Music as a trait in evolutionary theory: A musicological perspective.

Although it can be straightforward to define the features of physical traits, complex cultural categories tend to elude widely accepted definitions that transcend cultural and historical context. Addressing papers by Mehr et al. and Savage et al., which both aim to explain music as an evolved trait, we discuss fundamental problems that arise from their conceptualizations of music.

Perception of Nigerian Dùndún Talking Drum Performances as Speech-Like vs. Music-Like: The Role of Familiarity and Acoustic Cues.

It seems trivial to identify sound sequences as music or speech, particularly when the sequences come from different sound sources, such as an orchestra and a human voice. Can we also easily distinguish these categories when the sequence comes from the same sound source? On the basis of which acoustic features? We investigated these questions by examining listeners' classification of sound sequences performed by an instrument intertwining both speech and music: the dùndún talking drum. The dùndún is commonly used in south-west Nigeria as a musical instrument but is also perfectly fit for linguistic usage in what has been described as speech surrogates in Africa. One hundred seven participants from diverse geographical locations (15 different mother tongues represented) took part in an online experiment. Fifty-one participants reported being familiar with the dùndún talking drum, 55% of those being speakers of Yorùbá. During the experiment, participants listened to 30 dùndún samples of about 7s long, performed either as music or Yorùbá speech surrogate (n = 15 each) by a professional musician, and were asked to classify each sample as music or speech-like. The classification task revealed the ability of the listeners to identify the samples as intended by the performer, particularly when they were familiar with the dùndún, though even unfamiliar participants performed above chance. A logistic regression predicting participants' classification of the samples from several acoustic features confirmed the perceptual relevance of intensity, pitch, timbre, and timing measures and their interaction with listener familiarity. In all, this study provides empirical evidence supporting the discriminating role of acoustic features and the modulatory role of familiarity in teasing apart speech and music.

Mockingbird Morphing Music: Structured Transitions in a Complex Bird Song.

The song of the northern mockingbird, Mimus polyglottos, is notable for its extensive length and inclusion of numerous imitations of several common North American bird species. Because of its complexity, it is not widely studied by birdsong scientists. When they do study it, the specific imitations are often noted, and the total number of varying phrases. What is rarely noted is the systematic way the bird changes from one syllable to the next, often with a subtle transition where one sound is gradually transformed into a related sound, revealing an audible and specific compositional mode. It resembles a common strategy in human composing, which can be described as variation of a theme. In this paper, we present our initial attempts to describe the specific compositional rules behind the mockingbird song, focusing on the way the bird transitions from one syllable type to the next. We find that more often than chance, syllables before and after the transition are spectrally related, i.e., transitions are gradual, which we describe as morphing. In our paper, we categorize four common modes of morphing: timbre change, pitch change, squeeze (shortening in time), and stretch (lengthening in time). This is the first time such transition rules in any complex birdsong have been specifically articulated.

What Makes Babies Musical? Conceptions of Musicality in Infants and Toddlers.

Despite major advances in research on musical ability in infants, relatively little attention has been paid to individual differences in general musicality in infants. A fundamental problem has been the lack of a clear definition of what constitutes "general musicality" or "musical ability" in infants and toddlers, resulting in a wide range of test procedures that rely on different models of musicality. However, musicality can be seen as a social construct that can take on different meanings across cultures, sub-groups, and individuals, and may be subject to change over time. Therefore, one way to get a clearer picture of infant musicality is to assess conceptions of musicality in the general population. Using this approach, we surveyed 174 German adults, asking about their view and conceptions regarding behaviors that characterize a musical child under 3 years. Based on previous studies on adult and child musicality, we designed a survey containing 41 statements describing musical behaviors in children. Participants were asked to rate how indicative these behaviors were of musicality in infants and toddlers. PCA analysis revealed 4 components of musical abilities and behaviors in under-3-year-olds: Musical Communication, Enthusiasm and Motivation, Adaptive Expressiveness, and Musical Abilities as traditionally defined. Professional background and musical expertise of the respondents did not significantly influence participants' conceptions. Our results suggest that, in order to capture musicality in young children, a wider range of skills and observable behaviors should be taken into account than those assessed by traditional musical ability tests for young children.

Categorical Rhythms Are Shared between Songbirds and Humans.

Rhythm is a prominent feature of music. Of the infinite possible ways of organizing events in time, musical rhythms are almost always distributed categorically. Such categories can facilitate the transmission of culture-a feature that songbirds and humans share. We compared rhythms of live performances of music to rhythms of wild thrush nightingale and domestic zebra finch songs. In nightingales, but not in zebra finches, we found universal rhythm categories, with patterns that were surprisingly similar to those of music. Isochronous 1:1 rhythms were similarly common. Interestingly, a bias toward small ratios (around 1:2 to 1:3), which is highly abundant in music, was observed also in thrush nightingale songs. Within that range, however, there was no statistically significant bias toward exact integer ratios (1:2 or 1:3) in the birds. High-ratio rhythms were abundant in the nightingale song and are structurally similar to fusion rhythms (ornaments) in music. In both species, preferred rhythms remained invariant over extended ranges of tempos, indicating natural categories. The number of rhythm categories decreased at higher tempos, with a threshold above which rhythm became highly stereotyped. In thrush nightingales, this threshold occurred at a tempo twice faster than in humans, indicating weaker structural constraints and a remarkable motor proficiency. Together, the results suggest that categorical rhythms reflect similar constraints on learning motor skills across species. The saliency of categorical rhythms across humans and thrush nightingales suggests that they promote, or emerge from, the cultural transmission of learned vocalizations. VIDEO ABSTRACT.

Listening to birdsong reveals basic features of rate perception and aesthetic judgements.

The timing of acoustic events is central to human speech and music. Tempo tends to be slower in aesthetic contexts: rates in poetic speech and music are slower than non-poetic, running speech. We tested whether a general aesthetic preference for slower rates can account for this, using birdsong as a stimulus: it structurally resembles human sequences but is unbiased by their production or processing constraints. When listeners selected the birdsong playback tempo that was most pleasing, they showed no bias towards any range of note rates. However, upon hearing a novel stimulus, listeners rapidly formed a robust, implicit memory of its temporal properties, and developed a stimulus-specific preference for the memorized tempo. Interestingly, tempo perception in birdsong stimuli was strongly determined by individual, internal preferences for rates of 1-2 Hz. This suggests that processing complex sound sequences relies on a default time window, while aesthetic appreciation appears flexible, experience-based and not determined by absolute event rates.

Multifractal analysis reveals music-like dynamic structure in songbird rhythms.

Music is thought to engage its listeners by driving feelings of surprise, tension, and relief through a dynamic mixture of predictable and unpredictable patterns, a property summarized here as "expressiveness". Birdsong shares with music the goal to attract its listeners' attention and might use similar strategies to achieve this. We here tested a thrush nightingale's (Luscinia luscinia) rhythm, as represented by song amplitude envelope (containing information on note timing, duration, and intensity), for evidence of expressiveness. We used multifractal analysis, which is designed to detect in a signal dynamic fluctuations between predictable and unpredictable states on multiple timescales (e.g. notes, subphrases, songs). Results show that rhythm is strongly multifractal, indicating fluctuations between predictable and unpredictable patterns. Moreover, comparing original songs with re-synthesized songs that lack all subtle deviations from the "standard" note envelopes, we find that deviations in note intensity and duration significantly contributed to multifractality. This suggests that birdsong is more dynamic due to subtle note timing patterns, often similar to musical operations like accelerando or crescendo. While different sources of these dynamics are conceivable, this study shows that multi-timescale rhythm fluctuations can be detected in birdsong, paving the path to studying mechanisms and function behind such patterns.

Long-distance retinoid signaling in the zebra finch brain.

All-trans retinoic acid (ATRA), the main active metabolite of vitamin A, is a powerful signaling molecule that regulates large-scale morphogenetic processes during vertebrate embryonic development, but is also involved post-natally in regulating neural plasticity and cognition. In songbirds, it plays an important role in the maturation of learned song. The distribution of the ATRA-synthesizing enzyme, zRalDH, and of ATRA receptors (RARs) have been described, but information on the distribution of other components of the retinoid signaling pathway is still lacking. To address this gap, we have determined the expression patterns of two obligatory RAR co-receptors, the retinoid X receptors (RXR) α and γ, and of the three ATRA-degrading cytochromes CYP26A1, CYP26B1, and CYP26C1. We have also studied the distribution of zRalDH protein using immunohistochemistry, and generated a refined map of ATRA localization, using a modified reporter cell assay to examine entire brain sections. Our results show that (1) ATRA is more broadly distributed in the brain than previously predicted by the spatially restricted distribution of zRalDH transcripts. This could be due to long-range transport of zRalDH enzyme between different nuclei of the song system: Experimental lesions of putative zRalDH peptide source regions diminish ATRA-induced transcription in target regions. (2) Four telencephalic song nuclei express different and specific subsets of retinoid-related receptors and could be targets of retinoid regulation; in the case of the lateral magnocellular nucleus of the anterior nidopallium (lMAN), receptor expression is dynamically regulated in a circadian and age-dependent manner. (3) High-order auditory areas exhibit a complex distribution of transcripts representing ATRA synthesizing and degrading enzymes and could also be a target of retinoid signaling. Together, our survey across multiple connected song nuclei and auditory brain regions underscores the prominent role of retinoid signaling in modulating the circuitry that underlies the acquisition and production of learned vocalizations.

Investigation of musicality in birdsong.

Songbirds spend much of their time learning, producing, and listening to complex vocal sequences we call songs. Songs are learned via cultural transmission, and singing, usually by males, has a strong impact on the behavioral state of the listeners, often promoting affiliation, pair bonding, or aggression. What is it in the acoustic structure of birdsong that makes it such a potent stimulus? We suggest that birdsong potency might be driven by principles similar to those that make music so effective in inducing emotional responses in humans: a combination of rhythms and pitches-and the transitions between acoustic states-affecting emotions through creating expectations, anticipations, tension, tension release, or surprise. Here we propose a framework for investigating how birdsong, like human music, employs the above "musical" features to affect the emotions of avian listeners. First we analyze songs of thrush nightingales (Luscinia luscinia) by examining their trajectories in terms of transitions in rhythm and pitch. These transitions show gradual escalations and graceful modifications, which are comparable to some aspects of human musicality. We then explore the feasibility of stripping such putative musical features from the songs and testing how this might affect patterns of auditory responses, focusing on fMRI data in songbirds that demonstrate the feasibility of such approaches. Finally, we explore ideas for investigating whether musical features of birdsong activate avian brains and affect avian behavior in manners comparable to music's effects on humans. In conclusion, we suggest that birdsong research would benefit from current advances in music theory by attempting to identify structures that are designed to elicit listeners' emotions and then testing for such effects experimentally. Birdsong research that takes into account the striking complexity of song structure in light of its more immediate function - to affect behavioral state in listeners - could provide a useful animal model for studying basic principles of music neuroscience in a system that is very accessible for investigation, and where developmental auditory and social experience can be tightly controlled.

The molecular receptive range of an olfactory receptor in vivo (Drosophila melanogaster Or22a).

Understanding how odors are coded within an olfactory system requires knowledge about its input. This is constituted by the molecular receptive ranges (MRR) of olfactory sensory neurons that converge in the glomeruli of the olfactory bulb (vertebrates) or the antennal lobe (AL, insects). Aiming at a comprehensive characterization of MRRs in Drosophila melanogaster we measured odor-evoked calcium responses in olfactory sensory neurons that express the olfactory receptor Or22a. We used an automated stimulus application system to screen [Ca(2+)] responses to 104 odors both in the antenna (sensory transduction) and in the AL (neuronal transmission). At 10(-2) (vol/vol) dilution, 39 odors elicited at least a half-maximal response. For these odorants we established dose-response relationships over their entire dynamic range. We tested 15 additional chemicals that are structurally related to the most efficient odors. Ethyl hexanoate and methyl hexanoate were the best stimuli, eliciting consistent responses at dilutions as low as 10(-9). Two substances led to calcium decrease, suggesting that Or22a might be constitutively active, and that these substances might act as inverse agonists, reminiscent of G-protein coupled receptors. There was no difference between the antennal and the AL MRR. Furthermore we show that Or22a has a broad yet selective MRR, and must be functionally described both as a specialist and a generalist. Both these descriptions are ecologically relevant. Given that adult Drosophila use approximately 43 ORs, a complete description of all MRRs appears now in reach.