipped with headphones in an identification task. Budgerigars were less sensitive than humans and cats, and more similar to rabbits, barn owls, and monkeys, in their abilities to lateralize dichotic signals. Threshold ITDs were relatively constant for pure tones below 4 kHz, and were immeasurable at higher frequencies. Threshold ILDs were relatively constant over a wide range of frequencies, similar to humans. Thresholds in both experiments were best for broadband noise stimuli. These lateralization results are generally consistent with the free field localization abilities of these birds, and add support to the idea that budgerigars may be able to enhance their cues to directional hearing (e.g., via connected interaural pathways) beyond what would be expected based on head size.     

Kanamycin induced low-frequency hearing loss in the budgerigar (Melopsittacus undulatus)

The chronic effects of kanamycin (KM) on hearing in the budgerigar were investigated by behavioral audiometry. The birds received a daily intramuscular injection of KM (100 mg/kg or 200 mg/kg) for 10 successive days, and absolute thresholds between pre- and post-treatment were compared. KM induced both transient and permanent low-frequency specific hearing loss; i.e., the elevation of threshold for frequencies below 1 kHz was greater than that for frequencies above 1 kHz. Moreover, the degree of hearing loss was dose dependent. The low-frequency selective effects of KM in the present study were contrary to the high-frequency specificity of aminoglycoside ototoxicity in mammals. To assess the effect of KM on auditory frequency resolution, critical ratios were estimated in pathological birds with low-frequency specific hearing loss. There was a linear relation between shift in critical ratio and shift in absolute threshold, suggesting that the increase in the critical ratio is due to a decrease in the efficiency of the detector mechanism rather than a change in the spectral resolving power of the birds.     

Vocal production mechanisms in the budgerigar (Melopsittacus undulatus): the presence and implications of amplitude modulation.

In this paper acoustic evidence is presented for the presence of amplitude modulation in budgerigar (Melopsittacus undulatus) contact calls and learned English vocalizations. Previously, acoustic analyses of budgerigar vocalizations have consisted solely of visual inspection of spectrograms or power spectra (derived from Fourier transformation). Such analyses have led researchers to conclude that budgerigar vocalizations are primarily frequency-modulated, harmonic vocalizations. Although budgerigar calls have been shown to contain regions that are modulated in amplitude, the implications of this fact have been largely ignored. Amplitude modulation, the nonlinear interaction between two separate signals that results in the creation of new, heterodyne (sum and difference) frequencies, can produce a very complex Fourier spectrum that may resemble that produced by a harmonic vocalization. In this paper, the acoustic principles necessary for identifying amplitude modulation present in signals are outlined, and followed by data demonstrating that amplitude modulation is a prominent feature not only of natural budgerigar contact calls, but also of their learned English vocalizations. It is illustrated how analyzing a vocalization that contains amplitude modulation as if it were harmonic can result in misinterpretations of the acoustic and physical properties of the sound and sound source. The implications of amplitude modulation for studies of the ontogenetic, physical, and neural basis of budgerigar vocalizations are discussed, and a potential model for how the budgerigar syrinx may function to produce amplitude modulation is proposed.     

Masking by harmonic complexes in budgerigars (Melopsittacus undulatus)

In humans, masking by harmonic complexes is dependent not only on the frequency content of the masker, but also its phase spectrum. Complexes that have highly modulated temporal waveforms due to the selection of their component phases usually provide less masking than those with flatter temporal envelopes. Moreover, harmonic complexes that are created with negative Schroeder phases (component phases monotonically decreasing with increasing harmonic frequency) may provide more masking than those created with positive Schroeder phases (monotonically increasing phase), even though both temporal envelopes are equally flat. To date, there has been little comparative work on the masking effectiveness of harmonic complexes. Using operant conditioning and the method of constant stimuli, masking of pure tones by harmonic complexes was examined in budgerigars at several different masker levels for complexes constructed with two different fundamental frequencies. In contrast to humans, thresholds in budgerigars differed very little for the two Schroeder-phase waveforms. Moreover, when there was a difference in masking by these two waveforms, the positive Schroeder was the more effective masker--the reverse of that described for humans. Control experiments showed that phase selection was relevant to the masking ability of harmonic complexes in budgerigars. Release from masking occurred when the components were in coherent phase, compared with a complex with random phases selected for each component. It is suggested that these psychoacoustic differences may emerge from structural and functional differences between the avian and mammalian peripheral auditory systems involving traveling wave mechanics and spectral tuning characteristics.     

Development of auditory sensitivity in budgerigars (Melopsittacus undulatus)

Auditory feedback influences the development of vocalizations in songbirds and parrots; however, little is known about the development of hearing in these birds. The auditory brainstem response was used to track the development of auditory sensitivity in budgerigars from hatch to 6 weeks of age. Responses were first obtained from 1-week-old at high stimulation levels at frequencies at or below 2 kHz, showing that budgerigars do not hear well at hatch. Over the next week, thresholds improved markedly, and responses were obtained for almost all test frequencies throughout the range of hearing by 14 days. By 3 weeks posthatch, birds' best sensitivity shifted from 2 to 2.86 kHz, and the shape of the auditory brainstem response (ABR) audiogram became similar to that of adult budgerigars. About a week before leaving the nest, ABR audiograms of young budgerigars are very similar to those of adult birds. These data complement what is known about vocal development in budgerigars and show that hearing is fully developed by the time that vocal learning begins.     

Vocal learning in Budgerigars (Melopsittacus undulatus): effects of an acoustic reference on vocal matching

Budgerigars were trained to produce specific vocalizations (calls) using operant conditioning and food reinforcement. The bird's call was compared to a digital representation of the call stored in a computer to determine a match. Once birds were responding at a high level of precision, we measured the effect of several manipulations upon the accuracy and the intensity of call production. Also, by differentially reinforcing other aspects of vocal behavior, budgerigars were trained to produce a call that matched another bird's contact call and to alter the latency of their vocal response. Both the accuracy of vocal matching and the intensity level of vocal production increased significantly when the bird could hear the template immediately before each trial. Moreover, manipulating the delay between the presentation of an acoustic reference and the onset of vocal production did not significantly affect either vocal intensity or matching accuracy. Interestingly, the vocalizations learned and reinforced in these operant experiments were only occasionally used in more natural communicative situations, such as when birds called back and forth to one another in their home cages.     

Detection of modulation in spectral envelopes and linear-rippled noises by budgerigars (Melopsittacus undulatus)

Budgerigars were trained to discriminate complex sounds with two different types of spectral profiles from flat-spectrum, wideband noise. In one case, complex sounds with a sinusoidal ripple in (log) amplitude across (log) frequency bandwidth were generated by combining 201 logarithmically spaced tones covering the frequency region from 500 Hz to 10 kHz. A second type of rippled stimulus was generated by delaying broadband noise and adding it to the original noise in an iterative fashion. In each case, thresholds for modulation depth (i.e., peak-to-valley in dB) were measured at several different ripple frequencies (i.e., cycles/octave for logarithmic profiles) or different repetition pitches (i.e., delay for ripple noises). Budgerigars were similar to humans in detecting ripple at low spatial frequencies, but were considerably more sensitive than humans in detecting ripples in log ripple spectra at high spatial frequencies. Budgerigars were also similar to humans in detecting linear ripple in broadband noise over a wide range of repetition pitches. Taken together, these data show that the avian auditory system is at least as good, if not better, than the human auditory system at detecting spectral ripples in noise despite gross anatomical differences in both the peripheral and central auditory nervous systems.     

Identification of auditory distance cues by zebra finches (Taeniopygia guttata) and budgerigars (Melopsittacus undulatus)

The present study examined auditory distance perception cues in a non-territorial songbird, the zebra finch (Taeniopygia guttata), and in a non-songbird, the budgerigar (Melopsittacus undulatus). Using operant conditioning procedures, three zebra finches and three budgerigars were trained to identify 1- (Near) and 75-m (Far) recordings of three budgerigar contact calls, one male zebra finch song, and one female zebra finch call. Once the birds were trained on these endpoint stimuli, other stimuli were introduced into the operant task. These stimuli included recordings at intermediate distances and artificially altered stimuli simulating changes in overall amplitude, high-frequency attenuation, reverberation, and all three cues combined. By examining distance cues (amplitude, high-frequency attenuation, and reverberation) separately, this study sought to determine which cue was the most salient for the birds. The results suggest that both species could scale the stimuli on a continuum from Near to Far and that amplitude was the most important cue for these birds in auditory distance perception, as in humans and other animals.     

Perception of complex sounds in budgerigars (Melopsittacus undulatus) with temporary hearing loss

Songbirds and parrots deafened as nestlings fail to develop normal vocalizations, while birds deafened as adults show a gradual deterioration in the quality and precision of vocal production. Beyond this, little is known about the effect of hearing loss on the perception of vocalizations. Here, we induced temporary hearing loss in budgerigars with kanamycin and tested several aspects of the hearing, including the perception of complex, species-specific vocalizations. The ability of these birds to discriminate among acoustically distinct vocalizations was not impaired but the ability to make fine-grain discriminations among acoustically similar vocalizations was affected, even weeks after the basilar papilla had been repopulated with new hair cells. Interestingly, these birds were initially unable to recognize previously familiar contact calls in a classification task-suggesting that previously familiar vocalizations sounded unfamiliar with new hair cells. Eventually, in spite of slightly elevated absolute thresholds, the performance of birds on discrimination and perceptual recognition of vocalizations tasks returned to original levels. Thus, even though vocalizations may initially sound different with new hair cells, there are only minimal long-term effects of temporary hearing loss on auditory perception, recognition of species-specific vocalizations, or other aspects of acoustic communication in these birds.     

The Franssen effect illusion in budgerigars (Melopsittacus undulatus) and zebra finches (Taeniopygia guttata)

The properties of the Franssen effect (FE) were measured in budgerigars and zebra finches. To elicit the FE, listeners are presented with a signal which has been split into a transient component, carrying an abrupt onset and ramped offset and separated in space from the sustained component which has a slowly rising onset and longer duration. When these two signals are played under certain conditions, the perception is that of a long-duration steady state tone being played at the location of the transient. The birds were trained using operant conditioning methods on a categorization task to peck a left key when presented with a stimulus from a left speaker and to peck a right key when presented with a stimulus from a right speaker. Once training was completed, FE stimuli were presented during a small proportion of trials. The FE was measured at speaker separations of 60 degrees and 180 degrees in both echoic and echoic-reduced conditions. Both species of birds exhibited the FE, although to varying degrees, across conditions. These results show that nonmammals also experience the FE illusion in confusing listening situations in a manner similar to mammals, suggestive of similar auditory processing mechanisms.     

Effects of deafening on the calls and warble song of adult budgerigars (Melopsittacus undulatus)

Budgerigars are small Australian parrots that learn new vocalizations throughout adulthood. Earlier work has shown that an external acoustic model and auditory feedback are necessary for the development of normal contact calls in this species. Here, the role of auditory feedback in the maintenance of species-typical contact calls and warble song in adult budgerigars is documented. Deafened adult birds (five male, one female) vocalized less frequently and showed both suprasegmental and segmental changes in their contact calls and warble song. Contact calls of all adult-deafened budgerigars showed abnormalities in acoustic structure within days to a few weeks following surgery. Within 6 months of surgery, nearly all contact calls produced by deafened birds were strikingly abnormal, showing highly variable patterns of frequency modulation and duration. The warble song of deafened male budgerigars also differed significantly from that of normal budgerigars on several acoustic measures. These results show that auditory feedback is necessary for the maintenance of a normal, species-typical vocal repertoire in budgerigars.     

Learned vocalizations in budgerigars (Melopsittacus undulatus): the relationship between contact calls and warble song

The budgerigar (Melopsittacus undulatus) has an extraordinarily complex, learned, vocal repertoire consisting of both the long rambling warble song of males and a number of short calls produced by both sexes. In warble, the most common elements (>30%) bear a strong resemblance to the highly frequency-modulated, learned contact calls that the birds produce as single utterances. However, aside from this apparent similarity, little else is known about the relationship between contact calls and warble call elements. Here, both types of calls were recorded from four male budgerigars. Signal analysis and psychophysical testing procedures showed that the acoustic features of these two vocalizations were acoustically different and perceived as distinctive vocalizations by birds. This suggests that warble call elements are not simple insertions of contact calls but are most likely different acoustic elements, created de novo, and used solely in warble. Results show that, like contact calls, warble call elements contain information about signaler identity. The fact that contact calls and warble call elements are acoustically and perceptually distinct suggests that they probably represent two phonological systems in the budgerigar vocal repertoire, both of which arise by production learning.     

Temporal coherence of acoustic signals in a fluctuating ocean

Temporal coherence of acoustic signals propagating in a fluctuating ocean is important for many practical applications and has been studied intensively experimentally. However, only a few theoretical formulations of temporal coherence exist. In the present paper, a three-dimensional (3D) modal theory of sound propagation in a fluctuating ocean is used to derive closed-form equations for the spatial-temporal coherence function of a broadband signal. The theory is applied to the analysis of the temporal coherence of a monochromatic signal propagating in an ocean perturbed by linear internal waves obeying the Garrett-Munk (G-M) spectral model. In particular, the temporal coherence function is calculated for propagation ranges up to 10(4) km and for five sound frequencies: 12, 25, 50, 75, and 100 Hz. Then, the dependence of the coherence time (i.e., the value of the time lag at which the temporal coherence decreases by a factor of e) on range and frequency is studied. The results obtained are compared with experimental data and predictions of the path-integral theory.     

Temporal integration in zebra finches (Poephila guttata)

Three zebra finches were trained with operant techniques to respond to pure tones. Absolute thresholds were obtained for nine durations of a 3-kHz tone and five durations of a 1-kHz tone. The temporal integration functions were described using the negative exponential function proposed by Plomp and Bouman [J. Acoust. Soc. Am. 31, 749-758 (1959)]. The time constants obtained for zebra finches are about 250 ms, which are similar to those reported for a number of species, including humans and other bird species.     

Light modulation by acoustic signals for high diffraction efficiency

A theoretical discussion is presented on strong Bragg acoustooptic interaction AOI of light beams in the dynamic field of an acoustic signal. A system of integrodifferential equations is formulated to describe the evolution of the angular and frequency spectra of the beams in the AOI region for a high level of acoustooptic coupling. The third-order approximation in the perturbation method is used to obtain an analytic solution. Calculations are presented on the modulation of monochromatic beams by acoustic pulses having rectangular envelopes and propagating in a lithium niobate crystal, and the same for a signal having linear frequency modulation LFM in a paratellurite crystal, which demonstrate the broadening of the beam spectrum as the depth of the acoustooptic coupling increases, together with the occurrence of an asymmetry specific to strong AOI in the response of the light field to the symmetrical acoustic signal. Tomsk State University for Control Systems and Electronics, Tomsk. Translated from Izvestiya Vysshikh Uchebnykh Zavedenii, Fizika, Vol. 42, No. 1, pp. 99–106, January, 1999.