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Review ARTICLE

Front. Hum. Neurosci., 26 August 2020 | https://doi.org/10.3389/fnhum.2020.00350

Links Between the Neurobiology of Oxytocin and Human Musicality

  • School of Human Sciences, The University of Western Australia, Perron Institute for Neurological and Translational Science, Perth, WA, Australia

The human species possesses two complementary, yet distinct, universal communication systems—language and music. Functional imaging studies have revealed that some core elements of these two systems are processed in closely related brain regions, but there are also clear differences in brain circuitry that likely underlie differences in functionality. Music affects many aspects of human behavior, especially in encouraging prosocial interactions and promoting trust and cooperation within groups of culturally compatible but not necessarily genetically related individuals. Music, presumably via its impact on the limbic system, is also rewarding and motivating, and music can facilitate aspects of learning and memory. In this review these special characteristics of music are considered in light of recent research on the neuroscience of the peptide oxytocin, a hormone that has both peripheral and central actions, that plays a role in many complex human behaviors, and whose expression has recently been reported to be affected by music-related activities. I will first briefly discuss what is currently known about the peptide’s physiological actions on neurons and its interactions with other neuromodulator systems, then summarize recent advances in our knowledge of the distribution of oxytocin and its receptor (OXTR) in the human brain. Next, the complex links between oxytocin and various social behaviors in humans are considered. First, how endogenous oxytocin levels relate to individual personality traits, and then how exogenous, intranasal application of oxytocin affects behaviors such as trust, empathy, reciprocity, group conformity, anxiety, and overall social decision making under different environmental conditions. It is argued that many of these characteristics of oxytocin biology closely mirror the diverse effects that music has on human cognition and emotion, providing a link to the important role music has played throughout human evolutionary history and helping to explain why music remains a special prosocial human asset. Finally, it is suggested that there is a potential synergy in combining oxytocin- and music-based strategies to improve general health and aid in the treatment of various neurological dysfunctions.

Introduction

The human species has evolved two universal systems of inter-personal communication, language, and music. These communication streams possess some common elements, for example, a requirement for processing certain aspects of pitch, rhythm, and syntax; however, there are also well-established differences in neural circuitry that are linked to differences in functionality. The possible evolutionary origin of musical behaviors in our species has been discussed elsewhere (e.g., Brown, 2000; Mithen, 2005; Fitch, 2006; Patel, 2008; Morley, 2013; Richter and Ostovar, 2016; Harvey, 2017) and is not considered in detail here. Language plays an essential role in cognition; it is the primary means by which modern humans communicate thoughts and ideas, it facilitates the sharing of learned information and knowledge within and between generations, it permits intuitive reasoning, foresight, and planning, and it likely co-evolved with our capacity to imagine times and places not personally experienced in our lifetime (Harvey, 2017). Language and the emergence and continued development of human culture seem to be closely intertwined, but then why do we also communicate and enjoy music and its partner dance? Why does music continue as a human universal and what is its significance to the species?

Music affects many aspects of human behavior, behaviors that may have had (and still have) adaptive benefits that presumably contribute to the ongoing existence of musicality in humans (e.g., Cross, 2009; Harvey, 2018). These benefits, which are by no means mutually exclusive, are thought to include the attraction and selection of mates, the facilitation of attachment between caregivers and preverbal infants, aiding the development of perceptual, cognitive and motor skills, and encouraging trust, social bonding, and mutual cooperation. In a group context music-related activities, including dance (Laland et al., 2016; Richter and Ostovar, 2016), encourage the formation of bigger social networks, help to define cultural identity, and may represent a “safe haven” in which individuals can interact and share experiences without revealing their innermost thoughts and fears. Evidence supporting the important role that music plays in promoting the development and maintenance of cooperative, prosocial behaviors comes from an increasing number of studies in children and in adults (Freeman, 2000; Kirschner and Tomasello, 2009; Tarr et al., 2014; Pearce et al., 2015; Schellenberg et al., 2015). Music, via its impact on various regions within the limbic system, is also rewarding, motivating, and facilitates aspects of learning and memory (Zatorre and Salimpoor, 2013; Koelsch, 2018). Lastly, and no less important, it is increasingly appreciated that musical activities are useful therapeutic tools, aiding in the treatment of some developmental disorders (Quintin, 2019), and capable of ameliorating behavioral and psychological symptoms in several neurodegenerative conditions (e.g., Abraha et al., 2017; Zhang et al., 2017; Särkämö and Sihvonen, 2018; Groussard et al., 2019; Pereira et al., 2019).

In this review article, these special characteristics of music are considered in light of recent research on the neurobiology of the peptide oxytocin. Oxytocin is a hormone, synthesized in the hypothalamus that has both peripheral and central actions. Peripherally, oxytocin has important roles before and after childbirth, acting on the uterus during labor and stimulating lactation. Centrally, oxytocinergic systems are thought to influence many complex human social behaviors including, for example, pair bonding, attachment and social memory, emotional empathy, trust and generosity, and suppression of anxiety. The first part of the review focuses on what is currently known about the physiological actions of oxytocin on cells in the mammalian central nervous system (CNS) and the peptide’s interactions with other neuromodulator systems including the closely related pituitary hormone arginine vasopressin (AVP), the stress-related hormone cortisol, and neurotransmitters such as dopamine and serotonin. The second section summarizes recent advances in our knowledge of the distribution of the peptide and its receptor in the human brain, the relationship between endogenous oxytocin levels and complex behavioral traits typical of Homo sapiens, and then reviews the diverse effects of intranasal oxytocin administration on human behavior. The final section discusses links between music-related activities and oxytocin expression, documenting the similarities between the generally prosocial behaviors engendered by oxytocin and the many positive effects that music has on human cognition, memory, and mental health. Oxytocin and music can also have beneficial effects on cardiovascular and immune systems, and it is argued that a better understanding of the multiple actions of the oxytocinergic system may lead to its synergistic use with music in a range of therapeutic applications in psychology and neurology.

The Neuroscience of Oxytocin—Animal Studies

Oxytocin is a nine amino-acid peptide that is enzymatically derived from a larger peptide precursor made from the oxytocin gene. This peptide, or closely related versions of it, is involved in reproductive functions across almost all vertebrate species (Carter, 2014; Ebitz and Platt, 2014; Grinevich et al., 2016; Feldman, 2017; Jurek and Neumann, 2018) and its peripheral and central actions have been the subject of increasing interest in recent years (Jurek and Neumann, 2018)—as of 1st April 2020 there were more than 27,000 articles, including 3,700 reviews, listed on the NIH PubMed search engine.

In the mammalian brain, oxytocin is synthesized predominantly by magnocellular neurons in the supraoptic (SON) and paraventricular (PVN) nuclei of the hypothalamus, by some parvocellular neurons in PVN, and in accessory magnocellular nuclei of the hypothalamus. There is also some expression in peripheral tissues such as the gonads, kidney, and pancreas although the oxytocin generated there is unlikely to enter the CNS (Jirikowski, 2019). Many oxytocin-expressing neurons likely bifurcate, with a “traditional” endocrine-related projection to the posterior pituitary for systemic release into the bloodstream and a second central branch projecting to about 50 brain regions, including the sensory and prefrontal cortex, nucleus accumbens in the ventral striatum, amygdala, hippocampus, hypothalamus and ventral tegmentum (Grinevich and Stoop, 2018). In the circulation, oxytocin has a half-life of only a few minutes before it is metabolized in the liver and kidneys. This is an important point that will be returned to later in this review when discussing how best to measure and interpret peripheral oxytocin levels in humans.

In animal models, a variety of methods have been used to investigate the molecular and cellular properties of oxytocin and its receptor (OXTR), and to better understand the nature of the relationship between the physiology and pharmacology of oxytocin signaling and overt behavior (Jurek and Neumann, 2018; Mitre et al., 2018; Cilz et al., 2019; Neumann and Landgraf, 2019; Tan et al., 2019; Raam, 2020). These methods include neuroanatomical pathway tracing, immunohistochemistry, electron microscopy. receptor autoradiography, in situ hybridization, electrophysiology, microdialysis, and functional magnetic resonance imaging (fMRI). Experimental interventions have also been used to perturb the oxytocinergic system such as intracerebral infusion of the peptide, the use of receptor agonists or antagonists, antisense methods, optogenetic and chemogenetic stimulation of oxytocinergic neurons, conditional deletion of OXTR, and the use of genetically engineered reporter mice.

The Oxytocin Receptor

Oxytocin binds with high affinity to its specific receptor OXTR and can initiate an array of intracellular signaling cascades and transcriptional events (Chatterjee et al., 2016; Busnelli and Chini, 2017; Jurek and Neumann, 2018). There is also significant crosstalk with structurally related AVP receptors, most particularly the AVP1a receptor (AVPR1a; Bakos et al., 2018; Grinevich and Stoop, 2018; Song and Albers, 2018). In turn, AVP can also bind to OXTR, however, the specificity of action largely remains, probably due to differences in the distribution of oxytocin vs. AVP containing axons (Grinevich and Stoop, 2018; Rogers et al., 2018; Song and Albers, 2018; Pekarek et al., 2020). Nonetheless, there are potential sites of interaction in some brain regions (Smith et al., 2019), and there is evidence of functionally relevant spillover of oxytocin into the extracellular space beyond traditional synaptic sites (Busnelli and Chini, 2017; Chini et al., 2017; Song and Albers, 2018). OXTRs are widely distributed and found in many neuronal types, expressed on cell bodies, dendrites, and axon terminals, and the receptor is also expressed by astrocytes (Wang et al., 2017; Bakos et al., 2018; Young and Song, 2020). In different species, the receptor seems to be specifically enriched in those sensory/perceptual systems that are most relevant to conspecific maternal as well as more general socially interactive behaviors (Grinevich and Stoop, 2018; Pekarek et al., 2020).

The Physiology of Oxytocin

In the CNS, oxytocin can affect various ion channels, increase intracellular calcium ion concentrations, alter membrane excitability and enhance long-term potentiation (LTP) in neurons (Tomizawa et al., 2003; Lee et al., 2015; Lin and Hsu, 2018; Tirko et al., 2018). Oxytocin signaling also increases the expression of neurotrophic factors such as brain-derived neurotrophic factor (BDNF; Bakos et al., 2018; Zhang et al., 2020), of relevance to later discussion focussed on oxytocin, social learning/memory, and hippocampal function. The peptide can also act presynaptically to affect neurotransmitter secretion (Dölen et al., 2013; Bakos et al., 2018). Overall, from a physiological perspective, oxytocin influences cell viability, synaptic and structural plasticity in neurons (Bakos et al., 2018; Jurek and Neumann, 2018; Pekarek et al., 2020), and modulates the balance of excitatory and inhibitory activity in regions such as the cerebral cortex and hippocampus (e.g., Mitre et al., 2016; Grinevich and Stoop, 2018; Lin and Hsu, 2018; Lopatina et al., 2018; Tirko et al., 2018; Cilz et al., 2019; Maniezzi et al., 2019; Tan et al., 2019), amygdala (Crane et al., 2020), and nucleus accumbens (Moaddab et al., 2015; Cox et al., 2017). Rodents lacking oxytocin or OXTR display impaired sociability and social memory (Ferguson et al., 2000). and conditional deletion of OXTR in the hippocampus negatively affects LTP and impairs long-term social recognition memory (Lin et al., 2018).

Likely increasing its diversity of action, oxytocin also interacts with several other receptors and neuromodulatory systems. For example, the peptide: (i) potentiates excitatory dopamine-mediated synaptic transmission (Li et al., 2020); (ii) interacts with a class of serotonin receptor (Chruścicka et al., 2019) and affects serotonin release (Yoshida et al., 2009); (iii) modulates signaling mediated by opioid receptors (dal Monte et al., 2017; Meguro et al., 2018; Salighedar et al., 2019); and (iv) activates TRPV2 channels (Van den Burg et al., 2015). There are also dynamic interactions with steroids (Jirikowski et al., 2018) and oxytocin levels are negatively correlated with cortisol, significantly modifying responses to stress (Lee et al., 2015; Schladt et al., 2017; Latt et al., 2018; Masis-Calvo et al., 2018; Neumann and Landgraf, 2019).

The foregoing section has, of necessity, over-simplified the physiological effects of oxytocin on neural tissue in animals, and more in-depth reviews are available (e.g., Bakos et al., 2018; Jurek and Neumann, 2018; Mitre et al., 2018; Neumann and Landgraf, 2019). However, some discussion of animal-based research is warranted because, given that the peptide is highly conserved in evolution it is likely that similar wide-ranging molecular and cellular mechanisms are operative in the human brain (see also Grinevich and Neumann, 2020). From a behavioral perspective, animal studies reveal that oxytocin has an important role in pair-bonding and maternal attachment, in moderating affiliative behaviors and conspecific social recognition, and in modulating the formation and maintenance of episodic memories, whether they be positive or negative. The next section will show that oxytocin has generally similar effects on human social behavior, but these effects would seem to be more subtle and complex in cognitively advanced members of Homo sapiens, extending to personality traits, emotional empathy, trust, altruism, reciprocity, group conformity, social decision making and so on.

Oxytocin in Humans

Oxytocinergic Networks

In humans, immunoreactive oxytocinergic fibers are sparse but present in all cortical layers of the orbitofrontal cortex and anterior cingulate (Rogers et al., 2018). The fibers were found to have large varicosities usually associated with en passant boutons—likely sites of oxytocin release into the surrounding neuropil (Busnelli and Chini, 2017; Chini et al., 2017; Song and Albers, 2018). Fibers immunoreactive for AVP were also seen in these cortical regions and in the insular and olfactory cortices. Using antibodies to the receptor, OXTR was first identified in parts of the amygdala, anterior cingulate cortex, hypothalamus, and preoptic area, olfactory nucleus, and some brainstem nuclei (Boccia et al., 2013). A more recent extensive survey of the oxytocin system analyzed the distribution of the gene encoding OXTR as well as the gene encoding the oxytocin prepropeptide and the gene encoding CD38, a transmembrane protein needed for oxytocin secretion (Quintana et al., 2019). OXTR gene expression was widespread throughout the brain, significantly higher in olfactory bulbs, but also higher in the caudate, putamen, pallidum, and hypothalamus; levels were also greater than average in the hippocampus, parahippocampal region, amygdala, parts of the temporal lobe and anterior cingulate cortex. Expression of the gene was “reproducible, regardless of individual differences, such as ethnicity and sex” (Quintana et al., 2019).

The pattern of expression was essentially similar for the CD38 gene, with significantly increased expression in caudate, putamen, pallidum, thalamus, and anterior cingulum. For both genes, expression was significantly lower in the cerebellum. Interestingly, there was co-expression with several genes involved in dopaminergic and muscarinic cholinergic signaling, suggesting potential pathway interactions perhaps similar to those suggested for the opioids (dal Monte et al., 2017). Co-expression with genes involved in the regulation of metabolism and appetite was also seen. According to Quintana et al. (2019), “the oxytocin pathway gene maps correspond with the processing of anticipatory, appetitive, and aversive cognitive states.” Interaction with dopaminergic and cholinergic systems is likely to add to the broad impact of oxytocin on social behaviors, motivation, reward, desire, anxiety, and the processing of emotions.

Receptor Polymorphisms and Behavior

In children, adolescents, and adults, genetic variants of the OXTR gene are linked to an individual’s response to stress (Rodrigues et al., 2009) and altered prosocial/affiliative behaviors and empathy. The need for pleasant social company is increased after a stressful event, a need that varies depending on which alleles of OXTR are present (Sicorello et al., 2020). Anatomically, there are subtle changes in structure and inter-connectivity of hypothalamus and parts of the limbic system, and mutations have been implicated in a range of highly maladaptive, sometimes psychopathic traits (e.g., Israel et al., 2008; Tost et al., 2010; Dadds et al., 2014; Aspé-Sánchez et al., 2016; Feldman et al., 2016; Gedeon et al., 2019; Poore and Waldman, 2020). A recent neuroimaging study examining the effect of OXTR alleles on resting-state networks reported that receptor genotype affected connectivity between the right hippocampus, medial prefrontal cortex, dorsal anterior cingulate cortex, amygdala, basal ganglia and thalamus (Luo et al., 2020). The functional impact that alleles of OXTR have on social behavior is however complex and not always consistent across studies, and is affected by factors such as gender, age, upbringing, and culture (Tost et al., 2010; Feldman et al., 2016; Fujiwara et al., 2019; Plasencia et al., 2019; Poore and Waldman, 2020). Environmental epigenetic influences on the OXTR function that influence social interactions must also be considered (Chen et al., 2020), and as described earlier there may be differential interactions with other neuromodulatory systems such as AVP, the opioids, steroids, and various catecholamines.

Measurement of Endogenous Oxytocin

As yet it has not proved possible to measure oxytocin levels in the living human brain, thus endogenous oxytocin measurements are obtained from either plasma, saliva, or urine. Interpretation of these peripheral measures of oxytocin is however difficult for several reasons (Ebstein et al., 2012; Leng and Ludwig, 2016; Mitre et al., 2016; Valstad et al., 2017; Jurek and Neumann, 2018). First, peripheral oxytocin levels are related to the release of the peptide from the posterior pituitary and do not necessarily reflect levels of the peptide within specific regions of the brain that contain neurons expressing OXTR. Second, even when undertaking peripheral measurements, compared to saliva there is, in animals at least, a more consistent relationship between blood plasma levels of oxytocin and levels of the peptide found in cerebrospinal fluid (Valstad et al., 2017). Third, as pointed out by Jurek and Neumann (2018): “basal plasma or brain oxytocin levels might strongly depend on individual events occurring within the last hour(s) before sampling (e.g., fear of hospital or laboratory, prior eating, rushing to the laboratory, or sex) or on the time of the day.” And all is compounded by the fact that circulating levels of oxytocin are normally low, even exogenous peptide is rapidly eliminated 1–2 h after intranasal delivery, and to measure native (unbound) oxytocin levels requires sophisticated techniques for specificity and accuracy of analysis (Franke et al., 2019, 2020). Nonetheless, and given these caveats, some intriguing and important observations have come from peripheral endogenous oxytocin measurements in humans.

Nurturing and Bonding

Plasma and/or salivary oxytocin levels rise postpartum when mothers interact and bond with their infants (Matthiesen et al., 2001; Feldman et al., 2007; Feldman, 2012; Gordon et al., 2010). These interactive behaviors include gaze, facial expression, vocalizing using the preverbal maternal-infant communication known as “motherese,” affectionate touch, and so on (Kerr et al., 2019). The increase in serum oxytocin when mothers interact and bond with their own smiling/happy infants is higher in mothers rated as having “secure” attachment to their offspring (Strathearn et al., 2009) and with a sensitive temperament (Strathearn et al., 2012). In these mothers, fMRI revealed greater activity in the hypothalamus/pituitary region and in reward centers in the ventral striatum. The rise in oxytocin is, at least in part, related to the amount of maternal gaze directed towards the child (Kim et al., 2014). Intranasal oxytocin also increases a father’s neural response to images of their young children, with increased activity in caudate, anterior cingulate, and visual cortex (Li et al., 2017a).

Lullabies are a universal way of soothing infants (Mehr et al., 2018), and it is thus of interest that vocalization by mothers increases levels of salivary oxytocin (and reduces cortisol) in their children, although admittedly these were older girls aged between 7 and 12 years old (Seltzer et al., 2010). In comparing maternal vs. paternal changes in endogenous oxytocin during early parent-infant bonding, both mothers and fathers showed increases but there were dimorphic differences that depended on the type of interaction (Gordon et al., 2010). The development of affiliative behaviors between caregiver and infant, linked especially to oxytocin, leads to plasticity and adaptations in both the parent and infant (Feldman, 2015). Remarkably, the basal level of oxytocin measured in the saliva, and certain polymorphisms in the OXTR gene, are transgenerationally associated with the type of parental care that is given, influencing affiliative and social behaviors across as many as three generations within a family (Fujiwara et al., 2019).

Oxytocin in Adolescents and Adults

Endogenous plasma oxytocin concentrations vary with age and there are differences between males and females, young women having the highest and old men the lowest levels (Plasencia et al., 2019). Experimentally, levels of salivary oxytocin were higher when female subjects were confronted with a novel situation, associated with reduced stress and greater trust, compared with a later familiarization session (Tops et al., 2013). Higher levels of plasma oxytocin in women but not men were linked to indicators of relationship stress and attachment anxiety (Taylor et al., 2010; Weisman et al., 2013; Moons et al., 2014). In young males, lower urinary oxytocin (but not AVP) levels were linked to lower measures of empathy and trust, presumably associated with a greater propensity for aggressive behavior (Malik et al., 2012; Weisman et al., 2013; de Jong and Neumann, 2018; Berends et al., 2019), and in males social cognitive ability was correlated with plasma oxytocin concentrations (Deuse et al., 2019; Strauss et al., 2019).

Effects of Exogenous Administration of Oxytocin

Exogenous delivery of oxytocin affects neural processing and has consistently been reported to influence a wide range of interactive human behaviors. These behaviors have been described in different ways and using different terminologies. They include pair bonding, attachment, and social learning/memory, social salience and emotional empathy, recognition and interpretation of emotions, behavioral synchrony, familiarization and within group co-operation, altruism, generosity and trust, reward sensitivity, calmness and reduction of stress, and amelioration of anxiety (anxiolytic effects; e.g., Kosfeld et al., 2005; Baumgartner et al., 2008; Ditzen et al., 2009; Strathearn et al., 2009; Hurlemann et al., 2010; De Dreu, 2012; Fischer-Shofty et al., 2012; Tops et al., 2013; Bethlehem et al., 2014; Preckel et al., 2014; Shamay-Tsoory and Abu-Akel, 2016; Feldman, 2017; Fineberg and Ross, 2017; Leppanen et al., 2017; Wang et al., 2017; Ellenbogen, 2018; Geng et al., 2018; Jurek and Neumann, 2018; Rilling et al., 2018; Alos-Ferrer and Farolfi, 2019; Liu et al., 2019; Tillman et al., 2019; Sicorello et al., 2020; Wu et al., 2020). Analyses of the impact of oxytocin on neural activity imaged in the brains of healthy subjects generally reflect this, with altered activity in interconnected structures associated with valence, salience, trust, prosocial behavior and mentalizing, including the amygdala, insula, nucleus accumbens, lateral septum, anterior cingulate, hippocampus, caudate, tempero-parietal cortex, dorsomedial and dorsolateral prefrontal cortex (e.g., Kirsch et al., 2005; Rilling and Sanfey, 2011; Bethlehem et al., 2013; Eckstein et al., 2017; Wang et al., 2017; Rilling et al., 2018; Kumar et al., 2020; Wu et al., 2020).

The great majority of studies emphasize positive, prosocial behavioral outcomes after exogenous oxytocin administration; however, it is important to emphasize that not all studies describe these effects (Keech et al., 2018; Tabak et al., 2019; Erdozain and Peñagarikano, 2020) and some antisocial outcomes have been reported, including increased competitive and aggressive tendencies, particularly in males (Fischer-Shofty et al., 2012; Alcorn et al., 2015; Ne’eman et al., 2016; de Jong and Neumann, 2018; Gedeon et al., 2019). Others have also reported differential effects of exogenous oxytocin on women compared to men (e.g., Rilling et al., 2012, 2018; Preckel et al., 2014; Feng et al., 2015; Chen et al., 2016; Bredewold and Veenema, 2018; Bartz et al., 2019; Xu et al., 2020).

Overall, variation in the effects of administered oxytocin is linked to: (i) gender differences; (ii) whether individuals possess intrinsic pro- or antisocial personality traits—sometimes related to polymorphisms in, or the methylation state of, OXTR; (iii) early environmental experience; and/or (iv) the social cues and psychological context when testing is undertaken (e.g., Guastella and MacLeod, 2012; Evans et al., 2014; Nishina et al., 2015; Chen et al., 2016; Feldman et al., 2016; Lambert et al., 2017; Aydogan et al., 2018; de Jong and Neumann, 2018; Wagner and Echterhoff, 2018; Fragkaki and Cima, 2019; Gedeon et al., 2019; Liu et al., 2019; Sicorello et al., 2020). As mentioned earlier, oxytocin and AVP can activate each other’s receptors, although the differential distribution of fibers and receptors may limit crosstalk (Rogers et al., 2018; Song and Albers, 2018). There may be little interaction under normal conditions of endogenous release, but perhaps there is more crosstalk after intranasal application of higher concentrations of oxytocin which may contribute to some of the complexity of behavioral outcomes.

The Links Between Oxytocin and Music

Oxytocin is an ancient peptide, in mammals universally involved in reproductive biology, modulating social learning and affiliative behaviors, as well as modifying responses to adverse conditions (Ebitz and Platt, 2014; Feldman et al., 2016; de Jong and Neumann, 2018). In Homo sapiens, it is conceivable that the unique prosocial, harmonizing activities of music and dance incorporated, perhaps even required, elements of this pre-existing oxytocinergic network. Music encourages affiliative interactions in infancy and adulthood, aids in the development of perceptual, cognitive, and motor skills, promotes trust and reduces a sense of social vulnerability, is rewarding and motivating, and has a beneficial effect on aspects of learning and memory. Music and its evolutionary partner dance (Richter and Ostovar, 2016) also promote synchrony and social interaction, contribute to cultural identity, and encourage the formation of cooperative networks. Based on the experimental work described above, it should be apparent that many of these musical influences on human behavior are also characteristic of many of the psychological and sociological effects of oxytocin. These associations become even clearer when comparing the neural networks that are: (i) activated when listening to music perceived as being rewarding and pleasurable with; (ii) regions that process behaviors that involve social cooperation, empathy and altruism; and (iii) the distribution of oxytocinergic fibers and OXTR in the human brain (Figure 1).

FIGURE 1
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Figure 1. Two schematic views of the brain showing many of the regions in common that are reported to be involved in: (i) processing subjective/arousing/rewarding/emotional aspects of human musicality; (ii) performing or responding to a range of interactive social tasks; and (iii) oxytocinergic processing. Two other potentially relevant regions are shown in italics. Amygdala not shown. PFC, the prefrontal cortex. Both diagrams modified from Harvey (2017).

Music Networks

As described in detail elsewhere (Harvey, 2017), and acknowledging that overlap in activity maps does not, a priori, mean that the same circuits are involved (Peretz et al., 2015), some of the basic elements of language and music such as pitch and rhythm share similar neural substrates, but clearer differences become apparent when more extended processing networks are considered. In right-handers at least, there is a left hemisphere bias for language and speech while the right hemisphere is biased more for music, and separate processing areas specific for these two communication streams have now been identified within secondary auditory regions in the superior temporal gyrus (Angulo-Perkins et al., 2014; Norman-Haignere et al., 2015). Most relevant to the present discussion are the limbic pathways and multiple cortical regions known to be activated by the overall subjective experience and emotional impact of music (Koelsch, 2018).

The limbic system, which includes the hippocampus, parahippocampal gyrus, amygdala, and cingulate cortex, is involved in several functions including learning, memory, motivation and emotional responsiveness. Music can induce activity in all these regions, while music that is perceived as arousing and is appreciated also drives dopaminergic activity in nucleus accumbens in the ventral striatum, an anticipatory and reward center (Blood and Zatorre, 2001; Menon and Levitin, 2005; Boso et al., 2006; Salimpoor et al., 2011; Zatorre and Salimpoor, 2013; Mueller et al., 2015; Ferreri et al., 2019; Gold et al., 2019; Shany et al., 2019). Music that evokes strong emotional valence is associated with altered activity not only in the superior temporal gyrus but also in the caudate nucleus, insula, thalamus, cingulate cortex, orbitofrontal, dorsomedial, dorsolateral and ventromedial prefrontal cortex, inferior frontal cortex and supplementary motor area (e.g., Blood and Zatorre, 2001; Koelsch et al., 2006; Mitterschiffthaler et al., 2007; Chapin et al., 2010; Brattico et al., 2011; Pereira et al., 2011; Khant et al., 2012; Altenmüller et al., 2014; Koelsch, 2018; Särkämö and Sihvonen, 2018; Sachs et al., 2019). Of course, music involves more than just listening, and imaging of people—alone or with others—creating and improvising jazz, rap or rock music has also revealed increased neural activity in the medial frontal lobe and altered, usually decreased, activity in the dorsolateral prefrontal cortex when compared to the same subjects playing or singing “formulaic sequences” (Limb and Braun, 2008; Liu et al., 2012; Donnay et al., 2014; Tachibana et al., 2019).

Behavioral Networks

Numerous studies have used fMRI to image healthy subjects performing and responding to interactive social tasks that may involve altruism, empathy, trust and cooperation, norm-abiding behavior, mentalizing, and/or an appreciation of nuanced social context (Figure 1). These studies generally find increased functional activity in the nucleus accumbens, amygdala, parahippocampal gyrus, caudate nucleus, insula, anterior cingulate cortex, superior temporal cortex, tempero-parietal junction, and several regions in the prefrontal cortex (medial orbitofrontal, medial prefrontal, ventromedial, dorsolateral, dorsomedial; e.g., O’Doherty, 2004; Völlm et al., 2006; Rilling et al., 2008; Cooper et al., 2010; Rilling and Sanfey, 2011; Rushworth et al., 2011; Carter et al., 2012; Korn et al., 2012; Fukuda et al., 2019).

Oxytocinergic Systems

Oxytocin fibers have been identified in the orbitofrontal cortex and anterior cingulate cortex (Rogers et al., 2018), and there are high levels of the receptor in the olfactory bulb, amygdala, hippocampus, parahippocampal gyrus, regions in the temporal lobe, anterior cingulate cortex, hypothalamus and preoptic area, and some brainstem nuclei (Boccia et al., 2013; Quintana et al., 2019). Intranasal oxytocin administration alters neural activity in structures such as the amygdala, insula, nucleus accumbens, anterior cingulate cortex, hippocampus, caudate, tempero-parietal cortex, dorsomedial and dorsolateral prefrontal cortex (e.g., Kirsch et al., 2005; Lischke et al., 2012; Bethlehem et al., 2013; Eckstein et al., 2017; Wang et al., 2017; Rilling et al., 2018; Kumar et al., 2020; Wu et al., 2020). Finally, altered OXTR genotypes have been found to correlate with altered local network metrics and functional connectivity between the hippocampus, medial prefrontal cortex, dorsal anterior cingulate cortex, amygdala, basal ganglia and thalamus (Luo et al., 2020).

Clearly then, musicality, cooperative prosocial interactions, and the oxytocinergic system are linked to neural activity in several common regions and interconnected networks in the brain of modern humans; the most consistently involved components being the hippocampus, parahippocampal gyrus, amygdala and anterior cingulate cortex, the caudate nucleus and nucleus accumbens, insula, superior temporal gyrus and orbitofrontal, ventromedial, dorsomedial and dorsolateral prefrontal cortex (Figure 1). Numerous examples of the close interrelationship between oxytocin and prosocial human behaviors have been presented, promoting group empathy and the participation in collective decision making, all involving a shift from personal concerns to more communal interests, including a willingness to learn from others (Zak and Berroza, 2013; Shalvi and De Dreu, 2014; De Dreu and Kret, 2016; De Wilde et al., 2017; Ten Velden et al., 2017; Schiller et al., 2020; Xu et al., 2020). But to what extent does oxytocin provide a nexus between these behaviors and music? What impact does music have on peripheral oxytocin release and OXTR expressing networks in the human brain, and can exogenous oxytocin administration synergistically affect performance and responsiveness to music?

Experimental Studies on Music and Oxytocin

Only a few studies have directly examined the impact of music on oxytocin expression, in solo or ensemble settings. As described earlier, comforting maternal vocalizations—which can have music-like properties—by themselves have been shown to increase oxytocin levels and reduce cortisol in young daughters (Seltzer et al., 2010). It was recently reported that salivary oxytocin levels are reduced in maltreated children (Suzuki et al., 2020) and it is therefore of interest that, following a program of group drumming sessions for “emotionally disturbed” children, salivary oxytocin concentrations were increased in both boys and girls, significantly so when comparing practice and free play sessions performed by boys aged 8–12 years (Yuhi et al., 2017). In adults, salivary oxytocin levels were also found to be raised after a singing lesson, amateur singers, in particular, expressing a heightened sense of well-being (Grape et al., 2003), and raised levels were also reported after choral singing (Kreutz, 2014). In one sensory study, it was found that listening to slow relaxing music was associated with raised salivary oxytocin levels and lower heart rate, whereas fast music had little impact on oxytocin but reduced cortisol levels and increased arousal (Ooishi et al., 2017). The effect of relaxing music on moderating salivary cortisol levels after the stress has also been noted (Khalfa et al., 2003).

The nature of the musical activity is important because an increase in plasma oxytocin levels in members of a vocal jazz group was only recorded when singers were improvising together (Keeler et al., 2015), likely due to altered activity in the prefrontal cortex and enhanced affiliative and prosocial interactions (Limb and Braun, 2008; Liu et al., 2012; Donnay et al., 2014; Tachibana et al., 2019). Schladt et al. (2017) reported that salivary oxytocin levels slightly increased when subjects were solo singing but were decreased when singing in a choir. In that same study, cortisol levels were reduced in both situations, but choral participants described greater feelings of happiness and reduced worry. Of course, performing music can be stressful, perhaps especially in a solo compared to an ensemble/choral situation. Indeed, the intranasal application of oxytocin has recently been shown to increase positive interactions between performers and reduce performance anxiety (Sabino et al., 2020). The reported variability in measured levels of oxytocin and markers of stress such as cortisol reflects the complex, and highly interactive, sexually dimorphic systems that are involved (Brown et al., 2016). The other issue that should be borne in mind, discussed in detail earlier, is that measurement of salivary or urinary oxytocin levels do not necessarily reflect the concentration of the peptide in OXTR expressing regions in the brain (Leng and Ludwig, 2016; Jurek and Neumann, 2018), although there is a closer relationship between plasma oxytocin levels and those in cerebrospinal fluid (Valstad et al., 2017).

Overall, whilst there is a clear trend for increased endogenous oxytocin and reduced cortisol in subjects involved in musical activities, more controlled trials are needed in this area because communal music experiences are prime examples of human social engagement. From a physiological and psychosocial perspective, group music-making such as choral singing increases connectedness, heightens empathy, reduces depression and improves mood, is arousing and stimulates cognition, and has systemic health benefits including improved immune competency, reduced cytokine and inflammatory markers, lowered blood pressure and reduced cortisol and ACTH levels (Kuhn, 2002; Khalfa et al., 2003; Kreutz et al., 2004; Dunbar et al., 2012; Fancourt et al., 2014; Keeler et al., 2015; Pearce et al., 2015; Stewart and Lonsdale, 2016; Johnson et al., 2017; Ooishi et al., 2017; Finn and Fancourt, 2018; Kang et al., 2018; Moss et al., 2018; Perkins et al., 2018; Walker et al., 2019). The impact of exogenous oxytocin is relevant here because of the positive effect that it has on individual stress levels and the promotion of group empathy, reciprocal trust and collective social decision making, all involving a shift from personal to group agency (Zak and Berroza, 2013; Chen et al., 2016; De Dreu and Kret, 2016; De Wilde et al., 2017; Ten Velden et al., 2017; Sicorello et al., 2020; Xu et al., 2020). Music and the community associated with it may be especially important to individuals who are lonely and/or who have lower emotional empathy and exhibit fewer prosocial traits (e.g., Berends et al., 2019; Fragkaki and Cima, 2019; Liu et al., 2019; Johnson et al., 2020; Schiller et al., 2020).

Links to Other Neuromodulatory Systems

Rhythm in music induces bodily movement and akin to music, dance is a universal human behavior (Levitin et al., 2018). From an evolutionary perspective, it has been argued that dance advantages humans “by contributing to sexual reproduction signaling, cooperation, social bonding, infant care, violence avoidance as well as embodied individual and social communication and memorization” (Richter and Ostovar, 2016). To my knowledge, there have not, to date, been any substantive reports on how oxytocin levels are affected by solo or group dance. Yet the impact of dance, especially in a coordinated group context, on increased empathy (Gujing et al., 2019) social bonding (Tarr et al., 2015, 2016), cognitive performance, general fitness and well-being (Kattenstroth et al., 2010, 2013; Zilidou et al., 2018; Douka et al., 2019) is clear. Choral singing and dance have both been reported to increase pain threshold, viewed as a surrogate for levels of circulating β-endorphin (Dunbar et al., 2012; Tarr et al., 2015; Weinstein et al., 2016). This peptide binds to μ-opioid receptors and plays a role in social networking and maintaining social bonds (Pearce et al., 2017). These observations are important, but it should be noted that many other factors can influence the perception and processing of pain (Millan, 2002), including oxytocin (Gamal-Eltrabily et al., 2020; Hilfiger et al., 2020; Schneider et al., 2020), which as described earlier positively modulates signaling mediated by opioid receptors (dal Monte et al., 2017; Meguro et al., 2018; Salighedar et al., 2019). Furthermore, depending on age and health status, the perception of pain does not necessarily reflect circulating β-endorphin levels (Bruehl et al., 2017; Ahn et al., 2019).

AVP is also thought to influence human behavior in many ways (Neumann and Landgraf, 2012; Benarroch, 2013) although significant effects are not always evident (Tabak et al., 2019). AVP and oxytocin may interact with each other to influence prosocial vs. antisocial behaviors, trust vs. aggression, fear and so on (Huber et al., 2005; Veenema and Neumann, 2018; Ebstein et al., 2012; Rilling et al., 2012; Jurek and Neumann, 2018; Song and Albers, 2018; Berends et al., 2019), at least some of which are sexually dimorphic (Rilling et al., 2014; Feng et al., 2015; Bredewold and Veenema, 2018). Comparison of endogenous AVP and oxytocin levels in plasma from young and old men and women revealed a negative correlation between all groups, higher AVP levels associated with greater “attachment anxiety” (Plasencia et al., 2019) and pair-bond distress in men (Taylor et al., 2010). Polymorphisms in the AVPR1a receptor have been linked to variability in aggression, response to stress, trust, and altruistic behaviors (Israel et al., 2008; Moons et al., 2014; Aspé-Sánchez et al., 2016; Nishina et al., 2019). Whilst there is as yet no evidence that endogenous levels of circulating AVP are altered by musical activity, AVPR1a receptor polymorphisms have been linked to musical aptitude (Pulli et al., 2008; Ukkola et al., 2009; Liu et al., 2016; Mariath et al., 2017), memory (Granot et al., 2007, 2013) and appreciation (Ukkola-Vuoti et al., 2011), as well as music and dance creativity (Bachner-Melman et al., 2005; Israel et al., 2008; Oikkonen et al., 2016). On the other hand, receptor polymorphisms were not more common in choral singers compared with people designated as non-musicians (Morley et al., 2012).

Anxiety, Extinction, and PTSD

Participation in music is rewarding; it encourages prosocial interactions, facilitates social cognition, and promotes cooperation within groups of culturally compatible but not necessarily genetically related individuals. Ensemble music-making, and communal choral and dance activities, involve synchronized and coordinated activity with the special attribute of allowing individuals to be subsumed within a greater, living whole. Perhaps most importantly, the generally ambiguous, non-propositional nature of music provides a safe, usually risk-free space where individual thoughts and emotions, personal autobiographical memories and ambitions, can exist in a cooperative and interactive social context. Participation in musical activities can help individuals who lack self-confidence, who lack trust and may feel socially excluded, reduces fear and a sense of vulnerability, and can diminish potential conflict: “Music allows participants to explore the prospective consequences of their actions and attitudes toward others within a temporal framework that promotes the alignment of participants’ sense of goals” (Cross, 2009).

This putative “safe haven” aspect of human musicality is similar to some of the behavioral effects elicited by oxytocin and further supports the proposed close links between music and oxytocinergic systems. Although not evident in all trials (Donadon et al., 2018), many studies have reported that exogenous delivery of oxytocin has anxiolytic and calming effects on human behavior (Neumann and Slattery, 2016; Wang et al., 2017; Lancaster et al., 2018; Yoon and Kim, 2020), enhancing the detection of threat (Lischke et al., 2012; Bredewold and Veenema, 2018) and facilitating the extinction of fearful or distressing memories (Kirsch et al., 2005; Hu et al., 2019; Koch et al., 2019; Triana-Del Río et al., 2019). Indeed, endogenous oxytocin levels are reduced in individuals with emotional trauma and in sufferers of posttraumatic stress disorder (PTSD; e.g., Frijling et al., 2015) and the administration of oxytocin may prove to be a useful therapeutic strategy (Giovanna et al., 2020). The acquisition and processing of autobiographical experiences, including fear and extinction, involves the hippocampus and ventromedial prefrontal cortex (Bonnici and Maguire, 2018; Dunsmoor et al., 2019) as well as interactions with the amygdala (Dunsmoor et al., 2019; Hasan et al., 2019). Activity in all these regions is associated with aspects of both musical and oxytocinergic processing. Concerning extinction, reducing the emotional impact of remembering fearful and threatening events involves substitution with novel, less impactful memories during the retrieval and reconsolidation process, a process facilitated by oxytocin (Hu et al., 2019; Triana-Del Río et al., 2019) and one that may also be aided by participation in the safe, neutral and motivating mental space evoked by communal music-related activities. In this context, music therapy has been suggested as a possible treatment for PTSD (Beck et al., 2018), and its use in association with oxytocin administration may prove even more beneficial.

Of the many endogenous opioids, β-endorphin—which may be raised by social music-making—has been implicated in resilience, stress, and PTSD (Bali et al., 2015) as has the neuropeptide nociceptin (Tollefson et al., 2017; Narendran et al., 2019). Nociceptin receptor polymorphisms have been linked to the severity of PTSD, but unlike β-endorphin, no relationship to music has been examined. Finally. there is also evidence of an important role for dopamine in the pathophysiology of PTSD (Lee et al., 2017; Torrisi et al., 2019) with potential interaction with oxytocinergic systems (Zhang et al., 2019), further strengthening the suggestion about the potential usefulness of music therapy given the known impact that music has on dopaminergic motivation and reward systems in the human brain (Chanda and Levitin, 2013; Zatorre and Salimpoor, 2013; Ferreri et al., 2019).

Learning, Social Memory, and Hippocampal Plasticity

In children, some degree of music training has a significant impact on brain structure and plasticity as well as having a positive influence on social, empathic, cognitive and academic development (e.g., Schlaug et al., 2009; Kirschner and Tomasello, 2009; Schellenberg et al., 2015; Habibi et al., 2018; Sachs et al., 2018; de Manzano and Ullén, 2018; Guhn et al., 2020). Learning to play an instrument requires the recruitment of many sensorimotor systems and circuits, and many studies have reported that music training has beneficial effects on various executive functions and some types of memory, benefits that are maintained throughout a person’s lifetime and may be protective against cognitive decline (Talamini et al., 2017; Mansens et al., 2018). Once again there are several intriguing and potentially important links between music training, music-related activities, and the neuroscience of oxytocin, in this case, the links that are relevant to memory and aging, with dance and exercise adding an additional dimension to the discussion. Oxytocin’s effects on social recognition, learning, and memory are associated with activity in the hippocampus, amygdala, nucleus accumbens, and prefrontal cortex (e.g., Ferguson et al., 2002; Hurlemann et al., 2010; Mitre et al., 2016; Grinevich and Stoop, 2018; Jurek and Neumann, 2018; Lin and Hsu, 2018; Lin et al., 2018; Lopatina et al., 2018; Cilz et al., 2019; Tan et al., 2019; Raam, 2020; Xu et al., 2020). In the following discussion, the focus is primarily on the hippocampus, given its role in consolidating, integrating and retrieving personal autobiographical memories (Bonnici and Maguire, 2018; Sheldon et al., 2019). There is a huge literature on hippocampal connectivity and plasticity related to these dynamic and transformational processes—the emphasis here will be limited to several aspects of social learning and memory perhaps most relevant to a review of music and oxytocin.

Music activates diverse regions and circuits within the CNS including, depending on context and emotional valence, essentially the same limbic structures that are responsive to oxytocin (Boso et al., 2006; Koelsch, 2014, 2018). Music training and practice improves memory (Talamini et al., 2017; Mansens et al., 2018) and affects the architecture and organization of both gray and white matter in the brain (de Manzano and Ullén, 2018). Of particular relevance here is the positive effect that music training has on gray matter volume and plasticity in the hippocampus (Herdener et al., 2010), and whether this may be in some way related to increased endogenous oxytocin and reduced cortisol levels in individuals involved in musical activities—by what mechanisms could music, memory and oxytocin be linked? Acting through its receptor, oxytocin can act both pre-and postsynaptically to enhance LTP, alter the balance of excitatory and inhibitory activity, and modulate synaptic plasticity (Tomizawa et al., 2003; Lee et al., 2015; Bakos et al., 2018; Lin and Hsu, 2018; Tirko et al., 2018), These are all critical elements during the process of learning, socialization and memory consolidation (Ferguson et al., 2000; Lin et al., 2018), and intranasal application of the peptide at low doses is known to enhance social memory in human subjects (Jurek and Neumann, 2018).

Neurogenesis

The hippocampal dentate gyrus appears to be one of the few sites in the adult mammalian CNS where new neurons are born throughout life (neurogenesis). This ongoing process is thought to be important in learning and in facilitating the addition of new memories onto similar previous experiences and knowledge, minimizing overlap in the resultant patterns of activity so that particular events can be discriminated from each other (e.g., Conçalves et al., 2016; França et al., 2017; Alam et al., 2018; Toda and Gage, 2018; Licht et al., 2020). In animals, experimental disruption of neurogenesis impairs social memory and coping with stress (Clelland et al., 2009; Garrett et al., 2015; Alam et al., 2018). Social interactions enhance new neuronal birth (Hsiao et al., 2014) whereas social isolation and stress-related changes that include increased cortisol levels lead to a reduction in neurogenesis (McEwen, 1999; Cinini et al., 2014; Opendak et al., 2016; Snyder and Drew, 2020), negatively affecting cognition, learning, and memory (Ouanes and Popp, 2019).

Oxytocin protects the hippocampus from stress-related effects including the negative impact of corticosterone treatment and directly induces neurogenesis in the adult rodent dentate gyrus (Lee et al., 2015; Sánchez-Vidaña et al., 2016; Lin et al., 2017; Lin and Hsu, 2018). The peptide also promotes the differentiation and dendritic maturation of these new neurons with associated effects on social behavior (Sánchez-Vidaña et al., 2016). This influence of oxytocin on hippocampal neurogenesis and social learning is indirectly enhanced by the peptide’s actions in increasing BDNF expression (Dayi et al., 2015; Havranek et al., 2015; Zhang et al., 2020). This neurotrophin plays a key role in hippocampal plasticity and neurogenesis (Miranda et al., 2019). Its expression in the hippocampus is negatively affected by stress (Bennett and Lagopoulos, 2014; Dayi et al., 2015) but is significantly increased by physical exercise (Ding et al., 2011). In animals, increased BDNF levels are correlated with increased neurogenesis, the greater the amount of exercise the greater the proliferation of new neurons (reviewed in Liu and Nusslock, 2018). Indeed, it was recently shown that intense physical activity releases breakdown products from the muscle that act on promoters to increase BDNF gene expression and protein (Sleiman et al., 2016; Stephan and Sleiman, 2019).

There remains some controversy as to whether new neurons are born and survive within the adult human dentate gyrus (Sorrells et al., 2018; Duque and Spector, 2019); however, the weight of evidence and opinion is that neurogenesis and neuronal turnover does occur (Spalding et al., 2013; Boldrini et al., 2018; Kempermann et al., 2018; Kuhn et al., 2018; Cope and Gould, 2019; Horgusluoglu-Moloch et al., 2019; Lima and Gomes-Leal, 2019; Petrik and Encinas, 2019; Tobin et al., 2019; Lucassen et al., 2020), although estimates of the number of neurons born each day vary, and numbers may decline with age and disease (Moreno-Jiménez et al., 2019; Snyder, 2019). It is however clear that exercise and cardiovascular fitness are correlated with increased hippocampal volume and improved cognitive function (Erickson et al., 2011; Stillman et al., 2016). Furthermore, while it is not yet known if such changes are associated with enhanced neurogenesis, hippocampal size in humans is correlated with plasma BDNF levels (Erickson et al., 2011).

It is well established that neural activity in the hippocampus and other parts of the limbic system is altered by listening to music. Given this, and what is known about the effects of music on hormones such as oxytocin and cortisol, it will be of interest to determine if participation in musical activities influences human hippocampal neurogenesis (Fukui and Toyoshima, 2008), and how this might relate to the known beneficial effects of music on memory and cognition. Such activities should include movement and dance which are entrained within the diverse neural networks responsive to music (e.g., Brown et al., 2006; Phillips-Silver and Trainor, 2007; Nozaradan et al., 2011). Dance not only increases cooperation and group synchrony (Reddish et al., 2013; Karpati et al., 2016; Chauvigné et al., 2019) but improves fitness in the elderly (Douka et al., 2019). Measurement of oxytocin, cortisol, and BDNF in dancers seems warranted, and it may well be that, in addition to potential oxytocin-mediated effects, exercise and cardiovascular fitness associated with dancing are capable of adding an important extra dimension to the social, physical and mental health benefits of music appreciation and music-related activities, perhaps especially in the elderly.

Systemic Effects—Further Links Between Oxytocin and Musicality

In addition to its physiological effects on CNS function, oxytocin has been reported to have even broader health benefits. The peptide decreases the progression of atherosclerosis and protects against cardiovascular disease (Reiss et al., 2019; Wang et al., 2019; Buemann and Uvnäs-Moberg, 2020), in association with social engagement (Ulmer-Yaniv et al., 2016; Walker et al., 2019) it has beneficial effects on the immune system, and it lowers cytokine levels and inhibits inflammation (Li et al., 2017b; Reiss et al., 2019). Oxytocin has also been reported to regulate appetite and food intake (Lawson et al., 2019; Onaka and Takayanagi, 2019; Quintana et al., 2019). Associated with these multiple beneficial effects, the peptide has been found to lower blood pressure and assist in maintaining glucose homeostasis, potentially useful as a therapeutic tool in the treatment of type 2 diabetes and obesity (Reiss et al., 2019). Again, many of these systemic oxytocinergic effects overlap those that can be elicited by listening to and/or performing music, including a reduction in blood pressure, the modification of immune responses and inflammatory markers, reduction of anxiety and stress and a moderating effect on blood glucose levels (Koelsch and Jäncke, 2015; Finn and Fancourt, 2018; Kang et al., 2018). The potentially additional benefits of music-related exercises such as dance have already been alluded to in the preceding paragraphs.

Conclusion and Therapeutic Implications

Given what is increasingly becoming known about the neurological and systemic effects of oxytocin, it is important to analyze further how music and dance influence this peptide and its downstream pathways, and the extent to which social musical activities drive some of the interactions between oxytocin and other neuromodulatory systems such as dopamine, BDNF, and the various endogenous opioids. From an evolutionary perspective, it may clarify the extent to which evolving musical capabilities in modern humans took advantage of the ancient oxytocinergic network to facilitate prosocial interactions, promote trust and reciprocal affiliative behaviors, and help reduce levels of anxiety and individual insecurity throughout life. It will also contribute to a better understanding of the mechanisms that underlie the mental and general health benefits of music, its remarkable emotional and mnemonic power, it’s capacity to alter brain architecture, and its ability to revitalize episodic memories (Zhang et al., 2017; Särkämö and Sihvonen, 2018), especially vulnerable in early stages of Alzheimer’s disease (Groussard et al., 2019; Slattery et al., 2019).

From a therapeutic perspective, dancing is already a useful tool in the treatment of Parkinson’s disease (Pereira et al., 2019); perhaps for some dementia sufferers, in addition to singing, dancing to favorite tunes may be even more beneficial when promoting social interactions with partners and unlocking autobiographical memories. Similarly, the use of intranasal oxytocin as a therapeutic tool in conditions such as PTSD shows promise (Giovanna et al., 2020) and may be even more effective when used with other treatments including combination with appropriate prosocial music-related activities. As another example, the combined use of music and the anti-nociceptive properties of oxytocin may enhance the therapeutic efficacy of strategies aimed at reducing the perception of chronic (Garza-Villarreal et al., 2017; Hilfiger et al., 2020; Schneider et al., 2020) or peri-operative pain (Nilsson et al., 2005, 2009; Nilsson, 2009; Van der Heijden et al., 2015). Last but not least, the use of oxytocin in the treatment of autism spectrum disorders (Yamasue and Domes, 2018) and other psychiatric conditions (Peled-Avron et al., 2020) may benefit from synergistic application with appropriate music-related therapeutic strategies (Quintin, 2019).

Author’s Note

During revision of this manuscript, a review was published that emphasized the importance of studying the biology of oxytocin systems in animals to better understand the translational potential of this peptide in psychiatry and mental health (Grinevich and Neumann, 2020). The reader is encouraged to access this review to complement the more focussed emphasis on human musicality described herein.

Author Contributions

The author confirms being the sole contributor of this work and has approved it for publication.

Conflict of Interest

The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

References

Abraha, I., Rimland, J. M., Trotta, F. M., Dell’Aquila, G., Cruz-Jentoft, A., Petrovic, M., et al. (2017). Systematic review of systematic reviews of non-pharmacological interventions to treat behavioural disturbances in older patients with dementia. The SENATOR-OnTop series. BMJ Open 7:e012759. doi: 10.1136/bmjopen-2016-012759

PubMed Abstract | CrossRef Full Text | Google Scholar

Ahn, H., La, J. H., Chung, J. M., Miao, H., Zhong, C., Kim, M., et al. (2019). The relationship between β-endorphin and experimental pain sensitivity in older adults with knee osteoarthritis. Biol. Res. Nurs. 21, 400–406. doi: 10.1177/1099800419853633

PubMed Abstract | CrossRef Full Text | Google Scholar

Alam, M. J., Kitamura, T., Saitoh, Y., Ohkawa, N., Kondo, T., and Inokuchi, K. (2018). Adult neurogenesis conserves hippocampal memory capacity. J. Neurosci. 38, 6854–6863. doi: 10.1523/jneurosci.2976-17.2018

PubMed Abstract | CrossRef Full Text | Google Scholar

Alcorn, J. L. III., Green, C. E., Schmitz, J., and Lane, S. D. (2015). Effects of oxytocin on aggressive responding in healthy adult men. Behav. Pharmacol. 8, 798–804. doi: 10.1097/fbp.0000000000000173

PubMed Abstract | CrossRef Full Text | Google Scholar

Alos-Ferrer, C., and Farolfi, F. (2019). Trust games and beyond. Front. Neurosci. 13:887. doi: 10.3389/fnins.2019.00887

PubMed Abstract | CrossRef Full Text | Google Scholar

Altenmüller, E., Siggel, S., Mohammadi, B., Samii, A., and Münte, T. F. (2014). Play it again, Sam: brain correlates of emotional music recognition. Front. Psychol. 5:114. doi: 10.3389/fpsyg.2014.00114

PubMed Abstract | CrossRef Full Text | Google Scholar

Angulo-Perkins, A., Aubé, W., Peretz, I., Barios, F. A., Armony, J. L., and Concha, L. (2014). Music listening engages specific cortical regions within the temporal lobes: differences between musicians and non-musicians. Cortex 59, 126–137. doi: 10.1016/j.cortex.2014.07.013

PubMed Abstract | CrossRef Full Text | Google Scholar

Aspé-Sánchez, M., Moreno, M., Rivera, M. I., Rossi, A., and Ewer, J. (2016). Oxytocin and vasopressin receptor gene polymorphisms: role in social and psychiatric traits. Front. Neurosci. 9:510. doi: 10.3389/fnins.2015.00510

PubMed Abstract | CrossRef Full Text | Google Scholar

Aydogan, G., Jobst, A., Loy, F., Dehning, S., Zill, P., Müller, N., et al. (2018). The effect of oxytocin on group formation and strategic thinking in men. Horm. Behav. 100, 100–106. doi: 10.1016/j.yhbeh.2018.02.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Bachner-Melman, R., Dina, C., Zohar, A. H., Constantini, N., Lerer, E., Hoch, S., et al. (2005). AVPR1a and SLC6A4 gene polymorphisms are associated with creative dance performance. PLoS Genet. 1:e42. doi: 10.1371/journal.pgen.0010042

PubMed Abstract | CrossRef Full Text | Google Scholar

Bakos, J., Srancikova, A., Havranek, T., and Bacova, Z. (2018). Molecular mechanisms of oxytocin signaling at the synaptic connection. Neural Plast. 2018:4864107. doi: 10.1155/2018/4864107

PubMed Abstract | CrossRef Full Text | Google Scholar

Bali, A., Randhawa, P. K., and Jaggi, A. S. (2015). Stress and opioids: role of opioids in modulating stress-related behavior and effect of stress on morphine conditioned place preference. Neurosci. Biobehav. Rev. 51, 138–150. doi: 10.1016/j.neubiorev.2014.12.018

PubMed Abstract | CrossRef Full Text | Google Scholar

Bartz, J. A., Nitschke, J. P., Krol, S. A., and Tellier, P. P. (2019). Oxytocin selectively improves empathic accuracy: a replication in men and novel insights in women. Biol. Psychiatry Cogn. Neurosci. Neuroimaging 4, 1042–1048. doi: 10.1016/j.bpsc.2019.01.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Baumgartner, T., Heinrichs, M., Vonlanthen, A., Fischbacher, U., and Fehr, E. (2008). Oxytocin shapes the neural circuitry of trust and adaptation in humans. Neuron 58, 639–650. doi: 10.1016/j.neuron.2008.04.009

PubMed Abstract | CrossRef Full Text | Google Scholar

Beck, B. D., Lund, S. T., Søgaard, U., Simonsen, E., Tellier, T. C., Cordtz, T. O., et al. (2018). Music therapy versus treatment as usual for refugees diagnosed with posttraumatic stress disorder (PTSD): study protocol for a randomized controlled trial. Trials 19:301. doi: 10.1186/s13063-018-2662-z

PubMed Abstract | CrossRef Full Text | Google Scholar

Benarroch, E. E. (2013). Oxytocin and vasopressin: social neuropeptides with complex modulatory functions. Neurology 80, 1521–1528. doi: 10.1212/wnl.0b013e31828cfb15

PubMed Abstract | CrossRef Full Text | Google Scholar

Bennett, M. R., and Lagopoulos, J. (2014). Stress and trauma: BDNF control of dendritic-spine formation and regression. Prog. Neurobiol. 112, 80–99. doi: 10.1016/j.pneurobio.2013.10.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Berends, Y. R., Tulen, J. H. M., Wierdsma, A. I., van Pelt, J., Kushner, S. A., and van Marle, H. J. C. (2019). Oxytocin, vasopressin and trust: associations with aggressive behavior in healthy young males. Physiol. Behav. 204, 180–185. doi: 10.1016/j.physbeh.2019.02.027

PubMed Abstract | CrossRef Full Text | Google Scholar

Bethlehem, R. A., Baron-Cohen, S., Van Honk, J., Auyeung, B., and Bos, P. A. (2014). The oxytocin paradox. Front. Behav. Neurosci. 8:48. doi: 10.3389/fnbeh.2014.00048

PubMed Abstract | CrossRef Full Text | Google Scholar

Bethlehem, R. A., van Honk, J., Auyeung, B., and Baron-Cohen, S. (2013). Oxytocin, brain physiology and functional connectivity: a review of intranasal oxytocin fMRI studies. Psychoneuroendocrinology 38, 962–974. doi: 10.1016/j.psyneuen.2012.10.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Blood, A. J., and Zatorre, R. L. (2001). Intensely pleasurable responses to music correlate with activity in brain regions implicated in reward and emotion. Proc. Natl. Acad. Sci. U S A 98, 11818–11823. doi: 10.1073/pnas.191355898

PubMed Abstract | CrossRef Full Text | Google Scholar

Boccia, M. L., Petrusz, P., Suzuki, K., Marson, L., and Pederson, C. A. (2013). Immunohistochemical localization of oxytocin receptors in human brain. Neuroscience 253, 155–164. doi: 10.1016/j.neuroscience.2013.08.048

PubMed Abstract | CrossRef Full Text | Google Scholar

Boldrini, M., Fulmore, C. A., Tartt, A. N., Simeon, L. R., Pavlova, I., Poposka, V., et al. (2018). Human hippocampal neurogenesis persists throughout aging. Cell Stem Cell 22, 589.e5–599.e5. doi: 10.1016/j.stem.2018.03.015

PubMed Abstract | CrossRef Full Text | Google Scholar

Bonnici, H. M., and Maguire, E. A. (2018). Two years later - revisiting autobiographical memory representations in vmPFC and hippocampus. Neuropsychologia 110, 159–169. doi: 10.1016/j.neuropsychologia.2017.05.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Boso, M., Politi, P., Barale, F., and Enzo, E. (2006). Neurophysiology and neurobiology of the musical experience. Funct. Neurol. 21, 187–191.

PubMed Abstract | Google Scholar

Brattico, E., Alluri, V., Bogert, B., Jacobsen, T., Vartiainen, N., Nieminen, S., et al. (2011). A functional MRI study of happy and sad emotions in music with and without lyrics. Front. Psychol. 2:308. doi: 10.3389/fpsyg.2011.00308

PubMed Abstract | CrossRef Full Text | Google Scholar

Bredewold, R., and Veenema, R. H. (2018). Sex differences in the regulation of social and anxiety-related behaviors: insights from vasopressin and oxytocin brain systems. Curr. Opin. Neurobiol. 49, 132–140. doi: 10.1016/j.conb.2018.02.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Brown, S. (2000). “The ‘musilanguage’ model of human evolution,” in Origins of Music, eds N. Wallin, B. Merker and S. Brown (Cambridge, MA: MIT Press), 271–300.

Google Scholar

Brown, C. A., Cardoso, C., and Ellenbogen, M. A. (2016). A meta-analytic review of the correlation between peripheral oxytocin and cortisol concentrations. Front. Neuroendocrinol. 43, 19–27. doi: 10.1016/j.yfrne.2016.11.001

PubMed Abstract | CrossRef Full Text | Google Scholar

Brown, S., Martinez, M. J., and Parsons, L. M. (2006). The neural basis of dance. Cereb. Cortex 16, 1157–1167. doi: 10.1093/cercor/bhj057

PubMed Abstract | CrossRef Full Text | Google Scholar

Bruehl, S., Burns, J. W., Gupta, R., Buvanendran, A., Chont, M., Orlowska, D., et al. (2017). Do resting plasma β-endorphin levels predict responses to opioid analgesics? Clin. J. Pain. 33, 12–20. doi: 10.1097/AJP.0000000000000389

PubMed Abstract | CrossRef Full Text | Google Scholar

Buemann, B., and Uvnäs-Moberg, K. (2020). Oxytocin may have a therapeutical potential against cardiovascular disease. Possible pharmaceutical and behavioral approaches. Med. Hypoth. 138:109597. doi: 10.1016/j.mehy.2020.109597

PubMed Abstract | CrossRef Full Text | Google Scholar

Busnelli, M., and Chini, B. (2017). “Molecular basis of oxytocin receptor signalling in the brain: what we know and what we need to know,” in Behavioral Pharmacology of Neuropeptides: Oxytocin. Current Topics in Behavioral Neurosciences. (Vol. 35), eds R. Hurlemann and V. Grinevich (Cham: Springer). doi: 10.1007/7854_2017_6

CrossRef Full Text | Google Scholar

Carter, C. S. (2014). Oxytocin pathways and the evolution of human behavior. Annu. Rev. Psychol. 65, 17–39. doi: 10.1146/annurev-psych-010213-115110

PubMed Abstract | CrossRef Full Text | Google Scholar

Carter, R. M., Bowling, D. L., Reeck, C., and Huettel, S. A. (2012). A distinct role of the temporal-parietal junction in predicting socially guided behaviour. Science 337, 109–111. doi: 10.1126/science.1219681

PubMed Abstract | CrossRef Full Text | Google Scholar

Chanda, M. L., and Levitin, D. J. (2013). The neurochemistry of music. Trends Cogn. Sci. 17, 179–193. doi: 10.1016/j.tics.2013.02.007

PubMed Abstract | CrossRef Full Text | Google Scholar

Chapin, H., Jantzen, K., Kelso, J. A., Steinberg, F., and Large, E. (2010). Dynamic emotional and neural responses to music depend on performance expression and listener experience. PLoS One 5:e13812. doi: 10.1371/journal.pone.0013812

PubMed Abstract | CrossRef Full Text | Google Scholar

Chatterjee, O., Patil, K., Sahu, A., Gopalakrishnan, L., Mol, P., Advani, J., et al. (2016). An overview of the oxytocin-oxytocin receptor signaling network. J. Cell Commun. Signal. 10, 355–360. doi: 10.1007/s12079-016-0353-7

PubMed Abstract | CrossRef Full Text | Google Scholar

Chauvigné, L. A. S., Walton, A., Richardson, M. J., and Brown, S. (2019). Multi-person and multisensory synchronization during group dancing. Hum. Mov. Sci. 63, 199–208. doi: 10.1016/j.humov.2018.12.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Chen, X., Hackett, P. D., DeMarco, A. C., Feng, C., Stair, S., Haroon, E., et al. (2016). Effects of oxytocin and vasopressin on the neural response to unreciprocated cooperation within brain regions involved in stress and anxiety in men and women. Brain Imaging Behav. 10, 581–593. doi: 10.1007/s11682-015-9411-7

PubMed Abstract | CrossRef Full Text | Google Scholar

Chen, X., Nishitani, S., Haroon, E., Smith, A. K., and Rilling, J. K. (2020). OXTR methylation modulates exogenous oxytocin effects on human brain activity during social interaction. Genes Brain Behav. 19:e12555. doi: 10.1111/gbb.12555

PubMed Abstract | CrossRef Full Text | Google Scholar

Chini, B., Verhage, M., and Grinevich, V. (2017). The action radius of oxytocin release in the mammalian CNS: from single vesicles to behavior. Trends Pharmacol. Sci. 38, 982–991. doi: 10.1016/j.tips.2017.08.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Chruścicka, B., Wallace Fitzsimons, S. E., Borroto-Escuela, D. O., Druelle, C., Stamou, P., Nally, K., et al. (2019). Attenuation of oxytocin and serotonin 2A receptor signaling through novel heteroreceptor formation. ACS Chem. Neurosci. 10, 3225–3240. doi: 10.1021/acschemneuro.8b00665

PubMed Abstract | CrossRef Full Text | Google Scholar

Cilz, N. I., Cymerblit-Sabba, A., and Young, W. S. (2019). Oxytocin and vasopressin in the rodent hippocampus. Genes Brain Behav. 18:e12535. doi: 10.1111/gbb.12535

PubMed Abstract | CrossRef Full Text | Google Scholar

Cinini, S. M., Barnabe, G. F., Galváo-Coelho, N., de Medeiros, M. A., Perez-Mendes, P., Sousa, M. B. C., et al. (2014). Social isolation disrupts hippocampal neurogenesis in young non-human primates. Front. Neurosci. 8:45. doi: 10.3389/fnins.2014.00045

PubMed Abstract | CrossRef Full Text | Google Scholar

Clelland, C. D., Choi, M., Romberg, C., Clemenson, G. D. Jr., Fragniere, A., Tyers, P., et al. (2009). A functional role for adult hippocampal neurogenesis in spatial pattern separation. Science 325, 210–213. doi: 10.1126/science.1173215

PubMed Abstract | CrossRef Full Text | Google Scholar

Conçalves, J. T., Schafer, S. T., and Gage, F. H. (2016). Adult neurogenesis in the hippocampus: from stem cells to behavior. Cell 167, 897–914. doi: 10.1016/j.cell.2016.10.021

PubMed Abstract | CrossRef Full Text | Google Scholar

Cooper, J. C., Kreps, T. A., Wiebe, T., Pirkl, T., and Knutson, B. (2010). When giving is good: ventromedial prefrontal cortex activation for others’ intentions. Neuron 67, 511–521. doi: 10.1016/j.neuron.2010.06.030

PubMed Abstract | CrossRef Full Text | Google Scholar

Cope, E. C., and Gould, E. (2019). Adult neurogenesis, glia and the extracellular matrix. Cell Stem Cell 24, 690–705. doi: 10.1016/j.stem.2019.03.023

PubMed Abstract | CrossRef Full Text | Google Scholar

Cox, B. M., Bentzley, B. S., Regen-Tuero, H., See, R. E., Reichel, C. M., and Aston-Jones, G. (2017). Oxytocin acts in nucleus accumbens to attenuate methamphetamine seeking and demand. Biol. Psych. 81, 949–958. doi: 10.1016/j.biopsych.2016.11.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Crane, J. W., Holmes, N. M., Fam, J., Westbrook, R. F., and Delaney, A. J. (2020). Oxytocin increases inhibitory synaptic transmission and blocks the development of long-term potentiation in the lateral amygdala. J. Neurophysiol. 123, 587–599. doi: 10.1152/jn.00571.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Cross, I. (2009). The evolutionary nature of musical meaning. Music Sci. 13, 179–200. doi: 10.1177/1029864909013002091

CrossRef Full Text | Google Scholar

Dadds, M. R., Moul, C., Cauchi, A., Dobson-Stone, C., Hawes, D. J., Brennan, J., et al. (2014). Polymorphisms in the oxytocin receptor gene are associated with the development of psychopathy. Dev. Psychopathol. 26, 21–31. doi: 10.1017/s0954579413000485

PubMed Abstract | CrossRef Full Text | Google Scholar

dal Monte, O., Piva, M., Anderson, K. M., Tringides, M., Holmes, A. J., and Chang, S. W. C. (2017). Oxytocin under opioid antagonism leads to supralinear enhancement of social attention. Proc. Natl. Acad. Sci. U S A 114, 5247–5252. doi: 10.1073/pnas.1702725114

PubMed Abstract | CrossRef Full Text | Google Scholar

Dayi, A., Cetin, F., Sisman, A. R., Aksu, I., Tas, A., Gönenc, S., et al. (2015). The effects of oxytocin on cognitive defect caused by chronic restraint stress applied to adolescent rats and on hippocampal VEGF and BDNF levels. Med. Sci. Monit. 21, 69–75. doi: 10.12659/msm.893159

PubMed Abstract | CrossRef Full Text | Google Scholar

De Dreu, C. K. (2012). Oxytocin modulates cooperation within and competition between groups: an integrative review and research agenda. Horm. Behav. 61, 419–428. doi: 10.1016/j.yhbeh.2011.12.009

PubMed Abstract | CrossRef Full Text | Google Scholar

De Dreu, C. K., and Kret, M. E. (2016). Oxytocin conditions intergroup relations through upregulated in-group empathy, cooperation, conformity, and defense. Biol. Psychiatry 79, 165–173. doi: 10.1016/j.biopsych.2015.03.020

PubMed Abstract | CrossRef Full Text | Google Scholar

de Jong, T. R., and Neumann, I. D. (2018). Oxytocin and aggression. Curr. Top. Behav. Neurosci. 35, 175–192. doi: 10.1007/7854_2017_13

PubMed Abstract | CrossRef Full Text | Google Scholar

de Manzano, Ö., and Ullén, F. (2018). Same genes, different brains: neuroanatomical differences between monozygotic twins discordant for musical training. Cereb. Cortex 28, 387–394. doi: 10.1093/cercor/bhx299

PubMed Abstract | CrossRef Full Text | Google Scholar

Deuse, L., Wudarczyk, O., Rademacher, L., Kaleta, P., Karges, W., Kacheva, S., et al. (2019). Peripheral oxytocin predicts higher-level social cognition in men regardless of empathy quotient. Pharmacopsychiatry 52, 148–154. doi: 10.1055/a-0590-4850

PubMed Abstract | CrossRef Full Text | Google Scholar

De Wilde, T. R., Ten Velden, F. S., and De Dreu, C. K. (2017). The neuropeptide oxytocin enhances information sharing and group decision making quality. Sci. Rep. 7:40622. doi: 10.1038/srep40622

PubMed Abstract | CrossRef Full Text | Google Scholar

Ding, Q., Ying, Z., and Gómez-Pinilla, F. (2011). Exercise influences hippocampal plasticity by modulating brain-derived neurotrophic factor processing. Neuroscience 192, 773–780. doi: 10.1016/j.neuroscience.2011.06.032

PubMed Abstract | CrossRef Full Text | Google Scholar

Ditzen, B., Schaer, M., Gabriel, B., Bodenmann, G., Ehlert, U., and Heinrichs, M. (2009). Intranasal oxytocin increases positive communication and reduces cortisol levels during couple conflict. Biol. Psychiatry 65, 728–731. doi: 10.1016/j.biopsych.2008.10.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Dölen, G., Darvishzadeh, A., Huang, K. W., and Malenka, R. C. (2013). Social reward requires coordinated activity of nucleus accumbens oxytocin and serotonin. Nature 510, 179–184. doi: 10.1038/nature12518

PubMed Abstract | CrossRef Full Text | Google Scholar

Donadon, M. F., Martin-Santos, R., and Osório, F. L. (2018). The associations between oxytocin and trauma in humans: a systematic review. Front. Pharmacol. 9:154. doi: 10.3389/fphar.2018.00154

PubMed Abstract | CrossRef Full Text | Google Scholar

Donnay, G. F., Rankin, S. K., Lopez-Gonzalez, M., Jiradejvong, P., and Limb, C. J. (2014). Neural substrates of interactive musical improvisation: an fMRI study of ‘trading fours’ in jazz. PLoS One 9:e88665. doi: 10.1371/journal.pone.0088665

PubMed Abstract | CrossRef Full Text | Google Scholar

Douka, S., Zilidou, V. I., Lilou, O., and Manou, V. (2019). Traditional dance improves the physical fitness and well-being of the elderly. Front. Aging Neurosci. 11:75. doi: 10.3389/fnagi.2019.00075

PubMed Abstract | CrossRef Full Text | Google Scholar

Dunbar, R. I. M., Kaskatis, K., MacDonald, I., and Barra, V. (2012). Performance of music elevates pain threshold and positive affect: implications for the evolutionary function of music. Evol. Psychol. 10, 688–702. doi: 10.1177/147470491201000403

PubMed Abstract | CrossRef Full Text | Google Scholar

Dunsmoor, J. E., Kroes, M. C. W., Li, J., Daw, N. D., Simpson, H. B., and Phelps, E. A. (2019). Role of human ventromedial prefrontal cortex in learning and recall of enhanced extinction. J. Neurosci. 39, 3264–3276. doi: 10.1523/JNEUROSCI.2713-18.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Duque, A., and Spector, R. (2019). A balanced evaluation of the evidence for adult neurogenesis in humans: implication for neuropsychiatric disorders. Brain Struct. Funct. 224, 2281–2295. doi: 10.1007/s00429-019-01917-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Ebitz, R. B., and Platt, M. L. (2014). An evolutionary perspective on the behavioral consequences of exogenous oxytocin application. Front. Behav. Neurosci. 7:225. doi: 10.3389/fnbeh.2013.00225

PubMed Abstract | CrossRef Full Text | Google Scholar

Ebstein, R. P., Knafo, A., Mankuta, D., Chew, S. H., and Lai, P. S. (2012). The contributions of oxytocin and vasopressin pathway genes to human behavior. Horm. Behav. 61, 359–379. doi: 10.1016/j.yhbeh.2011.12.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Eckstein, M., Markett, S., Kendrick, K. M., Ditzen, B., Liu, F., Hurlemann, R., et al. (2017). Oxytocin differentially alters resting state functional connectivity between amygdala subregions and emotional control networks: inverse correlation with depressive traits. NeuroImage 149, 458–467. doi: 10.1016/j.neuroimage.2017.01.078

PubMed Abstract | CrossRef Full Text | Google Scholar

Ellenbogen, M. A. (2018). Oxytocin and facial emotion recognition. Curr. Top. Behav. Neurosci. 35, 349–374. doi: 10.1007/7854_2017_20

PubMed Abstract | CrossRef Full Text | Google Scholar

Erdozain, A. M., and Peñagarikano, O. (2020). Oxytocin as a treatment for social cognition, not there yet. Front. Psychiatry 10:930. doi: 10.3389/fpsyt.2019.00930

PubMed Abstract | CrossRef Full Text | Google Scholar

Erickson, K. I., Miller, D. L., and Roecklein, K. A. (2011). The aging hippocampus: interactions between exercise, depression and BDNF. Neuroscientist 18, 82–97. doi: 10.1177/1073858410397054

PubMed Abstract | CrossRef Full Text | Google Scholar

Evans, S. L., Dal Monte, O., Noble, P., and Averbeck, B. B. (2014). Intranasal oxytocin effects on social cognition: a critique. Brain Res. 1580, 69–77. doi: 10.1016/j.brainres.2013.11.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Fancourt, D., Ockelford, A., and Belai, A. (2014). The psychoneuroimmunological effects of music: a systematic review and a new model. Brain Behav. Immun. 36, 15–26. doi: 10.1016/j.bbi.2013.10.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Feldman, R. (2012). Oxytocin and social affiliation in humans. Horm. Behav. 61, 380–391. doi: 10.1016/j.yhbeh.2012.01.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Feldman, R. (2015). The adaptive human parental brain: implications for children’s social development. Trends Neurosci. 38, 387–399. doi: 10.1016/j.tins.2015.04.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Feldman, R. (2017). The neurobiology of human attachments. Trends Cogn. Sci. 21, 80–99. doi: 10.1016/j.tics.2016.11.007

PubMed Abstract | CrossRef Full Text | Google Scholar

Feldman, R., Monakhov, M., Pratt, M., and Epstein, R. P. (2016). Oxytocin pathway genes: evolutionary ancient system impacting on human affiliation, sociality and psychopathology. Biol. Psychiatry 79, 174–184. doi: 10.1016/j.biopsych.2015.08.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Feldman, R., Weller, A., Zagoory-Sharon, O., and Levine, A. (2007). Evidence for a neuroendocrinological foundation of human affiliation: plasma oxytocin levels across pregnancy and the postpartum period predict mother-infant bonding. Psychol. Sci. 18, 965–970. doi: 10.1111/j.1467-9280.2007.02010.x

PubMed Abstract | CrossRef Full Text | Google Scholar

Feng, C., Hackett, P. D., DeMarco, A. C., Chen, X., Stair, S., Haroon, E., et al. (2015). Oxytocin and vasopressin effects on the neural response to social cooperation are modulated by sex in humans. Brain Imaging Behav. 9, 754–764. doi: 10.1007/s11682-014-9333-9

PubMed Abstract | CrossRef Full Text | Google Scholar

Ferguson, J. N., Young, L. J., Hearn, E. F., Matzuk, M. M., Insel, T. R., and Winslow, J. T. (2000). Social amnesia in mice lacking the oxytocin gene. Nat. Genet. 25, 284–288. doi: 10.1038/77040

PubMed Abstract | CrossRef Full Text | Google Scholar

Ferguson, J. N., Young, L. J., and Insel, T. R. (2002). The neuroendocrine basis of social recognition. Front. Neuroendocrinol. 23, 200–224. doi: 10.1006/frne.2002.0229

PubMed Abstract | CrossRef Full Text | Google Scholar

Ferreri, L., Mas-Herrero, E., Zatorre, R. J., Ripollés, P., Gomez-Andres, A., Alicart, H., et al. (2019). Dopamine modulates the reward experiences elicited by music. Proc. Natl. Acad. Sci. U S A 116, 3793–3798. doi: 10.1073/pnas.1811878116

PubMed Abstract | CrossRef Full Text | Google Scholar

Fineberg, S. K., and Ross, D. A. (2017). Oxytocin and the social brain. Biol. Psychiatry 81, e19–e21. doi: 10.1016/j.biopsych.2016.11.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Finn, S., and Fancourt, D. (2018). The biological impact of listening to music in clinical and nonclinical settings: a systematic review. Prog. Brain Res. 237, 173–200. doi: 10.1016/bs.pbr.2018.03.007

PubMed Abstract | CrossRef Full Text | Google Scholar

Fischer-Shofty, M., Levkovitz, Y., and Shamay-Tsoory, S. G. (2012). Oxytocin facilitates accurate perception of competition in men and kinship in women. Soc. Cogn. Affect. Neurosci. 8, 313–317. doi: 10.1093/scan/nsr100

PubMed Abstract | CrossRef Full Text | Google Scholar

Fitch, T. W. (2006). The biology and evolution of music: a comparative perspective. Cognition 100, 173–215. doi: 10.1016/j.cognition.2005.11.009

PubMed Abstract | CrossRef Full Text | Google Scholar

Fragkaki, I., and Cima, M. (2019). The effect of oxytocin administration on empathy and emotion recognition in residential youth: a randomized, within-subjects trial. Horm. Behav. 114:104561. doi: 10.1016/j.yhbeh.2019.104561

PubMed Abstract | CrossRef Full Text | Google Scholar

França, T. F. A., Bitencourt, A. M., Maximilla, N. R., Barros, D. M., and Monserrat, J. M. (2017). Hippocampal neurogenesis and pattern separation: a meta-analysis of behavioral data. Hippocampus 27, 937–950. doi: 10.1002/hipo.22746

PubMed Abstract | CrossRef Full Text | Google Scholar

Franke, A. A., Li, X., Dabalos, C., and Lai, J. F. (2020). Improved oxytocin analysis from human serum and urine by orbitrap ESI-LC-HRAM-MS. Drug Test Anal. 12, 846–852. doi: 10.1002/dta.2783

PubMed Abstract | CrossRef Full Text | Google Scholar

Franke, A. A., Li, X., Menden, A., Lee, M. R., and Lai, J. F. (2019). Oxytocin analysis from human serum, urine and saliva by orbitrap liquid chromatography-mass spectrometry. Drug Test Anal. 11, 119–128. doi: 10.1002/dta.2475

PubMed Abstract | CrossRef Full Text | Google Scholar

Freeman, W. (2000). “A neurological role of music in social bonding,” in The Origins of Music, eds N. L. Wallin, B. Merker and S. Brown (Cambridge, MA: MIT Press), 411–424.

Google Scholar

Frijling, J. L., van Zuiden, M., Nawijn, L., Koch, S. B., Neumann, I. D., Veltman, D. J., et al. (2015). Salivary oxytocin and vasopressin levels in police officers with and without post-traumatic stress disorder. J. Neuroendocrinol. 27, 743–751. doi: 10.1111/jne.12300

PubMed Abstract | CrossRef Full Text | Google Scholar

Fujiwara, T., Weisman, O., Ochi, M., Shirai, K., Matsumoto, K., Noguchi, E., et al. (2019). Genetic and peripheral markers of the oxytocin system and parental care jointly support the cross-generational transmission of bonding across three generations. Psychoneuroendocrinology 102, 172–181. doi: 10.1016/j.psyneuen.2018.12.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Fukuda, H., Ma, N., Suzuki, S., Harasawa, N., Ueno, K., Gardner, J. L., et al. (2019). Computing social value conversion in the human brain. J. Neurosci. 39, 5153–5172. doi: 10.1523/JNEUROSCI.3117-18.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Fukui, H., and Toyoshima, K. (2008). Music facilitate the neurogenesis, regeneration and repair of neurons. Med. Hypotheses 71, 765–769. doi: 10.1016/j.mehy.2008.06.019

PubMed Abstract | CrossRef Full Text | Google Scholar

Gamal-Eltrabily, M., de los Monteros-Zúñiga, E. A., Manzano-García, A., Martínez-Lorenzana, G., Condés-Lara, M., and González-Hernández, A. (2020). The rostral agranular insular cortex, a new site of oxytocin to induce antinociception. J. Neurosci. 40, 5669–5680. doi: 10.1523/jneurosci.0962-20.2020

PubMed Abstract | CrossRef Full Text | Google Scholar

Garrett, L., Zhang, J., Zimprich, A., Niedermeier, K. M., Fuchs, H., Gailus-Durner, V., et al. (2015). Conditional reduction of adult born doublecortin-positive neurons reversibly impairs selective behaviors. Front. Behav. Neurosci. 9:302. doi: 10.3389/fnbeh.2015.00302

PubMed Abstract | CrossRef Full Text | Google Scholar

Garza-Villarreal, E. A., Pando, P., Vuust, P., and Parsons, C. (2017). Music-induced analgesia in chronic pain conditions: a systematic review and meta-analysis. Pain Physician 20, 597–610.

PubMed Abstract | Google Scholar

Gedeon, T., Parry, J., and Völlm, B. (2019). The role of oxytocin in antisocial personality disorders: a systematic review of the literature. Front. Psychiatry 10:76. doi: 10.3389/fpsyt.2019.00076

PubMed Abstract | CrossRef Full Text | Google Scholar

Geng, Y., Zhao, W., Zhou, F., Ma, X., Yao, S., Hurlemann, R., et al. (2018). Oxytocin enhancement of emotional empathy: generalization across cultures and effects on amygdala activity. Front. Neurosci. 12:512. doi: 10.3389/fnins.2018.00512

PubMed Abstract | CrossRef Full Text | Google Scholar

Giovanna, G., Damiani, S., Fusar-Poli, L., Rocchetti, M., Brondino, N., de Cagna, F., et al. (2020). Intranasal oxytocin as a potential therapeutic strategy in post-traumatic stress disorder: a systematic review. Psychoneuroendocrinology 115:104605. doi: 10.1016/j.psyneuen.2020.104605

PubMed Abstract | CrossRef Full Text | Google Scholar

Gold, B. P., Mas-Herrero, E., Zeighami, Y., Benovoy, M., Dagher, A., and Zatorre, R. J. (2019). Musical reward prediction errors engage the nucleus accumbens and motivate learning. Proc. Natl. Acad. Sci. U S A 116, 3310–3315. doi: 10.1073/pnas.1809855116

PubMed Abstract | CrossRef Full Text | Google Scholar

Gordon, I., Zagoory-Sharon, O., Leckman, J. F., and Feldman, R. (2010). Oxytocin and the development of parenting in humans. Biol. Psych. 68, 377–382. doi: 10.1016/j.biopsych.2010.02.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Granot, R., Frankel, Y., Gritsenko, V., Lerer, E., Gritsenko, I., Bachner-Melman, R., et al. (2007). Provisional evidence that the arginine vasopressin 1a receptor gene is associated with musical memory. Evol. Hum. Behav. 28, 313–318. doi: 10.1016/j.evolhumbehav.2007.05.003

CrossRef Full Text | Google Scholar

Granot, R. Y., Uzefovsky, F., Bogopolsky, H., and Ebstein, R. P. (2013). Effects of arginine vasopressin on musical working memory. Front. Psychol. 4:712. doi: 10.3389/fpsyg.2013.00712

PubMed Abstract | CrossRef Full Text | Google Scholar

Grape, C., Sandgren, M., Hansson, L. O., Ericson, M., and Theorell, T. (2003). Does singing promote well-being? An empirical study of professional and amateur singers during a singing lesson. Integr. Phys. Behav. Sci. 38, 65–74. doi: 10.1007/bf02734261

PubMed Abstract | CrossRef Full Text | Google Scholar

Grinevich, V., Knobloch-Bollmann, H. S., Eliava, M., Busnelli, M., and Chini, B. (2016). Assembling the puzzle: pathways of oxytocin signaling in the brain. Biol. Psychiatry 79, 155–164. doi: 10.1016/j.biopsych.2015.04.013

PubMed Abstract | CrossRef Full Text | Google Scholar

Grinevich, V., and Neumann, I. D. (2020). Brain oxytocin: how puzzle stones from animal studies translate into psychiatry. Mol. Psychiatry doi: 10.1038/s41380-020-0802-9 [Epub ahead of print].

PubMed Abstract | CrossRef Full Text | Google Scholar

Grinevich, V., and Stoop, R. (2018). Interplay between oxytocin and sensory systems in the orchestration of socio-emotional behaviors. Neuron 99, 887–904. doi: 10.1016/j.neuron.2018.07.016

PubMed Abstract | CrossRef Full Text | Google Scholar

Groussard, M., Chan, T. G., Coppalle, R., and Patel, H. (2019). Preservation of musical memory throughout the progression of Alzheimer’s Disease? Toward a reconciliation of theoretical, clinical and neuroimaging evidence. Alzheimers Dis. 68, 857–883. doi: 10.3233/jad-180474

PubMed Abstract | CrossRef Full Text | Google Scholar

Guastella, A. J., and MacLeod, C. (2012). A critical review of the influence of oxytocin nasal spray on social cognition in humans: evidence and future directions. Horm. Behav. 61, 410–418. doi: 10.1016/j.yhbeh.2012.01.002

PubMed Abstract | CrossRef Full Text | Google Scholar

Guhn, M., Emerson, S. D., and Gouzouasis, P. (2020). A population-level analysis of associations between school music participation and academic achievement. J. Educat. Psychol. 112, 308–328. doi: 10.1037/edu0000376

CrossRef Full Text | Google Scholar

Gujing, L., Hui, H., Xin, L., Lirong, Z., Yutong, Y., Guofeng, Y., et al. (2019). Increased insular connectivity and enhanced empathic ability associated with dance/music training. Neural Plast. 2019:9693109. doi: 10.1155/2019/9693109

PubMed Abstract | CrossRef Full Text | Google Scholar

Habibi, A., Damasio, A., Ilari, B., Elliott Sachs, M., and Damasio, H. (2018). Music training and child development: a review of recent findings from a longitudinal study. Ann. N Y Acad. Sci. doi: 10.1111/nyas.13606 [Epub ahead of print].

PubMed Abstract | CrossRef Full Text | Google Scholar

Harvey, A. R. (2017). Music, Evolution and the Harmony of Souls. Oxford: Oxford University Press.

Google Scholar

Harvey, A. R. (2018). Music and the meeting of human minds. Front. Psychol. 9:762. doi: 10.3389/fpsyg.2018.00762

PubMed Abstract | CrossRef Full Text | Google Scholar

Hasan, M. T., Althammer, F., Silva da Gouveia, M., Goyon, S., Eliava, M., Lefevre, A., et al. (2019). A fear memory engram and its plasticity in the hypothalamic oxytocin system. Neuron 103, 133.e8–146.e8. doi: 10.1016/j.neuron.2019.04.029

PubMed Abstract | CrossRef Full Text | Google Scholar

Havranek, T., Zatkova, M., Lestanova, Z., Bacova, Z., Mravec, B., Hodosy, J., et al. (2015). Intracerebroventricular oxytocin administration in rats enhances object recognition and increases expression of neurotrophins, microtubule-associated protein 2 and synapsin I. J. Neurosci. Res. 93, 893–901. doi: 10.1002/jnr.23559

PubMed Abstract | CrossRef Full Text | Google Scholar

Herdener, M., Esposito, F., di Salle, F., Boller, C., Hilti, C. C., Habermeyer, B., et al. (2010). Musical training induces functional plasticity in human hippocampus. J. Neurosci. 30, 1377–1384. doi: 10.1523/jneurosci.4513-09.2010

PubMed Abstract | CrossRef Full Text | Google Scholar

Hilfiger, L., Zhao, Q., Kerspern, D., Inquimbert, P., Andry, V., Goumon, Y., et al. (2020). A nonpeptide oxytocin receptor agonist for a durable relief of inflammatory pain. Sci. Rep. 10:3017. doi: 10.1038/s41598-020-59929-w

PubMed Abstract | CrossRef Full Text | Google Scholar

Horgusluoglu-Moloch, E., Risacher, S. L., Crane, P. K., Hibar, D., Thompson, P. M., Saykin, A. J., et al. (2019). Genome-wide association analysis of hippocampal volume identifies enrichment of neurogenesis-related pathways. Sci. Rep. 9:14498. doi: 10.1038/s41598-019-50507-3

PubMed Abstract | CrossRef Full Text | Google Scholar

Hsiao, Y.-H., Hung, H.-C., Chen, S.-H., and Gean, P.-W. (2014). Social interaction rescues memory deficit in an animal model of Alzheimer’s Disease by increasing BDNF-dependent hippocampal neurogenesis. J. Neurosci. 34, 16207–16219. doi: 10.1523/JNEUROSCI.0747-14.2014

PubMed Abstract | CrossRef Full Text | Google Scholar

Hu, J., Wang, Z., Feng, X., Long, C., and Schiller, D. (2019). Post-retrieval oxytocin facilitates next day extinction of threat memory in humans. Psychopharmacology 236, 293–301. doi: 10.1007/s00213-018-5074-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Huber, D., Veinante, P., and Stoop, R. (2005). Vasopressin and oxytocin excite distinct neuronal populations in the central amygdala. Science 308, 245–248. doi: 10.1126/science.1105636

PubMed Abstract | CrossRef Full Text | Google Scholar

Hurlemann, R., Patin, A., Onur, O. A., Cohen, M. X., Baumgartner, T., Metzler, S., et al. (2010). Oxytocin enhances amygdala-dependent, socially reinforced learning and emotional empathy in humans. J. Neurosci. 30, 4999–5007. doi: 10.1523/JNEUROSCI.5538-09.2010

PubMed Abstract | CrossRef Full Text | Google Scholar

Israel, S., Lerer, E., Shalev, I., Uzefovsky, F., Reibold, M., Bachner-Melman, R., et al. (2008). Molecular genetic studies of the arginine vasopressin 1a receptor (AVPR1a) and the oxytocin receptor (OXTR) in human behaviour: from autism to altruism with some notes in between. Prog. Brain Res. 170, 435–449. doi: 10.1016/S0079-6123(08)00434-2

PubMed Abstract | CrossRef Full Text | Google Scholar

Jirikowski, G. F. (2019). Diversity of central oxytocinergic projections. Cell Tissue Res. 375, 41–48. doi: 10.1007/s00441-018-2960-5

PubMed Abstract | CrossRef Full Text | Google Scholar

Jirikowski, G. F., Ochs, S. D., and Caldwell, J. D. (2018). Oxytocin and steroid actions. Curr. Top. Behav. Neurosci. 35, 77–95. doi: 10.1007/7854_2017_9

PubMed Abstract | CrossRef Full Text | Google Scholar

Johnson, J. K., Louhivuori, J., and Silijander, E. (2017). Comparison of well-being of older adult choir singers and the general population in Finland: a case-control study. Music Sci. 21, 178–194. doi: 10.1177/1029864916644486

PubMed Abstract | CrossRef Full Text | Google Scholar

Johnson, J. K., Stewart, A. L., Acree, M., Nápoles, A. M., Flatt, J. D., Max, W. B., et al. (2020). A community choir intervention to promote well-being among diverse older adults: results from the community of voices trial. J. Gerontol. B Psychol. Sci. Soc. Sci. 75, 549–559. doi: 10.1093/geronb/gby132

PubMed Abstract | CrossRef Full Text | Google Scholar

Jurek, B., and Neumann, I. D. (2018). The oxytocin receptor: from intracellular signaling to behavior. Physiol. Rev. 98, 1805–1908. doi: 10.1152/physrev.00031.2017

PubMed Abstract | CrossRef Full Text | Google Scholar

Kang, J., Scholp, A., and Jiang, J. J. (2018). A review of the physiological effects and mechanisms of singing. J. Voice 32, 390–395. doi: 10.1016/j.jvoice.2017.07.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Karpati, F. J., Giacosa, C., Foster, N. E., Penhune, V. B., and Hyde, K. L. (2016). Sensorimotor integration is enhanced in dancers and musicians. Exp. Brain Res. 234, 893–903. doi: 10.1007/s00221-015-4524-1

PubMed Abstract | CrossRef Full Text | Google Scholar

Kattenstroth, J. C., Kalisch, T., Holt, S., Tegenthoff, M., and Dinse, H. R. (2013). Six months of dance intervention enhances postural, sensorimotor and cognitive performance in elderly without affecting cardio-respiratory functions. Front. Aging Neurosci. 5:5. doi: 10.3389/fnagi.2013.00005

PubMed Abstract | CrossRef Full Text | Google Scholar

Kattenstroth, J. C., Kolankowska, I., Kalisch, T., and Dinse, H. R. (2010). Superior sensory, motor and cognitive performance in elderly individuals with multi-year dancing activities. Front. Aging Neurosci. 2:31. doi: 10.3389/fnagi.2010.00031

PubMed Abstract | CrossRef Full Text | Google Scholar

Keech, B., Crowe, S., and Hocking, D. R. (2018). Intranasal oxytocin, social cognition and neurodevelopmental disorders: a meta-analysis. Psychoneuroendocrinology 87, 9–19. doi: 10.1016/j.psyneuen.2017.09.022

PubMed Abstract | CrossRef Full Text | Google Scholar

Keeler, J. R., Roth, E. A., Neuser, B. L., Spitsbergen, J. M., Waters, D. J., and Vianney, J. M. (2015). The neurochemistry and social flow of singing: bonding and oxytocin. Front. Hum. Neurosci. 9:518. doi: 10.3389/fnhum.2015.00518

PubMed Abstract | CrossRef Full Text | Google Scholar

Kempermann, G., Gage, F. H., Aigner, L., Song, H., Curtis, M. A., Thuret, S., et al. (2018). Human adult neurogenesis: evidence and remaining questions. Cell Stem Cell 23, 25–30. doi: 10.1016/j.stem.2018.04.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Kerr, F., Wiechula, R., Feo, R., Schultz, T., and Kitson, A. (2019). Neurophysiology of human touch and eye gaze in therapeutic relationships and healing: a scoping review. JBI Database System Rev. Implement. Rep. 17, 209–247. doi: 10.11124/JBISRIR-2017-003549

PubMed Abstract | CrossRef Full Text | Google Scholar

Khalfa, S., Bella, S. D., Roy, M., Peretz, I., and Lupien, S. J. (2003). Effects of relaxing music on salivary cortisol level after psychological stress. Ann. N Y Acad. Sci. 999, 374–376. doi: 10.1196/annals.1284.045

PubMed Abstract | CrossRef Full Text | Google Scholar

Khant, T., Chang, L. J., Park, S. Q., Heinzle, J., and Haynes, J.-D. (2012). Connectivity-based parcellation of the human orbitofrontal cortex. J. Neurosci. 32, 6240–6250. doi: 10.1523/JNEUROSCI.0257-12.2012

PubMed Abstract | CrossRef Full Text | Google Scholar

Kim, S., Fonagy, P., Koos, O., Dorsett, K., and Strathearn, L. (2014). Maternal oxytocin response predicts mother-to-infant gaze. Brain Res. 1580, 133–142. doi: 10.1016/j.brainres.2013.10.050

PubMed Abstract | CrossRef Full Text | Google Scholar

Kirsch, P., Esslinger, C., Chen, Q., Mier, D., Lis, S., Siddhanti, S., et al. (2005). Oxytocin modulates neural circuitry for social cognition and fear in humans. J. Neurosci. 25, 11489–11493. doi: 10.1523/JNEUROSCI.3984-05.2005

PubMed Abstract | CrossRef Full Text | Google Scholar

Kirschner, S., and Tomasello, M. (2009). Joint drumming: social context facilitates synchronization in preschool children. J. Exp. Child. Psychol. 102, 299–314. doi: 10.1016/j.jecp.2008.07.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Koch, S. B. J., van Zuiden, M., Nawijn, L., Frijling, J. L., Veltman, D. J., and Olff, M. (2019). Effects of intranasal oxytocin on distraction as emotion regulation strategy in patients with post-traumatic stress disorder. Eur. Neuropsychopharmacol. 29, 266–277. doi: 10.1016/j.euroneuro.2018.12.002

PubMed Abstract | CrossRef Full Text | Google Scholar

Koelsch, S. (2014). Brain correlates of music-evoked emotions. Nat. Rev. Neurosci. 15, 170–180. doi: 10.1038/nrn3666

PubMed Abstract | CrossRef Full Text | Google Scholar

Koelsch, S. (2018). Investigating the neural encoding of emotion with music. Neuron 98, 1075–1079. doi: 10.1016/j.neuron.2018.04.029

PubMed Abstract | CrossRef Full Text | Google Scholar

Koelsch, S., Fritz, T., v Cramon, D. Y., Müller, K., and Friederici, A. D. (2006). Investigating emotion with music: an fMRI study. Hum. Brain Mapp. 27, 239–250. doi: 10.1002/hbm.20180

PubMed Abstract | CrossRef Full Text | Google Scholar

Koelsch, S., and Jäncke, L. (2015). Music and the heart. Eur. Heart J. 36, 3043–3049. doi: 10.1093/eurheartj/ehv430

PubMed Abstract | CrossRef Full Text | Google Scholar

Korn, C. W., Prehn, K., Park, S. Q., Walter, H., and Heekeren, H. R. (2012). Positively based processing of self-relevant social feedback. J. Neurosci. 32, 16832–16844. doi: 10.1523/JNEUROSCI.3016-12.2012

CrossRef Full Text | Google Scholar

Kosfeld, M., Heinrichs, M., Zak, P. J., Fischbacher, U., and Fehr, E. (2005). Oxytocin increases trust in humans. Nature 435, 673–676. doi: 10.1038/nature03701

PubMed Abstract | CrossRef Full Text | Google Scholar

Kreutz, G. (2014). Singing and social bonding. Music Med. 6, 51–60.

Google Scholar

Kreutz, G., Bongard, S., Rohrmann, S., Hodapp, V., and Grebe, D. (2004). Effects of choir singing or listening on secretory immunoglobulin A, cortisol, and emotional state. J. Behav. Med. 27, 623–635. doi: 10.1007/s10865-004-0006-9

PubMed Abstract | CrossRef Full Text | Google Scholar

Kuhn, D. (2002). The effects of active and passive participation in musical activity on the immune system as measured by salivary immunoglobulin A (SIgA). J. Music Ther. 39, 30–39. doi: 10.1093/jmt/39.1.30

PubMed Abstract | CrossRef Full Text | Google Scholar

Kuhn, H. G., Toda, T., and Gage, F. H. (2018). Adult hippocampal neurogenesis: a coming of age story. J. Neurosci. 38, 10401–10410. doi: 10.1523/JNEUROSCI.2144-18.2018

PubMed Abstract | CrossRef Full Text | Google Scholar

Kumar, J., Iwabuchi, S. J., Völlm, B. A., and Palaniyappan, L. (2020). Oxytocin modulates the effective connectivity between the precuneus and the dorsolateral prefrontal cortex. Eur. Arch. Psychiatry Clin. Neurosci. 270, 567–576. doi: 10.1007/s00406-019-00989-z

PubMed Abstract | CrossRef Full Text | Google Scholar

Laland, K., Wilkins, C., and Clayton, N. (2016). The evolution of dance. Curr. Biol. 26, R5–R9. doi: 10.1016/j.cub.2015.11.031

PubMed Abstract | CrossRef Full Text | Google Scholar

Lambert, B., Declerck, C. H., Boone, C., and Parizel, P. M. (2017). A functional MRI study on how oxytocin affects decision making in social dilemmas: cooperate as long as it pays off, aggress only when you think you can win. Horm. Behav. 94, 145–152. doi: 10.1016/j.yhbeh.2017.06.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Lancaster, K., Goldbeck, L., Pournajafi-Nazarloo, H., Connelly, J. J., Carter, C. S., and Morris, J. P. (2018). The role of endogenous oxytocin in anxiolysis: structural and functional correlates. Biol. Psychiatry Cogn. Neurosci. Neuroimaging 3, 618–625. doi: 10.1016/j.bpsc.2017.10.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Latt, H. M., Matsushita, H., Morino, M., Koga, Y., Michiue, H., Nishiki, T., et al. (2018). Oxytocin inhibits corticosterone-induced apoptosis in primary hippocampal neurons. Neuroscience 379, 383–389. doi: 10.1016/j.neuroscience.2018.03.025

PubMed Abstract | CrossRef Full Text | Google Scholar

Lawson, E. A., Olszewski, P. K., Weller, A., and Blevins, J. E. (2019). The role of oxytocin in regulation of appetitive behaviour, body weight and glucose homeostasis. J. Neuroendocrinol. 28:e12805. doi: 10.1111/jne.12805

PubMed Abstract | CrossRef Full Text | Google Scholar

Lee, J. H., Lee, S., and Kim, J. H. (2017). Amygdala circuits for fear memory: a key role for dopamine regulation. Neuroscientist 23, 542–553. doi: 10.1177/1073858416679936

PubMed Abstract | CrossRef Full Text | Google Scholar

Lee, S. Y., Park, S. H., Chung, C., Kim, J. J., Choi, S. Y., and Han, J. S. (2015). Oxytocin protects hippocampal memory and plasticity from uncontrollable stress. Sci. Rep. 5:18540. doi: 10.1038/srep18540

PubMed Abstract | CrossRef Full Text | Google Scholar

Leng, G., and Ludwig, M. (2016). Intranasal oxytocin: myths and delusions. Biol. Psychiatry 79, 243–250. doi: 10.1016/j.biopsych.2015.05.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Leppanen, J., Ng, K. W., Tchanturia, K., and Treasure, J. (2017). Meta-analysis of the effects of intranasal oxytocin on interpretation and expression of emotions. Neurosci. Biobehav. Rev. 78, 125–144. doi: 10.1016/j.neubiorev.2017.04.010

PubMed Abstract | CrossRef Full Text | Google Scholar

Levitin, D. J., Grahn, J. A., and London, J. (2018). The psychology of music: rhythm and movement. Annu. Rev. Psychol. 69, 51–75. doi: 10.1146/annurev-psych-122216-011740

PubMed Abstract | CrossRef Full Text | Google Scholar

Li, T., Chen, X., Mascaro, J., Haroon, E., and Rilling, J. K. (2017a). Intranasal oxytocin, but not vasopressin, augments neural responses to toddlers in human fathers. Horm. Behav. 93, 193–202. doi: 10.1016/j.yhbeh.2017.01.006

PubMed Abstract | CrossRef Full Text | Google Scholar

Li, T., Wang, P., Wang, S. C., and Wang, Y. F. (2017b). Approaches mediating oxytocin regulation of the immune system. Front. Immunol. 7:693. doi: 10.3389/fimmu.2016.00693

PubMed Abstract | CrossRef Full Text | Google Scholar

Li, Q., Zhang, B., Cao, H., Liu, W., Guo, F., Shen, F., et al. (2020). Oxytocin exerts antidepressant-like effect by potentiating dopaminergic synaptic transmission in the mPFC. Neuropharmacology 162:107836. doi: 10.1016/j.neuropharm.2019.107836

PubMed Abstract | CrossRef Full Text | Google Scholar

Licht, T., Kreisel, T., Biala, Y., Mohan, S., Yaari, Y., Anisimov, A., et al. (2020). Age-dependent remarkable regenerative potential of the dentate gyrus provided by intrinsic stem cells. J. Neurosci. 40, 974–995. doi: 10.1523/jneurosci.1010-19.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Lima, S. M. A., and Gomes-Leal, W. (2019). Neurogenesis in the hippocampus of adult humans: controversy “fixed” at last. Neural Regen. Res. 14, 1917–1918. doi: 10.4103/1673-5374.259616

PubMed Abstract | CrossRef Full Text | Google Scholar

Limb, C. J., and Braun, A. R. (2008). Neural substrates of spontaneous musical performance: an FMRI study of jazz improvisation. PLoS One 3:e1679. doi: 10.1371/journal.pone.0001679

PubMed Abstract | CrossRef Full Text | Google Scholar

Lin, Y. T., and Hsu, K. S. (2018). Oxytocin receptor signaling in the hippocampus: role in regulating neuronal excitability, network oscillatory activity, synaptic plasticity and social memory. Prog. Neurobiol. 171, 1–14. doi: 10.1016/j.pneurobio.2018.10.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Lin, Y. T., Chen, C. C., Huang, C. C., Nishimori, K., and Hsu, K. S. (2017). Oxytocin stimulates hippocampal neurogenesis via oxytocin receptor expressed in CA3 pyramidal neurons. Nat. Commun. 8:537. doi: 10.1038/s41467-017-00675-5

PubMed Abstract | CrossRef Full Text | Google Scholar

Lin, Y.-T., Hsieh, T.-Y., Tsai, T.-C., Chen, C.-C., Huang, C.-C., and Hsu, K.-S. (2018). Conditional deletion of hippocampal CA2/CA3a oxytocin receptors impairs the persistence of long-term social recognition memory in mice. J. Neurosci. 38, 1218–1231. doi: 10.1523/jneurosci.1896-17.2017

PubMed Abstract | CrossRef Full Text | Google Scholar

Lischke, A., Gamer, M., Berger, C., Grossmann, A., Hauenstein, K., Heinrichs, M., et al. (2012). Oxytocin increases amygdala reactivity to threatening scenes in females. Psychoneuroendocrinology 37, 1431–1438. doi: 10.1016/j.psyneuen.2012.01.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, S., Chow, H. M., Xu, Y., Erkkinen, M. G., Swett, K. E., Eagle, M. W., et al. (2012). Neural correlates of lyrical improvisation: an FMRI study of freestyle rap. Sci. Rep. 2:834. doi: 10.1038/srep00834

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, X., Kanduri, C., Oikkonen, J., Karma, K., Raijas, P., Ukkola-Vuoti, L., et al. (2016). Detecting signatures of positive selection associated with musical aptitude in the human genome. Sci. Rep. 6:21198. doi: 10.1038/srep21198

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, Y., Li, S., Lin, W., Li, W., Yan, X., Wang, X., et al. (2019). Oxytocin modulates social value representations in the amygdala. Nat. Neurosci. 22, 633–641. doi: 10.1038/s41593-019-0351-1

PubMed Abstract | CrossRef Full Text | Google Scholar

Liu, P. Z., and Nusslock, R. (2018). Exercise-mediated neurogenesis in the hippocampus via BDNF. Front. Neurosci. 12:52. doi: 10.3389/fnins.2018.00052

PubMed Abstract | CrossRef Full Text | Google Scholar

Lopatina, O. L., Komleva, Y. K., Gorina, Y. V., Olovyannikova, R. Y., Trufanova, L. V., Hashimoto, T., et al. (2018). Oxytocin and excitation/inhibition balance in social recognition. Neuropeptides 72, 1–11. doi: 10.1016/j.npep.2018.09.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Lucassen, P. J., Fitzsimonsm, C. P., Salta, E., and Maletic-Savatic, M. (2020). Adult neurogenesis, human after all (again): classic, optimized and future approaches. Behav. Brain Res. 381:112458. doi: 10.1016/j.bbr.2019.112458

PubMed Abstract | CrossRef Full Text | Google Scholar

Luo, S., Zhu, Y., Fan, L., Gao, D., and Han, S. (2020). Resting-state brain network properties mediate the association between the oxytocin receptor gene and interdependence. Soc. Neurosci. 15, 296–310. doi: 10.1080/17470919.2020.1714718

PubMed Abstract | CrossRef Full Text | Google Scholar

Malik, A. I., Zai, C. C., Abu, Z., Nowrouzi, B., and Beitchman, J. H. (2012). The role of oxytocin and oxytocin receptor gene variants in childhood-onset aggression. Genes Brain Behav. 11, 545–551. doi: 10.1111/j.1601-183x.2012.00776.x

PubMed Abstract | CrossRef Full Text | Google Scholar

Maniezzi, C., Talpo, F., Spaiardi, P., Toselli, M., and Biella, G. (2019). Oxytocin increases phasic and tonic GABAergic transmission in CA1 region of mouse hippocampus. Front. Cell. Neurosci. 13:178. doi: 10.3389/fncel.2019.00178

PubMed Abstract | CrossRef Full Text | Google Scholar

Mansens, D., Deeg, D. J. H., and Comijs, H. C. (2018). The association between singing and/or playing a musical instrument and cognitive functions in older adults. Aging Ment. Health 22, 964–971. doi: 10.1080/13607863.2017.1328481

PubMed Abstract | CrossRef Full Text | Google Scholar

Mariath, L. M., Silva, A. M. D., Kowalski, T. W., Gattino, G. S., Araujo, G. A., Figueiredo, F. G., et al. (2017). Music genetics research: association with musicality of a polymorphism in the AVPRA gene. Genet. Mol. Biol. 40, 421–429. doi: 10.1590/1678-4685-gmb-2016-0021

PubMed Abstract | CrossRef Full Text | Google Scholar

Masis-Calvo, M., Schmidtner, A. K., de Moura Oliveira, V. E., Grossmann, C. P., de Jong, T. R., and Neumann, I. D. (2018). Animal models of social stress: the dark side of social interactions. Stress 21, 417–432. doi: 10.1080/10253890.2018.1462327

PubMed Abstract | CrossRef Full Text | Google Scholar

Matthiesen, A. S., Ransjo-Arvidson, A. B., Nissen, E., and Uvnas- Moberg, K. (2001). Postpartum maternal oxytocin release by newborns: effects of infant hand massage and sucking. Birth 28, 13–19. doi: 10.1046/j.1523-536x.2001.00013.x

PubMed Abstract | CrossRef Full Text | Google Scholar

McEwen, B. S. (1999). Stress and hippocampal plasticity. Annu. Rev. Neurosci. 12, 105–122.

Google Scholar

Meguro, Y., Miyano, K., Hirayama, S., Yoshida, Y., Ishibashi, N., Ogino, T., et al. (2018). Neuropeptide oxytocin enhances μ opioid receptor signaling as a positive allosteric modulator. J. Pharmacol. Sci. 137, 67–75. doi: 10.1016/j.jphs.2018.04.002

PubMed Abstract | CrossRef Full Text | Google Scholar

Mehr, S. A., Singh, M., York, H., Glowacki, L., and Krasnow, M. M. (2018). Form and function in human song. Curr. Biol. 28, 356.e5–368.e5. doi: 10.1016/j.cub.2017.12.042

PubMed Abstract | CrossRef Full Text | Google Scholar

Menon, V., and Levitin, D. J. (2005). The rewards of music listening: response and physiological connectivity of the mesolimbic system. NeuroImage 28, 175–184. doi: 10.1016/j.neuroimage.2005.05.053

PubMed Abstract | CrossRef Full Text | Google Scholar

Millan, M. J. (2002). Descending control of pain. Prog. Neurobiol. 66, 355–474. doi: 10.1016/s0301-0082(02)00009-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Miranda, M., Morici, J. F., Zanoni, M. B., and Bekinschtein, P. (2019). Brain-derived neurotrophic factor: a key molecule for memory in the healthy and the pathological brain. Front. Cell. Neurosci. 13:363. doi: 10.3389/fncel.2019.00363

PubMed Abstract | CrossRef Full Text | Google Scholar

Mithen, S. (2005). The Singing Neanderthals: The Origins of Music, Language, Mind and Body. London: Orion.

Google Scholar

Mitre, M., Marlin, B. J., Schiavo, J. K., Morina, E., Norden, S. E., Hackett, T. A., et al. (2016). A distributed network for social cognition enriched for oxytocin receptors. J. Neurosci. 36, 2517–2535. doi: 10.1523/jneurosci.2409-15.2016

PubMed Abstract | CrossRef Full Text | Google Scholar

Mitre, M., Minder, J., Morina, E. X., Chao, M. V., and Froemke, R. C. (2018). Oxytocin modulation of neural circuits. Curr. Top. Behav. Neurosci. 35, 31–53. doi: 10.1007/7854_2017_7

PubMed Abstract | CrossRef Full Text | Google Scholar

Mitterschiffthaler, M. T., Fu, C. H. Y., Dalton, J. A., Andrew, C. M., and Williams, S. C. R. (2007). A functional MRI study of happy and sad affective states induced by classical music. Hum. Brain Mapp. 28, 1150–1162. doi: 10.1002/hbm.20337

PubMed Abstract | CrossRef Full Text | Google Scholar

Moaddab, M., Hyland, B. I., and Brown, C. H. (2015). Oxytocin excites nucleus accumbens shell neurons in vivo. Mol. Cell. Neurosci. 68, 323–330. doi: 10.1016/j.mcn.2015.08.013

PubMed Abstract | CrossRef Full Text | Google Scholar

Moons, W. G., Way, B. M., and Taylor, S. E. (2014). Oxytocin and vasopressin receptor polymorphisms interact with circulating neuropeptides to predict human emotional reactions to stress. Emotion 14, 562–572. doi: 10.1037/a0035503

PubMed Abstract | CrossRef Full Text | Google Scholar

Moreno-Jiménez, E. P., Flor-García, M., Terreros-Roncal, J., Rábano, A., Cafini, F., Pallas-Bazarra, N., et al. (2019). Adult hippocampal neurogenesis is abundant in neurologically healthy subjects and drops sharply in patients with Alzheimer’s disease. Nat. Med. 25, 554–560. doi: 10.1038/s41591-019-0375-9

PubMed Abstract | CrossRef Full Text | Google Scholar

Morley, I. (2013). The Prehistory of Music: Human Evolution, Archaeology and the Origins of Musicality. Oxford: Oxford University Press.

Google Scholar

Morley, A. P., Narayanan, M., Mines, R., Molokhia, A., Baxter, S., Craig, G., et al. (2012). AVPR1A and SLC6A4 polymorphisms in choral singers and non-musicians: a gene association study. PLoS One 7:e31763. doi: 10.1371/journal.pone.0031763

PubMed Abstract | CrossRef Full Text | Google Scholar

Moss, H., Lynch, J., and O’Donoghue, J. (2018). Exploring the perceived health benefits of singing in a choir: an international cross-sectional mixed-methods study. Perspect. Public Health 138, 160–168. doi: 10.1177/1757913917739652

PubMed Abstract | CrossRef Full Text | Google Scholar

Mueller, K., Fritz, T., Mildner, T., Richter, M., Schulze, K., Lepsien, J., et al. (2015). Investigating the dynamics of the brain response to music: a central role of the ventral striatum/nucleus accumbens. NeuroImage 116, 68–79. doi: 10.1016/j.neuroimage.2015.05.006

PubMed Abstract | CrossRef Full Text | Google Scholar

Narendran, R., Tollefson, S., Fasenmyer, K., Paris, J., Himes, M. L., Lopresti, B., et al. (2019). Decreased nociceptin receptors are related to resilience and recovery in college women who have experienced sexual violence: therapeutic implications for posttraumatic stress disorder. Biol. Psych. 85, 1056–1064. doi: 10.1016/j.biopsych.2019.02.017

PubMed Abstract | CrossRef Full Text | Google Scholar

Ne’eman, R., Perach-Barzilay, N., Fischer-Shofty, M., Atias, A., and Shamay-Tsoory, S. G. (2016). Intranasal administration of oxytocin increases human aggressive behaviour. Horm. Behav. 80, 125–131. doi: 10.1016/j.yhbeh.2016.01.015

PubMed Abstract | CrossRef Full Text | Google Scholar

Neumann, I. D., and Landgraf, R. (2012). Balance of brain oxytocin and vasopressin: implications for anxiety, depression, and social behaviors. Trends Neurosci. 35, 649–659. doi: 10.1016/j.tins.2012.08.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Neumann, I. D., and Landgraf, R. (2019). Tracking oxytocin functions in the rodent brain during the last 30 years: from push-pull perfusion to chemogenetic silencing. J. Neuroendocrinol. 31:e12695. doi: 10.1111/jne.12695

PubMed Abstract | CrossRef Full Text | Google Scholar

Neumann, I. D., and Slattery, D. A. (2016). Oxytocin in general anxiety and social fear: a translational approach. Biol. Psych. 79, 213–221. doi: 10.1016/j.biopsych.2015.06.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Nilsson, U. (2009). Soothing music can increase oxytocin levels during bed rest after open-heart surgery: a randomised control trial. J. Clin. Nurs. 18, 2153–2161. doi: 10.1111/j.1365-2702.2008.02718.x

PubMed Abstract | CrossRef Full Text | Google Scholar

Nilsson, S., Kokinsky, E., Nilsson, U., Sidenvall, B., and Enskär, K. (2009). School-aged children’s experiences of postoperative music medicine on pain, distress and anxiety. Paediatr. Anaesth. 19, 1184–1190. doi: 10.1111/j.1460-9592.2009.03180.x

PubMed Abstract | CrossRef Full Text | Google Scholar

Nishina, K., Takagishi, H., Inoue-Murayama, M., Takahashi, H., and Yamagishi, T. (2015). Polymorphism of the oxytocin receptor gene modulates behavioral and attitudinal trust among men but not women. PLoS One 10:e0137089. doi: 10.1371/journal.pone.0137089

PubMed Abstract | CrossRef Full Text | Google Scholar

Nishina, K., Takagishi, H., Takahashi, H., Sakagami, M., and Inoue-Murayama, M. (2019). Association of polymorphism of arginine-vasopressin receptor 1A (AVPR1a) gene with trust and reciprocity. Front. Hum. Neurosci. 13:230. doi: 10.3389/fnhum.2019.00230

PubMed Abstract | CrossRef Full Text | Google Scholar

Nilsson, U., Unosson, M., and Rawal, N. (2005). Stress reduction and analgesia in patients exposed to calming music postoperatively: a randomized controlled trial. Eur. J. Anaesthesiol. 22, 96–102. doi: 10.1017/s0265021505000189

PubMed Abstract | CrossRef Full Text | Google Scholar

Norman-Haignere, S., Kanwisher, N. G., and McDermott, J. H. (2015). Distinct cortical pathways for music and speech revealed by hypothesis-free voxel decomposition. Neuron 88, 1281–1296. doi: 10.1016/j.neuron.2015.11.035

PubMed Abstract | CrossRef Full Text | Google Scholar

Nozaradan, S., Peretz, I., Missal, M., and Mouraux, A. (2011). Tagging the neuronal entrainment to beat and meter. J. Neurosci. 31, 10234–10240. doi: 10.1523/jneurosci.0411-11.2011

PubMed Abstract | CrossRef Full Text | Google Scholar

O’Doherty, J. P. (2004). Reward representations and reward-related learning in the human brain: insights from neuroimaging. Curr. Opin. Neurobiol. 14, 769–776. doi: 10.1016/j.conb.2004.10.016

PubMed Abstract | CrossRef Full Text | Google Scholar

Oikkonen, J., Kuusi, T., Peltonen, P., Raijas, P., Ukkola-Vuoti, L., Karma, K., et al. (2016). Creative activities in music—a genome-wide linkage analysis. PLoS One 11:e0148679. doi: 10.1371/journal.pone.0148679

PubMed Abstract | CrossRef Full Text | Google Scholar

Onaka, T., and Takayanagi, Y. (2019). Role of oxytocin in the control of stress and food intake. J. Neuroendocrinol. 31:e12700. doi: 10.1111/jne.12700

PubMed Abstract | CrossRef Full Text | Google Scholar

Ooishi, Y., Mukai, H., Watanabe, K., Kawato, S., and Kashino, M. (2017). Increase in salivary oxytocin and decrease in salivary cortisol after listening to relaxing slow-tempo and exciting fast-tempo music. PLoS One 12:e0189075. doi: 10.1371/journal.pone.0189075

PubMed Abstract | CrossRef Full Text | Google Scholar

Opendak, M., Offit, L., Monari, P., Schoenfeld, T. J., Sonti, A. N., Cameron, H. A., et al. (2016). Lasting adaptations in social behavior produced by social disruption and inhibition of adult neurogenesis. J. Neurosci. 36, 7027–7038. doi: 10.1523/jneurosci.4435-15.2016

PubMed Abstract | CrossRef Full Text | Google Scholar

Ouanes, S., and Popp, J. (2019). High cortisol and the risk of dementia and Alzheimer’s disease: a review of the literature. Front. Aging Neurosci. 11:43. doi: 10.3389/fnagi.2019.00043

PubMed Abstract | CrossRef Full Text | Google Scholar

Patel, A. D. (2008). Music, Language and the Brain. New York, NY: Oxford University Press.

Google Scholar

Pearce, E., Launay, J., and Dunbar, R. I. M. (2015). The ice-breaker effect: singing mediates fast social bonding. R. Soc. Open Sci. 2:150221. doi: 10.1098/rsos.150221

PubMed Abstract | CrossRef Full Text | Google Scholar

Pearce, E., Wlodarski, R., Machin, A., and Dunbar, R. I. M. (2017). Variation in the β-endorphin, oxytocin and dopamine receptor genes is associated with different dimensions of human sociality. Proc. Natl. Acad. Sci. U S A 114, 5300–5305. doi: 10.1073/pnas.1700712114

PubMed Abstract | CrossRef Full Text | Google Scholar

Pekarek, B. T., Hunt, P. J., and Arenkiel, B. R. (2020). Oxytocin and sensory network plasticity. Front. Neurosci. 14:30. doi: 10.3389/fnins.2020.00030

PubMed Abstract | CrossRef Full Text | Google Scholar

Peled-Avron, L., Abu-Akel, A., and Shamay-Tsoory, S. (2020). Exogenous effects of oxytocin in five psychiatric disorders: a systematic review, meta-analyses and a personalized approach through the lens of the social salience hypothesis. Neurosci. Biobehav. Rev. 114, 70–95. doi: 10.1016/j.neubiorev.2020.04.023

PubMed Abstract | CrossRef Full Text | Google Scholar

Pereira, A. P. S., Marinho, V., Gupta, D., Magalhães, F., Ayres, C., and Teixeira, S. (2019). Music therapy and dance as gait rehabilitation in patients with Parkinson disease: a review of evidence. J. Geriatr. Psychiatry Neurol. 32, 49–56. doi: 10.1177/0891988718819858

PubMed Abstract | CrossRef Full Text | Google Scholar

Pereira, C. S., Teixeira, J., Figueiredo, P., Xavier, J., Castro, S. L., and Brattico, E. (2011). Music and emotions in the brain: familiarity matters. PLoS One 6:e27241. doi: 10.1371/journal.pone.0027241

PubMed Abstract | CrossRef Full Text | Google Scholar

Peretz, I., Vuvan, D., Lagrois, M-E., and Armony, J. L. (2015). Neural overlap in processing music and speech. Philos. Trans. R. Soc. Lond. B Biol. Sci. 370:20140090. doi: 10.1098/rstb.2014.0090

PubMed Abstract | CrossRef Full Text | Google Scholar

Perkins, R., Yorke, S., and Fancourt, D. (2018). How group singing facilitates recovery from the symptoms of postnatal depression: a comparative qualitative study. BMC Psychol. 6:41. doi: 10.1186/s40359-018-0253-0

PubMed Abstract | CrossRef Full Text | Google Scholar

Petrik, D., and Encinas, J. M. (2019). Perspective: of mice and men - how widespread is adult neurogenesis? Front. Neurosci. 13:923. doi: 10.3389/fnins.2019.00923

PubMed Abstract | CrossRef Full Text | Google Scholar

Phillips-Silver, J., and Trainor, L. J. (2007). Hearing what the body feels: auditory encoding of rhythmic movement. Cognition 105, 533–546. doi: 10.1016/j.cognition.2006.11.006

PubMed Abstract | CrossRef Full Text | Google Scholar

Plasencia, G., Luedicke, J. M., Nazarloo, H. P., Carter, C. S., and Ebner, N. C. (2019). Plasma oxytocin and vasopressin levels in young and older men and women: functional relationships with attachment and cognition. Psychoneuroendocrinology 110:104419. doi: 10.1016/j.psyneuen.2019.104419

PubMed Abstract | CrossRef Full Text | Google Scholar

Poore, H. E., and Waldman, I. D. (2020). The association of oxytocin receptor gene (OXTR) polymorphisms with antisocial behavior: a meta-analysis. Behav. Genet. 50:174. doi: 10.1007/s10519-020-09998-4

PubMed Abstract | CrossRef Full Text | Google Scholar

Preckel, K., Scheele, D., Kendrick, K. M., Maier, W., and Hurlemann, R. (2014). Oxytocin facilitates social approach behaviour in women. Front. Behav. Neurosci. 8:191. doi: 10.3389/fnbeh.2014.00191

CrossRef Full Text | Google Scholar

Pulli, K., Karma, K., Norio, R., Sistonen, P., Göring, H. H., and Järvelä, I. (2008). Genome-wide linkage scan for loci of musical aptitude in Finnish families: evidence for a major locus at 4q22. J. Med. Genet. 45, 451–456. doi: 10.1136/jmg.2007.056366

PubMed Abstract | CrossRef Full Text | Google Scholar

Quintana, D. S., Rokicki, J., van der Meer, D., Alnæs, D., Kaufmann, T., Córdova-Palomera, A., et al. (2019). Oxytocin pathway gene networks in the human brain. Nat. Commun. 10:668. doi: 10.1038/s41467-019-08503-8

PubMed Abstract | CrossRef Full Text | Google Scholar

Quintin, E. M. (2019). Music-evoked reward and emotion: relative strengths and response to intervention of people with ASD. Front. Neural Circuits 13:49. doi: 10.3389/fncir.2019.00049

PubMed Abstract | CrossRef Full Text | Google Scholar

Raam, T. (2020). Oxytocin-sensitive neurons in prefrontal cortex gate social recognition memory. J. Neurosci. 40, 1194–1196. doi: 10.1523/jneurosci.1348-19.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Reddish, P., Fischer, R., and Bulbalia, L. (2013). Let’s dance together: synchrony, shared intentionality and cooperation. PLoS One 8:e71182. doi: 10.1371/journal.pone.0071182

PubMed Abstract | CrossRef Full Text | Google Scholar

Reiss, A. B., Glass, D. S., Lam, E., Glass, A. D., De Leon, J., and Kasselman, L. J. (2019). Oxytocin: potential to mitigate cardiovascular risk. Peptides 117:170089. doi: 10.1016/j.peptides.2019.05.001

PubMed Abstract | CrossRef Full Text | Google Scholar

Richter, J., and Ostovar, R. (2016). “It don‘t mean a thing if it ain’t got that swing”- an alternative concept for understanding the evolution of dance and music in human beings. Front. Hum. Neurosci. 10:485. doi: 10.3389/fnhum.2016.00485

PubMed Abstract | CrossRef Full Text | Google Scholar

Rilling, J. K., Chen, X., Chen, X., and Haroon, E. (2018). Intranasal oxytocin modulates neural functional connectivity during human social interaction. Am. J. Primatol. 80:e22740. doi: 10.1002/ajp.22740

PubMed Abstract | CrossRef Full Text | Google Scholar

Rilling, J. K., DeMarco, A. C., Hackett, P. D., Chen, X., Gautam, P., Stair, S., et al. (2014). Sex differences in the neural and behavioral response to intranasal oxytocin and vasopressin during human social interaction. Psychoneuroendocrinology 39, 237–248. doi: 10.1016/j.psyneuen.2013.09.022

PubMed Abstract | CrossRef Full Text | Google Scholar

Rilling, J. K., DeMarco, A. C., Hackett, P. D., Thompson, R., Ditzen, B., Patel, R., et al. (2012). Effects of intranasal oxytocin and vasopressin on cooperative behavior and associated brain activity in men. Psychoneuroendocrinology 37, 447–461. doi: 10.1016/j.psyneuen.2011.07.013

PubMed Abstract | CrossRef Full Text | Google Scholar

Rilling, J. K., King-Cass, B., and Sanfey, A. G. (2008). The neurobiology of social decision-making. Curr. Opin. Neurobiol. 18, 159–165. doi: 10.1016/j.conb.2008.06.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Rilling, J. K., and Sanfey, A. G. (2011). The neuroscience of social decision-making. Annu. Rev. Psychol. 62, 23–48. doi: 10.1146/annurev.psych.121208.131647

PubMed Abstract | CrossRef Full Text | Google Scholar

Rodrigues, S. M., Saslow, L. R., Garcia, N., John, O. P., and Keltner, D. (2009). Oxytocin receptor genetic variation relates to empathy and stress reactivity in humans. Proc. Natl. Acad. Sci. U S A 106, 21437–21441. doi: 10.1073/pnas.0909579106

PubMed Abstract | CrossRef Full Text | Google Scholar

Rogers, C. N., Ross, A. P., Sahu, S. P., Siegel, E. R., Dooyema, J. M., Cree, M. A., et al. (2018). Oxytocin- and arginine vasopressin-containing fibers in the cortex of humans, chimpanzees and rhesus macaques. Am. J. Primatol. 80:e22875. doi: 10.1002/ajp.22875

PubMed Abstract | CrossRef Full Text | Google Scholar

Rushworth, M. F. S., Noonan, M. P., Boorman, E. D., Walton, M. E., and Behrens, T. E. (2011). Frontal cortex and reward-guided learning and decision-making. Neuron 70, 1054–1069. doi: 10.1016/j.neuron.2011.05.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Sabino, A. D. V., Chagas, M. H. N., and Osório, F. L. (2020). Acute effects of oxytocin in music performance anxiety: a crossover, randomized, placebo-controlled trial. Psychopharmacology 237, 1757–1767. doi: 10.1007/s00213-020-05493-0

PubMed Abstract | CrossRef Full Text | Google Scholar

Sachs, M. E., Habibi, A., and Damasio, A. (2018). Reflections on music, affect and sociality. Prog. Brain Res. 237, 153–172. doi: 10.1016/bs.pbr.2018.03.009

PubMed Abstract | CrossRef Full Text | Google Scholar

Sachs, M. E., Habibi, A., Damasio, A., and Kaplan, J. T. (2019). Dynamic intersubject neural synchronization reflects affective responses to sad music. NeuroImage 218:116512. doi: 10.1016/j.neuroimage.2019.116512

PubMed Abstract | CrossRef Full Text | Google Scholar

Salighedar, R., Erfanparast, A., Tamaddonfard, E., and Soltanalinejad, F. (2019). Medial prefrontal cortex oxytocin-opioid receptors interaction in spatial memory processing in rats. Physiol. Behav. 209:112599. doi: 10.1016/j.physbeh.2019.112599

PubMed Abstract | CrossRef Full Text | Google Scholar

Salimpoor, V. N., Benovoy, M., Larcher, K., Dagher, A., and Zatorre, R. J. (2011). Anatomically distinct dopamine release during anticipation and experience of peak emotion in music. Nat. Neurosci. 14, 257–264. doi: 10.1038/nn.2726

PubMed Abstract | CrossRef Full Text | Google Scholar

Sánchez-Vidaña, D. I., Chan, N. M., Chan, A. H., Hui, K. K., Lee, S., Chan, H. Y., et al. (2016). Repeated treatment with oxytocin promotes hippocampal cell proliferation, dendritic maturation and affects socio-emotional behavior. Neuroscience 333, 65–77. doi: 10.1016/j.neuroscience.2016.07.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Särkämö, T., and Sihvonen, A. J. (2018). Golden oldies and silver brains: deficits, preservation, learning and rehabilitation effects of music in ageing-related neurological disorders. Cortex 109, 104–123. doi: 10.1016/j.cortex.2018.08.034

PubMed Abstract | CrossRef Full Text | Google Scholar

Schellenberg, E. G., Corrigall, K. A., Dys, S. P., and Malti, T. (2015). Group music training and children’s prosocial skills. PLoS One 10:e0141449. doi: 10.1371/journal.pone.0141449

PubMed Abstract | CrossRef Full Text | Google Scholar

Schiller, B., Domes, G., and Heinrichs, M. (2020). Oxytocin changes behavior and spatio-temporal brain dynamics underlying inter-group conflict in humans. Eur. Neuropsychopharmacol. 31, 119–130. doi: 10.1016/j.euroneuro.2019.12.109

PubMed Abstract | CrossRef Full Text | Google Scholar

Schladt, T. M., Nordmann, G. C., Emilius, R., Kudielka, B. M., de Jong, T. R., and Neumann, I. D. (2017). Choir versus solo singing: effects on mood and salivary oxytocin and cortisol concentrations. Front. Hum. Neurosci. 11:430. doi: 10.3389/fnhum.2017.00430

PubMed Abstract | CrossRef Full Text | Google Scholar

Schlaug, G., Forgeard, M., Zhu, L., Norton, A., Norton, A., and Winner, E. (2009). Training-induced neuroplasticity in young children. The neurosciences and music III - disorders and plasticity. Ann. N Y Acad. Sci. 1169, 205–208. doi: 10.1111/j.1749-6632.2009.04842.x

CrossRef Full Text | Google Scholar

Schneider, I., Schmitgen, M. M., Boll, S., Roth, C., Nees, F., Usai, K., et al. (2020). Oxytocin modulates intrinsic neural activity in patients with chronic low back pain. Eur. J. Pain 24, 945–955. doi: 10.1002/ejp.1543

PubMed Abstract | CrossRef Full Text | Google Scholar

Seltzer, L. J., Ziegler, T. E., and Pollak, S. D. (2010). Social vocalizations can release oxytocin in humans. Proc. Biol. Sci. 277, 2661–2666. doi: 10.1098/rspb.2010.0567

PubMed Abstract | CrossRef Full Text | Google Scholar

Shalvi, S., and De Dreu, C. K. (2014). Oxytocin promotes group-serving dishonesty. Proc. Natl. Acad. Sci. U S A 111, 5503–5507. doi: 10.1073/pnas.1400724111

PubMed Abstract | CrossRef Full Text | Google Scholar

Shamay-Tsoory, S. G., and Abu-Akel, A. (2016). The social salience hypothesis of oxytocin. Biol. Psych. 79, 194–202. doi: 10.1016/j.biopsych.2015.07.020

PubMed Abstract | CrossRef Full Text | Google Scholar

Shany, O., Singer, N., Gold, B. P., Jacoby, N., Tarrasch, R., Hendler, T., et al. (2019). Surprise-related activation in the nucleus accumbens interacts with music-induced pleasantness. Soc. Cogn. Affect. Neurosci. 14, 459–470. doi: 10.1093/scan/nsz019

PubMed Abstract | CrossRef Full Text | Google Scholar

Sheldon, S., Fenerci, C., and Gurguryan, L. (2019). A neurocognitive perspective on the forms and functions of autobiographical memory retrieval. Front. Syst. Neurosci. 13:4. doi: 10.3389/fnsys.2019.00004

PubMed Abstract | CrossRef Full Text | Google Scholar

Sicorello, M., Dieckmann, L., Moser, D., Lux, V., Luhmann, M., Schlotz, W., et al. (2020). Oxytocin and the stress buffering effect of social company: a genetic study in daily life. Soc. Cogn. Affect. Neurosci. 15, 293–301. doi: 10.1093/scan/nsaa034

PubMed Abstract | CrossRef Full Text | Google Scholar

Slattery, C. F., Agustus, J. L., Paterson, R. W., McCallion, O., Foulkes, A. J. M., Macpherson, K., et al. (2019). The functional neuroanatomy of musical memory in Alzheimer’s disease. Cortex 115, 357–370. doi: 10.1016/j.cortex.2019.02.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Sleiman, S. F., Henry, J., Al-Haddad, R., El Hayek, L., Abou Haidar, E., Stringer, T., et al. (2016). Exercise promotes the expression of brain derived neurotrophic factor (BDNF) through the action of the ketone body β-hydroxybutyrate. Elife 5:e15092. doi: 10.7554/elife.15092

PubMed Abstract | CrossRef Full Text | Google Scholar

Smith, C. J. W., Di Benedictis, B. T., and Veenema, A. H. (2019). Comparing vasopressin and oxytocin fiber and receptor density patterns in the social behavior neural network: implications for cross-system signaling. Front. Neuroendocrinol. 53:100737. doi: 10.1016/j.yfrne.2019.02.001

PubMed Abstract | CrossRef Full Text | Google Scholar

Snyder, J. S. (2019). Recalibrating the relevance of adult neurogenesis. Trends Neurosci. 42, 164–178. doi: 10.1016/j.tins.2018.12.001

PubMed Abstract | CrossRef Full Text | Google Scholar

Snyder, J. S., and Drew, M. R. (2020). Functional neurogenesis over the years. Behav. Brain Res. 382:112470. doi: 10.1016/j.bbr.2020.112470

PubMed Abstract | CrossRef Full Text | Google Scholar

Song, Z., and Albers, H. E. (2018). Cross-talk among oxytocin and arginine-vasopressin receptors: relevance for basic and clinical studies of the brain and periphery. Front. Neuroendocrinol. 51, 14–24. doi: 10.1016/j.yfrne.2017.10.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Sorrells, S. F., Paredes, M. F., Cebrian-Silla, A., Sandoval, K., Qi, D., Kelley, K. W., et al. (2018). Human hippocampal neurogenesis drops sharply in children to undetectable levels in adults. Nature 555, 377–381. doi: 10.1038/nature25975

PubMed Abstract | CrossRef Full Text | Google Scholar

Spalding, K. L., Bergmann, O., Alkass, K., Bernard, S., Salehpour, M., Huttner, H. B., et al. (2013). Dynamics of hippocampal neurogenesis in adult humans. Cell 153, 1219–1227. doi: 10.1016/j.cell.2013.05.002

PubMed Abstract | CrossRef Full Text | Google Scholar

Stephan, J. S., and Sleiman, S. F. (2019). Exercise factors as potential mediators of cognitive rehabilitation following traumatic brain injury. Curr. Opin. Neurol. 32, 808–814. doi: 10.1097/wco.0000000000000754

PubMed Abstract | CrossRef Full Text | Google Scholar

Stewart, N. A. J., and Lonsdale, A. J. (2016). It’s better together: the psychological benefits of singing in a choir. Psychol. Music 44, 1240–1254. doi: 10.1177/0305735615624976

CrossRef Full Text | Google Scholar

Stillman, C. M., Cohen, J., Lehman, M. E., and Erickson, K. I. (2016). Mediators of physical activity on neurocognitive function: a review at multiple levels of analysis. Front. Hum. Neurosci. 10:626. doi: 10.3389/fnhum.2016.00626

PubMed Abstract | CrossRef Full Text | Google Scholar

Strathearn, L., Fonagy, P., Amico, J., and Montague, P. R. (2009). Adult attachment predicts maternal brain and oxytocin response to infant cues. Neuropsychopharmacol. 34, 2655–2666. doi: 10.1038/npp.2009.103

PubMed Abstract | CrossRef Full Text | Google Scholar

Strathearn, L., Iyengar, U., Fonagy, P., and Kim, S. (2012). Maternal oxytocin response during mother-infant interaction: associations with adult temperament. Horm. Behav. 61, 429–435. doi: 10.1016/j.yhbeh.2012.01.014

PubMed Abstract | CrossRef Full Text | Google Scholar

Strauss, G. P., Chapman, H. C., Keller, W. R., Koenig, J. I., Gold, J. M., Carpenter, W. T., et al. (2019). Endogenous oxytocin levels are associated with impaired social cognition and neurocognition in schizophrenia. J. Psychiatr. Res. 112, 38–43. doi: 10.1016/j.jpsychires.2019.02.017

PubMed Abstract | CrossRef Full Text | Google Scholar

Suzuki, S., Fujisawa, T. X., Sakakibara, N., Fujioka, T., Takiguchi, S., and Tomoda, A. (2020). Development of social attention and oxytocin levels in maltreated children. Sci. Rep. 10:7407. doi: 10.1038/s41598-020-64297-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Tabak, B. A., Teed, A. R., Castle, E., Dutcher, J. M., Meyer, M. L., Bryan, R., et al. (2019). Null results of oxytocin and vasopressin administration across a range of social cognitive and behavioral paradigms: evidence from a randomized controlled trial. Psychoneuroendocrinology 107, 124–132. doi: 10.1016/j.psyneuen.2019.04.019

PubMed Abstract | CrossRef Full Text | Google Scholar

Tachibana, A., Noah, J. A., Ono, Y., Taguchi, D., and Ueda, S. (2019). Prefrontal activation related to spontaneous creativity with rock music improvisation: a functional near-infrared spectroscopy study. Sci. Rep. 9:16044. doi: 10.1038/s41598-019-52348-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Talamini, F., Altoè, G., Carretti, B., and Grassi, M. (2017). Musicians have better memory than nonmusicians: a meta-analysis. PLoS One 2:e0186773. doi: 10.1371/journal.pone.0186773

PubMed Abstract | CrossRef Full Text | Google Scholar

Tan, Y., Singhal, S. M., Harden, S. W., Cahill, K. M., Nguyen, D. M., Colon-Perez, L. M., et al. (2019). Oxytocin receptors are expressed by glutamatergic prefrontal cortical neurons that selectively modulate social recognition. J. Neurosci. 39, 3249–3263. doi: 10.1523/jneurosci.2944-18.2019

PubMed Abstract | CrossRef Full Text | Google Scholar

Tarr, B., Launay, J., Cohen, E., and Dunbar, R. (2015). Synchrony and exertion during dance independently raise pain threshold and encourage social bonding. Biol. Lett. 11:20150767. doi: 10.1098/rsbl.2015.0767

PubMed Abstract | CrossRef Full Text | Google Scholar

Tarr, B., Launay, J., and Dunbar, R. I. M. (2014). Music and social bonding: “self-other” merging and neurohormonal mechanisms. Front. Psychol. 5:1096. doi: 10.3389/fpsyg.2014.01096

PubMed Abstract | CrossRef Full Text | Google Scholar

Tarr, B., Launay, J., and Dunbar, R. I. (2016). Silent disco: dancing in synchrony leads to elevated pain thresholds and social closeness. Evol. Hum. Behav. 37, 343–349. doi: 10.1016/j.evolhumbehav.2016.02.004

PubMed Abstract | CrossRef Full Text | Google Scholar

Taylor, S. E., Saphire-Bernstein, S., and Seeman, T. E. (2010). Are plasma oxytocin in women and plasma vasopressin in men biomarkers of distressed pair-bond relationships? Psychol. Sci. 21, 3–7. doi: 10.1177/0956797609356507

PubMed Abstract | CrossRef Full Text | Google Scholar

Ten Velden, F. S., Daughters, K., and De Dreu, C. K. W. (2017). Oxytocin promotes intuitive rather than deliberated cooperation with the in-group. Horm. Behav. 92, 164–171. doi: 10.1016/j.yhbeh.2016.06.005

PubMed Abstract | CrossRef Full Text | Google Scholar

Tillman, R., Gordon, I., Naples, A., Rolison, M., Leckman, J. F., Feldman, R., et al. (2019). Oxytocin enhances the neural efficiency of social perception. Front. Hum. Neurosci. 13:71. doi: 10.3389/fnhum.2019.00071

PubMed Abstract | CrossRef Full Text | Google Scholar

Tirko, N. N., Eyring, K. W., Carcea, I., Mitre, M., Chao, M. V., Froemke, R. C., et al. (2018). Oxytocin transforms firing mode of CA2 hippocampal neurons. Neuron 100, 593.e3–608.e3. doi: 10.1016/j.neuron.2018.09.008

PubMed Abstract | CrossRef Full Text | Google Scholar

Tobin, M. K., Musaraca, K., Disouky, A., Shetti, A., Bheri, A., Honer, W. G., et al. (2019). Human hippocampal neurogenesis persists in aged adults and Alzheimer’s disease patients. Cell Stem Cell 24, 974.e3–982.e3. doi: 10.1016/j.stem.2019.05.003

PubMed Abstract | CrossRef Full Text | Google Scholar

Toda, T., and Gage, F. H. (2018). Review: adult neurogenesis contributes to hippocampal plasticity. Cell Tissue Res. 373, 693–709. doi: 10.1007/s00441-017-2735-4

PubMed Abstract | CrossRef Full Text | Google Scholar

Tollefson, S., Himes, M., and Narendran, R. (2017). Imaging corticotropin-releasing-factor and nociceptin in addiction and PTSD models. Int. Rev. Psych. 29, 567–579. doi: 10.1080/09540261.2017.1404445

PubMed Abstract | CrossRef Full Text | Google Scholar

Tomizawa, K., Iga, N., Lu, Y.-F., Moriwaki, A., Matsushita, M., Li, S.-T., et al. (2003). Oxytocin improves long-lasting spatial memory during motherhood through MAP kinase cascade. Nat. Neurosci. 6, 384–390. doi: 10.1038/nn1023

PubMed Abstract | CrossRef Full Text | Google Scholar

Tops, M., Huffmeijer, R., Linting, M., Grewen, K. M., Light, K. C., Koole, S. L., et al. (2013). The role of oxytocin in familiarization-habituation responses to social novelty. Front. Psychol. 4:761. doi: 10.3389/fpsyg.2013.00761

PubMed Abstract | CrossRef Full Text | Google Scholar

Torrisi, S. A., Leggio, G. M., Drago, F., and Salamone, S. (2019). Therapeutic challenges of post-traumatic stress disorder: focus on the dopaminergic system. Front. Pharmacol. 10:404. doi: 10.3389/fphar.2019.00404

PubMed Abstract | CrossRef Full Text | Google Scholar

Tost, H., Kolachana, B., Hakimi, S., Lemaitre, H., Verchinski, B. A., Mattay, V. S., et al. (2010). A common allele in the oxytocin receptor gene (OXTR) impacts prosocial temperament and human hypothalamic-limbic structure and function. Proc. Natl. Acad. Sci. U S A 107, 13936–13941. doi: 10.1073/pnas.1003296107

PubMed Abstract | CrossRef Full Text | Google Scholar

Triana-Del Río, R., van den Burg, E., Stoop, R., and Hegoburu, C. (2019). Acute and long-lasting effects of oxytocin in cortico-limbic circuits: consequences for fear recall and extinction. Psychopharmacology 236, 339–354. doi: 10.1007/s00213-018-5030-5

PubMed Abstract | CrossRef Full Text | Google Scholar

Ukkola, L. T., Onkamo, P., Raijas, P., Karma, K., and Järvelä, I. (2009). Musical aptitude is associated with AVPR1A-haplotypes. PLoS One 4:e5534. doi: 10.1371/journal.pone.0005534

PubMed Abstract | CrossRef Full Text | Google Scholar

Ukkola-Vuoti, L., Oikkonen, J., Onkamo, P., Karma, K., Raijas, P., and Järvelä, I. (2011). Association of the arginine vasopressin receptor 1A (AVPR1A) haplotypes with listening to music. J. Hum. Genet. 56, 324–329. doi: 10.1038/jhg.2011.13

PubMed Abstract | CrossRef Full Text | Google Scholar

Ulmer-Yaniv, A., Avitsur, R., Kanat-Maymon, Y., Schneiderman, I., Zagoory-Sharon, O., and Feldman, R. (2016). Affiliation, reward and immune biomarkers coalesce to support social synchrony during periods of bond formation in humans. Brain Behav. Immun. 56, 130–139. doi: 10.1016/j.bbi.2016.02.017

PubMed Abstract | CrossRef Full Text | Google Scholar

Valstad, M., Alvares, G. A., Egknud, M., Matziorinis, A. M., Andreassen, O. A., Westlye, L. T., et al. (2017). The correlation between central and peripheral oxytocin concentrations: a systematic review and meta-analysis. Neurosci. Biobehav. Rev. 78, 117–124. doi: 10.1016/j.neubiorev.2017.04.017

PubMed Abstract | CrossRef Full Text | Google Scholar

Van den Burg, E. H., Stindl, J., Grund, T., Neumann, I. D., and Strauss, O. (2015). Oxytocin stimulates extracellular Ca2+ influx through TRPV2 channels in hypothalamic neurons to exert its anxiolytic effects. Neuropsychopharmacol. 40, 2938–2947. doi: 10.1038/npp.2015.147

PubMed Abstract | CrossRef Full Text | Google Scholar

Van der Heijden, M. J. E., Araghi, S., van Dijk, M., Jeekel, J., and Hunink, M. G. M. (2015). The effects of perioperative music interventions in pediatric surgery: a systematic review and meta-analysis of randomized controlled trials. PLoS One 10:e0133608. doi: 10.1371/journal.pone.0133608

PubMed Abstract | CrossRef Full Text | Google Scholar

Veenema, Y. H., and Neumann, I. D. (2018). Central vasopressin and oxytocin release: regulation of complex social behaviours. Prog. Brain Res. 170, 261–276. doi: 10.1016/S0079-6123(08)00422-6

PubMed Abstract | CrossRef Full Text | Google Scholar

Völlm, B. A., Taylor, A. N., Richardson, P., Corcoran, R., Stirling, J., McKie, S., et al. (2006). Neuronal correlates of theory of mind and empathy: a functional magnetic resonance imaging study in a nonverbal task. NeuroImage 29, 90–98. doi: 10.1016/j.neuroimage.2005.07.022

PubMed Abstract | CrossRef Full Text | Google Scholar

Wagner, U., and Echterhoff, G. (2018). When does oxytocin affect human memory encoding? The role of social context and individual attachment style. Front. Hum. Neurosci. 12:349. doi: 10.3389/fnhum.2018.00349

PubMed Abstract | CrossRef Full Text | Google Scholar

Walker, E., Ploubidis, G., and Fancourt, D. (2019). Social engagement and loneliness are differentially associated with neuro-immune markers in older age: time-varying associations from the English Longitudinal Study of Ageing. Brain Behav. Immun. 82, 224–229. doi: 10.1016/j.bbi.2019.08.189

PubMed Abstract | CrossRef Full Text | Google Scholar

Wang, P., Wang, S. C., Yang, H., Lv, C., Jia, S., Liu, X., et al. (2019). Therapeutic ootential of oxytocin in atherosclerotic cardiovascular disease: mechanisms and signaling pathways. Front. Neurosci. 13:454. doi: 10.3389/fnins.2019.00454

PubMed Abstract | CrossRef Full Text | Google Scholar

Wang, D., Yan, X., Li, M., and Ma, Y. (2017). Neural substrates underlying the effects of oxytocin: a quantitative meta-analysis of pharmaco-imaging studies. Soc. Cogn. Affect. Neurosci. 12, 1565–1573. doi: 10.1093/scan/nsx085

PubMed Abstract | CrossRef Full Text | Google Scholar

Weinstein, D., Launay, J., Pearce, E., Dunbar, R. I., and Stewart, L. (2016). Group music performance causes elevated pain thresholds and social bonding in small and large groups of singers. Evol. Hum. Behav. 37, 152–158. doi: 10.1016/j.evolhumbehav.2015.10.002

PubMed Abstract | CrossRef Full Text | Google Scholar

Weisman, O., Zagoory-Sharon, O., Schneiderman, I., Gordon, I., and Feldman, R. (2013). Plasma oxytocin distributions in a large cohort of women and men and their gender specific associations with anxiety. Psychoneuroendocrinology 38, 694–701. doi: 10.1016/j.psyneuen.2012.08.011

PubMed Abstract | CrossRef Full Text | Google Scholar

Wu, H., Feng, C., Lu, X., Liu, X., and Liu, Q. (2020). Oxytocin effects on the resting-state mentalizing brain network. Brain Imaging Behav. doi: 10.1007/s11682-019-00205-5 [Epub ahead of print].

PubMed Abstract | CrossRef Full Text | Google Scholar

Xu, L., Becker, B., Luo, R., Zheng, X., Zhao, W., Zhang, Q., et al. (2020). Oxytocin amplifies sex differences in human mate choice. Psychoneuroendocrinology 112:104483. doi: 10.1016/j.psyneuen.2019.104483

PubMed Abstract | CrossRef Full Text | Google Scholar

Yamasue, H., and Domes, G. (2018). Oxytocin and autism spectrum disorders. Curr. Top. Behav. Neurosci. 35, 449–465. doi: 10.1007/7854_2017_24

PubMed Abstract | CrossRef Full Text | Google Scholar

Yoon, S., and Kim, Y. K. (2020). The role of the oxytocin system in anxiety disorders. Adv. Exp. Med. Biol. 1191, 103–120. doi: 10.1007/978-981-32-9705-0_7

PubMed Abstract | CrossRef Full Text | Google Scholar

Yoshida, M., Takayanagi, Y., Inoue, K., Kimura, T., Young, L. J., Onaka, T., et al. (2009). Evidence that oxytocin exerts anxiolytic effects via oxytocin receptor expressed in serotonergic neurons in mice. J. Neurosci. 29, 2259–2271. doi: 10.1523/jneurosci.5593-08.2009

PubMed Abstract | CrossRef Full Text | Google Scholar

Young, W. S., and Song, J. (2020). Characterization of oxytocin receptor expression within various neuronal populations of the mouse dorsal hippocampus. Front. Mol. Neurosci. 13:40. doi: 10.3389/fnmol.2020.00040

PubMed Abstract | CrossRef Full Text | Google Scholar

Yuhi, T., Kyuta, H., Mori, H. A., Murakami, C., Furuhara, K., Okuno, M., et al. (2017). Salivary oxytocin concentration changes during a group drumming intervention for maltreated school children. Brain Sci. 7:E152. doi: 10.3390/brainsci7110152

PubMed Abstract | CrossRef Full Text | Google Scholar

Zak, P. J., and Berroza, J. A. (2013). The neurobiology of collective action. Front. Neurosci. 7:211. doi: 10.3389/fnins.2013.00211

PubMed Abstract | CrossRef Full Text | Google Scholar

Zatorre, R. J., and Salimpoor, V. N. (2013). From perception to pleasure: music and its neural substrates. Proc. Natl. Acad. Sci. U S A 110, 10430–10437. doi: 10.1073/pnas.1301228110

PubMed Abstract | CrossRef Full Text | Google Scholar

Zhang, K., Li, G., Wang, L., Cao, C., Fang, R., Luo, S., et al. (2019). An epistasis between dopaminergic and oxytocinergic systems confers risk of post-traumatic stress disorder in a traumatized Chinese cohort. Sci. Rep. 9:19252. doi: 10.1038/s41598-019-55936-8

PubMed Abstract | CrossRef Full Text | Google Scholar

Zhang, S., Liu, D., Ye, D., Li, H., and Chen, F. (2017). Can music-based movement therapy improve motor dysfunction in patients with Parkinson’s disease? Systematic review and meta-analysis. Neurol. Sci. 38, 1629–1636. doi: 10.1007/s10072-017-3020-8

PubMed Abstract | CrossRef Full Text | Google Scholar

Zhang, T. Y., Shahrokh, D., Hellstrom, I. C., Wen, X., Diorio, J., Breuillaud, L., et al. (2020). Brain-derived neurotrophic factor in the nucleus accumbens mediates individual differences in behavioral responses to a natural, social reward. Mol. Neurobiol. 57, 290–301. doi: 10.1007/s12035-019-01699-2

PubMed Abstract | CrossRef Full Text | Google Scholar

Zilidou, V. I., Frantzidis, C. A., Romanopoulou, E. D., Paraskevopoulos, E., Douka, S., and Bamidis, P. D. (2018). Functional re-organization of cortical networks of senior citizens after a 24-week traditional dance program. Front. Aging Neurosci. 10:422. doi: 10.3389/fnagi.2018.00422

PubMed Abstract | CrossRef Full Text | Google Scholar

Keywords: oxytocin, music, dance, reward, empathy, trust, therapy

Citation: Harvey AR (2020) Links Between the Neurobiology of Oxytocin and Human Musicality. Front. Hum. Neurosci. 14:350. doi: 10.3389/fnhum.2020.00350

Received: 01 May 2020; Accepted: 04 August 2020;
Published: 26 August 2020.

Edited by:

Franco Delogu, Lawrence Technological University, United States

Reviewed by:

Kai Lutz, Center for Neurology and Rehabilitation (CERENEO), Switzerland
Madhavi Rangaswamy, Christ University, India

Copyright © 2020 Harvey. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Alan R. Harvey, alan.harvey@uwa.edu.au