My research aims to translate progress in the speech and music sciences into improved diagnostics and treatments for affective and social dimensions of mental health, including disorders of mood, anxiety, and sociality. See and

My doctoral research in Neurobiology at Duke University School of Medicine focused on the biology of emotional expression in speech and music. My postdoctoral work at the University of Vienna focused on bioacoustics, interpersonal synchrony, and social bonding. I have completed certificate courses in Cognitive Neuroscience and Translational Medicine, as well as undergraduate degrees in Biological Psychology and Neurophilosophy.

I have authored 40 scientific articles in top journals including Science, PNAS, Molecular Psychiatry, Translational Psychiatry, PLoS Biology, Trends in Cognitive Science, and Physics of Life Reviews. My work has been supported by grants from the National Institutes of Health, the Austrian Science Foundation, the Faculty of Life Sciences at the University of Vienna, and the Wu Tsai Neuroscience Institute at Stanford University.

Academic Appointments

All Publications

  • Biological principles for music and mental health. Translational psychiatry Bowling, D. L. 2023; 13 (1): 374


    Efforts to integrate music into healthcare systems and wellness practices are accelerating but the biological foundations supporting these initiatives remain underappreciated. As a result, music-based interventions are often sidelined in medicine. Here, I bring together advances in music research from neuroscience, psychology, and psychiatry to bridge music's specific foundations in human biology with its specific therapeutic applications. The framework I propose organizes the neurophysiological effects of music around four core elements of human musicality: tonality, rhythm, reward, and sociality. For each, I review key concepts, biological bases, and evidence of clinical benefits. Within this framework, I outline a strategy to increase music's impact on health based on standardizing treatments and their alignment with individual differences in responsivity to these musical elements. I propose that an integrated biological understanding of human musicality-describing each element's functional origins, development, phylogeny, and neural bases-is critical to advancing rational applications of music in mental health and wellness.

    View details for DOI 10.1038/s41398-023-02671-4

    View details for PubMedID 38049408

    View details for PubMedCentralID 6688399

  • ACOUSTICALLY-DRIVEN PHONEME REMOVAL THAT PRESERVES VOCAL AFFECT CUES. Proceedings of the ... IEEE International Conference on Acoustics, Speech, and Signal Processing. ICASSP (Conference) Noufi, C., Berger, J., Frank, M., Parker, K., Bowling, D. L. 2023; 2023


    In this paper, we propose a method for removing linguistic information from speech for the purpose of isolating paralinguistic indicators of affect. The immediate utility of this method lies in clinical tests of sensitivity to vocal affect that are not confounded by language, which is impaired in a variety of clinical populations. The method is based on simultaneous recordings of speech audio and electroglotto-graphic (EGG) signals. The speech audio signal is used to estimate the average vocal tract filter response and amplitude envelop. The EGG signal supplies a direct correlate of voice source activity that is mostly independent of phonetic articulation. These signals are used to create a third signal designed to capture as much paralinguistic information from the vocal production system as possible-maximizing the retention of bioacoustic cues to affect-while eliminating phonetic cues to verbal meaning. To evaluate the success of this method, we studied the perception of corresponding speech audio and transformed EGG signals in an affect rating experiment with online listeners. The results show a high degree of similarity in the perceived affect of matched signals, indicating that our method is effective.

    View details for DOI 10.1109/icassp49357.2023.10095942

    View details for PubMedID 37701064

    View details for PubMedCentralID PMC10495117

  • Vocal similarity theory and the biology of musical tonality. Physics of life reviews Bowling, D. L. 2023; 46: 46-51

    View details for DOI 10.1016/j.plrev.2023.05.006

    View details for PubMedID 37244152

  • Oxytocin and the social facilitation of placebo effects. Molecular psychiatry Itskovich, E., Bowling, D. L., Garner, J. P., Parker, K. J. 2022


    Significant clinical improvement is often observed in patients who receive placebo treatment in randomized double-blind placebo-controlled trials. While a proportion of this "improvement" reflects experimental design limitations (e.g., reliance on subjective outcomes, unbalanced groups, reporting biases), some of it reflects genuine improvement corroborated by physiological change. Converging evidence across diverse medical conditions suggests that clinically-relevant benefits from placebo treatment are associated with the activation of brain reward circuits. In parallel, evidence has accumulated showing that such benefits are facilitated by clinicians that demonstrate warmth and proficiency during interactions with patients. Here, we integrate research on these neural and social aspects of placebo effects with evidence linking oxytocin and social reward to advance a neurobiological account for the social facilitation of placebo effects. This account frames oxytocin as a key mediator of treatment success across a wide-spectrum of interventions that increase social connectedness, therebyproviding a biological basis for assessing this fundamental non-specific element of medical care.

    View details for DOI 10.1038/s41380-022-01515-9

    View details for PubMedID 35338314

  • Endogenous oxytocin, cortisol, and testosterone in response to group singing. Hormones and behavior Bowling, D. L., Gahr, J., Ancochea, P. G., Hoeschele, M., Canoine, V., Fusani, L., Fitch, W. T. 1800; 139: 105105


    Humans have sung together for thousands of years. Today, regular participation in group singing is associated with benefits across psychological and biological dimensions of human health. Here we examine the hypothesis that a portion of these benefits stem from changes in endocrine activity associated with affiliation and social bonding. Working with a young adult choir (n=71), we measured changes salivary concentrations of oxytocin, cortisol, and testosterone from before and after four experimental conditions crossing two factors: vocal production mode (singing vs. speaking) and social context (together vs. alone). Salivary oxytocin and cortisol decreased from before to after the experimental manipulations. For oxytocin the magnitude of this decrease was significantly smaller after singing compared to speaking, resulting in concentrations that were significantly elevated after singing together compared to speaking together, after controlling for baseline differences. In contrast, the magnitude of the salivary cortisol decreases was the same across experimental manipulations, and although large, could not be separated from diurnal cycling. No significant effects were found in a low-powered exploratory evaluation of testosterone (tested only in males). At a psychological level, we found that singing stimulates greater positive shifts in self-perceived affect compared to speaking-particularly when performed together-and that singing together enhances feelings of social connection more than speaking together. Finally, measurements of heart rate made for a subset of participants provide preliminary evidence regarding physical exertion levels across conditions. These results are discussed in the context of a growing multidisciplinary literature on the endocrinological correlates of musical behavior. We conclude that singing together can have biological and psychological effects associated with affiliation and social bonding, and that these effects extend beyond comparable but non-musical group activities. However, we also note that these effects appear heavily influenced by broader contextual factors that shape social dynamics, such as stress levels, the intimacy of interactions, and the status of existing relationships.

    View details for DOI 10.1016/j.yhbeh.2021.105105

    View details for PubMedID 34999566

  • Progress without exclusion in the search for an evolutionary basis of music. The Behavioral and brain sciences Bowling, D. L., Hoeschele, M., Dunn, J. C. 2021; 44: e97


    Mehr et al.'s hypothesis that the origins of music lie in credible signaling emerges here as a strong contender to explain early adaptive functions of music. Its integration with evolutionary biology and its specificity mark important contributions. However, much of the paper is dedicated to the exclusion of popular alternative hypotheses, which we argue is unjustified and premature.

    View details for DOI 10.1017/S0140525X20001466

    View details for PubMedID 34588062

  • Harmonicity and Roughness in the Biology of Tonal Aesthetics. Music perception Bowling, D. L. 2021; 38 (3): 331-334


    Evidence supporting a link between harmoni-city and the attractiveness of simultaneous tone combinations has emerged from an experiment designed to mitigate effects of musical enculturation. I examine the analysis undertaken to produce this evidence and clarify its relation to an account of tonal aesthetics based on the biology of auditory-vocal communication.

    View details for DOI 10.1525/mp.2021.38.3.331

    View details for PubMedID 34566250

    View details for PubMedCentralID PMC8460127

  • Selection on vocal output affects laryngeal morphology in rats. Journal of anatomy Lesch, R., Schwaha, T., Orozco, A., Shilling, M., Brunelli, S., Hofer, M., Bowling, D. L., Zimmerberg, B., Fitch, W. T. 2021


    Although laryngeal morphology often reflects adaptations for vocalization, the structural consequences of selection for particular aspects of vocal behavior remain poorly understood. In this study, we investigated the effects of increased ultrasonic calling in pups on the adult larynx morphology in selectively bred rat lines. Laryngeal morphology was assessed using multiple techniques: mineralized cartilage volumes were compared in 3D-models derived from microCT scans, internal structure was compared using clearing and staining procedures combined with microscopy, cellular structure was compared using histology and microscopy, and element composition was assessed with scanning energy dispersive X-ray spectroscopy. Our results show that adult rats from lines bred to produce ultrasonic calls at higher rates as pups have shorter vocal folds and a more mineralized thyroid cartilage compared to rats bred to produce ultrasonic calls at lower rates. The change in vocal fold length appears to account for differences in low-frequency calls in these two rat lines. We suggest that the observed increases in mineralization of the thyroid cartilage in the high-ultrasound lineage provide increased reinforcement of the laryngeal structure during ultrasonic call production. Our findings therefore demonstrate an effect of selection for vocal behavior on laryngeal morphology, with acoustic consequences.

    View details for DOI 10.1111/joa.13366

    View details for PubMedID 33480050

  • Is consonance attractive to budgerigars? No evidence from a place preference study. Animal cognition Wagner, B., Bowling, D. L., Hoeschele, M. 2020


    Consonant tone combinations occur naturally in the overtone series of harmonic sounds. These include sounds that many non-human animals produce to communicate. As such, non-human animals may be attracted to consonant intervals, interpreting them, e.g., as a feature of important social stimuli. There is preliminary evidence of attraction to consonance in various bird species in the wild, but few experimental studies with birds. We tested budgerigars (Melopsittacus undulatus) for attraction to consonant over dissonant intervals in two experiments. In Experiment 1, we tested humans and budgerigars using a place preference paradigm in which individuals could explore an environment with multiple sound sources. Both species were tested with consonant and dissonant versions of a previously studied piano melody, and we recorded time spent with each stimulus as a measure of attraction. Human females spent more time with consonant than dissonant stimuli in this experiment, but human males spent equal time with both consonant and dissonant stimuli. Neither male nor female budgerigars spent more time with either stimulus type. In Experiment 2, we tested budgerigars with more ecologically relevant stimuli comprised of sampled budgerigar vocalizations arranged into consonant or dissonant chords. These stimuli, however, also failed to produce any evidence of preference in budgerigar responses. We discuss these results in the context of ongoing research on the study of consonance as a potential general feature of auditory perception in animals with harmonic vocalizations, with respect to similarities and differences between human and budgerigar vocal behaviour, and future methodological directions.

    View details for DOI 10.1007/s10071-020-01404-0

    View details for PubMedID 32572655

  • Claims of categorical primacy for musical affect are confounded by using language as a measure. Proceedings of the National Academy of Sciences of the United States of America Bowling, D. L. 2020

    View details for DOI 10.1073/pnas.2001689117

    View details for PubMedID 32291344

  • Rapid evolution of the primate larynx? PLoS biology Bowling, D. L., Dunn, J. C., Smaers, J. B., Garcia, M. n., Sato, A. n., Hantke, G. n., Handschuh, S. n., Dengg, S. n., Kerney, M. n., Kitchener, A. C., Gumpenberger, M. n., Fitch, W. T. 2020; 18 (8): e3000764


    Tissue vibrations in the larynx produce most sounds that comprise vocal communication in mammals. Larynx morphology is thus predicted to be a key target for selection, particularly in species with highly developed vocal communication systems. Here, we present a novel database of digitally modeled scanned larynges from 55 different mammalian species, representing a wide range of body sizes in the primate and carnivoran orders. Using phylogenetic comparative methods, we demonstrate that the primate larynx has evolved more rapidly than the carnivoran larynx, resulting in a pattern of larger size and increased deviation from expected allometry with body size. These results imply fundamental differences between primates and carnivorans in the balance of selective forces that constrain larynx size and highlight an evolutionary flexibility in primates that may help explain why we have developed complex and diverse uses of the vocal organ for communication.

    View details for DOI 10.1371/journal.pbio.3000764

    View details for PubMedID 32780733

  • Pupillometry of Groove: Evidence for Noradrenergic Arousal in the Link Between Music and Movement FRONTIERS IN NEUROSCIENCE Bowling, D. L., Ancochea, P., Hove, M. J., Fitch, W. 2019; 12
  • Temporal modulation in speech, music, and animal vocal communication: evidence of conserved function. Annals of the New York Academy of Sciences Filippi, P. n., Hoeschele, M. n., Spierings, M. n., Bowling, D. L. 2019


    Speech is a distinctive feature of our species. It is the default channel for language and constitutes our primary mode of social communication. Determining the evolutionary origins of speech is a challenging prospect, in large part because it appears to be unique in the animal kingdom. However, direct comparisons between speech and other forms of acoustic communication, both in humans (music) and animals (vocalization), suggest that important components of speech are shared across domains and species. In this review, we focus on a single aspect of speech-temporal patterning-examining similarities and differences across speech, music, and animal vocalization. Additional structure is provided by focusing on three specific functions of temporal patterning across domains: (1) emotional expression, (2) social interaction, and (3) unit identification. We hypothesize an evolutionary trajectory wherein the ability to identify units within a continuous stream of vocal sounds derives from social vocal interaction, which, in turn, derives from vocal emotional communication. This hypothesis implies that unit identification has parallels in music and precursors in animal vocal communication. Accordingly, we demonstrate the potential of comparisons between fundamental domains of biological acoustic communication to provide insight into the evolution of language.

    View details for DOI 10.1111/nyas.14228

    View details for PubMedID 31482571

  • Comparing Chalk With Cheese-The EGG Contact Quotient Is Only a Limited Surrogate of the Closed Quotient JOURNAL OF VOICE Herbst, C. T., Schutte, H. K., Bowling, D. L., Svec, J. G. 2017; 31 (4): 401–9


    The electroglottographic (EGG) contact quotient (CQegg), an estimate of the relative duration of vocal fold contact per vibratory cycle, is the most commonly used quantitative analysis parameter in EGG. The purpose of this study is to quantify the CQegg's relation to the closed quotient, a measure more directly related to glottal width changes during vocal fold vibration and the respective sound generation events. Thirteen singers (six females) phonated in four extreme phonation types while independently varying the degree of breathiness and vocal register. EGG recordings were complemented by simultaneous videokymographic (VKG) endoscopy, which allows for calculation of the VKG closed quotient (CQvkg). The CQegg was computed with five different algorithms, all used in previous research. All CQegg algorithms produced CQegg values that clearly differed from the respective CQvkg, with standard deviations around 20% of cycle duration. The difference between CQvkg and CQegg was generally greater for phonations with lower CQvkg. The largest differences were found for low-quality EGG signals with a signal-to-noise ratio below 10 dB, typically stemming from phonations with incomplete glottal closure. Disregarding those low-quality signals, we found the best match between CQegg and CQvkg for a CQegg algorithm operating on the first derivative of the EGG signal. These results show that the terms "closed quotient" and "contact quotient" should not be used interchangeably. They relate to different physiological phenomena. Phonations with incomplete glottal closure having an EGG signal-to-noise ratio below 10 dB are not suited for CQegg analysis.

    View details for DOI 10.1016/j.jvoice.2016.11.007

    View details for Web of Science ID 000406147000002

    View details for PubMedID 28017461