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Music in the brain

Imagery and Memory

Rebecca S. Schaefer

The experience of music, like many other perceptual and cognitive processes, can be argued to largely take place internally; air pressure waves are not necessarily music until they hit the ear, get processed in the brain and body, and are interpreted as music. A special but recognizable case of an internal musical experience is one we generate ourselves as a musical image in the absence of sound, either deliberately or spontaneously. The music in our heads can come from an effortfully initiated and sustained mental action, but often also arises automatically, either with or without some contextual link to other stimuli or situations we are exposed to, and in certain cases as part, and possibly in support, of other cognitive functions. In this chapter, I will discuss different types of imagery and their interactions with other cognitive functions, specifically aspects of memory, and their neural signatures. I will argue that a supportive (also termed constructive ) form of imagery is of crucial importance to the generation of perceptual predictions, which can be considered to be central to the cognition of music listening. Finally, implications for future research are considered.

Characteristics of Imagery

Imagery Modality

A rich history of research focused on mental imagery has shown that when we imagine something, the related neural processes are at least partly shared with those of actually perceiving or performing the same stimulus or action (for an overview, see Kosslyn, Ganis, & Thompson, 2001). This means that depending on the sensory type, or modality of the imagery—referring to whether the imagined stimulus is visual, auditory, tactile, olfactory, or some sort of movement—the neural areas that are involved in that specific type of actual perception or action are often found to also be active during effortful mental imagery in that modality, albeit often to a lesser extent. Additionally, modality-independent neural activation patterns have also been identified that may be related to imagery vividness (e.g. Daselaar, Porat, Huijbers, & Pennartz, 2010), indicating that the task of initiating and sustaining a mental image also has commonalities between modalities.

Although some studies have found correlations between self-reported abilities across modalities, often visual and auditory, suggesting a general, modality-independent imagery ability (Gissurarson, 1992; Halpern, 2015), there are also clear indications that deliberate imagery abilities are increased generally in specific modalities with related expertise, such as auditory imagery (musical and non-musical) in musicians (e.g. Aleman, Nieuwenstein, Böcker, & De Haan, 2000), or movement imagery (Isaac & Marks, 1994) and spatial imagery (such as mental rotation, Ozel, Larue, & Molinaro, 2004) in trained athletes. It may also be the case that certain imagery modalities are more intricately related to each other than others. For instance, perceptual-imagery modalities may be more related to each other than to kinesthetic movement imagery, or certain modalities may share specific aspects (e.g., the time dimension necessarily present in both movement and auditory imagery; see Schaefer, 2014a for further discussion).

Reviewing the available experimental findings of imagery specific to varying kinds of musical stimuli, Hubbard (2010) concluded that many structural and temporal features of music appear to be preserved in musical images, and—importantly, in the context of the current discussion—can influence concurrent perception as well as expectations of upcoming stimuli. Crucially, the to-be-imagined stimuli need to be prescribed in experimental investigations of musical imagery quality in order to be able to make any kind of judgment about its structural properties, as improvised imagery is generally unpredictable and thus more difficult to evaluate in a lab setting.

Imagery Type

Although researching mental imagery is in itself a complicated endeavor because of difficulties in experimental measurement of subjective experience, the effort of clarifying its mechanism is also sometimes hampered by varying use of similar terminology. The many activities or tasks that all may be called “imagery” in the scientific literature include generating a maximally vivid image, but also the ability to manipulate an image, or to compare it—while keeping it in working memory—with perceptual input. Other, somewhat more abstract types of imagery include sensory expectations of actions; mental rehearsal in experts; perceptual judgments that are thought to involve some sort of inner representations and many more. Specifically relating to music, imagery processes have been put forward as being crucial to active music listening (Hargreaves, 2012), arguing that inner representations play a central role in the experience of perceiving music. Moreover, imagery skills are often mentioned in the context of musical aptitude (Gordon, 1965). However, in this last case, as well as in music listening, it is not always clear which kind of imagery is meant; and it is likely that multiple aspects of imagery are relevant to both music perception and production (see for instance the imagery thought to be crucial in joint music making, cf. Keller, 2012).

One way to be more precise about the specific role of imagery in these functions is to make distinctions between different types of imagery, which could potentially apply to any sensory modality. Following a taxonomy based on Strawson’s (1974) philosophical essay “Imagination and Perception,” and further developed for music imagery specifically by Moore (2010), one can argue that the type of imagery most often investigated experimentally is sensory imagery, where someone deliberately imagines a sound (or some other sensory experience). Conversely, a different type of imagery, referred to as constructive imagery, does not take place deliberately, but functions more as a process of perceptual organization, and appears to be more in line with how imagery processes are thought to support music perception. The idea that something like imagery is inherently involved in any perceptual process that involves prediction of upcoming stimuli is not exclusive to music listening (cf. Clark, 2012), and also fits with the concept of modality-specific grounding of conceptual knowledge (cf. Barsalou, Simmons, Barbey, & Wilson, 2003). Given its hypothesized involvement in actual perception, constructive imagery has been put forward as a likely explanation for the shared aspects of perception and sensory imagery represented by overlapping neural activation patterns that have been identified between these two tasks through brain imaging findings (for further discussion of this idea, see Schaefer, 2014b). As the concept of constructive imagery needs further exploration, this assertion remains speculative for now, but it is clear that more insights from research into the different imagery types will not only inform our knowledge of music processing but of cognitive functioning more generally.

Imagery Content

While the different modalities of imagery have long been distinguished, and the type of imagery is now also receiving more attention from researchers, the specific content of imagery itself within a certain modality is now also within reach of experimental paradigms, including those paradigms using brain measurements. Meticulous behavioral experiments have been able to isolate specific aspects of imagery content, for instance by looking specifically at pitch, contour, rhythm, timbre, loudness and so on in the context of music, either in isolation or combined, as a rich, holistic representation (cf. Hubbard, 2010). Using careful experimental design in combination with specific analysis methods that allow exploration of the information content of neural activity, we can now also distinguish brain activity patterns that are related to specific stimuli (or aspects of these stimuli) or actions. Examples come from different modalities, and include being able to distinguish between specific imagined visual images (e.g. cf. Cichy, Heinzle, & Haynes, 2012) or actions (Oosterhof, Tipper, & Downing, 2012), or isolating the neural activity related to specific melodic aspects of an imagined tune (e.g. Schaefer, Desain, & Suppes, 2009).

Spontaneous Imagery

Even though most experimental evidence focuses on effortful, deliberate imagery, there are many ways in which a mental image can surface into consciousness spontaneously. In the musical domain, a significant body of work focuses on spontaneously experienced music (also termed “earworms,” or INMI, for Involuntary Music Imagery), which has been investigated in a very large sample through a self–report questionnaire (Floridou, Williamson, Stewart, & Müllensiefen, 2015) and now also includes an exploration of brain differences for people who frequently experience spontaneous music imagery (Farrugia, Jakubowski, Cusack, & Stewart, 2015). Although early work on this phenomenon had already shown that music students experience music imagery more than a third of the time (Bailes, 2007), this experience is extremely widespread. In a sample of 2671 individuals, only 29 reported never experiencing INMI, giving an impression of the ubiquity of this experience. Unsurprisingly, INMI frequency was shown to correlate positively with musical behaviors (i.e. musical training, active engagement with music, perceptual and singing abilities, and emotional responses to music). Furthermore, INMI frequency showed substantial correlations with other kinds of spontaneous (or even intrusive) thought, such as mind wandering or daydreaming (Floridou et al., 2015).

Although there are indications that spontaneous imagery implicates similar brain areas as effortful imagery (Farrugia et al., 2015), it is likely that the cognitive aspects of involuntary imagery relate to low-attention states, and do not necessarily depend on deliberate concentration or attentional functions, or at least interact with these functions in a different way. Different aspects of the INMI experience, identified through the factor structure of the questionnaire created by Floridou et al. (2015), and found to have specific structural neural correlates (Farrugia et al., 2015), may also differentiate between the varying subjective experiences of INMI, from the annoyance of an unwanted song in your head to a helpful function that can support one’s movements, reflections, or focus. Considering imagery more broadly, there are indications that individual differences in frequency of spontaneous imagery occur in visualization as well (Nelis, Holmes, Griffith, & Raes, 2014).

Applications

When considering the potential applications of imagery, the most obvious areas of interest are clinical settings and pedagogy. In clinical settings, specifically visual imagery is increasingly utilized to achieve experiences that support healthy functioning, potentially through imagined rehearsal of that experience (for an overview, see Pearson, Naselaris, Holmes, & Kosslyn, 2015). Furthermore, there are indications that actively imagined music or singing may be useful in movement rehabilitation to regularize movement, effectively functioning as a self-generated movement cue (Satoh & Kazuhara, 2008; Schaefer, Morcom, Roberts & Overy, 2014).

Imagined rehearsal also supports learning in non-clinical settings, specifically acquiring motor skills such as those used in athletics and surgery (cf. Cocks, Moulton, Luu, & Cil, 2014), as well as music learning (cf. Holmes, 2005). Although the development of imagery techniques is not often explicitly taught in music pedagogy (Clark & Williamon, 2011), there are clear instances of expert musicians using, and benefiting from, imagery techniques that support memorization and performance (e.g. Davidson Kelly, Schaefer, Moran, & Overy, 2015). The way that imagery is approached in these applied settings— even though it does usually take the shape of deliberate, sensory imagery—differs from the way it is approached in most experimental research in several ways. Most saliently, it seems that in applied settings, rich, holistic, and thus multimodal images are encouraged, rather than focusing on a single stimulus aspect or modality. The goal of the strategy of employing rich images is to increase the effectiveness of the rehearsal method, thus directly tying in to learning and memory functions. The implication here is that the richer the image or rehearsed situation, the more effective the imagery intervention in terms of learning. As an example, consider the practice of creating a multimodal internal representation of a piano performance, which not only consists of an auditory memory of the piece but also includes visual aspects such as the score and the keyboard, and movement aspects of the actual performance. This approach, although requiring some extra memorization effort initially, can offer a much-reduced cognitive load to the pianist during performance, freeing up attentional resources for expressive communication, or other performance goals (Davidson Kelly et al., 2015).

Taken together, the main concepts to consider when discussing imagery are the sensory modality (or modalities) that are included in the image, the kind of imagery that is under discussion, and the content of the imagery, which is obviously related to the modality. These factors also determine the level of focus or engagement that is required to experience the imagined stimulus or action. Images can be made up of basic, low-level perceptual features up to complex, multimodal constructs or situations. Thus, imagery arguably does not exist as an isolated cognitive function, but is inherently related with other cognitive functions, as well as interrelated with perception and action.

Imagery and Memory

As already suggested above, memory processes can be argued to be a part of imagery processes, as well as vice versa. The former appears intuitive; to actively call up a vivid image requires the preexisting knowledge of the to-be-imagined stimulus and the capacity to retrieve and (internally) reproduce it, even when producing it spontaneously. Moreover, both long-term memory and working memory function are suggested to contribute to imagery vividness through increased representation strength (Baddeley & Andrade, 2000; Navarro Cebrian & Janata, 2010).

The other direction, where memory depends on imagery, is somewhat less straightforward, but is arguably dependent on the type of imagery and the type of memory under consideration. Although imagery strategies can drive memory encoding processes (Alonso et al., 2016), and possibly enhance learning through mental rehearsal (cf. Davidson Kelly et al., 2015), the imagery necessarily involved in long-term memory may be restricted to the moment of recall (and possibly dependent on modality, cf. Greenberg & Rubin, 2003), but thought of as crucial to vivid remembering (Huijbers, Pennartz, Rubin, & Daselaar, 2011). Looking more broadly at holistic, multimodal scenarios, findings have been reported indicating that remembering the past and imagining the future lead to activation in comparable brain networks (e.g. Schacter, Addis, & Buckner, 2007), in line with the idea that both remembering and simulating an experience involve mechanisms related to sensory imagery.

The commonalities between deliberate sensory imagery and short-term or working memory1 are further supported by more detailed research findings. For music specifically, similar neural activation features have been identified for effortful imagery and for modality-specific working memory in the electroencephalogram (EEG; cf. Schaefer, Vlek, & Desain, 2011) as well as in the brain responses measured by functional magnetic resonance imaging (fMRI; cf. Herholz, Halpern, & Zatorre, 2012). More recent results using visual and non-musical auditory stimuli indicate that the content of the memorized or imagined stimulus can be also detected from brain activity measurements (e.g. Linke & Cusack, 2015), although there appear to be differences in the extent to which low-level perceptual features are processed as compared to more abstract representations, with low-level features showing more overlap with actual perception than high-level features. This result is in line with an exploratory finding that the neural correlates for imagery of more complex musical material appear to overlap with perception of that same material to a lesser degree than for very simple stimuli (Schaefer, Desain, & Farquhar, 2013), suggesting that the more complex a stimulus is, the less its effortful imagination relies on the perceptual apparatus of the brain. Moreover, there are also distinct differences in neural activation patterns between imagery and short-term memory that indicate that, although there are great similarities in terms of the brain areas that are involved, there are also detectable differences between actively imagining a sound stimulus as vividly as possible, and keeping that stimulus in working memory for a change detection task (Linke & Cusack, 2015). Further investigations are necessary to better untangle the conceptual differences between active imagery and working memory, and to establish whether the same mechanisms apply to different modalities or complexity of imagery content.

Cognitive Overlap and Shared Brain Signatures

Focusing specifically on short-term memory rehearsal and effortful sensory imagination, we can identify clear commonalities in their functions and features, centered mainly on the conscious inner experience of a specific stimulus. This experience may emerge because we deliberately imagine it or, alternately, because we need to keep the information or stimulus active for later use, such as when rehearsing a phone number, while performing some sort of change detection task, or while manipulating information (for example, doing mental arithmetic). The key difference appears to be that in the second case, the imagery is conjured in support of a goal beyond simply simulating an experience. As such, the internal percept that subserves a memory function tends to be spontaneous, but conscious, whereas in deliberate imagery it is more effortfully initiated. In either situation, the core of the percept may have modality-specific and amodal aspects, i.e. characteristics related to the type of sensory information (or action), or to the concept of the object itself.

Research into the neural activation patterns related to amodal conceptual information (i.e. lacking a specific sensory modality), reports temporal gyrus and precuneus activation that follows the structure of behavioral ratings of similarity between concepts (Fairhall & Caramazza, 2013). This, however, differs from the activation patterns that are related to modality-independent components of active imagery, seen in frontal (superior middle frontal gyrus and medial prefrontal cortex) and parietal areas (lateral parietal and posterior cingulate, Daselaar et al., 2010). Although these two studies approached the analyses of brain activation patterns in quite different ways, and more findings on these kinds of tasks are needed before being able to reach more robust conclusions, these results suggest that mentally activating a concept for manipulation (as in the similarity judgment task used by Fairhall & Caramazza, 2013) and deliberately imagining a maximally vivid sound or visual image (as was required in Daselaar et al., 2010) do not necessarily involve very similar neural processes. Thus, although the activation of amodal concepts must be part of imagining, there may be different timecourses of activation depending on the task. As neither of these studies used musical material, these findings make little suggestion as to how the amodal aspects of music (i.e. representations of semantic or emotional content, episodic knowledge related to the musical piece, etc.) are combined with the auditory (and other sensory or motor) aspects of the imagined stimulus.

Future work on how imagery processes may support other cognitive functions, for instance as described for different types of memory, should yield illuminating results concerning the role of attention or focus during spontaneous or deliberate internal experiences, and whether inter-individual differences in deliberate imagery ability or spontaneous imagery affect these functions. In particular, the idea that spontaneous music imagery or visualization may support other functions, such as personal reflection or focus during low-attention states (as reported by Floridou et al., 2105) is very interesting. Here, the spontaneous inner experience may speculatively be thought to be mediated by individual traits relating to tendencies for spontaneous simulation, degrees in which associated concepts may trigger one another, specific strategies that have been acquired over time, and so on.

Considering musical imagery specifically, much debate has surrounded the findings of activity in brain areas traditionally known as motor areas during active imagery. The relatively stable findings that, in addition to activating secondary auditory areas, music imagery leads to activation in (pre-)supplementary motor areas (SMA) and sometimes premotor areas and cerebellum (Halpern & Zatorre, 1999; Herholz et al., 2012), was initially interpreted as covert movement imagery (Zatorre & Halpern, 2005). However, rather than music imagery necessarily including some form of movement representation, this may actually reflect a much broader function related to timing and sequencing, which are increasingly seen as more prominent functions of these neural areas, thus expanding their status from “only” motor areas to areas involved in general temporal organization of sensory as well as motor processing, specifically timing processes (See Henry and Grahn, this volume; also Schwartze, Rothermich, & Kotz, 2012; Teki, Grube, & Griffiths, 2011). As such, these areas may mainly be involved in processing temporal information relating to an imagined sound or action (Schaefer, 2014a). Of course this does not preclude the possibility that many cases of musical imagery involve both motor and sound aspects, either deliberately and consciously, or passively through neural coactivation, as tends to develop in expert musicians. Aspects of music processing as a whole, and indeed music imagery processes specifically, often do include some representation of movement; however, when the assumption of covert motor imagery in the form of subvocalisation was specifically tested, results did not support this conclusion and instead support a broader role for SMA than pure motor processing (Halpern, Zatorre, Bouffard, & Johnson, 2004). Moreover, this music imagery-related SMA activation is seen additionally to activation related to actual movement when imagining music while performing a simple wrist flexions (Schaefer et al., 2014), suggesting that this imagery-related activation of SMA takes place independently from movement-related activation, supporting Halpern et al.’s (2004) finding.

When we consider other functions that might recruit SMA during music imagery, there is actually a range of functions in addition to those associated with movement that are reported to be related to SMA activity, including sensory processing, word generation, and even working memory (cf. Chung, Han, Jeong, & Jack, 2005). As our concept of working memory is also still changing from that of an item-holding storage to an increasingly flexible function that is more dependent on salience, attention, and quality of the representation than thought before (e.g Ma, Husain, & Bays, 2014), our ideas of how imagery and memory interact will certainly develop further. As such, it can be argued that research into the interactions between imagery and memory allows not only for a paradigm for the investigation of amodal and modality-specific conceptual knowledge, but also a way to increase our understanding of the brain systems relating action and perception to cognitive function.

Mental Models and Perceptual Prediction

Predictive processes during music perception are thought to be at the core of our affective responses to music listening, and have enjoyed a rich history of theoretical development (cf. Meyer, 1956; Huron, 2006). The main concept which has been developed is that our expectations about where music may be going next—harmonically, melodically, rhythmically, etc.—relate directly to our emotional response to music, offering structure and regularity through predictable features, or surprise and potentially humor in more unpredictable transitions. More recent developments in thinking about predictive processing (or predictive coding) in the broader area of cognitive science (cf. Clark, 2013) can arguably be seen as an extension of these ideas, with specific implications for music processing (Schaefer, Overy, & Nelson, 2013).

At the basis of the framework of predictive coding as a theory of cognition, with clear parallels to the abovementioned ideas about music, lies the assumption that we create predictions, which are met or violated as new information comes in, which then lead to a response in the listener. In general perception, like in music, these predictions can be generated at multiple levels simultaneously, and vary greatly from low-level to high-level expectations. Arguably, a prediction can only be generated when an internal model exists, and while this internal model may be interpreted as constructive imagery (Schaefer, 2014b), there are many authors who frame this phenomenon as a learning mechanism, where statistical learning based on many stimuli creates a sense of what is predictable in the world around us and causes our predictions (musical and otherwise) to be based on what we have been exposed to before. Where deliberate, effortful imagery can be thought of as having direct commonalities with active working or short-term memory rehearsal, constructive imagery can be thought of as the outcome of statistical learning in the shape of internal models, on which perceptual predictions are based. Future investigations will have to further specify the potential differences between these functions, and the extent to which perceptual organization and statistical learning interact. However, the main implication is that imagery processes have been interpreted as impacting the same domains as not only explicit but also implicit memory and learning processes. Furthermore, the type of imagery considered to be crucial to the musical experience (cf. Hargreaves, 2012) can thus directly be interpreted as related to personal experience, mediated by all the associative processing gained through exposure to an auditory environment.

As mentioned previously, another way in which learning and memory functions may interact with implicit and explicit imagery processes is in specific types of expertise. As stated above, musical imagery abilities are often considered an important part of musicianship, and the relationship between modality-specific expertise and effortful imagery abilities also extends to other domains, such as athletics (cf. Ozel et al., 2004). When these findings on deliberate imagery are viewed as resulting from statistical learning (and potential concomitant increases in constructive imagery abilities), it may mean that constructive imagery and sensory imagery, although distinct functions, are actually part of the same system. From this perspective, the way that imagery is conceptualized and measured in an experiment becomes even more crucial in terms of interpreting the findings of effects of expertise on imagery abilities as result of a learning process. For instance, many musicians report that they imagine music while performing, and use this imagery as a guide for their performance, in terms of shaping their performance to produce a target sound, but also directing their attention to what is coming next, or to be aware of other parts performed by their collaborators (cf. Davidson Kelly et al., 2015; Keller, 2012). This imagery, which may have originated from deliberate practice, eventually happens spontaneously in experts, and supports expert performance.

Conclusions and Future Research

In sum, the available literature shows us that not only the modality and content of imagery, but also the type of imagery may vary, and that imagery functions interact with other cognitive functions. There are strong cognitive parallels between imagery functions and memory functions, where specific types of imagery closely align with specific types of memory. Although considerable overlap between the functional neural activation of various types of imagery and memory can be found in the literature, the studies directly comparing imagery to memory are relatively sparse. And while an exhaustive review of this literature was not the aim of this chapter, a clear need has been demonstrated for studies that delve deeper into the mechanisms of imagery to better understand the different imagery types, their interactions with other cognitive functions, and their applications. Topics such as the individual differences in imagery ability and potential effects of aging, the effects of focus or attention on imagery quality as well as the interactions with expertise, the degree to which modality-specific and amodal aspects of a construct impact the learning effects of mental rehearsal, and a range of other questions need to be clarified through future research in order to fully make use of the potential of imagery strategies for learning purposes. Thus, we may learn more about the impact that our internal images can have, on our daily experiences more broadly, and on musical processing specifically. A broad range of musical activities – listening, performing and composing – inherently make use of rich internal representations, and thus a deeper understanding of imagery processes promises to yield crucial insights regarding musical perception, action, and cognition.

Note

1. The terms “short-term memory” and “working memory” here refer to the maintenance and the maintenance plus manipulation of information, respectively.

Core Reading

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Herholz, S. C., Halpern, A. R., & Zatorre, R. J. (2012). Neuronal correlates of perception, imagery, and memory for familiar tunes. Journal of Cognitive Neuroscience, 24(6), 1382–1397.

Hubbard, T. L. (2010). Auditory imagery: Empirical findings. Psychological Bulletin, 136 (2), 302–329.

Keller, P. E. (2012). Mental imagery in music performance: Underlying mechanisms and potential benefits. Annals of the New York Academy of Sciences, 1252, 206–213.

Schaefer, R. S. (2014b). Mental representations in musical processing and their role in action-perception loops. Emprirical Musicology Review, 9 (3), 161–176.

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