Hemi-laryngeal Setup for Studying Vocal Fold Vibration in Three Dimensions

The voice of humans and most non-human mammals is generated in the larynx through self-sustaining oscillation of the vocal folds. Direct visual ...

University of Vienna

Université Lyon/Saint-Etienne, France

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags' responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.

Do red deer stags (Cervus elaphus) use roar fundamental frequency (F0) to assess rivals?

Red deer stags (Cervus elaphus) give two distinct types of roars during the breeding season, the "common roar" and the "harsh roar." Harsh roars are more frequent during contexts of intense competition, and characterized by a set of features that increase their perceptual salience, suggesting that they signal heightened arousal. While common roars have been shown to encode size information and mediate both male competition and female choice, to our knowledge, the specific function of harsh roars during male competition has not yet been studied. Here, we investigate the hypothesis that the specific structure of male harsh roars signals high arousal to competitors. We contrast the behavioral responses of free ranging, harem-holding stags to the playback of harsh roars from an unfamiliar competitor with their response to the playback of common roars from the same animal. We show that males react less strongly to sequences of harsh roars than to sequences of common roars, possibly because they are reluctant to escalate conflicts with highly motivated and threatening unfamiliar males in the absence of visual information. While future work should investigate the response of stags to harsh roars from familiar opponents, our observations remain consistent with the hypothesis that harsh roars may signal motivation during male competition, and illustrate how intrasexual selection can contribute to the diversification of male vocal signals.

Response of red deer stags (Cervus elaphus) to playback of harsh versus common roars.

Electroglottography (EGG) is a widely used noninvasive method that purports to measure changes in relative vocal fold contact area (VFCA) during phonation. Despite its broad application, the putative direct relation between the EGG waveform and VFCA has to date only been formally tested in a single study, suggesting an approximately linear relationship. However, in that study, flow-induced vocal fold (VF) vibration was not investigated. A rigorous empirical evaluation of EGG as a measure of VFCA under proper physiological conditions is therefore still needed.

Relationship Between the Electroglottographic Signal and Vocal Fold Contact Area.

The information conveyed in acoustic signals is a central topic in mammal vocal communication research. Body size is one form of information that can be encoded in calls. Acoustic allometry aims to identify the specific acoustic correlates of body size within the vocalizations of a given species, and formants are often a useful acoustic cue in this context. We conducted a longitudinal investigation of acoustic allometry in domestic piglets (Sus scrofa domesticus), asking whether formants of grunt vocalizations provide information concerning the caller's body size over time. On four occasions, we recorded grunts from 20 kunekune piglets, measured their vocal tract length by means of radiographs (X-rays) and weighed them. Controlling for effects of age and sex, we found that body weight strongly predicts vocal tract length, which in turn determines formant frequencies. We conclude that grunt formant frequencies could allow domestic pigs to assess a signaler's body size as it grows. Further research using playback experiments is needed to determine the perceptual role of formants in domestic pig communication.

Honest signaling in domestic piglets (Sus scrofa domesticus): vocal allometry and the information content of grunt calls.

Determining whether a species' vocal communication system is graded or discrete requires definition of its vocal repertoire. In this context, research on domestic pig (Sus scrofa domesticus) vocalizations, for example, has led to significant advances in our understanding of communicative functions. Despite their close relation to domestic pigs, little is known about wild boar (Sus scrofa) vocalizations. The few existing studies, conducted in the 1970s, relied on visual inspections of spectrograms to quantify acoustic parameters and lacked statistical analysis. Here, we use objective signal processing techniques and advanced statistical approaches to classify 616 calls recorded from semi-free ranging animals. Based on four spectral and temporal acoustic parameters-quartile Q25, duration, spectral flux, and spectral flatness-extracted from a multivariate analysis, we refine and extend the conclusions drawn from previous work and present a statistically validated classification of the wild boar vocal repertoire into four call types: grunts, grunt-squeals, squeals, and trumpets. While the majority of calls could be sorted into these categories using objective criteria, we also found evidence supporting a graded interpretation of some wild boar vocalizations as acoustically continuous, with the extremes representing discrete call types. The use of objective criteria based on modern techniques and statistics in respect to acoustic continuity advances our understanding of vocal variation. Integrating our findings with recent studies on domestic pig vocal behavior and emotions, we emphasize the importance of grunt-squeals for acoustic approaches to animal welfare and underline the need of further research investigating the role of domestication on animal vocal communication.

Structural Classification of Wild Boar (Sus scrofa) Vocalizations.

A fundamental issue in the evolution of communication is the degree to which signals convey accurate ("honest") information about the signaler. In bioacoustics, the assumption that fundamental frequency (f o) should correlate with the body size of the caller is widespread, but this belief has been challenged by various studies, possibly because larynx size and body size can vary independently. In the present comparative study, we conducted excised larynx experiments to investigate this hypothesis rigorously and explore the determinants of f o. Using specimens from eleven primate species, we carried out an inter-specific investigation, examining correlations between the minimum f o produced by the sound source, body size and vocal fold length (VFL). We found that, across species, VFL predicted minimum f o much better than body size, clearly demonstrating the potential for decoupling between larynx size and body size in primates. These findings shed new light on the diversity of primate vocalizations and vocal morphology, highlighting the importance of vocal physiology in understanding the evolution of mammal vocal communication.

Maxime Garcia

Laboratory of Bio-Acoustics,
Dept. of Cognitive Biology,
Laboratory of Bio-Acoustics, Dept. of Cognitive Biology

University of Vienna

Hemi-laryngeal Setup for Studying Vocal Fold Vibration in Three Dimensions

Maxime Garcia

.css-o250i3{font-size:18px;font-family:Open Sans,sans-serif;line-height:21.6px;}Laboratory of Bio-Acoustics,Dept. of Cognitive Biology,Laboratory of Bio-Acoustics, Dept. of Cognitive Biology

University of Vienna

Hemi-laryngeal Setup for Studying Vocal Fold Vibration in Three Dimensions

Laboratory of Bio-Acoustics,
Dept. of Cognitive Biology,
Laboratory of Bio-Acoustics, Dept. of Cognitive Biology