Auditory scene analysis by echolocation in bats.

Echolocating bats transmit ultrasonic vocalizations and use information contained in the reflected sounds to analyze the auditory scene. Auditory scene analysis, a phenomenon that applies broadly to all hearing vertebrates, involves the grouping and segregation of sounds to perceptually organize information about auditory objects. The perceptual organization of sound is influenced by the spectral and temporal characteristics of acoustic signals. In the case of the echolocating bat, its active control over the timing, duration, intensity, and bandwidth of sonar transmissions directly impacts its perception of the auditory objects that comprise the scene. Here, data are presented from perceptual experiments, laboratory insect capture studies, and field recordings of sonar behavior of different bat species, to illustrate principles of importance to auditory scene analysis by echolocation in bats. In the perceptual experiments, FM bats (Eptesicus fuscus) learned to discriminate between systematic and random delay sequences in echo playback sets. The results of these experiments demonstrate that the FM bat can assemble information about echo delay changes over time, a requirement for the analysis of a dynamic auditory scene. Laboratory insect capture experiments examined the vocal production patterns of flying E. fuscus taking tethered insects in a large room. In each trial, the bats consistently produced echolocation signal groups with a relatively stable repetition rate (within 5%). Similar temporal patterning of sonar vocalizations was also observed in the field recordings from E. fuscus, thus suggesting the importance of temporal control of vocal production for perceptually guided behavior. It is hypothesized that a stable sonar signal production rate facilitates the perceptual organization of echoes arriving from objects at different directions and distances as the bat flies through a dynamic auditory scene. Field recordings of E. fuscus, Noctilio albiventris, N. leporinus, Pippistrellus pippistrellus, and Cormura brevirostris revealed that spectral adjustments in sonar signals may also be important to permit tracking of echoes in a complex auditory scene.     

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.     

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.     

Coding of concurrent vocal signals by the auditory midbrain: effects of stimulus level and depth of modulation

The segregation of concurrent vocal signals is an auditory processing task faced by all vocal species. To segregate concurrent signals, the auditory system must encode the spectral and temporal features of the fused waveforms such that at least one signal can be individually detected. In the plainfin midshipman fish (Porichthys notatus), the overlapping mate calls of neighboring males produce acoustic beats with amplitude and phase modulations at the difference frequencies (dF) between spectral components. Prior studies in midshipman have shown that midbrain neurons provide a combinatorial code of the temporal and spectral characteristics of beats via synchronization of spike bursts to dF and changes in spike rate and interspike intervals with changes in spectral composition. In the present study we examine the effects of changes in signal parameters of beats (overall intensity level and depth of modulation) on the spike train outputs of midbrain neurons. The observed changes in spike train parameters further support the hypothesis that midbrain neurons provide a combinatorial code of the spectral and temporal features of concurrent vocal signals.     

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.     

Acoustic correlates of individuality in the vocal repertoire of a nocturnal primate (Microcebus murinus)

In mammals individual distinctiveness in vocalizations provides the basis for individual recognition and thus plays an important role in social behavior. In this study, first evidence is provided for a nocturnal primate that variation in individual distinctiveness across the vocal repertoire is to some extent determined by the context and the acoustic structure of the call types. Individual distinctiveness was investigated across call types in the gray mouse lemur, a nocturnal primate, living in a dispersed multi-male multi-female social system. To explore to what degree context and acoustic structure predict variations in individual distinctiveness, four major call types were examined (grunts, tsaks, short whistles, and trills). Call types differed in context and acoustic structure and were recorded under controlled experimental conditions. A discriminant function analysis revealed that all call types are individually distinct, but not to the same degree. The findings suggest that variations in individual distinctiveness can to some extent be explained by the context and the acoustic structure of the call types.     

An acoustic analysis of laughter produced by congenitally deaf and normally hearing college students

Relatively few empirical data are available concerning the role of auditory experience in nonverbal human vocal behavior, such as laughter production. This study compared the acoustic properties of laughter in 19 congenitally, bilaterally, and profoundly deaf college students and in 23 normally hearing control participants. Analyses focused on degree of voicing, mouth position, air-flow direction, temporal features, relative amplitude, fundamental frequency, and formant frequencies. Results showed that laughter produced by the deaf participants was fundamentally similar to that produced by the normally hearing individuals, which in turn was consistent with previously reported findings. Finding comparable acoustic properties in the sounds produced by deaf and hearing vocalizers confirms the presumption that laughter is importantly grounded in human biology, and that auditory experience with this vocalization is not necessary for it to emerge in species-typical form. Some differences were found between the laughter of deaf and hearing groups; the most important being that the deaf participants produced lower-amplitude and longer-duration laughs. These discrepancies are likely due to a combination of the physiological and social factors that routinely affect profoundly deaf individuals, including low overall rates of vocal fold use and pressure from the hearing world to suppress spontaneous vocalizations.     

Age-related differences in the acoustic characteristics of male leopard seals, Hydrurga leptonyx

During the breeding season, the underwater vocalizations and calling rates of adult male leopard seals are highly stereotyped. In contrast, sub-adult males have more variable acoustic behavior. Although adult males produce only five stereotyped broadcast calls as part of their long-range underwater breeding displays the sub-adults have a greater repertoire including the adult-like broadcast calls, as well as variants of these. Whether this extended repertoire has a social function is unknown due to the paucity of behavioral data for this species. The broadcast calls of the sub-adults are less stereotyped in their acoustic characteristics and they have a more variable calling rate. These age-related differences have major implications for geographic variation studies, where the acoustic behavior of different populations are compared, as well as for acoustic surveying studies, where numbers of calls are used to indicate numbers of individuals present. Sampling regimes which unknowingly include recordings from sub-adult animals will artificially exaggerate differences between populations and numbers of calling animals. The acoustic behavior of sub-adult and adult male leopard seals were significantly different and although this study does not show evidence that these differences reflect vocal learning in the male leopard seal it does suggest that contextual learning may be present.     

The relationship between acoustic structure and semantic information in Diana monkey alarm vocalization

Mammalian vocal production mechanisms are still poorly understood despite their significance for theories of human speech evolution. Particularly, it is still unclear to what degree mammals are capable of actively controlling vocal-tract filtering, a defining feature of human speech production. To address this issue, a detailed acoustic analysis on the alarm vocalization of free-ranging Diana monkeys was conducted. These vocalizations are especially interesting because they convey semantic information about two of the monkeys' natural predators, the leopard and the crowned eagle. Here, vocal tract and sound source parameter in Diana monkey alarm vocalizations are described. It is found that a vocalization-initial formant downward transition distinguishes most reliably between eagle and leopard alarm vocalization. This finding is discussed as an indication of articulation and alternatively as the result of a strong nasalization effect. It is suggested that the formant modulation is the result of active vocal filtering used by the monkeys to encode semantic information, an ability previously thought to be restricted to human speech.     

Rhesus macaques spontaneously perceive formants in conspecific vocalizations

We provide a direct demonstration that nonhuman primates spontaneously perceive changes in formant frequencies in their own species-typical vocalizations, without training or reinforcement. Formants are vocal tract resonances leading to distinctive spectral prominences in the vocal signal, and provide the acoustic determinant of many key phonetic distinctions in human languages. We developed algorithms for manipulating formants in rhesus macaque calls. Using the resulting computer-manipulated calls in a habituation/dishabituation paradigm, with blind video scoring, we show that rhesus macaques spontaneously respond to a change in formant frequencies within the normal macaque vocal range. Lack of dishabituation to a "synthetic replica" signal demonstrates that dishabituation was not due to an artificial quality of synthetic calls, but to the formant shift itself. These results indicate that formant perception, a significant component of human voice and speech perception, is a perceptual ability shared with other primates.     

The synergy between speech production and perception

Speech intelligibility is known to be relatively unaffected by certain deformations of the acoustic spectrum. These include translations, stretching or contracting dilations, and shearing of the spectrum (represented along the logarithmic frequency axis). It is argued here that such robustness reflects a synergy between vocal production and auditory perception. Thus, on the one hand, it is shown that these spectral distortions are produced by common and unavoidable variations among different speakers pertaining to the length, cross-sectional profile, and losses of their vocal tracts. On the other hand, it is argued that these spectral changes leave the auditory cortical representation of the spectrum largely unchanged except for translations along one of its representational axes. These assertions are supported by analyses of production and perception models. On the production side, a simplified sinusoidal model of the vocal tract is developed which analytically relates a few "articulatory" parameters, such as the extent and location of the vocal tract constriction, to the spectral peaks of the acoustic spectra synthesized from it. The model is evaluated by comparing the identification of synthesized sustained vowels to labeled natural vowels extracted from the TIMIT corpus. On the perception side a "multiscale" model of sound processing is utilized to elucidate the effects of the deformations on the representation of the acoustic spectrum in the primary auditory cortex. Finally, the implications of these results for the perception of generally identifiable classes of sound sources beyond the specific case of speech and the vocal tract are discussed.     

Acoustics of children's speech: developmental changes of temporal and spectral parameters

Changes in magnitude and variability of duration, fundamental frequency, formant frequencies, and spectral envelope of children's speech are investigated as a function of age and gender using data obtained from 436 children, ages 5 to 17 years, and 56 adults. The results confirm that the reduction in magnitude and within-subject variability of both temporal and spectral acoustic parameters with age is a major trend associated with speech development in normal children. Between ages 9 and 12, both magnitude and variability of segmental durations decrease significantly and rapidly, converging to adult levels around age 12. Within-subject fundamental frequency and formant-frequency variability, however, may reach adult range about 2 or 3 years later. Differentiation of male and female fundamental frequency and formant frequency patterns begins at around age 11, becoming fully established around age 15. During that time period, changes in vowel formant frequencies of male speakers is approximately linear with age, while such a linear trend is less obvious for female speakers. These results support the hypothesis of uniform axial growth of the vocal tract for male speakers. The study also shows evidence for an apparent overshoot in acoustic parameter values, somewhere between ages 13 and 15, before converging to the canonical levels for adults. For instance, teenagers around age 14 differ from adults in that, on average, they show shorter segmental durations and exhibit less within-subject variability in durations, fundamental frequency, and spectral envelope measures.     

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.     

Acoustic analyses of infant fricative and trill vocalizations

Closants, or consonantlike sounds in infant vocalizations, were described acoustically using 16-kHz spectrograms and LPC or FFT analyses based on waveforms sampled at 20 or 40 kHz. The two major closant types studied were fricatives and trills. Compared to similar fricative sounds in adult speech, the fricative sounds of the 3-, 6-, 9-, and 12-month-old infants had primary spectral components at higher frequencies, i.e., to and above 14 kHz. Trill rate varied from 16-180 Hz with a mean of about 100, approximately four times the mean trill rate reported for adult talkers. Acoustic features are described for various places of articulation for fricatives and trills. The discussion of the data emphasizes dimensions of acoustic contrast that appear in infant vocalizations during the first year of life, and implications of the spectral data for auditory and motor self-stimulation by normal-hearing and hearing-impaired infants.     

Nonlinear analysis of irregular animal vocalizations

Animal vocalizations range from almost periodic vocal-fold vibration to completely atonal turbulent noise. Between these two extremes, a variety of nonlinear dynamics such as limit cycles, subharmonics, biphonation, and chaotic episodes have been recently observed. These observations imply possible functional roles of nonlinear dynamics in animal acoustic communication. Nonlinear dynamics may also provide insight into the degree to which detailed features of vocalizations are under close neural control, as opposed to more directly reflecting biomechanical properties of the vibrating vocal folds themselves. So far, nonlinear dynamical structures of animal voices have been mainly studied with spectrograms. In this study, the deterministic versus stochastic (DVS) prediction technique was used to quantify the amount of nonlinearity in three animal vocalizations: macaque screams, piglet screams, and dog barks. Results showed that in vocalizations with pronounced harmonic components (adult macaque screams, certain piglet screams, and dog barks), deterministic nonlinear prediction was clearly more powerful than stochastic linear prediction. The difference, termed low-dimensional nonlinearity measure (LNM), indicates the presence of a low-dimensional attractor. In highly irregular signals such as juvenile macaque screams, piglet screams, and some dog barks, the detectable amount of nonlinearity was comparatively small. Analyzing 120 samples of dog barks, it was further shown that the harmonic-to-noise ratio (HNR) was positively correlated with LNM. It is concluded that nonlinear analysis is primarily useful in animal vocalizations with strong harmonic components (including subharmonics and biphonation) or low-dimensional chaos.     

Functional Analysis of Voice Using Simultaneous High-Speed Imaging and Acoustic Recordings

We present a comprehensive, functional analysis of clinical voice data derived from both high-speed digital imaging (HSDI) of the larynx and simultaneously acquired acoustic recordings. The goals of this study are to: (1) correlate dynamic characteristics of the vocal folds derived from direct laryngeal imaging with indirectly acquired acoustic measurements; (2) define the advantages of using a combined imaging/acoustic approach for the analysis of voice condition; and (3) identify new quantitative measures to evaluate the regularity of the vocal fold vibration and the complexity of the vocal output -- these measures will be key to successful diagnosis of vocal abnormalities. Image- and acoustic-based analyses are performed using an analytic phase plot approach previously introduced by our group (referred to as 'Nyquist' plot). Fast Fourier Transform (FFT) spectral analyses are performed on the same data for a comparison. Clinical HSDI and acoustic recordings from subjects having normal and specific voice pathologies, including muscular tension dysphonia (MTD) and recurrent respiratory papillomatosis (RRP) were analyzed using the Nyquist plot approach. The results of these analyses show that a combined imaging/acoustic analysis approach provides better characterization of the vibratory behavior of the vocal folds as it correlates with vocal output and pathology.     

Effects of noise on speech production: acoustic and perceptual analyses

Acoustical analyses were carried out on a set of utterances produced by two male speakers talking in quiet and in 80, 90, and 100 dB SPL of masking noise. In addition to replicating previous studies demonstrating increases in amplitude, duration, and vocal pitch while talking in noise, these analyses also found reliable differences in the formant frequencies and short-term spectra of vowels. Perceptual experiments were also conducted to assess the intelligibility of utterances produced in quiet and in noise when they were presented at equal S/N ratios for identification. In each experiment, utterances originally produced in noise were found to be more intelligible than utterances produced in the quiet. The results of the acoustic analyses showed clear and consistent differences in the acoustic-phonetic characteristics of speech produced in quiet versus noisy environments. Moreover, these accounts differences produced reliable effects on intelligibility. The findings are discussed in terms of: (1) the nature of the acoustic changes that taken place when speakers produce speech under adverse conditions such as noise, psychological stress, or high cognitive load: (2) the role of training and feedback in controlling and modifying a talker's speech to improve performance of current speech recognizers; and (3) the development of robust algorithms for recognition of speech in 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.     

Vocal aging and the impact on daily life: a longitudinal study

Longitudinal studies on vocal aging are scarce, and information on the impact of age-related voice changes on daily life is lacking. This longitudinal study reports on age-related voice changes and the impact on daily life over a time period of 5 years on 11 healthy male speakers, age ranging from 50 to 81 years. All males completed a questionnaire on vocal performance in daily life, and perceptual and acoustical analyses of vocal quality and analyses of maximum performance tasks of vocal function (voice range profile) were performed. Results showed a significant deterioration of the acoustic voice signal as well as increased ratings on vocal roughness judged by experts after the time period of 5 years. An increase of self-reported voice instability and the tendency to avoid social parties supported these findings. Smoking males had a lower speaking fundamental frequency compared with nonsmoking males, and this seemed reversible for males who stop smoking. This study suggests a normal gradual vocal aging process with clear consequences in daily life, which should be taken into consideration in clinical practice as well as in studies concerning communication in social life.     

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.