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.     

Cepstral coefficients and hidden Markov models reveal idiosyncratic voice characteristics in red deer (Cervus elaphus) stags

Bouts of vocalizations given by seven red deer stags were recorded over the rutting period, and homomorphic analysis and hidden Markov models (two techniques typically used for the automatic recognition of human speech utterances) were used to investigate whether the spectral envelope of the calls was individually distinctive. Bouts of common roars (the most common call type) were highly individually distinctive, with an average recognition percentage of 93.5%. A "temporal" split-sample approach indicated that although in most individuals these identity cues held over the rutting period, the ability of the models trained with the bouts of roars recorded early in the rut to correctly classify later vocalizations decreased as the recording date increased. When Markov models trained using the bouts of common roars were used to classify other call types according to their individual membership, the classification results indicated that the cues to identity contained in the common roars were also present in the other call types. This is the first demonstration in mammals other than primates that individuals have vocal cues to identity that are common to the different call types that compose their vocal repertoire.     

Acoustic correlates of caller identity and affect intensity in the vowel-like grunt vocalizations of baboons

Comparative, production-based research on animal vocalizations can allow assessments of continuity in vocal communication processes across species, including humans, and may aid in the development of general frameworks relating specific constitutional attributes of callers to acoustic-structural details of their vocal output. Analyses were undertaken on vowel-like baboon grunts to examine variation attributable to caller identity and the intensity of the affective state underlying call production. Six hundred six grunts from eight adult females were analyzed. Grunts derived from 128 bouts of calling in two behavioral contexts: concerted group movements and social interactions involving mothers and their young infants. Each context was subdivided into a high- and low-arousal condition. Thirteen acoustic features variously predicted to reflect variation in either caller identity or arousal intensity were measured for each grunt bout, including tempo-, source- and filter-related features. Grunt bouts were highly individually distinctive, differing in a variety of acoustic dimensions but with some indication that filter-related features contributed disproportionately to individual distinctiveness. In contrast, variation according to arousal condition was associated primarily with tempo- and source-related features, many matching those identified as vehicles of affect expression in other nonhuman primate species and in human speech and other nonverbal vocal signals.     

Affect cues in vocalizations of the bat, Megaderma lyra, during agonistic interactions

Some features of emotional prosody in human speech may be traced back to affect cues in mammalian vocalizations. The present study addresses the question whether affect intensity, as expressed by the intensity of behavioral displays, is encoded in vocal cues, i.e., changes in the structure of associated calls, in bats, a group evolutionarily remote from primates. A frame-by-frame video analysis of 109 dyadic agonistic interactions recorded in approach situations was performed to categorize displays into two intensity levels based on a cost-benefit estimate. M. lyra showed graded visual displays accompanied by specific calls and response calls of the second bat. A sound analysis revealed systematic changes of call sequence parameters with display level. At the high intensity level, total call duration, number of syllables within a call, and the number of calls within a sequence were increased, while intervals between call syllables were decreased for both call types. In addition, the latency of the response call was shorter, and its main syllable-type durations and fundamental frequency were increased. These systematic changes of vocal parameters with affect intensity correspond to prosodic changes in human speech, suggesting that emotion-related acoustic cues are a common feature of vocal communication in mammals.     

The discrimination of baboon grunt calls and human vowel sounds by baboons

The ability of baboons to discriminate changes in the formant structures of a synthetic baboon grunt call and an acoustically similar human vowel (/epsilon/) was examined to determine how comparable baboons are to humans in discriminating small changes in vowel sounds, and whether or not any species-specific advantage in discriminability might exist when baboons discriminate their own vocalizations. Baboons were trained to press and hold down a lever to produce a pulsed train of a standard sound (e.g., /epsilon/ or a baboon grunt call), and to release the lever only when a variant of the sound occurred. Synthetic variants of each sound had the same first and third through fifth formants (F1 and F3-5), but varied in the location of the second formant (F2). Thresholds for F2 frequency changes were 55 and 67 Hz for the grunt and vowel stimuli, respectively, and were not statistically different from one another. Baboons discriminated changes in vowel formant structures comparable to those discriminated by humans. No distinct advantages in discrimination performances were observed when the baboons discriminated these synthetic grunt vocalizations.     

Influence of anabolic steroid treatment associated to physical exercise in the ultrasonic vocalization of rats

Unlike humans, who communicate in frequency bands between 250 Hz and 6 kHz, rats can communicate in frequencies above 18 kHz. Their vocalization types depend on the context and are normally associated to subjective or emotional states. It was reported significant vocal changes due to administration of replacement testosterone in a trained tenor singer with hypogonadism. Speech-Language Pathology clinical practices are being sought by singers who sporadically use anabolic steroids associated with physical exercise. They report difficulties in reaching and keeping high notes, "breakage" in the passage of musical notes and post singing vocal fatigue. Those abnormalities could be raised by the association of anabolic steroids and physical exercise. Thus, in order to verify if this association could promote vocal changes, maximum, minimum and fundamental frequencies and call duration in rats treated with anabolic steroids and physically trained (10 weeks duration) were evaluated. The vocalizations were obtained by handling the animals. At the end of that period, rats treated and trained showed significant decrease in call duration, but not in other parameters. The decrease in call duration could be associated to functional alterations in the vocal folds of treated and trained animals due to a synergism between anabolic steroids and physical training.     

The effects of articulation on the acoustical structure of feline vocalizations

Feline isolation calls were analyzed, and a model was developed to relate the acoustical features of these calls to the physical processes used in their production. Fifty isolation calls were recorded from each of five cats for a total sample of 250 vocalizations. By combinations of Fourier transform, autocorrelation, and linear prediction methods, the fundamental frequency (glottal-pulse period) F0, the energy of F0, the frequency having maximum energy Fmax (not always F0), and the energy at this frequency were computed. Mean F0 ranged from 400-600 Hz for individual cats. For some cats F0 was consistent within calls, but for other cats sudden shifts in F0 occurred within calls. Here, Fmax was almost a harmonic of F0 and generally ranged from 1-2 kHz. For individual cats, the energy ratio E = (energy of Fmax/energy of F0) varied from 1 to 60 and the grand average E over the time course of the call varied from about 12 to 38. The mean rms call intensity was an inverted-U function of time. Measured jaw opening was strongly correlated with acoustical features of call. A Bessel-horn model with time-varying flare gave a good account of acoustical parameters such as Fmax. The presence of formantlike resonances in cat vocalizations and the important role of jaw movements (vocal gestures) in the production of these calls suggest that cats may provide a useful model for some aspects of human vocal behavior.     

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.     

Acoustic analysis of vocal development in a New World primate, the common marmoset (Callithrix jacchus)

In contrast to humans and songbirds, there is limited evidence of vocal learning in nonhuman primates. While previous studies suggested that primate vocalizations exhibit developmental changes, detailed analyses of the extent and time course of such changes across a species' vocal repertoire remain limited. In a highly vocal primate, the common marmoset (Callithrix jacchus), we studied developmental changes in the acoustic structure of species-specific communication sounds produced in a social setting. We performed detailed acoustic analyses of the spectral and temporal characteristics of marmoset vocalizations during development, comparing differences between genders and twin pairs, as well as with vocalizations from adult marmosets residing in the same colony. Our analyses revealed significant changes in spectral and temporal features as well as variability of particular call types over time. Infant and juvenile vocalizations changed progressively toward the vocalizations produced by adult marmosets. Call types observed early in development that were unique to infants disappeared gradually with age, while vocal exchanges with conspecifics emerged. Our observations clearly indicate that marmoset vocalizations undergo both qualitative and quantitative postnatal changes, establishing the basis for further studies to delineate contributions from maturation of the vocal apparatus and behavioral experience.     

Nonlinear phenomena in contemporary vocal music.

Complex and multiphonic voice signals of vocal improvisors are analyzed within the framework of nonlinear dynamics. Evidence is given that nonlinear phenomena are extensively used by performers associated with contemporary music. Narrow-band spectrograms of complex vocalizations are used to visualize the appearance of nonlinear phenomena (spectral bifurcation diagrams). Possible production mechanisms are discussed in connection with previous research, personal performance and pedagogical experience. Examples for period doubling, biphonation and irregular aperiodic phonation in vocal sonorities of contemporary vocal improvisors are given, and glottal whistle production encompassed with biphonation and triphonation is shown. Furthermore, coincidences of harmonics-formant matching associated with abrupt transitions to subharmonics and biphonation in the vocal output are provided. This also shows the recurrent use of nonlinear phenomena by performers. It is argued that mechanisms such as source-tract coupling or vocal fold desynchronization due to asymmetry are used in a reproducible way for musical tasks.     

Pitch Matching Accuracy of Trained Singers, Untrained Subjects with Talented Singing Voices, and Untrained Subjects with Nontalented Singing Voices in Conditions of Varying Feedback

At a physiological level, the act of singing involves control and coordination of several systems involved in the production of sound, including respiration, phonation, resonance, and afferent systems used to monitor production. The ability to produce a melodious singing voice (eg, in tune with accurate pitch) is dependent on control over these motor and sensory systems. To test this position, trained singers and untrained subjects with and without expressed singing talent were asked to match pitches of target pure tones. The ability to match pitch reflected the ability to accurately integrate sensory perception with motor planning and execution. Pitch-matching accuracy was measured at the onset of phonation (prephonatory set) before external feedback could be utilized to adjust the voiced source, during phonation when external auditory feedback could be utilized, and during phonation when external auditory feedback was masked. Results revealed trained singers and untrained subjects with singing talent were no different in their pitch-matching abilities when measured before or after external feedback could be utilized. The untrained subjects with singing talent were also significantly more accurate than the trained singers when external auditory feedback was masked. Both groups were significantly more accurate than the untrained subjects without singing talent.     

Vocal Expression of Emotions in Normally Hearing and Hearing-Impaired Infants

The vocalizations of seven normally hearing (NH) and seven severely hearing-impaired (HI) infants were compared to find out the influence of auditory feedback on preverbal utterances. It was tested whether there are general differences in vocalizations between NH and HI infants, and whether specific emotional states affect the vocal production of NH and HI infants in the same way. First, the acoustic structure of the three most common vocal types was analyzed; second, the composition of vocal sequences was examined. Vocal sequence composition turned out to be more affected by hearing impairment than the acoustic structure of single vocalizations. This result indicates that the acoustic structure of preverbal vocalizations is to a great extent predetermined, whereas the composition of vocal sequences is influenced by auditory input.     

Repetition patterns in Weddell seal (Leptonychotes weddellii) underwater multiple element calls

Many vocalizations produced by Weddell seals (Leptonychotes weddellii) are made up of repeated individual distinct sounds (elements). Patterning of multiple element calls was examined during the breeding season at Casey and Davis, Antarctica. Element and interval durations were measured from 405 calls all > 3 elements in length. The duration of the calls (22+/-16.6 s) did not seem to vary with an increasing number of elements (F4,404=1.83,p = 0.122) because element and interval durations decreased as the number of elements within a call increased. Underwater vocalizations showed seven distinct timing patterns of increasing, decreasing, or constant element and interval durations throughout the calls. One call type occurred with six rhythm patterns, although the majority exhibited only two rhythms. Some call types also displayed steady frequency changes as they progressed. Weddell seal multiple element calls are rhythmically repeated and thus the durations of the elements and intervals within a call occur in a regular manner. Rhythmical repetition used during vocal communication likely enhances the probability of a call being detected and has important implications for the extent to which the seals can successfully transmit information over long distances and during times of high level background noise.     

Preliminary evidence for signature vocalizations among free-ranging narwhals (Monodon monoceros)

Animal signature vocalizations that are distinctive at the individual or group level can facilitate recognition between conspecifics and re-establish contact with an animal that has become separated from its associates. In this study, the vocal behavior of two free-ranging adult male narwhals (Monodon monoceros) in Admiralty Inlet, Baffin Island was recorded using digital archival tags. These recording instruments were deployed when the animals were caught and held onshore to attach satellite tags, a protocol that separated them from their groups. The signature content of two vocal categories was considered: (1) combined tonal/pulsed signals, which contained synchronous pulsatile and tonal content; (2) whistles, or frequency modulated tonal signals with harmonic energy. Nonparametric comparisons of the temporal and spectral features of each vocal class revealed significant differences between the two individuals. A separate, cross-correlation measure conducted on the whistles that accounted for overall contour shape and absolute frequency content confirmed greater interindividual compared to intraindividual differences. These data are consistent with the hypothesis that narwhals produce signature vocalizations that may facilitate their reunion with group members once they become separated, but additional data are required to demonstrate this claim more rigorously.     

Sex differences in the acoustic structure of vowel-like grunt vocalizations in baboons and their perceptual discrimination by baboon listeners

This study quantifies sex differences in the acoustic structure of vowel-like grunt vocalizations in baboons (Papio spp.) and tests the basic perceptual discriminability of these differences to baboon listeners. Acoustic analyses were performed on 1028 grunts recorded from 27 adult baboons (11 males and 16 females) in southern Africa, focusing specifically on the fundamental frequency (F0) and formant frequencies. The mean F0 and the mean frequencies of the first three formants were all significantly lower in males than they were in females, more dramatically so for F0. Experiments using standard psychophysical procedures subsequently tested the discriminability of adult male and adult female grunts. After learning to discriminate the grunt of one male from that of one female, five baboon subjects subsequently generalized this discrimination both to new call tokens from the same individuals and to grunts from novel males and females. These results are discussed in the context of both the possible vocal anatomical basis for sex differences in call structure and the potential perceptual mechanisms involved in their processing by listeners, particularly as these relate to analogous issues in human speech production and perception.     

Variation in chick-a-dee calls of tufted titmice, Baeolophus bicolor: note type and individual distinctiveness

The chick-a-dee call of chickadee species (genus Poecile) has been the focus of much research. A great deal is known about the structural complexity and the meaning of variation in notes making up calls in these species. However, little is known about the likely homologous "chick-a-dee" call of the closely related tufted titmouse, Baeolophus bicolor. Tufted titmice are a prime candidate for comparative analyses of the call, because their vocal and social systems share many characteristics with those of chickadees. To address the paucity of data on the structure of chick-a-dee calls of tufted titmice, we recorded birds in field and aviary settings. Four main note types were identified in the call: Z, A, D(h), and D notes. Several acoustic parameters of each note type were measured, and statistical analyses revealed that the note types are acoustically distinct from one another. Furthermore, note types vary in the extent of individual distinctiveness reflected in their acoustic parameters. This first step towards understanding the chick-a-dee call of tufted titmice indicates that the call is comparable in structure and complexity to the calls of chickadees.     

Colony differences in auditory thresholds in the canary (Serinus canarius)

Adult canaries (Serinus canarius) from a closebred colony of the Belgian "Waterslager" strain were trained with operant techniques to respond to pure tones. A psychophysical tracking procedure was used to measure absolute auditory thresholds in quiet and in noise. Absolute thresholds in the middle- to high-frequency region of the audiogram were between 30 and 40 dB higher (4-5 standard deviations) than those typically reported for other song birds including canaries of other strains and Waterslagers tested some years ago from another colony. Thus the Millbrook colony of domestic canary--an oscine songbird which learns its vocalizations by reference to auditory information--shows unusually high absolute thresholds for pure tones.     

Phonetogram Changes for Trained Singers Over a Nine-Month Period of Vocal Training

Professional vocalists encounter demands requiring voluntary control of phonation, while utilizing a considerable range of frequency and intensity. These quantifiable acoustic events can be measured and represented in a phonetogram. Previous research has compared the phonetograms of trained and untrained voices and found significant differences between these groups. This study was designed to assess the effects of vocal training for singers over a period of nine months. Phonetogram contour changes were examined, with the primary focus on expansion of frequency range and/or intensity control. Twenty-one first-year, master's level, vocal music students, who were engaged in an intensive vocal performance curriculum, participated in this study. Following nine months of vocal training, significant differences were revealed in the subjects' mean frequency range and minimum vocal intensity across frequency levels. There was no significant difference for the mean maximum vocal intensity across frequency levels following vocal training.     

Finding good acoustic features for parrot vocalizations: the feature generation approach

A crucial step in the understanding of vocal behavior of birds is to be able to classify calls in the repertoire into meaningful types. Methods developed to this aim are limited either because of human subjectivity or because of methodological issues. The present study investigated whether a feature generation system could categorize vocalizations of a bird species automatically and effectively. This procedure was applied to vocalizations of African gray parrots, known for their capacity to reproduce almost any sound of their environment. Outcomes of the feature generation approach agreed well with a much more labor-intensive process of a human expert classifying based on spectrographic representation, while clearly out-performing other automated methods. The method brings significant improvements in precision over commonly used bioacoustical analyses. As such, the method enlarges the scope of automated, acoustics-based sound classification.     

An electroglottographic study of vocal vibrato

This study investigated vocal vibrato in trained singers through the use of electroglottography (EGG). Ten adult trained singers (7 women, 3 men) were each required to produce four singing tasks. The tasks involved the production of /a/ during an ascending interval of a third in both chest and falsetto registers, once with and once without vibrato. Audio and EGG output during these tasks were directly digitized for subsequent analysis. Results pertaining to changes in the EGG waveform as a function of the singer's gender, the vocal register and the vibrato condition are discussed. A major finding was the apparent absence of consistent, distinctive differentiation in the EGG waveforms when comparing the vibrato and vibrato-inhibited conditions for the majority of subjects. Possible explanations for the majority findings, as well as comment on the feasibility of the EGG for investigating vocal vibrato, are discussed.