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Timing, rhythm and synchrony in animal Signals (Week 10, 3 ECTS)

In this module we will introduce the major temporal features of acoustic signaling in arthropod and vertebrate animals, including acoustic expression in human language and music. Particular attention will be paid to the role of rhythm in organizing signal features occurring in the time domain.  Having established this general framework, we will examine the ways in which the signal rhythms of multiple individuals displaying within animal groups interact and generate temporal patterns ranging from synchrony to counter-singing (alternation) and more complex ‘orchestral’ arrangements.  Physiological, ecological and evolutionary bases of signaling rhythm and rhythm interaction will be addressed.  Practical sessions will encompass study of current research reported in the primary literature, quantitative and experimental methods for measuring rhythm and rhythm interaction, and modeling approaches used to simulate and test the functions of signaling interaction in animal groups.


Learning outcomes:


By the end of the course, a successful student should be able to:

1. Demonstrate familiarity with the temporal features of signaling and signal interaction in arthropod and vertebrate animals and in the expression of language and music in humans.

2. Demonstrate an understanding of how rhythm interaction may function and of the various factors which may account for such phenomena.

3. Measure the various temporal parameters, and their variation, of signals and signal interaction.

4. Understand how modeling can be used to simulate signal interactions and test how and why these interactions function.


2023 provisional Schedule

Monday 13th November

Morning : Time, rhythm and synchrony in bioacoustics, with emphases on acoustic insects and anurans

Who are the actors ? Phylogenetic perspectives. Sound, song and hearing from the perspective of physics.  Near field and far field features.  Sound vs substrate vibration. How do acoustic insects produce sounds ? Frequency multipliers and resonators.  How do they generate the salient features found in the time domain ? (Michael Greenfield).



Timing and rhythmicity at different temporal scales. Circadian rhythmicity : studies in plasticity. Ultradian rhythm (period < 10 s) ; neural oscillators.

Practical 1  (one hour): Focus on the time domain :  Measurement of temporal acoustic parameters

(Michael Greenfield).


Tuesday 14th November

Morning:  Ultradian rhythm:  sexual selection. Male-male competition. Mate recognition and mate choice. Ultradian rhythm:  recognition and discrimination of natural enemies (M. Greenfield).

Afternoon :  Anuran bioacoustics:  Two speaker playback tests, playback tests with calling males, temporal encoding of information in duets, transmission and signal degradation in the temporal scale, vibrational triggers and inhibitors of activity(Raphael Marquez)


Wednesday 15th November

Morning : Signal and rhythm interaction in the time domain. Rhythm coordination ; entrainment ; synchrony (from crickets to humans). Coupled oscillators. Adaptation and emergence M. Greenfield).

Afternoon: Practical 2  (one hour). Agent-based modeling of coupled oscillators ; Monte Carlo simulation. Phase delay difference models and phase oscillator differential (Kuramoto) models.


Thursday 16th November

Morning: Measuring rhythm; Visualizing rhythmic structure; Rhythm in human music an primates songs (Chiara De Gregorio)

Afternoon: 2 hours Practical: from Praat textgrids to rhythm calculation and visualization. Discussion of results (Chiara De Gregorio).


Friday 17th November

Oral presentations of scientific papers.



The final mark will be based on the oral presentation of a scientific paper (duration = 10 min + 5 min questions).


Organisers: Michael Greenfield & David Reby. Guest tutors: Raphael Marquez & Chiara De Gregorio

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