Abstract
It has been suggested that a key function of music during its development and spread amongst human populations was its capacity to create and strengthen social bonds amongst interacting group members. However, the mechanisms by which this occurs have not been fully discussed. In this paper we review evidence supporting two thus far independently investigated mechanisms for this social bonding effect: self-other merging as a consequence of inter-personal synchrony, and the release of endorphins during exertive rhythmic activities including musical interaction. In general, self-other merging has been experimentally investigated using dyads, which provide limited insight into large-scale musical activities. Given that music can provide an external rhythmic framework that facilitates synchrony, explanations of social bonding during group musical activities should include reference to endorphins, which are released during synchronized exertive movements. Endorphins (and the endogenous opioid system (EOS) in general) are involved in social bonding across primate species, and are associated with a number of human social behaviors (e.g., laughter, synchronized sports), as well as musical activities (e.g., singing and dancing). Furthermore, passively listening to music engages the EOS, so here we suggest that both self-other merging and the EOS are important in the social bonding effects of music. In order to investigate possible interactions between these two mechanisms, future experiments should recreate ecologically valid examples of musical activities.
INTRODUCTION
Music-making, and movement to music, are activities central to ritual, courtship, identity, and human expression cross-culturally. Based on this ubiquity, it is argued that music has played an important role during our evolutionary history (Cross, 2001; Huron, 2001; McDermott, 2008; Dunbar, 2012b; although see, Pinker, 1997 for an alternative perspective).Whilst sexual selection and courtship are proposed as partial explanations for the widespread appreciation and aptitude for music (Brown, 2000; Merker, 2000; Miller, 2000 for a critique), there are other suggestions regarding its positive role for human societies. In this review we focus on the fact that in almost all cultures globally, and throughout history, music is a social activity (Nettl, 1983, 2000) that involves movement to rhythmic sound and plays a significant role both in creating social bonds (Roederer, 1984; McNeill, 1995; Freeman, 2000; Dunbar, 2004) and indicating coalition strength (Hagen and Bryant, 2003). This effect of musical activity on “social bonding” (the psychological experience of increased social closeness, reflected in prosocial behaviors) may be responsible for the widespread occurrence of musical activities and may have played an important role in the evolution of human sociality (Dunbar, 2012a,b).
While there has been much interest in the relationship between music and social bonding, there is as yet no consensus about the mechanisms by which this might occur. Many aspects of music-making which make people feel socially close are not specific to music-based activities, such as sharing attention with co-actors (e.g., Reddy, 2003), working toward similar goals (e.g., Tomasello et al., 2005), and experiencing a sense of positivity after successful co-engagement (e.g., Isen, 1970). An important feature that distinguishes musical activities from other social behavior is the importance of shared rhythms, and the externalization of predictable rhythms that allow synchronization to occur between two or more people (e.g., Bispham, 2006; Merker et al., 2009). Furthermore, people attribute movement and human agency to musical sound (e.g., Cross, 2001), which influences how synchronization occurs (Launay et al., 2013, 2014) as well as impacting upon affective experience (e.g., Fritz et al., 2013a). Here we focus on two proposed mechanisms of social bonding: self-other merging as a consequence of interpersonal synchrony, and the release of endorphins during synchronized exertive movements. We bring together evidence that both pathways from music-making to social bonding are relevant, highlight connections between the two, and suggest that both should be included in any account of how people form and maintain social bonds through music-making.
Firstly, performing movements simultaneously with someone else, (i.e., synchronizing), is believed to cause some blurring of self and other via neural pathways that code for both action and perception (Overy and Molnar-Szakacs, 2009). Secondly, it has been argued that group music-making leads to social bonding due to the release of neurohormones, specifically oxytocin (e.g., Freeman, 2000; Huron, 2001; Grape et al., 2003). The oxytocin account relies on its action as a social neurohormone in a range of mammals (e.g., Insel, 2010), and the suggestion that music-making (which involves sensory overload, physical activity, strong emotional arousal and social behavior) is particularly conducive to oxytocin release (e.g., Freeman, 2000). Whilst elevated oxytocin levels has been linked to increased trust (Kosfeld et al., 2005; Zak et al., 2005), eye contact (Guastella et al., 2008), face memory (Savaskan et al., 2008), generosity (Zak et al., 2007), empathy and the ability to infer the mental state of others (Domes et al., 2007), the causal link between music-related physical experience and oxytocin described by Freeman is tenuous. Here we review the evidence that the endogenous opioid system (EOS), and particularly endorphins, play a central role in the maintenance of non-sexual, non-kinship social bonds (Machin and Dunbar, 2011) that are characteristic of group musical activities. Given that endorphins are argued to mediate the pleasure experienced when listening to music (e.g., Huron, 2006; Koelsch, 2010) and recent evidence demonstrates that endorphins are released during synchronized and exertive activity (Cohen et al., 2010; Sullivan and Rickers, 2013; Sullivan et al., 2014), we argue that this particular peptide is an important candidate for the neurohormonal underpinnings of social bonding during group musical activities.
To begin, we will explore the evidence linking synchronization and social bonding, and subsequently the particular role of self-other merging, which may occur via shared neural pathways for action and perception. Following this we review evidence of the EOS’s role in social bonding, and discuss the case for this mechanism in musical activities. Finally we highlight the importance of using ecologically valid musical contexts in future investigation into the possible relationship between the two mechanisms that underpin the relationship between music and social bonding.
SYNCHRONIZATION AND SOCIAL BONDING
Synchronization is often cited as an important mechanism by which social bonding can occur (Hove and Risen, 2009; Wiltermuth and Heath, 2009; Valdesolo and Desteno, 2011; Launay et al., 2013). This proposition builds in part on an identified relationship between mimicry (i.e., making a similar movement to another individual) and positive social behavior, such as self-reported rapport between two individuals (e.g., LaFrance and Broadbent, 1976; LaFrance, 1979). Mimicry improves rapport between people (Chartrand and Bargh, 1999; Lakin and Chartrand, 2003), which in turn influences the amount of mimicry that people perform (Van Baaren et al., 2004; Stel et al., 2010), thereby causing a positive feedback loop in which people can become increasingly socially close to one another through making similar movements, and more inclined to continue making similar movements once social closeness is established. Synchrony, like mimicry, involves simultaneous movements with another individual, with the additional element of rhythmically matched timing, which requires the prediction of movements of co-actors. Consequently, synchronization is likely to have similar or more pronounced effects on social bonding than mimicry.
People tend to spontaneously and unintentionally synchronize movements with one another, even to some extent when instructed not to do so (Issartel et al., 2007; Oullier et al., 2008; van Ulzen et al., 2008). Those with pro-social tendencies exhibit more spontaneous synchronization than those with pro-self tendencies (Lumsden et al., 2012), and the desirability of a partner can influence whether synchrony occurs (Miles et al., 2010, 2011), suggesting that this is a social behavior, rather than an automatic motor process. Perception of synchrony is also interpreted as a signal of rapport for both basic sounds (e.g., sound of people walking together: Miles et al., 2009; Lakens and Stel, 2011), and more complex musical stimuli (Hagen and Bryant, 2003).
More importantly, there is evidence that synchronization between people can influence their subsequent positive social feelings toward one another. This has been demonstrated in a number of experimental studies, involving participants tapping synchronously with an experimenter (Hove and Risen, 2009; Valdesolo and Desteno, 2011), walking in time with other people (Wiltermuth and Heath, 2009; Wiltermuth, 2012), dancing together (Reddish et al., 2013), and even when people have no visual access to one another but are synchronizing with the sounds of another person (Kokal et al., 2011; Launay et al., 2014).
The likely importance of social bonding via synchrony in music-based activities draws on the observation that beyond a tendency to synchronize with one another, humans have a culturally ubiquitous aptitude for entrainment to rhythmic beats (Clayton et al., 2005; Brown and Jordania, 2011), particularly those embedded in music (e.g., Demos et al., 2012). However, the source and context associated with those rhythms are paramount. For example, Kirschner and Tomasello (2009) demonstrated that children’s synchronization with a beat is improved in the presence of a person compared to when interacting solely with an isochronously beating drum. This suggests that from a young age, the awareness of agency related to perceived sound (and belief that the sound is produced by the intentional movements of another person) encourages synchronization with that sound, thereby likely influencing the social bonding effects of musical activities. Agent-driven sounds, and the associated perception of movement of another person, engage motor regions in the listener’s brain, potentially resulting in “self-other merging,” which has been argued to arise when individuals experience their movement simultaneously with another’s.
SELF-OTHER MERGING AND SOCIAL BONDING
When moving at the same time as others we experience some co-activation of neural networks that relate to movement of self (as action), and other (as perception; e.g., Overy and Molnar-Szakacs, 2009). There is much recent research investigating the relationship between perception and action (Buccino et al., 2001; Fadiga et al., 2002; Rizzolatti, 2005; Caetano et al., 2007), which has identified “mirror neurons” in macaques (Gallese et al., 1996; Rizzolatti et al., 1996) that selectively respond to the macaque’s own movement and perception of the goal-directed movement of others. While there is no evidence for neurons with equivalent selectivity in humans (Hickok, 2009), this research led to much interest into how perception of goal-directed movement can engage regions of the brain related to making similar movements (Rizzolatti, 2005). Importantly it is now well recognized that perceiving the actions of another person can lead to activation of the same neural motor networks involved in making those actions oneself (e.g., Fadiga et al., 1995).
When our own actions match those of another’s, it is possible that the intrinsic and extrinsic engagement of neural action-perception networks make it difficult to distinguish between self and perceived other, thus creating at least a transient bond between the two (Decety and Sommerville, 2003; Sebanz et al., 2006; Sommerville and Decety, 2006; Knoblich and Sebanz, 2008; Marsh et al., 2009; Overy and Molnar-Szakacs, 2009). A well replicated experimental example of this is the rubber hand illusion (Botvinick and Cohen, 1998). In this paradigm, a participant’s arm is hidden from sight, and a replacement rubber arm is visible where their own arm is expected. While they view the rubber hand being touched with a paintbrush, their own (hidden) hand is simultaneously touched with a paintbrush, with synchronized strokes. This matching of visual and tactile input leads to an increased subjective sense that the rubber hand is part of the participant’s body. The effect disappears when the two inputs are not synchronized. This provides evidence that self-other blurring is possible even with an inanimate object, and some aspects of this are likely to apply to human–human synchronized interaction. Indeed, behavioral synchrony has also been demonstrated to induce common neural signatures between interacting agents (Oullier et al., 2005; Tognoli et al., 2007; Lindenberger et al., 2009; Dumas et al., 2010). However, evidence for common neural signatures during synchronization should be interpreted with caution, as it can only indicate that similar cortical networks are involved in making the same movements for different people.
Researchers who argue that self-other merging is an important part of the bonding effects of synchronization primarily draw support from dyadic experiments in which participants’ actions are perceived to occur at the same time as one another. Theoretically, dyads are capable of achieving synchrony with relative ease simply because there is only one other person to keep track of. As such, synchrony is reasonably attainable, and associated self-other merging (and bonding) effects are likely to be achieved fairly easily.
Musical activities, on the other hand, are not limited to one-on-one interactions, and have historically involved groups (Nettl, 1983, 2000). With large numbers of people, it is difficult to simultaneously observe the movements of all the other participants, making self-other merging a less likely prospect. Rhythm provides an external, predictable scaffolding that can facilitate synchrony with both the music, and by extension, aids synchrony between individuals engaging in the same musical experience. A recent experiment involved people rocking on rocking chairs with one another, while music played in the room or did not (Demos et al., 2012). While self-reported rapport between co-actors correlated with synchronization achieved with the music, rapport did not correlate with synchronization that occurred between co-actors. This implies that externalizing the target of synchrony (e.g., to music) allows bonding with other people present, in the absence of explicit synchrony between those people. This finding has important implications given that group musical activities often involve non-identical movements between people (making self-other merging an unlikely prospect).
Given that the self-other merging account of social bonding relies on simultaneous, similar movements, it is likely that this mechanism does not provide a complete account for the bonding that arises in large group situations. Additional mechanisms need to be considered, in particular mechanisms that underpin the social bonding associated with musical activities. One likely mechanism involves the EOS, and particularly endorphins, which are released through synchronous and exertive activities, and during passive engagement with music, and play a central role in social bonding among primate species (e.g., Keverne et al., 1989).
ENDORPHINS AND SOCIAL BONDING
Investigation into the neuropeptide underpinnings of social bonding have implicated neurohormonal cascades involving oxytocin and vasopressin (e.g., Carter, 1998), dopamine and serotonin (e.g., Depue and Morrone-Strupinsky, 2005), and endorphins released by the EOS (Curley and Keverne, 2005; Dunbar, 2010). Recently, oxytocin has been promoted as the social neurohormone (Bartz et al., 2011; Meyer-Lindenberg et al., 2011), largely due to evidence from pair-bonding and mother-infant bonding (e.g., Atzil et al., 2011; Feldman, 2012). However, despite apparent interactions between opioids (specifically endorphins) in the bonding activity of oxytocin (e.g., Depue and Morrone-Strupinsky, 2005), and evidence of the EOS’s role in primate pair-bonding (Ragen et al., 2013), maternal care (Martel et al., 1993), as well as empirical evidence that increased opioid levels are associated with social grooming and affiliative behaviors in non-sexual, non-kin related conspecifics (Keverne et al., 1989; Schino and Troisi, 1992; Martel et al., 1995), the role of the EOS in social bonding remains relatively underexplored, possibly due to the difficulties in measuring endorphin titres directly (Dearman and Francis, 1983).
The EOS consists of opioid receptors and associated ligands distributed throughout the central nervous system and peripheral tissues, such as the nucleus accumbens (Fields, 2007; Trigo et al., 2010). The EOS is central in opioid-mediated reward (Koob, 1992; Olmstead and Franklin, 1997; Comings et al., 1999), social motivation (Chelnokova et al., 2014), and pleasure and pain perception (Janal et al., 1984; Leknes and Tracey, 2008). Elevated opioid levels are correlated with feelings of euphoria (Boecker et al., 2008), and Koepp et al. (2009) report activation of general opioid receptors in the hippocampus and amygdala in response to positive affect. Deactivation of certain opioid receptor sites has been associated with negative affect (Zubieta et al., 2003).
The possible role of the EOS in social bonding is formalized in the brain opioid theory of social attachment (BOTSA). BOTSA is based on evidence of behavioral and emotional similarities between those in intense relationships, and those addicted to narcotics (Insel, 2003). Furthermore, endogenous opioids, particularly endorphins, are related to social bonding in many non-human animals such as rhesus macaques (Schino and Troisi, 1992; Graves et al., 2002), other monkeys (Keverne et al., 1989; Martel et al., 1995; Ragen et al., 2013), voles (Resendez et al., 2013), puppies, rats and chicks (Panksepp et al., 1980), and mammals generally (Broad et al., 2006). Given the role of endorphins in bonding in other species, it is plausible that the EOS may also underpin human social bonds (Matthes et al., 1996; Moles et al., 2004; Depue and Morrone-Strupinsky, 2005; Dunbar, 2010).
Opioids are released in response to low levels of muscular and psychological stress (Howlett et al., 1984), for example during exercise (Harbach et al., 2000). Positron emission tomography (PET) scans have confirmed the euphoric state that follows exercise (termed “runner’s high”) is due to endogenous opioids (Boecker et al., 2008). Further to the effect on mood, opioids have an analgesic effect (Van Ree et al., 2000), and much evidence suggests that endorphins are central in the pain management system (D’Amato and Pavone, 1993; Benedetti, 1996; Zubieta et al., 2001; Fields, 2007; Bodnar, 2008; Dishman and O’Connor, 2009; Mueller et al., 2010). Given that direct measures of endogenous opioids are costly and invasive (Dearman and Francis, 1983), pain threshold is a commonly used proxy measure of endorphin release, and this has been operationalised using the length of time holding a hand in ice water (Dunbar et al., 2012a,b), a ski exercise (maintaining a squat position with legs at right angles: Dunbar et al., 2012a), an electrocutaneous simulator (Jamner and Leigh, 1999), pressure produced using a blood pressure cuff (Cogan et al., 1987; Cohen et al., 2010; Dunbar et al., 2012a,b), and the amount of pain medication requested by patients (Zillmann et al., 1993).
According to pain threshold assays, various exertive human social bonding activities, such as laughter (Dezecache and Dunbar, 2012; Dunbar et al., 2012a), group synchronized sport (Cohen et al., 2010; Sullivan and Rickers, 2013), and singing and dance (Dunbar et al., 2012b), trigger endorphin release. Specifically, synchronized exertive activity (such as rowing) elevates pain thresholds significantly more than non-synchronized exertion (Sullivan and Rickers, 2013; Sullivan et al., 2014), suggesting that rhythmic, music-based activities may similarly facilitate endorphin release.
ENDORPHINS AND MUSIC
Based on the association between exertion and endorphin release, a number of studies have investigated the effect of active engagement in musical activities (i.e., involving overt movement) and the EOS (see Table 1). For example, sufficiently vigorous singing, dancing, and drumming trigger a significantly larger increase in pain threshold and positive affect compared to listening to music and engaging in low energy musical activities (Dunbar et al., 2012b). In a recent set of studies, exercise machines were linked to musical output software such that individuals “created” music as they exerted themselves (Fritz et al., 2013a,b). These experiments demonstrated that when movement (during group exercise) results in musical feedback, participants perceived exertion to be lower (Fritz et al., 2013b), reported enhanced mood, and felt a greater desire to exert themselves further (Fritz et al., 2013a), in comparison to when they were exercising whilst listening (passively) to independently provided music. As such, perception of agency in a musical setting is associated with greater endorphin activation and may therefore lead to greater effects in terms of mood and ability to withstand strenuous exercise.
Table 1
| Passive listening | Active engagement | |
|---|---|---|
| Pain threshold, pain management | Post-operative pain: Koch et al. (1998), Allen et al. (2001), Good et al. (2001), Lepage et al. (2001), Nilsson et al. (2001, 2003), Nilsson (2008), Bernatzky et al. (2011) for a review see Cepeda et al. (2006) | Singing, drumming, dance: Dunbar et al. (2012b) |
| Brain activation regions | EOS, pleasure, and reward circuits: Blood and Zatorre (2001), Stefano et al. (2004), Menon and Levitin (2005) Nucleus accumbens and pleasure states: Koelsch (2014) | |
| Emotions and mood | Techno-music: Gerra et al. (1998) Emotional effects of music: Koelsch (2010) Positive affect: Huron (2006) | Increased positive affect: Dunbar et al. (2012b) Enhanced mood: Fritz et al. (2013a) |
| Health | Lower blood pressure and relaxation: Chiu and Kumar (2003), Stefano et al. (2004) Anxiolytic music: McKinney et al. (1997b) | |
| Other | Musical “thrills”: Goldstein (1980), Panksepp (1995), Menon and Levitin (2005) | Perception of exertion and desire to exert oneself: Fritz et al. (2013b) |
Summary of studies providing evidence for the role of EOS in music-related activities.
However, activation of the EOS through music is not limited exclusively to situations involving exertion (see Table 1). Listening to music reportedly helps to manage pre-operative hypertension and psychological stress (Allen et al., 2001), reduces sedative requirements during spinal anesthesia (Lepage et al., 2001) and other surgical procedures (Koch et al., 1998), decreases perception of pain (Good et al., 2001; Nilsson et al., 2003) thereby diminishing the need for opioid agonists following operative care (Cepeda et al., 2006; Bernatzky et al., 2011), and improves post-operative recovery (Nilsson et al., 2001). Many of the experiments in this area directly attribute these results to the EOS, and given the strong role of opioid receptor activation in analgesia (Leknes and Tracey, 2008), the body of work linking music and pain may generally be considered convincing evidence of the role of opioid activation.
Positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) research also provide evidence that passive listening to music activates the EOS and brain areas associated with pleasure and reward (Blood and Zatorre, 2001; Stefano et al., 2004). For example, recent evidence that music listening is associated with activation in areas such as the nucleus accumbens (Brown et al., 2004; Menon and Levitin, 2005; Koelsch, 2014), the high number of opioid receptors in this region (Fields, 2007; Trigo et al., 2010) and the role of opioids in mood and pleasure states (Berridge and Kringelbach, 2008) provide support for the theory that the EOS is involved in music listening.
The importance of the EOS in regulating affective experiences in response to music (Zubieta et al., 2003) is further supported by evidence linking music induced “thrills” to endorphin activation (Goldstein, 1980), and the EOS’s association with reward circuits (Menon and Levitin, 2005). In addition, the sense of elation that arises when engaging in musical activities has been attributed to endorphin release (Chiu and Kumar, 2003; Huron, 2006; Dunbar, 2009). Calming music is thought to act via the EOS by buffering the effect of stressful events (see McKinney et al., 1997b for a review), and relaxation following music listening is also linked to the EOS (Stefano et al., 2004). Gerra et al. (1998) report that listening to techno-music significantly changes emotional states (and increases beta-endorphin levels), due to its strong rhythmic beat and engagement of motor regions of the brain. Activation of the EOS, and its role in various affective, calming and analgesic effects, is therefore evident in cases of passive music listening, although a systematic investigation of this effect is still lacking.
It is important to note that there is also some evidence indicating that neurohormones other than endorphins are involved during music-based activities (e.g., Grape et al., 2003; Bachner-Melman et al., 2005; Chanda and Levitin, 2013). In a recent review, Chanda and Levitin (2013) highlight evidence suggesting that stress and arousal effects associated with music-based activities can be linked to cortisol, corticotrophin-releasing hormone and adrenocorticotropic hormone (e.g., McKinney et al., 1997a; Gerra et al., 1998). Various immunity benefits of music have been attributed to, inter alia, cortisol (e.g., Beck et al., 2000; Kuhn, 2002), cytokinin (e.g., Stefano et al., 2004), and growth hormones (e.g., Gerra et al., 1998). Finally, dopamine is key in reward and motivation circuits during musical activities (e.g., Salimpoor et al., 2011), which are likely to interact synergistically with the EOS in mediating the pleasure states associated with music (Chanda and Levitin, 2013). While we argue for further investigation of the EOS as a potential mediator of the positive social effects of musical engagement, it may be difficult to separate out the role of this hormone from other neurochemicals involved in these experiences.
As indicated by the evidence reviewed above, the way that we experience music, whether during passive listening or active engagement, appears to involve the EOS, and endorphins specifically. In the following section we discuss how both self-other merging and the EOS mechanisms might underpin our musical experiences.
FROM MUSIC TO SOCIAL BONDING
Both self-other merging and the EOS help explain the subjective experience of social bonding that can arise during musical activities, as illustrated by Figure 1. However, as these two mechanisms have thus far been independently investigated, the interplay between them remains unclear.
FIGURE 1
Schematic diagram illustrating possible direction of causality, and relationship between, mechanisms behind the social bonding effects of music.
As mentioned previously, dyads are capable of achieving synchrony with relative ease (even without music), while in larger groups, synchrony is facilitated via rhythmic scaffolding. Additionally, as music encourages movement (Janata and Grafton, 2003; Madison, 2006; Madison et al., 2011; Janata et al., 2012) by engaging motor regions of the brain (Levitin and Menon, 2003), we might expect engagement with musical sounds to be more exertive than with non-musical sounds. The combination of larger movements and the externalization of the target of synchrony likely facilitates synchronization.
Exertive movements cause affiliative sentiments and behaviors (e.g., Mueller et al., 2003), have effects on mood and emotion (e.g., Karageorghis and Terry, 2009), and, in combination with synchrony, can elevate pain thresholds (e.g., Cohen et al., 2010). These phenomena are all strongly associated with the EOS (e.g., Dishman and O’Connor, 2009). Accordingly, we propose that self-other matching and activation of the EOS are interconnected in explaining the bonding effects that arise during active engagement in group music-based activities, with a possibility that the EOS underpins the psychological experience of self-other merging.
In terms of passive listening to music, the literature reviewed here suggests that the EOS is likely to play a role also in the absence of explicit movement and self-other merging during synchrony. Dynamic attending theory (e.g., Jones and Boltz, 1989) suggests that through monitoring of events occurring with predictable temporal patterns we can become entrained to those events. This rhythmic predictability has been suggested to play a key role in the pleasure experienced when listening to music, which may be mediated by the release of endorphins (Huron, 2006; Margulis, 2013). The effect of tempo on arousal (Husain et al., 2002) and the strong ability for music to alter mood (Thayer et al., 1994) and motivational states (Frijda and Zeelenberg, 2001) are both congruous with evidence that EOS activation occurs in the brain when listening and entraining to music (Blood and Zatorre, 2001; Stefano et al., 2004; Menon and Levitin, 2005). Sievers et al. (2013) demonstrate that movement and music are processed cross-modally, as are the emotions expressed through movement and music. Elements of music significantly affect various dimensions of imagery relating to motion (Eitan and Granot, 2006), and listening to music may itself induce thoughts about movement, whether conscious or subconscious (Chen et al., 2008; Levitin and Tirovolas, 2009; Clarke, 2011). Through activation of motor regions of the brain during music listening (Levitin and Menon, 2003), passive engagement with music likely triggers the same neural pathways involved in active engagement (i.e. movement) to music, including pathways implicating the EOS. This activity in motor regions of the brain during music perception is likely to underlie the self-reported experience of “embodied movement” even when listening and not moving to music (e.g., Peters, 2010).
CONCLUSION
While most accounts of the relationship between music and social bonding have focused separately on self-other merging via synchrony or neurohormonal mechanisms, here we suggest that associations between the two need to be considered, especially when assessing large-scale musical activities. Future work should be directed toward ecologically valid musical experiences involving groups of people interacting with one another rather than dyadic interaction, exertive movements rather than small movements, and movements that are temporally co-ordinated rather than synchronized per se. Using these forms of musical activity it will become possible to explore the relative importance of self-other matching and EOS in music-based activities (including passive listening). Given that humans have significantly larger and more complex social networks than our primate cousins, research in this field will elucidate the means by which our species has the capacity to bond with large groups of conspecifics at the same time. It is likely that some combination of endorphin release and self-other merging lead to the social bonding effects of music, although the relationship between the two mechanisms remains to be sufficiently explored.
Statements
Acknowledgments
We would like to thank the European Research Council (Grant Number 295663) for its funding support during the writing of this article, and Cole Robertson for commenting on a draft of this manuscript.
Conflict of interest
The authors declare 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
1
AllenK.GoldenL. H.IzzoJ. L.ChingM. I.ForrestA.NilesC. R.et al (2001). Normalization of hypertensive responses during ambulatory surgical stress by perioperative music.Psychosom. Med.63487–492. 10.1097/00006842-200105000-00019
2
AtzilS.HendlerT.FeldmanR. (2011). Specifying the neurobiological basis of human attachment: brain, hormones, and behavior in synchronous and intrusive mothers.Neuropsychopharmacology362603–2615. 10.1038/npp.2011.172
3
Bachner-MelmanR.DinaC.ZoharA. H.ConstantiniN.LererE.HochS.et al (2005). AVPR1a and SLC6A4 gene polymorphisms are associated with creative dance performance.PLoS Genet.1:e42. 10.1371/journal.pgen.0010042
4
BartzJ. A.SimeonD.HamiltonH.KimS.CrystalS.BraunA.et al (2011). Oxytocin can hinder trust and cooperation in borderline personality disorder.Soc. Cogn. Affect. Neurosci.6556–563. 10.1093/scan/nsq085
5
BeckR. J.CesarioT. C.YousefiA.EnamotoH. (2000). Choral singing, performance perception, and immune system changes in alivary immunoglobulin a and cortisol.Music Percept. Interdiscip. J.1887–106. 10.2307/40285902
6
BenedettiF. (1996). The opposite effects of the opiate antagonist naloxone and the cholecystokinin antagonist proglumide on placebo analgesia.Pain64535–543. 10.1016/0304-3959(95)00179-4
7
BernatzkyG.PreschM.AndersonM.PankseppJ. (2011). Emotional foundations of music as a non-pharmacological pain management tool in modern medicine.Neurosci. Biobehav. Rev.351989–1999. 10.1016/j.neubiorev.2011.06.005
8
BerridgeK. C.KringelbachM. L. (2008). Affective neuroscience of pleasure: reward in humans and animals.Psychopharmacology (Berl.) 199457–480. 10.1007/s00213-008-1099-6
9
BisphamJ. (2006). Rhythm in music: what is it? Who has it? And why?Music Percept.24125–134. 10.1525/mp.2006.24.2.125
10
BloodA. J.ZatorreR. J. (2001). Intensely pleasurable responses to music correlate with activity in brain regions implicated in reward and emotion.Proc. Natl. Acad. Sci. U.S.A.9811818–11823. 10.1073/pnas.191355898
11
BodnarR. J. (2008). Endogenous opiates and behavior: 2007.Peptides292292–2375. 10.1016/j.peptides.2008.09.007
12
BoeckerH.SprengerT.SpilkerM. E.HenriksenG.KoppenhoeferM.WagnerK. J.et al (2008). The runner’s high: opioidergic mechanisms in the human brain.Cereb. Cortex182523–2531. 10.1093/cercor/bhn013
13
BotvinickM.CohenJ. (1998). Rubber hands “feel” touch that eyes see.Nature391:756. 10.1038/35784
14
BroadK. D.CurleyJ. P.KeverneE. B. (2006). Mother-infant bonding and the evolution of mammalian social relationships.Philos. Trans. R. Soc. Lond. B Biol. Sci.3612199–2214. 10.1098/rstb.2006.1940
15
BrownS. (2000). “Evolutionary models of music: from sexual selection to group selection,” inPerspectives in EthologyedsTonneauF.ThompsonN. S. (New York: Plenum) 221–281.
16
BrownS.JordaniaJ. (2011). Universals in the world’s musics.Psychol. Music41229–248. 10.1177/0305735611425896
17
BrownS.MartinezM. J.ParsonsL. M. (2004). Passive music listening spontaneously engages limbic and paralimbic systems.Neuroreport152033–2037. 10.1097/00001756-200409150-00008
18
BuccinoG.BinkofskiF.FinkG. R.FadigaL.FogassiL.GalleseV.et al (2001). Action observation activates premotor and parietal areas in a somatotopic manner: an fMRI study.Eur. J. Neurosci.13400–404. 10.1111/j.1460-9568.2001.01385.x
19
CaetanoG.JousmäkiV.HariR. (2007). Actor’s and observer’s primary motor cortices stabilize similarly after seen or heard motor actions.Proc. Natl. Acad. Sci. U.S.A.1049058–9062. 10.1073/pnas.0702453104
20
CarterC. S. (1998). Neuroendocrine perspectives on social attachment and love.Psychoneuroendocrinology23779–818. 10.1016/S0306-4530(98)00055-9
21
CepedaM. S.CarrD. B.LauJ.AlvarezH. (2006). Music for pain relief (review).Cochrane Database Syst. Rev.CD004843. 10.1002/14651858.CD004843.pub2
22
ChandaM. L.LevitinD. J. (2013). The neurochemistry of music.Trends Cogn. Sci.17179–193. 10.1016/j.tics.2013.02.007
23
ChartrandT. L.BarghJ. (1999). The chameleon effect: the perception-behavior link and social interaction.J. Pers. Soc. Psychol.76893–910. 10.1037/0022-3514.76.6.893
24
ChelnokovaO.LaengB.EikemoM.RiegelsJ.LøsethG.MaurudH.et al (2014). Rewards of beauty: the opioid system mediates social motivation in humans.Mol. Psychiatry19746–747. 10.1038/mp.2014.1
25
ChenJ. L.PenhuneV. B.ZatorreR. J. (2008). Listening to musical rhythms recruits motor regions of the brain.Cereb. Cortex182844–2854. 10.1093/cercor/bhn042
26
ChiuP.KumarA. (2003). Music therapy: loud noise or soothing notes.Int. Pediatr.18204–208.
27
ClarkeE. F. (2011). “Music perception and musical consciousness,” inMusic and Consciousness: Philosophical, Psychological, and Cultural Perspectives,edsClarkeD.ClarkeE. F. (New York: Oxford University Press), 193–213.
28
ClaytonM.SagerR.WillU. (2005). In time with the music: the concept of entrainment and its significance for ethnomusicology.Eur. J. Soc. Psychol.113–142.
29
CoganR.CoganD.WaltzW.McCueM. (1987). Effects of laughter and relaxation on discomfort thresholds.J. Behav. Med.10139–144. 10.1007/BF00846422
30
CohenE. E. A.Ejsmond-FreyR.KnightN.DunbarR. I. M. (2010). Rowers’ high: behavioural synchrony is correlated with elevated pain thresholds.Biol. Lett.6106–108. 10.1098/rsbl.2009.0670
31
ComingsD. E.BlakeH.DietzG.Gade-AndavoluR.LegroR. S.SaucierG.et al (1999). The proenkephalin gene (PENK) and opioid dependence.Neuroreport101133–1135. 10.1097/00001756-199904060-00042
32
CrossI. (2001). Music, cognition, culture, and evolution.Ann. N. Y. Acad. Sci.93028–42. 10.1111/j.1749-6632.2001.tb05723.x
33
CurleyJ. P.KeverneE. B. (2005). Genes, brains and mammalian social bonds.Trends Ecol. Evol.20561–567. 10.1016/j.tree.2005.05.018
34
D’AmatoF. R.PavoneF. (1993). Endogenous opioids: a proximate reward mechanism for kin selection?Behav. Neural Biol.6079–83. 10.1016/0163-1047(93)90768-D
35
DearmanJ.FrancisK. T. (1983). Plasma levels of catecholamines, cortisol, and beta-endorphins in male athletes after running 26.2, 6, and 2 miles.J. Sports Med. Phys. Fitness2330–38.
36
DecetyJ.SommervilleJ. A. (2003). Shared representations between self and other: a social cognitive neuroscience view.Trends Cogn. Sci.7527–533. 10.1016/j.tics.2003.10.004
37
DemosA. P.ChaffinR.BegoshK. T.DanielsJ. R.MarshK. L. (2012). Rocking to the beat: effects of music and partner’s movements on spontaneous interpersonal coordination.J. Exp. Psychol. Gen.14149–53. 10.1037/a0023843
38
DepueR. A.Morrone-StrupinskyJ. V. (2005). A neurobehavioral model of affiliative bonding: implications for conceptualizing a human trait of affiliation.Behav. Brain Sci.28313–395. 10.1017/S0140525X05000063
39
DezecacheG.DunbarR. I. M. (2012). Sharing the joke: the size of natural laughter groups.Evol. Hum. Behav.33775–779. 10.1016/j.evolhumbehav.2012.07.002
40
DishmanR. K.O’ConnorP. J. (2009). Lessons in exercise neurobiology: the case of endorphins.Ment. Health Phys. Act.24–9. 10.1016/j.mhpa.2009.01.002
41
DomesG.HeinrichsM.MichelA.BergerC.HerpertzS. C. (2007). Oxytocin improves “mind-reading” in humans.Biol. Psychiatry61731–733. 10.1016/j.biopsych.2006.07.015
42
DumasG.NadelJ.SoussignanR.MartinerieJ.GarneroL. (2010). Inter-brain synchronization during social interaction.PLoS ONE5:e12166. 10.1371/journal.pone.0012166
43
DunbarR. I. M. (2004). “Language, music, and laughter in evolutionary perspective,” inEvolution of Communication Systems: A Comparative Approach,edsOllerD. K.GriebelU. (Cambridge, MA: MIT Press) 257–274.
44
DunbarR. I. M. (2009). “Mind the bonding gap: constraints on the evolution of hominin societies,” inPattern and Process in Cultural Evolutioned.ShennanS. (Berkeley, CA: University of California Press) 223–234.
45
DunbarR. I. M. (2010). The social role of touch in humans and primates: behavioural function and neurobiological mechanisms.Neurosci. Biobehav. Rev.34260–268. 10.1016/j.neubiorev.2008.07.001
46
DunbarR. I. M. (2012a). Bridging the bonding gap: the transition from primates to humans.Philos. Trans. R. Soc. Lond. B Biol. Sci.3671837–1846. 10.1098/rstb.2011.0217
47
DunbarR. I. M. (2012b). “On the evolutionary function of song and dance,” inMusic, Language and Human Evolution,edsBannanN.MithenS. (Oxford: Oxford University Press) 201–214.
48
DunbarR. I. M.BaronR.FrangouA.PearceE.Van LeeuwenE. J. C.StowJ.et al (2012a). Social laughter is correlated with an elevated pain threshold.Proc. Biol. Sci.2791161–1167. 10.1098/rspb.2011.1373
49
DunbarR. I. M.KaskatisK.MacDonaldI.BarraV. (2012b). Performance of music elevates pain threshold and positive affect: implications for the evolutionary function of music.Evol. Psychol.10688–702.
50
EitanZ.GranotR. Y. (2006). How music moves.Music Percept.23221–248. 10.1525/mp.2006.23.3.221
51
FadigaL.CraigheroL.BuccinoG.RizzolattiG. (2002). Speech listening specifically modulates the excitability of tongue muscles: a TMS study.Eur. J. Neurosci.15399–402. 10.1046/j.0953-816x.2001.01874.x
52
FadigaL.FogassiL.PavesiG.RizzolattiG. (1995). Motor facilitation during action observation: a magnetic stimulation study.J. Neurophysiol.732608–2611.
53
FeldmanR. (2012). Oxytocin and social affiliation in humans.Horm. Behav.61380–391. 10.1016/j.yhbeh.2012.01.008
54
FieldsH. L. (2007). Understanding how opioids contribute to reward and analgesia.Reg. Anesth. Pain Med.32242–246. 10.1016/j.rapm.2007.01.001
55
FreemanW. J.III. (2000). “A neurobiological role of music in social bonding,” inThe Origins of Music,edsWallinN.MerkurB.BrownS. (Cambridge, MA: MIT Press) 411–424.
56
FrijdaN.ZeelenbergM. (2001). “Appraisal: what is the dependent?” inAppraisal Processes in Emotion,edsDavidsonR. J.EkmanP.SchererK. R. (Oxford: Oxford University Press) 141–155.
57
FritzT. H.HalfpaapJ.GrahlS.KirklandA.VillringerA. (2013a). Musical feedback during exercise machine workout enhances mood.Front. Psychol.4:921. 10.3389/fpsyg.2013.00921
58
FritzT. H.HardikarS.DemoucronM.NiessenM.DemeyM.GiotO.et al (2013b). Musical agency reduces perceived exertion during strenuous physical performance.Proc. Natl. Acad. Sci. U.S.A.11017784–17789. 10.1073/pnas.1217252110
59
GalleseV.FadigaL.FogassiL.RizzolattiG. (1996). Action recognition in the premotor cortex.Brain119(Pt 2)593–609. 10.1093/brain/119.2.593
60
GerraG.ZaimovicA.FranchiniD.PalladinoM.GiucastroG.RealiN.et al (1998). Neuroendocrine responses of healthy volunteers to “techno-music”: relationships with personality traits and emotional state.Int. J. Psychophysiol.2899–111. 10.1016/S0167-8760(97)00071-8
61
GoldsteinA. (1980). Thrills in response to music and other stimuli.Physiol. Psychol.8126–129. 10.3758/BF03326460
62
GoodM.Stanton-HicksM.GrassJ. A.AndersonG. C.LaiH. L.RoykulcharoenV.et al (2001). Relaxation and music to reduce postsurgical pain.J. Adv. Nurs.33208–215. 10.1046/j.1365-2648.2001.01655.x
63
GrapeC.SandgrenM.HanssonL.-O.EricsonM.TheorellT. (2003). Does singing promote well-being?: an empirical study of professional and amateur singers during a singing lesson.Integr. Physiol. Behav. Sci.3865–74. 10.1007/BF02734261
64
GravesF. C.WallenK.MaestripieriD. (2002). Opioids and attachment in rhesus macaque (Macaca mulatta) abusive mothers.Behav. Neurosci.116489–493. 10.1037//0735-7044.116.3.489
65
GuastellaA. J.MitchellP. B.DaddsM. R. (2008). Oxytocin increases gaze to the eye region of human faces.Biol. Psychiatry633–5. 10.1016/j.biopsych.2007.06.026
66
HagenE. H.BryantG. A. (2003). Music and dance as a coalition signalling system.Hum. Nat.1421–51. 10.1007/s12110-003-1015-z
67
HarbachH.HellK.GramschC.KatzN.HempelmannG.TeschemacherH. (2000). Beta-endorphin (1-31) in the plasma of male volunteers undergoing physical exercise.Psychoneuroendocrinology25551–562. 10.1016/S0306-4530(00)00009-3
68
HickokG. (2009). Eight problems for the mirror neuron theory of action understanding in monkeys and humans.J. Cogn. Neurosci.211229–1243. 10.1162/jocn.2009.21189
69
HoveM. J.RisenJ. L. (2009). It’s all in the timing: interpersonal synchrony increases affiliation.Soc. Cogn.27949–961. 10.1521/soco.2009.27.6.949
70
HowlettT. A.TomlinS.NgahfoongL.ReesL. H.BullenB. A.SkrinarG. S.et al (1984). Release of beta endorphin and met-enkephalin during exercise in normal women: response to training.Br. Med. J.2881950–1952. 10.1136/bmj.288.6435.1950
71
HuronD. (2001). Is music an evolutionary adaptation?Ann. N. Y. Acad. Sci.93043–61. 10.1111/j.1749-6632.2001.tb05724.x
72
HuronD. (2006). Sweet Anticipation: Music and the Psychology of Expectation.Cambridge, MA: MIT Press.
73
HusainG.ThompsonW. F.SchellenbergE. G. (2002). Effects of musical tempo and mode on arousal, mood, and spatial abilities.Music Percept.20151–171. 10.1525/mp.2002.20.2.151
74
InselT. R. (2003). Is social attachment an addictive disorder?Physiol. Behav.79351–357. 10.1016/S0031-9384(03)00148-3
75
InselT. R. (2010). The challenge of translation in social neuroscience: a review of oxytocin, vasopressin, and affiliative behavior.Neuron65768–779. 10.1016/j.neuron.2010.03.005
76
IsenA. M. (1970). Success, failure, attention, and reaction to others: the warm glow of success.J. Pers. Soc. Psychol.15294–301. 10.1037/h0029610
77
IssartelJ.MarinL.CadopiM. (2007). Unintended interpersonal co-ordination: “can we march to the beat of our own drum?”Neurosci. Lett.411174–179. 10.1016/j.neulet.2006.09.086
78
JamnerL. D.LeighH. (1999). Repressive/defensive coping, endogenous opioids and health: how a life so perfect can make you sick.Psychiatry Res.8517–31. 10.1016/S0165-1781(98)00134-6
79
JanalM. N.ColtE. W.ClarkW. C.GlusmanM. (1984). Pain sensitivity, mood and plasma endocrine levels in man following long-distance running: effects of naloxone.Pain1913–25. 10.1016/0304-3959(84)90061-7
80
JanataP.GraftonS. T. (2003). Swinging in the brain: shared neural substrates for behaviors related to sequencing and music.Nat. Neurosci.6682–687. 10.1038/nn1081
81
JanataP.TomicS. T.HabermanJ. M. (2012). Sensorimotor coupling in music and the psychology of the groove.J. Exp. Psychol. Gen.14154–75. 10.1037/a0024208
82
JonesM. R.BoltzM. (1989). Dynamic attending and responses to time.Psychol. Rev.96459–491. 10.1037/0033-295X.96.3.459
83
KarageorghisC. I.TerryP. C. (2009). “The psychological, psychophysical, and ergogenic effects of music in sport: a review and synthesis,” inSporting Sounds: Relationships Between Sport and Music,edsBatemanA.BaleJ. (London: Routeledge) 13–36.
84
KeverneE. B.MartenszN. D.TuiteB. (1989). Beta-endorphin concentrations in cerebrospinal fluid of monkeys are influenced by grooming relationships.Psychoneuroendocrinology14155–161. 10.1016/0306-4530(89)90065-6
85
KirschnerS.TomaselloM. (2009). Joint drumming: social context facilitates synchronization in preschool children.J. Exp. Child Psychol.102299–314. 10.1016/j.jecp.2008.07.005
86
KnoblichG.SebanzN. (2008). Evolving intentions for social interaction: from entrainment to joint action.Philos. Trans. R. Soc. Lond. B Biol. Sci.3632021–2031. 10.1098/rstb.2008.0006
87
KochM. E.KainZ. N.AyoubC.RosenbaumS. H. (1998). The sedative and analgesic sparing effect of music.Anesthesiology89300–306. 10.1097/00000542-199808000-00005
88
KoelschS. (2010). Towards a neural basis of music-evoked emotions.Trends Cogn. Sci.14131–137. 10.1016/j.tics.2010.01.002
89
KoelschS. (2014). Brain correlates of music-evoked emotions.Nat. Rev. Neurosci.15170–180. 10.1038/nrn3666
90
KoeppM. J.HammersA.LawrenceA. D.AsselinM. C.GrasbyP. M.BenchC. J. (2009). Evidence for endogenous opioid release in the amygdala during positive emotion.Neuroimage44252–256. 10.1016/j.neuroimage.2008.08.032
91
KokalI.EngelA.KirschnerS.KeysersC. (2011). Synchronized drumming enhances activity in the caudate and facilitates prosocial commitment-if the rhythm comes easily.PLoS ONE6:e27272. 10.1371/journal.pone.0027272
92
KoobG. F. (1992). Drugs of abuse: anatomy, pharmacology and function of reward pathways.Trends Pharmacol. Sci.13177–184. 10.1016/0165-6147(92)90060-J
93
KosfeldM.HeinrichsM.ZakP. J.FischbacherU.FehrE. (2005). Oxytocin increases trust in humans.Nature435673–676. 10.1038/nature03701
94
KuhnD. (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.3930–39. 10.1093/jmt/39.1.30
95
LaFranceM. (1979). Nonverbal synchrony and rapport: analysis by the cross-lag panel technique.Soc. Psychol. Q.4266–70. 10.2307/3033875
96
LaFranceM.BroadbentM. (1976). Group rapport: posture sharing as a nonverbal indicator.Group Organ. Stud.1328–333. 10.1177/105960117600100307
97
LakensD.StelM. (2011). If they move in sync, they must feel in sync: movement synchrony leads to attributions of rapport and entitativity.Soc. Cogn.291–14. 10.1521/soco.2011.29.1.1
98
LakinJ. L.ChartrandT. L. (2003). Using nonconscious behavioral mimicry to create affiliation and rapport.Psychol. Sci.14334–339. 10.1111/1467-9280.14481
99
LaunayJ.DeanR. T.BailesF. (2013). Synchronization can influence trust following virtual interaction.Exp. Psychol.6053–63. 10.1027/1618-3169/a000173
100
LaunayJ.DeanR. T.BailesF. (2014). Synchronising movements with the sounds of a virtual partner enhances partner likeability.Cogn. Process.10.1007/s10339-014-0618-0[Epub ahead of print].
101
LeknesS.TraceyI. (2008). A common neurobiology for pain and pleasure.Nat. Rev. Neurosci.9314–320. 10.1038/nrn2333
102
LepageC.DroletP.GirardM.GrenierY.De GagnéR. (2001). Music decreases sedative requirements during spinal anesthesia.Anesth. Analg.93912–916. 10.1097/00000539-200110000-00022
103
LevitinD. J.MenonV. (2003). Musical structure is processed in “language” areas of the brain: a possible role for Brodmann Area 47 in temporal coherence.Neuroimage202142–2152. 10.1016/j.neuroimage.2003.08.016
104
LevitinD. J.TirovolasA. K. (2009). Current advances in the cognitive neuroscience of music.Ann. N. Y. Acad. Sci.1156211–231. 10.1111/j.1749-6632.2009.04417.x
105
LindenbergerU.LiS.-C.GruberW.MüllerV. (2009). Brains swinging in concert: cortical phase synchronization while playing guitar.BMC Neurosci.10:22. 10.1186/1471-2202-10-22
106
LumsdenJ.MilesL. K.RichardsonM. J.SmithC. A.MacraeC. N. (2012). Who syncs? Social motives and interpersonal coordination.J. Exp. Soc. Psychol.48746–751. 10.1016/j.jesp.2011.12.007
107
MachinA. J.DunbarR. I. M. (2011). The brain opioid theory of social attachment: a review of the evidence.Behaviour148985–1025. 10.1163/000579511X596624
108
MadisonG. S. (2006). Experiencing groove induced by music: consistency and phenomenology.Music Percept. Interdiscip. J.24201–208. 10.1525/mp.2006.24.2.201
109
MadisonG. S.GouyonF.UllénF.HörnströmK. (2011). Modeling the tendency for music to induce movement in humans: first correlations with low-level audio descriptors across music genres.J. Exp. Psychol. Hum. Percept. Perform.371578–1594. 10.1037/a0024323
110
MargulisE. H. (2013). On Repeat: How Music Plays the Mind.New York: Oxford University Press.
111
MarshK. L.RichardsonM. J.SchmidtR. C. (2009). Social connection through joint action and interpersonal coordination.Top. Cogn. Sci.1320–339. 10.1111/j.1756-8765.2009.01022.x
112
MartelF. L.NevisonC. M.RaymentD.SimpsonM. J.KeverneE. B. (1993). Opioid receptor blockade reduces maternal affect and social grooming in rheses monkeys.Psychoneuroendocrinology18307–321. 10.1016/0306-4530(93)90027-I
113
MartelF. L.NevisonC. M.SimpsonM. J.KeverneE. B. (1995). Effects of opioid receptor blockade on the social behavior of rhesus monkeys living in large family groups.Dev. Psychobiol.2871–84. 10.1002/dev.420280202
114
MatthesH. W. D.MaldonadoR.SimoninF.ValverdeO.SloweS.KitchenI.et al (1996). Loss of morphine-induced analgesia, reward effect and withdrawal symptoms in mice lacking the mu-opioid-receptor gene.Nature383819–823. 10.1038/383819a0
115
McDermottJ. (2008). The evolution of music.Nature453287–288. 10.1038/453287a
116
McKinneyC. H.AntoniM. H.KumarM.RimsF. C.McCabeP. M. (1997a). Effects of guided imagery and music (GIM) therapy on mood and cortisol in healthy adults.Health Psychol.16390–400. 10.1037/0278-6133.16.4.390
117
McKinneyC. H.TimsF. C.KumarA. M.KumarM. (1997b). The effect of selected classical music and spontaneous imagery on plasma beta-endorphin.J. Behav. Med.2085–99. 10.1023/A:1025543330939
118
McNeillW. H. (1995). Keeping Together in Time.Cambridge, MA: Harvard University Press.
119
MenonV.LevitinD. J. (2005). The rewards of music listening: response and physiological connectivity of the mesolimbic system.Neuroimage28175–184. 10.1016/j.neuroimage.2005.05.053
120
MerkerB. (2000). “Synchronous chorusing and human origins,” inThe Origins of MusicedsWallinN.MerkerB. H.BrownS. (Cambridge, MA: MIT Press) 315–327.
121
MerkerB.MadisonG. S.EckerdalP. (2009). On the role and origin of isochrony in human rhythmic entrainment.Cortex454–17. 10.1016/j.cortex.2008.06.011
122
Meyer-LindenbergA.DomesG.KirschP.HeinrichsM. (2011). Oxytocin and vasopressin in the human brain: social neuropeptides for translational medicine.Nat. Rev. Neurosci.12524–538. 10.1038/nrn3044
123
MilesL. K.GriffithsJ. L.RichardsonM. J.MacraeC. N. (2010). Too late to coordinate: contextual influences on behavioral synchrony.Eur. J. Soc. Psychol.4052–60. 10.1002/ejsp.721
124
MilesL. K.LumsdenJ.RichardsonM. J.Neil MacraeC. (2011). Do birds of a feather move together? Group membership and behavioral synchrony.Exp. Brain Res.211495–503. 10.1007/s00221-011-2641-z
125
MilesL. K.NindL. K.MacraeC. N. (2009). The rhythm of rapport: interpersonal synchrony and social perception.J. Exp. Soc. Psychol.45585–589. 10.1016/j.jesp.2009.02.002
126
MillerG. F. (2000). The Mating Mind: How Sexual Choice Shaped the Evolution of the Human Nature.New York, NY: Random House Inc.
127
MolesA.KiefferB. L.D’AmatoF. R. (2004). Deficit in attachment behavior in mice lacking the mu-opioid receptor gene.Science3041983–1986. 10.1126/science.1095943
128
MuellerC.KlegaA.BuchholzH.-G.RolkeR.MagerlW.SchirrmacherR.et al (2010). Basal opioid receptor binding is associated with differences in sensory perception in healthy human subjects: a [18F]diprenorphine PET study.Neuroimage49731–737. 10.1016/j.neuroimage.2009.08.033
129
MuellerF.AgamanolisS.PicardR. (2003). “Exertion interfaces: sports over a distance for social bonding and fun,” inProceedings of the SIGCHI Conference on Human Factors in Computing Systems,Lauderdale, FL.
130
NettlB. (1983). The Study of Ethnomusicology: Twenty-Nine Issues and Concepts.Urbana, IL: University of Illinois Press.
131
NettlB. (2000). “An ethnomusicologist contemplates universals in musical sound and musical culture,” inThe Origins of Music,edsWallinN. L.MerkerB.BrownS. (Cambridge, MA: MIT Press) 463–472.
132
NilssonU. (2008). Effects of music interventions: a systematic review.AORN J.87780–807. 10.1016/j.aorn.2007.09.013
133
NilssonU.RawalN.EnqvistB.UnossonM. (2003). Analgesia following music and therapeutic suggestions in the PACU in ambulatory surgery; a randomized controlled trial.Acta Anaesthesiol. Scand.47278–283. 10.1034/j.1399-6576.2003.00064.x
134
NilssonU.RawalN.UneståhlL. E.ZetterbergC.UnossonM. (2001). Improved recovery after music and therapeutic suggestions during general anaesthesia: a double-blind randomised controlled trial.Acta Anaesthesiol. Scand.45812–817. 10.1034/j.1399-6576.2001.045007812.x
135
OlmsteadM. C.FranklinK. B. (1997). The development of a conditioned place preference to morphine: effects of lesions of various CNS sites.Behav. Neurosci.1111313–1323. 10.1037/0735-7044.111.6.1313
136
OullierO.De GuzmanG. C.JantzenK. J.LagardeJ.KelsoJ. A S. (2008). Social coordination dynamics: measuring human bonding.Soc. Neurosci.3178–192. 10.1080/17470910701563392
137
OullierO.JantzenK. J.SteinbergF. L.KelsoJ. A S. (2005). Neural substrates of real and imagined sensorimotor coordination.Cereb. Cortex15975–985. 10.1093/cercor/bhh198
138
OveryK.Molnar-SzakacsI. (2009). Being together in time: musical experience and the mirror neuron system.Music Percept.26489–504. 10.1525/mp.2009.26.5.489
139
PankseppJ. (1995). The emotional sources of “chills” induced by music.Music Percept. Interdiscip. J.13171–207. 10.2307/40285693
140
PankseppJ.HermanB. H.VilbergT.BishopP.De EskinaziF. G. (1980). Endogenous opioids and social behavior.Neurosci. Biobehav. Rev.4473–487. 10.1016/0149-7634(80)90036-6
141
PetersD. (2010). Enactment in listening: intermedial dance in EGM sonic scenarios and the bodily grounding of the listening experience.Perform. Res.15:3. 10.1080/13528165.2010.527211
142
PinkerS. (1997). How the Mind Works.New York: Norton.
143
RagenB. J.ManingerN.MendozaS. P.JarchoM. R.BalesK. L. (2013). Presence of a pair-mate regulates the behavioral and physiological effects of opioid manipulation in the monogamous titi monkey (Callicebus cupreus).Psychoneuroendocrinology382448–2461. 10.1016/j.psyneuen.2013.05.009
144
ReddishP.FischerR.BulbuliaJ. (2013). Let’s dance together: synchrony, shared intentionality and cooperation.PLoS ONE8:e71182. 10.1371/journal.pone.0071182
145
ReddyV. (2003). On being the object of attention: implications for self-other consciousness.Trends Cogn. Sci.7397–402. 10.1016/S1364-6613(03)00191-8
146
ResendezS. L.DomeM.GormleyG.FrancoD.NevárezN.HamidA. A.et al (2013). μ-Opioid receptors within subregions of the striatum mediate pair bond formation through parallel yet distinct reward mechanisms.J. Neurosci.339140–9149. 10.1523/JNEUROSCI.4123-12.2013
147
RizzolattiG. (2005). The mirror neuron system and its function in humans.Anat. Embryol. (Berl.) 210419–421. 10.1007/s00429-005-0039-z
148
RizzolattiG.FadigaL.GalleseV.FogassiL. (1996). Premotor cortex and the recognition of motor actions.Brain Res. Cogn. Brain Res.3131–141. 10.1016/0926-6410(95)00038-0
149
RoedererJ. G. (1984). The search for a survival value of music.Music Percept.1350–356. 10.2307/40285265
150
SalimpoorV. N.BenovoyM.LarcherK.DagherA.ZatorreR. J. (2011). Anatomically distinct dopamine release during anticipation and experience of peak emotion to music.Nat. Neurosci.14257–262. 10.1038/nn.2726
151
SavaskanE.EhrhardtR.SchulzA.WalterM.SchächingerH. (2008). Post-learning intranasal oxytocin modulates human memory for facial identity.Psychoneuroendocrinology33368–374. 10.1016/j.psyneuen.2007.12.004
152
SchinoG.TroisiA. (1992). Opiate receptor blockade in juvenile macaques: effect on affiliative interactions with their mothers and group companions.Brain Res.576125–130. 10.1016/0006-8993(92)90617-I
153
SebanzN.BekkeringH.KnoblichG. (2006). Joint action: bodies and minds moving together.Trends Cogn. Sci.1070–76. 10.1016/j.tics.2005.12.009
154
SieversB.PolanskyL.CaseyM.WheatleyT. (2013). Music and movement share a dynamic structure that supports universal expressions of emotion.Proc. Natl. Acad. Sci. U.S.A.11070–75. 10.1073/pnas.1209023110
155
SommervilleJ. A.DecetyJ. (2006). Weaving the fabric of social interaction: articulating developmental psychology and cognitive neuroscience in the domain of motor cognition.Psychon. Bull. Rev.13179–200. 10.3758/BF03193831
156
StefanoG. B.ZhuW.CadetP.SalamonE.MantioneK. J. (2004). Music alters constitutively expressed opiate and cytokine processes in listeners.Med. Sci. Monit.10 MS18–MS27.
157
StelM.BlascovichJ.MccallC.MastopJ.Van baarenR. B.VonkR. (2010). Mimicking disliked others: effects of a priori liking on the mimicry-liking link.Eur. J. Soc. Psychol.40867–880. 10.1002/ejsp.655
158
SullivanP.RickersK. (2013). The effect of behavioral synchrony in groups of teammates and strangers.Int. J. Sport Exerc. Psychol.11286–291. 10.1080/1612197X.2013.750139
159
SullivanP. J.RickersK.GammageK. L. (2014). The effect of different phases of synchrony on pain threshold.Gr. Dyn.18122–128. 10.1037/gdn0000001
160
ThayerR. E.NewmanJ. R.McclainT. M. (1994). Self-regulation self-regulation of mood: strategies for changing a bad mood, raising energy, and reducing tension search.J. Pers. Soc. Psychol.64910–925. 10.1037/0022-3514.67.5.910
161
TognoliE.LagardeJ.De GuzmanG. C.KelsoJ. A S. (2007). The phi complex as a neuromarker of human social coordination.Proc. Natl. Acad. Sci. U.S.A.1048190–8195. 10.1073/pnas.0611453104
162
TomaselloM.CarpenterM.CallJ.BehneT.MollH. (2005). Understanding and sharing intentions: the origins of cultural cognition.Behav. Brain Sci.28675–691; discussion 691–735. 10.1017/S0140525X05000129
163
TrigoJ. M.Martin-GarcíaE.BerrenderoF.RobledoP.MaldonadoR. (2010). The endogenous opioid system: a common substrate in drug addiction.Drug Alcohol Depend.108183–194. 10.1016/j.drugalcdep.2009.10.011
164
ValdesoloP.DestenoD. (2011). Synchrony and the social tuning of compassion.Emotion11262–266. 10.1037/a0021302
165
Van BaarenR. B.HollandR. W.KawakamiK.Van KnippenbergA. (2004). Mimicry and prosocial behavior.Psychol. Sci.1571–74. 10.1111/j.0963-7214.2004.01501012.x
166
Van ReeJ. M.NiesinkR. J.Van WolfswinkelL.RamseyN. F.KornetM. M.Van FurthW. R.et al (2000). Endogenous opioids and reward.Eur. J. Pharmacol.40589–101. 10.1016/S0014-2999(00)00544-6
167
van UlzenN. R.LamothC. J. C.DaffertshoferA.SeminG. R.BeekP. J. (2008). Characteristics of instructed and uninstructed interpersonal coordination while walking side-by-side.Neurosci. Lett.43288–93. 10.1016/j.neulet.2007.11.070
168
WiltermuthS. S. (2012). Synchronous activity boosts compliance with requests to aggress.J. Exp. Soc. Psychol.48453–456. 10.1016/j.jesp.2011.10.007
169
WiltermuthS. S.HeathC. (2009). Synchrony and cooperation.Psychol. Sci.201–5. 10.1111/j.1467-9280.2008.02253.x
170
ZakP. J.KurzbanR.MatznerW. T. (2005). Oxytocin is associated with human trustworthiness.Horm. Behav.48522–527. 10.1016/j.yhbeh.2005.07.009
171
ZakP. J.StantonA. A.AhmadiS. (2007). Oxytocin increases generosity in humans.PLoS ONE2:e1128. 10.1371/journal.pone.0001128
172
ZillmannD.RockwellS.SchweitzerK.SundarS. S. (1993). Does humor facilitate coping with physical discomfort?Motiv. Emot.171–21. 10.1007/BF00995204
173
ZubietaJ.-K.KetterT. A.BuellerJ. A.XuY.KilbournM. R.YoungE. A.et al (2003). Regulation of human affective responses by anterior cingulate and limbic mu-opioid neurotransmission.Arch. Gen. Psychiatry601145–1153. 10.1001/archpsyc.60.11.1145
174
ZubietaJ. K.SmithY. R.BuellerJ. A.XuY.KilbournM. R.JewettD. M.et al (2001). Regional mu opioid receptor regulation of sensory and affective dimensions of pain.Science293311–315. 10.1126/science.1060952
Summary
Keywords
music, rhythm, social bonding, endorphins, self-other merging, synchrony
Citation
Tarr B, Launay J and Dunbar RIM (2014) Music and social bonding: “self-other” merging and neurohormonal mechanisms. Front. Psychol. 5:1096. doi: 10.3389/fpsyg.2014.01096
Received
01 June 2014
Accepted
10 September 2014
Published
30 September 2014
Volume
5 - 2014
Edited by
Guy Madison, Umeå University, Sweden
Reviewed by
Elizabeth Hellmuth Margulis, University of Arkansas, USA; Guy Madison, Umeå University, Sweden
Copyright
© 2014 Tarr, Launay and Dunbar.
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) or licensor 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: Bronwyn Tarr, Social and Evolutionary Neuroscience Research Group, Department of Experimental Psychology, University of Oxford, Tinbergen Building, Oxford OX1 3UD, UK e-mail: bronwyn.tarr@psy.ox.ac.uk
This article was submitted to Auditory Cognitive Neuroscience, a section of the journal Frontiers in Psychology.
Disclaimer
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher.