What sort of culture would evolve in an island colony of naive founders? This question cannot be studied experimentally in humans. We performed the analogous experiment using socially learned birdsong. Culture is typically viewed as consisting of traits inherited epigenetically, via social learning. However, cultural diversity has species-typical constraints1, presumably of genetic origin. A celebrated, if contentious, example is whether a universal grammar constrains syntactic diversity in human languages2. Oscine songbirds exhibit song learning and provide biologically tractable models of culture: members of a species show individual variation in song3 and geographically separated groups have local song dialects 4,5. Different species exhibit distinct song cultures6,7, suggestive of genetic constraints8,9. Absent such constraints, innovations and copying errors should cause unbounded variation over multiple generations or geographical distance, contrary to observations9. We asked if wild-type song culture might emerge over multiple generations in an isolated colony founded by isolates, and if so, how this might happen and what type of social environment is required10. Zebra finch isolates, unexposed to singing males during development, produce song with characteristics that differ from the wild-type song found in laboratory11 or natural colonies. In tutoring lineages starting from isolate founders, we quantified alterations in song across tutoring generations in two social environments: tutor-pupil pairs in sound-isolated chambers and an isolated semi-natural colony. In both settings, juveniles imitated the isolate tutors, but changed certain characteristics of the songs. These alterations accumulated over learning generations. Consequently, songs evolved toward the wild-type in 3–4 generations. Thus, species-typical song culture can appear de novo. Our study has parallels with language change and evolution12,13. In analogy to models in quantitative genetics14,15, we model song culture as a multi-generational phenotype, partly encoded genetically in an isolate founding population, influenced by environmental variables, and taking multiple generations to emerge.
The developmental trajectory of nervous system dynamics shows hierarchical structure on time scales spanning ten orders of magnitude from milliseconds to years. Analyzing and characterizing this structure poses significant signal processing challenges. In the context of birdsong development, we have previously proposed that an effective way to do this is to use the dynamic spectrum or spectrogram, a classical signal processing tool, computed at multiple time scales in a nested fashion. Temporal structure on the millisecond timescale is normally captured using a short time Fourier analysis, and structure on the second timescale using song spectrograms. Here we use the dynamic spectrum on time series of song features to study the development of rhythm in juvenile zebra finch. The method is able to detect rhythmic structure in juvenile song in contrast to previous characterizations of such song as unstructured. We show that the method can be used to examine song development, the accuracy with which rhythm is imitated, and the variability of rhythms across different renditions of a song. We hope that this technique will provide a standard, automated method for measuring and characterizing song rhythm.
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