Linking the genomic signatures of human beat synchronization and learned song in birds.

The development of rhythmicity is foundational to communicative and social behaviours in humans and many other species, and mechanisms of synchrony could be conserved across species. The goal of the current paper is to explore evolutionary hypotheses linking vocal learning and beat synchronization through genomic approaches, testing the prediction that genetic underpinnings of birdsong also contribute to the aetiology of human interactions with musical beat structure. We combined state-of-the-art-genomic datasets that account for underlying polygenicity of these traits: birdsong genome-wide transcriptomics linked to singing in zebra finches, and a human genome-wide association study of beat synchronization. Results of competitive gene set analysis revealed that the genetic architecture of human beat synchronization is significantly enriched for birdsong genes expressed in songbird Area X (a key nucleus for vocal learning, and homologous to human basal ganglia). These findings complement ethological and neural evidence of the relationship between vocal learning and beat synchronization, supporting a framework of some degree of common genomic substrates underlying rhythm-related behaviours in two clades, humans and songbirds (the largest evolutionary radiation of vocal learners). Future cross-species approaches investigating the genetic underpinnings of beat synchronization in a broad evolutionary context are discussed. This article is part of the theme issue 'Synchrony and rhythm interaction: from the brain to behavioural ecology'.

Regularities in zebra finch song beyond the repeated motif.

The proliferation of birdsong research into the neural mechanisms of vocal learning is indebted to the remarkable stereotypy of the zebra finch's song motif. Motifs are composed of several syllables, which birds learn to produce in a fixed order. But at a higher level of organization-the bout-zebra finch song is no longer stereotyped. Song bouts include several repetitions of the motif, which are often linked by a variable number of short "connector" vocalizations. In this conceptual methods paper, we show that combinatorial analysis alone yields an incomplete description of this bout-level structure. In contrast, studying birdsong as a time-varying analog signal can reveal patterns of flexibility in the rhythmic organization of song bouts. Visualizing large song-samples in sorted raster plots shows that motifs are strung together via two distinct categories of connections: tight or loose. Loose connections allow considerable timing variation across renditions. Even among co-tutored birds that acquired similar motifs, we observe strong individual variability in rhythms and temporal plasticity of song bouts. These findings suggest that vocal flexibility could potentially allow individuals to express a variety of behavioral states through their songs, even in species that sing only a single stereotyped motif.

Sexual dimorphism in striatal dopaminergic responses promotes monogamy in social songbirds.

In many songbird species, males sing to attract females and repel rivals. How can gregarious, non-territorial songbirds such as zebra finches, where females have access to numerous males, sustain monogamy? We found that the dopaminergic reward circuitry of zebra finches can simultaneously promote social cohesion and breeding boundaries. Surprisingly, in unmated males but not in females, striatal dopamine neurotransmission was elevated after hearing songs. Behaviorally too, unmated males but not females persistently exchanged mild punishments in return for songs. Song reinforcement diminished when dopamine receptors were blocked. In females, we observed song reinforcement exclusively to the mate's song, although their striatal dopamine neurotransmission was only slightly elevated. These findings suggest that song-triggered dopaminergic activation serves a dual function in social songbirds: as low-threshold social reinforcement in males and as ultra-selective sexual reinforcement in females. Co-evolution of sexually dimorphic reinforcement systems can explain the coexistence of gregariousness and monogamy.

Acoustic sequences in non-human animals: a tutorial review and prospectus.

Animal acoustic communication often takes the form of complex sequences, made up of multiple distinct acoustic units. Apart from the well-known example of birdsong, other animals such as insects, amphibians, and mammals (including bats, rodents, primates, and cetaceans) also generate complex acoustic sequences. Occasionally, such as with birdsong, the adaptive role of these sequences seems clear (e.g. mate attraction and territorial defence). More often however, researchers have only begun to characterise - let alone understand - the significance and meaning of acoustic sequences. Hypotheses abound, but there is little agreement as to how sequences should be defined and analysed. Our review aims to outline suitable methods for testing these hypotheses, and to describe the major limitations to our current and near-future knowledge on questions of acoustic sequences. This review and prospectus is the result of a collaborative effort between 43 scientists from the fields of animal behaviour, ecology and evolution, signal processing, machine learning, quantitative linguistics, and information theory, who gathered for a 2013 workshop entitled, 'Analysing vocal sequences in animals'. Our goal is to present not just a review of the state of the art, but to propose a methodological framework that summarises what we suggest are the best practices for research in this field, across taxa and across disciplines. We also provide a tutorial-style introduction to some of the most promising algorithmic approaches for analysing sequences. We divide our review into three sections: identifying the distinct units of an acoustic sequence, describing the different ways that information can be contained within a sequence, and analysing the structure of that sequence. Each of these sections is further subdivided to address the key questions and approaches in that area. We propose a uniform, systematic, and comprehensive approach to studying sequences, with the goal of clarifying research terms used in different fields, and facilitating collaboration and comparative studies. Allowing greater interdisciplinary collaboration will facilitate the investigation of many important questions in the evolution of communication and sociality.

New measures to establish the evidence base for medical education: identifying educationally sensitive patient outcomes.

Researchers lack the rich evidence base and benchmark patient outcomes needed to evaluate the effectiveness of medical education practice and guide policy. The authors offer a framework for medical education research that focuses on physician-influenced patient outcomes that are potentially sensitive to medical education. Adapting the concept of ambulatory care sensitive conditions, which provided traction to health services research by defining benchmark patient outcomes to measure health system performance, the authors introduce the concept and propose the adoption of educationally sensitive patient outcomes and suggest two measures: patient activation and clinical microsystem activation. They assert that the ultimate goal of medical education is to ensure that measurement of future physicians' competence and skills is based not only on biomedical knowledge and critical clinical skills but also on the ability to translate these competencies into effective patient- and systems-level outcomes. The authors consider methodological approaches and challenges to measuring such outcomes and argue for large, multiinstitutional, prospective cohort studies and the development of a national Database for Research in Education in Academic Medicine to provide the needed infrastructure. They advocate taking the next steps to establish an educational evidence base to guide the academic medical centers of the 21st century in aligning medical education practice with health care delivery that meets the needs of individuals and populations.