Publications

Abstract

The appreciation of music is a universal trait of humankind.1,2,3 Evidence supporting this notion includes the ubiquity of music across cultures4,5,6,7 and the natural predisposition toward music that humans display early in development.8,9,10 Are we musical animals because of species-specific predispositions? This question cannot be answered by relying on cross-cultural or developmental studies alone, as these cannot rule out enculturation.11 Instead, it calls for cross-species experiments testing whether homologous neural mechanisms underlying music perception are present in non-human primates. We present music to two rhesus monkeys, reared without musical exposure, while recording electroencephalography (EEG) and pupillometry. Monkeys exhibit higher engagement and neural encoding of expectations based on the previously seeded musical context when passively listening to real music as opposed to shuffled controls. We then compare human and monkey neural responses to the same stimuli and find a species-dependent contribution of two fundamental musical features—pitch and timing12—in generating expectations: while timing- and pitch-based expectations13 are similarly weighted in humans, monkeys rely on timing rather than pitch. Together, these results shed light on the phylogeny of music perception. They highlight monkeys’ capacity for processing temporal structures beyond plain acoustic processing, and they identify a species-dependent contribution of time- and pitch-related features to the neural encoding of musical expectations.

Abstract

Bioacoustics increasingly relies on large datasets and computational methods. The need to batch-process large amounts of data and the increased focus on algorithmic processing require software tools. To optimally assist in a bioacoustician’s workflow, software tools need to be as simple and effective as possible. Five years ago, the Python package Parselmouth was released to provide easy and intuitive access to all functionality in the Praat software. Whereas Praat is principally designed for phonetics and speech processing, plenty of bioacoustics studies have used its advanced acoustic algorithms. Here, we evaluate existing usage of Parselmouth and discuss in detail several studies which used the software library. We argue that Parselmouth has the potential to be used even more in bioacoustics research, and suggest future directions to be pursued with the help of Parselmouth.

Abstract

Multimodal rhythmic signalling abounds across animal taxa. Studying its mechanisms and functions can highlight adaptive components in highly complex rhythmic behaviours, like dance and music. Pinnipeds, such as the harbour seal, Phoca vitulina, are excellent comparative models to assess rhythmic capacities. Harbour seals engage in rhythmic percussive behaviours which, until now, have not been described in detail. In our study, eight zoo-housed harbour seals (two pups, two juveniles and four adults) were passively monitored by audio and video during their pupping/breeding season. All juvenile and adult animals performed percussive signalling with their fore flippers in agonistic conditions, both on land and in water. Flipper slap sequences produced on the ground or on the seals' bodies were often highly regular in their interval duration, that is, were quasi-isochronous, at a 200–600 beats/min pace. Three animals also showed significant lateralization in slapping. In contrast to slapping on land, display slapping in water, performed only by adult males, showed slower tempo by one order of magnitude, and a rather motivic temporal structure. Our work highlights that percussive communication is a significant part of harbour seals' behavioural repertoire. We hypothesize that its forms of rhythm production may reflect adaptive functions such as regulating internal states and advertising individual traits.

Abstract

Recursive procedures that allow placing a vocal signal inside another of a similar kind provide a neuro-computational blueprint for syntax and phonology in spoken language and human song. There are, however, no known vocal sequences among nonhuman primates arranged in self-embedded patterns that evince vocal recursion or potential incipient or evolutionary transitional forms thereof, suggesting a neuro-cognitive transformation exclusive to humans. Here, we uncover that wild flanged male orangutan long calls feature rhythmically isochronous call sequences nested within isochronous call sequences, consistent with two hierarchical strata. Remarkably, three temporally and acoustically distinct call rhythms in the lower stratum were not related to the overarching rhythm at the higher stratum by any low multiples, which suggests that these recursive structures were neither the result of parallel non-hierarchical procedures nor anatomical artifacts of bodily constraints or resonances. Findings represent a case of temporally recursive hominid vocal combinatorics in the absence of syntax, semantics, phonology, or music. Second-order combinatorics, ‘sequences within sequences’, involving hierarchically organized and cyclically structured vocal sounds in ancient hominids may have preluded the evolution of recursion in modern language-able humans.

Abstract

Tail wagging is a conspicuous behaviour in domestic dogs (Canis familiaris). Despite how much meaning humans attribute to this display, its quantitative description and evolutionary history are rarely studied. We summarize what is known about the mechanism, ontogeny, function and evolution of this behaviour. We suggest two hypotheses to explain its increased occurrence and frequency in dogs compared to other canids. During the domestication process, enhanced rhythmic tail wagging behaviour could have (i) arisen as a by-product of selection for other traits, such as docility and tameness, or (ii) been directly selected by humans, due to our proclivity for rhythmic stimuli. We invite testing of these hypotheses through neurobiological and ethological experiments, which will shed light on one of the most readily observed yet understudied animal behaviours. Targeted tail wagging research can be a window into both canine ethology and the evolutionary history of characteristic human traits, such as our ability to perceive and produce rhythmic behaviours.

Abstract

Animal vocal communication often relies on call sequences. The temporal patterns of such sequences can be adjusted to other callers, follow complex rhythmic structures or exhibit a metronome-like pattern (i.e., isochronous). How regular are the temporal patterns in animal signals, and what influences their precision? If present, are rhythms already there early in ontogeny? Here, we describe an exploratory study of Cape fur seal (Arctocephalus pusillus pusillus) barks—a vocalisation type produced across many pinniped species in rhythmic, percussive bouts. This study is the first quantitative description of barking in Cape fur seal pups. We analysed the rhythmic structures of spontaneous barking bouts of pups and adult females from the breeding colony in Cape Cross, Namibia. Barks of adult females exhibited isochrony, that is they were produced at fairly regular points in time. Instead, intervals between pup barks were more variable, that is skipping a bark in the isochronous series occasionally. In both age classes, beat precision, that is how well the barks followed a perfect template, was worse when barking at higher rates. Differences could be explained by physiological factors, such as respiration or arousal. Whether, and how, isochrony develops in this species remains an open question. This study provides evidence towards a rhythmic production of barks in Cape fur seal pups and lays the groundwork for future studies to investigate the development of rhythm using multidimensional metrics.

Abstract

thebeat is a Python package for working with temporal sequences and rhythms in the behavioral and cognitive sciences, as well as in bioacoustics. It provides functionality for creating experimental stimuli, and for visualizing and analyzing temporal data. Sequences, sounds, and experimental trials can be generated using single lines of code. thebeat contains functions for calculating common rhythmic measures, such as interval ratios, and for producing plots, such as circular histograms. thebeat saves researchers time when creating experiments, and provides the first steps in collecting widely accepted methods for use in timing research. thebeat is an open-source, on-going, and collaborative project, and can be extended for use in specialized subfields. thebeat integrates easily with the existing Python ecosystem, allowing one to combine our tested code with custom-made scripts. The package was specifically designed to be useful for both skilled and novice programmers. thebeat provides a foundation for working with temporal sequences onto which additional functionality can be built. This combination of specificity and plasticity should facilitate research in multiple research contexts and fields of study.

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

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.

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.

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.

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.

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.

Abstract

Sensitivity to dependencies (correspondences between distant items) in sensory stimuli plays a crucial role in human music and language. Here, we show that squirrel monkeys (Saimiri sciureus) can detect abstract, non-adjacent dependencies in auditory stimuli. Monkeys discriminated between tone sequences containing a dependency and those lacking it, and generalized to previously unheard pitch classes and novel dependency distances. This constitutes the first pattern learning study where artificial stimuli were designed with the species' communication system in mind. These results suggest that the ability to recognize dependencies represents a capability that had already evolved in humans’ last common ancestor with squirrel monkeys, and perhaps before.

Abstract

The possibility of achieving experimentally controlled, non-vocal acoustic production in non-human primates is a key step to enable the testing of a number of hypotheses on primate behavior and cognition. However, no device or solution is currently available, with the use of sensors in non-human animals being almost exclusively devoted to applications in food industry and animal surveillance. Specifically, no device exists which simultaneously allows: (i) spontaneous production of sound or music by non-human animals via object manipulation, (ii) systematical recording of data sensed from these movements, (iii) the possibility to alter the acoustic feedback properties of the object using remote control. We present two prototypes we developed for application with chimpanzees (Pan troglodytes) which, while fulfilling the aforementioned requirements, allow to arbitrarily associate sounds to physical object movements. The prototypes differ in sensing technology, costs, intended use and construction requirements. One prototype uses four piezoelectric elements embedded between layers of Plexiglas and foam. Strain data is sent to a computer running Python through an Arduino board. A second prototype consists in a modified Wii Remote contained in a gum toy. Acceleration data is sent via Bluetooth to a computer running Max/MSP. We successfully pilot tested the first device with a group of chimpanzees. We foresee using these devices for a range of cognitive experiments.

Abstract

Music is a pervasive phenomenon in human culture, and musical
rhythm is virtually present in all musical traditions. Research
on the evolution and cognitive underpinnings of rhythm
can benefit from a number of approaches. We outline key concepts
and definitions, allowing fine-grained analysis of rhythmic
cognition in experimental studies. We advocate comparative
animal research as a useful approach to answer questions
about human music cognition and review experimental evidence
from different species. Finally, we suggest future directions
for research on the cognitive basis of rhythm. Apart from
research in semi-natural setups, possibly allowed by “drum set
for chimpanzees” prototypes presented here for the first time,
mathematical modeling and systematic use of circular statistics
may allow promising advances.