Publications

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.

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.

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.

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)’.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Abstract

This editorial serves a number of purposes. First, it aims at summarizing and discussing 33 accepted contributions to the special issue “The evolution of rhythm cognition: Timing in music and speech.” The major focus of the issue is the cognitive neuroscience of rhythm, intended as a neurobehavioral trait undergoing an evolutionary process. Second, this editorial provides the interested reader with a guide to navigate the interdisciplinary contributions to this special issue. For this purpose, we have compiled Table 1, where methods, topics, and study species are summarized and related across contributions. Third, we also briefly highlight research relevant to the evolution of rhythm that has appeared in other journals while this special issue was compiled. Altogether, this editorial constitutes a summary of rhythm research in music and speech spanning two years, from mid-2015 until mid-2017

Abstract

Vocal communication is a crucial aspect of animal behavior. The mechanism which most mammals use to vocalize relies on three anatomical components. First, air overpressure is generated inside the lower vocal tract. Second, as the airstream goes through the glottis, sound is produced via vocal fold vibration. Third, this sound is further filtered by the geometry and length of the upper vocal tract. Evidence from mammalian anatomy and bioacoustics suggests that some of these three components may covary with an animal’s body size. The framework provided by acoustic allometry suggests that, because vocal tract length (VTL) is more strongly constrained by the growth of the body than vocal fold length (VFL), VTL generates more reliable acoustic cues to an animal’s size. This hypothesis is often tested acoustically but rarely anatomically, especially in pinnipeds. Here, we test the anatomical bases of the acoustic allometry hypothesis in harbor seal pups Phoca vitulina. We dissected and measured vocal tract, vocal folds, and other anatomical features of 15 harbor seals post-mortem. We found that, while VTL correlates with body size, VFL does not. This suggests that, while body growth puts anatomical constraints on how vocalizations are filtered by harbor seals’ vocal tract, no such constraints appear to exist on vocal folds, at least during puppyhood. It is particularly interesting to find anatomical constraints on harbor seals’ vocal tracts, the same anatomical region partially enabling pups to produce individually distinctive vocalizations.

Abstract

Research on the evolution of human speech and phonology benefits from the comparative approach: structural, spectral, and temporal features can be extracted and compared across species in an attempt to reconstruct the evolutionary history of human speech. Here we focus on analytical tools to measure and compare temporal structure in human speech and animal vocalizations. We introduce the reader to a range of statistical methods usable, on the one hand, to quantify rhythmic complexity in single vocalizations, and on the other hand, to compare rhythmic structure between multiple vocalizations. These methods include: time series analysis, distributional measures, variability metrics, Fourier transform, auto- and cross-correlation, phase portraits, and circular statistics. Using computer-generated data, we apply a range of techniques, walking the reader through the necessary software and its functions. We describe which techniques are most appropriate to test particular hypotheses on rhythmic structure, and provide possible interpretations of the tests. These techniques can be equally well applied to find rhythmic structure in gesture, movement, and any other behavior developing over time, when the research focus lies on its temporal structure. This introduction to quantitative techniques for rhythm and timing analysis will hopefully spur additional comparative research, and will produce comparable results across all disciplines working on the evolution of speech, ultimately advancing the field.

Abstract

Isochrony is crucial to the rhythm of human music. Some neural, behavioral and anatomical traits underlying rhythm perception and production are shared with a broad range of species. These may either have a common evolutionary origin, or have evolved into similar traits under different evolutionary pressures. Other traits underlying rhythm are rare across species, only found in humans and few other animals. Isochrony, or stable periodicity, is common to most human music, but isochronous behaviors are also found in many species. It appears paradoxical that humans are particularly good at producing and perceiving isochronous patterns, although this ability does not conceivably confer any evolutionary advantage to modern humans. This article will attempt to solve this conundrum. To this end, we define the concept of isochrony from the present functional perspective of physiology, cognitive neuroscience, signal processing, and interactive behavior, and review available evidence on isochrony in the signals of humans and other animals. We then attempt to resolve the paradox of isochrony by expanding an evolutionary hypothesis about the function that isochronous behavior may have had in early hominids. Finally, we propose avenues for empirical research to examine this hypothesis and to understand the evolutionary origin of isochrony in general.

Abstract

STRUCTURE IN MUSICAL RHYTHM CAN BE MEASURED using a number of analytical techniques. While some techniques—like circular statistics or grammar induction—rely on strong top-down assumptions, assumption-free techniques can only provide limited insights on higher-order rhythmic structure. I suggest that research in music perception and performance can benefit from systematically adopting phase space plots, a visualization technique originally developed in mathematical physics that overcomes the aforementioned limitations. By jointly plotting adjacent interonset intervals (IOI), the motivic rhythmic structure of musical phrases, if present, is visualized geometrically without making any a priori assumptions concerning isochrony, beat induction, or metrical hierarchies. I provide visual examples and describe how particular features of rhythmic patterns correspond to geometrical shapes in phase space plots. I argue that research on music perception and systematic musicology stands to benefit from this descriptive tool, particularly in comparative analyses of rhythm production. Phase space plots can be employed as an initial assumption-free diagnostic to find higher order structures (i.e., beyond distributional regularities) before proceeding to more specific, theory-driven analyses.