On the embodied meaning of emotional responses to music
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On the embodied meaning of emotional responses to music: A semiotic perspective


Semiotica 231 (2019), 225-244.


Previous attempts to find meaning in emotional responses to music often begin with analysis of dynamic tonal patterns, with observation of the emotional behavior of listeners or with self-reports of emotional feelings. In this study, we begin with a somewhat detailed description of physical processes in the human auditory system that lead to the activation of processes in the autonomic nervous system, which produce embodied emotional responses to environmental challenges. We then propose an answer to the question: Why were some of the same embodied responses that were originally adapted to meet the challenges of self-preservation and self-perpetuation in the course of human evolution coopted to serve as responses to perceived dynamic patterns in music? We find that a likely answer to this question involves uncertainties in the possible outcomes of antecedent or consequent musical events.

Keywords: autonomic nervous system, emotional sign, embodied meaning, exaptation, musical implication, Leonard B.Meyer

1. Introduction

It is common knowledge that humans respond to external and internal stimuli both cognitively and emotionally. Cognitive responses include thought, speech and voluntary behavior, whereas emotional responses include feelings, vocalizations and involuntary behavior. Both cognitive and emotional responses are grounded in physiology, i.e., physical and chemical bodily functions. From a phaneroscopic perspective, thought and feelings are private responses, while speech, vocalizations and behavior are public responses that may present as visual or auditory signs. In this paper, our approach to the meaning of emotional responses to music is based upon functional and evolutionary principles. In sections 2 and 3, we define the major semiotic, anatomic and functional terms that are relevant to the subsequent argument. In sections 4, 5 and 6, we briefly sketch the anatomic and functional pathways involved in the processing of cognitive and emotional responses to auditory stimuli. In sections 7 and 8, we provide a detailed account of the biological functions of the autonomic nervous system and their role in the formation of emotional responses to external stimuli. In section 9, we summarize the results of empirical studies of emotional responses to musical stimuli. In sections 10 and 11, we propose an evolutionary explanation for the observed commonalities between biological and aesthetic responses to external stimuli, as mediated by the autonomic nervous system. In section 12, we present a simplified version of Leonard B. Meyer’s theory of implicative meaning in music. In section 13, we present a plausible explanation for emotional responses to music, specifically those involving ‘thrills’ and tears. In conclusion, we summarize the argument that characterizes the embodied meaning of emotional responses to music as based upon uncertainties in the possible outcomes of anticipated musical events or in the possible consequences of experienced musical events.

2. Categories and signs

The argument of this paper refers to semiotic principles that were originally conceived by Charles S. Peirce. Peirce postulated a triadic categorization of the mental operations involved in thought processes. In turn, Peirce observed that these operations determine the Categories of Being that are accessible to thought. Accordingly, a 1st Category process involves attribution, a 2nd Category process involves opposition, and a 3rd Category process involves mediation of an opposition (c.f. Peirce c1890: 1.356). From this scheme, Peirce derived a triadic concept of ‘sign’ or ‘representation.’ To facilitate its use in applications, we restate the Peircean concept of ‘sign’ as follows: A sign is a 3-part relation in which a 1st relate is an attribution by perception, a 2nd relate is in opposition to the 1st relate by recollection and a 3rd relate mediates between the two by conception. In triadic signs, we shall refer to the 1st relate as a representamen, the 2nd relate as a significate and the 3rd relate as an interpretant. As an intuitive restatement, we conceive of a sign as some thing that stands for some other thing in some way to some one (c.f. Peirce c1897: 2.228).

3. Emotional and anatomic terms

In general, emotions may be defined as changes in bodily states in response to perceived or imagined stimuli (c.f. Damasio 1999:282). Such changes may be expressed as feelings or behavior. Characteristically, emotions are of brief duration as opposed to moods that are of relatively prolonged duration (Damasio 1999:341). Both emotions and moods are referred to as affects. The term ‘emotional expression’ refers to the origin of emotional feelings as subjective states that have emerged into conscious awareness from preconscious neuro-physiological processes. In contrast, the term ‘behavioral expression’ refers to a process that involves bodily movements or a process that may be perceived by an external observer. We assume with Damasio (2003:44) that the bodily sensations that accompany the functions of daily life, to which we are accustomed, constitute the feelings of a background or baseline emotional ‘state of being.’ We understand that perceived bodily threats and experienced physical needs elicit involuntary emotional feelings and behavior that interrupt this ongoing background emotional state. Such responses to emotion-inducing stimuli involve parallel changes in both the body and the mind. It is in this sense that we think of emotional responses as being embodied or ‘expressed’ by the body. We observe that embodied expressions of emotion have meaning, i.e., that emotional responses to extrinsic threats and intrinsic needs have embodied meaning for both the subject and the public (section 6). According to Damasio’s “somatic marker hypothesis,” the embodied meaning of emotional expressions or signs is marked by feelings of positive or negative valence. Such signs automatically indicate the importance of emotional stimuli for ongoing life-management (Damasio 2010:175). It has been observed that there is no single center for the processing of emotions in the brain. However, patterns of activity in well-defined regions of the brain have been associated with specific emotions (Levenson 2003; Damasio 1999:62).

3.1 Anatomic terms

The nervous system of communication and control of bodily functions is divided topographically into central and peripheral subsystems. By convention, the central nervous system is divided into three parts: the brain, the brainstem and the spinal cord. The brainstem mediates between the brain and spinal cord, both anatomically and functionally. The peripheral nervous system is subdivided functionally into somatic, autonomic and neuroendocrine systems. The somatic nervous system controls the voluntary actions of striated (skeletal) muscle. The autonomic nervous system controls the involuntary actions of smooth (visceral) muscle. It also innervates the neuroendocrine system, comprised of exocrine glands that release clear fluids onto body surfaces and endocrine glands that release hormones (molecular messengers) into the bloodstream for wide distribution throughout the body (Jänig 2003:140).

3.2 The central nervous system

The brain may be thought of as a tripartite structure consisting of the cerebrum located above the brainstem, the cerebellum located behind the brainstem, and midbrain formations that mediate among the three. The cerebrum is divided by a surface fissure into bilaterally symmetric and intercommunicating hemispheres. Topographically, each cerebral hemisphere is subdivided by a surface fissure into an anterior part (the frontal lobe) and a posterior part that is itself subdivided into three lobes: the parietal and occipital lobes arranged from front to back, and a temporal lobe that extends anterolaterally. The frontal and temporal lobes play fundamental roles in the production of cognitive and emotional responses to auditory stimuli (sections 5 and 6). Each cerebral hemisphere is covered by a thin, multilayered structure termed cerebral cortex. Each layer of the cortex is made up of nerve cells (neurons) of a predominant type that communicate with other neurons in the same layer and in other layers to form a vast three-dimensional neuronal network (Fuster 2003:62). The thalamus is a midline-symmetric subcortical structure that relays auditory signals to emotional and cognitive processing stations in the temporal and frontal lobes (sections 5 and 6). The hypothalamus is also a midline subcortical formation located below the thalamus and above the brainstem. This signal processing station relays emotive signals to the adrenal and pituitary glands that participate in the formation of emotional responses by the release of emotion motivating hormones into the bloodstream (section 6.1). Bilateral symmetry prevails throughout the central nervous system for both cortical and subcortical structures. Hence, in speaking of bilateral neural structures and pathways, it is customary to use singular terms. However, gross anatomic symmetry frequently coexists with functional asymmetry. For example, there is left cerebral hemisphere dominance for the control of speech (Fuster 2003:184).

3.3 The autonomic nervous system

The autonomic nervous system is functionally divided into at least two subsystems: the sympathetic and parasympathetic nervous systems. It is commonly thought that these two systems evolved to have antagonistic functions (Jänig 2003:142). Specifically, many believe that the sympathetic system evolved to provide automatic defensive responses to imminent threats, whereas the parasympathetic system evolved to neutralize such responses. However, in section 8, we propose a functional-evolutionary model, based upon empirical findings, in which the sympathetic and parasympathetic systems function as complementary mechanisms of defense against bodily threats that are perceived either at a distance from or in contact with the body. In fact, the autonomic nervous system, as a whole, is a mechanism for both self-preservation from physical threats and for the satisfaction of physical needs. Ultimately, this concept of self-maintenance and defense will contribute to an understanding of emotional responses to music!

4. The auditory detection pathway

Music is organized sound that is produced by the selective vibration of material objects which excite pressure waves in the air. These pressure waves are then perceived as sound by a complex biophysical process involving the transformation of continuous sound waves into discrete electrochemical signals in the brain. In this section, we present a survey of the physical processes involved in the detection of auditory stimuli. Sound waves entering the external auditory canal (the outer ear) excite sympathetic vibrations in a membrane (the eardrum) that occludes its inner end within the temporal bone of the skull. These vibrations are then transmitted, modulated, and focused by a chain of ossicles contained in an adjacent chamber within the temporal bone (the middle ear). This mechanical linkage, in turn, excites similar vibrations in a membrane that covers a ‘window’ into the cochlea, a structure contained in a deeper chamber (the inner ear) within the temporal bone. The cochlea is essentially a tapered membranous tube that is coiled, filled with fluid, and enclosed within a bony shell. The only openings in the shell are two membrane-covered ‘windows’ at its base. One window transmits vibrations from the ossicle chain in the middle ear to the cochlea by creating pressure waves in the cochlear fluid. The other window prevents pressure overload within the cochlear shell. The fluid-filled space within the cochlea is partitioned by a longitudinal membranous band (the basilar membrane), with its basilar attachment between the two cochlear windows. Pressure waves in the cochlear fluid excite surface waves on the basilar membrane such that each point along the length of the membrane vibrates at a characteristic frequency. This frequency varies from a maximum near its basilar attachment to a minimum at its apex. Hence, the cochlea functions as a “mechanical frequency analyzer” that separates and orders the different frequency components of the incoming auditory signal (Schnupp, et. al. 2011:57). The next phase in the processing of auditory stimuli involves the transduction of the mechanical vibrations of the basilar membrane into neural signals, i.e., the transformation of a continuous process into a discrete process. Specialized cells (hair cells) within a functional unit attached to the basilar membrane “…convert the mechanical vibrations of the basilar membrane into a pattern of excitation that can be encoded [as signals] by sensory neurons in the spiral ganglion of the inner ear for transmission to the brain” (Schnupp, et. al. 2011:64). The axons of these neurons form the auditory nerve that runs through a canal (the internal auditory canal) within the temporal bone to the brainstem. In the brainstem, an ascending auditory pathway is formed by a sequence of bilateral intercommunicating signal processing stations that lead to bilateral relay stations within the auditory regions of the thalamus (Schnupp, et. al. 2011:89). By the account of LeDoux (1996:164), sensory signals that reach the thalamus are relayed along parallel and intercommunicating pathways for cognitive and emotive processing in both cerebral hemispheres. The frequency-place or tonotopic organization in the basilar membrane of the cochlea is preserved in the ascending auditory pathway and extends at least as far as the auditory cortex of the temporal lobe (Schnupp, et. al. 2011:89-90).

5. The cognitive auditory pathway

Following the detection phase of auditory perception, neural signals are relayed directly from the thalamus to the auditory cortex of the temporal lobe for cognitive processing. Parallel pathways for the localization and identification of the sources of auditory stimuli that originate in the auditory cortex ultimately converge on the prefrontal cortex that covers the anterior surface of the frontal lobe (Schnupp, et. al. 2011:170-171; Alain, et.al. 2001). Analogous localization and identification pathways that follow a detection pathway have been described for visual perception (c.f. Cantor 2014, for references). It is thought that all sensory information that arrives at the prefrontal cortex is integrated in short term (working) memory before it is represented in conscious awareness, i.e., in immediate consciousness (LeDoux 1996:278-280; 2002:192-193). Following this, information may be relayed to the somatomotor cortex in the frontal lobe that controls voluntary body movements or to specific areas in the frontal and parietal lobes that are involved in speech production (Broca’s area) and speech comprehension (Wernicke’s area) (Schnupp, et. al. 2011:160). In this way, overt cognitive responses to musical stimuli may be expressed vocally or behaviorally.

5.1 Overt cognitive responses to music

Overt cognitive responses to musical stimuli tend to be imitative representations of perceived musical qualities. Meyer (2001:352) has referred to this type of behavioral response as “performative empathy, a kind of imitative identification with the qualities and patterns of the music.” Musical beat (pulse) may be represented by regularly repeating hand or arm movements, head nodding, body swaying or foot tapping, depending on situational constraints. Musical rhythm may be represented by repeating patterns of accented and unaccented body movements, as in dancing. Melodic contours may be represented by coordinated hand and arm movements as in ‘conducting.’ A listener may also attempt to represent musical qualities by vocalization, singing or metaphoric speech, as in music criticism and theory.

5.2 Covert cognitive responses to music

The concept that conscious awareness emerges spontaneously following the integration of sensory information in working memory suggests that there exists an ongoing preconscious or unconscious state of ‘awareness.’ The process by which such covert awareness spontaneously and involuntarily enters conscious awareness has been variously referred to in different contexts as “abduction” (Peirce 5.181), “illumination” (Hadamard 1954), “tacit knowledge” (Polanyi 2009), etc. It is likely that covert preconscious cognitive processing is a precursor of all conscious thought and, in the present context, of cognitive responses to musical stimuli.

6. Emotive auditory pathways

At least two parallel and intercommunicating pathways are involved in the processing of signals representing emotive auditory stimuli (LeDoux 2002:206-207). One pathway produces emotional reactions (objective emotional responses) while the other produces emotional feelings (subjective emotional responses). Objective emotional responses are displayed publicly as behavior and physiology. Subjective emotional responses are experienced privately. Both objective and subjective responses are spontaneous and involuntary. Both types of response result from the activation of a neural pathway leading directly from the auditory thalamus to the amygdala, a subcortical formation in the medial aspect of the temporal lobe that contains the hub of an emotion processing network. This processing center plays a major role in expressions of fear (LeDoux 1996:174).

6.1 Emotional reactions

Emotional reactions are controlled by the autonomic nervous system in coordination with the neuroendocrine system. These two communication and control systems regulate all involuntary bodily functions (Jänig 2003:136). The autonomic nervous system is activated by emotive signals from the amygdala that are relayed by the hypothalamus to target organs throughout the body.

6.2 Emotional valuation

Emotional valuation is a subjective judgment of the relevance of perceived emotional stimuli to bodily defenses and needs. Emotional valence is usually represented on a binary scale, commonly expressed as negative (unpleasant) or positive (pleasant). There are indications that the processing of strong negatively valenced stimuli occurs preferentially in the amygdala, whereas the processing of strong positively valenced stimuli occurs preferentially in the nucleus accumbens, a formation located in the base of the frontal lobe, in proximity to the amygdala. This signal processing station plays a major role in behavior directed toward appetitive or incentive stimuli, i.e., in expressions of desire or need (Zatorre & Salimpoor 2013; Lane 2000:362). Characteristically, behavioral responses in emotions with positive valence involve approach or seeking. Behavioral responses in emotions with negative valence tend to involve withdrawal, hiding or aggression.

6.3 Emotional meaning

Emotional behavior, feelings and valuation pathways, together with long-term memory pathways, ultimately converge in the prefrontal cortex where they provide inputs to working memory and immediate consciousness (c.f. LeDoux 1996:201). According to Joseph LeDoux (2002:206): “…information received by sensory systems activate emotion processing circuits, which evaluate the meaning of the stimulus input and initiate specific emotional responses…” Hence, from a semiotic perspective, emotional responses to musical stimuli are embodied expressions that constitute the representamina of emotional signs with valuations as emotional interpretants.

7. Objective emotional responses

We have seen that emotional responses to sensory stimuli interrupt the state of background emotional awareness of the perceiver (section 3). Such changes are initiated by surges of autonomic activity. We recall that the autonomic division of the peripheral nervous system has at least two sub-divisions: the sympathetic and parasympathetic nervous systems. Generally, the effects of sympathetic activation are widespread, whereas the effects of parasympathetic activation tend to be localized (Jänig 2003:141).

7.1 Motor responses to emotional stimuli

The two autonomic divisions have opposite effects on cardiac pacemaker cells. Sympathetic activation produces a rapid increase in heart rate, whereas parasympathetic activation produces a decrease. These effects are accompanied by similar changes in breathing rate as regulated by a pneumotaxic control center in the brainstem. In some target organs, the two autonomic sub-divisions have opposite motor effects, i.e., contraction and relaxation. Such effects are produced in the walls and sphincters of most anatomic conduits (the airways and gastrointestinal tract, as well as in the blood vessels of the face and erectile tissues (Jänig 2003:141). However, in other target organs, motor units are affected in only one way by only one autonomic division. Such effects occur in the dilator and constrictor muscles of the pupil, in most skeletal muscles, in piloerector (hair-raising) muscles, as well as in the coronary arteries, blood vessels in the skin of the trunk and limbs, in the viscera and in the reproductive organs). Hence, all rapid and involuntary motor responses to emotional stimuli are mediated by the autonomic nervous system.

7.2 Exocrine responses to emotional stimuli

Parasympathetic stimulation of exocrine glands (lachrymal, salivary, nasopharyngeal, digestive and intestinal glands) generates watery secretions that are released onto external surfaces of the body and internal surfaces of hollow viscera (Jänig 2003:140).

7.3 Regulatory responses to emotional stimuli

It has been proposed that transient emotional reactions and subsequent emotional actions are regulated by distinct arousal and motivational systems. Neural outputs of the amygdala that elicit emotional responses are potentiated by an arousal system. This purely neural activating system is controlled by specialized cells that are located in the brainstem, which modulate neural activity in widespread regions of the cerebral cortex (LeDoux 1996:287-289; 202:58). Neural outputs from the amygdala that elicit emotional actions, e.g., ‘fight’ or ‘flight’ behavior, are regulated by a motivational system consisting of two distinct neuroendocrine subsystems with control centers in the hypothalamus (section 3). These systems stimulate the adrenal glands to release functionally specific hormones into the blood stream. These hormones modulate autonomic defensive reactions to emotional stimuli and regulate the metabolic processes that sustain voluntary emotional actions (c.f. LeDoux 1996:208; 2002:223).

8. Functional-evolutionary perspectives

We have seen that the autonomic nervous system rapidly mobilizes defensive responses to threatening stimuli and activates the pursuit of appetitive stimuli. Hence, the two divisions of the autonomic nervous system (the sympathetic and parasympathetic systems) sustain the drive for self-preservation, i.e., for the defense, maintenance and reproduction of the body. Responses to autonomic activation present as embodied signs. Defensive and appetitive emotional signs are communicated by the body to self-awareness as valence or feelings, and to the awareness of others as behavior or functions. Interactions between the body and its environment result in a continuous adjustment of ongoing autonomic activity to meet the demands of self-preservation (Öhman & Wiens 2003:256). Defensive responses to external threats may involve removal of the body from the threat (withdrawal), removal of the threat from the body (expulsion), or neutralization of the threat by deception or destruction. All body surfaces that are exposed to the environment have protective coverings: skin for external surfaces and membranes for internal surfaces. The skin is directly exposed to environmental influences. The corneal membrane of the eye and the mucosal membranes that cover the inner surfaces of the digestive, respiratory and excretory tracts are intermittently or indirectly exposed to environmental influences.

8.1 Sympathetic responses

So-called ‘goose bumps’ on the skin (usually accompanied by subjective ‘thrills’) are produced by contraction of muscle units attached to hair follicles in response to surges in sympathetic neural activity. In fur-bearing animals, this pilomotor reflex enlarges the visual silhouette, thus simulating a looming counter-threat (c.f. Darwin 1998:106). In humans, ‘goose bumps’ and subjective ‘thrills’ occur as expressions of fear, in anticipation of seen or unseen threats. Defensive autonomic functions also involve the removal of noxious agents and foreign material from external and internal surfaces of the body. Activation of the sudomotor (sweat activation) reflex by sympathetic neural activity releases sweat onto skin surfaces where its evaporation dissipates excess body heat, either absorbed from the environment or generated by the increased metabolic activity that sustains intense motor activity. It is likely that the primary biological function of the piloerection reflex is to enhance the insulating characteristics of fur or hair by the formation of an insulating layer of air, which covers the skin. However, in the course of human evolution, development of the sudomotor reflex for the dissipation of excess body heat was probably accompanied by the loss of body hair, since the presence of hair retards the process of sweat evaporation, which limits its effectiveness. In compensation for loss of the insulating properties of body hair, reflex constriction of blood vessels in the skin became the primary means of conserving body heat. We note that all the above reflexes, which are involved in the maintenance of thermal equilibrium within the body, are activated by the sympathetic nervous system. Hence, these reflexes originated as defenses against threats to disrupt the thermal equilibrium within the body, either from the outside or the inside. This accounts for the fact that, in humans, the same sympathetic response to external threats (‘goose bumps’) is also provoked by exposure to a cold environment. In response to imminent threats, surges in sympathetic neural activity produce increased cardiac output accompanied by constriction of blood vessels in the skin (with resultant blanching), which diverts blood flow to the brain and skeletal muscles in preparation for defensive action.

8.2 Parasympathetic responses

In response to internal threats, parasympathetic neural activity initiates the release of clear secretions onto the surfaces of all visceral conduits from underlying exocrine glands, i.e., from lachrymal, salivary, tracheobronchial, digestive and intestinal glands. Parasympathetic activity also stimulates the motility of visceral conduits. Hence, there is a synergy between parasympathetic motor and exocrine responses that facilitates the removal of injurious agents from exposed internal surfaces of the body. The defensive function of exocrine secretions is enhanced by the antibiotic action of lysozyme that is present in tears, saliva and gastrointestinal secretions (c.f. Provine 2012, on emotional tearing). Hence, in response to aversive stimuli, both the sympathetic and parasympathetic nervous systems function as involuntary defense mechanisms. Also, in strong responses to appetitive stimuli, e.g., arousal of the desire for food or sex, both autonomic systems produce their expected cardiovascular, muscular or exocrine effects. However, the production of saliva and blood flow to erectile tissues are selectively increased by a surge in parasympathetic activity that facilitates self-maintenance and ultimately self-propagation. Similarities in behavioral expressions of rage and lust are explained by their common origin in augmented parasympathetic activity.

9. Autonomic responses to musical stimuli

In a comprehensive review of emotional responses to musical stimuli, Hodges (2010) notes that such responses may be classified as either overt or covert, depending upon their accessibility to direct or instrumentally mediated perception. Examination of these findings suggests that the occurrence of emotional responses to music depends largely upon the attentional focus, the emotional state and the native emotional sensibility of the listener, i.e., extra-musical factors. There are commonalities between reported emotional responses to musical stimuli and autonomic responses to environmental stimuli.

9.1 Overt autonomic responses to music

Overt sympathetic responses to music include: preprogrammed facial expressions (smiling, frowning) due to selective contraction of facial muscles, bodily ‘freezing’ that accompanies intensely focused attention, expressive body movements with rhythmic entrainment (c.f. McNeill 1995), and ‘goose bumps’ accompanied by tingling sensations variously referred to as ‘thrills,’ ‘shivers,’ or ‘chills.’ Overt parasympathetic responses to music include: blushing of facial skin and emotional tearing or weeping.

9.2 Covert autonomic responses to music

Covert sympathetic responses to music include: increased heart and pulse rate with stimulative music, increased tension in skeletal muscles in preparation for movement, decreased skin temperature in hands and feet due to vasoconstriction, and increased skin conductance (electrodermal activity) secondary to the sudomotor reflex. Covert parasympathetic responses to music include: decreased heart and pulse rate with sedative music, and a sensation of ‘lump in the throat’ or choking due to transient contraction of smooth muscle in the upper airway.

9.3 Strong emotional responses to music

In an exhaustive review of the literature on ‘strong’ experiences with music, Gabrielsson (2010) reported relative frequencies of objective emotional responses by the listener, i.e., responses to musical stimuli that present as observable signs or self-reports of subjective signs. In his study, Gabrielsson reported that the most frequent responses were observed as tears and felt as ‘thrills.’ Behavioral responses were either active or inactive. Active responses included: clapping the hands, rhythmic movements of the body, dancing and singing. In contrast, an inactive reaction was to become motionless and silent. This was interpreted as being due to either intensely focused attention or an attempt to conceal strong feelings in a social setting. In the following sections, we propose an explanation for these phenomena that is based upon principles of biological evolution, autonomic physiology and musical semiotics.

10. Exaptation of autonomic responses

Biologists traditionally use the term adaptation in reference to features of organisms that were shaped by natural selection for their current use. In contrast, Gould and Vrba (1982) propose the term exaptation in reference to ‘a character shaped by evolution for a particular use (an adaptation) that is coopted for a different use.’ As examples of this process, these authors suggest:

Another example, which pertains specifically to this study, involves the sensory and motor systems that control finger, hand and arm movements. These systems were initially adaptations for the performance of human survival functions (self-maintenance and self-defense) that later became exaptations for performance of the complex movements involved in playing musical instruments. Based on these examples, it is not unreasonable to suspect that the surges of autonomic activity which produce strong emotional responses to music are exaptations of autonomic functions that were initially shaped by evolution in response to aversive and appetitive stimuli. Hence, increased and decreased heart rate, blanching and blushing of facial skin, ‘thrills’ and tears, etc., in response to musical stimuli are probably exaptations from phylogenetically determined primary emotions, which are produced by sympathetic and parasympathetic autonomic activity. We note that emotive exaptation is fundamentally a semiotic process within the embodied self that is used for deception as well as for information in response to novel or unexpected environmental stimuli. In subsequent sections, we attempt to account for commonalities of expression and meaning between emotional responses to musical stimuli and emotional responses to aversive or appetitive stimuli.

11. On the meaning of emotional responses to music

In his classic work, “Emotion and meaning in music,” Leonard B. Meyer (1956) presents a theory of emotion based upon a concept derived from the thought of John Dewey. Meyer assumes a “law of affect” in stating that “Emotion or affect is aroused when a tendency to respond [to a stimulus] is arrested or inhibited” (Meyer 1956:14). Furthermore, he states that music “…activates tendencies, inhibits them, and provides meaningful resolutions for them” (Meyer 1956:31). For Meyer, ‘tendencies’ may be thought of as unconscious habits or conscious expectations (Meyer 1956:30). He observes that music may be experienced as “fraught with powerful uncertainty” (Meyer 1956:28), that things acquire meaning only in relation to other things (Meyer 1956:34), and that “Embodied musical meaning is [fundamentally] a product of expectation” (Meyer 1956:35) . In a later development of his thesis, he emphasizes the fundamental role played by uncertainty in the formation of emotional responses, in general (Meyer 2001:15.2.2).

In section 9, we have seen that strong emotional responses to music, i.e., responses that present as objective or overt signs are produced by surges of activity in the autonomic nervous system. However, the association between musical stimuli and autonomic responses remains to be explained. It has been demonstrated that activation of the autonomic nervous system produces automatic, i.e., involuntary motor and exocrine responses that prepare the body for defense against imminent threats and for the satisfaction of bodily needs (section 8). Hence, the autonomic nervous system has evolved as a means of internal communication and control that sustains the most essential bodily functions: those of self-preservation and self-propagation. Such preprogrammed responses are always accompanied by an uncertainty of outcome. They may be referred to as anticipatory signs of ‘fear’ and ‘desire.’ It is likely that the feelings of emotions evolved as subjective signs that enable the subject to discriminate between emotional stimuli and at the same time interpret their significance by an emotional assessment. Such emotional discriminations and valuations serve as preparations for subsequent conscious behavioral responses. From a biological perspective, the emotion of ‘fear’ is provoked by the anticipation of a possible physical injury. In contrast, the emotion of ‘desire’ is provoked by the experience of a physical need. We are now prepared to ask: What is the commonality between listening to music and facing the challenges of self-preservation that led to the exaptation (section 10) of autonomic responses from the domain of biology to that of aesthetics? From what we have seen, the commonality appears to involve uncertainties in the possible outcomes of anticipated or experienced events.

12. A simplified approach to Meyer’s theory of meaning in music

In a theoretical study of conceptual and emotional meaning in music, Leonard B. Meyer (1973) identified three types of tonal-temporal relation as sources of intrinsic meaning in music. Meyer referred to these relations as conformant, hierarchic and implicative. Conformant relations are based on similarity, hierarchic relations on containment (c.f. Simon 1996:184), and implicative relations on stylistic convention (Meyer 1973:44, 80, 110). There are obvious similarities between Meyer’s typology of meaningful musical relations and each of the following three typologies:

We note that the concepts of resemblance and similarity entail a commonality of features, the concepts of contiguity and containment entail commonality of place, and the concepts of custom, imputation and habit entail commonality by convention. Hence, all four typologies may be regarded as different presentations in different contexts of Peirce’s universal typology of Categorical relations which, for simplicity, may be expressed as: similarity, opponency and conventionality.

In Meyer’s typology of musical relations, an implicative relation in tonal music was defined using the terminology of deductive inference that suggests a relation of necessity, which may lead to misunderstanding. Here, we propose a simplified concept of musical implication that has been derived from Meyer’s original statements. This simplification will then be used to account for the likely processes by which strong emotional responses to music evolved from autonomic functions that originated as adaptations to cope with environmental challenges:

Furthermore, “[Musical] implications are guesses (feelings) about how present patterns will be continued and perhaps reach closure” (Meyer 2001:346). Assuming this concept of prospective musical implication, if an antecedent event induces a degree of uncertainty in the listener, then its consequent may induce a greater or lesser degree of uncertainty.

13. Speculations on ‘thrills’ and tears

Uncertainty may be used as the basis for a unified understanding of some embodied emotions. Humans possess a native drive to preserve a steady state of being, both in body and in mind. The capacity to anticipate future outcomes of present events serves to maintain this steady state. Awareness of uncertainty in anticipated outcomes of perceived events perturbs the background emotional state of the individual, which is briefly replaced by an emotional response appropriate for the situation. Previously, we have seen that uncertainties in the outcomes of anticipated events that are perceived to be threats may induce a surge in sympathetic autonomic activity that is expressed by the pilomotor reflex. This change of state may be felt as a ‘thrill’ or seen as ‘goose bumps,’ facial pallor, etc. Such autonomic responses occur before the outcome of an anticipated threat. They tend to occur in situations involving risk or danger, and are provoked by a native fear of the unexpected. In contrast, uncertainties in the possible consequences of experienced events may induce a surge in parasympathetic activity that tends to be expressed as emotional tears or weeping, facial blushing, etc. Such autonomic responses tend to occur after the experience or recollection of an event that provokes fear of an enduring or irreversible change of state of the self or (by empathy) of others. For example, intolerable pain (anguish), irretrievable loss (despair), frustrated intention (anger), and unexpected incongruity (humor) may induce emotional tearing or weeping. It is also possible that weeping as an expression of joy, e.g., with the fulfilment of a strong desire, is an exaptation of the parasympathetic response to strong appetitive stimuli (c.f. section 8). We recall that sympathetic autonomic responses originally evolved as defenses against emotional stimuli perceived at a distance from the body, whereas parasympathetic responses originally evolved as defenses against emotional stimuli in contact with or experienced by the body (section 3.3). It follows that emotional ‘thrills’ and tears function as nonspecific signs of uncertainties in possible outcomes of threatening events or uncertainties in possible consequences of experienced events.

14. Conclusion

We have seen that responses of the body to perceived threats are expressed at body surfaces, i.e., at interfaces between the body and the outside world. Such expressions are involuntary functions of the autonomic nervous system. Defensive responses at cutaneous surfaces are produced by the sympathetic nervous system. Defensive responses at surfaces of the eyes and at inner surfaces of the alimentary and respiratory tracts, both of which communicate with the environment, are produced by the parasympathetic nervous system. Hence, these two divisions of the autonomic nervous system play complementary roles in enabling the body to rapidly mobilize defenses against imminent external threats, as well as to prepare for the satisfaction of bodily needs. We recall that embodied emotions are subjectively experienced as feelings and objectively perceived as behavioral or physiological signs. For economy of terminology, we have referred to the category of emotional responses to threats as ‘fear’ and to the category of emotional responses to bodily needs as ‘desire,’ although responses in specific situations have specific names. From a semiotic perspective, emotions of fear and desire are expressed as embodied signs that may be interpreted either privately or publicly. Such signs are ‘anticipatory’ in that their interpretants involve possible outcomes of anticipated events or possible consequences of experienced events. Of particular interest in this study is the basic emotion of fear. It is understood that fear is a nonspecific response to uncertainty. It is recognized that embodied signs of emotion may present during events that appear to be unrelated to bodily threats. For example, strong emotional responses, as listed in section 9.3, may be provoked by any of the temporal arts, which include music, theater, film, dance, spectator sports as well as in moments of discovery. Based upon a functional-evolutionary argument, it is likely that embodied emotional responses to musical stimuli are produced by surges of autonomic activity, i.e., mechanisms that originally evolved for the purpose of defense against perceived threats. We have seen that strong emotional responses are provoked by the perception of threats at a distance from the body and expressed by surges in sympathetic neural activity. In contrast, strong emotional responses are also provoked by the perception of threats in contact with the body and expressed by surges in parasympathetic neural activity. We propose that at a later stage of human evolution, autonomic functions were exapted as responses to events with uncertain outcomes in general, including non-life-threatening events. By the exaptation of autonomic neural responses to non-life-threatening events, strong sympathetic emotional responses are provoked by uncertainties in the possible outcomes of anticipated events. In contrast, strong parasympathetic emotional responses are provoked by uncertainties in the possible outcomes of experienced events.

In this study, we have traced the embodied meaning of emotional responses to musical stimuli to its likely evolutionary origin: the exaptation of preprogrammed responses to environmental challenges. Hence, it is likely that embodied emotional responses to music, such as emotional ‘thrills’ and tears, are provoked by uncertainties in the possible outcomes of antecedent or consequent musical events.


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