Vocal Signals Research Articles

Cues to Androgens and Quality in Male Gibbon Songs

Animal vocal signals may provide information about senders and mediate important social interactions like sexual competition, territory maintenance and mate selection. Hence, it is important to understand whether vocal signals provide accurate information about animal attributes or status. Gibbons are non-human primates that produce loud, distinctive and melodious vocalizations resembling more those of birds than of other non-human primates. Wild gibbons are characterized by flexibility in social organization (i.e., pairs and multimale units) as well as in mating system (i.e., monogamy and polyandry). Such features make them a suitable model to investigate whether the physiology (hormonal status) and socio-demographic features find their correspondence in the structure of their songs. By combining male solo song recordings, endocrine outputs using non-invasive fecal androgen measures and behavioral observations, we studied 14 groups (10 pair-living, 4 multimale) of wild white-handed gibbons (Hylobates lar) residing at Khao Yai National Park, Thailand. We collected a total of 322 fecal samples and recorded 48 songs from 18 adult animals. Our results confirmed inter-individuality in male gibbon songs, and showed a clear correlation between androgen levels and song structures. Gibbons with higher androgen levels produced calls having higher pitch, and similarly adult individuals produced longer calls than senior males. Thus, it is plausible that gibbon vocalizations provide receivers with information about singers' attributes.

The Ascending Mesolimbic Cholinergic System—A Specific Division of the Reticular Activating System Involved in the Initiation of Negative Emotional States

The review summarizes evidences from extensive studies suggesting that ascending mesolimbic cholinergic system (AMCS) that terminates in vast areas of forebrain and diencephalic limbic areas is responsible for specific generation of aversive arousal and aversive emotional state. This state is accompanied by emission of threatening and/or alarming vocalizations that served as a quantitative measure of the emotional response. The AMCS originates from the cholinergic neurons within the laterodorsal tegmental nucleus that have widespread and diffuse ascending connections. Activity of the AMCS induced by activation of the muscarinic cholinergic receptors in the terminal fields of this system, or by glutamate stimulation of neurons of the laterodorsal tegmental nucleus, brings about aversive state with alarming vocalizations. It is postulated that release of acetylcholine from the terminals of the AMCS in the vast areas of the forebrain and diencephalon serves as the initiator of the aversive emotional state with concomitant manifestations and alarming vocal signaling. It is concluded that the AMCS serves as a specific physiological, psychological, and social arousing and alarming system.

Individual vocal signatures in barn owl nestlings: does individual recognition have an adaptive role in sibling vocal competition?

To compete over limited parental resources, young animals communicate with their parents and siblings by producing honest vocal signals of need. Components of begging calls that are sensitive to food deprivation may honestly signal need, whereas other components may be associated with individual-specific attributes that do not change with time such as identity, sex, absolute age and hierarchy. In a sib-sib communication system where barn owl (Tyto alba) nestlings vocally negotiate priority access to food resources, we show that calls have individual signatures that are used by nestlings to recognize which siblings are motivated to compete, even if most vocalization features vary with hunger level. Nestlings were more identifiable when food-deprived than food-satiated, suggesting that vocal identity is emphasized when the benefit of winning a vocal contest is higher. In broods where siblings interact iteratively, we speculate that individual-specific signature permits siblings to verify that the most vocal individual in the absence of parents is the one that indeed perceived the food brought by parents. Individual recognition may also allow nestlings to associate identity with individual-specific characteristics such as position in the within-brood dominance hierarchy. Calls indeed revealed age hierarchy and to a lower extent sex and absolute age. Using a cross-fostering experimental design, we show that most acoustic features were related to the nest of origin (but not the nest of rearing), suggesting a genetic or an early developmental effect on the ontogeny of vocal signatures. To conclude, our study suggests that sibling competition has promoted the evolution of vocal behaviours that signal not only hunger level but also intrinsic individual characteristics such as identity, family, sex and age.

The acoustic signature of the male northern elephant seal: Individual variation supports recognition during competitive interactions

Northern elephant seals (Mirounga angustirostris) have a polygynous breeding system in which adult males establish dominance hierarchies that determine access to females. Acoustic signaling plays an important role in settling fights between males, as stereotyped displays elicit appropriate behavioral responses from individuals without contact during an energetically demanding breeding season. To determine whether reliable differences exist in the acoustic displays of individuals and whether these differences function to convey identity, we behaviorally and acoustically sampled male seals during the breeding season. Vocalizations were recorded during competitive interactions and analyzed for spectral, temporal, and amplitude characteristics. A cross-validated discriminant function analysis revealed small differences within—and significant differences between—the calls produced by 17 adult males of known dominance status. To determine whether acoustic displays serve as individual signatures that males learn to recognize during the breeding season, we conducted playback experiments to test if having prior experience with a particular caller would influence the approach or avoidance response of the listener. Our findings reveal that these unique acoustic signals serve as individual vocal signatures, and males likely remember the identity of their rivals based on call features that have been associated with the outcome of previous competitive interactions.

하모닉 정보를 이용한 SAOC의 보컬 신호 제거 방법에 관한 연구

IAS는 대게 사용자가 자신의 취향에 맞는 음악을 직접 제작 및 편집 가능한 기능을 제공하는 서비스이다. SAOC는 낮은 전송률로 IAS가 가능한 다객체 오디오 코딩 기술이다. 하지만 SAOC 기법은 특정 객체를 제거하는 경우, 특히 보컬 객체를 제거하는 경우 배경음악에 보컬 객체의 하모닉이 남아있는 문제점이 있다. 그래서 본 논문은 하모닉 추출과 제거를 사용한 보컬 객체 제거 기법을 제안한다. 제안 하는 기법은 부호화기에서 추출한 하모닉 정보를 이용하여 복호화기에서 보컬 객체 신호를 다운믹스 신호에서 제거하는 기법이다. 하모닉 정보로써, 기본 주파수, MVF, 하모닉 크기를 사용한다. 성능평가로 객관적, 주관적 실험을 수행하였으며 모든 실험 결과를 통해 SAOC 기법보다 제안하는 기법이 우수함을 확인한다. Interactive audio service provide with audio generating and editing functionality according to user's preference. A spatial audio object coding (SAOC) scheme is audio coding technology that can support the interactive audio service with relatively low bit-rate. However, when the SAOC scheme remove the specific one object such as vocal object signal for Karaoke mode, the scheme support poor quality because the removed vocal object remain in the SAOC-decoded background music. Thus, we propose a new SAOC vocal harmonic extranction and elimination technique to improve the background music quality in the Karaoke service. Namely, utilizing the harmonic information of the vocal object, we removed the harmonics of the vocal object remaining in the background music. As harmonic parameters, we utilize the pitch, MVF(maximum voiced frequency), and harmonic amplitude. To evaluate the performance of the proposed scheme, we perform the objective and subjective evaluation. As our experimental results, we can confirm that the background music quality is improved by the proposed scheme comparing with the SAOC scheme.

Entropy rate as a measure of animal vocal complexity

Vocal complexity is an important concept for investigating the role and evolution of animal communication and sociality. However, no one definition of ‘complexity’ appears to be appropriate for all uses. Repertoire size has been used to quantify complexity in many bird and some mammalian studies, but is impractical in cases where vocalizations are highly diverse, and repertoire size is essentially non-limited at realistic sample sizes. Some researchers have used information-theoretic measures such as Shannon entropy, to describe vocal complexity, but these techniques are descriptive only, as they do not address hypotheses of the cognitive mechanisms behind vocal signal generation. In addition, it can be shown that simple measures of entropy, in particular, do not capture syntactic structure. In this work, I demonstrate the use of an alternative information-theoretic measure, the Markov entropy rate, which quantifies the diversity of transitions in a vocal sequence, and thus is capable of distinguishing sequences with syntactic structure from those generated by random, statistically independent processes. I use artificial sequences generated from different stochastic mechanisms, as well as real data from the vocalizations of the rock hyrax Procavia capensis, to show how different complexity metrics scale differently with sample size. I show that entropy rate provides a good measure of complexity for Markov processes and converges faster than repertoire size estimates, such as the Lempel–Ziv metric. The commonly used Shannon entropy performs poorly in quantifying complexity.

Chimpanzee alarm call production meets key criteria for intentionality.

Determining the intentionality of primate communication is critical to understanding the evolution of human language. Although intentional signalling has been claimed for some great ape gestural signals, comparable evidence is currently lacking for their vocal signals. We presented wild chimpanzees with a python model and found that two of three alarm call types exhibited characteristics previously used to argue for intentionality in gestural communication. These alarm calls were: (i) socially directed and given to the arrival of friends, (ii) associated with visual monitoring of the audience and gaze alternations, and (iii) goal directed, as calling only stopped when recipients were safe from the predator. Our results demonstrate that certain vocalisations of our closest living relatives qualify as intentional signals, in a directly comparable way to many great ape gestures. We conclude that our results undermine a central argument of gestural theories of language evolution and instead support a multimodal origin of human language.

The vocal repertoire of adult male blue monkeys (Cercopithecus mitis stulmanni): A quantitative analysis of acoustic structure

Vocal signals are key elements in understanding species' behavior, sociality, and evolution. Quantified repertoires serve as foundations for investigating usage and function of particular signals, and also provide a basis for comparative analyses among individuals, populations, and taxa to explore how entire signal systems evolve. This study presents a descriptive catalogue of all vocal signals used by adult male blue monkeys (Cercopithecus mitis stuhlmanni). During 12 months in the Kakamega Forest, Kenya, I observed and digitally recorded vocal behavior of 32 adult males across a variety of socioecological contexts. From recordings, I measured 18 temporal-frequency parameters. Undirected ordination and hierarchical cluster analysis identified six distinct call types regularly used by males: ant, boom, ka, katrain, nasal scream, and pyow. Cross-validated discriminant function analysis supported the classifications. The repertoire is best described as discrete, though some gradation occurs between pyows and ants. Summary of acoustic structure and exemplar spectrograms are provided for each call type, along with preliminary examination of socioecological contexts in which they were produced. Discussion addresses repertoire structure, similarity to other taxa, and potential for functional inferences.

Masculine Men Articulate Less Clearly

In previous research, acoustic characteristics of the male voice have been shown to signal various aspects of mate quality and threat potential. But the human voice is also a medium of linguistic communication. The present study explores whether physical and vocal indicators of male mate quality and threat potential are linked to effective communicative behaviors such as vowel differentiation and use of more salient phonetic variants of consonants. We show that physical and vocal indicators of male threat potential, height and formant position, are negatively linked to vowel space size, and that height and levels of circulating testosterone are negatively linked to the use of the aspirated variant of the alveolar stop consonant /t/. Thus, taller, more masculine men display less clarity in their speech and prefer phonetic variants that may be associated with masculine attributes such as toughness. These findings suggest that vocal signals of men's mate quality and/or dominance are not confined to the realm of voice acoustics but extend to other aspects of communicative behavior, even if this means a trade-off with speech patterns that are considered communicatively advantageous, such as clarity and indexical cues to higher social class.

Escalation of aggressive vocal signals: a sequential playback study

Rival conspecifics often produce stereotyped sequences of signals as agonistic interactions escalate. Successive signals in sequence are thought to convey increasingly pronounced levels of aggressive motivation. Here, we propose and test a model of aggressive escalation in black-throated blue warblers, presenting subjects with two sequential and increasingly elevated levels of threat. From a speaker outside the territorial boundary, we initiated an interaction (low-threat level), and from a second speaker inside the territory, accompanied by a taxidermic mount, we subsequently simulated a territorial intrusion (escalated threat level). Our two main predictions were that signalling behaviours in response to low-threat boundary playback would predict signalling responses to the escalated within-territory threat, and that these latter signalling behaviours would in turn reliably predict attack. We find clear support for both predictions: (i) specific song types (type II songs) produced early in the simulated interaction, in response to boundary playback, predicted later use of low-amplitude 'soft' song, in response to within-territory playback; and (ii) soft song, in turn, predicted attack of the mount. Unexpectedly, use of the early-stage signal (type II song) itself did not predict attack, despite its apparent role in aggressive escalation. This raises the intriguing question of whether type II song can actually be considered a reliable aggressive signal. Overall, our results provide new empirical insights into how songbirds may use progressive vocal signalling to convey increasing levels of threat.

Ultrasonic communication in Gliridae (Rodentia): the hazel dormouse (Muscardinus avellanarius) as a case study

Ultrasonic vocalizations (USVs) have been detected in a range of small mammals, particularly rodents, and are associated not only with different social behaviours, e.g. sexual behaviour and aggression, but also with non-social behaviour in several species of mammals, including echolocation and as a by-product of respiration. Rodents in the Gliridae family are known to broadcast a variety of signals, but the behavioural relevance of the latter is poorly known. The aim of our study was to describe the vocal signals of Muscardinus avellanarius and to assess their function. By recording vocal and non-vocal behaviour of captive individuals in different social contexts, we demonstrate that M. avellanarius emits USVs and provide contextual evidence that such sounds are largely used for social communication. We identified six different vocalizations with an overall frequency range between 6.5 and 52.1 kHz, five out of six being ultrasonic (>18 kHz). The vocal repertoire in M. avellanarius was associated with social behaviours, e.g. mother–infant reunion, isolation and paired opposite-sex courtship, all activities involving individuals that are out of mutual visual contact. We therefore infer that this repertoire constitutes a medium-distance (in the range of a few metres) communication system. Further research testing different social contexts both in nature and in captivity is needed for a more complete assessment of the vocal repertoire of this species and its function.

Vocal Recruitment for Joint Travel in Wild Chimpanzees

Joint travel is a common social activity of many group-living animals, which requires some degree of coordination, sometimes through communication signals. Here, we studied the use of an acoustically distinct vocalisation in chimpanzees, the ‘travel hoo’, a signal given specifically in the travel context. We were interested in how this call type was produced to coordinate travel, whether it was aimed at specific individuals and how recipients responded. We found that ‘travel hoos’ were regularly given prior to impending departures and that silent travel initiations were less successful in recruiting than vocal initiations. Other behaviours associated with departure were unrelated to recruitment, suggesting that ‘travel hoos’ facilitated joint travel. Crucially, ‘travel hoos’ were more often produced in the presence of allies than other individuals, with high rates of recruitment success. We discuss these findings as evidence for how motivation to perform a specific social activity can lead to the production of a vocal signal that qualifies as ‘intentional’ according to most definitions, suggesting that a key psychological component of human language may have already been present in the common ancestor of chimpanzees and humans.

Vocal signature in capybara, Hydrochoerus hydrochaeris

Social groups of capybaras are stable and cohesive. The species’ vocal communication is complex and mediates social interaction. The click call is emitted in a variety of contexts by animals from all age groups, but differs among groups; its attributed function is to keep contact among animals. To evaluate the presence of individual characteristics in the click call of capybaras, we recorded the vocalizations emitted spontaneously by six adults kept either solitary or in groups. We selected and measured the acoustic parameters of 300 click call phrases, 50 per individual. The parameters were submitted to a discriminant function analysis that revealed a classification accuracy of 76.8 %. A General Linear Model analysis revealed significant differences among the six individuals, and post hoc results showed that differences between a given pair were different from those of any other pair. The acoustic parameters that most contributed to discriminate the individual calls were click interval duration and click duration, suggesting that temporal parameters are more important than frequency parameters for individuals’ discrimination. The findings of individual characteristics in the click calls indicate that these vocalizations can be used as vocal signatures during social interactions.

Mother–offspring recognition in the bat Carollia perspicillata

Parental care is crucial for offspring survival in many taxa but its burden and costs are often not equally distributed between the sexes. In bats, the majority of parental care is provided by females, making examples of paternal support towards pups exceedingly rare. One exception to this general pattern seemed to be the polygynous Seba's short-tailed fruit bat Carollia perspicillata; two earlier studies suggested that paternal care occurs, i.e. that harem males prompt females to retrieve vocalizing pups. To corroborate this suggestion, we investigated the occurrence of maternal and paternal care in reaction to pup isolation calls in C. perspicillata. Acoustic measurements of 905 isolation calls from 17 pups revealed sufficient interindividual variation to encode an individual vocal signature. Correspondingly, mothers were capable of using this individual signature to discriminate between their own pups and age-matched pups from other females belonging to their colony. Maternal experience was positively correlated with the strength of response behaviour during playbacks. Thus, our results indicate that pup isolation calls were used to elicit maternal care and that mothers recognize their pups based on an individual signature in isolation calls. However, in contrast to the previous studies mentioned above, we found no evidence that harem males reacted to pup isolation calls by prompting the respective mothers to retrieve their vocalizing pups. Instead, our results demonstrate that harem males engaged in courtship activities that were unaffected by pup isolation calls.

Understanding Vocalization Might Help to Assess Stressful Conditions in Piglets.

Simple SummaryThis research aimed to analyze the possibility of assessing piglets’ welfare using the records of their vocalization. The trial was done in a pig commercial farm, and we recorded the vocal signals from piglets in several stressful exposure situations. Data mining techniques were applied to the processed signals in order to obtain a stress classification using the recorded data. We found that, using the piglets’ vocalization, it was possible to identify the most frequent stressful conditions at the farrowing phase, namely: pain, cold and hunger.Assessing pigs’ welfare is one of the most challenging subjects in intensive pig farming. Animal vocalization analysis is a noninvasive procedure and may be used as a tool for assessing animal welfare status. The objective of this research was to identify stress conditions in piglets reared in farrowing pens through their vocalization. Vocal signals were collected from 40 animals under the following situations: normal (baseline), feeling cold, in pain, and feeling hunger. A unidirectional microphone positioned about 15 cm from the animals’ mouth was used for recording the acoustic signals. The microphone was connected to a digital recorder, where the signals were digitized at the 44,100 Hz frequency. The collected sounds were edited and analyzed. The J48 decision tree algorithm available at the Weka® data mining software was used for stress classification. It was possible to categorize diverse conditions from the piglets’ vocalization during the farrowing phase (pain, cold and hunger), with an accuracy rate of 81.12%. Results indicated that vocalization might be an effective welfare indicator, and it could be applied for assessing distress from pain, cold and hunger in farrowing piglets.

Brief Report: Atypical Expression of Distress During the Separation Phase of the Strange Situation Procedure in Infant Siblings at High Risk for ASD

Previous studies have provided preliminary evidence that disruptions in cry acoustics may be part of an atypical vocal signature of autism early in life. We examined the acoustic characteristics of cries extracted from the separation phase of the strange situation procedure in a sample of toddler of younger siblings of a child with autism spectrum disorder-autism spectrum disorders (ASD) (high risk, HR) and a low risk (LR) group. Cry samples derived from vocal recordings of 15-month-old HR (n=13) and LR infants (n=14) were subjected to acoustic analyses. HR toddlers, compared to those with LR, produced cries that were shorter and had a higher fundamental frequency (F0). Three HR toddlers later classified with an ASD at 36months (autistic disorder in all cases) produced cries that had among the highest F0 and shortest durations. Taken together these results indicate that toddlers at high risk for ASD (and those with an ASD) express atypical patterns of distress in response a social stressor. Implications for early diagnosis and parenting are discussed.

Predicting the Effect of Urban Noise on the Active Space of Avian Vocal Signals

Urbanization changes the physical environment of nonhuman species but also markedly changes their acoustic environment. Urban noise interferes with acoustic communication in a range of animals, including birds, with potentially profound impacts on fitness. However, a mechanistic theory to predict which species of birds will be most affected by urban noise is lacking. We develop a mathematical model to predict the decrease in the active space of avian vocal signals after moving from quiet forest habitats to noisy urban habitats. We find that the magnitude of the decrease is largely a function of signal frequency. However, this relationship is not monotonic. A metaregression of observed increases in the frequency of birdsong in urban noise supports the model's predictions for signals with frequencies between 1.5 and 4 kHz. Using results of the metaregression and the model described above, we show that the expected gain in active space following observed frequency shifts is up to 12% and greatest for birds with signals at the lower end of this frequency range. Our generally applicable model, along with three predictions regarding the behavioral and population-level responses of birds to urban noise, represents an important step toward a theory of acoustic communication in urban habitats.

Thinking one thing, saying another: The behavioral correlates of mind-wandering while reading aloud

Although mind-wandering during silent reading is well documented, to date no research has investigated whether similar processes occur during reading aloud. In the present study, participants read a passage either silently or aloud while periodically being probed about mind-wandering. Although their comprehension accuracies were similar for both reading conditions, participants reported more mind-wandering while they were reading aloud. These episodes of mindless reading were associated with nearly normal prosody, but were nevertheless distinguished by subtle fluctuations in volume that were predictive of both overall comprehension accuracy and individual sentence comprehension. Together, these findings reveal that previously hidden within the common activity of reading aloud lies: (1) a demonstration of the remarkable automaticity of speech, (2) a situation that is surprisingly conducive to mind-wandering, (3) subtle vocal signatures of mind-wandering and comprehension accuracy, and (4) the promise of developing useful interventions to improve reading.

Biasing the perception of ambiguous vocal affect: a TMS study on frontal asymmetry

Several sources of evidence point toward a link between asymmetry of prefrontal brain activity and approach-withdrawal tendencies. Here, we tested the causal nature of this link and examined if the categorization of an ambiguous approach- or withdrawal-related vocal signal can be biased by manipulating left and right frontal neural activity. We used voice morphing of affective non-verbal vocalizations to create individually tailored affectively ambiguous stimuli on an Anger-Fear continuum-two emotions that represent extremes on the approach-withdrawal dimension. We tested perception of these stimuli after 10 min of low-frequency repetitive transcranial magnetic stimulation over left or right dorsolateral prefrontal cortex or over the vertex (control), a technique that has transient inhibitory effects on the targeted brain region. As expected, ambiguous stimuli were more likely perceived as expressing Anger (approach) than Fear (withdrawal) after right prefrontal compared with left prefrontal or control stimulation. These results provide the first evidence that the manipulation of asymmetrical activity in prefrontal cortex can change the explicit categorization of ambiguous emotional signals.

Female Putty-Nosed Monkeys Use Experimentally Altered Contextual Information to Disambiguate the Cause of Male Alarm Calls

Many animal vocal signals are given in a wide range of contexts which can sometimes have little in common. Yet, to respond adaptively, listeners must find ways to identify the cause of a signal, or at least rule out alternatives. Here, we investigate the nature of this process in putty-nosed monkeys, a forest primate. In this species, adult males have a very restricted repertoire of vocalizations which are given in response to a wide variety of events occurring under conditions of limited visibility. We carried out a series of field playback experiments on females (N = 6) in a habituated group in Gashaka Gumti National Park, Nigeria, in which male alarm/loud calls were presented either alone, or following acoustic information that simulated the occurrence of natural disturbances. We demonstrate that listeners appear to integrate contextual information in order to distinguish among possible causes of calls. We conclude that, in many cases, pragmatic aspects of communication play a crucial role in call interpretation and place a premium on listeners' abilities to integrate information from different sources.