Andrea Ravignani

Publications

Displaying 1 - 37 of 37
  • Anichini, M., de Reus, K., Hersh, T. A., Valente, D., Salazar-Casals, A., Berry, C., Keller, P. E., & Ravignani, A. (2023). Measuring rhythms of vocal interactions: A proof of principle in harbour seal pups. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 378(1875): 20210477. doi:10.1098/rstb.2021.0477.

    Abstract

    Rhythmic patterns in interactive contexts characterize human behaviours such as conversational turn-taking. These timed patterns are also present in other animals, and often described as rhythm. Understanding fine-grained temporal adjustments in interaction requires complementary quantitative methodologies. Here, we showcase how vocal interactive rhythmicity in a non-human animal can be quantified using a multi-method approach. We record vocal interactions in harbour seal pups (Phoca vitulina) under controlled conditions. We analyse these data by combining analytical approaches, namely categorical rhythm analysis, circular statistics and time series analyses. We test whether pups' vocal rhythmicity varies across behavioural contexts depending on the absence or presence of a calling partner. Four research questions illustrate which analytical approaches are complementary versus orthogonal. For our data, circular statistics and categorical rhythms suggest that a calling partner affects a pup's call timing. Granger causality suggests that pups predictively adjust their call timing when interacting with a real partner. Lastly, the ADaptation and Anticipation Model estimates statistical parameters for a potential mechanism of temporal adaptation and anticipation. Our analytical complementary approach constitutes a proof of concept; it shows feasibility in applying typically unrelated techniques to seals to quantify vocal rhythmic interactivity across behavioural contexts.

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  • Düngen, D., Fitch, W. T., & Ravignani, A. (2023). Hoover the talking seal [quick guide]. Current Biology, 33, R50-R52. doi:10.1016/j.cub.2022.12.023.
  • Düngen, D., & Ravignani, A. (2023). The paradox of learned song in a semi-solitary mammal. Ethology, 129(9), 445-497. doi:10.1111/eth.13385.

    Abstract

    Learning can occur via trial and error; however, learning from conspecifics is faster and more efficient. Social animals can easily learn from conspecifics, but how do less social species learn? In particular, birds provide astonishing examples of social learning of vocalizations, while vocal learning from conspecifics is much less understood in mammals. We present a hypothesis aimed at solving an apparent paradox: how can harbor seals (Phoca vitulina) learn their song when their whole lives are marked by loose conspecific social contact? Harbor seal pups are raised individually by their mostly silent mothers. Pups' first few weeks of life show developed vocal plasticity; these weeks are followed by relatively silent years until sexually mature individuals start singing. How can this rather solitary life lead to a learned song? Why do pups display vocal plasticity at a few weeks of age, when this is apparently not needed? Our hypothesis addresses these questions and tries to explain how vocal learning fits into the natural history of harbor seals, and potentially other less social mammals. We suggest that harbor seals learn during a sensitive period within puppyhood, where they are exposed to adult males singing. In particular, we hypothesize that, to make this learning possible, the following happens concurrently: (1) mothers give birth right before male singing starts, (2) pups enter a sensitive learning phase around weaning time, which (3) coincides with their foraging expeditions at sea which, (4) in turn, coincide with the peak singing activity of adult males. In other words, harbor seals show vocal learning as pups so they can acquire elements of their future song from adults, and solitary adults can sing because they have acquired these elements as pups. We review the available evidence and suggest that pups learn adult vocalizations because they are born exactly at the right time to eavesdrop on singing adults. We conclude by advancing empirical predictions and testable hypotheses for future work.
  • Düngen, D., Sarfati, M., & Ravignani, A. (2023). Cross-species research in biomusicality: Methods, pitfalls, and prospects. In E. H. Margulis, P. Loui, & D. Loughridge (Eds.), The science-music borderlands: Reckoning with the past and imagining the future (pp. 57-95). Cambridge, MA, USA: The MIT Press. doi:10.7551/mitpress/14186.003.0008.
  • Fiveash, A., Ferreri, L., Bouwer, F. L., Kösem, A., Moghimi, S., Ravignani, A., Keller, P. E., & Tillmann, B. (2023). Can rhythm-mediated reward boost learning, memory, and social connection? Perspectives for future research. Neuroscience and Biobehavioral Reviews, 149: 105153. doi:10.1016/j.neubiorev.2023.105153.

    Abstract

    Studies of rhythm processing and of reward have progressed separately, with little connection between the two. However, consistent links between rhythm and reward are beginning to surface, with research suggesting that synchronization to rhythm is rewarding, and that this rewarding element may in turn also boost this synchronization. The current mini review shows that the combined study of rhythm and reward can be beneficial to better understand their independent and combined roles across two central aspects of cognition: 1) learning and memory, and 2) social connection and interpersonal synchronization; which have so far been studied largely independently. From this basis, it is discussed how connections between rhythm and reward can be applied to learning and memory and social connection across different populations, taking into account individual differences, clinical populations, human development, and animal research. Future research will need to consider the rewarding nature of rhythm, and that rhythm can in turn boost reward, potentially enhancing other cognitive and social processes.
  • Gamba, M., Raimondi, T., De Gregorio, C., Valente, D., Carugati, F., Cristiano, W., Ferrario, V., Torti, V., Favaro, L., Friard, O., Giacoma, C., & Ravignani, A. (2023). Rhythmic categories across primate vocal displays. In A. Astolfi, F. Asdrubali, & L. Shtrepi (Eds.), Proceedings of the 10th Convention of the European Acoustics Association Forum Acusticum 2023 (pp. 3971-3974). Torino: European Acoustics Association.

    Abstract

    The last few years have revealed that several species may share the building blocks of Musicality with humans. The recognition of these building blocks (e.g., rhythm, frequency variation) was a necessary impetus for a new round of studies investigating rhythmic variation in animal vocal displays. Singing primates are a small group of primate species that produce modulated songs ranging from tens to thousands of vocal units. Previous studies showed that the indri, the only singing lemur, is currently the only known species that perform duet and choruses showing multiple rhythmic categories, as seen in human music. Rhythmic categories occur when temporal intervals between note onsets are not uniformly distributed, and rhythms with a small integer ratio between these intervals are typical of human music. Besides indris, white-handed gibbons and three crested gibbon species showed a prominent rhythmic category corresponding to a single small integer ratio, isochrony. This study reviews previous evidence on the co-occurrence of rhythmic categories in primates and focuses on the prospects for a comparative, multimodal study of rhythmicity in this clade.
  • De Gregorio, C., Raimondi, T., Bevilacqua, V., Pertosa, C., Valente, D., Carugati, F., Bandoli, F., Favaro, L., Lefaux, B., Ravignani, A., & Gamba, M. (2023). Isochronous singing in 3 crested gibbon species (Nomascusspp.). Current Zoology. Advance online publication. doi:10.1093/cz/zoad029.

    Abstract

    The search for common characteristics between the musical abilities of humans and other animal species is still taking its first steps. One of the most promising aspects from a comparative point of view is the analysis of rhythmic components, which are crucial features of human communicative performance but also well-identifiable patterns in the vocal displays of other species. Therefore, the study of rhythm is becoming essential to understand the mechanisms of singing behavior and the evolution of human communication. Recent findings provided evidence that particular rhythmic structures occur in human music and some singing animal species, such as birds and rock hyraxes, but only 2 species of nonhuman primates have been investigated so far (Indri indri and Hylobates lar). Therefore, our study aims to consistently broaden the list of species studied regarding the presence of rhythmic categories. We investigated the temporal organization in the singing of 3 species of crested gibbons (Nomascus gabriellae, Nomascus leucogenys, and Nomascus siki) and found that the most prominent rhythmic category was isochrony. Moreover, we found slight variation in songs’ tempo among species, with N. gabriellae and N. siki singing with a temporal pattern involving a gradually increasing tempo (a musical accelerando), and N. leucogenys with a more regular pattern. Here, we show how the prominence of a peak at the isochrony establishes itself as a shared characteristic in the small apes considered so far.

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  • Hersh, T. A., Ravignani, A., & Burchardt, L. (2023). Robust rhythm reporting will advance ecological and evolutionary research. Methods in Ecology and Evolution, 14(6), 1398-1407. doi:10.1111/2041-210X.14118.

    Abstract


    Rhythmicity in the millisecond to second range is a fundamental building block of communication and coordinated movement. But how widespread are rhythmic capacities across species, and how did they evolve under different environmental pressures? Comparative research is necessary to answer these questions but has been hindered by limited crosstalk and comparability among results from different study species.
    Most acoustics studies do not explicitly focus on characterising or quantifying rhythm, but many are just a few scrapes away from contributing to and advancing the field of comparative rhythm research. Here, we present an eight-level rhythm reporting framework which details actionable steps researchers can take to report rhythm-relevant metrics. Levels fall into two categories: metric reporting and data sharing. Metric reporting levels include defining rhythm-relevant metrics, providing point estimates of temporal interval variability, reporting interval distributions, and conducting rhythm analyses. Data sharing levels are: sharing audio recordings, sharing interval durations, sharing sound element start and end times, and sharing audio recordings with sound element start/end times.
    Using sounds recorded from a sperm whale as a case study, we demonstrate how each reporting framework level can be implemented on real data. We also highlight existing best practice examples from recent research spanning multiple species. We clearly detail how engagement with our framework can be tailored case-by-case based on how much time and effort researchers are willing to contribute. Finally, we illustrate how reporting at any of the suggested levels will help advance comparative rhythm research.
    This framework will actively facilitate a comparative approach to acoustic rhythms while also promoting cooperation and data sustainability. By quantifying and reporting rhythm metrics more consistently and broadly, new avenues of inquiry and several long-standing, big picture research questions become more tractable. These lines of research can inform not only about the behavioural ecology of animals but also about the evolution of rhythm-relevant phenomena and the behavioural neuroscience of rhythm production and perception. Rhythm is clearly an emergent feature of life; adopting our framework, researchers from different fields and with different study species can help understand why.

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  • Jadoul, Y., & Ravignani, A. (2023). Modelling the emergence of synchrony from decentralized rhythmic interactions in animal communication. Proceedings of the Royal Society B: Biological Sciences, 290(2003). doi:10.1098/rspb.2023.0876.

    Abstract

    To communicate, an animal's strategic timing of rhythmic signals is crucial. Evolutionary, game-theoretical, and dynamical systems models can shed light on the interaction between individuals and the associated costs and benefits of signalling at a specific time. Mathematical models that study rhythmic interactions from a strategic or evolutionary perspective are rare in animal communication research. But new inspiration may come from a recent game theory model of how group synchrony emerges from local interactions of oscillatory neurons. In the study, the authors analyse when the benefit of joint synchronization outweighs the cost of individual neurons sending electrical signals to each other. They postulate there is a benefit for pairs of neurons to fire together and a cost for a neuron to communicate. The resulting model delivers a variant of a classical dynamical system, the Kuramoto model. Here, we present an accessible overview of the Kuramoto model and evolutionary game theory, and of the 'oscillatory neurons' model. We interpret the model's results and discuss the advantages and limitations of using this particular model in the context of animal rhythmic communication. Finally, we sketch potential future directions and discuss the need to further combine evolutionary dynamics, game theory and rhythmic processes in animal communication studies.
  • Jadoul, Y., Düngen, D., & Ravignani, A. (2023). PyGellermann: a Python tool to generate pseudorandom series for human and non-human animal behavioural experiments. BMC Research Notes, 16: 135. doi:10.1186/s13104-023-06396-x.

    Abstract

    Objective

    Researchers in animal cognition, psychophysics, and experimental psychology need to randomise the presentation order of trials in experimental sessions. In many paradigms, for each trial, one of two responses can be correct, and the trials need to be ordered such that the participant’s responses are a fair assessment of their performance. Specifically, in some cases, especially for low numbers of trials, randomised trial orders need to be excluded if they contain simple patterns which a participant could accidentally match and so succeed at the task without learning.
    Results

    We present and distribute a simple Python software package and tool to produce pseudorandom sequences following the Gellermann series. This series has been proposed to pre-empt simple heuristics and avoid inflated performance rates via false positive responses. Our tool allows users to choose the sequence length and outputs a .csv file with newly and randomly generated sequences. This allows behavioural researchers to produce, in a few seconds, a pseudorandom sequence for their specific experiment. PyGellermann is available at https://github.com/YannickJadoul/PyGellermann.
  • Jadoul, Y., Düngen, D., & Ravignani, A. (2023). Live-tracking acoustic parameters in animal behavioural experiments: Interactive bioacoustics with parselmouth. In A. Astolfi, F. Asdrubali, & L. Shtrepi (Eds.), Proceedings of the 10th Convention of the European Acoustics Association Forum Acusticum 2023 (pp. 4675-4678). Torino: European Acoustics Association.

    Abstract

    Most bioacoustics software is used to analyse the already collected acoustics data in batch, i.e., after the data-collecting phase of a scientific study. However, experiments based on animal training require immediate and precise reactions from the experimenter, and thus do not easily dovetail with a typical bioacoustics workflow. Bridging this methodological gap, we have developed a custom application to live-monitor the vocal development of harbour seals in a behavioural experiment. In each trial, the application records and automatically detects an animal's call, and immediately measures duration and acoustic measures such as intensity, fundamental frequency, or formant frequencies. It then displays a spectrogram of the recording and the acoustic measurements, allowing the experimenter to instantly evaluate whether or not to reinforce the animal's vocalisation. From a technical perspective, the rapid and easy development of this custom software was made possible by combining multiple open-source software projects. Here, we integrated the acoustic analyses from Parselmouth, a Python library for Praat, together with PyAudio and Matplotlib's recording and plotting functionality, into a custom graphical user interface created with PyQt. This flexible recombination of different open-source Python libraries allows the whole program to be written in a mere couple of hundred lines of code
  • Lumaca, M., Bonetti, L., Brattico, E., Baggio, G., Ravignani, A., & Vuust, P. (2023). High-fidelity transmission of auditory symbolic material is associated with reduced right–left neuroanatomical asymmetry between primary auditory regions. Cerebral Cortex, 33(11), 6902-6919. doi:10.1093/cercor/bhad009.

    Abstract

    The intergenerational stability of auditory symbolic systems, such as music, is thought to rely on brain processes that allow the faithful transmission of complex sounds. Little is known about the functional and structural aspects of the human brain which support this ability, with a few studies pointing to the bilateral organization of auditory networks as a putative neural substrate. Here, we further tested this hypothesis by examining the role of left–right neuroanatomical asymmetries between auditory cortices. We collected neuroanatomical images from a large sample of participants (nonmusicians) and analyzed them with Freesurfer’s surface-based morphometry method. Weeks after scanning, the same individuals participated in a laboratory experiment that simulated music transmission: the signaling games. We found that high accuracy in the intergenerational transmission of an artificial tone system was associated with reduced rightward asymmetry of cortical thickness in Heschl’s sulcus. Our study suggests that the high-fidelity copying of melodic material may rely on the extent to which computational neuronal resources are distributed across hemispheres. Our data further support the role of interhemispheric brain organization in the cultural transmission and evolution of auditory symbolic systems.
  • Raimondi, T., Di Panfilo, G., Pasquali, M., Zarantonello, M., Favaro, L., Savini, T., Gamba, M., & Ravignani, A. (2023). Isochrony and rhythmic interaction in ape duetting. Proceedings of the Royal Society B: Biological Sciences, 290: 20222244. doi:10.1098/rspb.2022.2244.

    Abstract

    How did rhythm originate in humans, and other species? One cross-cultural universal, frequently found in human music, is isochrony: when note onsets repeat regularly like the ticking of a clock. Another universal consists in synchrony (e.g. when individuals coordinate their notes so that they are sung at the same time). An approach to biomusicology focuses on similarities and differences across species, trying to build phylogenies of musical traits. Here we test for the presence of, and a link between, isochrony and synchrony in a non-human animal. We focus on the songs of one of the few singing primates, the lar gibbon (Hylobates lar), extracting temporal features from their solo songs and duets. We show that another ape exhibits one rhythmic feature at the core of human musicality: isochrony. We show that an enhanced call rate overall boosts isochrony, suggesting that respiratory physiological constraints play a role in determining the song's rhythmic structure. However, call rate alone cannot explain the flexible isochrony we witness. Isochrony is plastic and modulated depending on the context of emission: gibbons are more isochronous when duetting than singing solo. We present evidence for rhythmic interaction: we find statistical causality between one individual's note onsets and the co-singer's onsets, and a higher than chance degree of synchrony in the duets. Finally, we find a sex-specific trade-off between individual isochrony and synchrony. Gibbon's plasticity for isochrony and rhythmic overlap may suggest a potential shared selective pressure for interactive vocal displays in singing primates. This pressure may have convergently shaped human and gibbon musicality while acting on a common neural primate substrate. Beyond humans, singing primates are promising models to understand how music and, specifically, a sense of rhythm originated in the primate phylogeny.
  • Ravignani, A., & Herbst, C. T. (2023). Voices in the ocean: Toothed whales evolved a third way of making sounds similar to that of land mammals and birds. Science, 379(6635), 881-882. doi:10.1126/science.adg5256.
  • Tomasek, M., Ravignani, A., Boucherie, P. H., Van Meyel, S., & Dufour, V. (2023). Spontaneous vocal coordination of vocalizations to water noise in rooks (Corvus frugilegus): An exploratory study. Ecology and Evolution, 13(2): e9791. doi:10.1002/ece3.9791.

    Abstract

    The ability to control one's vocal production is a major advantage in acoustic communication. Yet, not all species have the same level of control over their vocal output. Several bird species can interrupt their song upon hearing an external stimulus, but there is no evidence how flexible this behavior is. Most research on corvids focuses on their cognitive abilities, but few studies explore their vocal aptitudes. Recent research shows that crows can be experimentally trained to vocalize in response to a brief visual stimulus. Our study investigated vocal control abilities with a more ecologically embedded approach in rooks. We show that two rooks could spontaneously coordinate their vocalizations to a long-lasting stimulus (the sound of their small bathing pool being filled with a water hose), one of them adjusting roughly (in the second range) its vocalizations as the stimuli began and stopped. This exploratory study adds to the literature showing that corvids, a group of species capable of cognitive prowess, are indeed able to display good vocal control abilities.
  • Verga, L., Kotz, S. A., & Ravignani, A. (2023). The evolution of social timing. Physics of Life Reviews, 46, 131-151. doi:10.1016/j.plrev.2023.06.006.

    Abstract

    Sociality and timing are tightly interrelated in human interaction as seen in turn-taking or synchronised dance movements. Sociality and timing also show in communicative acts of other species that might be pleasurable, but also necessary for survival. Sociality and timing often co-occur, but their shared phylogenetic trajectory is unknown: How, when, and why did they become so tightly linked? Answering these questions is complicated by several constraints; these include the use of divergent operational definitions across fields and species, the focus on diverse mechanistic explanations (e.g., physiological, neural, or cognitive), and the frequent adoption of anthropocentric theories and methodologies in comparative research. These limitations hinder the development of an integrative framework on the evolutionary trajectory of social timing and make comparative studies not as fruitful as they could be. Here, we outline a theoretical and empirical framework to test contrasting hypotheses on the evolution of social timing with species-appropriate paradigms and consistent definitions. To facilitate future research, we introduce an initial set of representative species and empirical hypotheses. The proposed framework aims at building and contrasting evolutionary trees of social timing toward and beyond the crucial branch represented by our own lineage. Given the integration of cross-species and quantitative approaches, this research line might lead to an integrated empirical-theoretical paradigm and, as a long-term goal, explain why humans are such socially coordinated animals.
  • Gamba, M., De Gregorio, C., Valente, D., Raimondi, T., Torti, V., Miaretsoa, L., Carugati, F., Friard, O., Giacoma, C., & Ravignani, A. (2022). Primate rhythmic categories analyzed on an individual basis. In A. Ravignani, R. Asano, D. Valente, F. Ferretti, S. Hartmann, M. Hayashi, Y. Jadoul, M. Martins, Y. Oseki, E. D. Rodrigues, O. Vasileva, & S. Wacewicz (Eds.), The evolution of language: Proceedings of the Joint Conference on Language Evolution (JCoLE) (pp. 229-236). Nijmegen: Joint Conference on Language Evolution (JCoLE).

    Abstract

    Rhythm is a fundamental feature characterizing communicative displays, and recent studies showed that primate songs encompass categorical rhythms falling on small integer ratios observed in humans. We individually assessed the presence and sexual dimorphism of rhythmic categories, analyzing songs emitted by 39 wild indris. Considering the intervals between the units given during each song, we extracted 13556 interval ratios and found three peaks (at around 0.33, 0.47, and 0.70). Two peaks indicated rhythmic categories corresponding to small integer ratios (1:1, 2:1). All individuals showed a peak at 0.70, and
    most showed those at 0.47 and 0.33. In addition, we found sex differences in the peak at 0.47 only, with males showing lower values than females. This work investigates the presence of individual rhythmic categories in a non-human species; further research may highlight the significance of rhythmicity and untie selective pressures that guided its evolution across species, including humans.
  • Gamba, M., Torti, V., De Gregorio, C., Raimondi, T., Miaretsoa, L., Carugati, F., Cristiano, W., Randrianarison, R. M., Bonadonna, G., Zanoli, A., Friard, O., Valente, D., Ravignani, A., & Giacoma, C. (2022). Caractéristiques rythmiques du chant de l'indri et nouvelles perspectives pour une évaluation comparative du rythme chez les primates non humains. Revue de primatologie, 13. doi:10.4000/primatologie.14989.

    Abstract

    Since the discovery that rhythmic abilities are universal in humans, temporal features of vocal communication have greatly interested researchers studying animal communication. Rhythmic patterns are a valuable tool for species discrimination, mate choice, and individual recognition. A recent study showed that bird songs and human music share rhythmic categories when a signal's temporal intervals are distributed categorically rather than uniformly. Following that study, we aimed to investigate whether songs of indris (Indri indri), the only singing lemur, may show similar features. We measured the inter-onset intervals (tk), delimited by the onsets of two consecutive units, and the rhythmic ratios between these intervals (rk), calculated by dividing an interval by itself plus its adjacent, and finded a three-cluster distribution. Two clusters corresponded to rhythmic categories at 1:1 and 1:2, and the third approached a 2:1 ratio. Our results demonstrated for the first time that another primate besides humans produces categorical rhythms, an ability likely evolved convergently among singing species such as songbirds, indris, and humans. Understanding which communicative features are shared with other species is fundamental to understanding how they have evolved. In this regard, thanks to the simplicity of data processing and interpretation, our study relied on an accessible analytical approach that could open up new branches of the investigation into primate communication, leading the way to reconstruct a phylogeny of rhythm abilities across the entire order.
  • Oswald, J. N., Van Cise, A. M., Dassow, A., Elliott, T., Johnson, M. T., Ravignani, A., & Podos, J. (2022). A collection of best practices for the collection and analysis of bioacoustic data. Applied Sciences, 12(23): 12046. doi:10.3390/app122312046.

    Abstract

    The field of bioacoustics is rapidly developing and characterized by diverse methodologies, approaches and aims. For instance, bioacoustics encompasses studies on the perception of pure tones in meticulously controlled laboratory settings, documentation of species’ presence and activities using recordings from the field, and analyses of circadian calling patterns in animal choruses. Newcomers to the field are confronted with a vast and fragmented literature, and a lack of accessible reference papers or textbooks. In this paper we contribute towards filling this gap. Instead of a classical list of “dos” and “don’ts”, we review some key papers which, we believe, embody best practices in several bioacoustic subfields. In the first three case studies, we discuss how bioacoustics can help identify the ‘who’, ‘where’ and ‘how many’ of animals within a given ecosystem. Specifically, we review cases in which bioacoustic methods have been applied with success to draw inferences regarding species identification, population structure, and biodiversity. In fourth and fifth case studies, we highlight how structural properties in signal evolution can emerge via ecological constraints or cultural transmission. Finally, in a sixth example, we discuss acoustic methods that have been used to infer predator–prey dynamics in cases where direct observation was not feasible. Across all these examples, we emphasize the importance of appropriate recording parameters and experimental design. We conclude by highlighting common best practices across studies as well as caveats about our own overview. We hope our efforts spur a more general effort in standardizing best practices across the subareas we’ve highlighted in order to increase compatibility among bioacoustic studies and inspire cross-pollination across the discipline.
  • Ravignani, A., & Garcia, M. (2022). A cross-species framework to identify vocal learning abilities in mammals. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 377: 20200394. doi:10.1098/rstb.2020.0394.

    Abstract

    Vocal production learning (VPL) is the experience-driven ability to produce novel vocal signals through imitation or modification of existing vocalizations. A parallel strand of research investigates acoustic allometry, namely how information about body size is conveyed by acoustic signals. Recently, we proposed that deviation from acoustic allometry principles as a result of sexual selection may have been an intermediate step towards the evolution of vocal learning abilities in mammals. Adopting a more hypothesis-neutral stance, here we perform phylogenetic regressions and other analyses further testing a potential link between VPL and being an allometric outlier. We find that multiple species belonging to VPL clades deviate from allometric scaling but in the opposite direction to that expected from size exaggeration mechanisms. In other words, our correlational approach finds an association between VPL and being an allometric outlier. However, the direction of this association, contra our original hypothesis, may indicate that VPL did not necessarily emerge via sexual selection for size exaggeration: VPL clades show higher vocalization frequencies than expected. In addition, our approach allows us to identify species with potential for VPL abilities: we hypothesize that those outliers from acoustic allometry lying above the regression line may be VPL species. Our results may help better understand the cross-species diversity, variability and aetiology of VPL, which among other things is a key underpinning of speech in our species.

    This article is part of the theme issue ‘Voice modulation: from origin and mechanism to social impact (Part II)’.

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  • Ravignani, A., Asano, R., Valente, D., Ferretti, F., Hartmann, S., Hayashi, M., Jadoul, Y., Martins, M., Oseki, Y., Rodrigues, E. D., Vasileva, O., & Wacewicz, S. (Eds.). (2022). The evolution of language: Proceedings of the Joint Conference on Language Evolution (JCoLE). Nijmegen: Joint Conference on Language Evolution (JCoLE). doi:10.17617/2.3398549.
  • Ravignani, A. (2022). Language evolution: Sound meets gesture? [Review of the book From signal to symbol: The evolution of language by By R. Planer and K. Sterelny]. Evolutionary Anthropology, 31, 317-318. doi:10.1002/evan.21961.
  • de Reus, K., Carlson, D., Lowry, A., Gross, S., Garcia, M., Rubio-García, A., Salazar-Casals, A., & Ravignani, A. (2022). Body size predicts vocal tract size in a mammalian vocal learner. In A. Ravignani, R. Asano, D. Valente, F. Ferretti, S. Hartmann, M. Hayashi, Y. Jadoul, M. Martins, Y. Oseki, E. D. Rodrigues, O. Vasileva, & S. Wacewicz (Eds.), The evolution of language: Proceedings of the Joint Conference on Language Evolution (JCoLE) (pp. 154-156). Nijmegen: Joint Conference on Language Evolution (JCoLE).
  • de Reus, K., Carlson, D., Lowry, A., Gross, S., Garcia, M., Rubio-Garcia, A., Salazar-Casals, A., & Ravignani, A. (2022). Vocal tract allometry in a mammalian vocal learner. Journal of Experimental Biology, 225(8): jeb243766. doi:10.1242/jeb.243766.

    Abstract

    Acoustic allometry occurs when features of animal vocalisations can be predicted from body size measurements. Despite this being considered the norm, allometry sometimes breaks, resulting in species sounding smaller or larger than expected. A recent hypothesis suggests that allometry-breaking animals cluster into two groups: those with anatomical adaptations to their vocal tracts and those capable of learning new sounds (vocal learners). Here we test this hypothesis by probing vocal tract allometry in a proven mammalian vocal learner, the harbour seal (Phoca vitulina). We test whether vocal tract structures and body size scale allometrically in 68 individuals. We find that both body length and body weight accurately predict vocal tract length and one tracheal dimension. Independently, body length predicts vocal fold length while body weight predicts a second tracheal dimension. All vocal tract measures are larger in weaners than in pups and some structures are sexually dimorphic within age classes. We conclude that harbour seals do comply with allometric constraints, lending support to our hypothesis. However, allometry between body size and vocal fold length seems to emerge after puppyhood, suggesting that ontogeny may modulate the anatomy-learning distinction previously hypothesised as clear-cut. Species capable of producing non-allometric signals while their vocal tract scales allometrically, like seals, may then use non-morphological allometry-breaking mechanisms. We suggest that seals, and potentially other vocal learning mammals, may achieve allometry-breaking through developed neural control over their vocal organs.
  • Verga, L., Sroka, M. G. U., Varola, M., Villanueva, S., & Ravignani, A. (2022). Spontaneous rhythm discrimination in a mammalian vocal learner. Biology Letters, 18: 20220316. doi:10.1098/rsbl.2022.0316.

    Abstract

    Rhythm and vocal production learning are building blocks of human music and speech. Vocal learning has been hypothesized as a prerequisite for rhythmic capacities. Yet, no mammalian vocal learner but humans have shown the capacity to flexibly and spontaneously discriminate rhythmic patterns. Here we tested untrained rhythm discrimination in a mammalian vocal learning species, the harbour seal (Phoca vitulina). Twenty wild-born seals were exposed to music-like playbacks of conspecific call sequences varying in basic rhythmic properties. These properties were called length, sequence regularity, and overall tempo. All three features significantly influenced seals' reaction (number of looks and their duration), demonstrating spontaneous rhythm discrimination in a vocal learning mammal. This finding supports the rhythm–vocal learning hypothesis and showcases pinnipeds as promising models for comparative research on rhythmic phylogenies.
  • Delgado, T., Ravignani, A., Verhoef, T., Thompson, B., Grossi, T., & Kirby, S. (2018). Cultural transmission of melodic and rhythmic universals: Four experiments and a model. In C. Cuskley, M. Flaherty, H. Little, L. McCrohon, A. Ravignani, & T. Verhoef (Eds.), Proceedings of the 12th International Conference on the Evolution of Language (EVOLANG XII) (pp. 89-91). Toruń, Poland: NCU Press. doi:10.12775/3991-1.019.
  • Kotz, S. A., Ravignani, A., & Fitch, W. T. (2018). The evolution of rhythm processing. Trends in Cognitive Sciences, 22(10), 896-910. doi:10.1016/j.tics.2018.08.002.
  • Lumaca, M., Ravignani, A., & Baggio, G. (2018). Music evolution in the laboratory: Cultural transmission meets neurophysiology. Frontiers in Neuroscience, 12: 246. doi:10.3389%2Ffnins.2018.00246.

    Abstract

    In recent years, there has been renewed interest in the biological and cultural evolution of music, and specifically in the role played by perceptual and cognitive factors in shaping core features of musical systems, such as melody, harmony, and rhythm. One proposal originates in the language sciences. It holds that aspects of musical systems evolve by adapting gradually, in the course of successive generations, to the structural and functional characteristics of the sensory and memory systems of learners and “users” of music. This hypothesis has found initial support in laboratory experiments on music transmission. In this article, we first review some of the most important theoretical and empirical contributions to the field of music evolution. Next, we identify a major current limitation of these studies, i.e., the lack of direct neural support for the hypothesis of cognitive adaptation. Finally, we discuss a recent experiment in which this issue was addressed by using event-related potentials (ERPs). We suggest that the introduction of neurophysiology in cultural transmission research may provide novel insights on the micro-evolutionary origins of forms of variation observed in cultural systems.
  • Ravignani, A. (2018). Darwin, sexual selection, and the origins of music. Trends in Ecology and Evolution, 33(10), 716-719. doi:10.1016/j.tree.2018.07.006.

    Abstract

    Humans devote ample time to produce and perceive music. How and why this behavioral propensity originated in our species is unknown. For centuries, speculation dominated the study of the evolutionary origins of musicality. Following Darwin’s early intuitions, recent empirical research is opening a new chapter to tackle this mystery.
  • Ravignani, A. (2018). Comment on “Temporal and spatial variation in harbor seal (Phoca vitulina L.) roar calls from southern Scandinavia” [J. Acoust. Soc. Am. 141, 1824-1834 (2017)]. The Journal of the Acoustical Society of America, 143, 504-508. doi:10.1121/1.5021770.

    Abstract

    In their recent article, Sabinsky and colleagues investigated heterogeneity in harbor seals' vocalizations. The authors found seasonal and geographical variation in acoustic parameters, warning readers that recording conditions might account for some of their results. This paper expands on the temporal aspect of the encountered heterogeneity in harbor seals' vocalizations. Temporal information is the least susceptible to variable recording conditions. Hence geographical and seasonal variability in roar timing constitutes the most robust finding in the target article. In pinnipeds, evidence of timing and rhythm in the millisecond range—as opposed to circadian and seasonal rhythms—has theoretical and interdisciplinary relevance. In fact, the study of rhythm and timing in harbor seals is particularly decisive to support or confute a cross-species hypothesis, causally linking the evolution of vocal production learning and rhythm. The results by Sabinsky and colleagues can shed light on current scientific questions beyond pinniped bioacoustics, and help formulate empirically testable predictions.
  • Ravignani, A., Chiandetti, C., & Gamba, M. (2018). L'evoluzione del ritmo. Le Scienze, (04 maggio 2018).
  • Ravignani, A., Thompson, B., Grossi, T., Delgado, T., & Kirby, S. (2018). Evolving building blocks of rhythm: How human cognition creates music via cultural transmission. Annals of the New York Academy of Sciences, 1423(1), 176-187. doi:10.1111/nyas.13610.

    Abstract

    Why does musical rhythm have the structure it does? Musical rhythm, in all its cross-cultural diversity, exhibits
    commonalities across world cultures. Traditionally, music research has been split into two fields. Some scientists
    focused onmusicality, namely the human biocognitive predispositions formusic, with an emphasis on cross-cultural
    similarities. Other scholars investigatedmusic, seen as a cultural product, focusing on the variation in worldmusical
    cultures.Recent experiments founddeep connections betweenmusicandmusicality, reconciling theseopposing views.
    Here, we address the question of how individual cognitive biases affect the process of cultural evolution of music.
    Data from two experiments are analyzed using two complementary techniques. In the experiments, participants
    hear drumming patterns and imitate them. These patterns are then given to the same or another participant to
    imitate. The structure of these initially random patterns is tracked along experimental “generations.” Frequentist
    statistics show how participants’ biases are amplified by cultural transmission, making drumming patterns more
    structured. Structure is achieved faster in transmission within rather than between participants. A Bayesian model
    approximates the motif structures participants learned and created. Our data and models suggest that individual
    biases for musicality may shape the cultural transmission of musical rhythm.

    Additional information

    nyas13610-sup-0001-suppmat.pdf
  • Ravignani, A., Thompson, B., & Filippi, P. (2018). The evolution of musicality: What can be learned from language evolution research? Frontiers in Neuroscience, 12: 20. doi:10.3389/fnins.2018.00020.

    Abstract

    Language and music share many commonalities, both as natural phenomena and as subjects of intellectual inquiry. Rather than exhaustively reviewing these connections, we focus on potential cross-pollination of methodological inquiries and attitudes. We highlight areas in which scholarship on the evolution of language may inform the evolution of music. We focus on the value of coupled empirical and formal methodologies, and on the futility of mysterianism, the declining view that the nature, origins and evolution of language cannot be addressed empirically. We identify key areas in which the evolution of language as a discipline has flourished historically, and suggest ways in which these advances can be integrated into the study of the evolution of music.
  • Ravignani, A. (2018). Spontaneous rhythms in a harbor seal pup calls. BMC Research Notes, 11: 3. doi:10.1186/s13104-017-3107-6.

    Abstract

    Objectives: Timing and rhythm (i.e. temporal structure) are crucial, though historically neglected, dimensions of animal communication. When investigating these in non-human animals, it is often difficult to balance experimental control and ecological validity. Here I present the first step of an attempt to balance the two, focusing on the timing of vocal rhythms in a harbor seal pup (Phoca vitulina). Collection of this data had a clear aim: To find spontaneous vocal rhythms in this individual in order to design individually-adapted and ecologically-relevant stimuli for a later playback experiment. Data description: The calls of one seal pup were recorded. The audio recordings were annotated using Praat, a free software to analyze vocalizations in humans and other animals. The annotated onsets and offsets of vocalizations were then imported in a Python script. The script extracted three types of timing information: the duration of calls, the intervals between calls’ onsets, and the intervals between calls’ maximum-intensity peaks. Based on the annotated data, available to download, I provide simple descriptive statistics for these temporal measures, and compare their distributions.
  • Ravignani, A., Garcia, M., Gross, S., de Reus, K., Hoeksema, N., Rubio-Garcia, A., & de Boer, B. (2018). Pinnipeds have something to say about speech and rhythm. In C. Cuskley, M. Flaherty, H. Little, L. McCrohon, A. Ravignani, & T. Verhoef (Eds.), Proceedings of the 12th International Conference on the Evolution of Language (EVOLANG XII) (pp. 399-401). Toruń, Poland: NCU Press. doi:10.12775/3991-1.095.
  • Ravignani, A., & Verhoef, T. (2018). Which melodic universals emerge from repeated signaling games?: A Note on Lumaca and Baggio (2017). Artificial Life, 24(2), 149-153. doi:10.1162/ARTL_a_00259.

    Abstract

    Music is a peculiar human behavior, yet we still know little as to why and how music emerged. For centuries, the study of music has been the sole prerogative of the humanities. Lately, however, music is being increasingly investigated by psychologists, neuroscientists, biologists, and computer scientists. One approach to studying the origins of music is to empirically test hypotheses about the mechanisms behind this structured behavior. Recent lab experiments show how musical rhythm and melody can emerge via the process of cultural transmission. In particular, Lumaca and Baggio (2017) tested the emergence of a sound system at the boundary between music and language. In this study, participants were given random pairs of signal-meanings; when participants negotiated their meaning and played a “ game of telephone ” with them, these pairs became more structured and systematic. Over time, the small biases introduced in each artificial transmission step accumulated, displaying quantitative trends, including the emergence, over the course of artificial human generations, of features resembling properties of language and music. In this Note, we highlight the importance of Lumaca and Baggio ʼ s experiment, place it in the broader literature on the evolution of language and music, and suggest refinements for future experiments. We conclude that, while psychological evidence for the emergence of proto-musical features is accumulating, complementary work is needed: Mathematical modeling and computer simulations should be used to test the internal consistency of experimentally generated hypotheses and to make new predictions.
  • Ravignani, A., Thompson, B., Lumaca, M., & Grube, M. (2018). Why do durations in musical rhythms conform to small integer ratios? Frontiers in Computational Neuroscience, 12: 86. doi:10.3389/fncom.2018.00086.

    Abstract

    One curious aspect of human timing is the organization of rhythmic patterns in small integer ratios. Behavioral and neural research has shown that adjacent time intervals in rhythms tend to be perceived and reproduced as approximate fractions of small numbers (e.g., 3/2). Recent work on iterated learning and reproduction further supports this: given a randomly timed drum pattern to reproduce, participants subconsciously transform it toward small integer ratios. The mechanisms accounting for this “attractor” phenomenon are little understood, but might be explained by combining two theoretical frameworks from psychophysics. The scalar expectancy theory describes time interval perception and reproduction in terms of Weber's law: just detectable durational differences equal a constant fraction of the reference duration. The notion of categorical perception emphasizes the tendency to perceive time intervals in categories, i.e., “short” vs. “long.” In this piece, we put forward the hypothesis that the integer-ratio bias in rhythm perception and production might arise from the interaction of the scalar property of timing with the categorical perception of time intervals, and that neurally it can plausibly be related to oscillatory activity. We support our integrative approach with mathematical derivations to formalize assumptions and provide testable predictions. We present equations to calculate durational ratios by: (i) parameterizing the relationship between durational categories, (ii) assuming a scalar timing constant, and (iii) specifying one (of K) category of ratios. Our derivations provide the basis for future computational, behavioral, and neurophysiological work to test our model.

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