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Jadoul, Y., Hersh, T. A., Fernández Domingos, E., Gamba, M., Favaro, L., & Ravignani, A. (2025). An evolutionary model of rhythmic accelerando in animal vocal signalling. PLOS Computational Biology, 21(4): e1013011. doi:10.1371/journal.pcbi.1013011.
Abstract
Animal acoustic communication contains many structural features. Among these, temporal structure, or rhythmicity, is increasingly tested empirically and modelled quantitatively. Accelerando is a rhythmic structure which consists of temporal intervals increasing in rate over a sequence. Why this particular vocal behaviour is widespread in many different animal lineages, and how it evolved, is so far unknown. Here, we use evolutionary game theory and computer simulations to link two rhythmic aspects of animal communication, acceleration and overlap: We test whether rhythmic accelerando could evolve under a pressure for acoustic overlap in time. Our models show that higher acceleration values result in a higher payoff, driven by the higher relative overlap between sequences. The addition of a cost to the payoff matrix models a physiological disadvantage to high acceleration rates and introduces a divergence between an individual’s incentive and the overall payoff of the population. Analysis of the invasion dynamics of acceleration strategies shows a stable, non-invadable range of strategies for moderate acceleration levels. Our computational simulations confirm these results: A simple selective pressure to maximise the expected overlap, while minimising the associated physiological cost, causes an initially isochronous population to evolve towards producing increasingly accelerating sequences until a population-wide equilibrium of rhythmic accelerando is reached. These results are robust to a broad range of parameter values. Overall, our analyses show that if overlap is beneficial, emergent evolutionary dynamics allow a population to gradually start producing accelerating sequences and reach a stable state of moderate acceleration. Finally, our modelling results closely match empirical data recorded from an avian species showing rhythmic accelerando, the African penguin. This shows the productive interplay between theoretical and empirical biology. -
Filippi, P., Jadoul, Y., Ravignani, A., Thompson, B., & de Boer, B. (2016). Seeking Temporal Predictability in Speech: Comparing Statistical Approaches on 18 World Languages. Frontiers in Human Neuroscience, 10: 586. doi:10.3389/fnhum.2016.00586.
Abstract
Temporal regularities in speech, such as interdependencies in the timing of speech events, are thought to scaffold early acquisition of the building blocks in speech. By providing on-line clues to the location and duration of upcoming syllables, temporal structure may aid segmentation and clustering of continuous speech into separable units. This hypothesis tacitly assumes that learners exploit predictability in the temporal structure of speech. Existing measures of speech timing tend to focus on first-order regularities among adjacent units, and are overly sensitive to idiosyncrasies in the data they describe. Here, we compare several statistical methods on a sample of 18 languages, testing whether syllable occurrence is predictable over time. Rather than looking for differences between languages, we aim to find across languages (using clearly defined acoustic, rather than orthographic, measures), temporal predictability in the speech signal which could be exploited by a language learner. First, we analyse distributional regularities using two novel techniques: a Bayesian ideal learner analysis, and a simple distributional measure. Second, we model higher-order temporal structure—regularities arising in an ordered series of syllable timings—testing the hypothesis that non-adjacent temporal structures may explain the gap between subjectively-perceived temporal regularities, and the absence of universally-accepted lower-order objective measures. Together, our analyses provide limited evidence for predictability at different time scales, though higher-order predictability is difficult to reliably infer. We conclude that temporal predictability in speech may well arise from a combination of individually weak perceptual cues at multiple structural levels, but is challenging to pinpoint.Additional information
https://www.frontiersin.org/article/10.3389/fnhum.2016.00586/full#supplementary… -
Geambaşu, A., Ravignani, A., & Levelt, C. C. (2016). Preliminary experiments on human sensitivity to rhythmic structure in a grammar with recursive self-similarity. Frontiers in Neuroscience, 10: 281. doi:10.3389/fnins.2016.00281.
Abstract
We present the first rhythm detection experiment using a Lindenmayer grammar, a self-similar recursive grammar shown previously to be learnable by adults using speech stimuli. Results show that learners were unable to correctly accept or reject grammatical and ungrammatical strings at the group level, although five (of 40) participants were able to do so with detailed instructions before the exposure phase. -
Ravignani, A., Delgado, T., & Kirby, S. (2016). Musical evolution in the lab exhibits rhythmic universals. Nature Human Behaviour, 1: 0007. doi:10.1038/s41562-016-0007.
Abstract
Music exhibits some cross-cultural similarities, despite its variety across the world. Evidence from a broad range of human cultures suggests the existence of musical universals1, here defined as strong regularities emerging across cultures above chance. In particular, humans demonstrate a general proclivity for rhythm2, although little is known about why music is particularly rhythmic and why the same structural regularities are present in rhythms around the world. We empirically investigate the mechanisms underlying musical universals for rhythm, showing how music can evolve culturally from randomness. Human participants were asked to imitate sets of randomly generated drumming sequences and their imitation attempts became the training set for the next participants in independent transmission chains. By perceiving and imitating drumming sequences from each other, participants turned initially random sequences into rhythmically structured patterns. Drumming patterns developed into rhythms that are more structured, easier to learn, distinctive for each experimental cultural tradition and characterized by all six statistical universals found among world music1; the patterns appear to be adapted to human learning, memory and cognition. We conclude that musical rhythm partially arises from the influence of human cognitive and biological biases on the process of cultural evolution. -
Ravignani, A., & Cook, P. F. (2016). The evolutionary biology of dance without frills. Current Biology, 26(19), R878-R879. doi:10.1016/j.cub.2016.07.076.
Abstract
Recently psychologists have taken up the question of whether dance is reliant on unique human adaptations, or whether it is rooted in neural and cognitive mechanisms shared with other species 1, 2. In its full cultural complexity, human dance clearly has no direct analog in animal behavior. Most definitions of dance include the consistent production of movement sequences timed to an external rhythm. While not sufficient for dance, modes of auditory-motor timing, such as synchronization and entrainment, are experimentally tractable constructs that may be analyzed and compared between species. In an effort to assess the evolutionary precursors to entrainment and social features of human dance, Laland and colleagues [2] have suggested that dance may be an incidental byproduct of adaptations supporting vocal or motor imitation — referred to here as the ‘imitation and sequencing’ hypothesis. In support of this hypothesis, Laland and colleagues rely on four convergent lines of evidence drawn from behavioral and neurobiological research on dance behavior in humans and rhythmic behavior in other animals. Here, we propose a less cognitive, more parsimonious account for the evolution of dance. Our ‘timing and interaction’ hypothesis suggests that dance is scaffolded off of broadly conserved timing mechanisms allowing both cooperative and antagonistic social coordination.Additional information
Experimental Procedures and Two Tables -
Ravignani, A., Fitch, W. T., Hanke, F. D., Heinrich, T., Hurgitsch, B., Kotz, S. A., Scharff, C., Stoeger, A. S., & de Boer, B. (2016). What pinnipeds have to say about human speech, music, and the evolution of rhythm. Frontiers in Neuroscience, 10: 274. doi:10.3389/fnins.2016.00274.
Abstract
Research on the evolution of human speech and music benefits from hypotheses and data generated in a number of disciplines. The purpose of this article is to illustrate the high relevance of pinniped research for the study of speech, musical rhythm, and their origins, bridging and complementing current research on primates and birds. We briefly discuss speech, vocal learning, and rhythm from an evolutionary and comparative perspective. We review the current state of the art on pinniped communication and behavior relevant to the evolution of human speech and music, showing interesting parallels to hypotheses on rhythmic behavior in early hominids. We suggest future research directions in terms of species to test and empirical data needed.
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